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FOR THE YEAR

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VOL. LXXVI.

WITH FIFTEEN PLATES,
236 Text-figures.

SYDNEY:
PRINTED AND PUBLISHED FOR THE SOCIETY BY

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

and
SOLD BY THE SOCIETY.
1952.

CONTENTS OF PROCEEDINGS, 1951.

PARTS I-II (Nos. 353-354).
(Issued 31st May, 1951.)

Presidential Address, delivered at the Seventy-sixth Annual General Meeting,
28th March, 1951, by D. J. Lee, B.Se. The Problems of Insect Quarantine ..

Elections
Balance Sheet for the Year ending 28th February, 1951 ..

Serological Studies of the Root-nodule Bacteria. IV. Further Analysis of
Isolates from Trifolium and Medicago. By Hilary F. Purchase, J. M.
Vincent and Lawrie M. Ward ..

A Septoria Disease of Huphorbia peplus L. By Dorothy HE. Shaw. (Plate i and
one Text-figure. )

Preservation Techniques for Scarabaeid and other Insect Larvae. By P. B.
Carne

The Anatomy and Morphology of the Operculum in the Genus Eucalyptus.

Part I. The Occurrence of Petals in Hucalyptus gummifera (Gaertn.)
Hochr. By J. L. Willis. (Plates ii-iii and two Text-figures.)

Some Notes on Athrotaxvis. By Charles G. Elliott. (Communicated by Dr.
Patrick Brough.) (Sixteen Text-figures.)

The Paramphistomes (Trematoda) of Australian Ruminants. iPanvee
Systematics. By P. H. Durie. (Plates iv—v.)

Pages.

i-xix

xix

xx

1-6

1-25

26-30

36-40

41-48

CONTENTS.

PARTS III-IV (Nos. 355-356).
(Issued 17th September, 1951.)

A Review of the Australian Species of Sarcochilus (Orchidaceae). By H. M. R.
Rupp. (One Text-figure.)

Australian Rust Studies. VIII. Puccinia graminis lolii, an Undescribed Rust
of Lolium spp., and Other Grasses in Australia. By W. L. Waterhouse .

Celaenopsoides Gunther, 1942: A Synonym of Ophiomegistus Banks, 1914
(Acarina: Parasitidae). By Carl EH. M. Gunther

On Trombicula minor Berlese, 1905. By Carl E. M. Gunther. (Three Text-
figures. )

The EHvolution of the Radio-medial Area in the Wings of the Muscoidea
Acalyptrata (Diptera). By H. F. Lower. (Thirty Text-figures.)

Studies on Australian Marine Algae. VI. New Geographical Records of
Certain Species. By Valerie May ..

The Marine Algae of Brampton Island, Great Barrier Reef, off Mackay,
Queensland. By Valerie May ..

The Development of Bryozoan Faunas in the Upper Palaeozoic of Australia.
By Joan Crockford (Mrs. Beattie.) (Four Text-figures.)

Suecinoxidase of Potato Tuber. By Adele Millerd. (Two Text-figures.)

The Nomenclature of Heteronychus sanctae-helenae Blanchard (Coleoptera:
Scarabaeidae: Dynastinae). By EH. B. Britton. (Communicated by P. B.
Carne.)

Controlled Pollination of Hucalyptus. By L. D. Pryor. (Plate vi.)

A Genetic Analysis of some Hucalyptus Species. By L. D. Pryor. (Plates
vii—xi.)

Investigations on the Preference shown by Aédes (Stegomyia) aegypti Linn.,
and Culex (Culex) fatigans Wied., for Specific Types of Breeding Water.
By Tom Manefield

A Mite from a Beehive on Singapore Island (Acarina: Laelaptidae). By Carl
EH. M. Gunther. (Five Text-figures.)

ili

57-64

65

66-70

71-82

83-87

88-104

135-139

140-148

149-154

155-157

iv CONTENTS.

PARTS V-VI (Nos. 357-858).
(Issued 15th January, 1952.)

A Critical Consideration of ec-mitosis with Reference to the Effects of Nitro-
phenols. By Mary M. Hindmarsh, Linnean Macleay Fellow in Botany.
(Plate xiii.)

The Life-history of a Penaeid Prawn (Metapenaeus) breeding in a Coastal
Lake (Tuggerah, New South Wales). By Muriel C. Morris and Isobel
Bennett. (Plate xii and ninety-six Text-figures. )

The Use of Excised Shoots in Linseed Investigations. By H. B. Kerr.
(Plate xiv.>

Remarks on some Australian Laius Guér. (Col.: Malachiidae). By W.
Wittmer. (Communicated by J. W. T. Armstrong.) (One Text-figure. )

A Revision of Australian Species previously referred to the Genus Himpousca
(Cicadellidae, Homoptera). By Harold F. Lower. (Plate xv and seventy-

five Text-figures.)

Miscellaneous Notes on Australian Diptera. XV. Tabanus, Heteropsilopus.
By G. H. Hardy

Balance Sheets

Pages.

158-163

164-182

183-186

187-189

190-221

222-225

XXi-Xxli

Abstract of Proceedings BPE ks MRO WTS tet, RNIB ot We Cree Sle saa XX11i-xxvii

ASEOERAMEMDET Sh sacar a etater: Oe toa ee) eee MMO Riot AM Nes) epee VAC Stee ee ae CONV T < ReN ERG NT

List of New Genera and Species

List of Plates

XXXiil

XXXiil

Index saat Sash oath as, exe allteae ae) Oe Ea RRO 4 ee Se eee Vee

ANNUAL GENERAL MEETING.

WEDNESDAY, 28TH MArcH, 1951.

The Seventy-sixth Annual General Meeting was held in the Society’s Rooms, Science
House, Gloucester Street, Sydney, on Wednesday, 28th March, 1951.

Mr. D. J. Lee, B.Se., President, occupied the chair.

The Minutes of the Annual General Meeting held on 29th March, 1950, were read
and confirmed.

i PRESIDENTIAL ADDRESS.

As is usual, the first part of my address is devoted to a brief review of the Society’s
activities during the past year.

Volume 75 of the Society’s Proceedings was published during 1950. It consisted of
359 + xl pages, 12 plates and 442 text-figures, thus being a larger volume than the
previous one. Financial assistance towards the cost of publication of three papers was
received from Department of Anatomy, University of Sydney (£74 16s.), Commonwealth
Scientific Publications Committee (£55) and University of Melbourne (£25). The
annual grant from the State Government of £100 towards the cost of publication of the
Proceedings was increased to £150 during the year.

Exchanges received from scientific societies and institutions totalled 1,414. Loans
from the Library, particularly interstate inter-library loans, have been requested as in
the previous year. Additional steel shelving has been installed in the Library and some
duplicate books are still available for sale. New exchanges were commenced with:
Academy of the Popular Republic of Roumania; California Zoological Club; Hlisha
Mitchell Scientific Society; Estacao de Melhoramento de Plantas, Elvas, Portugal;
Société des Sciences Naturelles de Tunisie; University of Upsala (““Symbolae Botanicae
Upsalienses’”), and Victoria University College, Wellington, New Zealand. A number
of books from the library of the late Dr. F. G. Hardwick were presented to the Library
by his son, Mr. F. L. Hardwick.

Interesting programmes were given during the year at the following monthly
meetings:

June: A talk on “Coral Genera of Heron Island (Barrier Reef)”, by Mr. K. HK. W.
Salter and Miss M. A. Besley.

July: The following members contributed items of interest: Miss M. Hindmarsh—
a note on “Mitotic Poisons’; Mr. A. Musgrave—exhibition of specimens of Stephanitis
queenslandensis Hacker, 1927, a member of the family Tingidae (Lace Bugs); Mr.
A. K. O’Gower—exhibition of the eggs, larvae and pupae of cat fleas. ©

August—A series of talks was given by members of the New South Wales Depart-
ment of Agriculture, the speakers being: Mr. Grahame Edgar, Director of Veterinary
Research (investigations on problems of animal industry in the State); Dr. S. L.
Macindoe, Principal Research Agronomist (an account of the organization, progress and
results of plant research conducted primarily on the ten Experiment Farms and two
Agricultural Colleges of New South Wales); Dr. F. T. Bowman, Special Fruit Research
Officer (outline of the research work of the Division of Horticulture); Mr. S. L.
Allman, Senior Entomologist (description of the Entomological Branch and its work
with insect pest problems); and Dr. C. J. Magee, Chief Biologist (research in the
Biological Branch which is directed mainly at preventing losses from plant diseases).

September: Talks on “Recent Changes in the Vegetation of North-western New
South Wales’, by Dr. N. C. W. Beadle and Professor N. A. Burges.

October: Short talks by Professor P. D. F. Murray on “The Fusion of Long Bones”
and by Mr. B. R. A. O’Brien on “Some Aspects of Regeneration in Earthworms”’.

November: Talks on various aspects of Northern Australia were given by: Professor
J. Macdonald Holmes—“The Geographical Background to the Problems of North

A

ii PRESIDENTIAL ADDRESS.

Australia’; Dr. W. Kirkland—‘‘Medical Problems of North Australia”; and Dr. N. W. G.
Macintosh—“Comments on the Physical Types of the Aborigines of Arnhem Land”. A
talk prepared by Mr. W. Poggendort on “Problems in Agriculture” was given by
Mr. E. B. Furby.

We thank all who have contributed to these programmes.

Since the last Annual Meeting the names of fourteen membérs have been added to
the list, three members have been lost by death, and eighteen have resigned. The
number of members as at 1st March, 1951, is: Ordinary Members, 206; Life Members,
22: Honorary Member, 1; and Corresponding Members, 3—total 232.

As Dr. Dorothy Carroll tendered her resignation as Secretary to the Society as
from 22nd January, 1951, it was found necessary to make interim arrangements for
the successful prosecution of the business of the Society. To this end Council appointed
Dr. W. R. Browne as Honorary Secretary and Dr. A. B. Walkom as Honorary Editor
for the time being. It is hoped that Council will be able to decide on some more
permanent arrangement in the near future. Council’s intention at present is to employ
a part-time editor, if one can be found suitable and willing to undertake this responsi-
bility, and carry on the secretarial work with an honorary secretary and have the
assistant secretary in charge of the Society’s rooms and library. In view of a further
50% increase in the basic cost of publication of the Society’s Proceedings, Council has
felt that some economy in the cost of the secretarial activities of the Society may be
necessary in order to maintain the level of publication at one approximating that of
previous years.

Two Special General Meetings were held on 25th October and 29th November, 1950,
respectively, to make an addition to Rule VI as follows: “An Ordinary Member who has
paid the Annual Subscription for forty years shall be exempt from payment of further
subscriptions.” Council’s object was to relieve long-standing members of the burden
of their subscriptions, particularly in their years of retirement.

Dr. Ida A. Browne resigned as a member of Council in April, 1950. Council
expressed appreciation of her service on the Council during the past ten years.

Professor P. D. F. Murray was elected to. fill the vacancy on the Council caused
by the resignation of Dr. Ida Browne.

Dr. G. D. Osborne was granted leave of absence from the Council for six months
from ist January, 1951.

Dr. A. B. Walkom attended the Pan-Indian Ocean Science Congress at Bangalore
in January, 1951, as one of the Australian delegates.

Miss Mary Tindale was appointed the Society’s representative at the VIIth Inter-
national Botanical Congress in Stockholm, 1950, and accredited observer at the
International Union for the Protection of Nature Conference. Miss Tindale has informed
the Society of a number of points dealt with by the Nomenclature Committee and
offered to answer questions concerning special points in changes of the Rules.

The total net return from Science House for the year was £597. In future halt-
yearly meetings and half-yearly accounts and audits of the Science House Management
Committee will be abandoned and only annual meetings will take place. A survey of
the libraries of the owner-bodies of Science House has been made by the State Library
Board of New South Wales at the request of the Societies’ Library Committee, and a
report thereon was received.

Following a request from the Information Service, Commonwealth Scientific and_
Industrial Research Organization, which had resulted from one of the recommendations
of the Royal Society Scientific Information Conference (July, 1948), the Council
decided that a synopsis of each paper published in the Proceedings would appear below
the title of the paper. Such a synopsis will not interfere in any way with the arrange-
ment of the paper but will facilitate the reading of the papers in the Proceedings.
These authors’ synopses will be introduced in Volume 76 for 1951.

Preservation of Natural Areas—The Joint Committee of the Royal Zoological
Society and Linnean Society considered another survey, in this case at the Spencer’s
Creek Dam site. The Snowy Mountains Hydro-electric Authority offered to supply

PRESIDENTIAL ADDRESS. : iii

transport and accommodation for a field party for the projected Natural History survey
of this area. Under the leadership of Dr. W. R. Browne, and with the help of a grant
of £75 from Mr. BH. J. Hallstrom, a party of eight geologists and biologists visited the
Spencer’s Creek area during the period 17th to 30th January, 1951. A report on this
survey is now in course of preparation. It is considered that an initial survey of this
kind in any area shortly to be changed completely by the construction of a large dam
will provide valuable basic data for continued observation of the changes in the natural
history of the area which must inevitably take place during and after the construction
of the dam. It is hoped that further observations will be possible every year until the
new environment has reached a state of stability in relation to a changed fauna and
flora.

The Society again supported a request to renew the proclamation protecting certain
wild flowers and native plants for three years from ist July, 1950, and this has
been done.

The Society has also supported appeals against the possible resumption of land
within the Muogamarra Sanctuary. Similar action has been taken in a number of other
cases wherein it appeared that natural areas within reserves were to be thrown open
for development.

Commonwealth of Australia Jubilee Celebrations—The Society is co-operating with
the Royal Society of New South Wales and other scientific bodies in the planning of a
conversazione in the Great Hall of the University of Sydney on 18th April, 1951. An
exhibit dealing with the early natural history in the State will be our contribution.

We offer congratulations to: Dr. I. M. Mackerras, on the award of the Clarke
Memorial Medal of the Royal Society of New South Wales for 1950 for his work on
Diptera; Dr. T. B. Kiely, on obtaining the degree of D.Sc.Agr., and the award of the
Royal Society’s Medal for his work on the Black Spot of Citrus; Mr. Robert Endean,
Miss Janet Harker and Miss Jean Liddell, on obtaining the M.Sc. degree of the University
of Sydney; Miss Adele Millerd, on the award of a research fellowship which would
enable her to study at the California Institute of Technology; and Miss Muriel Morris,
on obtaining the degree of M.Sc. of the University of Sydney and receiving the award
of an International Federation of University Women Fellowship to study at the
University of Oxford.

Linnean Macleay Fellowships.

In 1949 the Council reappointed three Fellows for 1950, viz.: Miss M. Hindmarsh
(Botany), Miss A. Millerd (Biochemistry) and Miss M. Morris (Zoology).

Miss Hindmarsh continued her investigation of mitotic poisons during 1950. In
particular the effects of phosphates and nitrophenols were studied. Investigation of an
unexpected result obtained when using phosphate buffers to control pH showed that
phosphate inhibited mitosis in a manner similar to sulphanilamide inhibition. A similar
result had been reported recently by I. Galinsky in the Journal of Heredity. Further
work on this inhibition indicated that, although phosphate had a specific effect on cell
division, other salts in an unbalanced solution may show a similar effect. 2,4-dinitro-
phenol upsets cell division by delaying or inhibiting spindle formation and by inducing
stickiness in the chromosomes. It has been found, however, that dinitrophenol acts
as a slow fixative, gradually killing the cells, and not as a specific mitotic poison. The
action of mononitrophenols which have been reported to have a “colchicine-like’ action
on cell division is being investigated, since it is not known whether they too kill
the cells.

Miss Millerd continued her investigation of the succinoxidase and carboxylase of the
potato tuber. The investigation of the carboxylase activity of the potato tuber was
carried along various lines: (i) the preparation of the enzyme, (ii) the splitting and
reconstitution of the enzyme, and (iii) isolation of the product of reaction. Study of
the succinoxidase system had almost reached finality when Miss Millerd was awarded
a research fellowship which would enable her to study at the California Institute of
Technology, and resigned her Fellowship as from 30th September, 1950.

iv x PRESIDENTIAL ADDRESS.

Miss Morris completed a study of the life-history of one of our commercial prawns,
a new species of Metapenaeus. The interesting point about this work is the fact that
it demonstrates conclusively that there is a commercial prawn species breeding in our
coastal lakes. All other well-known species of commercial value definitely go to sea
to breed. It is hoped that this work will be published shortly. Miss Morris was
awarded an International Federation of University Women Fellowship and resigned her
Fellowship as from 14th July, 1950, to continue her studies on plankton at the University
of Oxford.

The Council reappointed Miss Mary Hindmarsh to a Fellowship in Botany for
1951 and appointed Mr. N. C. Stevens and Mr. T. G. Vallance to Fellowships in Geology
for 1951.

Miss Hindmarsh proposes to continue with the investigation of mitotic poisons on
plant cells. The work on the effects of phosphate and dinitrophenol on cell division
will be completed. Levan’s results will be checked by testing the cytological action
of nitrophenols and other related compounds, especially those which are not inhibitors
of phosphate transfer. -A further study of mitotic poisons of the spindle inhibitor type
will be made, to look for any similarity in action or true colchicine effects. Work on
the inhibition of cell division by sulphanilamide and its reversal by p-aminobenzoic
acid will be continued as suitable material becomes available.

The subject of Mr. Stevens’ studies will be the Geology and Petrology of Bathylithic
Intrusions in the Cowra-Gunning Region.

Mr. Vallance proposes geological investigations in the Ordovician Metamorphic Belt
of Central-western New South Wales.

We wish all three every success in their year’s work.

Macleay Bacteriologist.

Dr. Yao-tseng Tchan arrived from Paris and commenced his work as Macleay
Bacteriologist on ist August, 1950. An opportunity to meet Dr. Tchan was given to
members of Council and others at afternoon tea on 2nd August.

Dr. Tchan’s work on the Gilgai and red-brown earth soils has made some progress.
Generally speaking there is no N-fixation, but the addition of wheat straw modifies the
general microflora and as the result less N is lost as ammonia and the humus content
is increased. Experiments with wheat are not yet finished because the vegetative cycle
of the plants being used has not run its full course. Also, during the work, estimation
techniques for the N-fixation bacteria and the total soil flora have had to be modified.
Dr. Tchan gained some knowledge of Australian ecological conditions as the result of
an excursion with members of the Faculty of Agriculture. Dr. Tchan proposes to
continue work on the two soils mentioned above, as well as certain soils of the
Sydney area.

Obituaries.

It is recorded with regret that the following members died during the year:
Protessor W. J. Dakin, Dr. B. L. Middleton and Dr. G. A. Waterhouse.

William John Dakin, Emeritus Professor of Zoology in the University of Sydney,
died on 2nd April, 1950. Before coming to Sydney as Professor of Zoology in 1929
Professor Dakin had wide experience of research and teaching in Liverpool, Kiel,
Heligoland, Norway, Naples and Perth. Throughout his life he had a profound interest
in the problems of marine biology, which he pursued with far more than normal
vigour wherever he happened to be. He introduced this field of research to his
department at Sydney University and from humble beginnings gradually developed a
pattern of investigation which culminated in the foundation of the Commonwealth
Scientific and Industrial Research Organization Fisheries Laboratory at Cronulla.
During World War II Professor Dakin transferred his attention to problems of
camouflage and was appointed Technical Director of Camouflage for the Commonwealth
of Australia, an activity he continued until the end of the war. Throughout his life
Professor Dakin not only produced a considerable volume of research papers, but also
wrote several books, both text and documentary, and gained a widespread popularity

PRESIDENTIAL ADDRESS. Vv

as a radio speaker, expounding in understandable terms to a very wide audience the
problems of science he loved so dearly. Professor Dakin joined the Society in 1929,
was a member of Council from 1930 to 1943, and was our President for the 1934-35
session.

Bertram Lindsay Middleton died on 16th October, 1950. Dr. Middleton was a
graduate in Arts and Medicine of Trinity College, Dublin, who came to Australia
following a world tour many years ago. In 1912 he commenced practice in Murrurundi,
New South Wales, which he continued until the time of his death. Dr. Middleton had
a lifelong interest in Lepidoptera and amassed a most extensive collection of Australian
moths and butterflies amounting to some 10,000 specimens, which he bequeathed to the
Australian Museum. Dr. Middleton had been a member of this Society since 1937.

Gustavus Athol Waterhouse died on 29th July, 1950. Dr. Waterhouse was a
graduate in both engineering and science from the University of Sydney. He was
Assistant Assayer at the Sydney branch of the Royal Mint from 1900 until it closed
in 1926. His scientific work is best known from his long study of the Lepidoptera,
which culminated in the production of the books “The Butterflies of Australia” (in
collaboration with Mr. G. Lyell) and “What Butterfly is That?”. Many other contribu-
tions to our knowledge of Australasian Lepidoptera have been published in various
journals, and the collection of Australian Hesperiidae he assembled in the Australian
Museum is undoubtedly one of the world’s finest. Dr. Waterhouse was Keenly interested
in the affairs and management of a number of scientific societies, and it was in this
field that he made outstanding contributions to the organization of scientific work in
Australia. He first joined the Society in 1897 and was our President for the period
1921-23. The Society is particularly indebted to him for his supervision of our finances
during the period 1926-43, when he acted (except during 1928-29) as our Honorary
Treasurer. In 1943 ill health forced him to resign from the Council and he was elected
a Corresponding Member. The sound management of the Society’s finances carried out
by Dr. Waterhouse has become increasingly obvious since his retirement from active
participation in the Society’s affairs, and we are indeed grateful for the foresight he
displayed in building our finances to a level which makes it still possible to ‘carry on
without obvious retrenchment despite ever-rising costs for all our activities.

THE PROBLEMS OF INSECT QUARANTINE.
1. Definition and Introduction. - F
2. Insect Pests in Australia: (a) Exotic species now present in Australia; (b) Australian
insects of importance in other countries.
38. Recent Instances of the Spread of Exotic Pests within Australia.
4. Evidence of the Introduction of InsectS into other Countries.
5. Exotic Insect Pests not yet introduced into Australia.
6. The Reality of the Economic Consequences of Pest Insect Introductions.
7. Reecapitulation of the General Problem.
8. Can Quarantine Procedures be Justified?
9. The Development of Insect Quarantine Legislation.
10. The Broader Aspects of Insect Quarantine.

1. Definition and Introduction. |

_The concept which lately has been described as insect quarantine simply implies
considerations of the prevention of entry of noxious insects into areas where they are
not known to occur, whether such areas be geographically or politically limited. Initially
of course the enphasis is on the prevention of such entry into countries whether they
be continents, major geographical units of continents, or islands large or small. Despite
this initial emphasis, problems of prevention of spread of noxious insects within
geographically or politically limited areas also arise and are generally considered within
the field of insect quarantine. :

The basic problems are further complicated by the fact that certain of the harmless
or relatively harmless insects of one country may prove to be pests or even serious
hazards to the economy or health of another country, simply because a different set
of environmental conditions may make an enormous difference in the level of population

B

vi PRESIDENTIAL ADDRESS.

attained and in the ecological relationships of the insect and the plant or animal life
of the new environment. Furthermore, the insects themselves may not be intrinsically
noxious but may act as vectors of human, animal or plant diseases and, for this reason,
may be regarded as undesirable immigrants.

In actual fact there are natural ecological barriers of varying magnitude which,
in themselves, offer considerable opposition to the spread of insects from one country
to another, the most effective of these being the various oceans of the world. Deserts
and mountain ranges operate in the same way, and tracts of country ecologically
unsuitable to or otherwise uninhabited by the host plant or animal may also form
barriers.

Such natural barriers have not proved adequate to prevent the spread of insect
pests from country to country, even when separated by wide oceans, since the artificial
methods of transfer by ships or aircraft are readily available. The tempo of such
exchanges does seem to have increased very markedly with each increase in the speed
of transport. For instance, Deputy (1948) claims that few field insects of any
importance, other than the Hessian Fly and the Codling Moth, had established them-
selves in the United States of America prior’ to 1850. In his opinion, few insects other
than stored product pests were able to survive the lengthy journeys of the old sailing
vessels, and it was not until the advent of steam vessels that the majority of exotic
pest species now present in America were able to reach there. The story of recent
introductions of insects into Hawaii, particularly by aircraft, is one to be told at a
later stage, but, when unfolded, it does disclose an infinitely greater rate of exchange
of insect species than obtained up till a few years ago, when extensive intercontinental
air traffic was not in operation.

It is contended, then, that we are now, even more than previously, faced with the
necessity of devising artificial protective measures, and some, but not all, of these must
have a legislative content.

Most countries have varying enactments governing the entry of insect pests, and
such legislation is usually embodied in Plant Protection Acts or Quarantine Regulations
for Animal and Plant Diseases and Pests. Further, such legislation may be under the
administrative control of various authorities such as departments of health or of
agriculture.

As sometimes happens with a principle or, activity which has undergone gradual
development over a period of years, the arguments for and against these become obscure
and it may be found that the underlying reasons for certain actions appear to have
been accepted without being adequately stated. Such a position may arise particularly
when a system of legislation is built up item by item on specific issues, rather than
being the result of a full investigation of a general problem at any particular time.
Furthermore, when we are dealing with precautionary measures to prevent the entry
of undesirable insects, or in the closely related field, plant diseases, it is particularly
difficult, when such measures are successful, to establish that any real danger was
imminent. Perhaps the most important evidence we can offer is the result of lapsed
quarantine precautions due to the exigencies of World War II.

It is for these reasons that it has been felt worth while to review the activities
of insect quarantine measures and precautions, particularly as they affect Australia,
and endeavour to establish whether or not we have a great deal to gain by the
continuance of such activities. ’

Australia, with its distinctive fauna and flora, and complete lack of agriculture
and animal husbandry prior to the advent of the white man, and with her relatively
brief period of colonization, should be a very favourable area for the investigation of
the economic consequences of the introduction of noxious insects. Many of its insect
pests are obvious introductions; some are indigenous and, moreover, the results of
the emigration of some of its indigenous insects to other countries are well known.
Its disadvantages are its size and the complexity of its insect fauna, which remains
inadequately studied in most groups as compared with Hawaii, with a small island area
and relatively small and better known insect fauna. However, the diversity of the

PRESIDENTIAL ADDRESS. vii

environments presented within Australia, as well as the diversity of its primary
industries, may in some measure compensate for these disadvantages.

Finally, I should remark that in presenting this address I have in mind not the
entomologists or even biologists who may happen to hear it, but rather those whose
studies have followed different disciplines and yet who may feel that some elucidation
of this problem is worthy of a few minutes of their attention.

2. Insect Pests in Australia.
(a) Exotic species now present in Australia.

The noxious insects of Australia are partly indigenous species which have trans-
ferred themselves from native hosts, either animal or plant, to introduced animals or
to plants of agricultural or horticultural significance, and partly exotic species of
insects which have entered the Commonwealth since 1788. There is no question
concerning the relative importance of the two, the introduced pest species being far
more numerous and damaging.

It is difficult to learn much about those pests which became established during
the first hundred years or so of our history, and it is even difficult to pinpoint the
entry of those which have appeared within the last twenty years. On the other hand,
there is some information available concerning those pests which have been intercepted
at the occasion of entry, but here we find a regrettable tendency to believe that such
abortive introductions could not have established themselves in any case. That such
a belief has no basis in fact is evidenced by a selection of cases cited at a later stage
of this address.

However, let us consider those pests which may safely be considered to have
entered Australia within the past 180 odd years. In the case of those which have
been recorded from Australia over a long period, the evidence that they are exotic is
simply that they have been known at least as long, if not longer, elsewhere and that
they are associated with, particularly, plants which are themselves definite introductions
to Australia.

One of the earliest Aphids to arrive in Australia was the Woolly Aphis of Apple,
which was recorded as early as 1846 in Victoria. The Aphididae generally are of
special interest since no native species are known and yet some 15 or more introduced
species are established pests of a variety of plants. This is a remarkable record for
one family, but when we consider that most Aphids over-winter in the egg stage one
realizes how easily they might enter a new country on imported plants. Much the same
applies to the scale insects, which have a long period of attachment to the plant host
and hence considerable opportunity for transfer, with their hosts, from country to
country. Among the more important scale pests are the Red Scale, the San José
Scale, the Mussel Scale, and the White Wax Scale. All of these are definite introductions.

The Green Vegetable Bug has only been known in New South Wales since about
~ 1911, but has become a widespread pest in the intervening years. Similarly the Peach
Tip Moth first appeared in the Sydney district about 1909 and has since spread through-
out most coastal districts.

The Mediterranean Fruit Fly was first discovered in Western Australia in 1896
and the Brown Vegetable Weevil in Victoria in 1905. An earlier introduction was the
Maize and Tomato Caterpillar, the first record of which dates back to 1858.

Our domestic fleas are introductions despite the fact that there are many native
species. Most of our mosquitoes are indigenous, but the two most noteworthy domestic
species, Culex fatigans and Aédes aegypti are undoubtedly introduced and have been
here for many years.

It is safe to say that most, if not all, pests of stored products, grains and fabrics
have been introduced with certain of the materials they commonly infest. The same
applies in large measure to pests of stock and other domestic animals. Taken together
with the pests associated with the introduced plants of agriculture and horticulture,
the sum total of imsect pest species which have entered the Commonwealth from
elsewhere is a very formidable one.

viii PRESIDENTIAL ADDRESS.

(b) Australian insects of importance in other countries.

The earliest and most often quoted case of an indigenous Australian insect becoming
a pest elsewhere is that of the Cottony Cushion Scale. In Australia this scale appears
to be restricted to the members of the large genera, Acacia, Pittosporum, Casuarina,
Grevillea, Hakea and possibly other native Australian plants. When introduced into
other parts of the world it almost immediately extended its host range to include
Citrus, Prunus, Pyrus, Robinia, Vitis, Laurus, Magnolia, Quercus, Buxus and many other
surprisingly different food plants (Hssig, 1948).

The introduction of the Cottony Cushion Scale to California took place about 1868,
and it soon became one of the most serious pests of Citrus and later appeared on many
other fruit crops and ornamentals. By 1890 it had killed hundreds of thousands of
orange trees.*

Not so widely known are the cases cited by Miller (1948). ‘‘The Blue-gum Chalecid
(Rhicnopeltella .eucalypti)” is “a Tasmanian insect established in New Zealand, where
it is actually destroying, throughout the country, over a period of years, the Tasmanian
blue gum (Hucalyptus globulus) but does not attack any other species of eucalypt. On
the other hand the various species of Hucalyptus are attacked by the scale, Hriococcus
coriaceus, the Chrysomelid beetle, Paropsis dilatata, and the weevil, Gonipterus
scutellatus—all Australian species normal to the hosts. The scale actually kills the
trees, especially the more susceptible Hucalyptus globulus, but little harm is caused
by the beetles except that the weevil may produce a pronounced stunting of growth”.
Certain Australian termites, Coptotermes acinaciformes and a species of Porotermes,
have also been introduced to New Zealand within the last twenty years, and the eucalypt
weevil, Gonipterus, has also caused severe damage to eucalypts in South Africa
following its introduction there from Australia.

3. Recent Instances of the Spread of Exotic Pests within Australia.

The time of introduction of most exotic insect pests cannot be established for many
of our major pests, either because their early establishment passed unnoticed
some time during the last century, or because their importance as pests did not coincide
with their early distribution in the Commonwealth. In regard to the latter it is
possible that the Sheep Blowfly (Lucilia cuprina) did not achieve pest status until long
after its arrival, such status being conditional on changes in the type of sheep developed
in the country.

Recent introductions are few in number, and this may be a chance occurrence or a
tribute to the quarantines which have operated for the past forty odd years. Evidence
derived from elsewhere would almost conclusively establish the validity of the latter
viewpoint.

We must look particularly to these more recent cases for evidence of what actually -
happens following a new pest introduction, and one of the most illuminating is the case
of the Buffalo Fly.

In one respect the Buffalo Fly is falsely entered in this class, since it seems most
probable that it entered Australia as far back as 1825, when buffaloes came to the
Northern Territory from Timor. However, the pest remained confined to the Darwin
area for many years, and it was not until 1912 that Gilruth was the first to suggest
that this fly might eventually prove a major one of cattle. Extensive movements of
cattle in the north, which took place in succeeding years, distributed the fly over a
much wider area. In 1927 it extended from Broome in Western Australia to the
western border of Queensland, and in the following year it entered Queensland south
of Camooweal. The range of distribution of Buffalo Fly in Queensland advanced and
receded from time to time in the ensuing years in concert with cattle movements and
seasonal conditions. A series of wet seasons from 1939 to 1941 resulted in the fly

* Fortunately for our international reputation this pest was eventually brought under
complete control in California and in the other countries to which it had been introduced, by
the introduction from Australia of a predatory ladybird beetle.

PRESIDENTIAL ADDRESS. 1x

crossing the low rainfall hinterland of the Gulf of Carpentaria, and once across this
climatic, but intermittent, barrier its progress to the coast and southward down the
eastern coast of Queensland, almost continuously populated with cattle, has been
unimpeded (Belschner, 1946).

The history of Buffalo Fly in Australia provides a number of most interesting
lessons, although of course we have only gained our wisdom after the event. The
adult fly itself has an almost continuous association with cattle and only survives away
from its host for a matter of 24 hours. Hence it is most unlikely that adults could be
introduced unless cattle were being imported at the same time. The larval stages could
survive and develop in cattle dung, but again it is most unlikely that such material
would reach Australia without the importation of cattle. Given this knowledge,
prevention of its entry would have been quite a simple matter.

Again, for almost 90 years the pest had a restricted distribution in the Northern
Territory. At any time during this period an eradication programme might have been
attempted, although it must be admitted that this would have had greater chances
of success had modern insecticides then been available.

Further, the crossing of the intermittent climatic barrier south of the Gulf of
Carpentaria took place in spite of fairly vigorous quarantine precautions in north-
west Queensland, assisted, at least in part, by the lack of co-operation of those responsible
for cattle movements.

Finally, it is interesting to note that once across this barrier the fly quickly
extended its distribution to the hypothetical limits postulated for its distribution by
Handschin (1932).

Compared with the Buffalo Fly, the spread of the Cabbage White Butterfly following
its introduction, perhaps from New Zealand, was extremely rapid. It was first recorded
in a restricted area of Victoria in 1939, following which it was found at Albury in
1940 and in Sydney in 1941, and thence spread rapidly to Queensland. A case of this
nature emphasizes the urgent necessity for rapid decisions as to whether any attempts
should be made at eradication following the first discovery of such a pest. Delay may
mean the loss of all opportunity of eradication and leave us with just one more
perennial pest species to be subjected to measures of control year after year. Further,
the rapid spread through a number of States also emphasizes the tact that States
distant from the site of introduction may really be equally concerned in the arrival,
or in the eradication, of the pest within the area of early establishment in the country.

The Argentine Ant is again a recent introduction. It was first recognized in a
Melbourne suburb in 1939 and in Perth in 1941. The first Sydney findings were in
1950, but it seems likely that it has been present here for at least three years. It is
possible that three separate introductions were involved, but it is equally feasible that
this pest has been unknowingly transferred from place to place, perhaps in potplants.
There is no doubt, however, that this aggressive insect will continue to spread as
opportunity offers (it rarely flies), unless eradicated or most carefully controlled.

In these cases at least we have very tangible evidence of introduction and subsequent
spread within Australia. Similar events have taken place many times in the past; others
are perhaps still going on. But it is:such cases of rapid dispersion that make it
clear that widespread distribution following an introduction is possible and, indeed, is
almost inevitable, although in many cases the time necessary for obvious dispersion
may be a very variable factor.

' 4. Evidence of the Introduction of Insects into other Countries.

In the field of medical entomology clear evidence is available of the transfer of
pest insects between the continents of Africa and America across the formidable barrier
presented by the Atlantic Ocean.

The chigoe flea, Tunga penetrans, which occurs in the tropical and subtropical.
areas of North and South America, including the West Indies, has been established
in Africa for less than eighty years. It is not difficult to envisage how such an insect

x PRESIDENTIAL ADDRESS.

with a semi-permanent attachment to the body of man or pigs should effect its transfer
from America to Africa.

On the other hand, it is a far more difficult matter for a mosquito. to traverse the
same ocean, yet this has been accomplished by at least one non-domestic* species,
Anopheles gambiae. This potent African vector of malaria reached Natal in Brazil
about 1930 and in the ensuing years was responsible for epidemics of malaria of the
greatest importance.

We may say that these cases all come from the past and that all that is likely to
happen has already happened. A counter to this view comes from our knowledge of
what has happened in Hawaii in very recent years.

Swezey (1948) discusses 31 species of insects which became established in Hawaii
during and since World War II. Included are a variety of moths, wasps, beetles, bugs
and flies, and a grasshopper and a cricket. Some of these have already been shown to
be devastating pests, others are potentially dangerous, and some, on the other hand,
are beneficial. It is revealed that some of these must have come from U.S.A. and
others from various parts of the Western Pacific Zone.+ A significant number of these
introduced insects were first found in the vicinity of Pearl Harbour, “which could be
accounted for by reason of the excessive increase in shipping and airplane arrivals
at this military centre, and the consequent great difficulty of carrying on quarantine
inspections”. No attempt is made to include the large number of interceptions of
migrant insects made by entomologists in their quarantine inspections. Another point
of interest is the fact that-many of the first records of these introduced insects came
from light traps which have been operated in the Pearl Harbour area with the express
purpose of gaining prompt knowledge of the establishment of any anopheline mosquitoes,
an event which has not so far taken place.

It is perhaps of interest to mention two of these introductions which are not known
to be detrimental. One is the introduction of the wasp Humenes latreillei petiolaris
(Schulz) from New Guinea. This wasp has effected a considerable measure of control
on the moth Anacamptodes fragilaria (Grossbeck), which itself reached Pearl Harbour
from California only two years prior to the wasp which has proved such a valuable
addition to the fauna of Hawaii. Such a happy set of circumstances is unlikely to occur
with any frequency, but it is interesting to record that a chance introduction from the
east of a potentially destructive pest is brought under control, at least in part, by an
equally chance introduction from the west.

The other of special interest is the introduction of an apparently harmless wingless
grasshopper, Paraidemona mimica Scudder, from U.S.A., probably Texas. This is one
of those cases where it is difficult to imagine how a wingless insect, in the absence of
any host animal or plant or other material with which it might be closely associated,
should manage to join a ship or an aircraft in order to accomplish such a journey.

In an address to the Entomological Society of Queensland in September, 1950, a
guest speaker, Mr. C. Pemberton, stated that a total of 219 species of insects were
introduced into Hawaii during the years of World War II. The most important among
these was undoubtedly the Oriental Fruit Fly, Dacus dorsalis (Hendel), which appeared
first in 1945 and rapidly became established throughout the entire group. Its attacks
on avocardos and mangoes virtually ruined these industries until 1950, when the
work of a team of 40 men on a half-million dollar budget started to show results,
principally from the introduction of 15 different parasites gathered from various
countries of the Western Pacific Zone. The Oriental Fruit Fly is one of those particularly
devastating pests when divorced from its natural controls, and is capable of breeding
in all sorts of different fruits and nuts (more than a hundred different host plants

* Certain domestic species, e.g. Aédes aegypti, have become tropicopolitan or, Culex
fatigans, almost cosmopolitan. Such species which are likely to breed in drinking water
might easily have been distributed in water barrels in the days of sailing vessels.

+ Of the 31 species discussed, 11 have come from the U.S.A., 6 from Guam, 5 from the
Philippines, 5 from other Pacific islands or the Orient, while the origin of the remaining 4 species
is uncertain.

PRESIDENTIAL ADDRESS. xi

were recorded in Hawaii—Cooley, 1950) and should by no medns be considered as just
another fruit fly. :

We have mentioned before the claim that the tempo of insect transfers from
country to country underwent a marked increase with the advent of steam navigation.
Then, subsequent to the more or less general introduction of quarantine precautions
in the first decade of this century, there did appear to be a slowing up of such transfers.
For instance, only four major insect pests have become established in the U.S.A. in
the quarantine period 1912 to 1948 (Deputy, 1948). Of these, one, the Mediterranean
Fruit Fly, has been eradicated and two entered from Mexico, the Mexican Fruit Fly
and the Pink Bollworm. The fourth, the White Fringed Beetle, appears to have entered
from South America.

In contrast to this, some 30 major pests entered and became established in the
U.S.A. during the fifty years prior to the introduction of quarantine precautions.

How, then, do we reconcile the Hawaiian experience with such a trend? Admittedly,
quarantine services were in part disrupted during the war, but beyond this we saw
for the first time the real effect of extensive air transport. There seems no question
that the increased speed of air transport over all previous methods has again brought
in its train another and even greater increase in the tempo of insect transference
from country to country and has presented us with quite a new set of problems.

At this stage I feel it would be pertinent to review the role of aircraft in insect
pest introductions. To those who travel by aircraft, particularly in the northern half
of Australia, the frequency with which mosquitoes and house flies are encountered
inside passenger cabins is well known, but the extent to which other insects travel
inside aircraft is not generally appreciated.

A number of detailed surveys of the insect passengers in aircraft have been made
by various workers in the U.S.A. Hughes (1949) gives in considerable detail the
entomological findings from a large number of aircraft entering various airports in
the U.S.A. from other countries for the 10-year period 1937-47. Out of a total of
80,716 planes inspected during this period, 28,752, or 356% of the total, harboured
arthropods. From the same total 3,873 or 4:8% harboured mosquitoes. The total
number of arthropods found in this period was 106,106, of which 16,846 were alive,
and this in spite of the increasing use of disinsectization measures.

The same author records that 12,825 mosquitoes were found during this period,
representing 10 genera with a total of 73 species. Of these, 48 were indigenous to one
or more of the areas at which they were intercepted, but 25, or 34:2%, of the total number
of species, were truly exotic. Some of these were, of course, South American, but a
number must have travelled considerably further and nine of them undoubtedly
embarked on their flight at some point in the Western Pacific Zone. The distances
involved would necessarily be from 4,000 to 6,000 miles.

For one year (1945) the total arthropod records comprised “taxonomically deter-
mined categories consisting of 20 orders, 191 families, 680 genera and 524 species’, and
within the class Insecta, exclusive of biting mosquitoes, 17 orders, 188 families, 670
genera and 491 species were determined, not all catches being identified as far as the
species.

Admittedly many of these insects arrived at their destinations dead, but approxi-
mately 15% were alive, despite, I must emphasize, the current disinsectization measures
applied up till the time of disembarkation.

Another type of evidence of survival of insects in aircraft has been provided by
Laird (1948), who transported mosquitoes in cages by aircraft from New Zealand to
Japan and back. Of 75 female Aédes notoscriptus, 60% survived the 18-day journey of
more than 12,000 miles over a wide range of conditions, and the average life span of
those which survived was 61 days. Other details are given, but the above is surely
sufficient to establish the ability of mosquitoes to withstand successfully long journeys
by air.

xii PRESIDENTIAL ADDRESS.

5. Exotic Insect Pests not yet Introduced into Australia.

A system of insect quarantine regulations and the precautions that go with these
would not be justified if it could be shown that there were few, if any, major pests
still occurring elsewhere which had not so far reached and successfully established
themselves within the Commonwealth.

A little reflection on the part of entomologists will call to mind quite a formidable
list of insect pests, well known elsewhere in the world, which have not yet appeared
in Australia and yet would certainly occasion us serious trouble in the fields of agri-
culture, animal husbandry or health.

Among the more obvious plant pests are the European Corn Borer, the Hessian
Fly, the Oriental Fruit Fly, the Mexican Fruit Fly, the Boll Weevil, a number of Vine
Moths, certain Sugar Cane Borers, the Colorado Potato Beetle, the Apple Capsid, the
Pineapple Mealy Bug, Chinch Bugs, the Japanese Beetle, the Gypsy Moth, certain Rice
Borers, the European Red Mite, Sirex Wood Boring Wasps and Dry Wood Termites.

In the veterinary field obvious exotic pests, the exclusion of which is most
important, are the Warble Flies, the Screw Worm and certain species of ticks other
than those already in Australia.

From the viewpoint of human disease we would be most concerned in excluding
such potent vectors of malaria as Anopheles punctulatus punctulatus, Anopheles
sundaicus and Anopheles gambiae, although quite a number of other species would be
considered dangerous. Other arthropods to be excluded would be Tse-tse flies, certain
ticks (e.g., Ornithodorus) and blood-sucking bugs (Reduviidae).

Particularly in the agricultural field, closer study would reveal an infinitely greater
list of pest species one would wish to see prevented from entering the country. However,
in each field we have mentioned only the most obvious of the recognized undesirables,
and it must be emphasized that other problems exist of an equally serious nature.
These are worthy of specific citation.

(i) Phytophagous insects of little or no status as pests in their native environment
may be particularly devastating in a new environment or new host-relationship.

_ (ii) The danger of a plant insect pest does not always lie in the direct damage
caused, but such insects may introduce with them a new plant disease. On the other
hand, the disease may be present without an adequate vector species, which may find
an opportunity of entering at some later stage. Our major concerns here are with
plant-sucking bugs and virus diseases.

(iii) Again, in the case of human diseases particularly, we may find the vector well
established without the disease, as in the classic case of Yellow Fever. One method of
introducing the disease is in infected vectors, a possibility which increases with faster
means of transport and the diversification of transport routes. On the other hand we
may have the disease and certain of its vectors, but would still find a far more serious
problem developing from the introduction of more potent vectors of the same disease.

(iv) Further, it is worth emphasizing that species differences are often of the
greatest importance. The fact that we have present in the country certain ticks or
certain mosquitoes is no reason for complacency concerning the possibility of the intro-
duction of other species of these groups, and this applies time and time again in a
wide variety of groups and genera.

(v) Finally, even though a pest is already present in the country, it may not
necessarily be widely distributed and its reintroduction at other points may be just
as damaging as a de novo introduction. Such circumstances can readily be visualized
in the case of a pest of the nature of the Argentine Ant.

6. The Reality of the Consequences of Pest Insect Introductions.
Following the introduction of Anopheles gambiae into Brazil in 1930 the loss in
human lives directly attributable to this introduction was considerable, reaching an
estimated total of 14,000 to 20,000 people. For the State of Rio Grande do Norte the
estimated deaths exceeded 5,000 out of a population of 243,000 odd. (This State is only
a portion of the total area invaded by A. gambiae and the estimates were made in

PRESIDENTIAL ADDRESS. Xili

1938, a year before the intensive anti-gambiae campaign commenced). In the same
area more than 51,000 cases of malaria were estimated to occur and the economic losses
occasioned by the debilitated population were considerable. To these consequences
must be added the actual 3,000,000 dollar cost of the eradication of this species from
the invaded area.*

Again in 1944 Anopheles gambiae caused more than 32,000 primary cases of malaria,
of which almost 1,800 were fatal, after its introduction to Upper Egypt was first noticed
in 1942. Here also a successful eradication campaign cost a total of £1,000,000 sterling.

Estimates of the economic losses due to insect pests have been made from time to
time, but as they are usually based on reductions of yield for various crops they are
not necessarily convincing. Estimates of the losses due to Sheep Blowfly were estimated
to amount to £3,000,000 in this country during 1942, and this is rather more tangible
than a loss of a percentage of a crop before its actual maturity. Even more convincing
are figures given for control or eradication programmes. For instance, the Cattle Tick
eradication programme on the North Coast of New South Wales had cost £3,000,000 up
to the end of 1940.

Unequivocal evidence is also presented by those cases of almost complete annihilation
of particular primary industries as was occasioned by the Cottony Cushion Scale on
Citrus in California (see above) and more recently the ruin of the avocardo and mango
industries in Hawaii by the Oriental Fruit Fly. In the latter case a sum of 500,000
dollars was allocated for the investigation of control measures.

Apart from such tangible cases there is no doubt that considerable losses are
occasioned by most insect pests, even though they may be difficult to estimate con-
vineingly. On the other hand, the actual cost of having insect pests, apart from such
losses, is clearly indicated by the money spent on their control, and a partial estimate
of this (exclusive of labour costs) is presented by the value of insecticides used. No
accurate Australian estimates are available for insecticide consumption, but from the
limited data of production of certain basic insecticides, and the market has by no
means reached saturation for chlorine-containing insecticides at least, it would appear
that, at a conservative estimate, two to three million pounds worth of insecticides are
annually consumed in this country, and the real figure may be even higher.

Other figures on expenditure for individual pest control or eradication in American
cases are quoted by Annand (1950). For example, one fairly recently established pest,
the White Fringed Beetle, has occasioned the expenditure of about 1,200,000 dollars
during 1950, to which we can add a total of some 9,000,000 dollars spent prior to 1949.

Generally, a perusal of appropriate sources of information does give more and
more confirmation of the actual costs of insect pest introductions as being far-in excess
of the costs of administering quarantine precautions. Up to a point increases in
appropriations for such work do increase its efficiency, but beyond this it is a matter
of training and knowledge.

7. Recapitulation of the General Problem.
In reviewing the evidence presented above it is clear that the following points
have been established:
' (i) That a very considerable transfer of insect pests has gone on in the past
from country to country, even between those separated by wide oceans; and this has
been largely accomplished through the agency of modern methods of transport.

* Soper and Wilson (1943) discount the theory that A. gambiae arrived by aircraft. One
of the reasons (there are others) is that the first discovery, which had all the appearances
of revealing a very recent introduction, was made at a distance of seven kilometres from the
airfield with no foci of infestation closer to it. In this connection it is interesting to record
that Mr. Hegener, a Dutch colleague, captured a live specimen of Anopheles bancroftii on
the sixth floor of the Hotel Metropole, Sydney, in April, 1945. This species has not been
found south of Queensland and is certainly not known to occur in the Sydney district. It is
interesting to speculate just how this insect could have arrived at this location, a distance
of more than three miles from the flying boat terminal and considerably more from the
land plane airport.

xiv PRESIDENTIAL ADDRESS.

(ii) That, once introduced, insect pest species have proved themselves capable of
rapid or delayed spread throughout large tracts of the invaded country.

(iii) That insects of little or no importance in one country may become major
pests in another, or, in other words, it is not always possible to prejudge the importance
of a particular species should it appear outside its native environment.

(iv) That the possibilities of insect transfers have increased with each marked
improvement in methods of transportation, and that recent increases in the volume and
diversity of routes of air transport have enormously increased the possibilities of
further transfers.

(v) That serious economic losses are involved following the introduction of
important pest species which may also be accompanied by serious losses of human life
in the case of species of medical importance.

(vi) Finally, that even the reintroduction of existing species may have undesirable
consequences either by extending the existing range of distribution of the pest or by
the introduction of plant or animal diseases.

It seems obvious, then, that consideration should be given to the possibilities of
preventing the entry of insect pests into countries new to them, and we do, indeed, find
that insect pest quarantines have been in force in most countries for approximately
the last forty years.

8. Can Insect Quarantine Precautions be Justified?

Wardle (1929) is prepared to contend that “it is doubtful whether any pest which
is at present to the fore would have been prevented by quarantine Acts from gaining
admittance to the country” had U.S.A. instituted Plant Quarantine legislation fifty years
earlier than it did. He considers that “the possibility of legislative barriers really
fulfilling their intentions rests upon a perfection of circumstances which, in the present
imperfect condition of civilization, simply does not exist’”’.

Wardle raises these additional objections to legislative barriers: (q@) that it is rarely
possible to foresee that a potentially dangerous insect is likely to become introduced;
(db) that the cost of maintaining the required inspection forces may be out of all
proportion to the possible losses that insect introduction may bring about; and (c)
that such legislation may act in restriction of international trade and may be used as
a weapon by particular agricultural sections ‘that desire economic protection.

The statement by Strong (1923), ‘the fact that no pest of major importance has
become established in the United States since the passage of the Plant Quarantine Act
demonstrates in a graphic manner the value of plant quarantine to the United States
as a nation”, is discounted by Wardle on the grounds that the period of time which
had elapsed since the passing of the Act—ten years—was not long enough. We have
already quoted the results of thirty-six years of Insect Quarantine operation for the.
U.S.A. (Deputy, 1948), which is surely sufficient answer to this objection, and we have
also given the evidence provided by the wartime breakdown of quarantine precautions
in Hawaii. ‘

That the required perfection of circumstances does not exist is an indefinite criticism
implying, in part, lack of sufficient knowledge and lack of efficiency on the part of the
persons charged with the duties of administering the relevant quarantine procedures.
Neither of these difficulties is insuperable, but it is important that quarantine staffs
should be adequately trained and suited to their profession.

As experience is gained it becomes increasingly possible to foresee possibilities of
pest insect introduction, and as insect and related quarantines become more and more
international in their outlook the less credence can we give to this objection.

The critical objection surrounds the relative costs of the administration of such
quarantines as compared with the subsequent costs of control or eradication following
the establishment of a new pest. When we consider that quarantine is applied in a
limited number of ports to ships and aircraft and their cargoes instead of control
measures over wide areas of territory or large acreages of crops there seems little
room for comparison. Then, when we realize that the quarantine procedures cope with

PRESIDENTIAL ADDRESS. xXV

far more than just individual pest species at the one time, instead of separate control
measures for a number of different pests, there can be no doubt that precautionary
measures will prove far less expensive. The fact that eradication programmes are
seriously considered following the early discovery of a new insect pest is a further
denial of Wardle’s contention.

Finally, to say that insect quarantine legislation may be used as an economic
weapon is not a criticism of the principles of quarantine, but one of the integrity
of governments.

If these are the most serious objections to the promulgation of insect quarantine
precautions then it must be granted that we have strong reasons, along the lines which
have been previously discussed, for the maintenance of such precautions.

9. The Development of Insect Quarantine Legislation.

A full discussion of the history of the development of insect and plant quarantine
legislation, since the two usually go hand im hand, would be out of place in this
address. However, we should realize that such legislation is of comparatively recent
date. In California the first promulgations were issued in 1881, for the U.S.A. as a
whole not until 1912. ‘

In Australia the Federal Constitution of 1901 empowered the Commonwealth to
administer quarantine (Clause 9 of 51). Under the Constitution the first Quarantine
Act, which included a Plants Division, which in its turn dealt with insect pests, was
passed in 1908 and took effect on July 1, 1909. Until 1921 the Act was administered by
the Department of Trade and Customs, but in this year the Department of Health was
established by order of the Executive Council and the administration of quarantine
became a function of this department, wherein it still remains. The year 1927 saw the
foundation of the Divisions of Plant Quarantine and of Veterinary Hygiene as separate
divisions of the Department of Health.

Hxisting procedures do, of course, come under critical review from time to time,
and in the case of the Australian Act the most important contribution is found in
the Report of the Central Committee appointed by the Australian Institute of Agri-
cultural Science to deal with Overseas Plant Quarantine (Nicholson et al., 1941). Here
criticism of existing practice is combined with a detailed series of recommendations
concerning improvements in staff and regulations. These are too detailed to be
quoted here, but it should be pointed out that at least some of these recommendations
have been incorporated in the Regulations under the Quarantine Act, particularly by
Statutory Rules No. 92 of 1948 which gives wide powers for the complete destruction
of noxious animals or plants following their detection on entry, and by Statutory Rules
No. 78 of 1950, which requires the registration of approved authorities who may desire
to import nursery stock under permit. ;

What is of more immediate interest to us in the historical development of insect
quarantine precautions is the gradual change in approach that is evident in the legisla-
tion of most countries, including Australia. :

Harly quarantine legislation provided for embargoes against specific known pests,
at least from the countries where these pests were known to occur, or restricted entry,
subject to the fulfilment of particular provisions, of materials likely to harbour known
pests. Inspection on arrival was provided for and any materials subject to restrictions
would only be passed through quarantine if found free of the suspect insects, or
following suitable treatments to destroy them. Providing the inspecting officer has
been suitably trained and knows what to look for in any particular type of material,
or has ready access to sources of appropriate information and identification, such a
system works reasonably well. There are, of course, some cases wherein it is not
possible to detect the pest insect by inspection, e.g., Narcissus Fly in various bulbs.
To cope with such cases the material may be fumigated or subjected to heat treatment
before shipment, this being done at a time when the bulbs will suffer less damage
from such treatment. In other cases provision is now made for the, growing of suspect
material under quarantine. This may be done by a Federal authority, as in the U.S.A.,

Xvi PRESIDENTIAL ADDRESS.

or by the registered importers, as in Australia. A period of at least twelve months
is then available for the detection of any ‘pests, disease or insect that may be incapable
of detection on entry.

Various ways in which certain commodities, otherwise likely to be subject to
embargoes, may receive treatment prior to their consignment or during their period
of shipment are discussed by Annand (1950). Of particular interest is the shipment
of pre-cooled and refrigerated apples in specially equipped ships from South Africa
to the U.S.A., giving a complete mortality of fruit fly larvae, the most dangerous type
of pest likely to be contained therein. f

Undoubtedly the various quarantine provisions increase in complexity as time goes
on, but they do provide more and more protection against pest insects as they become
revealed as potential dangers. The tendency is to channel all imports of suspect
material through those agencies which would stand to lose most following the intro-
duction of a pest of one of the materials in which they are most interested. And there
can be little doubt that such restrictive precautions are ultimately understood to be
protective to the importer as well as to the country generally.

There are still cases in which material is imported which is of no interest to
anyone, and this applies to the various types of packing used in preparing goods for
shipment. In such contingencies the onus is entirely on the quarantine service to see
that such packing materials are free of pest insects.

Finally, following on the success of pre-flight treatment of aircraft leaving Hawaii
for the U.S.A., particularly to cope with the danger of the introduction of Oriental
Fruit Fly, similar arrangements are now the subject of negotiation between Australia,
New Zealand and Hawaii. It is considered, on sound evidence (Cooley, 1950), that
pre-flight inspection and treatment is a more effective way of dealing with possible
insect passengers than similar operations carried out at the end of the journey and,
moreover, they are more acceptable to airlines and passengers. There seems little
doubt that extensions of this type of operation, will come about in the future, especially
if reciprocal arrangements between countries can be agreed upon.

Except in the last case, we have dealt with problems usually associated with
shipping. Insects harbouring in their host material is one problem, but insects purely
as temporary passengers, .as they often are on aircraft, dissociated from any host
materials, is an entirely different one. Consequently, in recent years, quarantine laws
have had to cope with insects arriving in such a manner on aircraft. In the case
of Yellow Fever the problem is so serious that International Air Navigation Agreements
have been formulated.

Otherwise each country has added to its quarantine laws provisions regarding the
treatment of aircraft to ensure that they arrive in an insect-free condition. Further
precautions are often taken immediately after landing.

Annand (1950) foreshadows the wider use of point-of-origin inspection and clearance
in the future, since it offers greater protection at less cost. He cites the proposal made
in 1948 by the United States to the International Civil Aviation Organization.
“Contracting States should make arrangements whereby one State will permit another
State to station representatives of the public authorities concerned in its territory to
examine aircraft, passengers, crew, baggage, cargo, and documentation for customs,
immigration, public health and agricultural quarantine purposes, prior to departure for
the other State concerned, when such action will facilitate clearance upon arrival in
that State.”

Perhaps the most standard treatment in use at present for aircraft is the aerosol
formulation G—382, containing 3% DDT, 5% of pyrethrum extract (20% pyrethrins),
5% cyclohexanone, 2% lubricating oil and 85% of Freon-12 at a dosage of 5 gm. per
1,000 cubic feet. Admittedly this dosage is not fully effective and pre-flight treatments,
when carried out in the absence of passengers, are applied at four times this dosage
(Cooley, 1950). Other formulations are discussed by McBride et al. (1950), who
concluded that the best results were obtained by a combination of residual and aerosol
treatments.

PRESIDENTIAL ADDRESS. Xvii

In the case of aircraft the problem is not a legislative one but one of technique.
Considerable research has been undertaken to disclose a really effective insecticidal
technique for application to this special problem, but even though routine aerosol treat-
ments may cope with some of the more obvious Diptera in an aircraft, many other
insects remain unaffected. This does not mean that the solution will not be found;
eventually it probably will be. Automatic aerosol-dispensing systems for aircraft
have been designed and are under trial, and there is little doubt that these would
prove an improvement on hand application.

Another approach to the problem is contained in the Report on the Second Session
of the Expert Committee on Malaria of the Interim Commission (W.H.O., 1948). This
report includes the recommendation “that, whatever regulations be enforced regarding
the disinsectization of sea- or aircraft, rigid anti-mosquito sanitation should, as far as
practicable, be maintained within the mosquito flight-range of sea- or airports of the
country to be protected, so that no imported mosquitoes will be able to survive”. The
“reciprocal procedure, in the country of origin of such mosquitoes, should prove even
_more effective.

The present importance of aircraft in the transfer of insect species: has already
been emphasized. It is in this field particularly that uniform legislation and practice
is desirable for all countries and varying procedures in different countries can only
minimize the effectiveness of precautionary measures generally. The time available for
such measures is so brief, and aircraft are liable to pass through many countries, so
that changes in procedure will not only be confusing but are likely to evoke varying
degrees of co-operation. ;

We hope, then, that uniform practices, based on critical research, will eventually
be adopted on an international rather than a regional scale.

10. The Broader Aspects of Insect Quarantine.

We have discussed in some detail the overall background to the problem of insect
quarantine, principally because it is a wide field about which relatively few have
detailed .knowledge of all sections, even among entomologists, and also because its
wide scope, covering pests of all horticultural and agricultural plants, stored products,
the household, and pests of medical and veterinary significance is deserving of emphasis.

Some general discussion has been given to insect quarantine legislation, put a
little more should be said about the implementation of such legislation.

The fact that a port-of-entry inspection service is required has been mentioned, but
hand in hand with this must go an identification service for the less obvious of the
suspect insect pests that may be intercepted,-and efficient treatment facilities for infested
material. The identification service must be sufficiently skilled and knowledgeable to
make rapid determinations, and the treatment facilities should be sufficiently flexible
to cover a range of treatments for reasonable quantities of imported materials. Such
facilities do vary from place to place, and it seems only reasonable that those of the
Imajor ports of entry should be improved at least to the level of the most efficient.

Beyond this a quarantine service should also provide for prompt and effective
action in the way of an eradication programme following the early detection of an
insect importation which has escaped detection at entry. In the Australian Act this
has been provided for in Statutory Rules No. 92 of 1948. It is not necessary that all
the probable requirements for the implementation of such .a programme be held in
waiting for such an event, but it is necessary that we should know where and how
to marshal them when the occasion arises.

In the words of Annand (1947) “quarantines should have two functions, (i) to
prevent the entrance of new pests or those not widely distributed, and (ii) to facilitate
the movement of commerce by the application of treatments and other safeguards, when ‘
such’can be used, in lieu of embargoes. They are not intended to serve in place of
tariffs, and they must be maintained on a biologically sound basis. To be fully effective
they should be based on knowledge of the occurrence, distribution, abundance, hosts,
and habits of harmful pests at home and abroad.”

xvili PRESIDENTIAL ADDRESS.

This introduces the general background to the operation of quarantine legislation,
and Annand also stresses the need for exotic surveys of particularly important imports
and the areas from which they come, linked with the findings of inspectors after the
arrival of such imports.

In the detection of new pests Annand stresses the importance of initial surveys
to establish what is present, and claims that these need to be exhaustive and co-operative
and that greater allocations are required for the necessary taxonomic work.

Cooley (1947), writing particularly of the air traffic problem, defines three lines
of defence: (i) pest surveys in foreign countries so that we may become familiar with
species likely to be dangerous, (ii) inspections at airports, and (iii) continued surveys
in the vicinity of ports of entry. The same author also recommends that all vegetation
in the vicinity of international airports be continually. treated with DDT and other
insecticides to prevent establishment of any pests which may escape from arriving
planes and that various types of insect traps be maintained in the vicinity of all
important air terminals. Such recommendations presuppose the maintenance of an
entomological service at major air terminals.

In general, then, we are faced with the need for far more precise knowledge of
insect pests on an international basis. For each country to gain its knowledge of
exotic pests individually is more than a duplication of effort but, since few countries
document their insect pest problems in a form suitable for quarantine purposes, such
a procedure may be desirable, at least in special cases, for some time to come.
Ultimately, when the type of information required by quarantine services is more
widely realized, and there is co-operative effort on an international basis in the
preparation of such data and the early circulation of information on new pests the
need for exotic surveys will diminish.

But first of all we should know, and document, our own insect pest story, and
this applies equally to those which are to us of major and of minor importance. Perhaps
such information as we possess is available, but with six States in the Commonwealth
and an equal number of journals recording agricultural pest information, the task of
abstracting relevant details is a formidable one. The type of documentation envisaged
is only accomplished when a specific allocation of staff and money is made for such
a project, and it is one of our needs that the specific data on recorded pests, their
distributions, habits, life histories, and animals, plants or other materials attacked
should be gathered together in one series of publications.*

A prerequisite of such documentation is discussed by Hssig (1948), who stresses
the need for insect surveys of literature, of collections, and of insect occurrence in the
field on a scale sufficiently detailed to provide exact information for such areas with
which any particular survey unit can cope. Here again we reveal the need of a
wide basic taxonomic knowledge.

Finally, the efficient operation of quarantines depends not only on knowledge
within the professional field but it also requires a general public realization of its
purposes and a more specific knowledge of individual problems amongst those most
concerned with goods likely to be subject to quarantine inspections.

Im other words, a public relations programme is also necessary, since lack of
co-operation is usually based on ignorance.

In conclusion I should like to emphasize that the time has long since passed when
we regarded insect, and also plant, quarantines as restrictive to the commercial
exchange of various commodities. The basis on which they operate becomes biologically
more sound as time goes on, and the needs of international commerce continually

* This implies a central information service as envisaged by Nicholson et al. (1941). It
is pleasing to be able to record that for some years now the N.S.W. Department of
Agriculture, through its Entomological Branch, has been bringing out an ,annual insect-pest
survey which provides much useful information. For the past two years a section has also
been devoted to pests intercepted in quarantine and this provides information of considerable
significance.

PRESIDENTIAL ADDRESS. xix

demand that precautionary treatments and safeguards be developed. A. fuller realization
of the problems involved is particularly desirable and there is further scope for the
encouragement of co-operative effort amongst organizations not directly concerned in
the implementation of quarantines.

References.
ANNAND, P. N., 1947.—Preventive Entomology. J. Hcon. Ent., 40: 461-468.
: , 1950.—Today in Foreign Plant Quarantine. J. Hoon. Ent., 43:139-145.
BELSCHNER, H. G., 1946.—The Buffalo Fly—A Survey of the Position and Discussion of Control
: Measures. Agric. Gaz. N.S.W., 57:149-152, 211-214, 218.
Cooter, €. E., 1947.—International Air Commerce and Plant Quarantine. J. Hecon. Hnt.,

40: 129-132.
—, 1940.—Plant Quarantine Activities in the Pacific, with Particular Reference _to the

7)
Oriental Fruit Fly. Bull. State of Calif. Dept. Agric., 39:10-16. é
Deputy, O. D., 1948.—Are Foreign Plant Quarantines Worth While? J. Econ. Ent., 41: 528-531.
Essig, EH. O., 1948.—Insect Surveys in Relation to Quarantine and Control of Insect Pests.

J. Econ. Ent., 41:673-677.
HANDSCHIN, H., 1932.—Investigations on the Buffalo Fly and its Parasites. O.S.I.R. Pamphlet

No. 31 (Australia).
HuGHEs, J. H., 1949.—Aircraft and Public Health Service Foreign Quarantine Entomology.

U.S. Pub. Health Reps., Supplement 210:1-38.
LairpD, M., 1948.—Reactions of Mosquitoes to the Aircraft Environment. Trans. Roy. Soe.

N.Z., 77: 93-114.
McBribe. O. C., et al., 1950.—Treatment of Airplanes to Prevent the Transportation of Insects.

J. Hoon. Ent., 43: 66-70.
Miturr, D., 1948.—In Report of the Fifth Commonwealth Hntomological Conference, p. 68.

(New Zealand Pests.)
NicHotson, A. J., et al., 1941.—Plant Quarantine. J. Aust. Inst. Agric. Sci., 7: 143-146.
Soper, F. L., and WILSON, D. B., 1943.—Anopheles Gambiae in Brazil. The Rockefeller

Foundation. New York.
STRONG, L. A., 1923.—Western Views on Plant Quarantine. J. Econ. Ent., xvi: 266-270.
Swezey, O. H., 1948.—Insect Invaders in Hawaii During and Since World War II. J. Econ.

Eint., 41: 669-672.
WARDLE, R. A., 1929.—The Problems of Applied Entomology. Manchester University Press.

504-508.
WHO ExPerT COMMITTEE ON MALARIA, 1948.—Report on the 2nd Session of the Expert
Committee on Malaria of the Interim Commission. Bull. WHO, 1: 243.

s

The Honorary Treasurer, Dr. A. B. Walkom, presented the Balance Sheets for the
year ended 28th February, 1951, duly signed by the Auditor, Mr. S. J. Rayment, F.C.A.
(Aust.); and he moved that they be received and adopted, which was carried
unanimously.

A vote of thanks to the Acting Honorary Secretary (Dr. W. R. Browne) and the
Honorary Treasurer (Dr. A. B. Walkom) for their work for the Society was carried by
acclamation.

No nominations of other candidates having been received, the Chairman declared
the following elections for the ensuing year to be duly made:

President: Mr. A. N. Colefax, B.Sc.

Members of Council: Lilian Fraser, D.Sc.; Professor J. Macdonald Holmes, B.Sc.,
Ph.D., F.R.G.S., F.R.S.G.S.; Professor P. D. F. Murray, M.A., D.Se.; G. D. Osborne,
D.Se., Ph.D.; T. C. Roughley, B.Sc., F.R.Z.S., and A. B. Walkom, D.Sc.

Auditor: S. J. Rayment, F.C.A. (Aust.).

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

xx

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SEROLOGICAL STUDIES OF THE ROOT-NODULE BACTERIA.
Iv. FURTHER ANALYSIS OF ISOLATES FROM TRIFOLIUM AND MEDICAGO.

By Himary F. PURCHASE, J. M. VINCENT and LAWRIE M. WARD,
School of Agriculture, University of Sydney.

[Read 28th March, 1951.]

Synopsis.

The present paper summarizes serological data accumulated over a period of about ten
years in respect of reference strains of Rhizobium trifolii and Rh. meliloti. Over this time
the antigenic properties of the organisms have shown a high degree of stability and the
technique has proved useful for the typing of isolates in field and laboratory studies.

To obtain adequate typing it is necessary to distinguish flagellar and somatic reactions.
The latter on its own permits more strains to be distinguished than the former, but maximum
differentiation requires both to be taken into account.

Extending the method of analysis used in the earlier papers it has been found that the
description of the 12 isolates of Rh. trifolii requires at least 2 flagellar and 9 somatic antigens.
The corresponding figures for the 16 isolates of Rh. meliloti are 4 and 15.

INTRODUCTION.

Since the application of improved agglutination techniques to the study of root-
nodule bacteria (Vincent, 1941, 1942) fair use has been made of these criteria for the
classification and identification of serological strains in field and laboratory studies
(Hughes and Vincent, 1942; Kleczkowski and Thornton, 1944; Vincent, 1944 and 1945;
Purchase and Vincent, 1949; Read, 1950, private communication). Whilst there is no
apparent relationship between serological constitution and the organism’s behaviour in
association with the host, the method remains a useful technique for distinguishing
and grouping strains, a means of studying field distribution, and for ‘labelling’ material
for laboratory and field studies.

A comparison of our recent results with those reported in the earlier papers has
shown a high degree of stability in antigenic properties; few cases have occurred
where a culture has shown any significant change in this regard. The antigenic
constitution appears in fact more stable than other characteristics (cf., for example,
Kieczkowski and Thornton, 1944; Nutman, 1946).

The earlier papers from this laboratory included reasonably detailed analyses of
several strains of both species. The number of fully studied strains has been added
to in the intervening years so that a fair battery of testing sera is now available. It
has been thought worthwhile, therefore, to record these further studies which provide
the basis of our more recent investigations.

HE}XPERIMENTAL.

Organisms used for the development of antisera.—The isolates are identified by
collection numbers and are mostly described in the earlier papers: Kh. meliloti (Vincent,
1941) and Rh. trifolii (Vincent, 1942). No, 204 has been added to the clover strains,
having been obtained as “Clover F” strain from Dr. H. G. Thornton, Rothamsted
Experiment Station, England. Rh. meliloti No. 52 originated from Medicago hispida var.
denticulata growing at Warracknabeal, Victoria. Additionally it is worth noting that
strains 7, 8, 10 and 12 came to us from the United States, the first three from. the
collection of the University of Wisconsin, and the last via the N.S.W. State Depart-
ment of Agriculture.

Methods—The methods of obtaining cultures and using them as antigens for the
development and testing of antisera have been largely maintained as in the first paper
(Vincent, 1941). It has been generally advantageous to obtain an earlier reading for
‘flagellar (H) agglutination at about one hour and advisable to check somatic (0)
agglutination with heated antigen, particularly in the presence of flagellar agglutina-

Cc

2 SEROLOGICAL STUDIES OF THE ROOT-NODULE BACTERIA. IV,

tion. Occasional difficulties have been encountered with a less voluminous flagellar
agglutination but, provided a two-day motile culture of sufficient density is used, the
result has been almost always satisfactory. We have not found it necessary to
distinguish H by the use of O-absorbed sera or by the removal of O antibody by heat,
although some workers might well prefer this added check (Kleczkowski and Thornton,
1944). /

RESULTS.
Strains of Rh. trifolii.—Tables 1 and 2 give the results for flagellar and somatic
reactions respectively.

Two major groupings of flagellar antigens are revealed, one represented by strain 36,
and the other by strain 46. Following the 1942 treatment these have been classed A and
B respectively. The earlier paper also showed by absorption tests that the five members
of the first group tested on that occasion were identical. Flagellar reactivity has now
been found between 157, 161 and) 46—a result at variance with the earlier findings
but supported by the agreement between reciprocal tests.

As has been found generally in these species, somatic antigens provide more
groupings than do the flagellar. On the basis of what we now know of the somatic
cross reactions of these strains the minimum number of somatic antigens would have
to be extended from three to seven. Absorption tests have been applied in some detail
to the 94-160 group of Table 2, and make it necessary to bring the number of antigens
to nine. Some irregularities seem to be associated with antigen II of 94, 61, 111 and 161

TABLE 1.
Flagellar Cross Reactions of Rhizobium trifolii.

Antigens. Sera.
l
36 108 61 111 94 91 160 204 | 46 157 161 64
| |
36 58 63.8 28 PA 2 22 2 2 —_ = ae =
108 | 33 38 39 7, 2 33 2 2 ines als dea ae a
61 3 33 38 22 2 23 2 2 | —_ = = a
111 25 Be 33 23 2 22 2 2 — nna aG ae:
94 he 2 3 2 3 2 2 2 | —_ - Ree ee
91 33 33 3a 23 2 Be 2 2 | —_ = = Z
160 33 2 2 1 2 2 3 2 Pee ae eS =
204 3 2 2 2 2 2 1 3 ) = - aie eas
ey el emee) een vee ete eur bath) APSO a og.
a eel ee ee URE eb Th gen
[SEE ES ES eee | |
LGD roe = = = = = - - ! Pea Fe 3 0-1
ah Se Sa es Mea sey 2 p>
|

Note-—Numbers indicate highest dilutions showing agglutination, viz. 1=1/25 to 1/50, 2=1/100 to 1/200,
3=1/400 to 1/800, 4=1/1600 to 1/3200 or greater.
Top right-hand figure shows earlier result.

BY HILARY F. PURCHASE, J. M. VINCENT AND LAWRIE M. WARD. 3

and indicate the possibility that the antigen might be composite and its components
variable in their relative proportions between strains and in the one strain at different
times.

The findings can now be summarized:

Minimal Antigenic Composition.

Isolate. Flagellar. Somatic.
36, 108 ie ag Ae pd A I
94 ay es ah a A IN ay WARE
61, 111 Ha a we is A Oke WY

160 A IV, VII
91 A TII

204 A IX
64 B I

161 B LO TViveVali
46 B TIL

B VIII

157

Sera of the six strains underlined have been used for typing field isolates
(Purchase and Vincent, 1949) and include representatives of both flagellar types and
eight of the somatic antibodies.

t

TABLE 2.
Somatic Cross Reactions of Rhizobium trifolii.

Antigens. * Sera.
36 108 64 | 94 61 161 111 160 91 46 204 157
l
! pavietolats ivy.
36 44 44 2 - — - ae. = = 25 S =
108 | 44 43 4 OR es Sg Sra = 2 ara ae
| Py eel EER aM Se
64| 34 | 4 a= == = = - —\| - =
a eS ae Baers ea. ooh eam ane bem omen
Gy ne IE es 42 | 3 By We = = oe Ue RY;
161} — | - - 3 Ornate u 3 a ee eet ee
Melee (eee, |= alls = 2 aoa Ne EO ce ne Fee seeen eee
ame Or AIS cohuan Ion 7 gee ee wie a lea
———— ee
91 —_ — = = 44 AGS ak 3 =
ig | | nee (gee bee Yee uel ellia rar oy eae
ADs | se je = - => hie a een. Pca iat
|

Note.—Numbers indicate highest dilutions showing agglutination, viz, 1 =1/25 to 1/50, 2=1/100 to 1/200,
3=1/400 to 1/800, 4=1/1600 to 1/3200 or greater.
Top right-hand figure shows earlier result.

4 SEROLOGICAL STUDIES OF THE ROOT-NODULE BACTERIA. IV,

TABLE 3.

Antigens. Sera.

7 | 8 | 12 | 62 | 51 | 52 | .66 | 74 | 76 | 102 | 126 | 184! 10 | 47 | 101

~I
~
>
1
i

=
rs
1
i
cs
w
rs
i

102 4 4 4 Cees 4 ARN Ni Aes eS 44) 4 4 ul ite = I

126 4 4 4 44 4 4 4 4 4 4 4 4 il =T || =

1349 | ae ape lira lwp Pal la la 0 liege on lane iy |e eee
0) 2 a el ee es ee
|p | a | mf || | ==
AT Ape Dias oD, col Sian Cette th |e Sa hare = al 2 ea 54 oa gs AS er
i (| atts EW fe ee eee | Pe ga ee at ee
|

Note.— + =variable result at lowest titre ; code otherwise as for Table 1.

Strains of Rh. meliloti—Tables 3 and 4 give flagellar and somatic reactions of
sixteen strains.

Most of the flagellar reactions are those previously seen in strain 27, but strain 10
has the major antigen of 47, and 101 is different from both ‘these groups. Using the
notation of the first paper, 10 and 47 are described as Ab, the 27 group as bC and 101
is now characterized as D. The symbol Bb is used to describe a minor antigen that is
commonly shared by members of the first two groups.

Somatic antigens provide a basis for further division. Approximate groups are
represented by:

7, 27 and 62, 47
52, 102
126, 8, 12
134, 74
66
10, 76, 101
51
although there is a certain amount of cross reaction beyond these limits. Cross
absorption tests have shown 7, 27 and 62 to be identical; they have, however, an
antigen additional to those of 47 and the latter has an antigen not common to them.
No. 76 also appears to have the same antigenic constitution as 101, otherwise there are
differences even within the members of the groups set out. The group represented by
126, 8 and 12 shows a wide range of reactivity when they constitute the antigen, but
outside the group itself reciprocal tests are mostly negative. It seems that the cells of

BY HILARY F. PURCHASE, J. M. VINCENT AND LAWRIE M. WARD.

or

TABLE 4.
Somatic Cross Reactions of Rhizobium meliloti.

Antigens. = Sera.
27
7 62 47 52 | 102 | 126 8 12 | 134 | 74 66 10 76 | 101 | 51
oe ea aces eat 2) Stes Ee pe een bee he fire Nes (NS Pe a ae |e
cae, || ee ea NDEs eS [a ev ree [est (5 Oi Eg as eo (ee
A le ee Ree ea
BB Bt ce dc Hee Maes aks ea aie (RR Ps a a ee
103 |) SEE SSS. S| ae Pe ee an ee
126 ea, Sipe pk eae 2 “9s! a oe ea 2 MS A 8} 2 es 3
8 = Lf Be a 4 d 4 3 PRE OX 2 3 3
12 ae SF Gh Oy 1°] 4 ho 5, & ae iss 4 2 3 3
reas ee ee ee Ee a gs! ai) = | sf 2) =
——— — —————| el —
Ame el lear | oe |p |) ed 201 1a so joe fos Ps
; | [ees pees (| | pe | ea eee Po eo ge me
—— a fa es eS eS SE
10 = = habs, ls vil aa (eee = = ae i | 4 3 4 1
«0 | 2 26) ee ee ae ee ae eee eee
mms a lea | ie,
sat ts I a aes ee ae ae ee

Note.— + =variable result at lowest titre, code otherwise as for Table 1.

this group are relatively easy to agglutinate, perhaps by a less specific antibody, and
only reactions that have reciprocal tests in agreement have been used in stating somatic
antigens. Cases 52 against serum 7, 66 vs. 134, 10 vs. 52, in which the reciprocal test
is negative, have also been ignored. Antigen II, previously postulated as common to
66 and 74, would, on similar grounds, be regarded as a doubtful or minor antigen.

The data for cross reaction and absorption tests can be symbolized thus:

Minimal Antigenic Constitution.

Isolate. Flagellar. Somatic.
47 ms e 6 2 Ab I, Ill

10 ee = ie ve Ab IX, Si

Ys Ol, A> oe ae a oe be I, IV

52 25 ee = ae be III, V, XII
102 bc III, V, XIII
126 bC V, VIII, IX
Pe be VIII, XIV

12 is an ae a bc VIII, XV
134 - By , ba be AW, IDs

74 be II, VI

66 bc II, VII

76 bc x

51 Le es a 2 be XI

101 Ae Sp ef. ah D XK

Sera of the strains underlined have been used in typing field isolates (Purchase,
Vincent and Ward, 1950). These include the three flagellar groupings and all the
somatic antigens so far revealed.

6 SEROLOGICAL STUDIES OF THE ROOT-NODULE BACTERIA. IV.

DISCUSSION.

These results emphasize the serological heterogeneity that exists within a ‘‘species”
of Rhizobium. There are notably fewer differences in flagellar reaction than somatic,
and distinction between the two types is obviously desirable for the more specific
recognition of a strain. It is interesting to observe how, with both clover and medic
isolates, the one somatic antigen can occur with any of the flagellar groupings.

Acknowledgements.
The work reported in this paper has been supported in part by grants from the
Commonwealth Research Committee and the Rural Bank of New South Wales.

References.

HuGHEs, D. Q., and VINCENT, J. M., 1942.—Serological Studies of the root-nodule bacteria.
Ill. Tests of neighbouring strains of the same species. Proc. LINN. Soc. N.S.W., 67:
142-152.

KLECZKOWSKI, A., and THORNTON, H. G., 1944.—A serological study of root-nodule bacteria
from pea and clover inoculation groups. J. Bact., 48: 661-672.

NutTMAN, P. S., 1946.—Variation within strains of clover nodule bacteria in the size of nodules
produced and in the effectivity of the symbiosis. J. Bact., 51: 411-432.

PURCHASE, HILARY F., and VINCENT, J. M., 1949.—A detailed study of the field distribution of
strains of clover nodule bacteria. Proc. LINN. Soc. N.S.W., 74: 227-236. =
PURCHASE, HiLtary F., VINCENT, J. M., and WArp, LAWRIE M., 1950.—The field distribution of
strains of nodule bacteria from species of Medicago. Aust. J. Agr. Res., 1951 (in press).
VINCENT, J. M., 1941.—Serological studies of the root-nodule bacteria. I. Strains of Rhizobium

meliloti. Proc. LINN. Soc. N.S.W., 66: 145-154.

, 1942.—Serological studies of the root-nodule bacteria. II. Strains of Rhizobiwm
trifolii. Idem., 67: 82-86. 3

, 1944.—Variation in the nitrogen fixing property of Rhizobium trifolii. Natwre, 153:
496-497. :

, 1945.—Host specificity amongst root-nodule bacteria isolated from several clover
species. J. Aust. Inst. Agric. Sci., 11: 121-127.

sl

A SEPTORIA DISEASE OF HUPHORBIA PEPLUS L.

By DororHy EH. SHAw,
Faculty of Agriculture, University of Sydney.

(Plate i; one Text-figure. )
[Read 28th March, 1951.]

Synopsis.

A Septoria disease of Petty Spurge is described, and the Australian distribution is given.
The cultural characteristics and morphology of the causal organism are described. The results
of investigations concerning the host-parasite relations, the longevity of the spores, the search
for the perfect stage, and pathogenicity tests with other plants are given. The literature
describing species of Septoria parasitizing species of Huphorbia is examined, and the name
S. pepli, n. sp., is proposed for the causal organism.

INTRODUCTION.

A leaf and stem spot disease of Huphorbia peplus L., caused by a species of Septoria,
was pointed out to the writer by Professor W. L. Waterhouse in 1947. He had had it
under observation for some time, and noted that infection on Petty Spurge was quite
widespread around Sydney. At his suggestion an investigation of the disease was
carried out, with particular reference to the host range, as some serious diseases of
economic plants are caused by species of Septoria.

EconomMic IMPORTANCE.

Huphorbia peplus L., Petty Spurge, native to Hurope and Asia, occurs on the New
South Wales coast and tablelands, and in Queensland, Victoria, South Australia and
Western Australia, as recorded by Hurst (1942). She listed its reputed medicinal and
photographic properties, but to the writer’s knowledge it is not used commercially in
Australia. Hurst also recorded it as containing a poisonous principle, euphorbin. It
is a weed of gardens and waste places, but is easily eradicated. The disease is not of
economic importance.

REVIEW OF LITERATURE.
Australian Records.

No mention of a leaf and stem spot disease of H. peplus was made by Cooke (1892)
or by McAlpine (1895). The disease was noted by Waterhouse (unpublished data) in
June, 1921, in the Sydney area. Pieces of material collected by him and embedded in
wax in 1921 are filed at Sydney University. Also filed are pieces of diseased material
in wax from a collection made in July, 1932, again from the Sydney area. A record
of a Septoria leaf spot disease of H. peplus was made by Noble et al. (1934), without
either date or locality of occurrence. The herbarium specimen lodged at the Department
of Agriculture, Sydney, is also without date of occurrence, locality or collector’s name.
A Septoria species igs recorded occurring on ZH. peplus in Brittlebank’s catalogue of
Australian Fungi (unpublished), compiled between 10th May, 1937, and 2nd March,
1940, but no indication of the date or locality is given, and no specimen is filed in
the herbarium.*

\ Overseas Records.

Diseases caused by six species of Septoria and two species of Rhabdospora were
recorded by Saccardo (1884, 1892, 19138)), on eight species of Huwphorbia, but none on
#H. peplus. Oudemans (1921) listed species of Septoria, Phleospora and Rhabdospora on
various Huphorbia species in Hurope, but none on H. peplus. No Septoria was recorded
for any Huphorbia by Grove (1935), although a Rhabdospora was noted on one species
of Huphorbia in Britain.

* Personal communication from Mr. S. Fish, Government Biologist, Department of Agri-
culture, Victoria.

8 A SEPTORIA DISEASE OF EUPHORBIA PEPLUS L.,

In a personal communication, Dr. G. R. Bisby, of the Commonwealth Mycological
Institute, reported that he could find only one record of a species of Septoria on
E. peplus, and that was of Septoria euphorbiae Guep. in the Russian book “Key to Fungi,
Vol. 2. Fungi Imperfecti’, by A. A. Jaczewski (1917). I am indebted to Dr. Bisby
for his translation of the significant paragraph: “p. 102. S. euphorbiae Guep. on
Euphorbia amygdaloides, E. peplus. Round, olive-coloured spots. Stylospores 40 to 454
by 2 to 2:54 with 3 to 4 indistinct septa.” Dr. Bisby also reported that no Septoria
was recorded on Huphorbia spp. for North America by Seymour (Host Index of the
Fungi of North America. Cambridge, Mass., 1929), this volume being unavailable to the
writer.

No Septoria on any species of Huphorbia-was recorded for South Africa by Doidge
and Bottomley (1931), by Brien (1939) for New Zealand, or by Shigekatsu Hirayama
(1931) or Nakato Naito (1940) for Japan.

AUSTRALIAN OCCURRENCE.

The disease occurs on Petty Spurge over a wide area around Sydney, and collections
were made throughout the area extending from Mangrove Mountain, near Gosford, in
the north, Windsor in the west, and Mt. Keira, near Wollongong, in the south, in
1948 and 1949. In 1950, two further collections were made at Canberra, A.C.T., and
Wagga, N.S.W.

The disease has not been recorded for Western Australia, South Australia, or for
Queensland.* It is not known whether Brittlelank’s record was for Victorian occurrence.
In Tasmania in 1948 the writer collected diseased Petty Spurge plants at Launceston,
Hobart, and Port Arthur.

From the information available at present, the disease is known to occur around
Sydney, at Wagga, Canberra and throughout Tasmania.

APPEARANCE OF THE DISEASE.

Lesions first appear on the leaves as small, mostly circular areas, pale green in
colour, later spreading and turning pale yellow, then light brown, becoming papery
and covered with scattered black pycnidia. Many lesions per leaf were recorded (in
one case 14 apparently separate ones), but the majority of field specimens examined
showed that infection was from one centre only, although sometimes two were observed.
The pycnidia occur on both sides of the leaf, but mostly on the undersurface. A few
lesions examined had a total lack of pycnidia on the upper surface, although numerous
ones were found on the lower. In some cases a slight zonation of pycnidia occurs
radially from the centre of infection. The pycnidia are produced singly, only rarely
being grouped in twos or threes. As the necrotic tissue enlarges, slight puckering often
occurs between this area and the rest of the leaf. The edge of the necrotic area is
generally quite regular and definite, but without cicatrix formation. Pycnidial production
is usually confined to the necrotic area, but occasionally lesions were found where
pycnidia occurred at the edge of the lesions in advance of necrosis. Where several
lesions occur per leaf, the diseased areas later coalesce. In the advanced stages of the
disease premature leaf fall occurs.

Lesions on the stem occur less frequently than on the‘leaves. The leaf infection,
in many cases, extends to the petiole and thence to the stem. Pycnidia occur on both
petioles and stems. Stem infections were produced in the glasshouse, which were so
severe as to cause death to the uninfected upper part of the plants.

Pycnidia were recorded on the seed capsules on material from the field, as well as
on plants inoculated in the glasshouse. No pycnidia were noted on the seeds.

CULTURAL CHARACTERISTICS.
Isolation Methods.
Isolations of the causal organism were first made by the usual method of tissue
transplants, pieces of diseased material of about 3” x 3” being cut from the edge of

* Personal communications from Mr. W. P. Cass Smith, Government Plant Pathologist,
Western Australia, Mr. D. B. Adam, Department of Plant Pathology, University of Adelaide,
and Professor D. A. Herbert, University of Queensland.

~

BY DOROTHY E. SHAW. 9

the lesions, or just in front, and taken through 95% alcohol for 5 seconds, mercuric
chloride 1 in 1000 for 20 seconds, and three washings with sterile water, then transferred
with sterile forceps to freshly poured and cooled plates of potato dextrose agar. It
was found, however, that if a contaminant present had escaped the surface sterilization,
it usually grew at a rate much in, excess of the Septoria mycelium, which was sub-
sequently difficult to obtain in pure culture. The method later adopted was to allow
the spores to exude from the pycnidia in the leaf, into a drop of distilled water, and to
streak a loopful of the spore suspension over the surface of the P.D.A. plates. Bacterial
and fungal contaminants were sometimes present, but because of the scattered nature
of the Septoria spores isolations were not difficult. Transfers were then made to P.D.A.
slopes, of single spores or hyphal tips, as well as mass transfer of mycelium. Nine
isolations were made from collections, five from within the Sydney area, three from
Tasmania, and one from Wagga. Nineteen collections were made from many diseased
plants observed in the field, three of these being from Tasmania, one each from Wagga
and Canberra, and the rest from a wide area around Sydney.

Media.

The fungus grew on P.D.A., water agar, standard agar, maize meal agar, maize
husks and maize cobs, potato cubes, lucerne shoots, lima bean agar, dried pea agar,
lentil agar, soaked rye and peanut husks, also on Czapeck-Dox with marmite, and wort
agar. It also grew on a water extract of H. peplus leaves in agar, and on sterile shoots
and stems of H. peplus. Growth on water agar was very slow and meagre. Pyenidia,
mostly well formed, were produced on all the media, often with abundant pinkish
exudate of spores, especially on the sterile stems of Petty Spurge, and on the rye-peanut
husk media. P.D.A. proved the most satisfactory media for mycelium and spore
production and for maintaining cultures.

Appearance of Cultures.

Spores germinate on P.D.A. usually within twenty-four hours. After three to four
days the colonies are visible, being whitish in colour, and distinctly mucose, with a
smooth surface and an entire edge to the naked eye. As the colonies develop, the
edge becomes slightly irregular and the surface slightly ridged. At seven days most
colonies have become black in the centre, and at 17 days are all black, sometimes with
a tinge of greeny-grey, with a minute border of white at the edge.

On P.D.A., cultures later produce small greyish patches of very short, pile-like
aerial mycelium. Black carbonaceous pycnidia are embedded over the surface, and
these, in most cultures, give pink exudates of masses of spores. Where pycnidia occur
near the side of a tube, cirri can be easily distinguished under the low power of the
microscope. These spore masses are yeast-like in appearance to the naked eye. The
mycelial mass later becomes carbonaceous, piled and convoluted in the centre, and with
sub-surface hyphae at the edge. The hyphal mass remains very compact and difficult
to separate, with growth upwards nearly as great as lateral growth. Seldom is the
whole surface of the slope covered with mycelium, and often cracks occur in the agar.
After about three months small tufts of white cottony mycelium appear in patches
over the surface in some slopes. Colonies are usually non-sporulating about this time.
In some cultures more than four months old, hard carbonaceous bodies appeared on the
surface and along the cracks in the agar. The structures, when sectioned after fixing
in chrom-acetic and stained with gentian violet-orange G, were roughly circular to oval,
but measured anything from 200 to 5004 wide (measuring from the innermost walls).
The interior had no definite structure, but consisted of wispy strands of fungal material,
which could not be distinguished as hyphae. The walls consisted of 6-9 layers of very
dark brown, thick-walled, pseudoparenchymatous cells, which pass into a region of
brownish, twisted, strongly septate, clearly distinguished hyphae. No development
further than this sclerotial-like stage was observed.

Optimum Temperature.
Optimum temperature was determined by growing the fungus in small (24”) Petri
dishes on P.D.A., and incubating over a range of temperatures. Three tests were

D

10 A SEPTORIA DISEASE OF EUPHORBIA PEPLUS L.,

carried out at different times. In the first two tests duplicate plates were used, in the
third only one plate was inoculated for each temperature, except for 20°C., when dupli-
eate plates were used. All the tests gave approximately the same results for the
temperatures available. Inoculum was of the Pennant Hills isolate, approximately
2 mm. square, and readings were taken at four weeks. The readings for the third
test are given, as the range available at that time was alittle more comprehensive.

TABLE 1.
Growth of the Causal Organism at Various Temperatures.

Temperature. Colony Diameter. Spore Production and Character
LOR mm. of Growth.
3 DX 92 No growth discernible.
5 | Dire vl Very slight growth.
10 7x 9 Colony black with small white margin; no
spores.
15 13 x14 Colony similar to above ; no spores.
20 16x17 Colony black with greenish tinge; on the
(mean of 2 reverse side, clumps of pyenidia in con-
plates) centric circles; abundant spores.
25 10 x12 Colony black; no spores.
30 2x 2 No growth discernible.
Room temp. (light) 13 x16 Colony as at 20° C.; abundant spores.
Room temp. (dark) 12 x13 Colony as at 20° C.; very abundant spores.
(June-July)
|

Optimum temperature for mycelial growth and spore production is around 20°C.
Colonies at the other temperatures might have produced spores when older. No
measurement was made of the height of the colonies, but this increased with increase
in diameter. :

Optimum pH.

Optimum pH was determined by growing the fungus on plates of P.D.A. adjusted
with N/5 NaOH or N/5 HCI to give a series varying in hydrogen-ion concentration. Two
tests were conducted, and the pH was read on a Vane Electronic pH Meter. Duplicate
plates were used in the first test, triplicate plates, for the second. Readings of pH
were taken before inoculation. Inoculum was of the Sandy Bay (Tasmania) isolate,
approximately 2 mm. square. Plates were kept at room temperature during May and
June, and measured after seven weeks.

TABLE 2.
Growth of the Causal Organism at Various
Hydrogen-ion Concentrations. Z

|

Colony Diameter.
pH. mm.
(Mean of 3.)

3°6 6-0x 9:3
4°8 23°6 x 25:3
5°6 26:0 x 29°5
6°8 24°3 x 26°3
(eT 22-0 x 24-0
8-6 16:0 18-6

Optimum pH is about 5-6 or a little more alkaline. Specificity is not very marked
over a wide central range, the greatest effect being shown on the very acid side at the
concentrations taken.

BY DOROTHY E. SHAW. 11

TEST FOR PATHOGENICITY.

Isolations of the organism were made as outlined previously, and the fungus
maintained on P.D.A. slopes. Spores from pycnidia produced in culture were used to
inoculate leaves of Petty Spurge seedlings in the glasshouse as outlined under ‘Host-
parasite Relations, Method’. Typical symptoms of the disease appeared on the leaves,
with production of pycnidia. Isolations made from these lesions yielded the organism
which was similar in detail to that isolated initially.

MorPHOLOGY OF THE CAUSAL ORGANISM.
Mycelium.

The mycelium in young colonies consists of stellately radiating hyphae, which are
minutely guttulate, hyaline, septate, and branch profusely. Pigmentation occurs in
hyphae about six days old. Old hyphae (i.e., after a period of months), become very
nobbly in outline, olive in colour, often with large refractive globules, but usually
without any apparent contents. Young hyphae at the edge of colonies are very fine.
Mycelium in colonies 13 days old measured up to 3u in diameter, and at four months
measured about 4u in diameter. Hyphae in the plant tissues measure 2-3 in diameter...

Pycnidia. .

Pycnidia occur on both sides of the leaf, but mostly on the under-surface, scattered
over the centre and sometimes on the marginal green areas of the necrotic spots;
singly, only rarely in twos or threes; visible to the naked eye, black in colour, but
reddish-brown by transmitted light; globose, immersed but later erumpent; ostiole about
one-quarter the diameter of the pycnidium; pycnidial wall smooth, composed of 2-3
layers of pseudoparenchymatous cells; 85-135u (65—-160u), mean 110-194 + 18-98y.

Pycnidiospores.
Pyenidiospores hyaline, straight to very slightly curved, attenuated at one end, with
a varying number of guttulae; usually three septa, often two, sometimes one, rarely
aseptate or four-septate; 25—44u (17-51lu), mean 35-77u + 5-94u.

Cirrt.

Cirri were often found exuded from the ostioles in material examined straight from
the field, and practically always from material in the glasshouse, owing to the high
humidity. They are colourless under reflected light, and in strong sunlight can be
seen with the naked eye as a glistening whitish spot on the pycnidium. The horns
vary in length, nearly always curl over, and often adhere to the neighbouring horns.
On diseased material from the glasshouse, cirri were noted from quite small pycnidia.
Cirri were observed microscopically by placing diseased stems from the glasshouse
(with cirri still attached) into lacto-phenol cotton-blue, and by allowing pycnidia
produced in culture to exude spores into dilute lacto-phenol. The pycnidiospores are
oriented with their long axes parallel to the horn. When material with cirri already
exuded is placed into water the spores immediately separate from one another. When
pycnidia with unreleased spores are placed in water, the spores are ejected separately,
not exuded in cirri.

OBSERVATIONS ON PYCNIDIOSPORES.
. Spore Structure.

Pyenidiospores of all isolates examined were hyaline, the septa being indistinguish-
able unless stained. Stains used included cotton-blue lacto-phenol, aqueous gentian
violet, nigrosin, aceto-carmine, and gentian violet, orcein and carbol fuchsin in lacto-
phenol. Cotton-blue lacto-phenol proved -to be the best stain to show cell contents.
This stain acts quickly, the protoplasm staining blue of varying intensity. Guttulae
were of various sizes, sometimes two large ones appearing in each cell, one at each end
near the septa, sometimes one large guttula only, as well as numerous small ones. The
number of large guttulae was not constant per cell or per spore. Septa were unstained
with gentian violet, the spore contents appearing granular with large refractive drops
in most cells. In material fixed, sectioned and stained with gentian violet-orange G,

12 A SEPTORIA DISEASE OF EUPHORBIA PEPLUS L.,

nuclei were clearly distinguishable in the spores and conidiophores. Spores mounted
in water and examined with dark field illumination, showed the cell contents as
circular areas of light of varying size and intensity, as shown in Plate 1, B, by
MacMillan and Plunkett (1942).

Septation of Spores.

Because of the confusion in the literature regarding the number of septations in
most species of Septoria, this aspect of the causal organism was particularly observed.
MacMillan and Plunkett (1942), following on the work of Garman and Stevens* (cited
by the afore-mentioned but unavailable to the writer), have shown that there is a great
inadequacy on this point for most published descriptions of Septoria species. Sprague
(1944) and MacMillan and Plunkett consider that the number of septations of the
spores is usually 1, 3 or 7, depending on whether there are 1, 2, or 3 nuclear divisions
in the spores, and that there is simultaneous division of the end cells, after formation
of the primary septum. MacMillan and Plunkett concluded that the spores become
mature on attaining the 3-septate condition, but not before, and that an even number
of septa in the spores is anomalous. These authors found it difficult to account for
the very large number of observations in the literature for even-numbered septate
spores. Sprague (1944) found that some species infecting grasses are characterized by
2-septate spores, while five septa are common in others. He concluded that in some
species the number. of nuclear divisions evidently is dependent on the available cell
nutrients; that large spores produced in humid winter weather may have two or even
three nuclear divisions, while later in the season, when the weather is warmer and
drier, the same species may produce aseptate or 1-septate spores.

Septa are indistinguishable in unstained spores. Stained with cotton-blue lacto-
phenol, the septations are visible, but the granular nature of the cells does not make
for easy observation. The best method found was as follows: diseased material with
pycnidia was placed in a drop of water on a slide, and the spores allowed to exude.
After a minute or so, the tissue was lifted from the slide, and a drop of iodine in
potassium iodide in 80% alcohol was added, this being a modification of the method
used by Brodie and Neufeld (1942). The cover slip was lowered, and the septation
counts made immediately. The spores, with this method, appear a homogeneous golden-
brown, with the septa, under slightly reduced light, clearly distinguished.

TABLE 3.
Septation of Spores from Different Collections, at Various Intervals from Time of Collection, and from Culture.
|
Percentages of Spores in the Various Septate
No. of 1 } Classes.
Collection. Spores Time from. | be ty $
Examined. | Collection. |
| 0 1 2 3 4 | 5
weiss pian Ae | |
|
Mangrove Mtn. ue ie 200 2 weeks. — 10-0 28-5 60-5 1:0 —-
Mt. Keira cas 4S 200 2 months. (Nal 7) “al7/9t) 37-0 44-5 0-5 —
P. Arthur, Tas. .. ec 200 4 months. — | 19-0 46:5 33-0 1-0 0-5
Sandy Bay, Tas. .. “3 200 4 months. — 10°5 36-0 53°5 aS
Sandy Bay, Tas. .. Be 100 6 months. ae 6-0 15-0 79-0 — _—
*Sandy Bay, Tas. .. is 142 == [Pere Pea net) 46-0 52-1 say ih ce
Penshurst .. WG AG 100 | 11 months. 1:0 | 21-0 38-0 40-0 | — —_
| } ' |

* Spores from culture.

The results shown in Table 3 indicate that the maximum number of septa in
mature spores is three, practically no further division taking place until germination.
The first cell-division in the spore would appear to take place early, because of the
very small number of aseptate spores which appear in the counts. Spores in which

* Garman, P., and F. L. Stevens, 1920.—‘‘The genus Septoria.” Ill. State Acad. Sci. Trans.,
13° 2176-219.

BY DOROTHY E. SHAW. 7 a2:

two cell-divisions have taken place (giving four-celled spores) would most probably be
spores produced under optimum conditions. The large percentage of 2-septate spores
could only be explained where only one cell divided after the formation of the primary
septum. It is to be noted that a large number of 2-septate spores was also produced in
culture. Counts from all the above collections are of the same general type, i.e., 3>>2>1-
septate spores (except the Port Arthur isolate, where the 2-septate spores predominate),
with 0-, 4-, and 5-septate spores occurring only very rarely. The relative percentage of
3-, 2-, and l-septate spores is most likely conditioned by nutrition. Age, under the
conditions of storage of the material, did not appear to affect the relative percentages
greatly.
Germination of Spores.

1. Method and manner of germination.

Germinations were studied on plates of P.D.A., the media having been strained,
and only that amount which would just cover the bottom poured into Petri dishes.
Spores from a water suspension were streaked over the surface with a loop. With this
method, each streak of spores could be followed under the microscope with ease.
Germination conformed in general to that described by MacMillan and Plunkett (1942)
for 20 representative spores of NS. apii-graveolentis. These workers noted that, at the
end of four hours, the two end-cells had divided, and the tapered end of the spore had
increased in length more than the other cells. At twelve hours the end-cells had
divided, and at 24 hours the two original centre-cells had divided, and the outer cells
of this division had sent out tubes. The spores were originally all 3-septate, and the
germination was quite symmetrical.

In the present study, it was noted that germination was not always symmetrical,
and counts were made of the positions of emergence of germ tubes. Readings were
taken on 200 spores from the Penshurst collection, taken at random for both readings,
on P.D.A.

TABLE 4.
Germination of Spores at 24 and 50 Hours.

24 Hours. 50 Hours.
Type of Germination. % %
Not germinated .. Me aS ae Ee a Bs. 4 3
Germ tube at 1 end .. ais Ge Bs ss x 38 it
1 side ee is i ne ae 6-5 1
both ends sf <5 . ee 33 11
both sides Ne a =e a8 1:5 2
1 end, 1 side .. ae oe ff at 12-5 1
2 ends, 1 side .. 5 4 1V
1 end, 2 sides oe A oe 0-5 9
both ends and more than one side —- 55

. As indicated in Table 4, emergence of germ tubes was asymmetrical, although by
the second reading at 50 hours, emergence had proceeded towards the “normal” type
expected from a 3-septate spore, as shown by MacMillan and Plunkett. Spores from the
Penshurst collection included many with one and two septations, and the asymmetrical
emergences were most likely a reflection of this condition. The above workers considered
that spores with less than three septations were immature, but it is to be noted that
spores in this study which would be classed as “immature’’ were capable of germination,
as shown in Table 4, where 96% had germinated in 24 hours.

Germinations on water agar were slower than on P.D.A. (e.g., in one test, 94% had
germinated on P.D.A. in 48 hours, against 74-5% on water agar), and hyphae were
shorter, with fewer branches. Fer this reason, strained P.D.A. was used in preference
to water agar, although the latter is a little clearer. Germinations in water were
comparable with those on P.D.A. Spores retained their identity in most cases, up to
approximately 48 hours on P.D.A., but for a shorter time in water.

14 A SEPTORIA DISEASE OF EUPHORBIA PEPLUS L.,

No secondary spores borne directly on the mycelium were observed, as noted by
Weber (1922) and Sprague (1944) for several species of Septoria. Anastomoses
between germinating spores occurred in water, but none were observed on P.D.A. or
water agar.

2. Effect of Temperature on Germination. ‘

Several loopfuls of a spore suspension in water were placed on coverslips, which
were then inverted over van Tiegham cells containing several drops of water, and sealed
with vaseline. After seven days cotton-blue lacto-phenol was run in, to fix and stain the
hyphae, the coverslips were removed and placed on clean‘slides for observation.

TABLE 5.

Germination of Spores and Degree of Development at Different Temperatures.

} x
ey | | Room

ASIC a eee Sag Ce: Weal olG-cem eeenem ps | ZONC: BOSC | PeesiiaGs
ho) aiuikys | |
| | | |
| | |
| |

He srcn Been ceearest ar ules Ear gamer a ok - ~ ;

| | |

The results of the temperature test for germination agree fairly well with that for
vegetative growth, as in Table 1.

Spore Size.
1. Length.

Great discrepancy exists in the literature regarding methods employed for mounting
and measuring, and methods of recording measurements, and the position has been
reviewed by Bisby (1945) and Ramsbottom (1948). There are also wide differences in
opinion as to the number of items, e.g., spores, which are required to be measured.
Many workers have not recorded the number measured. Bisby (1945) suggested 20 or
so, to include spores at both ends of the range. Cochrane (1932) measured 1,000 spores
of each of the two species studied. Beach (1919) measured 200 spores for each test
when checking the effect of various microclimates on the length of Septoria spores.

In this study, spores were allowed to exude from pycnidia into a drop of water on
a slide, for not longer than two minutes, when lactophenol cotton-blue was added, to
prevent swelling and to stain the hyaline spores. One thousand spores were measured,
from different isolates and environments, and the results examined to see if there was
any difference between the spores of seven collections, between the Tasmanian and
Sydney area collections, and between the spores from diseased material and those from
culture. In some of the collections, material was limited.

Examination of the results in Tables 6 and 7 shows that the means range from
32u to 41u, and variation between groups is no greater than variation within: groups.
The greatest number of long spores was produced in culture. Other workers have
found that even slightly different environments affect the length of Septoria spores,
e.g., MacMillan and Plunkett (1942), found that spores of NS. apii-graveolentis from
pycnidia measured 36-8u (average of 50 spores), while spores from cirri measured 44-5u
(average of 20 spores). Moore, cited by Hughes (1949), found that the average length
of spores of S. lactucae increased from 27-5u to 35u after a week of dull or wet weather.
Beach (1919), measuring 200 spores for each test, with S. tritici and S. verbascicola,
found that size was affected by environmental conditions such as bright and dull
light, moist and dry culture conditions, and summer and winter development. While
the readings shown in Table 6 are not sufficiently comprehensive for a detailed
comparison, it is considered that the differences in length of spores of the various
isolates can be accounted for by differences in environment at time of spore production.

BY DOROTHY E. SHAW. 15

TABLE 6.
Length of Spores from Different Isolates.
| | Standard
Isolate. Month of Source.* No. } Mean. Deviation.
Collection. | Measured. | UL. UL.
| | |
| |
Sandy Bay, Tas. .. a iy .. | January. Ne 200 | 32-38 5-100
Sandy Bay, Tas. .. i ft .. | January. Cc. 100 | 33°46 4-676
Port Arthur, Tas. i, me .. | January. N. 200 35-60 5-821
Taunceston, Tas. .. as hs .. | January. N. 200 | 37-37 4-901
Pennant Hills (7) .. 3 Ka so |) IME C | 100 41-11 5-091
Rydalmere = nee sa ots .. | May. C. | 50 39-71 4-525
‘Penshurst .. ae ae a Sanlediutliys N. 100 33.52 6-225
Pennant Hills (9) .. ee be .. | August. | N 50 38-08 3:°994

* N.=nature, C.=culture.

TABLE 7.
Comparison between Groups.
Bulked
Tasmania Sydney Area | Mean Stan. Dev.
UL. 5 U- | U- U-
Nature .. ais 32°38 3 Snes 35-01 5-778
35-60 38-08
37-37 ; 41-11 |
Culture .. Ae 33°46 39-71 37°77 5:775
|
Bulked .. he 34-88 5-564 37°84 6-184 | 35-77 5-939
Mean Stan. Mean Stan. Mean Stan.
Dev. - Dev. Dev.

2. Width.

Spores were prepared for measuring as described under the previous section. Three
hundred spores were measured, at the widest part, using a 20x eyepiece with a calib-
rated ocular micrometer, and fifty were measured using a filar micrometer.

TABLE 8. “
Width of Spores.
Standard
Method. Isolate. — Source. Number Mean. Deviation.
‘ Measured. UL. (2.
Filar micrometer a .. | Penshurst. N. 50 1-8975 0-2204
Ocular micrometer .. .. | Sandy Bay. C. 100
Sandy Bay. N. 100
Launceston. N. 50 oe Ose
Pennant Hills. C. 50

No difference between isolates was detected. The use of the calibrated ocular
entailed making estimates for those spores whose widths fell between graduations.
Although greater accuracy is obtained by using the filar micrometer, whose graduations
are only 0-178u apart, the former method is quicker, and is sufficiently accurate to

16 A SEPTORIA DISEASE OF EUPHORBIA PEPLUS L.,

warrant its use under most circumstances, if a large number of spores are to be
measured.
SIze OF PYCNIDIA.

Diseased material was mounted in lactophenol to prevent shrinking and swelling,
and a coverslip gently lowered on top to avoid squashing. Measurements were made
from the outside of the pycnidial wall on those pycnidia whose ostiole was uppermost,
i.e., if the adaxial surface of the leaf was uppermost, only adaxial pyenidia were
measured. Readings for each collection, however, were made on pycnidia on both sides
of the leaf. After a preliminary examination of the pycnidia, it was decided that their
practically spherical formation required only one measurement. Where length and
width differed, the diameter measured was the one which happened to lie parallel with
the graduated scale of the eyepiece.

TABLE 9.
Size of Pycnidia from Different Isolates.
| iyi
No. of Standard
Tsolate. | Pycnidia Mean. | Deviation.
Measured. {L. {L.
| |
Sandy Bay, Tas. Eyeva. won ye 200 117-55 | 18-694
Port Arthur, Tas. ee an oF el| 100 107-44 | 18-712
Pennant Hills (7) ae hd Be 50 102-34 15-760
Rydalmere ve Bs eck Ke 100 110-33 17-847
Penshurst ue th ae sh. 50 104-04 18-241
Pennant Hills (9), 56 as ast 100 105-06 | 18-093
Bulked .. *. a ah al 600 110-19 i 18-982
a \

As shown in Table 9, the mean of the bulked readings was 110-194 = 18-982u.
Measurements of pycnidia grouped in twos and sometimes in threes fell within the range
of solitary pycnidia, and are not included in the above figures.

The mean of 50 measurements of ostioles of pycnidia on diseased leaves was 29-24u,
ranging from 17 to 47-6xu.

Host-PARASITE RELATIONS.
Method.

Seedlings of HE. peplus, raised from seed, or transplanted from the field, were grown
in pots in the glasshouse. Leaf surfaces were difficult to wet, and the best method
proved to be moistening with the fingers, followed by spraying from an atomizer.
Several loopfuls of inoculum were placed on both surfaces of the leaves, the inoculum
consisting of a water suspension of spores, either from pycnidia on leaves, or from
pycnidia in culture. Seedlings were incubated from 48 to 72 hours, then placed on
benches in the glasshouse.

Leaves were picked and fixed after 24 and 48 hours, and thereafter every two days
until pyenidia were abundant. Several methods of fixing and staining were tried,
including staining with Pianese 111) as recommended by Weber (1922), but the method
finally followed was as follows: leaves were fixed in Farmers (absolute alcohol and
glacial acetic acid 3:1) for 20 minutes, washed several times in 95% alcohol,
decolourized in 95% alcohol for 18—24 hours, and either mounted in lactophenol cotton-
blue permanently, or taken from it to lactophenol after ten minutes. This method was
found to be very satisfactory. The leaf tissue after treatment was quite colourless, and
the spores and hyphae on the leaf surface stained. blue, standing out vividly. Spores
in pycnidia stained in deepening amounts of blue, depending on the maturity. Hyphae
in the tissues were quite colourless when first mounted, but after several months
appeared faintly blue. Material mounted in this way and examined after fourteen
months was in good condition. The hyphae were clearly visible below the epidermis,

BY DOROTHY E. SHAW. iW

and although the chloroplasts by this time were stained blue, this assisted observations
by making cell boundaries easily discernible.

Stem tissue was stripped, fixed, stained and mounted as above.

Observations.

Spores had germinated on the leaf surface after 24 hours, usually from one end,
sometimes from both, more rarely from a side branch. The infection hyphae were
usually not much thinner than the width of the spore. Many germ tubes terminated
in small, appressoria-like bodies, usually at the junction of two epidermal cells. In
many cases hyphae were observed to pass alongside or over stomates. No entry was

Text-figure 1.

Camera-lucida drawings of spores germinating on the leaf surfaces. x 300. A, B and C,
spores germinating on the upper surface. D, fusion of spores germinating on the under
surface, with germ tube passing by stomate. BH, spore germinating om the under surface, with
germ tube passing over stomate. F, multiple fusion of spores germinating on the uppper leaf
surface.

observed through stomates, and there appeared to be no attraction for the hyphae to
do so, One case only was observed where the hypha ended at a stomate, and in that
ease there appeared to be further development.

Germination was similar on both leaf surfaces. Branching was observed only
rarely, and spores retained their identity longer than on P.D.A. The number of septa

18 A SEPTORIA DISEASE OF EUPHORBIA PEPLUS L.,

did not increase in most spores during germination. Anastomoses were frequently
observed between two spores, sometimes between three spores.

At a later stage, hyphae were observed in the leaf tissue, spreading out radially
from the infection centre, branching and passing beneath the epidermis, and around
the cells, mainly in the mesophyll region. Later, knots of hyphae were noted below
the stomates, sometimes not directly beneath, but always in the sub-stomatal cavity.
On no occasion did a knot develop on one hypha only: pyenidial formation therefore
is probably symphogenous.

On a few leaves, notably in one collection kept wrapped in moist paper for 24
hours, hyphae were observed issuing from the stomates above immature pycnidia. These
“aerial” hyphae measured approximately 3u wide, and were from several to 70u long.
In many cases not just one hypha projected from the stomate, but two or three, and
some cases were observed where bunches of five to eight short hyphal tips projected
through the stomates.

Pyenidia were apparent to the naked eye after about 14 days. The most mature
pycnidia occurred at the centre of the lesions, and radially from these were immature
pycnidia, aggregations of hyphae under stomates, and then hyphae ramifying through
the tissue. Hyphal tips were quite distinct, and measured about 3 across, being just
slightly wider than the older hyphae. Tips were located up to 500u in advance of
pycnidia formation. In the undiseased tissue, the chloroplasts appeared distributed
evenly around the cell walls, and remained so even with the advance hyphae passing
around them. In the region between the advance hyphae tips and the pyenidia, the
chloroplasfs lose their discreteness, and the cells appear collapsed. The boundary
between the normal-appearing cells and the collapsed cells was usually quite distinct.
Not all the stomates had pycnidia forming under them, but where a stomate nad been
missed in the first place, it often showed aggregations at a later stage, i.e., young
pycnidia could form in the zone of older mature pycnidia.

Strippings from stems showed a similar condition, with hyphae mostly running
up and down the stem, and pycnidia being more linearly placed, instead of zoned.
Pyenidial formation was symphogenous.

Examination of healthy Petty Spurge leaves showed that stomates occur on the
abaxial surface on the average of eight to a field (magnification x840), while very few
occur on the adaxial surface, most fields having none, or two or three, at certain parts,
e.g., along the sides of the main vein. This would account for the predominance of
pycnidia on the lower surface of the leaves.

TABLE 10.

Viability of Spores at Various Periods After Collection.

|

No. of |

Collection. Age at Test. Spores | Germinated.
Counted. | We
= = aoe

|
Mt. Keira Se 54 6 50 2 months. 170 94
Ban, 200 87
5 , 200 71
7 | 100 0
Mangrove Mtn. 7 ; | 200 | 12
Pennant Hills (9) 10 “ 200 0
Penshurst (ali is 300 0

Diseased tissue was also fixed in Flemming’s weaker solution and Farmers Fluid,
the former being stained with gentian violet-orange G, the latter with carbol fuchsin-
light green. Sections were cut at 8u and 10u. No cicatrix was ever observed at the
edge of the lesion. Hyphae ramified throughout the tissue, and the stages in pycnidial
formation were noted: hyphae loosely woven below the stomates, later developing into
a knot, followed by the development of the pyenidial wall. The pycnidia were subepi-

BY DOROTHY E. SHAW. 19

dermal, and later erumpent with a widened ostiole. Spores were produced on one-
celled pyenidiophores. Nuclei were clearly visible in sections stained with gentian
violet-orange G.

LONGEVITY OF SPORES.

Germination tests were made with spores from diseased material which, after the
initial test, was kept between paper at room temperature. Several loopfuls of the spore
suspensions were streaked across strained P.D.A. plates, and germination counts were
made at 48 hours. Spores from all isolates gave more than 95% germination at the
initial tests.

Spores retained within the pyenidium under the above conditions of storage were
viable for about six months.

SouRCE OF INOCULUM AND TRANSMISSION OF THE CAUSAL ORGANISM.

Diseased plants of Petty Spurge were found in the field’ throughout the year, with
the exception of late spring, and further search might have revealed infected plants
even during this period. These diseased plants, therefore, provide an immediate source
of inoculum. Pycnidia on fallen leaf and stem fragments could also liberate spores,
given favourable conditions of moisture, and provide an additional source of inoculum
in the absence of growing infected plants.

To determine whether the disease is seed-borne or not, tests were carried out as
follows:

A collection was made of the ripest capsules on heavily diseased plants at Penshurst.
In the laboratory, seeds (most of which had left the capsule cases) were picked aut
with forceps. The cases were floated and turned over in distilled water in watch
glasses and examined under low power. Many pycnidia were noted on them. The
cases were drawn up onto the sides of the watch glass with forceps, and the residue
water examined with reduced light. Many spores were noted. The seed was turned
over in distilled water in watch glasses and examined under low power. No pycnidia
were noted on the seed, but spores were found in the water residue. The seed was then
sown in pots containing sterilized soil, and the residue water with the spores sprayed
over the seed. The seedlings were examined every day after emergence, but no sign
of disease was apparent on the stems or leaves. Random seedlings were selected at
four weeks, washed in fast running tap water, and several times in distilled water, and
plated on P.D.A. No Septoria mycelium grew from the tissue. Capsules and seeds
from diseased plants at Allawah were examined as above. Pycnidia were found on the
cases, and spores in the washing, but no pycnidia were detected on the seeds.

Ripe seed capsules were collected from Petty Spurge plants in an isolated patch
at the University, the plants being apparently quite free from disease. The capsules
and seeds were examined as above, and as no pycnidia or spores were observed, the
seed was presumed to be clean. The seed was divided into four groups of about 40
seeds each, and treated as follows: 5

(a) Seed sown in pots in sterilized soil (control).

(6) Seed sown in pots in sterilized soil, and the surface of the soil sprayed
with a suspension of spores and mycelium of the Pennant Hills isolate.

(c) As (6), but from the Sandy Bay isolate.

(d@) Seed soaked for 24 hours in a spore and mycelium suspension, then sown
in pots in sterilized soil.

The seedlings were examined after emergence, up to a period of eight weeks, but
no sign of disease was observed on the stems or leaves. Random seedlings were washed
as above, and plated on P.D.A., but no Septoria mycelium grew from the tissue.

Seeds from. heavily diseased plants were surface sterilized and plated on P.D.A.,
but no Septoria mycelium developed.

Although the above tests are not conclusive, it seems unlikely that mycelium is
carried in the seed, and although spores are carried on the surface, no infection was
established from these. It must not be overlooked that spores in pyenidia on shed

@

20 A SEPTORIA DISEASE OF EUPHORBIA PEPLUS I., ©

capsule cases could constitute a source of inoculum, for example, if splashed up by the
rain on to the leaves of young seedlings.

SEARCH FOR THE PERFECT STAGE.

The perfect stage of some species of Septoria has been recorded, and the literature
was examined to note the environment favourable for perithecial production. Stevens
(1925) cited Klebahn, who recorded Mycosphaerella sentina as the perfect stage of
.S. piricola, occurring on over-wintered leaves of apple and pear. Stevens also recorded
Leptosphaeria phlogis as the perfect stage of S. phlogis. Roark (1921) showed that a
Mycosphaerella, occurring on over-wintered leaves and in pure culture (media not
given), was the perfect stage of S. rubri. Weber (1922) recorded Leptosphaeria
avenaria as the perfect stage of Septoria avenae Frank, perithecia occurring in oatmeal
agar and potato dextrose agar cultures. Stone (1916) showed that Mycosphaerella
grossulariae, occurring on dead over-wintered leaves of currant and gooseberry, was
the perfect stage of S. ribis, and that Mycosphaerella aurea, found on old leaves of
Ribis aureum, was the perfect stage of S. aurea. Thompson (1941) recorded
Mycosphaerella populorum as the perfect stage of S. musivad on poplars, and
Mycosphaerella populicola as the perfect stage of S. populicola, the ascigerous stages
occurring on over-wintered leaves. Ruggieri (1936) reported a Mycosphaerella as the
perfect stage of S. aqurantiorum, having obtained perithecia in pure culture. Klebahn
(1934) found that a Mycosphaerella on overwintered leaves of chestnut was the
perfect stage of S. castanicola. Wollenweber (1938) found perithecia of Sphaerella
linorum, the perfect stage of S. linicola, on flax straw in the Argentine. Johnson
(1947) found a Leptosphaeria on wheat, and rarely, barley leaves in Canada, and also
obtained perithecia on corn meal agar, this being the perfect stage of S. avenae
Frank F. sp. triticea. Cochrane (1932) could not find the perfect stages of S. a@pii or
S. apii-graveolentis, either in culture, on artificially wintered leaves, or in response
to treatment with ultra-violet light. 4

During the present study, all cultures on P.D.A. were periodically examined for
any evidence of the perfect stage, some slope cultures being kept for more than a year,
Cultures on the various types of media used, on P.D.A. in various environments, and
cultures used for temperature tests were examined, in case a necessary factor or
combination of factors giving optimum conditions for perithecial growth had been
supplied.

Cultures of the various isolates were opposed in all combinations on P.D.A., in
nearly all combinations on corn meal agar and on sterile lucerne shoots. These
cultures were kept for more than five months.

Heavily diseased leaves were placed on and just below the surface of soil
sterilized for four hours, in Petri dishes, and were (a@) held at approximately 2°C. for
four months, then at room temperature, or (0b) moistened periodically at room
temperature.

Diseased leaves on plants infected in the glasshouse were allowed to fall onto the
surface of the soil in the pots; some pots were kept in the glasshouse for six months,
others were put into the open, and the decomposing leaves examined from time to
time. A patch of heavily diseased plants was marked off in a garden at Hurstville,
and allowed to remain undisturbed. The diseased leaves shed on to the soil surface,
and later the stems, were examined. The Petty Spurge leaves are so delicate that they
do not retain their identity for long on the soil surface.

No evidence of a perfect stage was found under any of the above conditions.

PATHOGENICITY TESTS WITH OTHER PLANTS.

In order to determine the host range of the fungus, plants of the family
Euphorbiaceae, being species of Huphorbia and Ricinus, were inoculated in the glass-
house. The writer particularly desired to determine whether the species of Euphorbia
reported as hosts of species of Septoria were susceptible or not to the fungus under
study, but difficulty was experienced in obtaining seed, as most of the species
mentioned are of European-Asian habitat, and are not present in Australia. Black

BY DOROTHY KE. SHAW. 21

(1922) recorded H. exigua as present in South Australia, and the C.S.I.R. Bulletin
No. 156 (1942) listed H. exigua and H. Hsula as present in Australia, but seed of these
species was unobtainable at Sydney Botanic Gardens, Adelaide University (Waite
Institute), Melbourne Botanic Gardens and the Tasmanian University. Seed was also
requested through -the medium of the Australian Plant Disease Recorder, circulating
through all the States. Seed of H. Hsula, HE. exigua, EH. serrata, EH. silvatica, EH. palustris,
EH. aspera, H. angulata and EH. amygdaloides was requested from overseas institutions
by the Plant Introduction Office, C.S.I.R.O., and viable seed of H. Hsula and EH. exigua
was eventually obtained.

Plants not of the family Huphorbiaceae, but known to be hosts of other species of
Septoria, were also inoculated, together with other grasses, weeds and ornamentals.

The inoculum consisted of a suspension of spores, either from pycnidia on fresh
leaves, or from cultures of the various isolates. The plants were either transplanted
from the field, raised from seeds, or grown from cuttings. Seedlings of H. peplus

were inoculated at each test.

Plants inoculated were as follows:
Huphorbia helioscopia L., “Sun Spurge’.
E. Drummondii Boiss, “Caustic Weed’’.
E. Lathyris L., “Caper Spurge’.

EH. terracina L., “False Caper’’.

EH. splendens Bojer (H. Milli Desm.,
Sterigmanthe splendens Kl. et
Garcke), “Crown of Thorns”.

H. neriifolia L.

H. Bojeri Hook (Sterigmanthe Bojeri Kl.
et Garcke).

H. enigua lL.

H, Hsula Ll.

HE pulcherrima Willd. ex Klotzsch,
“Poinsettia”.

Ricinus communis L., “Castor Oil’.

Lycopersicon esculentum Mill., “Tomato”.

Linum usitatissimum L., “Flax”.

Pastinaca sativa L., “Parsnip’’.

Apium graveolens L., “Celery”.

Apium leptophyllum (DC) F. Muell.

Triticum vulgare Host. “Federation”
wheat.

Triticum monococcum L., “Hinkorn”
wheat.

Avena sativa L., “Richland” oats.

Hordeum vulgare L., “Kinver” barley.
Secale cereale L., “Open-pollinated rye’’.
Zea Mays L. (var. indentata).

Poa annua L., “Winter grass’.

Poa pratensis L., “Kentucky Blue grass’.

Bromus wunioloides H.B.K., “Prairie
grass”.

Lolium multiflorum Lam., “Italian rye
grass”.

Hordeum bulbosum lL.

Agrostis alba L.

Digitaria adscendens (H.B.K.) Henrard.

Dianthus spp., “Carnation”’’.

Malva_ parviflora L., “Small-flowered
Mallow’”’.

Sonchus oleraceus L., “Sow Thistle’.

Geranium sp., “Geranium”.

Geranium dissectum L.

Hrodium cygnorum Nees, “Blue-flowered
Crowsfoot”’.

Hrodium moschatum (l.) L’Hér., “Crows-
foot”.

Oxalis corniculata L.

Stellaria media (L.) Vill.

No infection was obtained in any of the above species, while the H. peplus

controls gave lesions and pycnidia.

The plants were kept under observation for

weeks after inoculation, in case development of the fungus was slower than in “Petty

Spurge’”’.

No Septoria disease was detected in the field on plants of H. helioscopia at Hobart,
Castle Hill and Hurstville, or on HE. Drummondii at Allawah and Sydney University,
or on plants of the latter species sent from Toowoomba, Queensland. At Hurstville,
several plants of H. helioscopia, growing in a patch of severely diseased H. peplus,

showed complete immunity.

Judged by the species tested, it appears as if the causal organism has a limited
host range. Specificity by Septoria species for one or a few hosts has been noted by
other workers, e.g., Beach (1919) working with 15 species, Weber (1922a, 1922b, 1923)
with eight species, Cochrane (1932) with two species, Thompson (1941) with two
species, and Sprague (1944) working with many species on grasses.

22 A SEPTORIA DISEASE OF KUPHORBIA PEPLUS L.,

NAME OF THE CAUSAL ORGANISM.

The history of the genus Septoria has been reviewed by Wakefield (1940) and
Sprague (1944). The former’s recommendation that Septoria Sace. (1884) described
from type species S. Cystisi Dem. be conserved against Septoria Fr. (1828), which
was based on non-pycnidial species, was upheld by Sprague. Phleospora Wallr. (1833),
based on non-pyenidial species, is an exact synonym of Septoria Fr. (1828). Grove
(1935), Clements and Shear (1944) and Ainsworth and Bisby (1945) followed
Saccardo’s use, employing Phleospora for the forms with incomplete pycnidia. The
writer, on the above authorities, retains the genus as described by Sacecardo (1884)
on page 474. 5

From examination of fresh diseased material and microtome sections, it is
evident that the causal organism of the disease of “Petty Spurge” conforms to
Septoria Sace. The filiform nature of the spores, their length in relation to their
width (ratio approximately 15-20:1), the yeasty, later carbonaceous, scanty mycelium
and distinct black pycnidia (to the naked eye), distinguish it from Stagnospora,
whose spores are typically cylindrical (although grading into more filiform types),
with a length: width ratio of less than 10:1, with a cottony appearance on P.D.A., and
with pale brown, less prominently distinguished pycnidia to the naked eye (Sprague,
1944; Clements and Shear, 1941). The pycnidia are too well-formed for Phleospora, _
and lesions occur too often on the leaves to consider Rhabdospora.

The only species of Septoria recorded on Huphorbia spp. are as follows:

1. S. bractearum Mont., 1849. The spore length is given as 50u. It was
described on H. serrata. (Saccardo, 1884.)

2. S. Kalchbrenneri Sacc. The type of this species is Rabenhorst Fungi
europaei 1854, issued as S. euphorbiae Kalchbrenner in Hedwigia, 1865,
p. 158, nec Guep. No spore measurements are given. It was described on
E. silvatica, E. palustris and E. aspera. (Saccardo, 1884.)

3. S. Huphorbiae Guep., 1879. The spore measurements are given as “40—45u x
2-23u, with 3-4 indistinct septa. It was described on H. EHsula and E£.
angulata. (Saceardo, 1884.)

The rest of the very brief descriptions of the above three species is more fe eR
the same.

Oudemans (1902), after examining the Exsiccati of Desmazieres, recommended
that S. bractearum Mont. should become S. Huphorbiae Desm., and that S. Huphorbiae
Guep. should yield place to S. Guepini Oud. S. Kalechbrenneri Sace. remained
unchanged.

4. §. media Sace. et Brun. A fairly full description is given. The spots are
described as having a dark reddish margin, with spores 50-55u x In. It
was described on EH. palustris. (Sacecardo, 1892.)

5. S. euphorbicola Hollos, 1910. The description of this species is fuller, and
the spots are given as 1 mm. in diameter, pycnidia 140-160u in diameter,
and spores 16-20u x 2-2-5u4. It was recorded on E. procera. (Saccardo,
1913.)

6. S. Hariotiana Sacc., 1906. A full description is given: the spots 1 mm.
in diameter, with a dark purple margin, pycnidia 120-1254 in diameter,
spores 3-4 septa, 30-324 x 34. It was recorded on EH. palustris. (Saccardo,
1913.)

Because of the inadequacy of the description of S. Kalchbrenneri Sace., a request
was made to the Commonwealth Mycological Institute for further information, if it
was available, and Dr. G. R. Bisby kindly supplied the following notes: “The Kew
specimen of this No. 584 consists of three leaves bearing spots 1-3 mm. in diameter,
roundish, visible on both sides of the leaf, at first brownish, then ashen, particularly
on the upper surface of the leaf, at all times surrounded by a slightly raised, distinct,
reddish brown margin; pyenidia circa 100—-200u wide, brown, with an ostiole which
becomes 35u or more wide; spores (25) 30-35u x 2—2-5u, somewhat elevate and tapering

BY DOROTHY E. SHAW. 23

to 1-1-5u at one end, hyaline, 0-3 septate, straight or somewhat curved. Most of the
pycnidia appear to open on the upper surface of the leaf.”

The only reference in the literature to a Septoria disease of H. peplus is in
Jaczewski’s “Key to Fungi’, Vol. 2, p. 107, 1917: “S. euphorbiae Guep. on HL. amygda-
loides, EH. peplus. Round, olive-coloured spots. Stylospores 40 to 454 by 2 to 2-5u
with 3 to 4 indistinct septa.” (Jaczewski was apparently unaware of the entry in
Revue Mycologique, 1902, whereby S. euphorbiae Guep. became S. Guepini Oud.). It
is not known whether Jaczewski was referring to H. amygdaloides Lam. or E. amygda-
loides L. Hooker and Jackson (1895) listed H. amygdaloides L. as being synonymous
with #. sylvatica L., on which a Septoria disease had already been described, namely,
S. Kalchbrenneri Sacc. Hooker and Jackson also listed H. amygdaloides Lam. as a
synonym of H. nicaeensis All., and H. nicaeensis (St. Am. Fl. Agen., 192) as a synonym
of H. Esula L. It was on H. Hsula L. that S. euphorbiae Guep. (now S. Guepini Oud.)
was originally described. It is not known whether the Russian organism was identified
on the grounds of morphological similarity. The spore sizes given in the Russian
text are the same as those given by Saccardo (1884).

’ The fungus under study, with spores 35-84 + 5-9u long by 1-9 + 0-17u wide and
pycnidia 110-2 + 19-0u and with unrestricted spots, differs in spore size from
S. Huphorbiae Desm. (once S. bractearum Mont.) (spores 50”), and from S. media
‘Sace. et Brun (spores 50-554 by lu); and in pycnidia and spore size from S. ewphor-
bicola Hollos (spores 16-204, pycnidia 140-1604). S. Huphorbiae Desm. was also
recorded by Oudemans (1921) on H. exigua, and no infection was obtained on this
plant with the organism from EH. peplus.

The fungus more closely resembles S. Hariotiana Sacc., S. Guepini Oud. and |
S. Kalchbrenneri Sace. The spores of S. Hariotiana, however, are given as 3u wide, and
the length of the spores of S. Guepini as 40—45u, and these are respectively wider and
longer than the spores under study. The pycnidia of S. Kalchbrenneri (100-2004) are
larger than those on Petty Spurge. S. Guwepini was recorded on H. Hsula, but no
infection was obtained on this plant with the organism under study. The appearance
of the lesions caused by S. Hariotiana and S. Kalchbrenneri on their hosts, differs
markedly from those caused by the Septoria on EH. peplus.

It is realized that the same organism can sometimes produce quite dissimilar
lesions on even closely related hosts, and that the difference in spore sizes of some of
the above species could, perhaps, be accounted for by natural variation or environment.
However, it is considered that the fungus is sufficiently different morphologically (as
far as can be determined from the brief description of some of the other species), and
with regard to host range, to be described as a new species. In correspondence with
the C.M.I., Dr. Bisby was of the opinion that this would be the best course to follow.
It is therefore proposed to name the fungus Septoria pepli, n. sp.

= SEPTORIA PEPLI, Nn. Sp.

Pycnidiis amphigenis, sed vulgo hypogenis, in orbicularibus non limitatis maculis;
sparsis aut raro aggregatis, atris, globosis, innatis vel dein erumptibus, ostiolatis, 85-135
(65-165); sporulis hyalinis, rectus vel leniter curvatis, sursum attenuatis, guttulatis,
3-septatis, saepe 2-, raro 0- vel 4-septatis, 25-444 (17—-51lu) x 1-5-2-0u (1-3-2-5y).

Hab. in foliis et in cauli H. peplus L. in N.S.W., A.C.T. et Tasmania.

Pycnidia on both sides of the leaves, but mostly on the undersurface, in circular,
unrestricted spots, singly or rarely aggregated in twos or threes, visible to the naked
eye, black, but reddish-brown by transmitted light, globose or slightly elongate, immersed
but later,erumpent; ostiole about one-quarter the diameter of the pycnidium; pycnidial
wall smooth, composed of 2-3 layers of pseudoparenchymatous cells; 85-1354 (65-165,) ;
spores hyaline, straight or slightly curved, tapering at one end, with a varying number
of guttulae, 3 septate, often 2-, rarely 0- or 4-septate;, 25-44u (17-5luw) x 1-5-2-0u
(1-3-2-5y).

Hab. on leaves and stems of E. peplus L., around Sydney, and at Wagga, N.S.W.,
-in the A.C.T. and Tasmania.

Type specimen collected at Pennant Hills, December, 1949.

24 A SEPTORIA DISEASE OF EUPHORBIA PEPLUS L.,

Acknowledgements.

I gratefully acknowledge the help of the following: Dr. G. R. Bisby of the C.M.I.
for assistance as mentioned specifically in the text; Dr. D. B. Duncan for advice on the
statistical analysis; Mr. 8. Fish, for two verbatim extracts from Revue Mycologique
which is housed at Melbourne; Mr. J. Humpoletz and Mr. C. Warner for assistance in
translating the extracts; Mr. J. Strang for the diseased material from Penshurst; the
Sydney Botanic Gardens, Adelaide University, University of Tasmania and the Plant
Introduction Office, C.S.I.R.O., Canberra, for cuttings and seed of the species of
Euphorbia tested; Professor W. L. Waterhouse and the Department of Medical Illus-
tration, Sydney University, for help with the photography. I am particularly grateful
for the help given at all times by Professor Waterhouse during the course of the study.

References.

AINSWORTH, G. C., and G. R. Bissy, 1945.—A Dictionary of the Fungi. Kew.

BeacH; W. S., 1919.—Biologiec Specialization in the Genus Septoria. Amer. Jour. Bot., 6: 1-33.

BissBy, G. R., 1945.—An Introduction to the Taxonomy and Nomenclature of Fungi. Kew,
Surrey.

Buack, J. M., 1922.—Flora of South Australia. Part 2. Adelaide.

BrIEN, R. M., 1939.—A List of Plant Diseases Recorded in New Zealand. Dept. S. and I.R. Bull.
No. 67.

Bropvift, H. J.. and C. C. NEUFELD, 1942.—The Development and Structure of the Conidia of
Erysiphe Polygoni D.C. and their Germination at Low Humidity. Can. Jour. Res., C. 20:
41-61.

CLEMENTS, F. E., and C. L. SHEAR, 1941.—The Genera of Fungi. New York.

COCHRANE, L. C., 1932.—A Study of Two Septoria Leaf Spots of Celery. Phytopath., 22: 791-812.

CooKE, M. C., 1892.—Handbook of Australian Fungi. London.

C.S.1.R., 1942.—Standardised Plant Names. Bull. No. 156. Melbourne.

DoipGE, E. M., and A. M. BoTTOMLEY, 1931.—A Revised List of Plant Diseases Occurring in
South Africa. Botanical Survey of South Africa, Mem. 11, Pretoria.

GROVE, W. B., 1935.—British Stem- and Leaf-fungi. Vol. 1. Cambridge.

Hooker, J. D., and B. D. JACKSON, 1895.—Index Kewensis Plantarum Phanerogamarum.
Tomus 1.

HuGuHeEs, 8S. J., 1949.—Studies on Some Diseases of Sainfoin (Onobrychis sativa), ii. The
Life History of Ramularia Onobrychidis Allescher. Trans. Br. Mycol. Soc., 32: 34-59.

Hurst., E., 1942.—The Poison Plants of N.S.W. Sydney.

JOHNSTON, T., 1947.—A Form of Leptosphaeria avenaria on Wheat in Canada. Can. Jour.
IMG (Ca 268 2470)

IKLEBAHN, H., 1934.—EHine Blattkrankheit und einige sie begleitende Pilze. JZeitschr. fur
Pflanzenkrankh. wu. Pflanzenschutz, 33, i, 1-23. Abstr. R.A.M., 13:407, 1934.

MACMILLAN, H. G., and O. A. PLUNKETT, 1942.—Structure and Germination of Septoria Spores.
Jour. Agr. Res., 64: 547-559.

MCALPINE, D., 1895.—Systematic Arrangement of Australian Fungi. Melbourne.

NAKATO NAITO, 1940.—Studies on Septorioses in Plants. New or Noteworthy Species of
Septoria Found in Japan. Mem. Coll. Agric. Kyoto, 47: 31-43.

NOBLE, R. J.. H. J. Hynes, F. C. McCurrery, and W. A. BIRMINGHAM, 1934.—Plant Diseases
recorded in N.S.W. Sci. Buwill., 46, Sydney.

OUDEMANS, C. A. J. A., 1921.—Enumeratio systematica fungorum. Vol. 3, p. 1071 et seq.

1902. Rectifications Systematiques, rédigées en ordre Alphabétique. Rev. Mycol.,
p: LZ: :

RAMsBOTTOM, J., 1948.—Presidential Address. Trans. Brit. Mycol. Soc., 30: 22-39 (pp. 33-34).

ROARK, E. W., 1921.—The Septoria Leaf Spot of Rubus. Phytopath., 11: 328-333.

RUGGIERI, G., 1933.—Lesions on Citrus sinensis Obeck caused by Mycosphaerella aurantiorum.
Boll. Staz. Pat. veg. Roma, N.S., 15, 2, 338-346, 1935. Abstr. R.A.M., 15:85, 1936.

SACCARDO, P. A., 1884.—Sylloge fungorum, 3: 474 and, 515.

—-— ——, 1892.—Sylloge fungorum, 10: 379.

, 1913.—Sylloge fungorum, 222: 1092.

SHIGEKATSU HIRAYAMA, 1931.—Studies of Septorioses of Plants. IV. New or Noteworthy
Species of Septoria Found in Japan.- Mem. Coll. Agric. Kyoto, 13: 33-39.

SPRAGUE, R., 1944.—Septoria Disease of Gramineae in Western United States. Oregon State
College, Oregon.

STEVENS, F. L., 1925.—The Fungi which Cause Plant Disease. New York.

STONE, R. E., 1916.—Studies in the Life Histories of Some Species of Septoria occurring on
Ribes. Phytopath., 6: 419-427.

THOMPSON, G. E., 1941.—Leaf Spot Diseases of Poplars caused by Septoria musiva and
S. populicola. Phytopath., 31: 241-254.

WAKEFIELD, E. M., 1940.—Nomina generica conservanda. Contributions from the Nomenclature
Committee of the British Mycological Society. 3. Trans. Brit. Mycol. Soc., 24: 245-293.

BY DOROTHY E. SHAW. 25

WEBER, G. F., 1922a@.—Septoria Diseases of Cereals. Phytopath., 12: 449-470.
, 19226.—Septoria Diseases of Cereals. Phytopath., 12: 5387-585.
, 1923.—Septoria Diseases of Cereals. 3. Phytopath., 13: 1-23.
WOLLENWEBER, H. W., 1938.—Sphaerella linicola n. sp., the Agent of the American Flax
Epidemic (‘“‘Spasm” or Septoria Disease). Rev. Bot. Inst. ‘Miguel Tillo’, ii, 2a, pp. 483-492,
1938. Abstr. R.A.M., 18: 111-112, 1939.

DESCRIPTION OF PLATE I.

Fig. 1.—Leaves of Huphorbia peplus L. showing lesions caused by Septoria pepli n. sp. x 2.

Fig. 2.—Six months old culture of the fungus on P.D.A., showing ‘‘staling’’. Note the
convoluted centre and the sub-surface hyphae at the edge. x 2.

Fig. 3.—Pycnidium produced in culture, on rye and peanut husks, with spores exuded in a
cirrus. Mounted in cotton-blue lactophenol. x 100.

Fig. 4.—Section through pycnidium showing filiform spores. Cut at 84% and stained with
gentian violet-orange G. x 400. :

26

PRESERVATION TECHNIQUES FOR SCARABAEID AND OTHER INSECT LARVAE.
By P. B. Carne, B.Agr.Se., Division of Entomology, C.S.I.R.0.
[Read 28th March, 1951.]

Synopsis. 4

Methods commonly employed in preserving insect larvae are compared and the use,
especially for scarabaeid larvae, of Peterson’s “KAAD” fixative is recommended. The various
fixatives in use are compared in regard to price; a modified cheaper form of Peterson’s
fixative, giving equally good results, is described.

The probable function of the fixative components is discussed and evidence recorded which
suggests that the discoloration of preserved larvae is due to tyrosinase activity; in vivo, the
tyrosinase is inhibited by a dehydrogenase system. The very rapid distension of larvae killed
in KAAD or its modification is discussed in the light of recent work on insect cuticle.

Detailed recommendations are given for the cleaning, fixing and storage of larval
Scarabaeidae.

INTRODUCTION.

The writer has experimented with a variety of methods of preserving insect larvae.
Much of his earlier collected material, and of that obtained from the-collections of
earlier workers, shows marked faults, resulting from the use of unsatisfactory methods
of preservation. While ethyl alcohol is used almost universally as a storage fluid,
previous fixation is essential with most species; larvae killed and stored directly in
alcohol become blackened and distorted and almost valueless for scientific study.

The purpose of this paper is to compare preservation techniques in common use, to
bring to notice a valuable new fixative devised by Alvah Peterson (Peterson, 1943, 1948),
and to discuss the functions of the constituents of the latter.

DEFINITION OF A SATISFACTORY FIXATIVE.

An ideal fixative would have the following characteristics:

(a) Larvae fixed in it should be distended and turgid, neither soft and flaccid, nor
hardened and shrivelled. (0b) The larvae so fixed should retain normal, or close to
normal, coloration (Lepidopterous larvae present much greater difficulties in this
regard than do Scarabaeid larvae), with no darkening of the softer parts, nor bleaching
of the head capsule. An increase in body opacity is desirable for morphological work
where setal patterns are to be studied. (c) The fixative fluid should be reasonably
inexpensive, as large quantities are used in the field collecting of larvae; and (d) the
fixative should be one which may be used cold, and in which the larvae may be held
for prolonged periods without deterioration. This is particularly desirable on collecting
expeditions, when regular transference of larvae from fixative to storage fluid is some-
times inconvenient. 2

PRESERVATION TECHNIQUES COMPARED.

The bulk of the larval collections seen by the writer have been treated by one or
other of the following methods. The following comments are based upon a critical
comparison of these methods, using series of larvae of Adoryphorus couloni Blackb.,
Semanopterus sp. and Sericesthis sp., and on the writer’s general observations on the
preservation of a wide range of species of soil-inhabiting larvae. The compositions of
the fixatives are given below.

a. By direct killing in a storage fluid of 70-95 per cent. ethyl alcohol or in methylated
spirits.

This technique, which appears to be frequently used, is most unsatisfactory for
scarab larvae. Darkening of the cuticle begins within 24 hours of killing, and older
specimens are completely blackened, and may become either very soft or hardened,
according to the strength of alcohol used.

BY P. B. CARNE. 27

b. By direct killing in a storage fluid of 4 per cent. formalin.

This technique is equally unsatisfactory, severe discoloration occurring in time.
Formaldehyde vapour from the specimens is a continual source of irritation to the eyes
during their examination.

c. By killing and fixing in boiling water, and subsequent storage in 70-95 per cent.
ethyl alcohol, or in 4 per cent. formalin.

Quite good results can be obtained by this method, and darkening on storage is
prevented to a great extent. Small larvae appear to respond better than large larvae,
which may fail to become, or to remain, turgid. Collapse is frequent if the larvae are
held in actively boiling water; much better results are obtained if the larvae are placed
in a beaker of boiling water, which is then allowed to cool. If the high temperature
is maintained, the fat lining the body wall is melted, and is deposited about the gut.
Any heat treatment is undesirable when larvae are to be dissected.

d. Killing and fixing in one of the acetic-formalin fixatives, e.g., Carls, Blés, or Bouin,
and transfer to 70 per cent. ethyl alcohol.

Carls’ or Blés’ fixatives can give good results in the laboratory where careful
attention can be given to time and temperature of fixation. Overfixation, resulting in
hardening and buckling of the body wall, is the common error. Such overfixation
makes examination of setal patterns extremely difficult, and special “renovation” tech-
niques may be necessary before determination of the larva is possible. These fixatives
can be used cold for longer periods, but larvae should be sorted into size groups,
because in the period required to ensure adequate fixation of large larvae, small larvae
in the same series becomes grossly overfixed. However, where collections are being
made at a number of localities, this introduces difficulties in that the number of
containers to be carried is multiplied.

Bouin easily results in overfixation, and imparts an undesirable picric staining to
the larvae, which is difficult to remove.

e. Killing and fixing in Carnoy’s fixative and storage in 90 per cent. ethyl alcohol.

Killing and fixing in cold Carnoy gives excellent results, although larvae become
somewhat soft and transparent if allowed to remain in the fixative for more than a few
days.

Larvae fixed by methods a—d invariably die with their mandibles closely opposed or
overlapping, and these cannot be moved apart, the articulations becoming rigid, so that
dissection of the mouthparts is difficult. On the other hand larvae fixed in Carnoy
die with the mandibles opposed, but the articulations remain quite flexible and are
easily dissected.

f. Killing and fixing in Peterson’s KAAD fixative, or derivatives thereof, and storing
in 95 per cent. ethyl alcohol.

The use of Peterson’s KAAD results in larvae in an ideal state of preservation.
The larvae are well distended, firm and completely free of any discoloration. There is
an increase in opacity of the body wall.

Although Peterson states that best results are obtained by fixing for periods not
longer than four hours, the writer has not observed any deterioration when larvae are
left in the fixative for periods of up to three weeks. Best results for scarab larvae are
given by fixation for not less than 2-3 hours.

As the writer’s use of KAAD began only two months prior to the drafting of this
paper, he has not seen larvae so preserved for periods longer than this. It is the
writer’s observation that, with all other preservation techniques, any tendency towards
deterioration becomes evident within a week of treatment. Peterson states that the
larvae retain their good condition for “a prolonged period”.

Larvae killed in KAAD almost invariably die with the mandibles apart, the
articulations remaining flexible. Important taxonomic characters occur on the
mandibles, and dissection may often be avoided when the mandibles remain apart.

The fixative is expensive, containing approximately 8 per cent. dioxane. For this
reason the writer has tried omitting this component, with results equal to those

28 PRESERVATION TECHNIQUES FOR SCARABAEID AND OTHER INSECT LARVAE,

obtained with the complete fixative. The acetic acid content may be reduced to 25 per
cent. of that recommended by Peterson, or may be omitted altogether if the larvae are
previously killed in hot water.

For purposes of gross dissection, larvae fixed in KAAD are considerably superior
to those prepared by any of the other methods described. Dr. M. F. Day, of the
Division of Entomology, C.S.I.R.O., has kindly examined mid-gut tissues of cetoniid
larvae so treated. He found that histologically the mid-gut was fairly well preserved,
considering the gross method of fixation employed, and that staining of the tissues was
satisfactory.

DISCUSSION.

The writer has rarely seen any discussion of the possible function of components
of fixative mixtures, and has attempted to gain some understanding of these functions.

Firstly, all fixatives known to the writer contain acetic acid and alcohol. It is
generally stated that acetic acid, together with the alcohol, is responsible for precipita-
tion (“fixation”) of the body protein, and that for most larvae its presence is necessary
to prevent subsequent blackening.

The fact that hot water treatment prevents discoloration suggests that the blacken-
ing is an enzymatic process. That the enzyme is probably tyrosinase is supported by
the following observations. Blackening is prevented by acids, suggesting that inactiva-
tion of the enzyme results from denaturation of the protein portion of the enzyme.
Tyrosinase is known to contain copper in its prosthetic group and to be inactivated by
cyanide. Larvae killed in cyanide do not blacken when stored in alcohol but will do
so if first placed in an alcoholic solution of cupric chloride. The period of immersion
in hot water necessary to destroy the enzyme varies from one minute at 100°C. to
30 minutes at 55°C. with Semanopterus sp.

The rapidity with which larvae blacken depends upon the treatment, which suggests
that there is some system present which prevents blackening. Dennell (1949) considers
that the tyrosinase in the larvae of Calliphora erythrocephala is prevented from acting
upon its substrate by the presence of a dehydrogenase system which maintains a low
redox potential in the insect tissues. Destruction of this system allows the redox
potential to rise and the tyrosinase to act upon its substrate. Chloroform inhibits
dehydrogenase systems and Adoryphorus and Sericesthis larvae placed in chloroform
vapour are completely blackened an hour after anesthesia. When killed in ethyl
acetate vapour these larvae show no trace of blackening in the same period of time,
which suggests the presence in these larvae of a tyrosinase-inhibiting dehydrogenase
system.

Hurst (1940) has observed that a polar substance of low dielectric constant such
as alcohol, is greatly assisted in its passage through the lipoid layer of insect cuticle
by the presence of a non-polar compound of high dielectric constant, such as kerosene.
The very rapid distension of larvae killed in Peterson’s “KAAD” appears to be due to
this phenomenon. Scarab larvae placed in either ethyl alcohol or kerosene die very
slowly, and distension takes place very slowly. Death occurs very rapidly in a
mixture of these two substances, and larvae are fully distended in less than an hour.
The rate of entry of alcohol may be reduced by lowering the proportion of kerosene in
the mixture, and Peterson found this necessary with some soft-bodied larvae, which
otherwise burst before equilibrium was established.

Some kerosenes are not completely miscible with alcohol-acetic acid mixtures, and
Peterson finds that adding dioxane results in complete miscibility. While Peterson
considers that the dioxane itself may improve the quality of some larvae, the writer
has found larvae preserved in KAA equally good. The KAA becomes cloudy during
fixation to a much greater extent than does KAAD, although the cloudiness may be
greatly reduced by the use of absolute rather than 95 per cent. alcohol in preparing
the fixative. The fixative may be filtered and used again a number of times,

BY P. B. CARNE. 29

COMPOSITION AND ESTIMATED COSTS OF FIXATIVES.*

Bouin.—Picrie sat. aqueous soln. 71 per cent., formalin 24 per cent., glacial acetic
3 per cent., approx. 7s. per gallon.

Carls.—Water 57 per cent., absolute alcohol 28 per cent., formalin 11 per cent.,
glacial acetic 4 per cent., approx. 4s. per gallon.

Blés.—70 per cent. alcohol 90 per cent., formalin 7 per cent., glacial acetic 3 per
cent., approx. 3s. 6d. per gallon.

Carnoy.—Absolute alcohol 60 per cent.,’ chloroform 30 per cent., glacial acetic 10
per cent., approx. £1 1s. per gallon.

‘KAAD.—Kerosene 8 per cent., 95 per cent. alcohol 70 per cent., glacial acetic 14 per
cent., dioxane 8 per cent., approx. £1 per gallon.

KAA (1).—Kerosene 8 per cent., 95 per cent. alcohol 77 per cent., glacial acetic acid
15 per cent., approx. 9s. per gallon.

KAA(2) (with acetic acid content reduced).—Kerosene 8 per cent., 95 per cent.
alcohol 87 per cent., glacial acetic 5 per cent., approx. 5s. per gallon.

While there is probably little to choose between Carnoy and KAA(2), it will be
seen that the price of the former is approximately four times that of the latter.

OTHER F‘AULTS IN PRESERVED LARVAE.

When larvae are immersed in any fluid other than actively boiling water, regurgita-
tion of part of the gut contents occurs. The black fluid coagulates on the mouthparts.
The latter possess taxonomically important structures, which must be examined in
detail for specific determination; the coagulated material must therefore first be
removed. This is a tedious operation, and one likely to damage delicate structures:
it may be made unnecessary either by starving the larvae for several days before
killing, so that the regurgitated fluid is colourless and leaves no deposit, or the larvae
may be anesthetized before placing in the fixative, when no regurgitation occurs.
Many scarab larvae are remarkably slow to succumb to cyanide, and the most rapid
anesthesia is brought about by carbon dioxide or chloroform. Deep anesthesia is
necessary, or the larvae recover sufficiently in cold fixative to regurgitate.

Tubes containing preserved larvae must be sealed to prevent evaporation, other-
wise the percentage alcohol content of the fluid decreases rapidly, due to the higher
volatility of aleohol over water in the mixture. Browning and changes of texture of
the larvae.then occur which are highly undesirable. If some evaporation does occur,
the tube should be topped up with an alcohol of higher strength than that originally
used. a :

RECOMMENDED PROCEDURE FOR PRESERVATION OF SCARABAEID LARVAE.f

(a) Handle larvae with blunt forceps, and remove superficial dirt by blowing, or
by gentle washing in cold water. With some cetoniid larvae, a soft brush may be
necessary to dislodge adhering soil. Such cleaning movements should be made in a
caudal direction to avoid damage to the setae clothing the body.

(bo) Anaesthetize the larvae deeply in carbon dioxide, or chloroform or ethyl
acetate vapours.

(c) Kill and fix in Peterson’s KAAD or KAA for not less than two hours.

(d@) Wash larvae in alcohol briefly and store in 95 per cent. alcohol. Not more
than two-thirds of the effective length of the tube should contain larvae, which should
always be well covered by alcohol.

LARVAE OTHER THAN SCARABABIDS.

Peterson’s KAAD or KAA has been tried with a number of larval types. Excellent
fixation was given with larvae of Carabidae, Elateridae, Cerambycidae, Chrysomelidae,
Asilidae, and Lepidoptera. The fixative appears to be unsatisfactory for some Calli-
phorid larvae.

* Based. on Australian prices as at July, 1950.

+ This refers to larvae killed in the laboratory. In the field, anesthesia may have to be
omitted, and cleaning postponed until transfer to storage fluid in the laboratory.

30 PRESERVATION TECHNIQUES FOR SCARABAEID AND OTHER INSECT LARVAE.
Acknowledgements. i
The writer is indebted to Dr. R. H. Hackman, Division of Industrial Chemistry, and
Drs. M. F. Day and K. H. L. Key of the Division of Entomology for advice and comment.

References.

DENNELL, R., 1949.—Weismann’s ring and the control of tyrosinase activity in the larva of
Calliphora erythrocephala. Proc. Roy. Soc. L., (B), 136: 94.

Hurst, H., 1940.—Permeability of insect cuticle. Nature, 145: 462.
PETERSON, ALVAH, 1943.—Some new killing fluids for larvae of insects. J. Hcon. Ent., 36: 115.
, 1948.—Larvae of insects, Lepidoptera and Hymenoptera. Pt. 1. Columbus, Ohio.

31

THE ANATOMY AND MORPHOLOGY OF THE OPERCULUM IN THE
GENUS HUCALYPTUS.

PART I. THE OCCURRENCE OF PETALS IN EUCALYPTUS GUMMIFERA (GAERTN.) HOCHR.

By J. L. WILLIS,
Museum of Applied Arts and Sciences, Sydney.

[Read 26th April, 1951.]

Synopsis.
The nature of the operculum present in the genus Hucalyptus has been for many years
the subject of a considerable amount of conjecture, and a number of conflicting interpretations
as to its morphological nature have been proposed.

In the young buds of H. gummifera, four minute imbricate petals have been found, which
gradually fuse together to form an inner corolline operculum. This inner operculum remains
quite distinct from the outer operculum which is probably calycine in origin.

There is, therefore, a close morphological relationship between the flowers of this species
and the two New Caledonian genera Piliocalyx and Acicalyptus, which may also indicate a close
phyletie relationship.

INTRODUCTION.
Although a considerable volume of literature dealing with the anatomy of the
genus Hucalyptus has accumulated, no investigations have yet been carried out on that
unique organ, the operculum, most attention having been focussed upon the wood
anatomy, leaf structure, etc. This is somewhat surprising, as the exact nature of the
operculum has been the subject of a considerable amount of conjecture since 1788 when
the genus was first described by L’Héritier.

HISTORICAL.

In his original description, and after consideration of the one species only
(#. obliqua), L’Héritier held the operculum to be corolline in nature. On the other
hand, Jussieu (1812) considered that it was formed by the fusion of two bracts. Robert
Brown (1814), however, came to the conclusion that the operculum had different origins
in different groups of species. He thought that in most species it represented a fusion
of the calyx and corolla; in those species with double opercula, the inner structure
represented the corolla, and the outer one the calyx; and in the genus Hudesmia R.Br.,
now Eucalyptus L’Hér., Series Hudesmiae (Blakely, 1934), the operculum was formed
from the corolla alone. Hooker (1860) concluded that the operculum was a combined
calyx and corolla, but Bentham (1866) was uncertain as to the correct interpretation
of the organ, although he considered it to be most likely corolline in nature. He noted
the presence of an additional outer operculum in some species, but regarded the nature
of this outer organ as doubtful. Bentham thought that this outer operculum would
eventually be found to be present in nearly all species but that it was deciduous so early
that it was not noticeable in most buds. Maiden (1923) regarded the operculum as
corolline in origin except in those species with double opercula where he considered
the inner one to be corolline and the outer one calycine. He predicted that eventually
all species would be found to have double opercula, the outer calycine one being
deciduous very early in most species. Naudin (1883), Deane (1900), Andrews (19138),
Hutchinson (1926), Blakely (1934), Rendle (1938), and Osborne (1947) all considered
the operculum to be formed from the fused petals, whereas von Mueller (1879-84)
concluded that the organ was nearly always calycine in nature. Hardy (1935, 1939),
however, thought the operculum was either a modified corolla or a fusion of both calyx
and corolla, but he considered the evidence insufficient to determine definitely which
interpretation was the correct one.

32 ANATOMY AND MORPHOLOGY OF THE OPERCULUM IN EUCALYPTUS. I,

In view of these conflicting opinions, it was thought that a study of the organogeny
of the operculum would definitely decide which of these interpretations should be |
adopted.

THE PRESENT STATUS OF THE PROBLEM.

It is apparent from the descriptions of the various species that there can be no
general interpretation of the morphological nature of the operculum, and that the
species fall roughly into three groups: (1) the Hudesmiae with four minute calyx teeth
surmounted by a single operculum; (2) those species with distinct double opercula
throughout most or all of the stages of bud development, notably the Corymbosae-
peltatae; and (3) those species with a single operculum only throughout most or all of
the bud’s development. This last group includes the majority of Eucalypt species.
These three groups coincide with those of Robert Brown (1814).

There are still good grounds for considering group (1) (Eudesmiae) to be a
separate genus closely related to Hucalyptus. Also, group (3) is probably derived from
group (2) by suppression of one of the opercula or by the fusion of both structures.
Consequently, an examination of the opercula in group (2) is the most likely to provide
information regarding the fundamental nature of the organ. It may also assist -in
interpreting the morphological nature of the single operculum in group (3).

MATERIALS AND METHODS.

The Bloodwood EL. gummifera was the first species to be examined as it is a common
tree on the Hawkesbury sandstone in the vicinity of Sydney, and so little difficulty was
experienced in collecting adequate material. The buds used in this study were collected
from three different localities—Avalon, Roseville, and National Park.

The buds at different stages were fixed under reduced pressure in F.A.A. or F.P.A.
and embedded in paraffin in the usual way. Staining was carried out with Safranin and
Delafield’s Haematoxylin using the method of Boke (1939).

The photographs were taken on Kodak Process Pan plates at a magnification of x 30,
and have been reproduced at the same magnification.

THE ANATOMY OF THE OPERCULUM.

An examination of the very young buds of H. gummifera revealed the presence of
four minute, imbricate petals, inserted between the staminal ring and outer operculum
(Plate ii, figs. 1 and 2). The petals are simple in construction with a uniform epidermis
and no cuticle, whilst a constant feature is the presence of many large, well-defined oil
glands (Fig. 2). The petals are attached by a broad base, and have a peculiar arrange-

|
Text-figures 1 and 2.

1. The petal arrangement in Hucalyptus gummifera.
2. The expected petal arrangement when the phyllotaxis is opposite and decussate.

ment in that they are not symmetrically imbricate. For example, in the case of three
of the petals, the right (or left) edge overlaps the left (or right) edge of the petal next
to it, whilst the fourth petal has both right and left edges enclosed by the petals on
either side (Text-fig. 1 and Plate iii, fig. 8). As Huealypts have basically a phyllotaxis
of one-half (Jacobs, 1986), one would expect a simple dimerous arrangement as shown
in Text-figure 2.

BY J. L. WILLIS. 33

This arrangement described above leads to one petal being folded and enclosed by
the other three, so that when the bud is cut transversely, the upper part of the folded
petal is sectioned somewhat longitudinally (Plate iii, fig. 7).

As the bud develops, the petals gradually fuse together to form a solid inner
operculum, which remains quite separate from the outer conspicuous operculum
(Plate iii, figs. 10 and 11). The calyx tube grows faster than both the operculum and
the fused petals, and as the proportional volume occupied by the fused petals becomes
less and less, they spread out to form an almost flat cap just under the operculum proper
(Plate iii, fig. 11). There is always a small space between the two structures and they
fall together when the stamens unfold. A comparison of the ratio opercular volume to
bud volume in Plate ii, fig. 1 and Plate iii, fig. 11, shows the much more rapid growth
of the calyx tube.

Hardy (1935, 1939) has pointed out that the operculum is not a solid mass of tissue
as most authors have assumed. Instead, a distinct lobing at or near the apex of the
operculum marks the:presence of a minute pore, which may either run right through
the organ, or only part of the way, according to the species under consideration.
Hardy found this pore or traces of it in 55 species, and there seems little doubt that it
will be found to be of general occurrence in the genus.

In H. gummifera this channel is lined with cuticularized elongated cells, and runs
completely through the organ (Plate iii, fig. 9). It nearly always emerges at one side
of the operculum near the apex, and rarely through the apex itself (Plate ii, fig. 3;
Plate iii, fig. 9). At its upper end the channel is usually three lobed (Plate ii, fig. 4)
but lower down it becomes a single elongated slit (Plate ii, fig. 5). It is eventually
obliterated as the bud matures.

DISCUSSION.

Angophora is generally considered to be the genus with the closest affinity to
Eucalyptus, the two genera being the only members of the Subtribe Eucalypteae
(Bentham and Hooker, 1862-67). However, in view of the occurrenee of four minute
petals in H. gummifera as described above, it now seems more likely that the genera
most closely related to Hucalyptus will be found to be the two genera Piliocalyx Brong.
and Gris., and Acicalyptus A. Gray. (Bentham and Hooker (1862-67) placed Piliocalyx
in the genera anomala, and Acicalyptus in the Subtribe Metrosidereae. )

Acicalyptus is confined to the Fiji Islands and New Caledonia. Its buds are
Eucalypt-like and four-angled, with a beaked operculum which falls off, revealing four
minute imbricate petals inserted on the margin of the calyx tube by broad bases. These
petals lightly cohere, thus forming a lid which falls in one piece when the operculum is
detached. There are numerous inflexed stamens but the ovary has only two loculi. The
fruit is unknown (Gray, 1854).

Piliocalyx also possesses an operculum covering four small, unequal imbricate
petals which cohere to form an inner operculum. The stamens are indefinite and the
fruit is unknown (Brongniart and Gris, 1865). One species is found on Lord Howe
Island (von Mueller, 1873), but otherwise the genus is known only from New Caledonia.
Both these genera, therefore, bear flowers having a very close morphological relation-
ship to those of H. gummifera, and a closer examination of them should prove of great
interest, as this similarity in floral morphology may well denote a close phyletic kinship.

It seems likely also that on investigation a situation similar to that described for
E. gummifera will be found to hold for all other species of Eucalyptus with double
opercula (mainly the Corymbosae-peltatae), and that by fusion or loss, the single
operculum of the majority of Eucalypts has been formed. However, the position of
-such species as H. camaldulensis Dehnh. (Series Exsertae) and #. microtheca F. Muell.
(Series Buxeales) described by Maiden (1923) as having double opercula, is still
obscure.

The presence of the opercular channel described above, may well indicate that the
operculum is calycine in nature, especially in view of the lobed nature of the channel
near the apex. In this connection it is significant that Hardy (1935, 1939) recorded
this channel in two other members of the Corymbosae-peltatae, H. calophylla R.Br. and

Fr

34 ANATOMY AND MORPHOLOGY OF THE OPERCULUM IN EUCALYPTUS. I,

E. ficifolia F. Muell., the former having four symmetrical lobes at the apex of the
operculum, and the latter either four, three or two lobes irregularly arranged, ‘but
approximating to a symmetrical arrangement of four about the axial point”. Although
Hardy considered the operculum, in the latter species at least, to be a combined calyx
and corolla, for simplicity he termed the lobes ‘“‘petaline vestiges’’.

However, it has been found (Willis, unpub.) that in #. ficifolia (and almost
certainly in #. calophylla) there are four imbricate petals similar to those reported
above for H#. gummifera, but larger and not completely concrescent. They are pressed
closely to the inside surface of the operculum but not fused to it, indicating that the
operculum is most likely calycine in nature and is not a composite structure.

The indistinct lobing or lack of lobing, and the incomplete nature of the opercular
channel found in such series as the Pachyphloiae and the Piperitales indicates that
fusion takes place earlier and earlier in the ontogeny as the species becomes more
advanced.

Jacobs (1936) has shown that the phyllotaxis of Hucalyptus is basically opposite
and decussate. The alignment of juvenile leaves in two rows and the varying arrange-
ments of mature leaves are caused by the growth in length of the stem between each
pair of leaves, the twisting of the petioles, and the twisting of the internodes between
each leaf pair. Consequently, the petals of a Hucalyptus flower would be expected to
show two dimerous whorls instead of the arrangement described. The arrangement of
the petals of Piliocalyx and Acicalyptus is at present unknown, but if they show an
arrangement similar to that shown by LH. gummifera, it will be additional confirmation
for the close morphological relationship to Hucalyptus postulated above for these two
genera.

SUMMARY.

1. The presence of four small imbricate petals in H. gummifera is demonstrated.

2. The petals have an unusual unsymmetrical arrangement instead of the expected
two dimerous whorls.

3. These petals fuse during the development of the bud forming an inner operculum.

4. The outer operculum has a distinct channel or slit running through it which is
obliterated before the bud reaches maturity.

5. The relationship between H. gummifera and the two genera Piliocalyx and
Acicalyptus is discussed.

6. There is some evidence indicating that the outer operculum represents a fusion
of the sepals.

ACKNOWLEDGEMENTS.
Thanks are due to the Trustees and the Director of the Museum of Applied Arts
and Sciences, Sydney, for permission to publish this work, and to Dr. P. Brough for
much encouragement and helpful criticism.

References. ,

ANDREWS, E. C., 1913.—The Development of the Natural Order Myrtaceae. Proc. LINN. Soc.
N.S.W., 38: 558.

BENTHAM, G., 1866.—Flora Australiensis, Vol. 3. Lovell Reeve and Co., London.

, and Hooxrmr, J. D., 1862-67.—Genera Plantarum, Vol. I. Reeve and Co., London.

BLAKELY, W. F., 1934.—A Key to the Eucalypts. The Worker Trustees, Sydney.

Boker, N. H., 1939.—Delafield’s Haematoxylin and Safranin for Staining Meristematic Tissues.
Stain Techn., 14: 129-131. ‘

BRONGNIART, AD., and Gris, A., 1865.—Observations sur Diverses Plantes Nouvelles ou peu
connues de la Nouvelle-Calédonie. Ann. Sci. Nat., Sér. V, 3: 225-227.

Brown, Ropert, 1814.—Appendix to Matthew Flinders’ Voyage to Terra Australia, Vol. 2.
Lovell Reeve and Co., London.

DBANB, H., 1900.—Observations on the Tertiary Flora of Australia, etc. Proc. LINN. Soc.
N.S.W., 25:471. Incorrectly cited by Maiden (1923) as 22:471 (1897).

Gray, ASA, 1854.—Wilkes’ U.S. Expl. Exped. Botany Phanerogamia. Vol. I and Atlas.
George Putnam and Co., New York.

Harpy, A. D., 1935.—Petaline Vestiges in Eucalyptus. Rept. Aust. N.Z. Assoc. Adv. Sci.,
22: 372-4.

, 1939.—Additional Notes on Petaline Vestiges in Eucalyptus. Proc. Roy. Soc. Vic., 51

(N.S.) : 215-18.

BY J. L. WILLIS. 35

Hooker, J. D., 1860.—Flora Tasmaniae. Vol. I. Dicotyledons. Lovell Reeve and Co., London.

HUTCHINSON, J., 1926.—The Families of Flowering Plants. Vol. I. Dicotyledons. Macmillan
and Co., London.

JACOBS, M. R., 1936.—The Primary and Secondary Leaf Bearing Systems of the Eucalypts and
their Sylvicultural Significance. Commonwealth Forestry Bureau. Bulletin No. 18. Com-
monwealth Government Printer, Canberra.

Jussieu, A. L. DE, 1812.—Note sur le Calice et la Corolle. Ann. Mus. Hist. Nat., 19: 431-2.

MaIpEN, J. H., 1923.—A Critical Revision of the Genus Hucalyptus. Vol. 6. Govt. Printer,
Sydney.

MUELLER, F. von, 1873.—Fragmenta Phytographiae Australiae. Vol. 8. Govt. Printer,
Melbourne. Fj

————,, 1879-84.—Hucalyptographia. Government Printer, Melbourne.

NAvDIN, C., 1883.—Mémoire sur les Hucalyptus Introduits dans la Région Méditerranéenne.
Ann. Sci. Nat., Sér. VI, 16: 344. :

OsBorN, T. G. B., 1947.—Book Review. New Phytol., 46: 289.

“RENDLE, A. B., 1938.—The Classification of Flowering Plants. Vol. 2. Dicotyledons. Cambridge
Univ. Press.

EXPLANATION OF PLATES II-III.
(All photographs x 30.)
Plate ii.

Fig. 1.—Longitudinal section through young bud showing insertion of petals.

Fig. 2.—Transverse section of young bud just below the level of the stigma showing the
four imbricate petals. Two of the petals are beginning to fuse.

Fig. 3.—Transverse section of operculum of young bud showing lateral position of the
entrance to the opercular channel.

Fig. 4.—Transverse section slightly lower than Fig. 3 showing triple nature of the channel
near the apex of the operculum.

Fig. 5.—Transverse section slightly lower than Fig. 4 showing the opercular channel.

Fig. 6.—Transverse section slightly lower~ than Fig. 5 showing three imbricate petal
segments.

?

Plate iii.

Fig. 7.—Transverse section slightly lower than Fig. 6 showing the four imbricate petals.
The characteristic infolding of the fourth petal beneath the other three is clearly shown.

Fig. 8.—Slightly oblique transverse section at the level of the stigma showing the four
imbricate petals. i

Fig. 9.—Longitudinal section of bud showing petals and opercular channel.

Fig. 10.—Longitudinal section showing gradual fusion of petal segments.

Fig. 11.—Longitudinal section of mature bud showing the outer calycine operculum, and
the inner operculum formed from the fusion of the petals. The irregularly pentagonal objects
just below the inner operculum are transverse sections through the filaments of inflexed
stamens.

36

SOME NOTES ON ATHROTAXIS.
By CHARLES G. ELLIOTT.
(Communicated by Dr. Patrick Brough.)
(Sixteen Text-figures. )

[Read 26th April, 1951.]

Synopsis.

This paper records some observations on Athrotaxvis made in Tasmania in 1946-47. The
relationships between the three species of the genus are briefly discussed. The pollen of
Athrotaxis differs from other Taxodiaceae in the absence of a germ pore or germinal papilla.
The intine is 2-layered. Female cones often contain larvae of a fly. Some features of the
gametophytes described by earlier workers are commented on in the light of more recent work
on Sequoia and Sequoiadendron. Athrotaxis differs from all other Taxodiaceae in the absence
of cleavage polyembryony. All the embryo initials contribute to a single dicotyledonous embryo.
The relationships between Athrotaxvis and Sequoia and Sequoiadendron are briefly discussed, but
no conclusions reached.

INTRODUCTION.

It is now more than 20 years since Saxton and Doyle (1929) published their
fragmentary account of the life history of Athrotaxis selaginoides. Since then a
description of the stem apex (Cross, 1943) is all that has been published on the
morphology of the genus. On the other hand, the life histories of Sequoia sempervirens
and Sequoia gigantea have been worked out (Looby and Doyle, 1937, 1942; Buchholz,
1939a, 1939b). Buchholz considers the two species sufficiently distinct to be placed in
different genera, and instead of Lindley’s invalid Wellingtonia has proposed the genus
Sequoiadendron for 8S. gigantea (Buchholz, 1939c). While some still consider the two
to belong to one genus (Doyle, 1945), many have accepted Sequoiadendron (see
especially Stebbins, 1948). This, together with the discovery in China of Metasequoia
glyptostroboides, the morphology and ecology of which are being investigated by a
number of botanists,* makes it highly desirable to know something more about
Athrotaxis. During 1946-47, at the University of Tasmania, I carried out some observa-
tions mainly on A. cupressoides Don, and though results were far from complete, the
following points seem worthy of record.

RELATIONSHIPS BETWEEN THE THREE SPECIES.

No one in Tasmania could agree with Dallimore and Jackson’s (1948, p. 207)
remark that the three forms ‘might well be regarded as gradations of one species”.
The two common species, A. cupressoides and A. selaginoides, are quite distinct, both
in the morphological characters used in taxonomy, such as the shape of the leaves and
cone-seales, and ecologically. A. cupressoides forms stands or occurs as single trees by
the side of tarns and along streams, while A. selaginoides is a taller forest tree
generally growing on sloping ground. It does sometimes grow beside tarns, but IL
have not observed it in such habitats when A. cuwpressoides is found there. Where the
two species occur near one another, their habitats are distinct, as has been noted by
Sutton (1928). <A. lawvifolia is rare, and does not form stands. Many people entertain
the possibility that it is a hybrid between the other two species, although there is little
substantial evidence for the view at present. However, many more trees than exist in
any one locality would be needed to show much segregation of characters.

MALE CONES AND POLLEN.
The male cones of Athrotaxis cupressoides can first be distinguished from vegetative
tips about mid-February, and their development proceeds until May, when spore mother
cells are found. Cones collected at Lake Dobson (Mt. Field National Park) on 25th

* A bibliography is given by Chu and Cooper (1950).

Led

BY CHARLES G. ELLIOTT. 37

August, 1946, and fixed in the field, contained microspores, but in some, the uppermost
sporangia had quadrinucleate protoplasts with the spindles of the second meiotic
division still present. Meiosis thus occurs about the second or third week of August.
The pollen is mature about a month later. I have not observed the shedding of pollen
in the field, but twigs with male cones collected at Lake Dobson on 9th September, 1946,
and placed in jars of water in the laboratory at Hobart shed pollen on 17th September,
and cones eollected at the Hugel Lakes (Lake St. Clair National Park) on 14th
September, 1947, shed pollen in Hobart on 18th September.

The pollen of Athrotaxis, in common with that of other Taxodiaceae and Cupres-
saceae, lacks air bladders, nor are there any male prothallial cells. As in other
members of these families, the exine is thin and is thrown off by the swelling of the
intine when the mature pollen grain comes into contact with water. The pollen of
A. cupressoides is spherical or subspherical, and has no trace of any germ pore or
germinal papilla. The diameter of acetolysed grains is 27-2 + 0-274. Three of the
layers of the exine recognized by Erdtman (1948) can be observed in Athrotazis. The
nexine, about 2u thick, is made up of an endonexine and an ectonexine of approximately
equal thickness. The sexine is represented by minute granulae, irregularly scattered,
and less than ly in diameter (Text-fig. 1).

Text-figures 1-6.—Athrotaxis cuwpressoides.

1. Acetolysed pollen grain. 2. Pollen, mounted in water, from which the exine has come
off, showing the protoplast surrounded by a gelatinous “halo’’. 38, 4. Pollen after four days in
water, showing enlargement of the protoplast within the ‘“‘halo’’. 5. Pollen after eight days in
M/4 sucrose. Protoplast with its wall (ii) emerging from the ‘‘halo” (oi). 6. Pollen mounted
in potash. es, exine; # and oi, inner and outer layers of the intine. Text-figure 1, x 640;
all others, x 400.

Text-figures 7-8.—Cones of Athrotaxis cupressoides. x 2.

7, about the time of fertilization, 14th December, 1946. 8, with nearly mature seed,
24th February, 1947.

The intine itself consists of two layers. When fresh pollen grains are mounted in
water, the grains first become turgid, and the average diameter is 30u. After the exine
is thrown off, the contents of the grain are seen to be surrounded by a spherical
gelatinous “halo”, the average diameter of the latter being 38u (Text-fig. 2). The
pear-shaped “cell” within the halo is closely surrounded by a wall of its own, especially
well seen in dry pollen mounted in potash (Text-fig. 6), which we may call the
“endintine”’ (ii), and the halo then is the “exintine”’ (oi). After four days in water
or sugar solutions the protoplast contained within the endintine has elongated consider-
ably, and although the shape of the protoplast is irregular, the exintine still preserves
a spherical form except whére it is actually forced out of position (Text-figs. 3, 4).
When the cell surrounded by the endintine emerges (Text-fig. 5), the exintine is seen
to have a definite thickness. A two-layered intine does not appear to have been

38 SOME NOTES ON ATHROTAXIS,

described before in a conifer, but since only the exine is preserved in fossils, it has
monopolized the attention of pollen morphologists to the exclusion of the intine.

The pollen of Athrotaxis differs markedly from that of Sequoia, Sequoiadendron,
and Metasequoia, in all of which there is a prominent germinal papilla, and it likewise
differs from Cryptomeria, Taxodium, and Glyptostrobus, which also have a germinal
papilla of some sort. It resembles more closely Cunninghamia and the Cupressaceae
(Wodehouse, 1935; Erdtman, 1943; Sterling, 1949).

FEMALE CONES.

Young female cones have not been distinguished from vegetative tips before mid-
September. In some localities production of cones varies from season to season. Cones
of A. cupressoides were found near Lake St. Clair both in 1946-47 and 1947-48. But in
the Mt. Field National Park, near the easterly limit of its distribution, A. cupressoides
formed no cones in 1946-47, although A. selaginoides several miles away did form cones
with fertile seeds. The same trees of A. cupressoides at Lake Dobson, however, bore
cones in 1947-48. Text-figure 7 is a sketch of a cone about the time of fertilization. A
fully grown cone is shown in Text-figure 8. Hames (1913, pp. 32, 33) has referred to
the arrangement of the vascular bundles in the cone scales. The ovules are in a single
row, and have their micropyles directed towards the cone axis. Cones of A. cupressoides
have frequently been found to contain larvae of a fly. Heavily infested cones were
deformed. The larvae appeared to be eating the ovules.

FEMALE GAMETOPHYTE.

Observations on the development of the female gametophyte are very incomplete.
As Saxton and Doyle (1929) reported, during the enlargement of the megaspore the
nucellus is soon consumed up to a thickness of one cell. In this respect Athrotazis
seems to be more advanced than Sequoia and Sequoiadendron. An important feature is
that in the free nuclear stages the nuclei are not evenly distributed round the embryo
sac but are congregated at the end away from the micropyle, where they are two deep
(Text-fig. 9). In this respect A. cupressoides resembles Sequoia sempervirens and differs
from Sequoiadendron giganteum (Looby and Doyle, 1942). Saxton and Doyle’s (1929)
Figure 8 of A. selaginoides shows nuclei distributed round the embryo sac in a single
layer, though more densely packed lower down. Looby and Doyle (1942) have shown
that in Sequoiadendron alveolation proceeds evenly all round the embryo sac. In
Sequoia, however, while alveoli are formed against the central vacuole, in the lower
part where nuclei are not in a single layer, walls form cutting out cells of irregular
arrangement. In A. selaginoides, Saxton and Doyle’s Figure 18 shows that wall forma-
tion takes place by the method in Sequoia, with alveoli being formed at the vacuolar |
edge of the basal portion.

Another feature reported by Saxton and Doyle in which Athrotaxis resembles
Sequoia and not Sequoiadendron is that the pollen tube grows over the surface of the
nucellus. This is apparently also the case in Metasequoia (Sterling, 1949).

Y EMBRYO.

Proembryo stages were not found. The earliest stage observed showed eight cells
in two tiers of four cells below the four-celled prosuspensor (Text-figs. 10, 13). The
early growth of these embryo cells (Text-figs. 11, 12) gives rise to a single embryonic
mass which may show some lobing (Text-fig. 14) and whose origin from four primary
cells is generally apparent for some time. However, cleavage of this embryo, such as
occurs in Sequoiadendron and Taxodium, was found never to take place.

There is no. primary suspensor. A massive secondary suspensor is produced by the
single embryo. An early stage of development of embryonal tubes is seen in Text-
figure 15. As the suspensor elongates and its upper part becomes folded, the nucellus
and rather indefinite prothallial tissue are included in the folds, and all together form
a compact mass at the micropylar end of the mature seed (Text-fig. 16). The embryo
has two cotyledons. Rosette cells are present in the early stages (Text-figs. 13, 14).
Nuclear divisions are numerous in the chalazal part of the gametophyte, but unfor-
tunately in no case could the chromosome number be counted; however, it does not

BY CHARLES G. ELLIOTT. 39

appear to be large. Embryo systems are frequently found in pairs, the result presumably
of the fertilization of two adjacent archegonia by the two male cells (Text-fig. 15).
Thus Athrotaxis exhibits only Simple Polyembryony as defined by Buchholz. In this
respect it is unique in the Taxodiaceae.

Text-figures 9-16.—Athrotaxis cwpressoides.

9. Longitudinal section of ovule at free nuclear stage, showing micropyle, integument, etc.
Embryo sac badly shrunken, but it shows nuclei congregated at lower end in two layers,
i4th December, 1946. x 60. 10. 8-celled embryo, 27th January, 1947. x 168. 11. Embryo with
+wo tiers of cells each of more than four cells, 27th January, 1947. x 168. 12. Three-tiered
embryo, 27th January, 1947. x 168. 13. Embryo system with two rosette cells, 4-celled
prosuspensor, and 8-celled embryo, similar to Text-figure 10. 27th January, 1947. x 72.
14. Embryo system with rosette cells and lobed multicellular embryo, 19th January, 1947. x 72.
15. Embryos of two different systems showing early stage in development of embryonal tubes,
26th January, 1947, x 72. (Material from which Text-figures 10-13 is taken is from a more
exposed locality at a higher altitude than those which gave Text-figures 14-15, hence the earlier
dates of the latter. Text-figures 10-15 from dissected embryos.) 16. Longitudinal section of
nearly mature seed, 24th February, 1947. x 24.

Free nuclear stages in the gametophyte were found in mid-December, and nearly
mature embryos late in February.

DISCUSSION.
Athrotaxis shows some significant differences from both Sequoia and Sequoiadendron
in its morphological features—in its pollen grain, simple polyembryony; also in its
one-cell-thick nucellus. It agrees with Sequoia in the method of wall formation in the

40 SOME NOTES ON ATHROTAXIS,

embryo sac, thin megaspore membrane, and two cotyledons. It resembles Sequoiadendron
in its mature leaves being of one type only, in the presence of a prosuspensor and
rosette cells, and one might expect in general features of proembryo development. It is
clearly impossible to derive the Athrotaxis type of embryogeny with its prosuspensor
from the Sequoia type which has none; but it may have been derived from that in
Sequoiadendron. Buchholz (1940, 1948) has rightly suggested that Athrotaxis, Sequoia-
dendron and Sequoia constitute a distinct subfamily, the Sequoidae. But it must be
borne in mind that Florin (1940) has shown that while Athrotaxvis was widely distributed
in the Southern Hemisphere in Tertiary times, “no representatives of this genus are
known with certainty from the Northern Hemisphere” (p. 90). Reports of the southern
occurrence of Sequoia, Florin shows, were the results of misdeterminations. On the other
hand, Sequoia was abundant in Europe and Metasequoia in North America (Chaney,
1948). It is clear that the position of Athrotaxvis is not merely intermediate between
Sequoia and Sequoiadendron, but we are not yet able to define their relationships more
precisely.

Acknowledgements.

I am greatly indebted to Professor Joseph Doyle, of University College, Dublin,
who agreed that some notes should be written on this work before his own observations
from trees growing in Ireland are published.

My thanks are also due to Drs. E. I. McLennan and I. C. Cookson, of the University
of Melbourne, and to Dr. P. Brough, of the University of Sydney, for their interest in
this work.

References.

BucHHo.uz, J. T., 1939a.—The Morphology and Embryogeny of Sequoia gigantea. Amer. J. Bot.,
26: 93-101.

, 1939b.—The Embryogeny of Sequoia sempervirens with a comparison of the Sequoias.
Amer. J. Bot., 26; 248-257.

, 1939ce.—The Generic Segregation of the Sequoias. Amer. J. Bot., 26: 535-8.
, 1940.—The Embryogeny of Cunninghamia. Amer. J. Bot., 27: 877-883.
, 1948.—Generic and Subgeneric Distribution of the Coniferales. Bot. Gaz., 110: 80-91.

CHANEY, R. W., 1948.—The bearing of the living Metasequoia on problems of Tertiary Palaeo-
botany. Proc. Nat. Acad. Sci. Wash., 34: 503-515.

CHu, K., and Cooper, W. S., 1950.—An ecological reconnaissance in the native home of
Metasequoia glyptostroboides. EHcology, 31: 260-279.

Cross, G. L., 1943.—The Shoot Apex of Athrotaxis and Taiwania. Bull. Torrey Bot. Cl., 70:
335-348.

DALLIMORE, W., and JAcKSON, A. B., 1948.—A Handbook of Coniferae. 3rd Edition. Arnold,
London.

Doy.LeE, J., 1945.—Naming of the Redwoods. Natwre, 155: 254-7.

EAMES, A. J., 1913.—The Morphology of Agathis australis. Ann. Bot., Lond., 27; 1-38.

ERDTMAN, G., 1943.—Introduction to Pollen Analysis. Chronica Botanica Co., Waltham.

, 1948.—Did Dicotyledonous Plants exist in early Jurassic Times? Geol. Foren. Stockh.
Forh., 70: 265-271.

FLORIN, R., 1940.—The Tertiary Fossil Conifers of South Chile and their phytogeographical
significance, with a Review of the fossil Conifers of Southern Lands.’ K. Svenska Vetensk.
Akad. Handl., 3rd Ser., 19 (2).

Loopy, W. J., and Doyue, J., 1937.—Fertilization and Proembryo Formation in Sequoia. Sci.
Proc. R. Dublin Soc., 21: 457-476.

———.,, 1942.—Formation of Gynospore, Female Gametophyte, and Archegonia in Sequoia.
Sci. Proc. R. Dublin Soc., 23: 35-54.

SAxTON, W. T., and Doytg, J., 1929.—The Ovule and Gametophytes of -Athrotaxis selaginoides.
Ann. Bot., Lond., 43: 833-840.

STEBBINS, G. L., 1948.—The Chromosomes and Relationships of Metasequoia and Sequoia.
Science, 108: 95-8.

STERLING, C., 1949.—Some Features in the Morphology of Metasequoia. Amer. J. Bot., 36:
461-471.

Sutton, C. S., 1928.—A Sketch of the Vegetation of the Cradle Mountain, Tasmania, and a
Census of its Plants. Pap. Roy. Soc. Tasm., 1928: 132-159.

WODEHOUSE, R. P., 1935.—Pollen Grains. McGraw Hill, New York.

41

THE PARAMPHISTOMES (TREMATODA) OF AUSTRALIAN RUMINANTS.
PART I. SYSTEMATICS.
By P. H. Durig£, B.Sc.,
Division of Animal Health and Production, C.S.I.R.O., Veterinary Parasitology
Laboratory, Yeerongpilly.
(Plates iv—v.)
[Read 26th April, 1951.]

Synopsis.

A study has been made of the species of Paramphistomidae occurring in the rumen and
reticulum of Australian cattle.

Identifications were based on the system devised by Nasmark 1937, involving an
examination of the acetabulum, genital atrium and pharynx, as seen in median sagittal sections.

The species previously known as Paramphistomum cervi, was found to consist of
Ceylonocotyle streptocoelium (Fischoeder, 1901) Nasmark, 1937, and Calicophoron calicophorum
(Fischoeder, 1901) N&smark, 1937, whilst the species previously known as P. cotylophorwm
was determined as P. ichikawat Fukui, 1922. A fourth species was present among the material
examined but as yet remains undetermined.

In view of the importance given by Edgar (1938), Roberts (1934), Ross and Gordon
(1936) to Paramphistomes as causal agents of parasitic gastro-enteritis in sheep and
cattle in Australia, investigations were commenced into their bionomics, pathogenicity,
and control. Three species of Amphistome flukes have been recorded from Australian
eattle (Seddon, 1947), namely, Paramphistomum cervi (Zeder, 1790), P. explanatum
(Creplin, 1847), and P. cotylophorum (¥ischoeder, 1901). P. cervi is recorded from
Queensland, New South Wales, Victoria, Western Australia and Tasmania; P. explanatum
from Queensland; and P. cotylophorum from Queensland and New South Wales. P. cervi
and P. cotylophorum are noted as occurring also in sheep.

The author (Durie, 1949) published a preliminary note implicating the Planorbid
snails, Glyptanisus gilberti and Segnitilia alphena as the intermediate hosts of P. cervi
and P. cotylophorum respectively. These observations were based on the recovery of
flukes, identified as those two species, from lambs fed cysts obtained from naturally
infested snails. Subsequent attempts to infest snails experimentally with miracidia,
considered to be those of P. cervi, were unsuccessful, and it seemed evident that the
flukes that had been identified as P. cervi probably consisted of more than one species.
A careful, taxonomic study of the Australian species was therefore considered essential
before further work on the bionomics of these parasites could be attempted.

MATERIALS AND METHODS.

The bulk of the material examined has been collected from the rumen and reticulum
of cattle slaughtered at abattoirs near Brisbane, Queensland, and drawn from the coastal
and sub-coastal areas of the State. Additional material has been obtained from other
States through the courtesy of slaughtering inspectors and veterinary research
laboratories.

It is evident from the literature, Fischoeder (1901), Stiles and Goldberger (1910),
Maplestone (1923), Travassos (1934), Dawes (1936, 1946) and Nasmark (1937), that the
family Paramphistomidae is a very difficult one from the systematic point of view.
This has been discussed fully by Nasmark (1937), who attempted to obtain a clearer
conception of the family’s classification by careful studies on serial sections, particularly
median sagittal sections, in which attention was largely directed to the muscle structures
of the acetabulum and pharynx.

Nasmark’s system has been used by the author in the studies reported here and
is considered a satisfactory one, at least, in so far as the Australian species are
concerned.

q

42 THE PARAMPHISTOMES OF AUSTRALIAN RUMINANTS. I,

The measurements of body length and breadth were made on entire specimens fixed
in Carnoy’s fluid, whereas all other measurements were made on median sagittal
sections. Ova dimensions were obtained from eggs deposited in physiological saline.

It was noted very early in these studies that Gigantocotyle (Paramphistomum)
explanatum (Creplin, 1847) Nasmark, 1937, was not represented in the Australian
material under examination. This species has a readily recognized and distinct
musculature of the acetabulum and pharynx (Nasmark, 1937), and specimens received
from Ceylon were easily identified on this character. G. explanatum occurs in the
gall bladder and bile ducts of bovines (Dawes, 1946; Nasmark, 1937). In view, therefore,
of the absence of reports from Australia of amphistomes occurring in these parts of
the body, together with the fact that the species was not represented among the
hundreds of amphistomes obtained for examination from various parts of Australia, it
is highly probable that G. explanatum is not present in this country and that records
of its occurrence by Roberts (1984) and Ross and Gordon (1936) are erroneous.

Two and possibly three species have been recognized among the flukes previously
identified as P. cervi. Calicophoron calicophorum (Fischoeder, 1901) Nasmark, 1937,
is very common and widespread and is probably the most prevalent amphistome in
Australian cattle. Ceylonocotyle streptocoelium (Fischoeder, 1901) Nasmark, 1937, is
the second species, whereas the third species as yet remains undetermined, but appears
to be closely related to C. calicophorum.

A re-examination of the species previously recorded in Queensland and New South
Wales as Paramphistomum cotylophorum has shown it to be Paramphistomum ichikawai
Fukui, 1922. A detailed description of each of these species is given below.

CEYLONOCOTYLE STREPTOCOELIUM (Fischoeder, 1901) Nasmark, 1937.

Host: Bos taurus—rumen and reticulum.

Distribution: Beaudesert, Queensland; Gympie, Queensland.

Length 4:55 mm. (3:0-5:0 mm.), breadth 2-4 mm. (2-3 mm.); D.V. measurement
1-4 mm. (1:1-1:8 mm.); dorsal line curved, curvature greater in the posterior third of
the body; ventral line plain to slightly concave. Acetabulum conforms to Nasmark’s
Streptocoelium type opening postero-ventrally, its maximum diameter 1:0 mm. (0-7-1-2
mm.), the ratio of its diameter to body length being 1:4:°5 (1:3:5-1:5:0). Pharynx
conforms to Nasmark’s Paramphistomum type, 0:40 mm. (0:30-0:46 mm.) in length;
ratio of pharynx length to body length 1:10:0 (1:9-1:12). Oesophagus 0:22 mm. (0-17-
0-25 mm.) in length; oesophageal sphincter present. Genital atrium with genital
sphincter, conforms to Nasmark’s Streptocoelium type; ratio of genital atrium to
acetabulum diameter 1:3:9 (1:3:5-1:4:5). Testes moderately lobed, oval to rectangular
in shape, situated tandem and measuring 0°-5 mm. in length and 1:0 mm. in D.V.
direction. Egg 0-148 x 0:074 mm. (0:154 x 0-070 — 0-145 x 0:078 mm.).

Acetabulum (Pl. iv, fig. 1).—The acetabulum conforms to Nasmark’s Streptocoelium
type. The circular muscle series are characterized by their relatively few muscle units,
with the interior series presenting a greater number than the exterior series. There is
no division into de, and de, circular. De and di circular correspond closely in structure
and in the number of units to ve and vi circular. - The de series shows a slight
variation in the size of the units, these being smaller externally and increasing in size
internally. The number of units present in 10 specimens is shown in Table 1.

Pharynz.—The pharynx conforms to Nasmark’s Paramphistomum type. The internal
circular muscle layer consists of small closely packed units. The internal surface of the
pharynx is smooth or with very small papillae. The interior longitudinal layer appears
as a clearly developed band and extends inwards for about one-quarter the width of
the pharynx. The middle circular layer is absent. The radial muscles are not strongly
developed but are clearly visible. The exterior circular layer is rather indistinct with
small units. The exterior longitudinal layer is indistinct and narrow. The basally
circular layer is strongly developed and the series appear to be arranged in two rows.
A slight trace of the anterior sphincter is present, but both the lip sphincter and
posterior sphincter are absent.

BY P. H. DURIE. 43

Genital Atrium (PI. iv, fig. 2)—The genital atrium of this species is much smaller
than that of P. ichikawai, and possesses both a genital sphincter and sphincter papillae.

The genital papilla is rather thick and variable in shape. Sphincter papillae,
although present, are not as strongly developed as described by Nasmark (1937). The
genital sphincter consists of several bundles of closely packed fibres. It is quite
conspicuous, but again not as strongly developed as described by Naésmark (1937).
The ventral atrium is only slightly developed.

TABLE 1.
C. streptocoelium: Unit Series of Circular Musculature of the Acetabulum from a
Median Sagittal Section.

de. di ; | vi ve
: <
12 | 32 37 15
14 | 30 25 15
17 32 | 29 15
15 30 27 13
15 30 30 14
14 31 22 12
12 28 25 13
14 29 26 14
13 31 28 14
ne 32 26 13
Discussion.

The specimens differ from Nasmark’s description mainly in body length (7:6 mm.),
but the measurements of the pharynx and other structures, except the oesophagus, agree
closely with his description. Consequently any ratios given by Nasmark involving
body length are greater than those given by the author.

Body length is regarded as a useful character to define the approximate size of a
species, and to differentiate species which vary greatly in size. However, the length ~
of the body in living specimens is an extremely variable one, as flukes actively elongate
and contract continually. In fixed specimens the body length may vary according to
the type of fixative and method of fixing employed and is therefore considered unsuitable
for determining critical ratios.

In specimens belonging to the genus Ceylonocotyle Laurer’s canal and the excretory
canal (PI. iv, fig. 3) do not cross one another, a character shared also by the genera
Nilocotyle (Nasmark, 1937) and Buzifrons (Nasmark, 1937). Ceylonocotyle may be
separated from these, however, by certain features of the acetabulum, and particularly
by the absence of strongly developed radial muscles. :

C. streptocoelium may be distinguished from other members of the genus by the
absence of a strongly developed oesophageal bulb and lip sphincter, and from the closely
allied ©. orthocoelium by the presence of a genital sphincter and an oesophageal
sphincter (Pl. iv, fig. 4). In the living state it may be separated from P. ichikawai,
which it resembles closely in size, colour and shape, by the absence of any noticeable
thickening around the genital pore. Both species are found in the rumen, with
C. streptocoelium in the reticulum as well.

The genital atrium in the specimens examined differed slightly from the description
of the Streptocoelium type given by Nasmark (1937), but this is probably due to the
fact that Nasmark’s material was poorly preserved.

The relationship between dimensions of the pharynx, genital atrium and acetabulum
is shown in Plate v, figure 5.

PARAMPHISTOMUM ICHIKAWAI Fukui, 1922,
Host: Bos taurus—rumen only.
Distribution; Coastal districts, Queensland. ,

44 THE PARAMPHISTOMES OF AUSTRALIAN RUMINANTS. I,

Length 5:7 mm. (3:12-7:00 mm.), breadth 2-7 mm. (2-3 mm.); D.V. measurement
1:80 mm. (1:40-2:17 mm.); dorsal line slightly curved, but more strongly so in posterior
two-thirds; ventral line plain to slightly concave (PI. v, fig. 6). Acetabulum conforms
to Nasmark’s Paramphistomum type, opening postero-ventrally, its maximum diameter
1:3 mm. (1:12-1:50 mm.), and the ratio to body length is 1:4:3 (1:3:8-1:4:7). Pharynx
conforms to Nasmark’s Paramphistomum type, 0°60 mm. (0:52-0:80 mm.) in length;
ratio of pharynx length to body length 1:9:5 (1:8-2-1:13:2), oesophagus 0-28 mm. (0:17—
0-50 mm.) in length. Genital atrium conforms to Nasmark’s ichikawai type, measures
0-53 mm. (0:45-0:62 mm.) in external diameter; ratio of genital atrium diameter to
acetabulum diameter 1:2:5 (1:2-1:2:9). Testes 0-75 mm. long by 1:0 mm. in D.V.
direction, lobed, oval, somewhat “shamrock” shaped, and ‘are situated tandem to each
other. Egg 0:143 x 0:064 mm. (0-129 x 0:067—0-148 x 0-059).

Acetabulum (Pl. v, fig. 7): The acetabulum conforms to Nasmark’s Paramphi-
stomum type. The dorsal, exterior, circular series is divided iato de, and de, circular.
The ventral exterior circular series corresponds to de, circular and shows no division
into two groups. The de, circular series is well developed and is strongest outwards.
The division between de, and de, circular is fairly clearly marked. The de. circular
series is weakly developed in comparison to de, circular with the units spaced at
irregular intervals. The di series is well developed, with units attaining their maximum
size in the centre. Vi circular corresponds closely in structure and number to di
circular, and ve circular corresponds similarly to de, circular. The number of units
in each series was found to be variable, especially in regard to de. circular, and, in
some cases, considerable difficulty was experienced in obtaining an accurate count.
The acetabulum, however, conformed to Nasmark’s Paramphistomum type plan in all
specimens examined, and the musculature was more strongly developed than in
C. streptocoelium. The number of unit series in each group taken from 10 specimens
is shown in Table 2.

TABLE 2.
P. ichikawai: Unit Series of Circular Musculature of the Acetabulum from a Median Sagittal Section.
|
de}. des. di vi. "ve.
23 6 40 48 17
19 9 43 42 14
19 8 46 48 15
20 8 43 43 16
20 8 39 43 17
20 5 36 37 17
23 12 43 44 | 18
19 9 40 46 | 15
19 ° 10 31 38 20
20 3 37 46 | 16

Pharynz: The pharynx conforms to Nasmark’s Paramphistomum type. The internal
circular muscle layer consists of small units increasing slightly in size towards the
anterior end. The internal surface of the pharynx is papillated slightly in the anterior
portion. The interior, longitudinal layer extends inwards to about one-third the width
of the pharynx; the interior margin is clearly defined. The middle circular layer is
absent. The radial musculature is clearly visible but not strongly developed. The
exterior circular layer is somewhat indistinct anteriorly but clearer towards the posterior
end. The exterior longitudinal layer is narrow but clearly visible. The basally
circular layer contains small units which lie in two rows towards the interior edge.
The anterior sphincter, lip sphincter and posterior sphincter are absent. '

Genital Atrium (PI. v, fig. 8): The genital atrium belongs to Nasmark’s Group II Bl,
which embraces forms with no genital sphincter, but with sphincter papillae. The
genital papillae are thick and clumsy. The genital atrium conforms to Nasmark’s

BY P. H. DURIE. 45

ichikawai type. The tissue of the walls of the genital atrium is thick and muscular,
with well-developed radial musculature, and could quite easily be mistaken for a sucker.
This is particularly the case in living specimens, where the structure shows a typical
“sucker-like”’ appearance. However, sections show that, although the atrium tissue is
clearly divided from the rest of the body tissue, a true delimiting membrane, charac-
teristic of the genus Cotylophoron, is lacking.

Specimens of this species in the living state are pink to white in colour and appear ,
to be confined to the rumen, as they have never been observed attached to the reticulum,
even when heavy infestations are present. They closely resemble C. streptocoelium in
size and colour, but may be distinguished from that species by the “sucker-like” genital
atrium.

Discussion.

The above description agrees closely with that of Nasmark for P. ichikawai but
differs slightly in body length dimensions. Measurements of other structures are in
close agreement. The remarks made in regard to body length in the description of
C. streptocoelium apply equally to this species. P. ichikawai exhibits a remarkably
constant ratio between the genital atrium diameter and acetabulum diameter, and
differs markedly from C. streptocoelium in this respect.

CALICOPHORON CALICOPHORUM (Fischoeder, 1901) Nasmark, 1937.

Host: Bos taurus—rumen and reticulum.

Distribution: Coastal and sub-coastal regions of Queensland, New South Wales,
Victoria, Western Australia, South Australia and Tasmania.

Length 11-1 mm. (i0-12 mm.), breadth 6-6 mm. (5:8-7-5 mm.); D.V. measurement
2°9 mm. (2:1-3:9 mm.); dorsal line evenly curved, truly conical in shape, with maximum
breadth at the posterior end; ventral surface slightly concave. Acetabulum conforms
to Nasmark’s Calicophoron type, 2-4 mm. (1:60—3:2 mm.) in diameter; ratio of acetabulum
diameter to body length 1:4. Pharynx conforms to Nasmark’s Calicophoron type, 1:4 mm.
(0-93-15 mm.) in length; ratio of pharynx length to body length 1:8. Oesophagus
0-9 mm. (approx.). Testes placed diagonally side by side, deeply lobed. Genital atrium
conforms to Nasmark’s Calicophoron type. Hegg 0:133 x 0:080 mm. (0-126 x 0:072—-0:147 x
0-090 mm.). '

Acetabulum: The acetabulum conforms to Nasmark’s Calicophoron type. The cir-
cular musculature is well developed and the dorsal, exterior, circular layer is not
divided into de, and de, circular. In this species the circular musculature of the dorsal
half of the acetabulum corresponds almost exactly to the musculature of the ventral
portion. The de circular layer is well developed and consists of large well-formed units,
largest in the centre and decreasing in size interiorly and exteriorly. The units of the
di circular layer reach their maximum size close to the exterior edge and gradually
decrease interiorly. Ve-and vi circular correspond to de and di series. The number
of units in the circular musculature is shown in Table 3.

Another series of units (Pl. v, fig. 9) is present in this type of acetabulum, which
has not been described in any of the other types of Nasmark (1937). This series consists
of small units, about 15 in number and about one-tenth the size of the largest units of
the other circular musculature. It connects the exterior ends of de and di circular in
the dorsal half with a corresponding series connecting ve and vi circular in the ventral
portion. As this series has not been previously described, the term lateral, circular
layer is proposed.

Pharynx: The ventral, circular, muscle layer consists of very small, closely packed
units, increasing slightly in size towards the posterior end. The interior, longitudinal
layer is narrow and rather indistinct in comparison with that in the Paramphistomum
type, and the interior border is not clearly marked. The middle, circular layer is
absent. The radial muscles are single fibres showing little ramification. The exterior,
circular layer is indistinct, with the units somewhat diffuse and not clearly visible.
The exterior, longitudinal layer is narrow but distinct. The basally circular layer is

46 THE PARAMPHISTOMES OF AUSTRALIAN RUMINANTS. I,

weakly developed and is confined to a single row of units. A trace of the anterior
sphincter is present, but the posterior sphincter and the lip sphincter are absent.

Genital Atrium (PI. v, fig. 10): The genital atrium conforms to Naismark’s Calico-
phoron type and is highly characteristic of the species (Nasmark, 1937), forming a
useful diagnostic character in living flukes. It is eversible and may either be found
protruding from the genital region of the body, surmounted on a genital pillar, or
retracted well within the body, as illustrated in Plate v, figure 10. Surrounding the
genital pore or pillar, depending on the degree of contraction of the atrium, is a
flattened circular area, bounded by a small circular ridge and, in living specimens,

TABLE 3
f-
C. calicophorum: Unit Series of Circular Musculature of the Acetabulum from a Median
Sagittal Section.

de. di. | vi. ve.
|
13 40 44 18
12 41 45 16
18 40 41 - 16
13 36 39 13
20 45 48 25
23 44 48 27
21 46 | 50 23

swift circular waves of contraction are visible passing around the area. The genital
sphincter is well marked and consists of closely packed units, the radial musculature
being strongly developed. The sphincter papilla is also well developed and appears
as a large closely packed area at the base of the genital papilla. Both the genital
sphincter and sphincter papillae are more strongly developed than in either C. strepto-.
coelium or P. ichikawdi.

Discussion.

Nasmark (1987) described two species in the genus Cdalicophoron, namely,
C. calicophorum and C. ijimai, which differ only in regard to the pharynx. Apart from
purely anatomical differences, he gives a range of pharynx length measurements for
both species which, in his opinion, are distinct enough to differentiate between them.
His measurements are 1:50-2:0 mm. for C. calicophorum and 0:95 mm. (average of six
specimens) for C. ijimai. Measurements of pharynx length made by the author on
Australian specimens identified as belonging to the genus Calicophoron gave a range
of 0-73 mm. to 150 mm. with lengths occurring throughout the range. This suggests
that the pharynx is of the type belonging to C. ijimai.

According to Nasmark (1937), the pharynx of both species varies anatomically
in several respects, particularly with regard to the interior, circular, muscle layer and
to the presence of a well-marked posterior sphincter. Specimens of C. ijimai have both
these characters strongly developed, while in C. calicophorum these structures are
poorly developed or absent. In the series examined by the author the pharynx of some
specimens varied in the size of the interior circular layer, but in no instance was a
well-developed posterior sphincter seen.

In view of this lack of agreement with Nasmark’s description of C. ijimai, and
also because the type material of OC. calicophorum was obtained from Queensland by
Fischoeder, it has been tentatively decided to consider the species as C. calicophorum
pending investigation into its life history.

SUMMARY.
A taxonomic study has been made of the Paramphistomidae occurring in domestic
ruminants in Australia,

BY P. H. DURIE. 47

Three species were recognized, namely, Ceylonocotyle streptocoelium (Fischoeder,
1901) Nasmark, 1937, Paramphistomum ichikawai Fukui, 1922, and Calicophoron calico-
phorum (Fischoeder, 1901) Nasmark, 1937. There is possibly a fourth species, as yet
undetermined.

Acknowledgements.

The author wishes to thank Dr. Ben Dawes, University College, London, and
Dr. F. W. Price, Zoological Division, Bureau of Animal Industry, Department of
Agriculture, Washington, D.C., U.S.A., for their interest and advice.

Thanks are also accorded the various people who forwarded specimens, particularly
Mr. W. Kearney, Inspector of Stock, Oxley.

This work was carried out as part of the research programme of the Division of
Animal Health and Production, C.S.I.R.O.

References.

Dawes, B., 1986.—On a collection of Paramphistomidae from Malaya with a Revision of the
Genera Paramphistomum FWischoeder 1901 and Gastrothylax Poirier 1883. Parasitology,
28 : 330-354.

, 1946.—The Trematoda. Cambridge Uni. Press, London.

Durig£, P. H., 1949.—A preliminary note on the life cycle of Paramphistomum cotylophorum
(Fischoeder, 1901) and P. cervi (Schrank, 1790) ete. Awst. Vet. J., 25 (9): 209.

Epe@ar, G., 1938.—Paramphistomiasis of young cattle. Awst. Vet. J., 14:27.

FISCHOEDER, F., 1901.—Die Paramphistomiden der Saéugethiere. Zool. Anz., 24 (646) : 367-375.

MAPLESTONE, P. A., 1923.—A revision of the Amphistomata of Mammals. Ann. Trop. Med.
Paras. Uni. of Liverpool, 27 (2) :113-212.

NAsMARK, K. E., 1937.—A revision of the Trematode Family Paramphistomidae. Zool. Bidrag
Uppsala, 16: 301-565. &

Ropers, F. H. S., 1934.—Worm parasites of domesticated animals in Queensland. Queensland
Agric. J., 41 (3) : 245-252.

Ross, I. C., and Gorpon, H. Mcl., 1936.—The Internal Parasites and Parasitic Diseases of
Sheep. Angus and Robertson, Sydney.

SEDDON, H. R., 1947.—Host check list of helminth and arthropod parasites present in domesti-
cated animals in Australia. C. of Aust. Dept. of Health, Service Publication No. 2.

STILES, W., and GOLDBERGER, J., 1910.—A study of the anatomy of Watsonius watsoni (n.g.)
of man of nineteen allied species of Mammalian Trematode worms of the superfamily
Paramphistomoidea. Hyg. Lab. Bull. No. 60:1-261.

TRAVASSOS, L., 1934.—Synopse dos Paramphistomoidea. Mem. Inst. Oswaldo Cruz., 29 (1):
19-178.

Witimort, S., 1950.—Gametogenesis and early development in Gigantocotyle bathycotyle
(Wischoeder, 1901) Nasmark, 1937. J. Helminth., 25 (1-2) :1-14.

EXPLANATION OF PLATES IV-V.

PLATE Iv.
Ceylonocotyle streptocoelium.

1. Median sagittal section through the acetabulum showing structure and arrangement. of
the circular musculature, x60. de, Dorsal exterior circular layer; di, Dorsal interior circular
layer ; Ra, Radial muscle fibres; vi, Ventral interior circular layer; ve, Ventral exterior circular
layer. f

2. Median sagittal section through the genital atrium, x184. GP, Genital papilla; GS,
Genital sphincter, SP, Sphincter papillae; <, ductus ejaculatorius; @, uterus.

3. Median sagittal section through Laurer’s canal and the excretory canal, x 60. KC,
Excretory canal; LC, Laurer’s canal.

4. Median sagittal section through part of the pharynx and oesophagus, showing a well-
developed oesophageal sphincter, x184. INT, Intestine; Oe, Oesophagus; OeS, Oesophageal
sphincter; Ph, Pharynx (portion of).

PLATD V.
Ceylonocotyle streptocoelium.

5. Median sagittal section, showing the position and size of the pharynx, genital atrium and
acetabulum, x 18. Ac, Acetabulum; DuE, Ductus ejaculatorius; EB, Excretory bladder; GA,
Genital atrium; Oe, Oesophagus; OV, Ovary; Ph, Pharynx; PM, Pars musculosa; T, Testes;
Ut, Uterus.

Paramphistomum ichikawai.

6. Median sagittal section, showing the position and size of the pharynx, genital atrium
and acetabulum, x15. Ac, Acetabulum; GA, Genital atrium; Oe, Oesophagus; Ph, Pharynx.

48 THE PARAMPHISTOMES OF AUSTRALIAN RUMINANTS. I,

7. Median sagittal section of the acetabulum showing structure and arrangement of the
circular musculature, x42. de,, Dorsal exterior circular layer 1; de,, Dorsal exterior circular
layer 2; di, Dorsal interior circular layer; Ra, Radial muscle fibres; ve, Ventral exterior
circular layer; vi, Ventral interior circular layer.

8. Median sagittal section through the genital atrium, x 60. GP, Genital papilla; SP,
Sphincter papillae; Ra, Radial muscle fibres; ¢&, Ductus ejaculatorius; 9, Uterus.
Calicophoron calicophorum.

9. Median sagittal section through portion of the acetabulum showing the lateral circular
muscle layer, x 180. 1, Lateral circular muscle layer; ve, Portion of, ventral exterior circular
layer; vi, Portion of ventral interior circular layer.

10. Median sagittal section through the genital atrium, x60. GP, Genital papilla; GS.
Genital sphincter; Ra, Radial muscle fibres; SP, Sphincter papillae.

Proc. Linn. Soc. N.S.W., 1951. PLATE I.

Septoria pepli on leaves of Euphorbia peplus.

Il.

7
yi

PLATE

N.S.W., 1951.

Soc.

IN.

Proc. Lin

alyplus gummifera.

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PLATE

»S.W., 1951.

Proc. LINN. Soc.

ane

yptus gummifera.

al

we

in #

culum

Oper

Proc. Linn. Soc. N.S.W.. 1951. PiPAni LV.

Ceylonocotyle s treptocoelium.

Proc. LINN. Soc. N.S.W., 1951. PLATE vy.

5, Ceylonocotyle streptocoeliumo. 6-8, Paramphistomwun ichikawai.
9-10, Calicophoron calicophorum.

*

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49

A REVIEW OF THE AUSTRALIAN SPECIES OF SARCOCHILUS (ORCHIDACEAE).
By the Rev. H. M. R. Rupp, B.A.
(One Text-figure.)

[Read 27th June, 1951.]

Synopsis.

Sixteen -valid species of the genus are recognized in this review, including one
(S. tricalliatus) newly admitted to specific rank. Nine excluded species are listed at the end
of the review.

Some different conceptions of the character of Sarcochilus are briefly discussed, and a
description of the genus as understood by the author is given. The remainder of the paper
consists of notes on the individual species. In the case of S. australis Lindl., which F. M.
Bailey recorded for Queensland under the synonym S. parviflorus, the author expresses the
view that there are no authentic records of this species occurring north of Gosford, N.S.W.,
and that the Queensland record should probably be applied to S. spathulatus Rogers.

Considerable difficulty is experienced in attempting to decide between. varying
interpretations of the genus Sarcochilus, which Pfitzer places between Grosourdya
Rehb. f. and Dendrocolla Bl., in the sub-tribe Aerideae of the tribe Sarcanthinae. The
late J. J. Smith, in Blumea, I, 1 (1930), pp. 194-215, described Sarcochilus as distin-
guished by a long straight column with a very short foot, and a lip with a very small
pit at the base. If this description is to be generally accepted, it seems to me that it
will mean the removal of all our Australian species to some other genus or genera.
For with the possible exception of S. australis Lindl., they are all distinguished by a
short column with a long foot, to which the lateral sepals are usually adnate; and in
most instances the “pit” or sac, situated below the mid-lobe of the labellum, is relatively
large. As I have not been able to ascertain whether J. J. Smith’s interpretation as
given above is generally acceptable, in this review I have retained the older conception
of the genus, in which the column is short with a long foot. So far as I am aware, no
one up to the present has proposed to remove the Australian species on the ground of
their incompatibility with J. J. Smith’s description. On other grounds, various authors
have from time to time transferred to other genera quite a number of Australian plants
previously includéd in Sarcochilus, as will be seen from the list of “Excluded Species”
at the end of this review. The latest of these removals is proposed by the present writer
in the Victorian Naturalist, Vol. 67 (1951), 206, where Mueller’s S. divitiflorus is made
the type of a new genus to be known as Rhinerrhiza. With the exclusion of this plant,
there remain sixteen Australian species to be reviewed, viz.: ;

1, S. Fitegeraldii; 2, S. Hartmannii; 3, S. falcatus; 4, S. Weinthalii; 5, S. australis;
6, S. spathulatus; 7, S. olivaceus; 8, S. Harriganae; 9, S. dilatatus; 10, 8S. Longmanii;
11, 8S. Bancroftii; 12, S. Ceciliae; 138, S. Hillii; 14, S. tricalliatus; 15, S. eriochilus;
16, S. minutiflos.

Before proceeding to review these in detail, however, I wish to submit the following
brief description of the genus itself as I understand it.

SARCOCHILUS R.Br.

Epiphytes or rock plants, usually rather small. Stems _ short. (Exception in
Australia, 8S. Fitezgeraldii.) Leaves from broadly lanceolate to linear, often more or less
faleate and usually distinctly channelled above. Racemes emerging below the leaves.
Flowers from a few mm. to 3 cm. in diameter, racemose, often numerous and showy,
in some species very fragrant. Sepals and petals approximately equal, free, relatively
broad at least in the distal portion, the lateral sepals usually more or less dilated at the
base and adnate to the foot of the column. Labellum articulate at the base of the
column-foot, spurless, trilobate. Lateral lobes erect, relatively large, curving inward;

H

50 THE AUSTRALIAN SPECIES OF SARCOCHILUS (ORCHIDACEAE),

mid-lobe usually very short, with a rather conspicuous sac or protuberance below.
projecting in front; disc with a few stalked calli. Column short, with a long foot;
anther operculate; stigma semicircular to oblong; pollinia 2.

An illuminating article on the genus by the late Dr. R. S. Rogers will be found in
the Australian Orchid Review, Vol. II, No. 3 (Sept., 1937), pp. 9-12. One or two
corrections are required. (The name is several times misspelt Sarchochilus, but this is.
probably a printer’s error.) On p. 10 the author states, “In habit the genus Sarcochilus
may be distinguished from Thrixspermum by the elongated inflorescence of the former,
on which all the flowers of a raceme open on the same day, or else begin to bloom
suddenly in series on a gradually lengthening inflorescence”. This is not correct,
at least so far as Australian species are concerned. The only Australian species with
elongated racemes on which all the flowers open on the same day is S. divitiflorus
F. Muell., now removed from the genus. I know of no species in which the flowers begin
to bloom suddenly in series on a gradually lengthening inflorescence, nor can I find any
observer who does. Dr. Rogers remarks that Thrixspermum had not been reported from
Australia at the time of writing; but Schlechter in 1911 (Orchis, v. 55) had removed
S. platystachys F. M. Bail. to that genus, and the transfer is not likely to be challenged.
Rogers gave the number of Australian species of Sarcochilus in 1937 as “17 (perhaps
18)”. Presumably he included S. phyllorhizus F. Muell. (now transferred to
Chiloschista) as well as S. platystachys. I do not know what his possible 18th species
was, but he felt some uncertainty as to the status of S. falcatus var. montanus, which
R. D. Fitzgerald originally published as S. montanus. Fitzgerald’s reduction of it to
a variety of S. falcatus, however, has been generally endorsed. The exclusion of
.S. phyllorhizus, S. platystachys, and S. divitifiorus would reduce the number given by
Rogers to 14; but to these are now added S. Harriganae and S. tricalliatus, the latter
being described here as a species for the first time.

A Key to the Species.
1. Sepals and petals from broad-lanceolate to almost orbicular.

2. Stems often rather long, scrambling on rocks, with somewhat flaccid leaves. Flowers
WANS Cre yoialke Wala, UMA lNOUOMWS soskogoasonsconasuocsoocdc S. Fitzgeraldii. 1.
2’. Stems always short.

a)

3. Robust plant; leaves in the typical form large, rigid, deeply channelled. Flowers white

With'-\ Maroon! sCeNbres <0: Sivshs cuscecus loss ieee ened. Gee ene ue ern cise pec Ro eieee S. Hartmannti. 2.
3’. Plants less robust than the last. Leaves shallowly channelled, often more or less
faleate.

4. Flowers white or pink.

5. Leaves moderately broad, light green, faleate. Flowers up to 3 em. in diameter
white with orange and reddish-purple markings on the lateral lobes of the
labellum and often! onthe labellar sac... ose ee ciee eiaeie S. faleatus. 3.

5’. Leaves linear, channelled. Small plants with small flowers.

6. Leaves more or less spotted, dull green.

7. Flowers bright pink, almost campanulate .................. SS) Geciliaes Al2:
7’. Flowers pink or white, expanding widely .................... Se Tahoe 183,
6’. Leaves not spotted, light green. Flowers white; labellum with three conspicuous
CELE ** 1:5, EE et Rr ae ie eta eR AO STR eee EC Ee Eee te ete ane S. tricalliatus. 14.

8. Lateral lobes of labellum densely pubescent or hairy ...... S. eriochilus. 15.

8’. Lateral lobes of labellum glabrous. Flowers very diminutive ..............
RE IORI SOON OCC IRR OZONE ORAMERG Caio cAO O Orcr ce chi S. minutiflos. 16.
4’. Flowers neither pink nor white.

9; Blowers. Brick Ted: Scenic koncss crept nenoueheac ekerel oer ea ld rk eee) vate S. Bancroftii. 11.
9’. Flowers never red.
10. Elowers cream, blotched with dull purple ....5..5.........% S. Weinthalii. 4.
1:02,- MIOWeERS VeEllOW? saws exeseseie oot. holla Peale ohcie lewells_ Sao Re aan meen te ce Ce tae S. Longmanii. 10.
1’. Sepals and petals narrow for their basal third, then expanding rather broadly.
11. Sepals dilated into a rhomb above the narrow basal portion .......... S. dilatatus. 9.

11’. Sepals expanding to lanceolate above the narrow basal portion.
12. Lateral lobes of the labellum spathulate.

13) Dabellar sac with bright purple “mankines eee kee eee ee S. spathulatus. 6

1e4n wabellarvsac) pale .2reeny ayy eeeie 1s eee end eer nee eer fs (oto S. Harriganae. 8.
12’. Lateral lobes of the labellum oblong to ovate.

{4.6 Mlowers#old-gold, or (OllVE=STECME ei kereu. ies eieledstneee nents eres ieae sv rel aaets S. olivaceus. 7.

14’. Flowers brownish-green; labellum white with purple and yellow markings ........
Lic: Re Sg a ES PE A AR TEETER he ya Cs ay Lis Sle leie pe os) | yee (CRUSTAL UIS ene

BY H. M. R. RUPP. 51

1. S. Fitegeraldii F. Muell., Fragm., vii, 1870, 115; Benth., Fl. Austr., vi, 1873, 293;
R. D. Fitzg., Austr. Orch., i, 3, 1877; Austr. Orch. Rev., i, No. 1, 1936, 10, and ii, No. 3,
1937, 9-10; Rupp, Orch. N.S.W., 19438, 136.

This is the largest and, in the opinion of many, the most beautiful, of all Australian
species of the genus. Mueller (l.c.) hints at the possibility of its proving to be a variety
of S. falcatus R.Br.; while Bentham, followed by F. M. Bailey, states, “Stem, foliage, and
general aspect of S. falcatus’’. It is difficult for anyone well acquainted with both plants
to perceive this supposed resemblance. Fitzgerald makes no allusion to it. S. falcatus
is a true epiphyte, always found on trees, short-stemmed, with light green leaves and
short racemes. S. Fitegeraldii is an extensively spreading, branching plant which
scrambles along cliffs and on rock-ledges in deep ravines; the leaves are dark green,
and the racemes are on long, stout peduncles. Bentham probably lacked a sufficiency
of good material, and of course he had never seen the plant growing; but it is curious
that Bailey, who must surely have seen it in southern Queensland, should have repeated
Bentham’s remark without comment.

The precise colouring of the flowers varies a good deal, but typically it may be
described as white with crimson or maroon blotches. Plants are occasionally found
with pure white flowers. The variety aqemulus (These PRrocrEpINGs, 69, 1944, p. 73)
was found by the present writer some 20 miles from the site of Fitzgerald’s original
discovery at the Naroo Falls on the Bellinger River, N.S.W. Its flowers are light
erimson with deep maroon blotches.

Although the species is not uncommon in gorges of the eastern slopes of the
Dividing Range in northern New South Wales and southern Queensland, its distribution
appears to be confined to these areas. There are’no definite records of its occurrence
south of the Hastings Valley in New South Wales, nor has it been found north of the
Brisbane River in Queensland. It responds fairly well to cultivation, provided the
requisite humidity and temperature are available; it does best in moderate shade, with
a rock to scramble on.

2. S. Hartmannii F. Muell., Fragm., viii, 1870, 248; F. M. Bailey, in Q’land Fl., v,
1902, 1551, plate Ixvii; Austr. Orch. Rev., i, No. 1, frontispiece; Rupp, Orch. N.S.W.,
1948, 137.

Although obviously closely related to the preceding species, this species-is quite
distinct, with a different habit. It grows on both rocks and trees, and likes a bright,
sunny situation. It is a more variable plant than S. Fitzgeraldii. In the typical form
the stem is erect, but short; the leaves are large, rigid, and deeply channelled. Bailey,
however, identified Fitzgerald’s S. rubicentrum (Austr. Orch., ii, 1, 1884) with
S. Hartmannii, and his view is generally accepted; but there is some difference between
the two forms, rubicentrum being less erect, with smaller leaves and flowers. There is
some mystery about the habitat of the plant figured by Fitzgerald; see footnote to the
description of S. Hartmannii in ‘Orch. N.S.W., Lc. Diligent search by several capable
collectors in the Cairns district during the past twenty years has failed to discover any
orchid resembling Fitzgzerald’s plate, and it seems probable that the plant he received
from E. Ramsay had been cultivated at Cairns. The experience of botanists and
collectors suggests that the habitat of S. Hartmannii is even more restricted than that
of S. Fitegeraldii, and is practically confined to the Macpherson Range in southern
Queensland, and one or two localities just on the New South Wales side of the border.

The flowers of S. Hartmannii vary a good deal in dimensions; sometimes they are
as large as those of S. Fitegeraldii, but more commonly they are smaller. They are in
a more compact raceme, borne on a long, stout peduncle. Usually they are white with
maroon spots at the base of the perianth; but pure white flowers are sometimes found.
The species is more easily cultivated than S. Fitegeraldii.

Synonymy.—Sarcochilus rubicentrum Fitzg., Austr. Orch., ii, 1, 1884.
3. 8. falcatus R.Br., Prodr., 1810, 332; Benth., Fl. Austr., vi, 1873, 293; Fitzg., Austr.

Orch. i, 5, 1879; Austr. Orch. Rev., i, 3, 1936, frontispiece; Rupp, Orch. N.S.W., 1943, 132;
Barrett and Nicholls, “Gems of the Bush’, 1934, 9.

on
bo

THE AUSTRALIAN SPECIES OF SARCOCHILUS (ORCHIDACEAE),

Generally known as the “Orange Blossom Orchid”, this attractive little species is a
familiar wildflower in many parts of the coastal belt of eastern Australia, and it ascends
to a considerable altitude on the Dividing Range and its spurs. As might be expected
in view of its extensive distribution, it varies considerably from the typical form;
but the variations rarely tend to obscure its identity. R. D. Fitzgerald, indeed, described
and figured one form (Austr. Orch., l.c.) as a distinct species (S. montanus); but
subsequently, in Moore and Betche’s “Handbook of the Flora of N.S.W.”, he reduced this
to a variety of S. falcatus. In the foothills of many mountain ranges, intermediates
between it and the type-form are so numerous that it is hardly desirable to perpetuate
the varietal status. One of the most distinctive forms known to me is found growing
on the Negrohead Beech (Nothofagus Moorei) near Barrington Tops, about sixty miles
north of Newcastle, N.S.W. This plant is more robust than usual, and the flowers are
of a rich cream colour, with a tuberose perfume. Another distinct form occurs on the
Atherton Tableland in North Queensland, more than a thousand miles away. In this
case the plant is very small, but the flowers are well above average size, heavily stained
with deep purple markings, and with a distinctive perfume.

As indicated above, the range of NS. falcatws is very extensive. From the Cann River
in eastern Victoria to the Atherton Tableland in North Queensland, the distance is
approximately 1,500 miles, but in and near the rain forests of many areas along this
great stretch of country, the “Orange Blossom” may be found.

Synonymy.—Thrixspermum falcatum (R.Br.) Rehb. f., Beitr. 46, and Xen. Orch.,
i, 122%

4. S. Weinthalii F. M. Bail., Q’land Agr. Journ., xiii, 1903, 346, and xxviii, 1912, 448;
also in Compr. Cat. Q’land Pl., p. 534; Rupp, Orch. N.S.W., 1943, 134.

This plant when not in bloom is hardly distinguishable from smaller forms of
S. falcatus. The flowers, numbering from 3 to about 12, are cream or white, blotched
with dull reddish-purple, and the lateral lobes of the labellum are very narrow, with
purple spots. The species seems to be very rare, being known only in one or two
localities in southern Queensland, and near Kyogle in the far north of New South
Wales.

5. S. australis (Lindl.) Rehb. f., Waip. Ann., vi, 501, and Xen. Orch., ii, 122; Rupp,
Orch. N.S.W., 1943, 1384, plate xxiii; Fitzg., Austr. Orch., i, 3, 1877 (as S. parviflorus).

This very attractive little orchid was for many years better known as S. parviflorus
Lindl. But Lindley had originally named it Gunnia australis, and the priority of the
latter specific name is beyond question. As it happens, it is a far more suitable name
than parviflorus; for while there are several species with smaller flowers, no other
extends so far south, this being one of the only two epiphytic orchids occurring in
Tasmania, where it was discovered by Ronald Gunn more than a century ago.

The species is recorded by F. M. Bailey for southern Queensland, but the record is
very vague, and no Queensland specimens are known at present. In my opinion,
S. australis does not extend farther north than the rain forests near Gosford in New
South Wales. I believe that the Queensland records should be applied to the allied
species S. spathulatus Rogers, which resembles S. australis both in habit and in super-
ficial appearance. It probably passed for the latter for many years, for it was not till
1927 that the late Dr. Rogers described and named SNS. spathulatus from specimens found
on Tamborine Mountain in southern Queensland. For more than twenty years I have
tried to trace S. australis at least as far north as the Hunter Valley in New South
Wales; but Gosford remains the nearest approach (50 miles south of the Hunter).
Unless direct evidence of the occurrence of this species in Queensland is forthcoming,
I think it should be deleted from the orchid flora of that State.

It is one of the daintiest and most attractive of the smaller Australian orchids.
Occasionally stems are elongated to as much as 20 cm.; but more usually they are quite
short. Racemes in well-developed plants are often numerous, each bearing from 5 to
about 14 very fragrant flowers, brownish-green with a white labellum, variably splashed
with purple, red, or yellow. Column relatively longer than in other species. The plant
is most frequently found growing on twigs of small trees and shrubs, in moist gullies.

BY H. M. R. RUPP. 53

In Tasmania it is confined to rain forests in the north and west of the State. In
Victoria, Mueller recorded it from Apollo Bay; but this is the only known record .
west of Port Phillip. It is not uncommon from the Dandenong Ranges eastward. In
New South Wales there are records from Braidwood, Picton, Campbelltown, the Blue
Mountains, National Park, northern arms of Port Jackson, Hawkesbury River, and
Gosford.

Synonymy—Gunnia australis Lindl., Bot. Reg., 1834, sub t. 1699; Hook. f., FI.
Tasm., ii, 1860, 338, t. 128; Sarcochilus parviflorus Lindl., Bot. Reg., 1838, Mise. 34;
Benth., Fl. Austr., vi, 1873, 294; Fitzgerald, Austr. Orch., i, 3, 1877; Sarcochilus Gunnii
F. Muell., Fragm., i, 1859, 90; Sarcochilus Barklyanus F. Muell., l.c., 89; Thrixspermum
_ parviflorum Rehb. f., Xen. Orch., ii, 122; Thrixspermum australe Rehb. f., l.c.

6. S. spathulatus Rogers in Trans. Roy. Soc. S. Austr., li, 1927, 1; Rupp, Orch.
N.S.W., 1943, 1386; Barrett and Nicholls, “Gems of the Bush”, 1934, 8.

I cannot agree at all with Murray Cox (Cultural Table of Orchidaceous Plants,
1946, p. 274) that this species in any way resembles S. Hillii; except perhaps in its
habit of growing on twigs. It does, however, closely resemble S. australis, and could
easily be mistaken for that species. The plant is, I think, consistently smaller, and
the stem is never elongated. The flowers are quite as large as those of S. australis,
and are somewhat similarly coloured, but they are never numerous. The specific name
was chosen in allusion to the conspicuously spathulate lateral lobes of the labellum.

The species was discovered on Tamborine Mountain, Queensland, in 1925, by Mrs.
H. Curtis; just a week later I found it in the southern foothills of Barrington Tops in
New South Wales. Between these two localities it is not uncommon in coastal rain
forests; it has not been seen south of the Hunter Valley, or north of the Brisbane River.

7. S. olivaceus Lindl., Bot. Reg., 1839, Misc. 32; Benth., FI. Austr., vi, 1873, 293;
Fitzg., Austr. Orch., i, 5, 1879; Rupp, Orch. N.S.W., 1943, 134.

Bentham remarks, ‘Sepals and petals of a duil pale purple or yellowish-brown”’.
This is a very unsatisfactory description; there is no purple at all. It is difficult to
define the colour; Fitzgerald’s plate does not do it justice. I think “old gold” is as
near as one can get to it—an uncommon shade of golden green. The flowers are
deliciously perfumed, and are more numerous than Bentham supposed—I have counted
11 on one raceme. When not in bloom this species is sometimes mistaken for
S. falcatus; but the leaves are thinner, and of a darker green. Bentham united Mueller’s
S. dilatatus with S. olivaceus, but there are ample grounds for specific distinction.
W. H. Nicholls has described a distinct form from North Queensland as var. borealis.
(N. Q’land Naturalist, Dec., 1939.) It has darker flowers, blotched with deep red-brown.

The species extends from at least as far south as the Shoalhaven River in New
South Wales, northward into the tropics of Queensland.

Synonymy.—Thrixspermum olivaceum Rehb. f., Xen. Orch., ii, 122. |

8. S. Harriganae Rupp in These Proc., |xiii, 1938, 128.

Unfortunately the type specimen of this species was inadvertently destroyed, and at
present no others are available. It is a rare plant, apparently confined to part of the
Dorrigo highlands in northern New South Wales. The plant itself somewhat resembles
a small S. falcatus, but the leaves are usually marked with irregular rows of dark dots.
The dull-green flowers almost suggest a natural hybrid S. olivaceus x S. spathulatus;
their general conformation agrees with the former, while the lateral lobes of the
labellum are definitely spathulate. The dorsal sepal is very broad.

9. S. dilatatus F. Muell., Fragm., i, 1859, 191; Rogers, Trans. Roy. Soc. S. Austr.,
li, 1927, 291.

As stated above, Bentham united this little species with S. olivaceus. Rogers, how-
ever, makes it clear that Mueller was justified in giving it specific rank. It never
attains the dimensions often reached by S. olivaceus, and the flowers (1 to 5) are
correspondingly smaller. The sepals and petals are pale green basally, shading to deep
brown at their distal ends. In the sepals the narrow claw forming the basal half
is dilated terminally into a rhomb; in the petals it merely becomes spathulate. The
whitish labellum is unlike that of S. olivaceus.

54 THE AUSTRALIAN SPECIES OF SARCOCHILUS (ORCHIDACEAE),

S. dilatatus has not yet been found outside southern Queensland. Mr. Trevor Hunt,
of Ipswich, writes, “I have found this species always in the thick, semi-dry vine scrubs
of the hilltops, at elevations up to 1,000 ft. These scrubs are in the nature of residuals
of the once dense scrubs covering hilly parts of the coast plain’. Recently I received
numerous specimens from Mr. W. W. Abell, of Durong State School, near Tingoora, on
the Kingaroy line.

10. S. Longmanii F. M. Bail., Q’land Agr. Journ., xxiii, 1909, 261; ibid., xxviii, 1912,
449; Compr. Cat. Q’land PIl., p. 532.

Bailey’s description of this species calls for no comment. The plant closely
resembles S. Weinthalii, but the flowers are a little smaller, and of a light yellow
colour. It is a rare species, being recorded only on the slopes of the main Dividing
Range in the Toowoomba district of southern Queensland.

11. S. Bancroftii F. M. Bail., Q’land Agr. Journ., xxviii, 1912, 450; Compr. Cat.
Q’land PIl., p. 532.

Nothing is known of this species at present beyond Bailey’s description and
figure. The latter shows a small plant similar to S. Weinthalii and S. Longmanii. The
only record is that of Bancroft from Hidsvold, Queensland, in 1912. It is most desirable
that efforts should be made to re-discover this orchid, the flowers of which are
described as “brick-red’”. In this respect it must present a striking contrast to all the
other species.

12. S. Ceciliae F. Muell., Fragm., v, 1865, 42, t. 42; Benth., Fl. Austr., vi, 1873,
294; Barrett and Nicholls, “Gems of the Bush’’, 1934, 8; Rupp, Orch. N.S.W., 1943, 137.

In Mueller’s plate the enlargements of an individual flower are excellent; but the
figure of the plant itself (natural size) with two racemes and several leaves, shows
the latter far more acuminate than is usual, and the flowers expanded too widely.
The photograph by W. H. Nicholls depicts the flowers more naturally; they are almost
campanulate, like inverted bells. (‘‘Fairy Bells” is a popular name for this species.)
The flowers are typically of a bright pink colour; a white-flowering form (var. albus
Hunt) has been found in south-east Queensland. S. Ceciliae is a very small plant, but
when growing, as it often does, on rock-ledges, it usually occurs in dense masses, the
numerous flowers being quite conspicuous. If growing on trees the plants are generally
solitary. The species is found in coastal forests and rocky gullies from the Macleay
River in New South Wales northward into the Queensland tropics, occasionally
extending to the ravines of the tablelands.

Synonymy.—Thrixzspermum Ceciliae (F. Muell.) Rehb. f., Beitr., 71.

13. S. Hillii F. Muell., Fragm., ii, 1860, 94; ibid., vii, 1870, 98; Benth., Fl. Austr.,
1873, 295; Fitzg., Austr. Orech., i, 3, 1877; Barrett and Nicholls, ‘Gems of the Bush”
(1934), 8; Rupp, Orch. N.S.W., 1943, 138.

When not in flower, this tiny plant could be mistaken for a small SNS. Ceciliae; but
it is strictly an epiphyte, and has not been recorded on rocks. Plants, however, are
sometimes crowded in great numbers on the trunks of small trees such as Backhousia
myrtifolia. The flowers expand more widely than those of S. Ceciliae; they are white
' or pink, and are very fragrant. The species has a fairly extensive range, from the
south coast of New South Wales at least as far north as Rockhampton in Queensland.

Synonymy—Dendrobium Hillii r. Muell., Fragm., i, 1859, 88; Thrixspermum Hillii
(EF. Muell.) Rehb. f., Beitr., 71.

14. S. tricalliatus (Rupp) Rupp, n. sp. (Text-fig. 1).

Planta parva caule brevissimo. Folia pallida, immaculata, linearia, canaliculata,
usque ad 7 em. longa. Flores 2-5, albi, inodorati. Sepala petalaque latissime lanceo-
lata, patentia, circa 5 mm. longa. Labelli lobi fere aequales, intus rubri; lobus inter-
medius marginibus aurantiacis, paulum pubescens. Discus callis clavatis conspicuis 3,
callus intermedius magnus, anceps. Columna non dentata.

A small plant with a very short stem. Leaves pale green, unspotted, linear,
channelled, up to 7 em. long. Flowers white, 2 to 5, without any perfume. Sepals
and petals very broadly lanceolate, about 5 mm. long, spreading. Labellum trilobate,

BY H. M. R. RUPP. 55D

the three lobes approximately equal; midlobe with orange margins, only slightly
pubescent. Disc with three rather stout clubbed calli, the middle one larger than
the others, and double-headed. Column devoid of teeth.

Specimens sent to me in 1935 from Mount Dryander, by Mr. K. MacPherson of
Proserpine, North Queensland, flowered the next year in February. At the time I
thought the plant could be regarded as a northern variety of S. Aillii, and I
provisionally named it S. Hillii var. tricalliatus (N.Q. Naturalist, May, 1936, 31). But
for some years past I have felt that the differences are sufficient to warrant the raising
of the Mount Dryander plant to specific rank. It has not been found in any other
locality. ‘

Text-fig. 1.

Sarcochilus tricalliatus, n. sp. Nat. size. A, the 3 calli on the dise of the labellum
(enlarged). :

15. 8. eriochilus R. D. Fitzg., Journ. Bot., xxix, 1891, 153; Rupp, Orch. N.S.W.,
1943, 137.

I regret that I have nothing to add to the remarks made on this species in the
work last cited. Apparently it is extremely rare. As the Tweed River, on the New
South Wales—Queensland border, is the only locality where it has been recorded, it is
possible that the clearing of the forests along that stream has exterminated it.

16. S. minutifilos F. M. Bail., Compr. Cat. Q’land PIl., pp. 845-847.

This species has been recorded only from Hidsvold in Queensland. There are
specimens in the Brisbane and Sydney Herbaria, but beyond these little is known of it.
Following is Bailey’s description: “On branchlets of shrubs and trees. Roots very
long and slender, white and more or less curled. Stem very short. Leaves several,
slender, 2—4 inches long and about 2 lines broad, sometimes dotted. Racemes mostly
very slender, from 2 to 6 inches long, sometimes forked, bearing throughout their
whole length, or nearly so, very numerous minute flowers. Flowers on slender
pedicels of about two lines, nearly globular from the incurving of the sepals and
petals, of a greenish white sometimes tinged with pink, and less than two lines in

56 THE AUSTRALIAN SPECIES OF SARCOCHILUS (ORCHIDACEAE).

diameter. Bracts minute. Sepals somewhat longer than the petals. Labellum small,
lateral lobes purplish, blunt, ovate-oblong, middle lobe stalked, for the greater part
composed of a globular mass of glandular. white hairs. Discal calli orange yellow.
Column short, anther lid stained with purple. Capsules narrow, straight, 2-23 in.
long.”’

Excluded Species.

S. Armitii F. Muell., Fragm., ix, 49 (Cleisostoma Armitii F. Muell., l.c.).

The rightful position of this plant has not been definitely settled. The type in
Mueller’s herbarium is too fragmentary to be of very much use, and no other specimens
have been clearly recognized as such. My own opinion now is that Saccolabium orbicu-
lare (Rupp) Rupp, in Vict. Nat., 67, 1941, 220, is really identical with Mueller’s plant,
and should in future be known as Saccolabium Armitii.

S. Baileyi F. Muell., Herb., is Taeniophyllum Muelleri Lindl., Herb., Benth., FI.
Austr., vi, 291.

S. Beckleri (F. Muell. ex Benth.) F. Muell., Cens. Austr. Pl., 1882, 111, is Sarcan-
thus Beckleri (F. Muell. ex Benth.) Rupp, Vict. Nat., l.c.

S. calcaratus (F. Muell.) F. Muell., Fragm., ii, 192, is Sarcanthus tridentatus
(Lindl.) Rupp, Vict. Nat., l.c., 218.

S. divitifiorus F. Muell. ex Benth., Fl. Austr., vi, 292, is Rhinerrhiza divitifiora
(F. Muell.) Rupp, Vict. Nat., Vol. 67, 1951, 206.

S. Newportii F. M. Bail., Q’land Fl. v, 2014, is Bulbophyllum Newportii (F. M.
Bail.) Rolfe, Orch. Rev. (London), xvii, 94.

S. phyllorhizus F. Muell., Fragm., v, 201, is Chiloschista phyllorhiza (F. Muell.)
Schlitr., Engl. Bot. Jahrb., lvi, 492.

S. platystachys F. M. Bail., 1st Suppl. Syn. Q’land FIl., 56, is Thrixzspermum
platystachys Schltr., ‘“Orchis’”, 1911, v. 155.

S. tridentatus (Lindl.) Rehb. f., Walp. Ann., vii, 98, is Sarcanthus tridentatus
(see S. calcaratus above).

I desire to acknowledge the assistance of Mr. Trevor E. Hunt, of Ipswich, Queens-
land, and Mr. H. K. C. Mair, B.Sec.,; of the New South Wales National Herbarium, who
have both read the MSS. of this review and have offered useful suggestions.

AUSTRALIAN RUST STUDIES.

VIII. PUCCINIA GRAMINIS LOLII, AN UNDESCRIBED RUST OF LOLIUM SPP.
AND OTHER GRASSES IN AUSTRALIA.

By W. L. WATERHOUSE,
The University of Sydney.
[Read 27th June, 1951.]

Synopsis.

A stem rust which attacks Loliwm spp., Dactylis glomerata Ll. and about 30 other grasses
is described. Aecidial, uredospore and teleutospore stages show that it is a type of Puccinia
graminis Pers. Spore morphology and cultural studies make it clear that it is not one of the
known subspecies of the cereal stem rusts, nor is it one of the described subspecies which
attack grasses.

Its host range has been studied and its distribution determined in all the States of
Australia and in New Zealand. The occurrence of physiologic races of the rust has been shown
by differential reactions of plants of Loliwm perenne L., L. rigidum Gaud., and Arrhenatherum
elatius (lu.) J. et C. Presl. In many grasses there are wide differences between the resistance
and susceptibility of individuals within the species.

The name Puccinia graminis lolit is proposed for the rust.

INTRODUCTION.

Stem rust of cereals and grasses is caused by the highly specialized fungus Puccinia
graminis Pers. The following subspecies (or varieties) are generally recognized: (i) a
group of three which attack cereals, viz., P. graminis tritici E. and H., P. graminis
avenae H. & H., and P. graminis secalis BE. and H.; and (ii) a group of subspecies which
attack grasses. Three of these have been studied intensively in U.S.A., viz., P. graminis
poae HK. and H., P. graminis agrostidis BE., and P. graminis phlei-pratensis (E. and H.)
Stak. and Piem. In addition, European workers have described four others, viz.,
P. graminis airae BE. and H., P. graminis calamagrostis Jacz., P. graminis aperae Jacz.,
and P. graminis arrhenatheri Jacz. Little is known of these.

McAlpine (1906) recorded P. graminis on cereals and a number of grasses, but made
no inoculation studies or determinations of the subspecies present. More recently
Australian investigations of P. graminis tritici and P. graminis avenae have been
carried out (Waterhouse, 1929). It has been shown that barley and rye are attacked
by the former: P. graminis secalis is not present in Australia. A number of grasses
were also found to be common hosts for one or other of these cereal rusts. But many
other stem rusts occur on grasses and very little is known about them.

Of these a common type which is specially notable for its attack of Lolium spp. has
been under observation since 1918. It quickly became apparent that it was not one of
the cereal rusts: no lesions were produced on wheat, very occasional resistant reactions
were shown on rye, tiny reactions occurred on a few varieties of oats at very favourable
temperatures, and 260 barley varieties showed sharp resistance with the exception of
two varieties in which a ‘2+’ reaction was given. In no case were the reactions
comparable with those caused by P. graminis tritici or P. graminis avenae. Detailed
studies were therefore undertaken to determine its identity, host range and distribution.

MorPHOLOGICAL STUDIES.

Large uredosori occur on stems and leaves, rupture of the epidermis being a notice-
able feature. Lesions on glumes and rachises are not uncommon. Those on the stems
frequently give place to black teleutosori late in the season, and sometimes these also
occur on the leaves. The aecidial stage has not been found in nature but has been
produced in the plant house on Berberis vulgaris L. from teleutospores on Lolium
perenne L. that had been exposed to winter conditions on the tablelands at Glen Innes,

58 AUSTRALIAN RUST STUDIES. VIII,

New South Wales. Inoculations with these aecidiospores produced uredosori on JL.
perenne and Dactylis glomerata L. but on none of the cereals. All spore forms show
characteristics which agree with those described for P. graminis. It has been shown
(Levine, 1923, and Waterhouse, 1930) that statistical examinations of the spore
morphology may reveal significant differences between subspecies.

Measurements were made of 200 aecidiospores and 200 uredospores lightly shaken
from leaves of the susceptible host, and of 200 teleutospores obtained by scraping sori
on the stems. The standard methods previously described (Waterhouse, 1930) were
used.

The results are set out in Table 1, in which the name P. graminis lolii is used for
the new rust, and the measurements are grouped with those determined for the
Australian subspecies P. graminis tritici and P. graminis avendae (Waterhouse, 1930),
and the U.S.A. determinations for the remaining grass subspecies and P. graminis
secalis (Levine, 1923).

TABLE 1.

Comparison of Measurements of Spore Forms of Several Subspecies of Puccinia graminis.

| Length in uw. | Width in w.

Rust Subspecies. |
Standard | | Standard |
| Mean. Deviation. Range. | Mean. Deviation. | Range.
| | |
Aecidiospores.
P. graminis lolii a ar 16-92 0-47 12-5-23-2 16-41 0-50 11-6-19°-3
P. graminis avenae r.1 ay 21-18 2-29 14-9-27-9 16-53 1-91 11-6-20-46
P. graminis tritici 1.46 ai 19-08 1-95 14-9-24-2 16°63 1:54 | 14-9-20-46
P. graminis secalis .. ae 17-10 2-0 12-0-22-0 13-46 1:17 11-0-16-0
P. graminis agrostidis a 16-46 1-93 | 12-0-22-0 | 12-98 0-97 | 11-0-15-0
P. graminis poae as #3: 15-07 1-35 12-0-18-0 13-23 0-97 =| 11-0-15-0
Uredospores.
P. graminis lolii as aus 27-61 0:56 22. -3-31-6 18-08 0-48 14-9-24-2
P. graminis avenae (all races) 29-99 3°14 20-5-39-1 18°33 1-59 13-1-22-3
P. graminis tritict (all races) 31-04 3°70 20-5-44:6 18-25 1-88 11 -2-22-3
P. graminis secalis .. 34 27-14 2-91 21-0-36-0 17°19 1-26 14-0-21-0
P. graminis agrostidis ne PIB 2-61 15-0-30-0 15-68 PotD 13-0-18-0
P. graminis phlei-pratensis .. 23-95 2°41 18-0-30-0 16°88 1-32 13 -0-20-0
P. graminis poae Bi AS 18-64 1:51 15-0-23-0 15:78 1-06 13-0-18-0
| | |
Teleutospores. b
sas E am Te .
P. graminis lolii as .. | 48°75 1:78 32-8-63 -7 16-93 | 0-56 11-6-21-2
I’. graminis avenae 1.1 Sinan| 46-95 6-54 29-8-67-0 15-27 1°85 } 11-2-20°5
P. graminis tritici (3 races) 44-32 7-40 29-8-74-4 15:90 | 2°20 11-2-22-3
P. graminis secalis .. ot 47-35 6-65 35-0-65-0 14-77 1-83 10-0-19-0
I. graminis agrostidis 3d 40-30 5:87 25-0-55-0 14-64 172 10-0-19-0
1’. graminis phlei-pratensis .. | 41-30 4-78 30:°0-55-0 15-63 1:46 12-0-19-0
P. graminis poae bi ae 36-90 9-13 20-0-60-0 15-52 2:07 11-0-21-0
|

The aecidiospores resemble those of P. graminis poae in being sub-spherical, but
are much larger. In making the measurements, the uniformity in outline was striking,
together with the tendency for the spores to remain in chains.

The uredospores are quite unlike those of P. graminis pode, resembling most closely
those of P. graminis secalis. They are much larger than those of any of the other grass
rusts.

BY W. L. WATERHOUSE. 59

The teleutospores agree in general with those of P. graminis avende and again are
quite unlike those of any of the grass rusts, being much larger. In the course of
making the measurements, teleutospores having either a single cell or a series of three
cells were noted. —

On these morphological grounds the rye grass rust is different from any of the
described subspecies.

CULTURAL STUDIES.

The recognized subspecies on grasses gave reactions in side-by-side comparisons
as shown in Table 2, in which the new rust is again listed under the name P. graminis
Jolii. The designations of the reactions are those proposed by Stakman and Levine
(1922).

TABLE 2.
Comparison of Reactions of Grass Subspecies of P. graminis on Grass Hosts.

|
| Grass Hosts and Their Reactions.

Rust.
Poa Agrostis Phlewm Lolium
compressa. alba. pratense. perenne.
|
P. graminis poae 4 0 | : ;
P. graminis agrostidis 0 | 4 ; ;
P. graminis phlei-pratensis 0 | (0) 4 0
P. graminis lolii | 0 0 : 4
|

On the basis of these grass reactions, P. graminis lolii is different from the recog-
nized subspecies.

Turning next to the subspecies on cereals, inoculation experiments gave the reactions
described hereunder:

P. graminis tritici (10 physiologic races) gave no attack on Lolium spp., and
P. graminis lolii produced no lesions on any of the very many varieties of all the
Triticum spp. tested. ;

P. graminis avenae (three races) attacked some individual plants of Loliwm spp.,
but with a low frequency: the usual reaction was a tiny fleck. P. graminis lolii gave
flecks on more than 100 varieties of oats: in the case of a few varieties like “Victory”
it gives a “2—” reaction at very favourable temperatures.

P. graminis secalis does not occur in Australia, but side-by-side comparisons using
three isolates of this subspecies which comprised two physiologic races showed that it.
is quite different from P. graminis lolii. Inoculations of 120 varieties of barley with
the P. graminis secalis cultures resulted in 17 of them giving large sémi-susceptible
reactions (‘‘2+’’), 41 gave semi-resistant (‘2’) reactions, whilst the remainder were
resistant (“;”) and (“1”). With P. graminis lolii, all gave resistant (“;” and ‘1’’)
reactions. An extensive series of inbred ryes which have been selfed for 17 years and
which were fully susceptible to the two races of P. graminis secalis gave resistant
reactions when inoculated with P. graminis lolii.

The cultural comparisons with the six subspecies of P. graminis. indicate that
P. graninis lolii must be regarded as a different subspecies.

DISTRIBUTION OF THE RUST.

Serious damage is.done to rye grass and cocksfoot when conditions are favourable
for rust development, and other grasses of lesser importance are also damaged. Stems
and leaves are attacked and not infrequently sori occur on the rachis and glumes.

In the course of the investigations the rust was isolated with the stated frequency
from the grasses set out in Table 3. Many of the specimens were sent in for study
by co-operators in scattered areas, whilst others were collected as opportunities offered.
But it has not been possible to make an exhaustive survey of its occurrence, which may
well be larger than indicated herein.

60 AUSTRALIAN RUST STUDIES. VIII,

The practice has been to test each grass rust isolate on the four cereals and Lolium
perenne: whenever an infection of the latter is shown, it is used to inoculate the
original grass host in order to check its pathogenicity.

It will be noted that a number of these grasses have already been recorded as
natural hosts for P. graminis avenae. This rust was found together with P. graminis
lolii many times on a number of the grasses listed above. Determinations showed
that it was either race 2 (not separated from race 1) or race 7 (not separated from

TABLE 3.
Naturally-occurring Grass Hosts of P. graminis lolii with the Number of Isolates Examined from Each.

Number of
Tribe. Grass. | Isolates.

Phalarideae .. .. | Phalaris tuberosa L.
P. minor Retz Me ve as Pec Ma
P. paradoxa UL. eae ne a Ae a rn |
P. aquatica L. Re Ap st Ne a ral
Agrostideae .. .. | Agrostis avenacea Gmel. 2 eos
Echinopogon caespitosus C. E. Hubbard
Dichelachne spp. ;

Amphibromus Neesii Steud.

Alopecurus pratensis L. ;
Gastridium ventricosum (Gonan) Schinz Si Thal.
Stipa variabilis Hughes

Lagurus ovatus L.

Aveneae My .. | Holeus lanatus L.

Aira cupaniana Guss.

Trisetum flavescens (L.) Beauv.

Trisetum sp.

Arrhenatherum lati! (L.) J. et C. prea
Festuceae .. .. | Festuca elatior var. pratensis (Huds.) A. Gray ..
F. elatior var. arundinacea (Schreb.) Wimm.
Scleropoa rigida (L.) Griseb. :

Vulpia bromoides (L.) 8S. F. Gray

V. Myuros (L.) Gmel.

Dactylis glomerata L.

Spartina alterniflora Loisel

Lamarckia aurea (L.) Moench.

Koeleria phleoides (Vill.) Pers.

Poa annua L. Pa

P. iridifolia Hauman.

Bromus Gussonii Parl.

B. Benekeni G. Beck.

| B. breviaristatus Buck). 2 ye Rs ese
Hordeae at .. | Lolium perenne L. .. ae Ss BY i tual 9
L. multiflorum Lam. é a a ae

LD. loliaceum (Bary et Chaub.) iy SA hare eel 3
| L. rigidum Gaud. .. ae ae Le nes 5 Stal
L. temulentum L. ; a 2 ae a aeel
| Hordeum leporinum Link. .. a ees at is |
|

Re

DWRrRrRrRPUrF RP RP NNN DOrR KE PO

e
bo be

>

~I
OSe Re Nr WOT OWwWwre

rs

race 3), and in one instance Lamarckia aurea (L.) Moench was infected by both these
oat rust races in addition to P. graminis lolii. The details of these occurrences are set
out in Table 4.

Apart from the joint occurrence on the one host of these two stem rusts, there were
numerous cases in which the grass host of P. graminis lolii was also attacked by its
leaf rust: Lolium spp. carrying P. coronata lolii E., and Bromus spp. attacked by
P. bromina E. were notable examples.

In addition to the natural occurrence of the rust on grass hosts, studies were made
of the plant house behaviour of many other grasses when inoculated under controlled
conditions. These have shown that there are susceptible members within the following
species, although in certain cases resistant members predominate.

BY W. L. WATERHOUSE. 61

Lolium remotum Schrank, Briza maxima L., Bromus racemosus L., B. hordeaceus L.,
B. madritensis L., Ehrharta longiflora Sm., Phleum pratense L., Aegilops ovata L.,
Hlymus paboanus Claus., H. canadensis L., E. junceus Fisch., Arrhenatherum elatius
(L.) J. et C. Presl., Hordeum marinum Huds., Poa annua L.

These plant house tests brought to light two important happenings, viz., that
physiologic specialization occurs within P. graminis lolii, and that there are striking
differences in the resistance and susceptibility of individuals within a species when
inoculated with the same rust isolate.

TABLE 4.

Frequency Occurrence of the Two Races of P. graminis avenae in Addition to
P. graminis lolii on Various Grasses.

+

Frequency of Occurrence of
Oat Stem Rust.
Grass Host.

|
Race 2. | Race 7.

Lolium perenne

L. multiflorum

LL. loliaceum

L. rigidum

Dactylis glomerata

Vulpia bromoides

V. Myuros Aa
Echinopogon caespitosus
Dichelachne spp. a
Festuca elatior var. pratensis ..
Phalaris tuberosa

P. minor

Agrostis avenacea

Koeleria phleoides

Lamarckia aurea

io)

Ww bv
Doe

NWONnPPNHr Wr OO WwW
Pp bw

PHYSIOLOGIC SPECIALIZATION IN P. GRAMINIS LOLII.

Ten clumps of Lolium perenne growing in the University grounds were potted and
tested with the stock culture of the rust. One gave sharp flecks (resistance), whilst
all the others gave ‘4’ reactions (susceptibility). The resistant plant and two of the
susceptibles were retained. Later the plants were inoculated with a different isolate:
the “resistant” clone produced ‘4’? reactions in a side-by-side comparison in which the
stock culture still gave flecks. A further interesting observation was that whilst the
“resistant” clone was heavily attacked by P. coronata loli, one of the two “susceptibles”
was quite resistant to this leaf rust. It is thus clear that the breeding of a type with the
combined resistance to stem and leaf rusts should not be difficult.

Further evidence of specialization came from L. rigidum. In a stand of rusted
Wimmera rye grass at Tichborne, New South Wales, Dr. I. A. Watson collected a plant
which was resistant. Seedlings from it segregated for resistance (fleck reactions) and
susceptibility (‘3 reactions) when inoculated with the stock culture. Typical plants
of each sort were grown to maturity without bagging, and seed saved for further testing.
These seedlings again segregated for resistance. Ten of these resistant ones were
inoculated with the different isolate of P. graminis loli and proved to be susceptible,
thus giving proof of specialization of the fungus.

In yet another case, 16 seedlings of Arrhenatherum elatius were grown to maturity
and each plant divided into separate entities for inoculation tests. Two of them, which
gave susceptible “3” reactions with the stock culture, produced resistant “1” reactions
with the second isolate.

\

AUSTRALIAN RUST STUDIES. VIII,

for)
bo

DIFFERENTIAL RESISTANCE OF INDIVIDUALS WITHIN SPECIES OF GRASSES.

In addition to the occurrence of resistance in Lolium spp., other cases of individual
plants showing resistance were found in the following: Agrostis avenacea Gmel,
Hordeum leporinum Link, H. marinum Huds., Briza maxima L., Phalaris tuberosa L.,
P. minor Retz., Bromus racemosus L., B. Gussonii Parl., B. hordeaceus L., B. madritensis
L., Lagurus ovatus L., Lamarckia aurea (L.) Moench., Lhrharta longiflora Sm., Phleum
pratense L., Arrhenatherum elatius var. bulbosum (Willd:) Spenner., Dactylis glomerata
L., Poa annua L., Lolium rigidum Gaud., L. multiflorum Lam.

These inoculations showed that in some of the cases, a pot of 30 to 40 seedlings
would give 3 or 4 resistant and the remainder susceptible plants, whereas the reverse
was found in other cases. As a safeguard, seedlings showing the unusual reaction were
grown to maturity in order to check their identity.

Inheritance studies were made in a few cases. In one, a collection was made ot
typical heads, nearing maturity, of Bromus madritensis growing in the Sydney Univer-
sity grounds. Pots of seedlings inoculated with the stock culture gave varying reactions:
some were ‘“X=’’, others a mixture of ‘X=’ to “X+’ and others “Xt”. Eleven of the
“X=” and 10 of the “Xz” were grown to maturity in the open.

In the next generation about 40 seedlings from each plant were tested. Of the 11
resistant (“X=”) plants, one gave ‘Xz’ (susceptible) reactions, five gave reactions
varying from “‘X=” to “X=” (heterozygous), and five gave “X=” (resistant) reactions. Of
the 10 original susceptible (‘Xi’) plants, five produced seedlings giving “X=” to
“Xz” reactions and five gave “X=” to “X” reactions—notably lower than the preceding.

For the next generation testing, representative resistant and susceptible plants from
each group—72 in all—were grown on. Seedling tests of their progenies showed that
three gave “Xz” (susceptible) reactions, nine gave “X=” to “X” (resistant) and the
remainder “X=” to “Xz” (heterozygous).

Pure-lining of the grass was not carried out, and observations at flowering time
indicate that natural crossing is to be expected. A genetical interpretation of the fore-
going results was not possible, but the evidence shows that within a species there are
marked differences between individuals in their resistance and susceptibility.

In the course of these seedling tests, the occurrence of albino seedlings was noted
in the following grasses: Lolium perenne, L. multiflorum, Agrostis avenacea, Hordeum
marinum, Phalaris minor, Vulpia bromoides, Koeleria phleoides, Dactylis glomerata,
Phleum pratense, and Agropyron trichophorum.

Variegation was found in seedlings of Lolium perenne and L. multiflorum. These
seedlings produced normal green leaves in later stages of growth, and their seedling
progenies in no case showed inheritance of chlorophyll deficiency.

RESISTANT GRASSES.

Apart from the occurrence of resistant and susceptible individuals within the
species mentioned, resistance only was found in many species. This was of two sorts.

In the first, small hypersensitive flecks were produced. The grasses concerned are
Aegilops ovata L. (some strains), Phleum pratense (some strains), Briza maxima
(some strains), Poa annua (some strains).

In the second, resistance was complete, since inoculation produced no reaction
whatever. These grasses were Briza maxima (some strains), B. minor, Phlewm pratense
(some strains), Secale montanum Guss., Agrostis gigantica Roth, Poa compressa L.,
Agropyron scabrum (Uabill.) Beauv. (long and short glumes), A. repens (L.) Beauv.

DISTRIBUTION OF P. GRAMINIS LOLTI.

In respect of time, the isolates examined were distributed as follows: 1918 (2),
1921 MUL) 1922) 4(2))),.. 1928) (3) 1924.5 (2) 925s | CA) 26) GS) al G27 (2a) eho eee
1929 (20), 1930 (24), 1931 (19), 19382 (18), 1933 (9), 1934 (45), 1935 (22), 1936 (21),
1937 (11), 1938 (19), 1939 (16), 1940 (9), 1941 (10), 1942 (2), 1943 (3), 1944 (2),
1945 (4), 1946 (4), 1947 (17),.1948 (9), 1949 (8), 1950 (6).

BY W. L. WATERHOUSE. 63

The source of the isolates was as follows: New South Wales, 249; Victoria, 4;
South Australia, 25; Queensland, 2; Tasmania, 30; Western Australia, 1; Australian
Capital Territory, 27; New Zealand, 31.

DISCUSSION.

Delimitation of the units of classification of living things is necessarily difficult.
It is particularly so in the organism under consideration because of the very wide
variations it shows. The characteristics of Puccinia graminis Pers. as a species are clear
as to general morphology and life history. Detailed studies, however, bring to light
differences which have led to the setting up of groups which are called “subspecies”
or “varieties”. Statistical studies of the spore morphology reveal differences between
them. Certain of them, e.g., P. graminis phlei-pratensis, may differ from others in their
capacity to attack the barberry. The parasitic behaviour shown on certain hosts further
emphasizes the differences between the subspecies and is, indeed, the main basis for
their determination.

Within these subspecies further separation is made into physiologic races. These
are characterized by the differential reactions they show upon groups of selected varieties
of the host plant. Thus P. graminis tritici is split into more than 200 physiologic races
which differ in the reactions produced on 12 particular varieties of wheat. The initial
choice of these differential varieties is an empirical one based upon numerous “trial and
error” tests. By more detailed study, as, for example, by using additional differential
host varieties, isolates which appear to be the same physiologic race are separable
further on the basis of the resistant or susceptible reactions now shown. These entities
have been styled “biotypes” and may be regarded as individuals or groups of individuals
which have the same genetic constitution.

In the present study no problem arises as to distinctions between biotypes and
physiologic races. The question has to be considered, however, whether the stem rust
on rye grass can rightly be considered as a physiologic race of one of the established
subspecies (or varieties) of P. graminis, or whether it should be regarded as a new
subspecies.

In his early work, Eriksson (1918) records as hosts of P. graminis avenae in
EKurope a number of grasses including Dactylis glomerata, Lamarckia aurea, Vulpia
bromoides, and Briza maxima. These all have been found in Australia infected by
P. graminis. In many of the cases the infection has been shown to be caused by one
or other of the recognized physiologic races of P. graminis avenae, e.g., race 2 and
race 7, of which the chief feature is their capacity to attack oats, even if some varieties
are resistant. In many other cases the stem rust present has been unable to infect oats,
or to do so very weakly in the case of onlv a few varieties in which a resistant reaction
is shown. In yet other instances both these rusts were present on the grass host:
one attacks oats and not Lolium perenne, and the other attacks L. perenne but not oats.
A detailed examination has shown that both types can infect the barberry. Differences
in the size of the spore forms is revealed by statistical examination of the measurements.
In itself this would not be sufficient to warrant establishing a new subspecies, since
significant differences are known to occur between certain well-recognized physiologic
races within a subspecies. But when, in addition, the grass rust is unable to attack the
cereal, then, instead of being regarded as a physiologic race of that cereal rust, it must
be considered to be more widely divergent and to belong to a different subspecies.

The question then arises whether it is one of the recognized subspecies of grass
stem rusts. Here again the rye grass rust is morphologically different when spore
measurements are examined, and it differs also in its capacity to infect the normal
hosts of these subspecies. Hence it must be regarded as a different subspecies.

Because of its frequent occurrence on Lolium spp. and the economic importance of
these grasses, the name P. graminis lolii is proposed for this rust.

Acknowledgements.

Valuable help has been given by Dr. I. A. Watson and Dr. E. P. Baker, and is
gratefully acknowledged. Thanks are due to workers in many places who have

.

64 AUSTRALIAN RUST STUDIES. VIII.

forwarded specimens for study, and to Miss Joyce W. Vickery of the National Herbarium,
for grass determinations. Financial assistance from the Commonwealth Research Grant
has been of the utmost value.

References.

[RiKSSON, J., 1918.—Fortgesetzte Studien tiber die Spezialisierung des Getreideschwarzrostes
(Puccinia graminis) in Sechweden und in anderen Landern. Centralb. Bakt. Il, Bd. 48:
349-417.

LEVINE, M. N., 1923.—A statistical study of the comparative morphology of biologic forms of
Puceinvia graminis. Jour. Agr. Res., 24: 539-567.

McALPINE, D., 1906.—The Rusts of Australia. yovt. Printer. Melbourne, pp. 349, plates 55.

STAKMAN, E. C., and LEvINE, M. N., 1922.—The determination of biologic forms of Puccinia
graminis on Triticum spp. Minn. Agr. Haupt. Sta., Tech. Bull. 8, pp. 10.

—, , 1924.—Puccinia graminis poae BE. & H. in the United States. Jour. Agr.
Res., 28: 541-548.

WATERHOUSE, W. L., 1929.—Australian Rust Studies. I. Proc. LANN. Soc N.S:W., 54:
616-680. {

————, 1930.—Australian Rust Studies. II. Biometrical studies of the morphology of

’

spore forms. Proc. LINN. Soc N.S.W., 50: 159-178.

65

CELAENOPSOIDES GUNTHER, 1942: A SYNONYM OF OPHIOMEGISTUS
BANKS, 1914 (ACARINA:PARASITIDAE).

By Cart BE. M. GuntHer, M.D., B.S., D.T.M. (Sydney), D.T.M.&H. (England),
Field Medical Officer, Bulolo Gold Dredging Limited, Bulolo, Territory
of Papua—New Guinea.

[Read /25th July, 1951.]

Synopsis.
Celaenopsoides buloloensis Gunther 1942, taken from rats at Bulolo, is congeneric with
Ophiomegistus luzonensis Banks 1914, from a snake in the Philippine Islands. This paper
records the consequent changes in nomenclature.

Mr. K. P. Lamb, of the New Zealand Department of Scientific and Industrial
Research, has very kindly pointed out (personal communication) that Celaenopsoides
buloloensis Gunther, 1942, is congeneric with Ohphiomegistus luzonensis Banks, 1914.

Banks’s original specimens were taken on a snake in the Philippine Islands; in
1947 Grant reported and redescribed the same species from a snake at Hollandia in
Dutch New Guinea. My specimens were taken from rats at Bulolo in the Mandated
Territory of New Guinea. Apart from the host-difference and certain specific differences,
there is one important distinction between O. luzonensis and C. buloloensis: the former
shows no parapodal plates, while the latter has long thin parapodal plates running back
to the tip of the abdomen. Banks’s key to the Parasitinae (1915) gives, in part:

“4. Venter with lateral chitinous plates, separate from the ventral plate, and extending

© [DOSISMMOIP TASia Oi Was) [NOCH “GoccocuososcocboanvusucadoggouccouudauLKo0Gob od 5
Venter without separate lateral plates reaching to the tip of the body ............ 6
DEPAMIAImApechurey in ~Ehemventrally plateiec ac | ieee isceeeencie e celctons = sie ise sh obese cle Caelenopsis [sic]
Analmapertunel may the: pOSt-=ventralesplater en aces cence since oierene chest ee eae Huzercon

6. Body nearly circular, legs short, first pair with quite long hairs at tip; hind legs

without teeth on the femora; male mandibles with a brush of long hair; male aperture

ENS Ste IVE SAE iui cles Pinay mh atrace egewtss See A ou eie eed gather open ecu Stouaa ahs Tribe Antennophorini.”

Later in the same publication Banks refers, in passing, to Ophiomegistus as belonging

to the Antennophorini. Now Ophiomegistus was erected aS a monotypical genus, and

O. luzonensis has no parapodal plates, but C. buloloensis is so clearly congeneric with

O. luzonensis that the presence or absence of parapodal plates is obviously of no

significance, and therefore Banks’s key is inadequate. However, that is not for
consideration here. The systematics of the genus are as follows:

‘Genus OPHIoMEGISTUS Banks, 1914.

Ophiomegistus Banks, N., 1914: J. Entom. and Zool., Pomona College, VI, 58; 1915: U.S.
Dept. Agric., Report 108, 87; Grant, C. D., 1947, Microentomology, xii, i, 22.

Celaenopsoides Gunther, 1942: Proc. Linn. Soc. N.S.W., Ixvii, 87.

1. O. luzonensis Banks, 1914 [loc. cit.]; Grant, 1947 [loc. cit.]. Type genus.

2. O. buloloensis (Gunther, 1942 [loc. cit.]) vice Celaenopsoides buloloensis Guuther,
1942 [loc. cit.].

66

ON TROMBICULA MINOR BERLESE, 1905.

By Cart E. M. Guntuer, M.D., B.S., D.T.M. (Sydney), D.T.M.&H. (England),
Field Medical Officer, Bulolo Gold Dredging Limited, Bulolo, Territory of
Papua—New Guinea.

(Three Text-figures. )
[Read 27th June, 1951.]

Synopsis.

Trombicula minor Berlese 1905 is the genotype of Trombicula, but unfortunately no
specimens of 7. minor are available. This paper records all that is known of the species; the
synonymy is discussed and a full list of references is given.

I. THE GENOTYPE.

The genus Trombicula Berlese, 1905, is of considerable medical importance, since
among its members are all the known vectors of tsutsugamushibyo (Japanese river fever,
mite typhus, scrub typhus). Trombicula minor Berlese, 1905, is the genotype of
Trombicula, and hence is of comparable acarological importance. But, unfortunately,
there are no specimens of J. minor at present available, and it has been deemed
advisable to assemble in one place all that is known of the species.

The original specimens were collected by Professor Kraepelin, then Director of the
Hamburg Museum, among bat guano in caves at Tjompea, in Java (Willmann, 1941,
p. 135: “Das Praparat von Trombicula minor ist mit folgender Fundortsangabe versehen:
‘Tjompea, Java, 19.111.1904, aus Hohlenguano gesiebt.’”). They were described by
Antonio Berlese (Berlese, 1905, p. 155: “‘Trombicula minor n. sp.’); it is apparent that
there were two complete specimens (Berlese, 1905, p. 156: “Duo vidi exempla collecta ad
Tjompea’”’, and Willmann, 1941, p. 133: “Im Praparat findet sich noch ein zweites
Exemplar ohne Hier’) and some fragments (Berlese, 1912, p. 94: “. . . io non possideo
che alcuni frammenti di questo acaro,...”).

The two complete specimens were lodged at the Hamburg Museum (Berlese, 1912,
p. 94: “I tipici si trovano al museo di Amburgo; .. .”’), and Berlese retained the
fragments himself (supra).

In 1941, Herr Carl Willmann, of Bremen, examined the Hamburg specimens and
redescribed them (Willmann, 1941, p. 132: “. . . habe ich eine Nachuntersuchung des
Typenexemplares vorgenommen. Das Praparat wurde mir vom zoologischen Museum in
Hamburg zu diesem Zwecke liebenswiirdigerweise bereitwilligst zur Verfltigung
gestellt.’”’). i

In 1950 I visited Hamburg, intending to study the genotype myself; Professor
F. Weyer, Director of the Bernhard-Nocht-Institut ftir Schiffs- und Tropenkrankheiten,
most kindly made inquiries for me at the Hamburg Museum and found that the two
specimens of 7. minor had been destroyed (Weyer, personal communication, 25th May,
1950: “Ich habe sofort mit dem Zoologischen Museum telephoniert und erfuhr, dass die
Typen und Paratypen yon Trombicula minor durch den Krieg zerstort sind.”).

I then consulted Herr Carl Willmann, of Bremen. He reviewed the above facts
and pointed out that 7. minor is unalterably the genotype of Trombicula, according to
Article 30c of the International Rules of Zodlogical Nomenclature (in Schenk and
McMasters, 1936, p. 34: “c) A genus proposed with a single original species takes that
species as its type.’’). He also pointed out that nothing but a suitable topotype would
serve to replace the lost genotype (cf. Schenk and McMasters, 1936, p. 8: “When the
original type material of a species is lost, the reviser of the species should choose a
neotype. The neotype must be a specimen from the typelocality of the species and
certainly must agree with the original figure and description.’’).

BY C. E. M. GUNTHER. 67

Later, I made inquiries at the Instituto Superiore de Sanita in Rome, but could
secure no information about the fragments retained by Berlese.

Dr. Cornelius B. Philip, of the Rocky Mountain Laboratory in Montana, very kindly
allowed me to see the correspondence he has had with Dr. A. Diakonoff, of the Zoological
Museum at Buitenzorg. Dr. Diakonoff, at Dr. Philip’s request, had secured some
material from the caves at Tjompea, and this was examined at the laboratory; unfor-
tunately there were no Trombiculae found. It is hoped that further attempts will be
made.

There the matter rests until such time as a suitable topotype is found and accepted.
Its acceptance must be based on its agreement with Berlese’s original description and
illustration (1905); but Berlese’s 1912 work, and especially the additional details
reported by Willmann in 1941, must also be given full consideration. Berlese’s fragments,
if they can be located, may be of some help, even though in 1912 he said of them:
«. . ho potuto rilevare solo i caratteri accenati, ma non si trova la base del capotorace

Text-figure 1.—Trombicula minor Berl. (Prona; 4a, corporis seta.).
(After Berlese, 1905, Pl. xv, figs. 4, 4a.)

coll’area sensilligera per potere vedere se esistono o meno gli occhi’. For completeness,
and for the convenience of future workers, I include here the relevant quotations and
the illustrations from both these writers:

Berlese, A., 1905: Redia, II, ii, 155: “Acari di Giava: Trombicula minor n. sp.: Albida
(?), elongata, in medio circiter corpore valde coarctata. Crista metopica posterius in
arcam subrhombicam desineno, utrinque longe unipilam (pili isti curte barbatuli).
Pedes antici caeteris longiores et robustiores, tarso longe conico. Palpi longiores, ungue
valido et longo, interne ad unguem spinis longis et robustis duabus, tentaculo autem sat
longo cylindrico. Derma corporis crasse et aeque granuloso-verruculosum ex verruca
quaque pilus exoritur qui sat brevis est, utrinque barbatulus. Pedes pilis conformibus
dense obtecti (Fig. 4). Ad 680 long.”

Berlese, A., 1912: Redia, VIII, i, 94: “Trombicula minor Berl. 1905. Albida. Pili
trunci.curtiores (20-25). Tarsi antici bene conici, acuti, tibiae vix curtiores, duplo et
dimidio longiores quam iati. Palpi graciles, longi, ungue exili et bene longo, falcato,
spinis internis duabus. Ad. 680 long.”

“Tnoltre anche il palpo é diverso [from 7. mediocris]. Esso é pit smilzo, con unghia
molto piu lunga e sottile, e con due sole spine alla sua radice, dal lato interno.”

“IT tarsi anteriori, ... sono conici e molto stretti nella parte apicale; misurano 100.
di lunghezza per 40m” di larghezza. Inoltre piccola é la differenza di lunghezza tra i
tarsi anteriori e le tibie, poiché queste sono lunghe 90 cioé piu corte (della decima
parte) del tarso.”

Willmann, C., 1941: ‘Zool. Anzeiger, CX XXIII, v/vi, 181: “Nach BERLESE (1905)
hat die Palptibia innen neben der Endkralle 2 starke Dornen.”

“Teh, konnte an dem Typenexemplar die Palpen leider nur in Dorsalansicht unter-
suchen, habe aber feststellen kénnen, dass BERLESE sich augenscheinlich geirrt hat, , , ,

68 ON TROMBICULA MINOR BERLESE, 1905,

An dem Typenexemplar ist trotz der ungtinstigen Lage deutlich zu erkennen, dass an
der Innenseiten 3 ‘Dornen’. .. vorhanden sind. Sie stehen aber so dicht nebeneinander,
dass es von oben aussieht, als wenn sie aus einem gemeinsamen Grundteil herauskamen.
Es ist aber anzunehmen, dass es sich in Wirklichkeit um 3 einzelne, starke Borsten
handelt, die so dicht neben- und etwas untereinander stehen, dass sie den Hindruck
einer dreifach gegabelten Borste hervorrufen. Hine dreizinkige Gabel an dieser Stelle
ware fiir die gesamte Familie der Trombidiidae etwas ganz Absonderliches. An der
Aussenseite der Endkralle steht eine lange, einfache Borst (Abb. 1b).”

“BERLESE war also selbst im Zweifel, ob die Spezies Augen habe oder nicht. Zu
einem einwandfreien Ergebnis hat die Untersuchung des Typenexemplares auch nicht
gefuhrt. Ich habe die Area sensilligera mit Immersionssystem untersucht und konnte
folgendes feststellen: Die lange, gerade Crista hat in der Mitte eine Langsrinne. Sie
erweitert sich hinten zu einer sechseckingen Area sensilligera (Abb. 1a). Die Haargruben,
Trichobothrien nach GRANDJEAN, liegen direkt am Aussenrande der Areole. Aus ihnen

2 3

Text-fig. 2.—Trombicula minor Berl. CAVE Fr 0.0) eee AG CAGSOMe mul bea melin
paio; B, palpo; C, apice del palpo molto piu ingrandito (lato interno). After
Berlese, 1912, p. 94.

Text-fig. 3.—Trombicula minor. a, area sensilligera; b, palptibia, dorsal.
After Willmann, 1941, p. 134.

regen die langen, steifen, feinbewimperten Sinneshaare heraus. Sie sind 107u lang,
wahrend die ganze Criste 127u lang ist. Diese deutlich zu erkennenden, von einem
Kreis umschlossenen, vertieften Haargruben sind nach aussen umgeben von einem stark
gewolbten, vollig glatten Wall, der etwa zwei Drittel eines Kreisumfanges einnimmt.
BERLESE zeichnet es 1905 so, als wenn neben dem schrag nach vorn gerichteten Seiten-
rande der Area eine starke, kreisformige Trichobothrie vorhanden sei, aus deren Mitte die
Sinneshaare hervortreten. Das ist aber nicht der Fall. Man sieht die Haargruben erst
bei tieferer Hinstellung des Mikroskoptubus. Sie liegen exzentrisch zu der Umwallung
und sind halb unter die Areole und den stark gewolbten dausseren Wall geschoben.
Dieser Wall jederseites des Seitenrandes der Area macht den Hindruck, als ob es sich
wohl um ein Auge handeln kénne. Mit Bestimmheit méchte ich das aber nicht behaupten.
Die Lage der Augen direkt am Seitenrande der Sinnesareole ware nicht besonders
auffallig, findet sich doch eine 4ahnliche Bildung bei der Gattung Trombiculoides JACOT,
bei der die als Augen angesprochenen Organe nur in einer starken Vorw6lbung des
Seitenrandes der Sinnesarea bestehen, also noch weniger hervortreten als hier. Das
Higenartige ist aber die Lage der Trichobothrien zu den ‘Augen’. Die Sinnesborsten
kommen genau auf der Grenze der Areole und der ‘Augen’ heraus, und die kreis-
formigen Umgrenzungen der Haargruben schieben sich, wie schon gesagt, sowohl unter
den Seitenrand der Areole als auch unter die vermeintlichen ‘Augen’ hinunter.”

Lest there be any doubt about the status of Berlese’s original specimens, Berlese
himself first started this hare (1912: “...e non 6 certo il caso di pensare ad individui -
giovani.’”), and Ewing followed him (1920 and 1938)—I include here Willmann’s state-

BY C. E. M. GUNTHER. 69

ment (1941): “Hs handelt sich bei dem Typenexemplar von Trombicula minor BERL. auf
keinen Fall um eine Nymphe, es ist ein aduites Tier, in dessen Korper deutlich ein
vollausgebildetes Hi zu erkennen ist.”

Il. THE SyNONYMY OF TROMBICULA MINOR.

I have been largely responsible for creating considerable confusion concerning the
taxonomy of ZT. minor and certain other Trombiculae, and it is now necessary for me
to explain and clear up the situation as far as I can.

In 1939 I described a larval mite from New Guinea, Trombicula hirsti var.
buloloensis, and stated that it was probably only a local variant of T. hirsti Sambon,
1927. In the same year I bred nymphs of this species and published (1939@) the
statement: “...as far as can be determined, these nymphs are identical with Trombicula
minor Berlese, 1904 [sic].”

Womersley (1939) considered certain collateral evidence from Queensland mites
and approved the above identification. He also discounted the slight differences between
T. hirsti Sambon and T. hirsti var. buloloensis, and pronounced them to be synonymous.

In 1940 I reviewed the literature and reached the following conclusions:

1. That 7. pseudoakamushi Hatori, 1919 (nec Tanaka, 1916), was synonymous with
T. mediocris Berlese, 1912.

2. That T. pseudoakamushi Hatori was also synonymous with 7. hirsti.

3. And therefore that, because of the assumption that 7. hirsti was a synonym of
T. minor, both JT. pseudoakamushi Hatori and YT. mediocris were also synonyms of
T. minor. 5

I used this synonymy in several subsequent papers (1939b, 1940a, 19400, and 1941).
It was adopted by many writers, but others did not accept it as proved, and Ewing
(1944), reviewing Willmann’s paper on the genotypes of 7. minor, concluded that
T. hirsti var. buloloensis was distinct from JT. minor.

In 1950 I took some nymphs of TJ. hirsti var. buloloensis to Herr Carl Willmann
in Bremen. He examined them, and from comparison with his notes, sketches, and
recollections of 7. minor he was able to state authoritatively that the two species were
quite distinct. This being so, the argument that JT. mediocris is synonymous with
T. minor falls down—there is no other evidence beyond Miyajima’s very vague sug-
gestion (1920). Thus JT. minor Berlese, 1905, stands alone, without any synonyms.

It is obviously necessary also at this stage to dispose of all those names which
have formerly been regarded as synonyms of 7. minor. That T. pseudoakamushi Hatori
is synonymous with 7. hirsti can hardly be doubted; Hirst (1929) and Gater (1932)
provide overwhelming evidence in support. That Radford (1942) listed 7. pseudo-
akamushi Hatori, 1919, as a valid species, without any reference whatever to
T. pseudoakamushi Tanaka, 1916, means nothing; Womersley and Heaslip (1948) list
“7 hatorii sp. nov.”, with T. pseudoakamushi Hatori as its synonym, but their discussion
is not convincing. d

But that T. pseudoakamushi Hatori is synonymous with 7. mediocris is probably
not so. The suggestion first came from Kawamura and Yamaguchi (1921), but there
is no supporting evidence available. Walch (1925) stated that his nymph of ‘7. pseudo-
akamushi (variatio deliensis ?)” resembled JT. mediocris, but his illustration of the
crista is clearly different from Berlese’s (1912). Thus it would seem that JT. mediocris
Berlese, 1912, must also be regarded as standing alone, without synonyms.

This leaves 7. hirsti. The situation here has become complicated because of the
work of Wharton (1946), Philip and Woodward (1946), and Philip and Kohls (1948).
They regard JT. hirsti var. buloloensis as Synonymous with 7. wichmanni (Oudemans,
1905) Hirst, 1917. This may well be—but the corollary would be that T. hirsti is
therefore also synonymous with TJ. wichmanni, for there is as little variation on the
one side as on the other. Moreover, Philip (1947) mentions that 7. hatorii (= T. pseudo-
akamushi Hatori, non Tanaka) is likely to prove a synonym of 7. wichmanni, which
would complete the-circle. However, I feel that this can wait until actual specimens

70 ON TROMBICULA MINOR BERLESE, 1905,

from all areas have been properly compared, and with the reservation that perhaps
T. hirsti and its synonyms will ultimately become in turn synonyms of 7. wichmanni,
I offer the following synonymy for 7. hirsti:

TROMBICULA HIRSTI Sambon, 1927.

(Sambon, L. W., 1927: Ann. Mag. Nat. Hist., 1X, xx, 157.)

. pseudoakamushi Hatori, 1919 (nec Tanaka, 1916).

. pseudoakamushi (variatio deliensis) Walch, 1924.

. pseudoakamushi (variatio deliensis ?) Walch, 1925.

. hirsti var. morobensis (nom. nud.) Gunther, 19388.

. hirsti var. buloloensis Gunther, 1939.

. hatorii Womersley and Heaslip, 1943.

. minor var. deliensis Walch, 1923, in Womersley and Heaslip, 1943.

. hirsti var. boloensis (laps. cal. J Farner and Katsampes, 1944.
. buloloensis Gunther, 1939, in Blake et al., 1945.

uibnomibieile buloloensis (Gunther, 1939) Wharton, 1946.

|} sels s| Sts) ly}

ACKNOWLEDGEMENTS.

My very grateful thanks are due to the many acarologists who have discussed
these matters with me and given me advice and help—in particular, Herr Carl Willmann,
who gave me so much of his time, and so courteously pronounced his opinions, and gave
me permission to quote him and use his illustrations; the late Dr. H. HE. Ewing, who,
though very sick, still exerted himself to offer advice; and Dr. C. B. Philip, who so
kindly read this paper, checked references, and gave much-appreciated advice.

References.
BERLESE, A., 1905.—Redia, II, ii, 154.
, 1912.—Tbid., VIII, i, 1.
BLAKE, F. G., Maxcy, K. F., SADUSK, J. F.. KoHLS, G. M., and BELL, E. J., 1945.—Amer. J..Hyg.,
XLI, iii, 243. “me
EwIneG, H. E., 1920.—Ann. Ent. Soc. Amer., xiii, 381.
, 1938.— J. Wash. Acad. Sci., xxviii, 292.
, 1944.—U.S. Nav. Med. Bull., xliii, 837.
FARNER, D. S., and Katsampss, C. P., 1944.—U.S. Nav. Med. Bull., xliii, 800.
GaTerR, B. A. R., 1932.—Parasitology, xxiv, 143.
GUNTHER, C. E. M., 1938.—Med. J. Aust., Aug. 6, 202.
—————, 1939.—Proc. LINN. Soc. N.S.W., lxiv, 73.
, 1939a:—Ibid., |lxiv, 466.
, 1939b.—Proc. 6th Pac. Sci. Cong., San Francisco.
, 1940.—PrRoc. LINN. Soc. N.S.W., lxv, 477.
, 1940a.—T bid. Ixv, 250.
, 1940b.—Med. J. Aust., Nov. 30, 564.
———,, 1941.—Proc. LINN. Soc. N.S.W., Ixvi, 391.
HATORI, J., 1919.—Ann. Trop. Med. Parasit., xiii, 233.
Hirst, S., 1917.—Brit. Mus. (Nat. Hist.) Heon. Series, vi.
, 1929.—Ann. Mag. Nat. Hist., III, x, 564. p
KAWAMURA, R., and YAMAGUCHI, M., 1921.—Kit. Arch. Exper. Med., iv, 169.
MIyAJIMA, M., 1920.—In Gater, 1932. ‘
OUDEMANS, A. C., 1905.—Hnt. Ber. ’s-Grav., I, xxii, 217.
PHILIP, C. B., 1947.—Am. J. Hyg., xlvi, 60.
, and Kouus, G. M., 1948.—Proc. 4th Int. Cong. Trop. Med. and Mal., Washington.
, and WOODWARD, T. E., 1946.—J. Parasit., xxxii, v, 502.
RADFORD, C. D., 1942.—Parasitology, xxxiv, 55.
SAMBON, L. W., 1927.—Ann. Mag. Nat. Hist., IX, xx, 157.
ScHENK, E. T., and McMaAsters, J. H., 1936.—Procedure in Taxonomy, Stanford University.
TANAKA, K., 1916.—IJkai Jiho, melxiv, 1701.
WatcH, E. W., 1924.—Trans. 5th Bienn. Cong. F.E. Ass. Trop. Med., Singapore, 1923, 601.
, 1925.—Kit. Arch. Exper. Med., VI, iii, 235.
WHARTON, G. W., 1946.—Proc. Ent. Soc. Wash., XLVIII, vii, 171.
WILLMANN, C., 1941.—Zool. Anzeiger, CXXXIII, v/vi, 131.
WoOMERSLEY, H., 1939.—Trans. Roy. Soc. S. Aust., LXIII, Wie We
, and HEASLIP, W. G., 1943,—Jbid., LXVII, i,

*

THE BEVOLUTION OF THE RADIO-MEDIAL AREA IN THE WINGS OF THE
MUSCOIDEA ACALYPTRATA (DIPTERA).

By Haroxtp F. Lower,

Systematic Entomologist, Waite Agricultural Research Institute,
University of Adelaide.

(Thirty Text-figures.)
F [Read 27th June, 1951.]

Synopsis.

The writer attempts to homologize the venation of the Muscoidea Acalyptrata with that
of the Nematocera and Orthorrhapha. As a basis for discussion of the changes involved, the
wings of two generalized Nematocera are considered in some detail. The medial field is then
traced through the higher Nematocera and Orthorrhapha to demonstrate that, in its essentials,
this field has remained unaitered since its first appearance in the Dolichopodidae, and evidence
in support of this view is presented. An essay is made to place the vena spuria and the vein
in the wings of the Syrphoidea and Muscoidea, usually referred to as M,, in their correct
relation to other parts of the venation, and to show that the vena spuria has exercised a
profound influence, even in the wings of the most advanced Diptera. Finally, in a series of
hypothetical figures, are set out the possible changes that have been necessary to evolve, from
the wing of an asilid-bombyliid-like ancestor, the wing of the Muscoidea Acalyptrata.

INTRODUCTION.

During the past hundred years the venation of the dipterous wing has claimed
the attention of many students. The need for a venational system early made
itself felt in that period of entomology when descriptive work was practically its sole
function. Through the efforts of Bates, Wallace, and others, great collections were being
built up, and the rapid naming of hosts of newly discovered insects was of prime
importance. In answer to this demand, the first venational systems were developed. Two
of these appeared in 1862, Loew’s system and Schiner’s. Both had one feature in
common: each was artificial, having as its object the supplying of a rapid and convenient
method of notation for descriptive purposes.

The first attempt at a study of wing venation, as contrasted with the mere giving
of names or numbers to the veins, was made in 1886, when Josef Redtenbacher
propounded his system of venational nomenclature. He realized that the six venational
fields were a common inheritance in the wings of all insects. To the principal vein in
each of these fields he gave the names accepted today, namely, Costa, Sub-costa, Radius,
Media, Cubitus, and the Anal group, and further, he adopted Adolph’s conception of the
alternation of convex and concave veins, a suggestion which then fell into abeyance until
its importance was again recognized by Lameere in 1922.

Using Redtenbacher’s ideas as a foundation, Comstock and Needham began their

work in 1898, but it was not till 1918 that Comstock, in his book “The Wings of

Insects”, gave a complete account of the now well-known Comstock-Needham system.
Based on morphology and homology, this system, modified as new evidence has become
available, has been universally accepted as the only scientific one. Some of the more
important modifications have resulted from the work of Tillyard, Alexander and, more
recently, Vignon. Tillyard’s researches have greatly clarified our ideas of the cubital
and anal fields; our present conception of the radial field is the result of Alexander’s
work. Vignon has made a new study of the dipterous wing as a whole, and while all

his conclusions are not yet accepted, there is much in his work that has real value.

In Australia, considerable research on the dipterous wing has been done by several
workers, but the outstanding contributions to our knowledge of the brachycerous
venation have come from Hardy.

=]
bo

EVOLUTION OF THE RADIO-MEDIAL AREA IN WINGS OF MUSCOIDEA ACALYPTRATA,

If evolution is to have any meaning for us we must regard the higher Diptera as
having developed from a more generalized ancestor having many features in common
with those of living Nematocera and Orthorrhapha, and it is to these that we must
look for aid in the interpretation of the venation of the wing in higher forms. This
paper makes no pretence to being a general study of the dipterous wing. I suggest
some modification of our concepts of the venation of the medial field and of the area
where that field and the radial are contiguous. The examples chosen for discussion are
those which appear to me to be our best guides to the interpretation of the venation
of this area of the wing in the Muscoidea Acalyptrata.

Evolutionary Trends in the Dipterous Wing.

The general trends of evolution have been to change the,contour of the wing and
to dispose the venation more effectively. From a long, narrow, petiolate wing, charac-
teristic of the Tipulidae (see Text-figure 1), a shorter, broader one has been developed
(see Text-figures 12, 14 and 15). Changes in venation have accompanied those in shape.
Whereas, in the primitive wing, numerous veins were disposed over the whole wing area,
in the higher Diptera their number is much reduced and they tend to be arranged
nearer the costal margin, where their strengthening has given to the wing the reinforce-
ment needed for sustained and rapid movement. To understand clearly these changes
and the stages through which they have passed, it will be necessary to discuss the
wing venation of some generalized types and then to follow the reduction through forms
of increasing complexity until the higher Diptera are met. By this means it will be
possible to homologize the veins of specialized forms with those of more generalized
types, so giving an insight as .to what the veins, present in the wings of the Muscoidea
Acalyptrata, really are.

Venation of Two Primitive Living Nematocera.

The most generalized types of venation, in living Diptera, are to be found in the two
nematocerous families, the Tanyderidae and the Psychodidae. As representative of
these, the wing of Nothoderus (Tanyderidae) and that of an undetermined psychodid
will serve to illustrate the archaic features of these wings. From the point of view
of venation, the Tanyderidae are the most primitive known living Diptera. The wing
of Nothoderus (Text-figure 1) is long and narrow, the veins being disposed evenly over
its whole area. Although considerable reduction has already occurred, it still shows a
very full venation. The subcosta still retains its primitive two-branched form, though
Se. is already in process of reduction. A complete radial field is present, R, and the
four branches of Rs all reaching the wing margin as independent veins. The medial
field, with its four independent branches, is complete, and the median cell, between
M, and M;, is small, elongate, sub-rectangular in outline, and closed by i-m. It is in the
cubital and anal fields that most departure from the wing of the archetype has occurred,
since considerable reduction in these fields is obvious. Cuj. and Cu,» have already
coalesced to form the single vein Cu,, while Cu, no longer reaches the anal margin.
The first anal vein alone is present, the second and third having been completely
suppressed.

The wing of Psychoda (Text-figure 2) shows features which indicate an advance on
that of Nothoderus. The venation is still of a generalized nature and is evenly disposed
over the whole wing area, but the length of the wing, relative to its width, is much
less. The subcosta is now represented by a single vein only, there being no trace of Sc..
The radial field is complete, as is the medial. The median cell is open, the crossvein i-m
having been lost. The cubital field has been further reduced by the complete loss of Cus,
while a single anal vein, IA (and that short and weak) has been retained.

In both these wings the characters to which I wish to draw attention are the
complete five-branched radial field, the complete four-branched medial field, and the small
median cell between M, and M;, in Nothoderus, closed by the cross vein i-m. This is the
primitive pattern of this part of the wing, a pattern shown in the wings of a very

BY H. F. LOWER. 73

small number of the older Nematocera only, and one which never reappears later.
It is the reduction which these fields undergo in the course of evolution that is the-
object of this study. ji

Text-figures 1-9.

i, 2—Generalized venation in the older Nematocera. 1, Wing of Nothoderus australiensis
Alexander (Tanyderidae) ; 2, Wing of Psychoda sp. (Psychodidae).

3.—Generalized venation in the older Asilidae; the wing of Leptogaster sp.

4-7.—Development of the medial field in the Asilidae. 4, Medial field of Leptogaster sp.
The small median cell closed by i-m, and the four free branches of the media are primitive
eharacters; 5, Medial field of Saropogon luteus. The anal turning of the termination of M,
is the first stage in the specialization of the medial field; 6, Medial field of Senobasis mendax.
Basad movement of the termination of M, has closed cell M,. Veins M, and M, have united at
the wing margin to form one vein, M,,,. Two closed cells in medial field; 7, Medial field of
Laphria sp. Basad movement along M, of point of union of M, and M, results in vein M,,,
becoming of considerable length.

8.—Wing of Laphria sp. (Asilidae).

9.—Medial field of Sphenoidoptera varipennis (Bombyliidae). The loss of the basal free
part of M, causes the first appearance of the cell M,+M,. The dotted line in the cell represents
the part of M, which has been suppressed.

J

74 EVOLUTION OF THE RADIO-MEDIAL AREA IN WINGS OF MUSCOIDEA ACALYPTRATA,

Venation of the Higher Nematocera.

Throughout the sub-order the above general disposition of the medial field is retained,
but great changes in the radial field, foreshadowing those by which this field has been
produced in the higher Diptera, have occurred in its more advanced members. Alexander
(1927 and 1928) demonstrated, in the Tipulidae, the manner in which evolution has
brought about the coalescence of R, and R, to form a single vein reaching the wing
margin as R,,.. He further showed that the changes applied not only to the Tipulidae,
but equally to all higher members of the order. R,,, is, therefore, a constant in the
wings of all Diptera except those few primitive species which still retain both veins as
separate entities. Where the union has occurred, R,, left as a separate prong of the
original R.,, fork, reaches the wing margin as an independent vein, and it, too, occurs
in all dipterous wings. In some of the higher Nematocera the R,,, fork, by moving
distad, has produced a single compound vein R,,;, but in many, R, and R,; retain their
separate identities so that a radial field of four veins, a disposition common in ‘the
Orthorrhapha, is formed. The veins are R,,., Rz, R, and R, (see Text-figures 3 and 8).

Venation of the Orthorrhapha.

Just as the more generalized examples of venation are to be found in the lower
Nematocera, so it is in the lower Orthorrhapha that is found what may be regarded
as the generalized venation of this division. From here onwards there is a strong
tendency towards the retention of a four-branched radial field, and it is the medial
field that becomes the focal point of reduction. The lower members of the Asilidae
form a good point of departure for a consideration of those changes which ultimately
lead to the type of medial field characteristic of the higher Diptera. The wing of
Leptogaster (Text-figure 3) illustrates those features of the venation common to many
of the lower Orthorrhapha: the subcosta is a single vein. The radial field is four-
branched, R,.., R;, R, and R,; all reaching the wing margin independently. Two cubital ©
branches, Cu, and Cu., are present and reach the wing margin. There are no anal
veins. The medial field (Text-figure 4) is of the primitive type, consisting of four
independent veins, M,, M:, M, and M,, all of which reach the wing margin. There is a
small, elongate, median cell, closed by i-m.

Specialization of the medial field begins in the Asilidae. An early stage of this is
shown in the medial field of Saropogon (Text-figure 5), where M,, instead of proceeding
directly to the wing margin, has its termination diverted towards the anal margin.
This tendency in M, becomes more pronounced in higher members of the family.

In Senobasis (Text-figure 6) further basad movement of the termination of M,
brings its tip into contact with that of M, at the wing margin, thereby closing cell Ms.
The medial field now includes two closed cells—me and cell Mz. The combination of
veins at the wing margin is Mg,,.

The highest of the Asilidae advance a stage further. The point of union of M, and
M, begins to move basad along M,, finally attaining the position shown in the medial
field of the wing of Laphria (Text-figure 7). There is thus brought about ‘a coalescence
of the two veins for a considerable part of their length.

Reduction of this field goes no further in the Asilidae, so that consideration of the
wing as a whole (Text-figure 8) will illustrate the highest type of venation occurring
in the family. The subcosta is a single vein and has considerably shortened. The radial
field consists of four branches, Ryo, Rz;, R, and R;. The union of R,,. with R, occurs
in the sub-families Laphriinae and Asilinae but, in the Dasypoginae, R,,. and R, both
end normally in the wing margin. R, and R, (the two small branches of R; are
aberrant) each attains the wing margin and shows a tendency to turn anteriorly towards
the wing apex. Of the medial field, three branches reach the wing margin. These are
M,, M, and M;,,. The median cell is small and elongate. Posterior to it lies the closed
triangular cell M;. The median cell and cell M, are closed by three veins or parts of
veins. Cell M, is closed by part of the vein M, while the median cell is closed by i-m
and part of M,. There are then, lying between M,,, and M,, two closed cells. In the
cubital field is seen the first stage of the reduction which gives rise later to the small

BY H. F. LOWER. 75

closed cubital cell of the higher Diptera; Cu,, by turning basad, has united with 1A, .
thereby closing cell Cu,. Cu, is vestigial, as also is 2A.

It is in the Bombyliidae that further stages in reduction are to be found. The
radial field remains relatively unchanged. It still retains its four branches but there
is a tendency for R, to turn apically, and, in the higher members of the family, this
tendency, in both R, and R,, is accentuated. In the medial field, however, very
significant reduction has occurred. That of Sphenoidoptera (Text-figure 9) has lost
the free basal part of M,, though three medial branches, M,, M, and M,,, still reach the
wing margin. The loss of the free basal part of M, results in the formation of a single,
large, closed, sub-triangular cell within the medial field. Anteriorly this cell is bounded
by M,,,, and posteriorly by M,. This large cell is not homologous with the median cell
of lower Diptera and should not be so referred to. It is a compound cell, M.+M,;. The
identity of the median cell is merged in that of the combined cell, and from here
onwards ceases to exist as an independent unit of the venation. Similarly, the “cross
vein” closing it is not the im of the lower Diptera. It is a serial vein composed of
three parts: one of these is the free basal part of M,, the second is the free terminal
part of M,, and these are linked in the middle by the true cross vein, i-m. The ‘cross
vein” so formed I propose to call the serial vein to indicate its true nature and to
notate as se. Hardy (1947) has shown that a similar course of events has produced
the compound cell in the wing of Lepidostola (Syrphidae). He is undecided as to
whether or not M, forms part of the “cross vein” since he states that “the apparent cross
vein may include the free basal part of M.’”. All the evidence, however, supports my
contention that the “cross vein” does include part of M,. A study of large numbers of
dipterous wings has convinced me that its inclusion, together with part of M,, is
certain in the wings of all Diptera which contain the cell M.+M,, that is, in all Diptera
above the Bombyliidae. It is largely due to the failure to recognize M, in its correct
position that has made impossible a satisfactory interpretation of the medial field in
the higher Diptera.

The three-branched medial field, enclosing cell M.+M;, is maintained throughout the
Orthorrhapha until the Dolichopodidae are reached, where a highly specialized field is
developed. In the higher dolichopodids and in all Diptera higher than this family the
terminal part of M, between the “cross vein” and the wing margin is suppressed and
_the medial field characteristic of the Cyclorrhapha makes its first appearance. In
Hydrophorus (Text-figure 10) the medial field is so disposed. Two branches only of
the media are evident; the anterior branch which reaches the wing margin is M,, the
posterior is M,. All that now remains of M,, as a separate entity, is that part of it
which enters into the formation of the vein, se. The only remnant of M, is that part
incorporated in the “cross vein” and the stub united with the end of M, as M,,,.

This advanced type of medial field is so important that some consideration of it
in detail is essential. If Text-figure 8 be again referred to, it will be obvious that, if
the basal part of M, and the distal part of M. be eliminated, a similar type of medial
field is the result. This has been done in Text-figure 11. The M,,. branch still persists,

but, of it, M, alone reaches the wing margin; M,,, reaches the wing margin, but the
free parts of M. and M, have lost their identities in linking with i-m to form the
serial vein. The important fact to note here is that both branches of M,,,. are accounted
for, M, terminating in the wing margin and M, in the anterior part of se. This is the
disposition of these two veins in all the higher Diptera. If such be correct, we should
expect te find, in the higher Diptera, “cross veins” of very irregular shape, indicative
of their mode of formation. This is exactly what does occur. This angular type of
“cross vein” is paralleled, time after time, in the wings of many Tachinidae which have
retained evidence of its origin more definitely than have other families of the higher
Diptera. It was this fact which, some years ago, caused me to begin the research,
the results of which are given in this paper. Such a cross vein is shown in the wing of
Prosenina (Tachinidae, Text-figure 12). The medial field of this wing is merely a
repetition of the hypothetical field of Text-figure 11, This is made eyen more evident

76 EVOLUTION OF THE RADIO-MEDIAL AREA IN WINGS OF MUSCOIDEA ACALYPTRATA,

if the two fields are superimposed, as has been done in Text-figure 13. The “cross
vein” and cell M.4+M, have the same shape because they have been brought about in
an identical manner. Throughout the Muscoidea Calyptrata every variation from the
highly angular (Prosenina, Tachinidae, see Text-figure 12), through the sinuous
(Cylindromyia, Tachinidae, Text-figure 14), to the almost straight (Wicrotropeza,
Tachinidae, Text-figure 15), cross vein exists. Even in the Muscoidea Acalyptrata (for
example, in the families Otitidae and Trypetidae) the sinuous se is not uncommon,
though in this group it is usually straight. This straightening of the “cross vein” has
hitherto tended to conceal its composite nature in those families in which it occurs.

Text-figures 10-15.

10.—Medial field of Hydrophorus sp. (Dolichopodidae).

The suppression of the free basal part of M, and the free terminal part of M, (shown by
dotted lines) has given rise to the earliest appearance of the medial field, typical of that of
the higher Diptera. The free parts of M, and M, are incorporated in the ‘cross vein”.

11.—The wing of Laphria sp. (as in Text-figure 8) with the free basal part of M, and the
free terminal part of M, (both shown dotted) eliminated to show how the medial field
characteristic of the Tachinidae has been developed. ,

12.—The wing of Prosenina sp. (Tachinidae) showing the serial nature of the angular
“cross vein” se. It is composed of the free basal part of M,, the free terminal part of M,;, and
the true cross vein i-m. Dotted lines represent the former courses of the suppressed parts of
M, and M,. X is the vein discussed later. (The cross vein notated as i-r should be r-m.)

13.—The medial field of Prosenina sp. (Tachinidae—solid lines) superimposed on that of
Laphria sp. (Asilidae—dotted lines) to show how the medial field of the higher Diptera has
evolved from one having asilid-like characteristics.

14.—Wing of Cylindromyia sp. (Tachinidae) showing sinuous form of vein se. Note stub
of M, directed towards wing margin.

15.—Wing of Microtropeza sp. (Tachinidae) showing straight form of vein se and the
stub of the suppressed free basal part of M,.

Venation of the Syrphoidea and Muscoidea Calyptrata.

The location of M, and M, in their true positions is essential to a correct interpreta-
tion of the medial field in the higher Diptera. Because this has not hitherto been
clearly understood needless confusion has arisen. A good illustration of this is the

BY H. F. LOWER. 1

vein which, branching from M,, is directed towards the wing apex (see X, Text-figure 12).
It is usually notated as M,, while M, itself is referred to as M,; the actual M, in the
serial vein is omitted altogether. Whatever this vein may be, it is certainly not M,.
Text-figures 12, 14 and 15 represent the wings of three species of Tachinidae. In these
the termination of M, is clearly visible as a stub vein projected towards the wing
margin. A fold in the wing membrane connects this stub vein with the margin, and
suitable staining defines its former course even more clearly. What has occurred in
the muscoid wing is that the termination of M, has been suppressed, its remnant being
that section of it lying between the serial vein and the end of the stub vein. M.,,
forming as it does part of se, cannot possibly occur in the position in which it is
represented.

A further fact which militates against vein X as being M, is that of its direction and
position. The occurrence of M, directed apically is highly exceptional, being found in
those families only whose venation is complex. These families are the Mydaidae,
Apioceridae and Nemestrinidae, and in the wings of each the presence of a four-branched
radial field leaves no doubt that the vein posterior to R; must be M,, or at least a
medial branch incorporating it. It would certainly conflict with the principles of
evolution of the insect wing if, in all the families of the Nematocera and the
Orthorrhapha, M, should tend to continue straight to the wing margin or be directed
anally, only suddenly to assume a completely new and abnormal position in the
Syrphoidea and Muscoidea Calyptrata, and then, equally suddenly, to revert to its
normal position in the Muscoidea Acalyptrata and the Hippoboscoidea. Yet this
violence to homology has been accepted despite the fact that in both the Syrphoidea
and the Muscoidea Calyptrata M, is clearly present in its normal position. The truth
is that, in the Diptera as a whole, M, is remarkably constant in both its position and
direction. Text-figures 16 and 17 represent the wing of Calliphora stygia Fabr. (Calli-
phoridae). Text-figure 16 is notated from Tillyard (1926); Text-figure 17 is notated to
show the correct positions of M, and M.. .

Vein X, then, not being M, as usually understood, there appear to be two possi-
bilities only as to its nature. It is either part of the anterior media, MA, or it is
part of the radial field. In the dipterous wing the balance of evidence at present
appears to favour the presence of MA, though Tillyard in 1926 (quoted from Hardy,
1947) stated that MA “appears to be entirely missing in most recent orders’, thereby
contradicting the opinion he had expressed in the previous year. Vignon (1932)
claimed MA to be present, stating that the vein usually referred to as Ry; was in
reality MA,; in the Syrphidae it is the vena spuria which he believes to be MA,. Hardy
(1947) considers that MA may be present in the Syrphidae, possibly incorporated in the
vena spuria. The matter is so important, however, that the presence of MA must be
demonstrated beyond all reasonable doubt before we can embody it in. our scheme of
venation, since our so doing will mean an entire recasting of the venation of the
dipterous wing, a course which, at this stage of our knowledge, I do not feel to be
warranted. Should MA be proven to be present, and Vignon’s suggestion that the
vena spuria is MA, be accepted, then I consider vein X to be the vestige of MA,, as I
believe it to be the functional part of the vena spuria. This would align my views
with those of Vignon, at least in so far as this part of the wing is concerned. Subject
to what has been said above, I shall regard MA as being absent from the dipterous
wing and consider the present medial field to have been derived from the original
postericr media, MP.

Provisionally rejecting the presence of MA, I believe vein X to be part of the
radial field and considerable evidence exists to support this view. The vein makes its
appearance in the wings of the Syrphoidea and the Muscoidea Calyptrata only, that is,
in those families which have the vena spuria either actually present or which retain
obvious vestiges of it. These two facts, considered together, suggest that there is some
relation between them. Examination of large numbers of syrphid wings shows that
the development of the vena spuria varies within wide limits among the species and

78 EVOLUTION OF THE RADIO-MEDIAL AREA IN WINGS OF MUSCOIDEA ACALYPTRATA,

even among individuals. Sometimes little more than the vestiges where it crosses
the so-called r-m can be observed. In other instances it attains a considerable length.
In some few examples it is present almost as a complete functional vein. In these
latter it appears to have its origin in Rs, and to be directed towards the termination
of M,, with which it may or may not coalesce. Even when incomplete, the stained
wing often shows connections with Rs and M, In Microdon (Text-figure 18) it is
practically complete, its distal end making contact with M, near the termination of the
latter. From here it continues towards the wing apex as a functional vein. My
interpretation, therefore, is that the vena spuria is R; and hence vein X is R,; (Text-
figure 18, and see Text-figures 14, 15 and 17). This means that in existing syrphids

Text-figures 16-20.

16, 17.—Wing of Calliphora stygia Fabr. (Calliphoridae). 16, Notated from Tillyard
(1926); 17, Notated to show correct positions of M, and M,. Rest of notation as explained
in text. (R.:+m, should be R,+M,.)

18.—Wing of Microdon fulgens (Syrphidae) showing the venational pattern from which
developed that of the higher Diptera. Note the very complete vena spuria (v.s.) and its con-
nections. The terminal part of the vena spwria turns apically after coming in contact with M,.

19.—Wing of Platypezoides diversa (Platypezidae) showing fusion of R, with M,.

20.—Wing of Chrysomyza aenea (Otitidae) showing the apical trend of R,+M,, an example
of the influence exerted on the combination by R,.

the functional part of the vena spuria is R;. Hardy (1947) questions the presence of
R,; in the wings of Syrphidae without denying the possibility of its occurrence. I
believe that, when the vena spuria has been more widely studied, my contention that it
is R, will be upheld. If the higher Diptera have developed from simpler forms allied
to the lower, it seems difficult to deny to the Syrphidae the presence of R;, which is a
constant in the wings of the Nematocera and the Orthorrhapha. This would mean that
the three radial branches in cyclorrhaphous wings are R,.., R, and R,, with the vestiges
of R; still retained as an independent vein in Syrphoidea and the Muscoidea Calyptrata.
The cross yein, usually referred to as r-m, therefore connects not R,,, with M,,., but R,

BY H. F. LOWER. 79

with M,,., passing across R; as it does so. The true r-m is the posterior section of
this vein.

The Muscoidea Calyptrata retain vestiges only of the vena spuria, but its former
course across “r-m’”’ is often defined by a noticeable angularity in “r-m’”’ with short stubs
of the original vein on either side (see Text-figures 14 and 15). When these are
present, folds in the wing membrane connect them with Rs basally and with the tip of
M, distally. In such cases the only part of R,; that remains is the vein X (see Text-
figure 12). The course of events, from the fully developed vena spuria to the final
result aS above, may have been brought about in the following manner: R; originally
left Rs as a separate vein directed towards the tip of M,, near where it turned sharply
in an apical direction as other branches of the radius tend to do. Ultimately, union
occurred between R,; and M,, and since M, now took over the functions previously
performed by that. section of R; between Rs and M,, disintegration. of this section began.
An early stage of the reduction is represented in the wings of those Syrphidae in
which the vena spuria is most complete (see Text-figure 18). With the passage of
time the whole section was eliminated, its only vestige occurring where the vein crossed

2|

Rise — R3

Text-figure 21.

Wing of Sapromyza ocellaris Malloch (Lauxaniidae) to show the system of notation used
in this paper for the wing of the Muscoidea Acalyptrata.

“r-m” (see Text-figures 14 and 15). In a later stage of development M, ceased to reach
the wing margin (see Text-figure 15) and the distad movement of the point of union
of R; and M, resulted in the coalescence of the two veins as R;+M, (see Text-figure 20).
The tendency for fusion to occur between R; and M, is early shown in the course of
evolution. In varying degrees of completeness it is found in the wings of many
Orthorrhapha. In the Tabanidae coalescence occurs, in most cases at the distal ends
of the two veins, near the wing margin. More advanced examples of fusion occur in
the Dolichopodidae, but it can best be seen in the wings of Platypezidae (Text-figure
19, Platypezoides) and Pipunculidae.

Venation of the Muscoidea Acalyptrata and Hippoboscoidea.

The culmination of the process of reduction is attained in these groups, and the
preceding discussion has been a necessary introduction to a satisfactory interpretation
of the venation, more particularly of the Muscoidea Acalyptrata. Further specialization
eliminates even the vestiges of R; as a separate vein, its identity being merged in that
of M,. Nevertheless, the influence of R; is still noticeable in certain of the Otitidae,
Tanypezidae and Calobatidae, in which a strong tendency is shown for R,;+M, still to
be directed apically, as in Chrysomyza (Otitidae, Text-figure 20). In the higher families,
such as the Lauxaniidae, this tendency is lost, R;+M, continuing more or less directly
to the wing margin. There is thus produced the venational plan which is characteristic
of the most advanced Diptera.

Sapromyza ocellaris Malloch (Lauxaniidae, Text-figure 21) exemplifies this type of
venation. The number of veins has been greatly reduced, they tend to be reinforced in
the anterior part of the wing, while large open cells occupy much of its posterior

é

80 EVOLUTION OF THE RADIO-MEDIAL AREA IN WINGS OF MUSCOIDEA ACALYPTRATA,

region. A noticeable feature is the small number of closed cells, the result of the loss
of cross veins, of which three only, “y-m”’, i-m incorporated in se, and m-cu remain. To
indicate the true nature of “r-m” which bridges not R,,; and M,,., but R, and M,,., I
notate it as r-m. The subcosta is always short, making contact with the costa less
than half-way along the margin. R,,. is also short, and in many families its tip occupies
the position formerly occupied by that of Se. This results in the elimination of Sc,
of which only a vestige remains near the wing base. If present, the sub-costal cell is
small, but the basad movement of the termination of R,,. often results in its obliteration.
Cell R. is attenuated. Rs gives rise to two branches. The anterior of these is Rs, a
long vein, more or less parallel to R,,., and terminating in the wing margin. The
posterior branch is R,, which continues independently to the wing margin. Of R; the
distal part only remains, fused with M,. The medial field consists of two branches.
The anterior branch is M,,, as far as the serial vein, where M, turns anally to form
part of that vein. M, united with the vestiges of R; reaches the wing margin as
R-+M,. The posterior branch of the media is M,, which unites with the vestige of M,
in the serial vein and reaches the wing margin as M,,, The large cell, bounded
anteriorly by M,,. and posteriorly by M,, is cell M.+M;. This is closed by the serial
vein se, composed of i-m, joined to parts of M, and M;. Cell M, is very large. Cu,
after forming the posterior boundary of cell M, turns anally and unites with 1A. The
vein Cu,+l1A, so produced, is always short, never reaching the wing margin. Vestiges
of Cu. and 2A sometimes occur near the wing base.

The venation of the Hippoboscoidea (Text-figure 22) is difficult of interpretation
in the medial field by reason of further reduction. The serial vein, sé, appears to have
been eliminated, leaving cell M.4M, open, but until I have a large collection of this
superfamily available for study my suggestions can only remain tentative. It is often
the exceptional individual which provides the key to difficulties, otherwise insuperable,
and such are more likely to occur when there is ample material for examination.
The effects of reduction have been to strengthen the veins of the radial field at the
expense of those of the medial, and in these circumstances the loss of se would be a
not unexpected result. Whether this has or has not happened will not affect the rest
of the venation, which is modelled on that of the Diptera Acalyptrata.

Conclusions and Discussion.

As pointed out earlier, discussion of the venation of the lower Diptera was not an
end in itself. I have used it in the elucidation of some difficulties of interpretation in
the radial and medial fields of the Muscoidea Acalyptrata, and my conclusions are those
stated above. I believe this venational type to have been derived from one haying
affinities with those of both the Asilidae and Bombyliidae. Such a wing may have been
not unlike that of Text-figure 23. Text-figures 24-30 represent possible changes that
occurred, culminating in the wing of Sapromyza (see Text-figure 21). The wing in
Text-figure 23 combines a complete bombyliid-like radial field with a complete asilid-
like medial field. In Text-figure 24 reduction has already resulted in the coalescence
of R, and R. and the closure of cell M;. Text-figure 25 shows the formation of the
combined cell M.+M, by the loss of the free basal part of M;, while R; is degenerating
into the vena spuria. In Text-figure 26 the loss of the terminal part of M. gives rise
to the final development of cell M.+M;, which is closed by se. The basal part of R; is
almost suppressed. In Text-figure 27 se is tending to straighten while only the free
terminal part of R; remains. M, is being reduced and no longer reaches the wing
margin. Text-figure 28 represents a further stage of Text-figure 27. In Text-figure 29,
M, has lost its identity by its coalescence with R;, the influence of which is still shown
by the direction taken by R;+M,. Text-figure 30 is the typical wing of the Muscoidea
Acalyptrata. A comparison with Text-figure 23 shows the very great reduction which
has occurred. These changes may be summarized as follows. Throughout the series
there has been a great reduction in the number of veins and their disposition has been
completely changed. Their general costad movement has produced large areas in the
posterior part of the wing membrane where venation is absent. The veins towards the

BY H. F. LOWER. 8]

h Sc Rise

Text-figures 22-30.

22. Wing of Ornithomyia sp. (Hippoboscidae) to show the maximum of reduction in the
dipterous wing. Note strengthening of anterior veins and corresponding weakening of the
veins in the posterior region of the wing. The dotted line shows the position of the
suppressed se. \

23-30.—Possible stages in the evolution of the wing of the Muscoidea Acalyptrata.
23, Wing of hypothetical orthorrhaphous ancestor. A five-branched bombyliid-like radial field
and a four-branched asilid-like medial field are present. The median cell is small and elongate.
24, R, and R, have fused to form R,,,. A four-branched radial field, R,,,, R,, R, and R,,
is produced. Basad movement of the termination of M, has closed cell M, and the combined
vein M,,, has been developed. This results in the reduction of the medial field to three
branches reaching the wing margin M,, M, and M,,,. 25, Following the coalescence of its
distal part with M,, the base of R, is disintegrating to form the vena spuria. Cell M,+M,
has been formed by the suppression of the free basal part of M,; the medial field still has
three branches reaching the wing margin. 26, Further disintegration of the vena spuria occurs;
the radial field still remains four-branched. The medial field has completed its development
by losing the free terminal part of M,. Two branches only of the media now reach the wing
margin M, and M,,,. The angular “cross vein” se, by its shape, still shows its components.
27, R,,. is shortening. M, now fails to reach the wing margin. The ‘‘cross vein” se is
Straightening. All trace of the vena spuria is lost except for the angularity in r,-m, where it
previously crossed. The fusion of R, and M, is proceeding. 28, R,,. is very short. Most of
the free terminal part of M, has been lost. Cell Cu, almost complete. 29, Influence of R,
still shown by apical turn in R,+M,. All trace of M, as an independent vein lost. Cell Cu,
complete. 30, The wing of the Muscoidea Acalyptrata. For complete notation see Text-
figure 21.

K

82 EVOLUTION OF THE RADIO-MEDIAL AREA IN WINGS OF MUSCOIDEA ACALYPTRATA,

costal margin are strengthened; those nearer the anal margin are weakened. I have
attempted to explain these changes logically, using as evidence in support of my
contentions the information available in the wings of living Diptera. With the exception
of the hypothetical development series (Text-figures 23-30), all wings figured are those
of actual insects.

Acknowledgements.

My thanks are due to Miss Helen M. Brookes, who assisted with the lettering of the
Text-figures, and to Miss Margaret EH. Morphett, who also helped with the lettering and
drew Text-figure 8.

Note on Staining Methods.

In clarifying doubtful parts of the venation, such as unions between veins, and the
courses previously taken by veins now suppressed, I have found suitable staining
invaluable. Various stains, and methods of using them, were tried before the following
method was adopted as being the most satisfactory. The stain used consists of one
gramme of basic fuchsin dissolved in 100 c.c. of 95% alcohol. Wings, removed from
freshly killed insects, were dropped into 95% alcohol, in which they were allowed to
stand for half an hour before transferring to the stain. Wings from dried insects were
put directly into the stain. Staining occurs very slowly, a desirable feature, since the
process is at all times completely under control. The bases of the long veins first
take up the dye, which then moves slowly along them until they are completely
stained. Very prolonged immersion is necessary before absorption by the wing membrane
begins, though areas in which veins have been recently suppressed absorb the dye
more easily than those which have not been veined or in which venation has long been
lost. Determined by what the operator is aiming at, staining, even for periods of four
to six weeks, may not be too long. Hvery few days the wings are removed, washed in
95% alcohol, and examined under the microscope to note the extent of the staining.
When this is sufficient for the purpose in view, they are immersed in absolute alcohol
for half an hour, then in xylol for the same length of time, and mounted. Since basic
fuchsin is sensitive to even, the slightest trace of acid, some neutral mountant must be
used. For permanence and best retention of colour “Sira’’ was found to give the best
results.

References.

ALEXANDER, C. P., 1927.—The Interpretation of the Radial Field of the Wing in the Nematocerous
Diptera, with Special Reference to the Tipulidae. Proc. LINN. Soc. N.S.W., 52, 42-72.

, 1928.—A Comparison of the Systems of Nomenclature that have been Applied to the
Radial Field of the Wing in Diptera. Proc. Fourth Intern. Cong. Entom. Cornell, U.S.A.,
700-707.

Harpy, G. H., 1947.—The Wing Venation of Syrphidae. Hntom. Month. Mag., 83, 142-144.

- —, 1947.—Miscellaneous Notes on Australian Diptera. XIII. The Origin of the Vena
Spuria. PRoc. LINN. Soc. N.S.W., 72, 229-232.

'TILLYARD, R. J., 1926.—The Insects of Australia and New Zealand, p. 369.

VIGNON, P., 1932.—Nomenclature des veines de l’aile chez les Diptéres. Encyc. Ent. Dipt., 6,
133-142.

STUDIES ON AUSTRALIAN MARINE ALGAE.
VI. NEW GEOGRAPHICAL RECORDS OF CERTAIN SPECIES.
By VALERIE May, M.Sc.,

National Herbarium of N.S.W., Sydney, Australia.
[Read 27th June, 1951.]

Synopsis.

In this paper it is shown that Australian records of Neomeris dumetosa refer to both that
species and N. van Bosseae.

Further, for the first time, Galaxaura fasciculata, G. glabriuscula and G. angustifrons are
recorded from Australia; Sargassum fallax and S. flavicans from New South Wales; Sargassum
cristatum and Hypnea cenomyce from both north and east Australia; and Hormophysa triquetra
and Hypnea cervicornis from Western Australia.

In addition, a list of Myxophyceae, determined by Dr. F. Drouet, and including algae new
to Australia, is presented.

INTRODUCTION.

This paper continues a series recording the occurrence of algae not previously
known from Australia, and extending the known geographic range here of other species.

The specimens cited are in the following herbaria: National Herbarium of N.S.W..,
Sydney (quoted as N.S.W.), the Lucas Collection now held at the above Herbarium
(quoted as A.H.S.L.), the Herbarium of the Botany School, University of Sydney
(quoted as SYD.), the herbarium of the Sydney University Biological Society, held
at the Botany School, University of Sydney (quoted as S.U.B.S.), the herbarium of the
North Queensland Naturalists’ Club, Cairns, Queensland (quoted as N.Q. Nats.) and my
own herbarium (quoted as V.M.).

CHLOROPHYCEAE.
NEOMERIS SPECIES IN AUSTRALIA.
Neomeris dumetosa Lamx. has been recorded from Australia.. The specimens cited
include both Neomeris dumetosa Lamx. and Neomeris van Bosseae Howe and are as

follows:
Neomeris van Bosseae Howe. Collected at Bowen, Qld., by Rainford, in Sept. 1927,
May 1928, and again in Jan. 1929 (all in Herb. A.H.S.L.). The first two:collections are
recorded by Lucas (1927) as N. dumetosa Lamx. The specimens agree with Harvey’s
Exsice. Alg. Friendly Islands No. 81 from Tonga, cited by Howe (1909) as being

N. van Bosseae.
N. dumetosa Lamx. Collected at Low Island, Qld., by Perrin and Lucas in May 1931

(in Herb. A.H.S.L.). Recorded by Lucas (1934) as N. (?) dumetosa Lamx. The
specimens are broken, but the bands of lime formed by cohering primary branches are

quite definite.
PHAEOPHYCEAE.
SARGASSUM FALLAX SOND.
New Record for New South Wales.
This species is described and illustrated by J. Agardh (1889); in Australia it is
already known from Western Australia and from Queensland.
Locality. Date. Herbarium. Notes.

Collaroy, nr. Sydney, N.S.W. 24,vili,1948 V.M. No. 377 Fertile, Drift,

84 STUDIES ON AUSTRALIAN MARINE ALGAE. VI,

SARGASSUM FLAVICANS (Mert.) J. Aq.
New Record for New South Wales.
This species is known from Queensland and Western Australia and it is now
recorded from New South Wales. All specimens quoted were kindly checked by Dr.
Tore Levring against original Agardh collections now at Lund, Sweden.

Locality. Date. Herbarium. Notes.
Near Boom, Botany Bay, N.S.W. 14.xii.1944 V.M. No. 235 Trawled.
Mouth of Gunnamatta Bay, Port 8.vi.1945 V.M. No. 811 Pp
Hacking, N.S.W.
Port Hacking, N.S.W. —.v.1944 V.M. No. 1007 Coll. by

R. Bouchier.
SARGASSUM CRISTATUM J. AG.
New Record for New South Wales and Queensland.

Previously known from the west and south coasts of Australia, this species is now
recorded, though only as drift, from New South Wales and from Queensland. Specimens
quoted were kindly checked by Dr. Tore Levring against original specimens in the
Agardh Herbarium now at Lund, Sweden.

Locality. Date. Herbarium. Notes.
Collaroy, nr. Sydney, N.S.W. 7.vii.1944 V.M. No. 274 Drift.
Mooloolabah, Qld. 24.1.1944 V.M. No. 1012 35

HORMOPHYSA TRIQUETRA (L.) KuErz.
New Record for Western Australia.

In the National Herbarium of New South Wales is a specimen collected by A. H. S.
Lucas at Port Stephens, N.S.W., in June, 1909, and identified as Hormosira ? articulata
(Forsk.) Zan. The label bears the additional information that this identification was
confirmed by Mrs. Gepp. This species was recorded from New South Wales by Lucas in
1913 (p. 51). The species is now known from the northern and eastern coasts of
Australia.

Osborn (1947) has recorded that the Australian plant known as Hormosira ?
articulata (Forsk.) Zan. is in fact Hormophysa triquetra (l.) Kuetz.

The present is the first record of this species from Western Australia.

Locality. Date. Herbarium.
Dongara, W.A. 15.xi.1942 V.M. No. 2689, 2698, 2699, 2700.
MyYXOPHYCEAE.

Dr. Francis Drouet, of the Chicago Natural History Museum, has kindly identified a
number of specimens of Myxophyceae for me. These include algae new to Australia. As
our blue-green flora is particularly poorly studied, and their records often unreliable, it
seems wise to publish all of these identifications by Dr. Drouet, which are therefore
listed below. An additional list will be included in a forthcoming publication on the
Marine Algae of Brampton Island, Qld. /

ANACYSTIS AERUGINOSA (ZAN.) DR. & DAtIry.
Locality. Date. Herbarium. Notes.
Curl Curl, nr. Sydney, N.S.W. 1.x11.1945 V.M. No. 1158 Black slime on

rocks nr. beach.
CALOTHRIX CRUSTACEA Born. & FLAH.

Corrimal Headland, N.S.W. 26.111.1945 V.M. No. 648

ENTOPHYSALIS CONFERTA (KueEtTz.) Dr. & Datty.
Wattamolla, nr. Sydney, N.S.W. 26.viii.1944 V.M. No. 214 On Lyngbya con-
fervoides Gom.
Growing in
rock pool,

BY VALERIE MAY. 85

HoORMOTHAMNION ENTEROMORPHOIDES GRUN. EX Born. & FLAH.

Heron Island, Great Barrier Reef, —.i.1948 V.M. and S.U.B.S. No. A18
Qld.
” 9 99 50 " 5 s No. A22
No. 37

” ” ” a ” ” 99 a9

HoRMOTHAMNION soLUTUM Born. & GRUN.
Floating, 25 miles EH. of Bedout Is., 23.x.1945 V.M. No. 2087 From surface scoopnet
W.A. from C.S.I.R. Fisheries
Investigation ship
“Tsobel’”’. Previously
recorded (May, 1946)
as Nodularia prob.
spumigena Mert.

HyYDROCOLEUM LYNGBYACEUM GoM.

Heron Island, Great Barrier Reef, —.i.1948 V.M. and S.U.B.S. No. 36
Qld.
LYNGBYA CONFERVOIDES GoM.
Wattamolla, nr. Sydney, N.S.W. 26.vili.1944 V.M. No. 214 Growing in rock pool.
LYNGBYA LUTEA GOM.
Corrimal Headland, N.S.W. 26.i11.1945 V.M. No. 648
LYNGBYA MAJUSCULA GOM.
Adams Beach, Stradbroke Is., More- 15.i.1944 V.M: No. 930
ton Bay, Qld.

LYNGBYA SEMIPLENA GOM.

North Curl Curl, nr. Sydney, N.S.W. = 21.vii.1947 V.M. No. 2392 On rocks near
; beach. Prevalent.

Curl Curl, nr. Sydney, N.S.W. 1.xii.1945 V.M. No. 1158 Black slime on
rocks nr. beach.

LYNGBYA SORDIDA GOM.
Headland, Palm Beach, nr. Sydney, 26.xi.1947 V.M. No. 2470 Small form.
N.S.W.

RIVULARIA FIRMA WOMERSL.
Twofold Bay, Eden, N.S.W. 24.1.1946 V.M. No. 2031 Coll. by EH. Pope.

SKUJAELLA HILDEBRANDTII (GOM.) J. DE TOoNt.
Joseph Bonaparte Gulf, W.A. 13.ix.1949 V.M. No. 2817 Surface trawl.

RHODOPHYCEAE.
GALAXAURA SPECIES.

The recent appearance of Chou’s two papers (1944, 1945) on Pacific species of
Galaxaura, containing keys to the species, assists greatly the identification of specimens
belonging to this difficult genus.

Chou, dealing with the Pacific species, records only five species as from Australia,
viz., G. spathulata, G. arborea, G. wmbellata, G. oblongata and G. elongata. Svedelius
(1945) gives important descriptions and comparisons of species, and records also from
here G. fastigiata and G. obtusata, if indeed the latter differs from G. wmbellata.

The following three species are now also recorded from Australia, while the
additional occurrence of one of these species, and of one other species, are to be
recorded in a forthcoming paper on the Marine Algae of Brampton Island, Qld,

86 STUDIES ON AUSTRALIAN MARINE ALGAE. VI.

GALAXAURA FASCICULATA KJELLM. EMEND CHOU.

Known from the Celebes and Philippine Islands, Tonga, Japan, Malayan Archi-
pelago, ete.

Locality. Date. Herbarium. Notes.
Whitsunday Reef, Hayman Island, —.viii.1946 V.M. No. 2284 Coll. Mrs. Bing-~
Qld. also SYD. ham.
Coll. F. Perrin
Low Island, N. Qld. ee N.S.W. } a Nee
9 ” ” —vi.1931 we

Lucas.

GALAXAURA GLABRIUSCULA KJELLM. EMEND CHOU.
Known from Tahiti, Hawaiian Islands, Java and the Bonin Islands.

Locality. Date. Herbarium. Notes. .
Coral Reef, Green Is., off Cairns, 30.viii.1936 V.M. & N.Q. Nats. Coll. by H.”
N. Qld. No. 3519 Flecker.
Arkhurst Reef, Hayman Is., Qld. —.viii.1946 V.M. No. 2283
also SYD.
Cape Tribulation, 30 m. south of 18.xi.1947 V.M. & N.Q. Nats.
Cooktown, Qld. No. 12117

GALAXAURA ANGUSTIFRONS KJELLM. EMEND CHOU.

This species is known from Brazil and Java, but the present is the first record of it
from Australia. The species has most likely been treated in Australia as Brachycladia
marginata, under which name G. arborea and G. spathulata also have been included.
Chou has illustrated and described these three species, and suggests that G. intermedia
and G. spathulata may be the corresponding sexual and non-sexual phases of the same
plant. It seems likely that G. angustifrons and G. arborea likewise correspond.

Locality. Date. Herbarium. Notes.
Collaroy, nr. Sydney, N.S.W. 19.xii.1944 V.M. No. 460 Drift.
5 ” es 7.vii.1944 V.M. No. 271 oH
Newport a a 16.xi.1944 V.M. No. 459 Growing in pool

under over-
i hanging rock.
Barrenjoey Hd., Palm Beach, N.S.W. = 25.11.1945 V.M. No. 513 Below L.T.L.

HYPNEA CENOMYCE J. Ag.
New Record for North and Hast Australia.

Previously known from Western Australia and Tasmania in Australia and from
Lord Howe Island, the present is the first record of this species from N.S.W. or
Queensland.

Dr. Tore Levring kindly checked my specimens No. 918 and 674 with original
Agardh material at Lund, Sweden, and he reports the identification as correct.

This species has a more compact habit than other local species ‘of this genus; it is
relatively coarse, much branched towards its apices and it occurs on ocean headlands,
while our common Hypneas are estuarine in habitat.

This is the species of Hypnea referred to (May, 1948, p. 35) as having been confused
sometimes with a species of Gracilaria (see below).

Locality. Date. Herbarium. Notes.
Rock Island, Long Reef, nr. 8.x1i.1944 V.M. No. 493 Sheet B is tetra
Sydney, N.S.W. sporic.
Far out rocks, Newport to Bilgola
Headland, nr. Sydney, N.S.W. 3.x1.1945 V.M. No. 918
Little Beach, Narrabeen, nr. Sydney,
N.S.W. 13.xii.1947 V.M. No. 2484

Extra L.T. Level, Whale Beach 29,xii,1947 V.M, No. 2501 Few tetraspores,
Headland, nr, Sydney, N,S,W,

BY VALERIE MAY. 87

Locality. Date. Herbarium. Notes.
Amity Beach, Stradbroke Is., 1.xi1.1943 V.M. No. 674 Tetrasporic.
Moreton Bay, Qld.

Green Is., N. Qld. 23.viii.1936 V.M. & N.Q. Nats. Coll. by A. B.

; No. 2191 Cumings.
Tetrasporic.

, Double Island Reef, N. Qld. 14.ix.1947 V.M. & N.Q, Nats. Coll. by H.

No. 11448 Flecker.

1 ScD Bud ea eT Tetrasporic.

—.vi.1931 A.H.S.L. Coll. by F. Perrin and
A. H. S. Lucas. Speci-
men labelled ‘“Graci-
laria’’.

Michaelmas Cay, off Cairns, Qld. —.vi.1926 A.H.S.L. Coll. by T. Iredale and

G. Whitley. Specimen

recorded (Lucas, 1927,

p. 155) as Gracilaria

taenioides J. Ag.

Low Island, N. Qld.

HYPNEA CERVICORNIS J. AG.
New Record for Western Australia.

This species is described and illustrated by Tanaka (1941, p. 240-2, fig. 13); it
differs from our common H. Valentiae (Turn.) Mont. in the absence of a persistent main
axis and in the great density of the lateral spinous branches. This species is wide-
spread in the warm Pacific and is already recorded from Queensland; the present
records extend its known range in Australia to Western Australia also.

Locality. Date. Herbarium. Notes.
Geraldton, W.A. —.vii.1928 N.S.W. Coll. by A. H. S.
‘ ; Lueas.
Rottnest Is., W.A. —.viii.1928 N.S.W. 3
On beach, North Is., Abrolhos 6.x11.1945 V.M. No. 2230 Coll. by G. P.
Group, W.A. Whitley.
Tetrasporic.
References.

AGARDH, J. G., 1889.—Species Sargassorum Australiae. Kongl. Svenska Vetenskaps-Akad., 23
(3): 1-183; Tab. 1-31. ;
CHou, RuTH CHEN-YING, 1944——Pacific Species of Galaxaura 1. Asexual Types. Pap. Mich.
Acad. Sci. Arts Lett., 30: 35-55, Pl. 1-11.
, 1945.—Pacific Species of Galaxaura 2. Sexual Types. Pap. Mich. Acad. Sci. Arts Lett.,
31: 3-24, Pl. 1-13.
Howe, M. A., 1909.—Phycological Studies. IV. The genus Neomeris and Notes on other
Siphonales. Bull. Torrey Bot. Club, 36: 75-104, Pl. 1-8. ;
Lueas, A. H. S., 1913.—Notes on Australian Marine Algae. I. Proc. Linn. Soe. N.S.W., 38
(1): 49-60, Pl. 1-5.
, 1927.—Notes on Australian Marine Algae. V. Proc. Linn. Soc. N.S.W., 52 (4):

555-562.
, 1934.—Notes on Australian Marine Algae. VII. Proc. LINN. Soc. N.S.W., 59 (5-6):

348-350.
May, VALERIE, 1946.—Studies on Australian Marine Algae. III. Proc. LINN. Soc. N.S.W., 71

(5-6) : 273-277, Pl. 19.
1948.—The Algal Genus Gracilaria in Australia. O.8.I.R. Bull. No. 235: 1-64, Pl.

1-15.

Osgporn, J. E. M., 1947.—The Structure and Life History of Hormosira Banksi (Turn.) Decne.
Trans. Roy. Soc. N.Z., 77 (1): 47-71, Pl. 1-3.

SvepELIus, N., 1945.—Critical Notes on Some Species of Galawaura from Ceylon. Ark. f. Bot.,
82 (6): 1-74, Pl. 1-9.

TANAKA, TAKESI, 1941.—The genus Hypnea from Japan. Sci. Pap. Inst. Algol. Research. Fac.
Sci. Hokkaido Imp. Univ., 2 (2): 227-250, Pl. 53-54.

io 2)
oo

THE MARINE ALGAE OF BRAMPTON ISLAND, GREAT BARRIER REEF,
OFF MACKAY, QUEENSLAND.

By VALERIE May, M.Sc.,
National Herbarium of New South Wales, Sydney, N.S.W.
[Read 27th June, 1951.]

Synopsis.

This paper describes the marine algal flora of Brampton Island. This flora was found to
be very rich and to include a number of species new to Queensland, and some new to Australia.

A comparison is made between the results obtained and other algal records from N.E.
Australia and also between the floras of the various areas of the island.

INTRODUCTION.

There are relatively few collections of, or studies on, the marine algae of any part
of the Great Barrier Reef of Australia. We have the records of the Great Barrier Reef
Expedition of 1928-1929, which include Lucas’s summary of algal species hitherto
recorded from N.E. Australia (1931) and the report by Stephenson and others (1931)
which includes a study of the ecology of the algae of the Low Islands. We have also
the detailed lists given by Lucas (1934) of the algae of the Low Islands. Other than
these the records are more fragmentary, the most detailed being by Sonder (1871).

The present paper presents the results of a collection made at Brampton Island,
in the Great Barrier Reef, off Mackay, Queensland, by the author during June, 1948
(specimens numbered 2660-2815), and also of a second collection from the same locality
made by members of the Sydney University Biological Society in January, 1949 (speci-
mens No. A-H). This second collection yielded two additional species (Sargassum
carpophyllum and Goniolithon ? Fosliei), while one other species (Sargassum flavicans),
which had been found by the author only as drift material, was found growing attached.
Obviously further collections at other seasons of the year are likely to increase the
number of species known from Brampton Island.

The marine algal flora of Brampton Island appears to be exceptionally varied. It
includes 85 species, aS compared with 74 from the Low Islands (Lucas, 1934), and 205
hitherto recorded (Lucas, 1931) from N.E. Australia. This list from N.E. Australia,
however, does not include the Myxophyceae, and is anyway not complete; for instance,
records by Grunow (1874) and even by Lucas himself (1927) are omitted. Recent
additions to the records of species occurring in Queensland have been made by the
present author (1946, 1948, 1949, 1951).

It is of interest to note that while the Low Islands are richer than is Brampton
Island in the number of species of Chlorophyceae recorded, they are poorer in the
number of species of Rhodophyceae recorded.

In the algae of Brampton Island the following species appear to be recorded from
Australia for the first time: Cladophoropsis ? membranacea, Vaughaniella rupicola,
Brachytrichia Quoyi, Calothrix pilosa, Sirocoleum guyanense, Galaxaura subfruticulosa,
Dasya pacifica, Endosiphonia spinuligera, Amphiroa prob. foliacea.

The following species have been recorded from other parts of Australia but are
not previously known from Queensland: Caulerpa papillosa, Entophysalis conferta,
Ceramium gracillimum, Polysiphonia implexa, Polysiphonia zostericola.

Brampton Island is a continental island with a tide range of about 15 feet. For
ecological comparison the sand and mud flats extending about the island have been
treated in this paper collectively as “Flats”, while the main reef developed of one side
of the island is designated “Reef”. One rocky headland—Pelican Island—appears
detached from the main island at high tide, but is linked by rock-strewn flats at low

BY VALERIE MAY. 89

tide; algae from this general location have been treated in this paper as from the
“Pelican Island Flats’. Carlyle Island also becomes land-linked to Brampton at
extremely low tides, and since a coral reef from this adjoining island was particularly
well developed, the collection from it has also been included in the present paper;
algae from this location are marked as from “Coral Reef”.

Apart from these four above described areas, two species were collected encrusting
rocks at high-tide level, and some drift material was also collected. These collections
are so labelled in the paper.

For a comparison of the floras of the various ecological areas the accompanying
table (Table 1) of species found in each location is presented. However, it is emphasized

TABLE 1.
The Species Collected from the Different Areas.

| Encrusting) Found
Flats. | Reef. Coral Reef. | Pelican Is. Flats. | Rocks at | Only as

| HTL. | Drift.
| |
CHLOROPHYCHAE.
Ulva lactuca. | |
Enteromorpha sp. Anadyomene brownii. | Valoniopsis pachy- | |
| nema.
Dictyosphaeria Ver- | D. favulosa. | D. favulosa.
sluysi. |
Struvea delicatula. | S. delicatula. S. delicatula.
| Boergesenia Forbesii.
| Cladophoropsis ? ae Bornetella oligospora.
| membranacea. |
Spongocladia vaucher- | S. vaucheriaeformis-
iaeformis.
Udotea flabellum. | U. flabellum.
| U. orientalis. U. orientalis.
Halimeda macroloba. | H. macroloba.
H. incrassata. | H. incrassata.
HA. opuntia.

| Codium spongiosum.
Caulerpasertularioides.
C. serrulata. C. serrulata. |

C. racemosa. C. racemosa.
C. papillosa.

No. of species :
11 13 f 4 3 0 0

PHAEOPHYCEAE. ©
Ectocarpus confer- | E. confervoides. E. confervoides.
voides.
Sphacelaria tribul-
oides.
Pocockiella nigrescens.
Stypopodium zonale.
Dictyota dichotoma.
Dictyopteris pardalis. D. pardalis.
Padina Commersonit. P. Commersonii. P. Commersonii.
Vaughaniella rupicola.
Hydroclathrus | H. clathratus.
clathratus.

Chnoospora obtus-
angula.
Cystophyllum = muri-
catum.
Hormophysa triquetra.
Sargassum scabripes. S. carpophyllum., S. lopho-
carpum.
S. flavicans.

Turbinaria ornata.

No. of species :
5 8 1 8 0 | 1

L

90

THE MARINE ALGAE OF BRAMPTON ISLAND,

TABLE 1.—Continued.

The Species Collected from the Different Areas Continued.

Flats.

MYXOPHYCEAE.

Brachytrichia Quoyt.

Lyngbya majuseula.
No. of species :
)

a

RHODOPHYCEAE.

Liagora ceranoides.
Galaxaura glabriuscula.

Desmia Kilneri.

Metagonolithon grani-
ferum.
Jania rubens.

Hypnea Valentiae.

Corallopsis minor.
C. Urvillei.
Ceratodictyon
giosum.
Champia parvula.
Ceramium gracillimum.
Centroceras clavulatum.

spon-

Dasya pacifica.

Polysiphonia implexa.

P. zostericola.

Digenea simplex.

Roschera glomerulata.

Endosiphonia spinu-
ligera.

Leveillea jungerman-
nioides.

Laurencia rigida.
L. obtusa.
Acanthophora
spicifera.
Chondria sp.
No. of species :
23

Total no. of species :
41

Reef.

Hydrocoleum lyng-
byaceum.

Sirocoleumguyanense.

bo

L. ceranoides.
Gelidiella acerosa.

Peyssonnelia Gun-

niana.

J. rubens.
Sarconema filiforme.
Eucheumamuricatum.

H. Valentiae.
H. cervicornis.

Spyridia filamentosa.

Dasya sp.

D. simplex.

EB. spinuligera.

LD. jungermannioides.
Amansia glomerata.

L. obtusa.

15

38

CC er ————————————————SeSSSSsSsSseseseseseseseb

Coral Reef.

G. subfruticulosa.
G. acerosa.

Mastophora plana.
Amphiroa prob.
foliacea.

Sebdenia ceylonica.

Plocamium hamatum.
Gracilaria lichenoides.

Dasya sp.
Dasya sp.

A. glomerata.

L. obtusa.

Chondria sp.

12

17

| Pelican Is. Flats.

solutum.

|

| L. majuseula.

2

P. Gunniana.

Goniolithon ? Fosliei.
Fosliella farinosa.

G. lichenoides.

D. pacifica.

D. simplex.
R. glomerulata.

Encrusting

| Hormothamnion

Rocks at |
1S tal BSI UE,

Ento-
physalis |

| conferta.

Calothrix
pilosa,

bo

Found
Only as
Drift.

BY VALERIE MAY. 91

that these figures are only approximate, the main purpose during the collecting being
to get a representative specimen of every species then growing at Brampton Island,
and only secondarily of all species in each location. The outstanding observation from
this table is the high percentage incidence of species of Chlorophyceae on the Reet
(34% of species occurring there), of Phaeophyceae in the Pelican Island Flats area
(40% of species occurring there) and of Rhodophyceae in the extreme Coral Reef
region (71% of species occurring there).

My sincere thanks are due to Dr. F. Drouet, of the Chicago Natural History Museum,
who kindly identified all the Myxophyceae recorded in the present paper.

CHLOROPHYCEAHE.
ULVALES.
Family ULVACEAE.
Utva Linnaeus.
ULvA LAcTUCA I, 1753, p. 11638; Borgs., 19389, pp. 57-58.
No. 2755, Flats.
Geogr. Distr.: Widespread. N., S. and E. Australia.

ENTEROMORPHA Link.
ENTEROMORPHA Sp.
Epiphytic on Cystophyllum muricatum. +
No. 2723, Flats.
SIPHONOCLADALES.
Family ANADYOMENACEAE.

ANADYOMENE Lamouroux.

ANADYOMENE BROWNIE (Gray) J. Ag., 1886, p. 127; De Toni, 1889, p. 370.—Calonema
Brownii Gray, 1866, p. 46, t. 44, f. 3.

No. 2786, 2787, Reef.
Geogr. Distr.: Celebes. N.E. Australia.

VALONIOPSIS Borgesen.

VALONIOPSIS PACHYNEMA (Martens) Borgs., 1934, pp. 10-17.—Bryopsis pachynema
Martens, 1866, p. 24, Pl. 4, fig. 2—Valonia confervoides Harv. Alg. Ceyl. Exsicc. No. 73;.
Alg. Friendly Is. Exsicc. No. 101; in J. Ag., 1886, p. 100; De Toni, 1889, p. 378.

No. 2769, Coral Reef.

Geogr. Distr.: Warm Atlantic and Indian Oceans, Ceylon, Friendly Is., Hawaiian
Archipelago. Lord Howe Is. N. and H. Australia.

Family VALONIACEAE.
DICTYOSPHAERIA Decaisne.

1. DICTYOSPHAERIA FAVULOSA Decne., 1842, p. 32; De Toni, 1889, p. 371.
No. 2790, Reef; D., Pelican Is. Flats.
Geogr. Distr.: Mexico, Red Sea, warm Indian and Pacific Oceans. N. Australia.
2. DiIcTYOSPHAERIA VERSLUYSI W.v.B., 1905, p. 114.
No. 2692, 2734, Flats.
Geogr. Distr.: Malayan Archipelago, Mexico. N.E. Australia.

Family SIPHONOCLADACEAE.
STRUVEA Sonder.
STRUVEA DELICATULA Kuetz., 1849-1869, t. 2, f. 2; De Toni, 1889, p. 366.
No. 2706, Flats; 2766, Coral Reef; 2793, Reef.
Geogr. Distr.: Ceylon, Island of Guadeloupe, Mexico. W. and N.E. Australia.

BOERGESENIA Feldmann.

BOERGESENIA FoRBESII (Harv.) Feldm., 1938, p. 18, figs. 8-5; Borgs., 1948, p. 21--22.—
Valonia Forbesii Harv., Alg. Ceylon Exsicc. No. 75; Friendly Islands Algae Exsice. No.
102; in J. Ag., 1886, p. 96; De Toni, 1889, p. 374.

No. 2676, 2782, Reef.

Geogr. Distr.: Red Sea, Indian and Pacific Oceans. N.E. Australia.

92 THE MARINE ALGAE OF BRAMPTON ISLAND,

CLADOPHOROPSIS Borgesen.

SLADOPHOROPSIS (?) MEMBRANACEA (Ag.) Borgs., 1905, p. 288-289, figs. 8-13.—
Cladophora membranacea (Ag.) Kuetz., 1849, p. 415.—Siphonocladus membranaceus
(Ag.) Born. in Hariot, 1887, p. 56; De Toni, 1889, pp. 358-359.—Conferva membranacea
Ag., 1824, p. 120.

No. 2797, Reef.

Geogr. Distr.: Antilles and Santa Cruz, New Zealand. Probable new record to
Australia.

SPONGOCLADIA Areschoug.

SPONGOCLADIA VAUCHERIAEFORMIS Aresch., 1853, p. 201; De Toni, 1889, p. 360.

No. 2675, Reef; 2724, Flats; 2777, Reef. ‘

Geogr. Distr.: Mauritius, Malayan Archipelago, New Guinea, Lord Howe Is., ete.
N.E. Australia.

DASYCLADALES.
Family DASYCLADACEAE.
BoRNETELLA Munier-Chalmas.

BORNETELLA OLIGOSPORA Solms-Laubach, 1893, p. 87, tab. 9, figs. 1-4, 6, 7.

No. 2800, Pelican Is. Flats.

Geogr. Distr.: Malayan Archipelago, New Guinea. N.H. Australia.

SIPHONALES.
Family CopIACEAE.
Uporea Lamouroux.

1. UDOTEA ORIENTALIS A. and EH. S. Gepp, 1911, pp. 119-120, Pl. 1, figs. 1, 4, Pl. 6,
figs. 47-48.

No. 2679, 2788, Reef: 2811, Pelican Is. Flats.

Geogr. Distr.: Indian and Pacific Oceans. N.EH. Australia.

2. UDOTEA FLABELLUM (Hllis and Sol.) Lamour., 1812, p. 186; Pap., 1944, pp. 337—338.—
Corallina Flabellum Ellis and Sol., 1786, p. 124, tab. 24, figs. A-C.

A. and E. S. Gepp (1911, pp. 131-133, Pl. 3, figs. 26-28) describe and figure this
plant under the name U. flabellum (Ellis and Sol.) Howe.

No. 2678, Reef; 2728, Flats.

Geogr. Distr.: Atlantic Ocean, Red Sea, Madras, Ceylon, Friendly Islands. N.E.
Australia.

HALIMEDA Lamouroux.

1. HALIMEDA OpuNTIA (L.) Lamour. emend. Bart., 1901, p. 18; Lamour., 1812, p.
186.—Corallina Opuntia L., 1765, p. 805 (in part).

No. 2677, Reef.

Geogr. Distr.: Widespread in warm seas. N. Australia.

2. HALIMEDA MACROLOBA (Decne.) Bart., 1901, p. 24; Decne., 1841, p. 118.

_ No. 2693, Flats; 2776, Reef.

Geogr. Distr.: Warm Indian and Pacific Oceans, Red Sea. W. and N. Australia.

3. HALIMEDA INCRASSATA (Ellis) Lamour. emend. Bart., 1901, p. 25; Lamour., 1812,
p. 186.—Corallina incrassata Hllis, 1755, p. 53, tab. 25, f. A.

No. 2713, Reef; 2721, Flats.

Geogr. Distr.: Atlantic, Indian and Pacific Oceans, W. Indies, Madagascar, Malayan
Archipelago, China Sea, Friendly Islands, ete. N.E. Australia.

Copium Stackhouse.
CopIUM SPONGIOSUM Harv., 1855, p. 565; De Toni, 1889, pp. 489-490.
No. 2667, drift; 2669, Reef.
Geogr. Distr.: W.N. and E. Australia.

Family CAULERPACEAE.
CAULERPA Lamouroux.
1. CAULERPA SERTULARIOIDES (Gmel.) Howe, 1905, p. 576.—Fucus sertularioides
Gmelin, 1768, tab. 15, fig. 4—Caulerpa plumaris (Forsk.) W.v.B., 1898, pp. 294-296.

BY VALERIE MAY. 93

No. 2727, Flats.
Geogr. Distr.: Tropical seas generally. N.E. and S. Australia.

2. CAULERPA SERRULATA (Forsk.) J. Ag. emend. Borgs., 1932; J. Ag. 1872, p. 19.—
Fucus serrulatus Forsk., 1775, p. 188—Caulerpa freycineti Ag., 1823-1828, p. 446;
W.v.B., 1898, pp. 310-318, Pl. 25, figs. 4-11, Pl. 26, figs. 1-6.

Var. DE BoryAana (Ag.) W.v.B. (loc. cit., pp. 315-316, Pl. 25, figs. 10-11).

No. 2703, Flats; 2711, Reef.

Geogr. Distr.: Red Sea, Gaudeloupe. N.H. Australia.

9

3. CAULERPA RACEMOSA (Forsk.) J. Ag. emend. W.v.B., 1898, pp. 357-373, Pl. 31, figs.
5-8, Pl. 32, figs. 1-7, and Pl. 33; J. Ag., 1872, p. 35, n. 51—Fucus racemosus Forsk., 1775,
joy JEL

Var. LAETEVIRENS (Mont.) W.v.B. (loc. cit., pp. 366-368, Pl. 33, fig. 8).

No. 2725, 2726, Flats.

Geogr. Distr.: Is. of Toud, Ceylon, Florida, Gaudeloupe. W. and N.E. Australia.

? var. NUMMULARIA (Harv.) W.v.B. (loc. cit., p. 376). A very small specimen.

No. 2768, Coral Reef.

Geogr. Distr.: Friendly Islands, etc. N.H. Australia.

4. CAULERPA PAPILLOSA J. Ag., 1872, p. 42, n. 61; W.v.B., 1898, pp. 383-384, Pl. 34,
fig. 8—An unexpected species.

No. 2764, Coral Reef.

Geogr. Distr.: W. and S. Australia.

PHAHOPHYCHAE.
ISOGENERATAE.
E\CTOCARPALES.
Family EcrocArRPpACcKEAE.
Ecrocareus Lyngbye.
ECTOCARPUS CONFERVOIDES (Roth.) Le Jol., 1863, p. 75; May, 1939, pp. 537-554.—
Ceramium confervoides Roth., 1797, pp. 151-152, Pl. 8, fig. 3.
Plurilocular reproductive structures do not bear hairs at their apices in these
specimens. —
No. 2759, Coral Reef; 2779, Reef; F., Pelican Is. Flats.
Geogr. Distr.: Widespread. All around Australia.

SPHACELARIALES.
Family SPHACELARIACEAE.
SPHACELARIA Lyngbye.
SPHACELARIA TRIBULOIDES Menegh., 1840, p. 2, n. 1; De Toni, 1895, pp. 502-503.
No. 2809, Pelican Is. Flats.
Geogr. Distr.: Mediterranean, Adriatic and Red Sea, Atlantic Ocean, Hawaii, ete.
W. and N. Australia.

DICTYOTALES.
Family DICTYOTACEAE.
POCOCKIELLA Papenfuss.
POCOCKIELLA NIGRESCENS (Sond.) Pap., 1943, p. 467—Zonaria nigrescens Sond., 1845,
Dp. 50.—Gymnosorus nigrescens (Sond.) J. Ag., 1894, p. 12; De Toni, 1895, pp. 228-229.
No. 2665, drift; 2807, Pelican Is. Flats.
Geogr. Distr.: N., S., EH. and W. Australia.

StyPoropiIum Kuetzing.
STYPOPODIUM ZONALE (Lamour.) Pap., 1940, pp. 205-206—Fucus zonalis Lamour.,
UPKU, Bis JEG 24H, wes, al
My grateful thanks are due to Miss C. I. Dickinson, of Reyal Botanic Gardens, Kew,
England, who has kindly compared my material with other specimens of this species.
No. 2789, Reef.
Geogr. Distr.: W. Indies, S. Africa, Japan, N.HE. Australia.

94 THE MARINE ALGAE OF BRAMPTON ISLAND,
DictyorA Lamouroux.
DicryoTA DICHOTOMA (Huds.) Lamour., 1809, p. 42; De Toni, 1895, pp. 263-264.—:

Ulva dichotoma Huds., 1778, p. 476.
Var. INTRICATA (Ag.) Grev., 1830, p. 58.
No. 2814, Pelican Is. Flats.
Geogr. Distr.: Widespread. N. and E. Australia and Tasmania.

DIcTYOPTERIS Lamouroux.
1947, p. 274.—Haliseris pardalis Harv. in

DICTYOPTERIS PARDALIS (Harv.) May,
Kuetz., 1849-1869 (Part 9), p. 24, t. 59, f. 2; De Toni, 1895, p. 258.

No. 2729, Flats; 2802, Pelican Is. Flats.

Geogr. Distr.: W., N. and EH. Australia.

PapInA Adanson.
De Toni, 1895, p. 244.---As

PADINA COMMERSONIIT Bory, 1828, n. 41, t. 21, f. 2;
Harvey’s Alg. Ceylon Exsice. No. 55 sub nomine P. Pavoniae.

No. 2715, Reef; 2750, Flats; C., Pelican Is. Flats.
Geogr. Distr.: Mauritius, Ceylon, Malaya, Japan, Friendly Is., etc. N., W. and S.

Australia.
VAUGHANIELLA Borgesen.

VAUGHANIELLA RUPICOLA Borgs., 1950, pp. 1-10, figs. 1-8.
No. 2751, Flats. On Padina sp.
Geogr. Distr.: Mauritius. New record to Australia.
HETEROGENERATAE.
PUNCTARIALES.
Family HNCOELIACEAE.

HyYDROCLATHRUS Bory.
Setchell and Gardner,

HYDROCLATHRUS CLATHRATUS (Bory) Howe, 1920, p. 590;
1925, p. 543.—Fucus clathratus Bory in Ag., 1823-1828.

Nos. 2683, 2722, Flats; 2778, Reef.
Geogr. Distr.: Widespread in tropical Atlantic and Pacific Oceans, Mediterranean

N., S., E. and W. Australia.

Cunoospora J. Agardh.
CHNOOSPORA OBTUSANGULA (Harv.) Sond., 1871, p. 45;. De Toni, 1895, p. 465.—

and Red Seas.

Dictyota obtusangula Harv., 1859, p. 329, n. 14.
As Harvey’s Friendly Island Algae Exsice. No. 4.

Nos. 2714, 2795, Reef.
Geogr. Distr.: Atlantic Ocean, Japan, Friendly Islands. N.E. Australia.
CYCLOSPOREFAE.
FUCALES.
Family SARGASSACEAE.
CYSTOPHYLLUM J. Agardh.
CYSTOPHYLLUM MURICATUM (Turn.) J. Ag., 1848, p. 231; De Toni, 1895, p. 154.—

Fucus muricatus Turn., 1808-1819 (vol. 2), p. 108, tab. 112.
Prevalent, but by accident there remains only a poor specimen.

No. 2723, Flats.
Geogr. Distr.: India, Admiralty Islands, ete. All around Australia.

HormMopHysa Kuetzing.
HORMOPHYSA TRIQUETRA (L.) Kuetz., 1843, p. 359; Borgs., 1939, p. 96—Fucus triqueter

L., 1771, p. 312.—-Cystoseira triquetra (L.) J. Ag., 1848, p. 215.
Osborn (1948, p. 48) records that the Australian plant known as Hormosira ?

articulata (Forsk.) Zan. is in fact Hormophysa triquetra.

No. 2664, drift; 2712, Reef.

Geogr. Distr.: India, Red Sea. W., N. and E. Australia.

BY VALERIE MAY. 95

Sargassum Agardh.

1. SARGASSUM SCABRIPES J. Ag., 1872, p. 52; 1889, p. 48, tab. 2.

No. 2660, Reef; 2661, drift.

Geogr. Distr.: N. Australia.

2. SARGASSUM CARPOPHYLLUM J. Ag., 1848, p. 304; 1889, p. 82, tae 25 (2).

No. A., Pelican Is. Flats.

Geogr. Distr.: Indian Ocean. N. and W. Australia.

38. SARGASSUM FLAVICANS (Mert.) Ag., 1823-1828, p. 18; J. Ag., 1889, pp. 82-83,
tab. 25 (3).— Fucus flavicans Mert., 1819, p. 8. .

No. 26638, drift; B., Pelican Is. Flats.

Geogr. Distr.: W., N. and EH. Australia.

4, SARGASSUM LOPHOCARPUM J. Ag., 1889, p. 93, tab. 27 (2).

No. 2662, drift.

Geogr. Distr.: N., S., H. and W. Australia.

TURBINARIA Lamouroux.
TURBINARIA ORNATA J. Ag., 1848, p. 266; De Toni, 1895, p. 128.—Fucus turbinatus
var. ornatus Turn., 1809-1819 (vol. 1), p. 50, tab. 24, figs. c—h.

No. 2668, Reef.
Geogr. Distr.:. Ceylon, New Zealand, Andaman Islands, Friendly Islands, Admiralty

Islands, Samoa, etc. N. Australia.

MY XOPHYCEAE.
Family CHAMAESIPHONACEAE.
ENTOPHYSALIS Kuetzing.
ENTOPHYSALIS CONFERTA (Kuetz.) Dr. and Dailey, 1948, p. 79.—Palmella conferta
Kuetz., 1845, p. 149.
No. 2701, encrusting rocks at high tide level.
Geogr. Distr.: Widespread. HE. Australia.

Family STIGONEMATACEAE.
BRACHYTRICHIA (Zanardini) Bornet and Flahault.
BRACHYTRICHIA Quoy1 (Ag.) Born. and Flah., (Part 2) 1886-1888, p. 73.—Nostoc
Quoyi Ag., 1824, p. 22.
Nos. 2736, 2757, Flats.
Geogr. Distr.: Widespread. New record to Australia.

Family NOSTOCACEAE.
HORMOTHAMNION (Grunow) Bornet and Flahault.
HORMOTHAMNION SOLUTUM Born. and Grun. ex Born. and Flah. .. (Part 4) 1886-1888,
p. 259.
Nos. 2803, 2813, Pelican Is. Flats.
Geogr. Distr.:. Widespread. W. and N.H. Australia.

Family RIVULARIACEAE.
CALoTHRIx (Agardh) Bornet and Flahault.
CALOTHRIX PILOSA Harv. ex Born. and Flah., (Part 1) 1886-1888, p. 363.
No. 2701, encrusting rocks at high tide level.
Geogr. Distr.: Red Sea, Mauritius, Friendly Is., Jamaica, Mexico, Brazil, etc. New
record to Australia.
Family OSCILLATORIACEAE.
HyprocoLeEuM (Kuetzing) Gomont.

HYDROCOLEUM LYNGBYACEUM Kuetz. ex Gom., (Part 15) 1892, p. 337, Pl. 12, f. 8, 9, 10.
No. 2671, Reef.
Geogr. Distr.: Widespread. N.E. Australia.

96 THE MARINE ALGAE OF BRAMPTON ISLAND,

SrROocOLEUM (Kuetzing) Gomont.
SIROCOLEUM GUYANENSE Kuetz. ex Gom., (Part 15) 1892, p. 348.
No. 2671, Reef.

Geogr. Distr.: Widespread. New record to Australia.
A

LyneByAa (Agardh) Gomont.
LYNGBYA MAJUSCULA Gom., (Part 16) 1892, p. 131, Pl. 3, figs. 3 and 4.
No. 2742, growing on Digenea simplez.
Nos. 2707, 2735, 2742, Flats; 2812, Pelican Is. Flats.
Geogr. Distr.: Widespread. N.E. Australia.

RHODOPHYCEAE.
FLORIDEAE.
NEMALIONALES.
Family HELMINTHOCLADIACEAE.
Liacora Lamouroux.
LIAGORA CERANOIDES Lamour., 1816, p. 239; Borg., 1939, p. 104.—Liagora leprosa J. Ag.
1847, p. 8.
No. 2697, Flats; 2716, 2796, Reef.
Geogr. Distr.: West Indies, Red Sea, Mauritius, Indian Ocean, Malayan Archipelago,
Japan, etc. N.E. Australia.

Family CHAETANGIACEAE,
GALAXAURA Lamouroux.
1. GALAXAURA SUBFRUTICULOSA Chou, 1944, pp. 41-44, Pl. 2, fig. 6, Pl. 8, fig. 2.
In this specimen the tumid cell is very large, usually being half as wide again as
the terminal cell; Chou reports the size of this cell is variable.
No. 2763, Coral Reef.
Geogr. Distr.: Mexico, prob. Japan and China. New record to Australia.
2. GALAXAURA GLABRIUSCULA Kjellm. emend. Chou, 1945, pp. 11-13, Pl. 4, figs. 14-20,
Pl. 10, fig. 1; Kjellm., 1900, pp. 56-77, Pl. 7, figs. 1-2, Pl. 20, fig. 26.
No. 2710, Flats.
Geogr. Distr.: Tahiti, Hawaii, Bonin Islands, Java. N.E. Australia.

GELIDIALES.
Family GELIDIACEAE.
GELIDIELLA Feldmann and Hamel.
GELIDIELLA ACEROSA (Forsk.) Feldm. and Hamel, 1934, t. 46, p. 533; Borg., 1939,
p. 107.—Fucus acerosus Forsk., 1775, p. 190.
No. 2771, Coral Reef; 2780, Reef.
Geogr. Distr.: Warm seas generally. N.E. Australia.

CRYPTONEMIALES.
Family RHIZOPHYLLIDACEAE.
Desmi1A Lyngbye.
DESMIA KiLNeERI J. Ag., 1876, p. 355—Chondrococcus Kilneri (J. Ag.) De Toni, 1897-
1905 (pt. 4), p. 1676.
Papenfuss (1940, p. 218) discusses the use of the generic name Desmia in preference
to Chondrococcus.
No. 2704, Flats.
Geogr. Distr.: N. Australia.

Family SQUAMARIACEAE.
PEYSSONNELIA Decaisne.
PEYSSONNELIA GUNNIANA J. Ag., 1876, p. 387; De Toni, 1924, p. 591.
Adherent to undersides of rocks.
No. 2673, Reef; 2806, Pelican Is. Flats.
Geogr. Distr.: Malayan Archipelago. N., S. and E. Australia, Tasmania.

BY VALERIE MAY. 97

Family CoORALLINACEAE.
GONIOLITHON Foslie.

GoNIOLITHON ? FosxLier (Heydr.) Fosl., 1903, p. 470, t. 25, f. 3; W.v.B. and Foslie,
1904, pp. 46-48, fig. 19, Pl. 9, figs. 1-5.—Lithothamnion Fosliei Heydr., 1897, p. 58 ex
parte.

Insufficient material for sure determination.

No. H., Pelican Is. Flats.

Geogr. Distr.: Maldive and Laccadive Islands, Red Sea, Zanzibar, Malayan Archi-
pelago. Murray Island, N. Australia.

FOSLIELLA Howe.
FOSLIELLA FARINOSA (Lamour.) Howe, 1920—Melobesia farinosa Lamour., 1816,
p. 315, tab. 12, fig. 3.
Specimen epiphytic on Sargassum sp.
No. 2815, Pelican Is. Flats.
Geogr. Distr.: Widespread. N., S., HE. and W. Australia.

MAsSToPHORA Decaisne.
MASTOPHORA PLANA (Sond.) Harv., 1847, p. 108; De Toni, 1924, p. 695.—Melobesia
plana Sond., 1845, p. 55.
No. 2758, Coral Reef.
Geogr. Distr.: Marianas Is. W. and N.E. Australia.

AMPHIROA Lamouroux.
AMPHIROA prob. FOLIACEA Lamour., in Freyc., 1824-1826, p. 628, t. 93, f. 2-31; W.v.B.
and Foslie, 1904, pp. 92-93, tab. 14, figs. 1-11.
A poor specimen, insufficient for definite determination. The plant bears a minute
epiphyte, Dasya sp. (No. 2772).
No. 2773, Coral Reef.
Geogr. Distr.: India, Malaya, Marianas. Not previously recorded from Australia.

METAGONIOLITHON Weber van Bosse.

METAGONIOLITHON GRANIFERUM (Hary.) W.v.B. and Foslie, 1904, p. 103, tab. 15, figs.
10, 12; De Toni, 1924, p. 704—Amphiroa granifera Harv. Syn. in 1858-1863, p. xxx,
n. 362.

No. 2691, Flats.

Geogr. Distr.: Peru. W., N., S. and E. Australia.

JANIA Lamouroux.
JANIA RUBENS (L.) Lamour., 1816, p. 272; De Toni, 1924, p. 709—Corallina rubens L.,
1765, p. 1305.
As Harvey’s Alg. Exsice. Friendly Is. No. 30.

Nos. 2681, 2781, Reef; 2702, Flats; 2741, Flats as epiphyte on Digenea simplex
(No. 2739).

Geogr. Distr.: Temperate and warm seas, Lord Howe Island. N., E. and S. Australia.

GIGARTINALES.
Family SEBDENIACEAE.
SEBDENIA Berthold.
SEBDENIA CEYLONICA (Harv.) Heydr., 1892, p. 477, t. 26, f. 16-17; De Toni, 1924, p.

297.—Halymena ceylonica Harv., Alg. Ceyl. Exsice. No. 39; in Kuetz. 1849-1869 (Part 16),
tis GB

- No. 2760, Coral Reef.
Geogr. Distr.: Red Sea, Ceylon, New Guinea, Samoa. N.E. Australia.

98 THE MARINE ALGAE OF BRAMPTON ISLAND,

Family SOLIERIACEAE.
SARCONEMA Zanardini.
SARCONEMA FILIFORME (Sond.) Kylin, 1932, pp. 22-23.—Dicranema filiforme Sond.,
1845, p. 56.
No. 2799, Reef. :
Geogr. Distr.: All around Australia.

EucHrEumMA J. Agardh.

EUCHEUMA MURICATUM (Gmel.) W.v.B., 1913-1928 (part d.), pp. 413-415, Pl. 12,
figs. 1-5 and fig. 164—Fucus muricatus Gmel., 1768, p. 111, Pl. 6—Huchewma spinosum
(L.) J. Ag., 1852-1863, p. 626; De Toni, 1897-1905 (Part 1), pp. 369-370.

No. 2794, Reef.

Geogr. Distr.: S. Africa, Indian Ccean, Sumatra, New Guinea, Lord Howe Is., Japan.
N. and W. Australia.

Family HyYpPNACEAE.

HypneA Lamouroux.
1. HYPNEA CERVICORNIS J. Ag., 1852-1863, p. 451; Tanaka, 1941, pp. 240-242, fig. 18.
No. 2785, Reef.

Geogr. Distr.: Brazil, Mexico, W. Indies, Atlantic and Indian Oceans. W. and N.E.
Australia.

2. HYPNEA VALENTIAE (Turn.) Mont., 1840, p. 161; Hauck, 1887, p. 20; Borg., 1934a,
pp. 17-18; 1943, pp. 58-59.—Fucus Valentiae Turn., 1808-1819, Pl. 78—Hypnea charoides
Lamour., 1813, p. 44, Pl. 10, figs. 1-3; Tanaka, 1941, pp. 243-244, fig. 16; Borg., 1943,
pp. 56-58 —Hypnea seticulosa J. Ag., 1852-1863.

This species appears to be both prevalent and variable. No. 2680 is very hirsute,
while No. 2696 is nearer H. nidifica J. Ag. Neither specimen bears stellate bulbils, but
other Australian collections have led me to the opinion that the presence or absence
of these bulbils is not of specific importance so I follow Hauck (1887, p. 20) and include
H. charoides among the synonyms of H. Valentiae.

No. 2680, Reef; 2696, Flats, on Cymodocea sp.

Geogr. Distr.: Indian Ocean, Japan, S. Pacific. All around Australia.

Family PLOCAMIACEAE.
PLocAmMiumM (Lamouroux) Lyngbye.
PLOCAMIUM HAMATUM J. Ag., 1876, p. 338; De Toni, 1897-1905 (Part 1), p. 589.
No. 2767, Coral Reef.
Geogr. Distr.: Norfolk Is. N. and EH. Australia.

Family GRACILARIACEAE.
GRACILARIA Greville.
GRACILARIA LICHENOWES (L. in Turn.) Harv., 1844, p. 445; May, 1948, pp. 27-39.--
Fucus lichenoides L. in Turn., 1808-1819, tab. 1184 (excl. var. and synon.).
Specimens are very small, so are not treated in subspecific units.
No. 2775, Coral Reef; 2808, Pelican Is. Flats.
Geogr. Distr.: Hast Indian to Pacific Ocean. N. and E. Australia.

CORALLOPSIS Greville.

1. CORALLOPSIS MINOR (Sond.) J. Ag., 1876, p. 409; De Toni, 1897-1905 (part 1),
p. 459.—Corallopsis salicornia var. minor Sond., 1871, p. 24, t. 3, f. 6-11.

Nos. 2666, 2748, Flats.

Geogr. Distr.: Marianas Islands. N. Australia.

2. CORALLOPSIS Urvi~ier (Mont.) J. Ag., 1852-1863, p. 583; De Toni, 1924, p. 276.--
Hydropuntia Urvillei Mont., 1842, p. 7.

No. 2731, Flats.

Geogr. Distr.: China Sea. N. Australia.

BY VALERIE MAY. 99

CERATODICTYON Zanardini.

CERATODICTYON SPONGIOSUM Zan., 1878, n. 8; De Toni, 1924, p. 243.
Attached to pebbles in mud.

Nos. 2670, 2682, Flats.

Geogr. Distr.: Warm Pacific and Indian Oceans. N.H. Australia.

RHODYMENIALES.
Family CHAMPIACEAE.
CHAMPIA Desveaux.
CHAMPIA PARVULA (Ag.) J. Ag., 1876, p. 303; De Toni, 1924, p. 307.--Chondria purvulu
Ag., 1824, p. 207.
No. 2732, Flats.
Geogr. Distr.: Common in warm seas. All around Australia.

CERAMIALES.
Family CERAMIACEAE.
CERAMIUM (Roth) Lyngbye.

CERAMIUM GRACILLIMUmM Griff. and Harv. in Harv., 1846-1851, t. 206; De Toni,
1924, p. 515.

No. 2708 was growing on Desmia Kilneri J. Ag., No. 2704. The distribution of this
species in Australia seems surprising.

Nos. 2694, 2708, 2737, 2744, Flats.

Geogr. Distr.: Atlantic Ocean, Mediterranean and Adriatic Seas, West Indian Ocean,
Japan, etc. S. Australia and Tasmania. :

CENTROCERAS Kuetzing.

CENTKOCERAS CLAVULATUM Mont., 1840-1850, p. 140; Feldmann-Mazoyer, 1940, pp.
337-341, figs. 128-129.

No. 2695 growing on Cymodocea sp.; No. 2740 on Digenea simplex.

Nos. 2695, 2740, 2753, Flats.

Geogr. Distr.: Mediterranean and Adriatic Seas, Atlantic and Pacific Oceans. All
around Australia.

SPYRIDIA Harvey.

SPYRIDIA FILAMENTOSA (Wulf.) Harv. in Hook., 1833, p. 336; Feldmann-Mazoyer,
1940, pp. 348-351, fig. 133.—Fucus filamentosus Wulf., 1803, p. 64.

No. 2720, Reef.

Geogr. Distr.: Atlantic coasts of Europe, Africa and America, Mediterranean and
Red Seas, Indian Ocean, Japan. N., S. and EH. Australia and Tasmania.

Family DASYACEAE.
Dasya Agardh.

1. DasyaA pPAciFIcA Harv., Friendly Is. Alg. No. 12; in J. Ag., 1852-1863, p. 1223;
De Toni, 1897-1905 (Part 3), p. 1207.

Nos. 2748, 2754, Flats; 2804, Pelican Is. Flats.

Geogr. Distr.: Friendly Islands. New record to Australia.

2. DASYA Sp.

A prevalent and distinctive species, probably new. The plant is dendroid, about
3 cm. high, bears dichotomous monosiphonous hairs from near the apex, has five peri-
central cells and is much corticated. It has not been found fertile.

Nos. 2718, 2783, 2784, Reef; 2770, Coral Reef.

Geogr. Distr.: I have also collected this species at Woolgoolga, northern N.S.W.

3. DASYA sp.

A small amount of a little epiphyte, No. 2772, was found growing on Amphiroa prob.
foliacea, No. 2723, from the Coral Reef.

100 THE MARINE ALGAE OF BRAMPTON ISLAND,

Family RHODOMELACEAE,
POLYSIPHONIA Greville.
1. PoLySIPHONIA IMPLEXA H. and H., 1845, n. 59; De Toni, 1897-1905 (Part 3), pp.
889-890.
As Lucas’s specimens from Lord Howe Is. (Lucas, 1935). A surprising distribution.
Nos. 2684, 2749, Flats.
Geogr. Distr.: Lord Howe Is. S.W. Australia.
2. POLYSIPHONIA ZOSTERICOLA Lucas, 1919, p. 177.
Checked against type material in Lucas’s Herbarium (housed at "National Herbarium
of N.S.W., Sydney, Aus.).
No. 2687, Flats.
Geogr. Distr.: E. Australia.

DIGENEA Agardh.
DIGENEA SIMPLEX (Wulf.) Ag., 1823-1828, p. 389; De Toni, 1924, p. 404.—Conferva
simplex Wulf., 1803, p. 17, n. 16.
Specimens of this species were usually found bearing many epiphytes.
No. 2739, Flats; 2792, Reef; E., Pelican Is. Flats.
Geogr. Distr.: Most warm seas. N. and W. Australia.

RoOsScHERA Sonder.

ROSCHERA GLOMERULATA (Ag.) W.v.B., 1914, p. 289; De ‘Toni, 1924, pp. 404—-405.—
Tolypiocladia glomerulata (Ag.) Schmitz. in Engl. and Prantl, 1897, p. 442.—Hutchinsia
glomerulata Ag., 1823-1828, p. 102.

Plant size is variable.

Nos. 2685, 2686, 2733, Flats; 2805, Pelican Is. Flats.

Geogr. Distr.: E. Africa, W. India, Ceylon, Japan, Friendly and Philippine Islands,
New Zealand, etc. N. Australia.

ENDOSIPHONIA Zanardini.

ENDOSIPHONIA SPINULIGERA Zan., 1878, iD, ot, in, ZiS ION IO, to, Hal tis we} i Ive
W.v.B., 1913-1928 (Part c), p. 354.

A species but rarely collected previously.

Weber van Bosse (loc. cit.) points out that her specimen (Siboga material) differs
from the type material in that there is only one layer of cells next to the pericentral
cells of the same length as these pericentral cells, whereas on description there should be
two or more such layers. She concludes, however, that both specimens represent the
same species. The Australian material in L.S. of the thallus, shows two rows of cells
external to, and of the same length as, the pericentral cells.

No. 2719, Reef; 2746, Flats.

Geogr. Distr.: New Guinea, E. Indian Ocean. New record toe Australia.

LEVEILLEA Decaisne.
LEVEILLEA JUNGERMANNIOIDES (Mart. and Hering) Harv., 1855, p. 539; Borg., 1939,
p. 132.—Amansia jungermannioides Mart. and Hering in Mart., 1836, p. 485.
No. 2791 growing on Digenea simplex.
No. 2745, 2752, Flats; 2791, Reef.
Geogr. Distr.: Red Sea, Indian Ocean, Japan, Norfolk Island. W. and N.E. Australia.

AMANSIA Lamouroux.
AMANSIA GLOMERATA Ag., 1824, p. 247; De Toni, 1924, p. 426.
No. 2674, Pool in Reef; 2765, Coral Reef.
Geogr. Distr.: Warm Pacific and Indian Oceans. N.H. Australia.

LAURENCIA Lamouroux.
1. LAURENCIA RIGIDA J. Ag., 1876, p. 651; De Toni, 1897-1905 (Part 3), p. 789.
No. 2747, Flats.
Geogr. Distr.: Warm Indian Ocean, Java, Korea. N. and S. Australia.

BY VALERIE MAY. 101

2. LAURENCIA OBTUSA (Huds.) Lamour., 1813, p. 42; De Toni, 1924, p. 371.—Fucus
obtusus Huds., 1778, p. 586.

This seems to be a prevalent species of rather varying habit.

No. 2688, Flats; 2717, Reef; 2762, Coral Reef.

Geogr. Distr.: Widéspread in warm seas. All around Australia.

ACANTHOPHORA Lamouroux.
ACANTHOPHORA SPICIFERA (Vahl) Borg., 1910, p. 201, figs. 18-19; 1945, p. 61—Fucus
spiciferus Vahl, 1802, p. 44.
No. 2730 is a poor specimen.
Nos. 2730, 2738, Flats.
Geogr. Distr.: Widespread in tropical seas. N.E. Australia.

CHONDRIA Harvey.
CHONDRIA SD.
I have insufficient comparative material to determine the species.
No. 2709, Flats; 2761, Coral Reef.

References.

AGARDH, C. A., 1823-1828.—Species algarum rite cognitae. Vols. 1-2. Gryphiswaldiae.

, 1824.—Systema algarum. Lund.

AGARDH, J. G:, 1847.—Nya alger fran Mexico (= Algae Liebmanniae). Ofversigt K. Vet. Akad.
Fordhandlingar. Stockholm.

, 1848.—Species Genera et Ordines Algarum. Vol. 1. Lund.

, 1852-1863.—Species Genera et Ordines Algarum. Vol. 2 (2-3). Lund.

, 1872.—Till. Algernes Systematik I-III. -Lunds Univ. Arsskrift, 9. Lund.

, 1876.—Species Genera et Ordines Algarum. Vol. 3 (1), Epicrisis. Leipzig.
———. 1886.-—Till. Algernes Systematik 8 Siphoneae. Lunds Univ. Arsskrift, 23. Lund.
———_—, 1889.—-Species Sargassorum Australiae. Kongl. Svenska Vetensk. Akad. Handl., 23

(3), W=18e, “deo, abegile

, 1894.—Analeeta algologica contin. I. Lwnds Univ. Arsskrift, 29.

ARESCHOUG, J. E. (1853).—Spongocladia ett nytt algslagte. HKongl. Vetensk. Akad. Fordhl., 10.
Stockholm. ;

Barton, E#., 1901.—The genus Halimeda. Monogr. 60. Siboga Expedition, 1-30, Pl. 1-4. Leiden.*

BorGESON, F., 1905.—Contributions 4 la connaissance du genre Siphonocladus Schmitz. Oversigt.
Kgl. Danske Videnskabernes Selskabs. Fordhandlingar, 3, 259-291.

, 1910.—Some new or little Known West Indian Florideae IJ. Botanisk Tidsskrift, 30,
177-207.

, 1932.—A Revision of Forskaals Algae menticned in Flora Aegyptiaco-Arabica.
Dansk. Bot. Arkiv, 8 (2).

, 1934.—Some Marine Algae from the Northern part of the Arabian Sea with Remarks
on their Geographicai Distribution. Det. Kgl. Danske Videnskabernes Selskab. Biol.
Meddelelser, 11 (6), 1-72, Pl. 1-2. or

—-—, 1934a.—Some Indian Rhodophyceae especially from the shores of the Presidency of
Bombay. IV. Hew Bull.

, 1939.—Marine Algae from the Iranian Gulf. Danish Scientific Investigations in Iran,
Part 1, 47-141. Copenhagen.

, 1943.—Some Marine Algae from Mauritius. III. Rhodophyceae, Part 2. Gelidiales
Cryptonemiales Gigartinales. Det. Kgl. Danske Videnskabernes Selskab. Biol. Meddelelser,
18) (i), WaSsy, JPW, i,

——_—,, 1945.—Some Marine Algae from Mauritius. III. Rhodophyceae, Part 4. Ceramiales.
Det. Kgl. Danske Videnskabernes Selskab. Biol. Meddelelser, 19 (10), 1-68.

, 1948.—Some Marine Algae from Mauritius, Additional Lists. Det. Kgl. Danske
Videnskabernes Selskab. Biol. Meddelelser, 20 (12), 1-55, Pl. 1-2.

, 1950.—Vaughaniella, a new genus of Dictyotaceae. Det. Kgl. Danske Videnskabernes
Selskab. Biol. Meddelelser, 18 (8), 1-10.

Bornet, J. B. H., and FLAHAULT, C. H. M., 1886-1888.—Revision des Nostocacées heterocystees,
i-tV. Ann: Sei. Nat, 7.

Bory DE 8S. VINCENT, J. B., 1828.—Cryptogamie in Duperrey, L. J., Voyage autour du monde
sur la Coquille. Paris.

CHOU, RUTH CHEN-YING, 1944.—Pacific Species of Galaxaura I. Asexual Types. Pap. Mich.
Acad. Sci. Arts and Letters, 30, 35-55, Pl. 1-11.

* The full reference to Siboga Expedition is: “Uitkomsten op zoologisch, botanisch, oceano-
graphisch en geologisch Gebied verzameld in Nederlandsch Oost-Indié 1899-1900 aan boord
H.M. “Siboga’’ onder commando van Luitenant ter Zee 1° kl. G. F. Tydeman.”

102 THE MARINE ALGAE OF BRAMPTON ISLAND,

CHAN, RuTH CHEN-YING, 1945.—Pacific Species of Galavauwra II. Sexual Types. Pap. Mich.
Acad. Sci. Arts and Letters, 31, 1-24, Pl. 1-13. (Publ. 1947.)
DECAISNE, J., 1841.—Corall. Archiv Mus. Hist. Nat., 2. Paris.
, 1842.—Essai sur une Classif. des Algues et des polypiers calciferes. Mémoire sur
les Corallines. Ann. Sc. Nat., 2nd series, 1-7. Paris.
De Toni, J. B., 1889.—Sylloge Algarwm. Vol. I. Padua.
, 1895.—Sylloge Algarum. Vol. III. Padua.
1897-1905.—Sylloge Algarum. Vol. IV (1-4). Padua.
, 1924.—Sylloge Algarum. VI. Padua.
DrRoveT, F., and Dainty, W. A., 1948.—Nomenclatural Transfers among (‘occoid Algae. Lloydia,
ik. WniSUee
Ewuis, J. B., 1755.—Natural History of Corallines. London.
, and SoLANDER, D., 1786.—The Natural History of many curious and uncommon
zoophytes collected from various parts of the globe. London.
ENGLER, A., and PRANTL, K., 1897.—Die Nattrlichen Pflanzenfamilien. Leipzig.
FALKENBERG, P., 1901.—Fauna und Flora des Golfes von Neapel. Monograph 26. Xhodo-
melaceen. Berlin.
FELDMANN, J., 1938.—Sur la classification de lVorde des Siphonocladales. Rev. Générale de
Botanique, 50. Paris.
, and HAMEL, G., i1934.—Observations sur quelques Gelidiacées. Rev. Générale de
Botanique, 46. Paris.
FELDMANN-MAZzOYER, G., 1940.—Recherches sur les Céramiacees de la Méditerranée occidentale.
Thesis presentées a la Faculté des Sciences de l’université de Paris. 1-510, Pl. 1-4. Alger.
ForsKAAL, P., 1775.—Flora Aegyptiaco-Arabica. Hauniae.
FOsuIn, M., 1903.—The Lithothamnia of the Maldive and Laccadive Islands. Fauna and Geog.
of the Mald. and Lace. Archipelago, 1 (28).
FREYCINET, M. L. De, 1824-1826.—Voyage autour du monde exécuté sur l’Uranie . . . Zoologie
par Quoy et Gaimard (Incl. Polypier Flexibles by Lamouroux).
Gepp, A., and H. S., 1911.—The Codiaceae of the Siboga Expedition. Monogr. 62. Siboga
Expedition. Leiden.*
GMELIN, S. G., 1768.—Historia Fucorum. 239 pages, 33 Pl. Petropoli.
Gomont, M., 1892.—-Monographie des Oscillariées. Ann. Sci. Nat., 7, Bot. 15 and 16.
Gray, J. E., 1866.—On Anadymene and Microdictyon. Lond. Jour. Bot.
GREVILLE, R. K., 1830.—Algae Britannicae. Edinburgh and London.
Grunow, A., 1874.—Algen der Fidischi, Tonga- und Samoa-Inseln, gesammelt von Dr. E.
Graeffe. Jour. des Mus. Godeffroy, 3 (6), 23-50.
HariotT, P., 1887.—Algues Magellaniques nouvelles. Jour. de Botanique, I.
Harvry, W. H., 1844-—Algae of Tasmania. London Jour. Bot., 3, 428-454.
, 1846-1851.—Phycologia Britannica. 4 vols. London.
—, 1847.—Nereis Australis. London.
, 1855.—Some account of the marine botany of the colony cf Western Australia.
Trans. Irish Acad. Dublin, 22.
, 1858-1863.—Phycologia Australica. 5 vols. London.

-—————,, 1859.—Characters of new algae chiefly from Japan and adjacent regions. Proc.
Amer. Acad., 4. =
Hauck, D., 1887.—Uber ein v. J. M. Hildebrandt in Rothen Meere. u.. Ind. Ocean gesamm.
Algen. Hedwigea, 26. Dresden.
HEYDRICH, F., 1892.—Beitrage zur Kentniss der Algenflora Kaiser-Wilhelms-Land (Deutsch-

Neu-Guinea). Ber. der Deutschen botan. Gesellsch., 10 (8), 458-486, Tab. 24-26.
, 1897.—Corallinaceae, insbesondere Melobesieae. Ber. Deutsch Bot. Geselisch., 15
MDs Star, anki. 3h;
Hooker, W. J., 1833.—British Flora. Vol. II. London. ;
Hooker, J. D., and Harvey, W. H., 1845.—Algae Novae Zelandiae. Lond. Jowr. Bot., 4. London.
Howe, M. A., 1905.—Phycological Studies, 2. New Chlorophyceae, new Rhodophyceae and
miscellaneous notes. Contr. N.Y. Bot. Garden, 72.
, 1920.—Algae in Britton and Millspaugh, The Bahama Flora, 553-618. New York.
Hupson, G., 1778.—Flora Anglica. Vols. 1-2, 2nd ed. London.
KJELLMAN, F. R., 1900.—Om, Floride-Slagtet Galazaura, dess organografi och systematik.
Kongl. Svenska Vetenskaps-Akad. Handlingar, 33 (1), 1-110, Pls. 1-20. Stockholm.
IXUETZING, F. T., 1843.—Phycologia generalis. Leipzig.
, 1845.—Phycologia germanica. Nordhausen.
, 1849.—Species Algarum. Leipzig.
, 1849-1869.—Tabulae Phycologicae, 1-19. Nordhausen.
IXYLIN, HARALD, 1932.—Die Florideenordnung Gigartinales.. Lunds Univ. Arsskrift, 28 (8),
1-88, Pl. 1-28.

’

* The full reference to Siboga Expedition is: ‘“‘Uitkomsten op zoologisch, botanisch, oceano-
graphisch en geologisch Gebied verzameld in Nederlandsch Oost-Indié 1899-1900 aan boord
H.M. “Siboga’ onder commando van Luitenant ter Zee 1° kl. G. F. Tydeman.”

BY VALERIE MAY. 103

Lamouroux, J. V. F., 1805.-—Dissertations sur Plusiers Espéces de Fucus, ete. Paris.
, 1809.—Exposition des caractéres du genre Dictyota et tableau des espéces qu'il
renferme. Desv. Journ. de Botanique, 2.
, 1812.—Zoophytes flexibles ou coralligénes non entirement pierreaux. Nouv. Bull. des
Sciences par la Société Philomatique de Paris, 3, 182-188.
, 1813.—Essai de classification sur les Genres de la fam. des Thalassiophytes non
articulées. Mem. du Museum, 20). Paris
, 1816.—Histoire des polypiers coralligenes flexibles vulgairement nommés Zoophytes.
Caen.
Le Jours, A., 1863.—lListes des Algues Marines de Cherbourg. Mem. Soc. Imp. Sci. Nat. de
Cherbourg, 10 (1). Paris.
LINNAEUS, C., 1753.—Species Plantarum. Vol. 2. Stockholm.
1765.—Systema Naturae. Vol. 1, ed. 12. Holmiae.

- , 1771.—Mantissa plantarum. Ed. 2. Stockholm.
Lucas, A. H. §., 1919.—Notes on Australian Marine Algae. 2. Proc. LINN: Soc. N.S.W., 44 (1),
174-179.
, 1927.—Notes on Australian Marine Algae. V. PRoc. LINN. Soc. N.S.W., 52 (4),
555-562, Pl. 41-48.
, 1931.—The Marine Algae hitherto recorded from North-East Australia. Reports of
the Great Barrier Reef Committee, Vol. 3, No. 6, Pl. 47-57.
, 1934.—Notes on Australian Marine Algae. VII. The Algae of the Low Islands.
Proc. LINN. Soc. N.S.W., 59 (5-6), 348-350.
—————, 1935.—The Marine Algae of Lord Howe Island. Proc. LINN. Soc. N.S.W., 60 (3-4),
194-231, Pl. 5-9.
MARTENS, G. v., 1836.—Amansia jungermannioides. Flora.
, L866. —Die Preussische Expedition nach Ostasien. Bot. Theil. Die Tange. Berlin.
May, VALERIE, 1939.—Hctocarpus confervoides (Roth) Le Jol. Proc. LINN. Soc. N.S.W., 64
(5-6), 5387-554.
, 1947.—Studies on Australian Marine Algae. III. PRoc. LINN. Soc. N.S.W., 71 (5-6),
BUSA, IeAl, sles
————, 1948.—The algal genus Gracilaria in Australia. CSIR. Bull. 235, 1-61, Pl. 1-15.

3)
, 1948.—Studies on Australian Marine Algae. IV. Proc. LINN. Soc. N.S.W., 73 (5-6),

292-297.

, 1949.—Studies on Australian Marine Algae. V. Proc. LINN. Soc. N.S.W., 74 (3-4),

196-202. -

, 1951.—Studies on Australian Marine Algae. VI. Proc. LINN. Soc. N.S.W., 76, 83-87.

MENEGHINI, G., 1840.—Lettera al Dott. J. Corinaldi a Pisa. Padova.

MERTENS, H., 1819.—Mémoire sur plusieurs espéces de Fucus nouvelles ou peu connues, observées
dans la collection du Muséum. Mem. Mus. Hist. Nat., 5.

MONTAGNE, C., 1840:—Plants cellulaires in Webb and Berthelot Histoire Naturelle des Iles
Canaries, 3 (2). Phytographia Canariensis, Sectio Ultima. Paris.

, 1840-1850.—Flore d’Algérie. Phyceae in Bory de Saint Vincent et Durieu de
Maisonneuve, Exploration Scientifique de l’Algérie. Bot., 1-197, Pl. 1-16. Paris.

a , 1842.—Prodromus generum specierumque Phycearum novarum in itinere ad polum
antarticum. Paris.

OsporN, J. E. M., 1948.—The Structure and Life History of Hormosira banksii (Turner)
Decaisne. Trans. Roy. Soc. N.Z., 77 (1), 47-71.

PAPENFUSS, G. F., 1940.—Notes on South African Marine Algae. I. Botaniska Notiser. Lund,
200-226. ; ¢

————,, 1943.—Notes on Algal Nomenclature. II. Gymnosorus J. Agardh. Amer. Jour. Bot.,
30 (7), 463-468.

, 1944.—Notes on Algal Nomenclature. III. Miscellaneous Species of Chlorophyceae,
Phaeophyceae and Rhodophyceae. Farlowia, 1 (3), 337-346.

RotuH, A. G., 1797.—Catalecta Botanica. Fase. 1. Leipzig.

SETCHELL, W. A., and GARDNER, N. L. (1925).—The Marine Algae of the Pacific Coast of
North America. 3, Melanophyceae. Univ. Calif. Publ. in Bot., 8 (3), 383-898, Pl. 34-107.

SOLMS-LAUBACH, H. GRAFEN ZU, 1893.—Uber die Algengenera Cymopolia, Neomeris und
Bornetella. Ann. Jard. Bot. Buitenzorg, II, 61-97, Tab. 8-9. Leiden.

SONDER, W., 1845.—Nova algarum genera et species, quas in itinere ad oras occidentales Novae
Hollandiae, collegit L. Preiss. Bot. Zeit., 3. Berlin.

ee, 187i =D Algen des tropischen Australiens. Hamburg.

STEPHENSON, T. A., and A., TANDY, G., and SPENDER, M., 1931.—The Structure and Ecology of
Low Isles and other Reefs. Sci. Reps. Gt. Barrier Reef Hxapedition, 1928-29, 3 (2). British
Museum Natural History.

TANAKA, TAKESI, 1941.—-The Genus Hypnea from Japan. Sci. Pap. Inst. Algol. Research Fac.
Sci. Hokkaido Imp. Univ., 2 (2), 227-250, Pl. 53-54.

TURNER, O., 1808-1819.—Historia Fucorum. Vols. 1-4, 258 Tab. London.

VAHL, A., 1802.—Hndeel cryptogamiske planter (Fuci) fra St. Croix. Natur. Hist. Selsk.
Skrift., 5 Bind, 2 Heft. Copenhagen.

104 THE MARINE ALGAE OF BRAMPTON ISLAND,

WEBER VAN Bossk, A., 1898.—Monographie des Caulerpes. Ann. de Jardin Botanique de
Buitenzorg, 15, 243-401, Pl. 20-34.
—, 1905.—Note sur le genre Dictyosphaeria Dec. Nuova Notarisia, 16, 142-144.
, 1913-1928.—Liste des Algues du Siboga. Monogr. 59, a, b, ¢, d. Siboga Expedition.
Leiden. *
, 1914.—Rhodophyceae. Percy Sladen Trust Expedition to the Indian Ocean in 1905.
Trans. Linn. Soc. London, 16 (3).
— , and Fosuie, M., 1904.—The Corallinaceae of the Siboga Expedition. Monogr. 61.
Siboga Expedition. lLeiden.*
WULFEN, F. X., 1803.—Cryptogama aquatica. Roemers Arch. f. Bot., 3. Leipzig.
ZANARDINI, G., 1878.—Phyceae papuanae novae vel minus cognitae. Nuovo giornale botanico
italiano, 10, 34-40.

An alphabetical list and index of the species recorded in the present paper.

CHLOROPHYCEAE.
Page. ae Page. Page.
Anadyomene brownii ..... 91 Caulerpa sertularioides ... 92 Halimeda Opuntia ........ 92
Boergesenia Forbesii ..... 91 Cladophoropsis (?) mem- Spongocladia vaucheriae-
Bornetella oligospora ..... 92 PANACEA NTA retsheacts oteinseee 92 LOTS screech eres 92
Caulerpa papillosa ........ 93 Codium spongiosum ...... 92 Struvea delicatula ........ 91
TACCINOSAM meee cb eiere etree 3 Dictyosphaeria favulosa ... 91 Udotea flabellum .....:... 92
var. laetevirens ...... 93 IWieTSIUY SIN oe ceercas coeheencneis 91 OLI€MtaliSs, yam <r 92
var nummulariay a...) Io Hinteromornphay sp. Seemecae Sal WIA. JEVCUUIGE, soocuasao cobs 91
SEricblleneh Baoeadoas oogac 93 Halimeda incrassata ...... 92 Valoniopsis pachynema ... 91
var. de Boryana ..... 3 TANEKOIANIOY ooccoocoau00s 92
PHAEOPHYCEAE.
Chnoospora obtusangula .. 94 Hormophysa triquetra .... 94 Sargassum lophocarpum .. 95
Cystophyllum muricatum .. 94 Hydroclathrus clathratus .. 94 Scabnripes: 5 2 ea saseree ee 95
Dictyopteris pardalis ..... 94 Padina Commersonii ..... 94 Sphacelaria tribuloides .... 938
Dretyota, dichotomay As. : 94 Pocockiella nigrescens .... 93 Stypopodium zonale ...... a3
Wee, inneKCRNE, o500500000 94 Sargassum carpophyllum .. 95 Turbinaria ornata ........ 95
Ectocarpus confervoides .. 93 Hawa Cams. watence.c Nore aie 95 Vaughaniella rupicola .... 94
MYxXOPHYCEAE.
Brachytrichia Quoyi ...... 95 Entophysalis conferta .... 95 Hydrocoleum lyngbyaceum 95
Chilowaab:< Hl Goocoosoue 95 Hormothamnion solutum .. 95 Lyngbya majuscula ....... 96
Sirocoleum guyanense .... 96
RHODOPHYCEAE.
Acanthophora spicifera ... 101 Desmia Kilneri ........... 96 haurencia obtusa ..2.05.- 101
Amansia glomerata ....... HOO Diseneaysimplexy =. es nee 100 CUM: - Ae ie ctohene eran 100
Amphiroa prob. foliacea .. 97 Endosiphonia spinuligera .. 100 Leveillea jungermannioides 100
Centroceras clavulatum ... 99 Hucheuma muricatum sss os Miacora "ceranoides! sense: 96
Ceramium gracilliimum .... 99 Fosliella farinosa ........ 97 Mastophora plana ........ 97
Ceratodictyon spongiosum.. 99 Galaxaura glabriuscula ... 96 Metagoniolithon graniferum 97
Champilan vamvulaneee sen 99 SUbERUTICULOSaIy erin 96 Peyssonnelia Gunniana ... 96
@hondria ssp. Was semester 101 Gelidiella acerosa ........ 96 Plocamium hamatum ..... 98
Sone VIO SHS Tawney? Goononac 98 Goniolithon ? Fosliei ..... 97 Polysiphonia implexa ..... 100
TUM VAL TST Wisisrsyciulbevada, eeeviewes ere 98 Gracilaria lichenoides ..... 98 ZOSTETICOl Ax ak Ren ree 100
IDEISAYE), iSENGWIICEY G5.q oo bo Ulb os 99 Hypnea cervicornis ....... 98 Roschera glomerulata .... 100
SD Ew Paretewe let ooh favcvocer eee eae 99 Valentiaer aioe see wie 98 Sarconema filiforme ...... 98
SD. eee raisin con ae ete 99) Jiamniay muben'S! ta aenceanrlers trains 97 Sebdenia ceylonica ....... Siri
Spyridia filamentosa ...... 99

* The full reference to Siboga Expedition is: “Uitkomsten op zoologisch, botanisch, oceano-
graphisch en geologisch Gebied verzameld in Nederlandsch Oost-Indié 1899-1900 aan boord
H.M. “Siboga’”’ onder commando van Luitenant ter Zee 1° kl. G. F. Tydeman.”

105

THE DEVELOPMENT OF BRYOZOAN FAUNAS IN THE UPPER PALAEOZOIC
OF AUSTRALIA.

By JoANn CrockrorD, D.Sc. (Mrs. Beattie) .*

(Four Text-figures. )
[Read 25th July, 1951.]

Synopsis.

The development, sequence, and stratigraphic significance of bryozoan faunas in Australia
during the Carboniferous and Permian are described in as much detail as is possible from the
research so far carried out on this group. The distribution of bryozoan faunas during each
period is first summarized; then the occurrence of any forms of stratigraphic significance in
each area and the correlations which they suggest are considered; in conclusion, the evolution
of the broad faunal provinces found in the late Palaeozoic of Australia, and the probable
sourees and relationships of the bryozoan faunas of. these provinces are discussed.

LowrER CARBONIFEROUS BRYOZOAN FAUNAS.

Lower Carboniferous strata extend from the Hunter River in New South Wales to
beyond Rockhampton in Queensland; small collections of Bryozoa from several localities
within this area (Text-fig. 1) have yielded a comparatively large and varied fauna. The
early Carboniferous saw the influx of new families and subfamilies of Bryozoa, and the
disappearance of some of the older families or, where these families persisted, new
genera of a distinctly Carboniferous type were evolved at this time; many of these new
and specialized genera rapidly appeared in the faunas here (Tables 1 and 2). Correlation
of Australian Lower Carboniferous horizons has been made by comparison of goniatite,
coral, and brachiopod faunas with those of the Tournaisian and Viséan of Belgium or
of England and Scotland; as few published descriptions of Bryozoa from western
EHuropean faunas of this age are available, comparison of the bryozoan faunas is best
made with those of the United States and Russia, and the Tournaisian-Viséan boundary
of Europe is here regarded as approximating to the Osage-Meramac boundary of the
United States, and the Viséan-Namurian boundary as lying at or near the top of the
Chester Series (Cheyney et al., 1945).

THe AGE OF BRYOZOAN FAUNAS FROM THE BURINDI AND LOWER KUTTUNG SERIES.

The marine Burindi Series ranges from the base of the Tournaisian to the top of
the Viséan, and is partly contemporaneous with the predominantly freshwater Lower
Kuttung Series; the freshwater glacial beds of the Upper Kuttung are Upper
Carboniferous (Text-fig. 2). ; :

The main bryozoan horizon in the Burindi is that found at Glen William and
Hilldale, and is some 200 to 400 feet below the top of the Lower Burindi Series at these
localities. The occurrence of Hvactinopora, Fistulamina, Goniocladia, and Streblotrypa
all indicate an early Mississippian age for this fauna. Hvactinopora first occurs in the
Burlington and Keokuk Limestones in the United States; #H. trifoliata is a distinctly
more primitive form than any described from these Limestones and fixes the probable
upward limit to the age of this fauna. Fistulamina (as Meekopora ? aperta Ulrich, 1890)
also occurs in the Keokuk; typical species of Streblotrypa readily distinguished from
the more primitive Devonian species appear in abundance in the early part of the Osage
group; Goniocladia appears at the base of the Viséan in Russia and in the Lower
Carboniferous of Scotland; and Hemitrypa is particularly abundant in the Keokuk, and
H. clarkei closely resembles two Keokuk and Warsaw species. The fenestrate and
pinnate Bryozoa from this horizon also bear a strong general resemblance to those
described by McCoy in 1845 from the Lower Carboniferous limestone of Ireland, and
several Burindi species, though now considered distinct, were originally identified by
de Koninck (1878) with these Irish species. The occurrence of this group of genera in

't This work was commenced during the tenure of a Linnean Macleay Fellowship and
continued under a Commonwealth Research Fellowship at the University of Sydney.

M

106 BRYOZOAN FAUNAS IN THE UPPER PALAEOZOIC OF AUSTRALIA,

the top beds of the Lower Burindi Series, and the stage of development reached by
species representing each of them, suggests an age equivalent to an early part of the
Osage Series of the United States, and so to an horizon within the upper part of the
Tournaisian.

er
ol

TEXT — FIG.1 . DISTRIBUTION OF UPPER PALAEOZOIC BRYOZOAN FAUNAS

IN AUSTRALIA.

Text-figure 1.—Distribution of Upper Palaeozoic Bryozoan Faunas in Australia. The
principal localities to which reference is made in the text are indicated as follows:
Lower Carboniferous (/\): 1, Glen William and Hilldale; 2, Rouchel Brook; 3, Barrington:
4, Taree; 5, Mundubbera and Mundowran; 6, Cannindah; 7, Stanwell and Rockhampton.
Upper Carboniferous (JM): 8, Stroud; 9, Mt. Barney; 1% Stanwell and Rockhampton.
Permian (©): 11, Hobart district; 12, Maria Island; 13, Haglehawk Neck; 14, Fitzgerald;
, Marlborough; 16, Ulladulla; 17, Gerringong; 18, Wollongong; 19, Bundanoon; 20, Rylstone ;
, Hunter River Coalfield; 22, Lake’s Creek; 23, Springsure; 24, Bowen River Coalfield;
Port Keats; 26, Nooncanbah, West Kimberley; 27, Minilya Rivér, North-West Basin ;
, Lyons and Gascoyne Rivers, North-West Basin; 29, Irwin River.

toh we
co oF Ol

The fauna of the marine intercalation near the base of the Lower Kuttung at
Rouchel Brook and Back Creek is not far separated (by coarse marine and volcanic
sediments) from this Lower Burindi horizon; its fauna, though less varied, shows no
distinctive differences from the Lower Burindi fauna and is assumed to be of closely
similar age. é

The bryozoan fauna occurring in the Burindi Series (undivided) at Barrington
includes species of “Batostomella” and Dichotrypa. “Batostomella’, congeneric with
B. spinulosa Ulrich, 1890, is first recorded from the Ste. Genevieve Limestone of the
Meramac Group in the United States, and in Russia this genus is listed (Nikiforova,
1933) from the Viséan, but is not known to occur in definitely Tournaisian localities.
Dichotrypa, though one species occurs in the Middle Devonian, is particularly abundant
in the St. Louis Limestone of the Meramac, and occurs also in the Viséan in Russia.

BY JOAN CROCKFORD. 107

TABLE 1.

Distribution of Species in the Lower Carboniferous of N.S.W.
(The forms listed were described as new species by de Koninck, 1878, and Crockford, 1947.)

Upper
Lower Burindi. | Lower Kuttung. | Burindi | Upper
(?). | Burindi.
Species. ; Aa ONE) ” ieee) ? Wi
| | |
Glen | Rouchel | Back Bar-
William. | Hilldale. Brook. | Creek. | rington. | ‘Taree.
| |
_ Fistulipora mirari we he SER x x |

Dybowskiella rhomboidea.... ui AN | x
Evactinopora trifoliata a Ma aay x | | |
Fistulamina inornata oe fs Bie x x2 x
Goniocladia laxa se ANG nes Bs x (1)
G. parva F a ae aus x
Ramipora (Ramiporalia) bifurcata .. is x |
“ Batostomella ’’ lineata x
Fenestella* propinqua Ae at i 58 x | x x
F. acarinata .. us o i At x s<
FF. cribriformis nan as Bic te x
F. roucheli aes ts oa aS ae 4
F. barringtonensis .. ae ee He x
F. cellulosa .. up Ee 2 ne | x
Hemitrypa clarkei es ay re ee x SK
Ptilopora konincki .., ot ae We x x
Dendricopora hardyi Nes ei bse (2)
Penniretepora osbornet ee te = x
Streblotrypa parallela. . ae sa BY x x x x
Dichotrypa fragilis .. ine aa ui : x

(1) Original locality Burragood and Colo Colo.

(2) Original locality Burragood.

* The generic name Fenestella Lonsdale, 1839, is here used in preference to Fenestrellina d’Orbigny, 1849: an
application for suspension of the Rules of Zoological Nomenclature for the generic name Fenestella has been submitted
to the International Commission on Zoological Nomenclature by G. E. Condra and M. K. Elias (J. Paleont., 15, 1941 .
565-6). - E Be C

TABLE 2..
Distribution of Species in the Lower Carboniferous of Queensland.
(The forms listed were described as new species by Crockford, 1947.)

Oolitic Lst. Lion Creek Riverleigh Cannindah
Species. Mundowran. Lst. Lst. ; Lst.
Fistulipora etheridgei a Ace fs ae x
Dybowskiella crescentica .. ate wet a x
Evactinopora irregularis .. Bis wee me x
Fistulamina malmoensis pA oe eee - 5
Ramipora (Ramiporella) flexuosa .. age ae x
Leioclema porosa Be HS aie Se na x
Stenodiscus stanwellensis .. a. oh fae x
Fenestella yarrolensis as ae Me ites x
F., sp. indet. x
Polypora sulcifera x
Arcihmedes regina x
A. spiralis .. ie BG
Penniretepora fragilis 2 Us x
‘Rhabdomeson, sp. indet. .. Ds ie 3 x
Streblotrypa, sp. indet. Be: ee 43 a | x

108 BRYOZOAN FAUNAS IN THE UPPER PALAEOZOIC OF AUSTRALIA,

There is no direct field evidence of the stratigraphic position of this fauna; the presence
of these two genera at Barrington, and their absence from the Hilldale-Glen William
horizon, suggest that the Barrington fauna is slightly younger and should be correlated
with part of the Meramac, rather than the Osage, Group of the United States, and is
therefore of early Viséan age.

Dybowskiella, associated with Fistulamina and Fistulipora, occurs in the Upper
Burindi Limestone at: Taree; the coral fauna of this limestone is Upper Viséan in age.

THE AGE OF BRYOZOAN FAUNAS FROM THE LOWER CARBONIFEROUS OF QUEENSLAND.

- Bryozoa have been described from the Lower Carboniferous at four localities in
Queensland (Text-fig. 1; Table 2). The earliest of these faunas seems to be that of the
oolitic limestone at Mundowran in the same district as the Riverleigh Limestone; the
field and stratigraphic relationships of these limestones to each other are not known
(Hill, 1934, 105). As well as Hvactinopora, Fenestella, and Rhabdomeson, this oolitic
limestone contains abundant fragments of specifically indeterminate Bryozoa,
Fistulamina, Fistulipora, and batostomellids having been recorded from it. Hvactinopora
irregularis is more advanced than EH. trifoliata of the Lower Burindi, and more closely
approaches North American Burlington and Keokuk species; its zoarium was probably
attached and its growth form is slightly irregular, so that it is probably no younger a
species than the Burlington-Keokuk forms, in which the zoaria are free and their

Bryozoan horizon, Stroud.
Neerkoel Series Queensland.

Upper Kuttung

Upper Burindi Taree ,
Lion CK. and Riverleigh Limestones.

=
=
ra
2
a
L

and
Lower Kuttung

ii je ? Barrington’

me _ ?@ Oolitic Let Mundowran

Uppe

>. Pie TO a5 oe Lot Marine intercalation in Lower
aw ee eres.

seed Kuttung, Rouchel Brook and Back CK.

-> >. Bryozean horizon , Hilldale
and Glen William .

Text-fig. 2.—The Carboniferous Sequence of the Northern Hunter Valley
area of New South Wales, and the relative positions of the principal
Bryozoan horizons.

growth form more stabilized. The presence of this form thus suggests that this horizon
is of earlier age than the Riverleigh Limestone, and is equivalent to some part of the
Osage Series and so to the topmost beds of the Tournaisian; but equivalent to a slightly
higher horizon than the top part of the Lower Burindi, in which the more primitive
E. trifoliata occurs.

The coral faunas of the Riverleigh and Lion Creek Limestones are Upper Viséan
(Zone D2) in age or very slightly younger (Hill, 1943, 62). Two species of Archimedes

BY JOAN CROCKFORD. 109

occur in the Riverleigh Limestone; though this genus first occurs in the earlier Keokuk-
Warsaw beds in the United States, the greatest number of species occurs in the Chester
Series, equivalent in age to the top of the Viséan; Archimedes first occurs in the
western United States and Russia in the Lower Pennsylvanian. Ramiporella flexuosa
is a more primitive form than the species of this subgenus described from the Upper
Carboniferous of Russia. Polypora also first occurs in Australia in this limestone, and
Stenodiscus appears in the Lion Creek Limestone, but each of these genera appears in
an earlier part of the Lower Carboniferous in Europe and the United States.

UPPER CARBONIFEROUS BRYOZOAN FAUNAS.
Bryozoa are abundant in the Neerkol Series at Stanwell, Rockhampton and Mt.
Barney, and in a thin marine intercalation in the Upper Kuttung at Stroud (Table 3).

TABLE 3.

Distribution of Species in the Upper Carboniferous of Queensland and New South Wales.
(The forms listed were described as new species by Crockford, 1949.)

!
1
I

! Neerkol Series.

‘ i ak vee i Marine
| | | | Intercalation
Species. | Ridge | Malchi | | Mt. Barney, in Upper
above Creek, Par. 2v, | Mt. Barney, | Pors. 127v Kuttung
Lion Creek. | Rock- Par. | Pors. 193-4, and 202, Series,
| hampton. | Stanwell. | Par. Palen. | Par. Palen. Stroud.
|
|
ase eels | | | |
Fistulamina frondescens | | x
F. dispersa .. ae oe: x | < s
Leioeclema ? sp. we oo x | |
Fenestella matchi ee tal | x x | x | <
F. osbornei... Ys oe x | x x | 5 |
F. micropora .. ee os x | \
F. barneyi | | x x |
F. cincta he a i | x | |
Polypora pustulosa .. Be | x
P. neerkolensis fi ae x | x | x
P. palenensis ae a | | | | x
P. tenuirama .. is ae | x | |
Penniretepora spp. .. ER x x |
| |

Fenestellids have been recorded from the Kullatine Series in the Manning River district
and the Emu Creek Series at Drake, but no specimens suitable for description have been
collected. These Upper Carboniferous faunas (Crockford, 1949, 419) are characterized
by the association of Fistulamina with abundant Fenestella and Polypora; they lack
the variety found in Lower Carboniferous faunas, especially in the absence of fistuli-
poroids other than Fistulamina, and are distinguished from succeeding, Permian faunas
by the virtual absence of batostomellids and also the absence of Protoretepora and
Minilya; while it is possible that faunas described so far have come only from zones
in which fenestrate forms predominate and that other facies have been passed over
in collecting, the collections have been made from many widely scattered localities and
at present are regarded as representative.

STRATIGRAPHIC USE OF THE BRYOZOAN FAUNAS.

Generalized correlation between the Neerkol Series at the type locality near
Rockhampton and at Mt. Barney and the Upper Kuttung is indicated by the bryozoan
faunas (Crockford, 1949, 428). No species described from the Upper Carboniferous here
is known to occur elsewhere, and all the genera present are known earlier as well
as in the Upper Carboniferous; thus there is, unfortunately, no possibility at present
of using these faunas for wider correlations. The Pennsylvanian of the United States
and the Upper Carboniferous of Europe saw the introduction of many new forms with

110 BRYOZOAN FAUNAS IN THE UPPER PALAEOZOIC OF AUSTRALIA,

TABLE 4.

Distribution of Species in the Permian of Eastern Australia.

———————— a fe = s

New South Wales. } Tasmania. Queensland.

Hunter Valley.

|
| South 4 |
| Coast. |

Lower Upper Marine
Marine. Series.

Species.

Maria Island and Porter’s Hill.
Eaglehawk Neck and Fitzgerald

Dilly Stage, Springsure.

Lake’s Creek Quarry.
Middle Bowen Series.

Berriedale Lst.
Grange Quarry.

Fenestella
Muree Stage.
Mudstones.
Wollongong &
Gerringong.
Marlborough.

Rutherford
Shales.

Allandale
Stage.

Stage.
Ulladulla

Mulbring
Stage.

Gympie Beds.

Batostomella cylindrica Crockford x
Dyscritella porosa Crockford .. x
D. restis Crockford ey ae <
Stenopora johnstoni Etheridge . . x
S. etheridgei Crockford .. oe x
S. spiculata Crockford ae x
S. rugosa Crockford .. ae x
S. contigua Crockford os x
S. erinita Lonsdale ie es x x
S. gracilis (Dana)
S. nigris Crockford
S. frondescens Crockford
S. tasmaniensis Lonsdale... (1)
S. ovata Lonsdale oe Ba (1)
S. pustulosa Crockford . . oe x
S. hirsuta Crockford x
S. parallela Crockford .. aA x
S. grantonensis Crockford <
S. australis Nicholson and
Etheridge oe e. aie
S. jackii Nicholson and Etheridge x
S. leichardtii Nicholson and
Etheridge a we =H : x
S. gimpiensis Etheridge
Stenodiseus moniliformis Crock-
ford xe gs sad
Fenestella fossula Lonsdale... x x
FF’. dispersa (Crockford) He x x
FI. exserta Laseron
F. granulifera (Crockford)
FE. altacarinata (Crockford)
EF. quinquecella (Crockford) .. x x
I. canthariformis (Crockford)
EF. horologia Bretnall .. -4 Xx
FE. aspratilis Bassler
FE. simulatria (Crockford)
F. sparsinodata (Crockford)
EF. rockhamptonensis (Crockford)
EF. sparsa (Crockford) ae
I’. spinifera. (Crockford) NS
F. expansa (Crockford)
FF. expansa var. nodulifera
(Crockford) .. by 35el| |
Hemitrypa sexangula Lonsdale.. | (2)
Minilya duplaris (Crockford) | x
M. bituberculata (Crockford) .. <
Polypora pertinax Laseron a x x 1
P. virga Laseron a pete ge)| x | ex x

x
x

xX KX XK X

x

XX XK XK XK
x

x
x
x

I, IK OS

x

BY JOAN CROCKEFORD. TLaL3L

TABLE 4.—Continued.
Distribution of Species in the Permian of Eastern Australia —Continued.

|
New South Wales. Tasmania. | Queensland.
|
|
South 2 |
Hunter Valley. Coast. | = = |
a &
7) §
5 Sy :
Lower Upper Marine = | Vitibe ees B é
7 4 er (c) | 2 = | S n
Marine. Series. a | a S| oa | 2
| z A Fa
: = Pelee ean aml ers
Species. | 3S -s P| 2 ie Sh a i
Gal Ae al ele I EUW el eee
6 ) a o 4 D
eta le a giae) a | Plo|/Bl|¥! a) 2] 6] a
© 6 |= +S | ow x SS) a 5 |= oS = a | faa]
ae |S |3 Mm |e |S es) ws Ss qe | 8 a se | 6
is) “|o -|ean o ‘mo ens =] f= 2 br © cat Rn el o om
BO) ao| Oo) §$ | 59\/ da] C5) 8 2 ae op o x ~ = =
go) sso) Sa) 2 |S) esl Sel a | =F Fa ef aera [eae et || Si liar
SS/ 58/55/64 SS S2/So) s |e) 3 |b |e) SB) eS] &
2n\Gn| an) = |SniSSleo Ss Sia lola /al/saleisa
| | | |
| |
= |
P. woodsi (Etheridge) Be x x x SZ x x
P. montuosa (Laseron) ov x
P. dichotoma Crockford ae i x
P, triseriata Crockford .. ae | x
P. multinedata Crockford ore x |
P. magnafenestrata Crockford x x x x
P. linea Crockford A 35 x
P. minuta Crockford .. me x
P. smithii Etheridge .. Eps x
|
Protoretepora ampla (Lonsdale) x (2)
P. konincki Etheridge .. ie, 5 x
Ptilopora carinata Crockford .. | (3)
Rhombopora filiformis Crockford x
R. laxa (Etheridge) a5 ere . x

(1) Mt. Wellington, Mt. Dromedary, and Norfolk Plains (Strzelecki).
(2) Originally described from an unknown horizon, Tasmania.
(3) Described from Crinoidal Stage, Shoalhaven Heads.

Permian affinities which existed under varied climatic conditions over a large part of
the world; the absence of any of these new forms from the Upper Carboniferous here
suggests that the eastern Australian geosyncline was already cut off by the beginning
of the Upper Carboniferous from the free communication with other areas of Carboni-
ferous sedimentation which was so evident in the Lower Carboniferous.

PERMIAN BRYOZOAN FAUNAS.

Permian marine sediments containing abundant bryozoan faunas are widespread in
Australia; their occurrences fall into two distinct provinces in Hastern and Western
Australia. Within these two broad provinces the faunas in the various basins of
deposition show individual characteristics, but there are comparatively minor differences;
some slight mingling of the two types of fauna occurs, mainly in northern Queensland.

THE BASTERN AUSTRALIAN PERMIAN FAUNAS.

Bryozoan faunas have been described from Hobart and Marlborough in Tasmania,
the South Coast, Western, and Hunter River coalfields in New South Wales, and
Rockhampton, Springsure, and the Bowen River in Queensland; faunas of the same
general type occur in many other parts of these three States, but their faunas are
less well known and are given only passing reference here. These Permian sediments
extend for more than a thousand miles, but in spite of this and their great thickness
and varied facies, the generic composition of the bryozoan faunas is remarkably
uniform, and most genera present are long-ranged forms (Table 4); a small collection

112 BRYOZOAN FAUNAS IN THE UPPER PALAEOZOIC OF AUSTRALIA,

of Bryozoa from the Permian of New Zealand in the Australian Museum collections
belongs to the same general faunal type; no Bryozoa have been described from the
Permian of New Zealand. The sequence in the Hunter River coalfield and the relative
positions of bryozoan horizons in other districts are summarized in Text-figure 3.

Upper Coal Measures

we me = ee

Raging Vote mene Steaepora crinitar horizen

Mulbring Sta See aad of Hunter R., South Coast
9 Qe tage ies mea ras Santas N-S-W-, Eaglehawk Neck, and

VY)
=
4
w
W
W
Z

Fit 25° vald Tasmania -

IES : = . 7 Be ozoan horizons iN
Muree Stage SE CIOS INTE Z8 - (HAS er RQ. distictand Rylstene
and Bundanoon NS Ww: pesstoly
Coral Ck : BewenR -,Q.

‘enestella Shales, and
Ulladulla Mud stones , M-Sav-

Branxton Stage

‘
Lower Coal Measures BEE Ck. Quarry, Q.
Farley Stage Soe aes ‘s = ; , are — Dilly Stage, Springsere, Q .
Rutherford Stage Fa | Jacksons till, Pxelbin
SS . Bryozoan horizons at
Allandale STage WJANNandale and Fela NSW,
Eurydesma or deturn rae eae atin es lis and Maria 5. and Porter's
Wi Tas

Lochinvar Stage

LOWER MARINE SERIES

Text-fig. 3.—Sequence in the Hunter River district, and the approximate
relative positions of Bryozoan horizons in Eastern Australia.

STRATIGRAPHIC USE OF THE BRYOZOAN F'AUNAS.

(a) Correlation within Eastern Australia. :

Distinct bryozoan faunas from several horizons within two areas, the Hunter River-
Western Coalfield-South Coast basin in New South Wales and the Hobart district in
Tasmania, are recognizable and their faunas can to some extent be correlated.

In the Hunter River district the Lower Marine contains few Bryozoa below the
base of the Allandale; specimens in the Lochinvar Stage, all fenestellids, are unfor-
tunately specifically unrecognizable. The Pecten horizon at the top of the Allandale
Stage contains abundant fenestellids, a few large stenoporids and fine ramose batosto-
mellids; at the base of the overlying Rutherford Stage these fine batostomellids locally
form bryozoal limestones. The stenoporids and other batostomellids of these horizons
are distinct from those found in higher stages (Table 4); Polypora pertinagr is also a
common form in both horizons and several associated fenestellids as yet undescribed
also appear to be of limited vertical range. Stenopora johnstoni and Polypora pertinax
should be of value as index fossils for this section of the Lower Marine Series. The
sandy sediments of the overlying Farley Stage are a facies unsuitable for Bryozoa;

BY JOAN CROCKFORD. 113
their virtual absence is unfortunate, as important bryozoan horizons in other States
occur in beds correlated with this stage.

Near the base of the Upper Marine, the Fenestella Shales contain an abundant
bryozoan fauna; several species are restricted to this one band within the Branxton
Stage (Table 4) and should be useful and easily identified zone fossils; on the South
Coast this horizon and its characteristic fauna occur at Ulladulla, some two hundred
miles distant. Stenoporids and other batostomellids are typically very rare throughout
this stage. The Muree Stage contains many fenestellids, coarse species of Polypora
being especially characteristic, but their preservation is usually too poor for description;
Protoretepora ampla occurs in this horizon in the Western and Southern coalfields, and
small zoaria of Stenopora crinita, in contrast to the very large zoaria developed later,
first appear in the Muree. The Mulbring Stage is characterized by abundant large zoaria
of S. crinita and of coarse ramose stenoporids, and by the extreme rarity of fenestellids
and of the tiny ramose batostomellids found in earlier horizons; the large zoaria of
S. crinita are distinctive and easily identified, and should be useful index fossils for
this stage.

Bryozoan faunas from five localities in Tasmania can be closely correlated with
these horizons. At Porter’s Hill and Maria Is. the occurrence of Stenopora johnstoni
indicates a horizon near the top of the Allandale, and at Maria Is. there is a Hurydesma
zone just below the beds in which S. johnstoni occurs. The fenestellids at Marlborough
suggest a horizon close to the Fenestella Shales. Large zoaria of Stenopora crinita
occurring at Eaglehawk Neck and Fitzgerald show that these horizons are equivalent
to the Mulbring Stage, and this agrees with the position of the beds at Haglehawk
Neck near the top of the marine Permian sequence there.

In the Hobart district the Permian is heavily faulted; faunas from Granton,

Collinsvale, and Rathbone’s Quarries (all in the Berriedale Limestone), and Glenorchy,
Newtown, and Mt. Wellington have common features which suggest they are on the same

TABLE 5.
Distribution of Species in the Hobart District.

“Mt. Wel-
Berriedale Limestone. lington ”*’
Huon Rd., of
Telosa Mt. Wel- | Strzelecki,
Species. Road, Newtown. | lington, | Strickland
Glenorchy. 1000 ft. Ave.
Granton | Collinsvale| Rathbone’s above Sea Track,
Quarry. Quarry. Quarry. Level. 1m. W. of
Cascade.
Stenopora tasman iensis x
S. ovata x
S. pustulosa x SG x x x
S. hirsuta x x x
S. parallela eS x x
S. grantonensis .. Ga S<
Stenodiscus moniliformi x
Fenestella fossula x
FE, dispersa x
F. granulifera Bs x
Polypora magnafenestrata x

horizon (Table 5); the Berriedale, Limestone has been considered to belong to the
Lower Marine Series; of Bryozoa occurring both in this horizon and in New South
Wales, only one is short ranged, and this form, Fenestella granulifera, though restricted
to the Fenestella Shales in the Hunter district, occurs also at the top of the Dilly

114 BRYOZOAN FAUNAS IN THE UPPER PALAEOZOIC OF AUSTRALIA,

Stage (correlated with the Farley Stage) in Queensland; its occurrence suggests a
horizon either within the topmost beds of the Lower Marine or the basal beds of the
Upper Marine Series. The rich fenestellid horizon of the Grange Quarry contains three
species which occur in New South Wales; of these, Polypora woodsi is found only, in ~
the Fenestella Shales here, but is longer ranged in Queensland and Western Australia.

The Lake’s Creek Quarry fauna in Queensland is Artinskian in age (Crockford,
1945, 125) but cannot at present be correlated with any one horizon in New South
Wales. The top of the Dilly Stage at Springsure contains numerous stenoporids as well
as the recorded fenestellids; this horizon has been correlated from other forms with the
Farley Stage; the bryozoan fauna resembles that of the slightly younger Fenestella
Shales, but the coarse sediments of the Farley Stage itself contain no recognizable
Bryozoa. Species of Protoretepora and Stenopora were described by Nicholson and
Etheridge from the Middle Bowen Series (Reid, 1930, 74; and Table 4); the stenoporids
differ from those found elsewhere in eastern Australia in the small diameter of their
tubes, only about half that generally found in eastern Australian species, irrespective
of the size or shape of the zoarium itself; in this respect the Middle Bowen stenoporids
strongly resemble those of the Permian of Western Australia. The presence of
Protoretepora upon this horizon suggests correlation with the Muree Series and with
the Nooncanbah Series in Western Australia.

(b) Correlation with other Permian Fauwnas.

The bryozoan faunas from the base of the Allandale Stage to the top of the Upper
Marine Series are of Artinskian age, possibly ranging into the base of the Kungurian;
there is no evidence from bryozoan faunas regarding the age of the Lochinvar Stage.
The general type of fauna which appeared well developed in the Allandale and in
rocks of equivalent age in Tasmania persisted without great change through almost
all of the Permian sequence; the stenoporids in particular of the Allandale Stage are
of a well-marked Permian type. Polypora woodsi indicates a Lower Permian age for the
Branxton Stage and equivalent horizons; P. woodsi is the Artinskian representative of
the P. elliptica gens (Hlias, 1937, 327), which shows well-marked progressive variation
from the base of the Carboniferous into the Permian. The occurrence of Minilya in
the Branxton Stage and of M. duplaris at Lake’s Creek also indicates a Lower Permian
age; Minilya first appears at the top of the Pennsylvanian, and M. duplaris is a wide-
spread Artinskian species. Fenestella horologia, which occurs in the Dilly Stage, is also
an Artinskian form. The occurrence of Protoretepora ampla in the Muree and in the
Nooncanbah Series of Western Australia supports correlations of these horizons; the
Nooncanbah Series probably lies at the top of the Lower Permian, and the Muree is
also probably upon or very near this horizon. Thus the Mulbring Stage and the
Eaglehawk Neck beds may be basal Kungurian; it is notable that in these beds
fenestellids are virtually absent and giant batostomellids such as Stenopora crinita have
become abundant; fenestellids rapidly disappear at about the top of the Artinskian
throughout the world and the stenoporids also disappear early in the Kungurian, the
development of giant forms being a typical final development of an evolutionary line
before extinction.

Many attempts have been made to correlate eastern Australian and Indian late
Palaeozoic sequences; in these attempts the occurrence of several Bryozoa has been
widely quoted, particularly the occurrence of forms described from Tasmania by
Lonsdale (1844, 1845). These identifications have been discussed in detail in the
revisions of each species described by Lonsdale; while none of the figured specimens
from India seem to be conspecific with the eastern Australian types, there is a
marked general similarity in the groups of genera present in the Anthracolithic Series
of Kashmir particularly and the eastern Australian Permian, and the stage of develop-
ment reached’ by several genera common in both areas, and especially of the stenoporids,
Fenestella and Protoretepora, suggests a generally similar time range for the sediments
in which they occur; but beyond this the bryozoan faunas do not at present give any
evidence for precise correlation between the various stages of the eastern Australian

BY JOAN CROCKFORD. 115

|
sequence and the Indian late Palaeozoic; closer comparison is at present possible
between the more varied faunas of the Western Australian province and the Salt Range
faunas of India.

THE WESTERN AUSTRALIAN PERMIAN FAUNAS.

Bryozoa are abundant in the Permian sediments of the Irwin River, North-West
Basin, and West Kimberley districts; the generalized sequence in each of these areas
is shown in Text-figure 4; the stratigraphy has been summarized by Teichert (1946).
Marine strata containing similar Bryozoa were cut in bores at Port Keats, on the west
coast of the Northern Territory; this fauna also is discussed here. These areas of

Permian deposition extend for some 1,000 miles from Port Keats south-west to the
Irwin River.

The Bryozoa so far described from the Western Australian Permian are listed and
their stratigraphic distribution summarized in Table 6; a number of specialized Permian
genera, aS well as long-ranged forms, occur, and though only a small proportion of the
species present are known elsewhere (Table 7), many of those described from Western

NORTH-WEST BAS/N

MINILYA R WEST KIMBERLEY

DISTRICT
etx e...: |Kennedy
feat Sandstone |... 029°) Erskine Series
binopreductus Z a =
Stage
Schizodus oy SA ols Upper Ferruginovs
Stage Wandagee. EN hee Series
Calroolisponagia y Ser Ba yok sales
Sue Series

Lingola 6
re Tar aR Pe Nooncanbah Series

Lower Ferruginous
Series

| Nura Nura Lst

us (eae "
-: a, |Grant Ra. Series
een

j “4

Scale of the order of Zfe0' 2 ”

Text-fig. 4—Generalized section of the Permian sequence in Western
Australia (after Teichert, 1941, 390).

Australia show clear affinity with Permian forms from other areas; the relationships
' of individual species have been discussed previously in descriptive papers. The fauna
of the Fossil Cliff horizon in the Irwin River district is not listed separately, since
descriptions of the main collections from this horizon, sent to the United States for
study by Moore and Bassler, are not yet published and only small collections at present
remain in Australia; in these the fauna appears generally similar to that found in
the Callytharra. Another group in the Western Australian faunas which has as yet

116

TABLE 6.
Distribution of Species in the Permian of Western Australia and the Northern Territory.

BRYOZOAN FAUNAS IN THE UPPER PALAEOZOIC OF AUSTRALIA,

|
Species.

Cally-
tharra.

Barra-
biddy
Shale
Series.

Cundlego.

Calceoli-
spongia
Stage,

Wandagee..

Lino-
productus
Stage,

Wandagee.

Noon-

canbah.

Port
Keats.

Fistulipora vacuolata Crockford

F. wadei Crockford

F. compacta Crockford

F. conica Crockford

F. gigantea

Dybowskiella geet Etheridge

D. crescens (Crockford)

Hexagonella australe (Bretnall)

H. dendroidea (Hudleston)

H. densa Crockford

H. nalbia Crockford

H. undulata Crockford F

H. bifida Crockford .. Lay

H. lineata Crockford

H. plana Crockford 26

Evactinopora crucialis Hudleston

Goinocladia timorensis Bassler. .

Ramipora ambrosoides (Bret:
nall)

* Sulcoretepora ’”’
(Etheridge) ate ae

Stenopora sp. ‘‘ A ’’ Etheridge

S. sp. ““ B’”’ Etheridge

S. sp. ‘‘C’’ Etheridge

Fenestella horologia Bretnall

. affluensa Bretnall ..

. chapmani (Crockford)

. sparsigemmata (Crockford)

alia (Crockford)

. disjecta (Crockford)

. columnaris (Crockford)

ruidacarinata (Crockford) ..

. valentis (Crockford)

. lennardi (Crockford)

. cacuminatis (Crockford)

Mi inilya duplaris (Crockford). .

M. princeps (Crockford)

M. amplia (Crockford)

Protoretepora ampla (Lonsdale)

P. australis (Hinde)

Lyropora erkosoides Etheridge

Polypora fovea Crockford

P. retificis Crockford ..

P. woodsi (Etheridge) . .

P. multiporifera Crockford

Penniretepora triporosa Crock-
ford

P. granulata Crockford ©

P. fossata Crockford

Septopora ornata Crockford

Synocladia spinosa Crockford

Rhabdomeson mammilata (Bret-
nall) .. Sy aol

Rhombopora tenuis (Hinde)

R. hindei Etheridge ..

Streblotrypa browni riences

S. marmionensis Etheridge

S. etheridgei Crockford ApeAl

Rhombocladia minor Crockford |

R. spinulifera Crockford

Streblocladia excavata Crockford

meridianus

See ee ee

> a> ny GD DG

x

xX X XX

x

XX KK XK XK x

x

BY JOAN CROCKFORD. Uae

received little attention is the Batostomellidae, which are abundantly represented by
Stenopora, Batostomella, and closely related forms; the wide distribution of batosto-
mellids in collections from most areas in Western Australia has been shown by the
numerous records of their occurrence in faunal lists in stratigraphic papers, but so far
this group has received little attention in descriptive work.

STRATIGRAPHIC USE OF THE BRYOZOAN FAUNAS.

(a) Correlation within Western Australia.

The occurrence in the Western Australian sequence of Penniretepora, Lyropora,
Septopora and Streblocladia is restricted to the Callytharra Series; not a single speci-
men of any of these genera has been found in large collections from other horizons.
The Callytharra is also characterized by the presence of numerous ribbon-like zoaria
of Hexagonella, and by a group of fenestellids which, though some are long-ranged,
contains a large proportion of species quite distinct from those found on higher horizons.
The apparent absence of Goniocladia, Stenopora, and Synocladia, and the comparative
rarity of Fistulipora, in the Callytharra is also notable.

Synocladia is common in and characteristic of the Cundlego Series and is not
known on any other horizon in Western Australia.

TABLE 7.
Distribution of Species Common to the Permian of Western Australia and Localities Elsewhere.

Species. Distribution in W.A. | Occurrences Elsewhere.
= | —
|
Goniocladia timorensis .. | Nooncanbah. Basleo Beds, Timor.
Ramipora ambrosoides .. | Callytharra to Wandagee and Noon- | Basleo Beds, Timor (as Acanthocladia
canbah. acuticostata Bassler.
Fenestella horologia. . os an 90 5 Bitaoeni to Basleo Beds, Timor, and

Permian, Vancouver Is. (as F. parvi-
uscula (Bassler)); Dilly Stage, Qld.
Minilya duplaris .. ae | 5 at a ? Middle Productus Lst., Salt Ra. (as.
F., perelegans Meek); Dilly Stage, and
Lake’s Ck., Qld.

Polypora woodsi .. .. | Callytharra and Nooncanbah. Bitaoeni Beds, Timor (as P. tripliseriata
Bassler) ; Dilly Stage and Lakes Ck.,
Qld. ; Upper Marine, N.S.W. ; Granton,

| Tas.
Protoretepora ampla | Nooncanbah. Muree Stage, N.S.W.
Rhabdomeson mammillata .. FA Anthracolithic Series, Southern Shan
a States (as R. shanse Reed).
Streblotrypa marmionensis Callytharra to Wandagee and Noon- | ? Basleo and Amarassi Beds, Timor
canbah. (?=8S. germana Bassler).

The Wandagee and Nooncanbah Series are characterized by abundant massive and
coarse ramose Fistulipora; by broad frond-like species of Hexagonella which, although
they do occur on other horizons, are particularly characteristic of the higher beds of
the Permian sequence; by the common occurrence of Goniocladia and the presence of
Protoretepora; and by the distinct species of other genera such as Fenestella, Polypora,
Minilya, Rhombopora and Rhabdomeson, in all of which, though they include some
long-ranged forms, there are several species which are restricted to these higher beds.
Massive and coarse ramose stenoporids are also very common in both series. These
faunas of the Wandagee Series of the North-West Basin and the Nooncanbah Series of
the West Kimberley district bear a strong general resemblance to each other, and as
more research is done the number of species common to both will be greatly increased.
The fact that, excluding long-ranged forms, only three species have been recorded as
common to both series does not give a true picture of their relationship, and reference
to published descriptions will show the similarity between several other species from
these two areas. Four stages within the Wandagee Series have been differentiated by

118 BRYOZOAN FAUNAS IN THE UPPER PALAEOZOIC OF AUSTRALIA,

Teichert (1946, 99); several described fenestellids and fistuliporoids are restricted to a
single stage within this series (Crockford, 1944, Table 1; 1946, 145-154).

The fauna described from Port Keats contains only two long-ranged species in
common with other localities in this province; no precise correlation of the Port Keats
horizon is possible at present, but in its general aspect this small fauna resembles
the Callytharra fauna much more closely than those of higher series.

(b) Correlation with Eastern Austraha.

The occurrences of several species common to both provinces (Table 7) and the
close similarity already noted between stenoporids from the Middle Bowen and Western
Australian species indicate that the faunas of these two provinces, despite the funda-
mental differences in their aspect, are of similar time range within the Permian. Only
one species appears to be of definite value in closer correlation: Protoretepora ampla,
the occurrence of which supports correlation of the Nooncanbah with the Muree Stage
of the Upper Marine Series.

(c) Correlation with other Permian Faunas.

Only seven species occurring in the Western Australian Permian are known from
localities outside Australia (Table 7), but the stage of development reached by many
of the genera present in the fauna and the occurrence of several rare and specialized>
forms give information of stratigraphic value. The Western Australian faunas show
most affinity with those described by Bassler (1929) from Timor and with the Middle
and Upper Productus Limestone faunas of the Salt Range; in addition, several
Callytharra species, and the fistuliporoids throughout the sequence, are similar to late
Pennsylvanian and Permian forms from midcontinental North America.

The Callytharra fauna shows a strong general resemblance to the fauna of the
Graham Formation of the Cisco Group at the top of the Pennsylvanian of Texas described
by Moore (1929), and of the Upper Pennsylvanian of Oklahoma described by Warthin
(1930); Raggatt and Fletcher (1936) have previously suggested correlation between
these horizons; they were probably deposited in very similar facies and at not widely
differing times, but no species are now considered common to these two areas. On the
other hand, the Callytharra contains at least five species (Table 6) in common with
the higher beds of the Western Australian sequence, and each of these five species
occurs in Artinskian but not in older strata elsewhere (Table 7). In addition,
Hezagonella australe and other ribbon-like species of this genus common in the
Callytharra represent a similar stage of development to H. turgida Bassler, 1929, from
the Basleo and Amarassi Beds of Timor, and to the genotype from the Middle and
Upper Productus Limestones of India, and Hexagonella is not known from pre-
Artinskian strata in any part of the world; Septopora ornata is more closely related
to species described from the Somohole and Bitaoeni Beds of Timor than to Russian
and North American late Carboniferous species; Streblotrypa marmionensis, which first
appears in the Callytharra, possesses the central bundle of small tubes characteristic
of Permian species of this genus, and is probably identical with a species described
from the Basleo and Amarassi Beds of Timor; and Polypora woodsi is a widespread
Artinskian species and is the Lower Permian representative of the P. elliptica genus of
Elias (1987, 327). HEvactinopora and Lyropora, both of which have not elsewhere been
recorded from post-Mississippian horizons, occur in the Callytharra, and Hvactinopora
also in higher beds of the Permian sequence; while the described species of Lyropora
is a typical form of this genus, Hvactinopora crucialis (Hudleston, 1883, 593: Etheridge,
1908, 9) is more advanced in zoarial form than any Mississippian species, and the
presence of these two genera does not suggest any correlation of the Callytharra with
Carboniferous horizons. The bryozoan faunas indicate that the Callytharra should be
correlated with the early part of the Artinskian, and probably, therefore, with the
Bitaoeni Beds of Timor (cf. Teichert, 1941, 399). Teichert also correlates the
Callytharra with the Lower Productus Limestone of the Salt Range; Bryozoa are
apparently uncommon in the Lower Productus Limestone; the Callytharra faunas show
some resemblance to those of the Middle and Upper Productus Limestones, but this

BY JOAN CROCKFORD. 119

resemblance is not so strong as that between the Wandagee and Nooncanbah faunas
and those of the Middle and Upper Productus Limestones.

Synociadia is restricted to the Cundlego Series; similar species of this genus also
appear in abundance in the Middle Productus Limestone and continue into the Upper
Productus Limestone, and other similar species have been described from the Permian
of England and Germany. The Cundlego fauna is of definite Artinskian age and
should probably be correlated with an early part of the Middle Productus Limestone.

The Wandagee and Nooncanbah faunas show their closest resemblance to those of
the Timor Permian; the resemblance is rather more marked in comparing the Wandagee
and Nooncanbah with the higher stages, the Basleo and Amarassi Beds, than with the
Bitaoeni Beds. This is particularly noticeable in comparing the abundance of massive
and.coarse ramose species of Fistulipora in the Wandagee and Nooncanbah with the
sudden abundance of similar forms with comparable internal structure in the Basleo
and Amarassi Beds, and it is also noticeable that Sphragiopora, which occurs only in
the Amarassi, and Goniocladia, which occurs only in the Basleo Beds, are both quite
abundant in the higher beds in Western Australia, but do not occur in the earlier
stages. Five species are known to be common to these higher beds in Western Australia
and the Timor Permian sequence (Table 7); but only one, Goniocladia timorensis, is of
restricted range in both areas, occurring only in the Nooncanbah and in the Basleo
Beds. Species of Hexagonella occurring in the Wandagee and Nooncanbah Series, and
also Streblotrypa etheridgei from the Nooncanbah, are distinctly more advanced species
than any described so far from Timor, and this is true also of coarse specimens of
Goniocladia which occur frequently in collections from the Wandagee and Nooncanbah.
The Bryozoa, therefore, suggest correlation of these two series with a slightly higher
horizon than that indicated by Teichert (1941, 399), and to be possibly slightly
younger, and at least no older, than the Basleo Beds.

Teichert (loc. cit.) also correlates the Wandagee and Nooncanbah with the top
part of the Lower Productus Limestone of India. The bryozoan faunas suggest that
the Western Australian sequence from at least the Cundlego Series upwards is younger
than the Lower Productus Limestone; the general aspect of the bryozoan fauna which
appeared in the Middle and persisted into the Upper Productus Limestone is closely
similar to that of the Wandagee and Nooncanbah. The similar stage of development
of internal structure in species of Fistulipora and Dybowskiella present in both areas,
and the affinity of species of Goniocladia is again noticeable, while the zoarial form in
species of Hexagonella from these higher beds in Western Australia seems to be of
later development.than that of any described Salt Range species; and among the
fenestellids, though so far only one species is probably common to both areas (Table 7),
there is distinct similarity between other species, especially in the occurrence in both
areas of Protoretepora s. str. It does not appear possible that the Wandagee and
Nooncanbah Series could represent a horizon earlier than one within the Middle
Productus Limestone. The bryozoan faunas of these two series therefore suggest
either a late Artinskian or an early Kungurian age for the horizons on which they occur.

THE DEVELOPMENT OF FAUNAL PROVINCES DURING THE UPPER PALAEOZOIC.

The bryozoan fauna of the Lower Carboniferous in eastern Australia migrated
freely and rapidly between this and other areas of early Carboniferous deposition.
This migration was from the north, the fauna being of the same general type as that
found contemporaneously in central North America and Europe. The mild, cold,
temperate climate during the Tournaisian favoured the development of the varied
fauna found; the seas commenced to withdraw to the north during the early Viséan and
the climate became sub-glacial, but rapid migration of faunas from this same Tethyan
source remained possible until the close of the Viséan, and this fauna quickly
re-established itself in the brief warmer intervals represented by the Upper Viséan
reef limestones.

The Upper Carboniferous was a period of intense glaciation in New South Wales,
where the freshwater facies of the Hunter River passes northwards into predominantly

120 BRYOZOAN FAUNAS IN THE UPPER PALAEOZOIC OF AUSTRALIA,

marine sediments in northern New South Wales and Queensland. The bryozoan faunas
appear to be the remnant, restricted by adverse climatic conditions, of the earlier
Carboniferous faunas; they appear, from the persistence of Fistulamina and the virtual
absence of batostomellids, to be more closely related to these earlier faunas than to
those of the succeeding Permian. By the Upper Carboniferous eastern Australia was
probably cut off from free migration of faunas from the north, for there
is no evidence of any one of the numerous genera of Bryozoa which were at that time
evolving in the North American and European seas.

In the Lower Permian an abundant Tethyan fauna resembling that of the Lower
Carboniferous of eastern States re-appeared in Western Australia. These faunas were
clearly derived from a common faunal source, the eariy Carboniferous derivatives
which migrated along the east coast being primitive representatives of genera which
much later, having reached a higher stage of their development, migrated into the
Permian basins of Western Australia. This widespread late Palaeozoic fauna did not
migrate as far as eastern Australia during the Permian.

The wide divergence between the eastern and Western Australian faunas is not
due to any difference in age, since the bryozoan faunas of Western Australia are the
Permian representatives of a general type of fauna which existed for a long part of the
Upper Palaeozoic, and the generally similar time range of the different areas of
Permian sedimentation in eastern and Western Australia has long been established;
nor is difference in facies sufficient reason for the distinct faunas, since Bryozoa are
abundant in many different types of facies in both areas.

The Western and eastern Australian faunas have frequently been referred to as
“warm-” and ‘cold-water’ faunas respectively, and difference in climate in part accounts
for the distinct faunas, but seems insufficient to do so fully. The type of fauna and
the groups of genera found in the Western Australian Permian are spread over so large
a part of the world that they must have been able to exist under a considerable
variety of climatic conditions; intermittently the climate of eastern Australia, especially
of New South Wales and Tasmania, was extremely cold during the Permian, and in
some cases glacial erratics are associated with abundant Bryozoa; but some of the
interglacial epochs were of long duration and the climate must then have been milder,
and earlier, during the Lower Carboniferous, a varied bryozoan fauna rapidly reappeared
during such milder intervals. In Western Australia also intermittent glaciation occurred
during the Permian, and one glacial horizon, the Nura Nura Limestone of the Kimberley
district, is correlated with the Callytharra Series of the North West Basin, which
contains an abundant and varied bryozoan fauna, especially fistuliporoids. The great
distance through which Permian deposits are distributed in eastern Australia must, like
the interglacial epochs, have afforded some appreciable difference in climate, but the
generic assemblage of Bryozoa is unvaried.

_ Limited migration between these provinces was possible during at least a part
of Permian time, as shown by the occurrence of a few identical species in Queensland,
Western Australia, and Timor, and to a lesser extent in comparing the faunas of New
South Wales and Tasmania with those of Western Australia (Tables 4, 7); the. occur-
rence of generally similar types of stenoporids in the Bowen River coalfield and in
Western Australia also suggests relationship between the faunas of these two areas.
This slight intermingling of faunas occurs earlier in Queensland, in the Dilly Stage,
than in New South Wales, where the first evidences of migration are seen in the
early part of the Upper Marine Series; but the fauna of each province retained without
appreciable modification its own characteristics until marine Permian sedimentation
ceased.

The eastern Australian Permian faunas may have been developed during the early
part of the Permian from the poor and restricted Neerkol faunas, or were more probably
(because of the absence of Fistulamina or any form derived from it, and because of the
abundance of batostomellids, which were virtually absent from the Neerkol) faunas
which migrated from a southern source northwards along the east coast of Australia.

BY JOAN CROCKFORD. 121

The strong resemblance between the Tasmanian faunas and those of the
Hunter Valley-South Coast Basin in New South Wales, and the gradual lessening of
these resemblances through the central north coast basin in New South Wales and
the areas of Permian deposition in Queensland also suggest that the eastern Australian
faunas as a whole migrated from the south northwards; such migration was possibly
influenced by cold south to north currents developed around the coastline of that time,
and such currents could have been an éffective barrier to any large-scale migration of
the Tethyan faunas into the eastern Australian province.

Bibliography.

BASSLER, R. S., 1929.—The Permian Bryozoa of Timor, Palaontologie von Timor, xvi Lief., xxviii.

BRETNALL, R. W., 1926.—Palaeontological Contributions to the Geology of Western Australia,
Series vii, No. xiii: Descriptions of some Western Australian Fossil Polyzoa. W. Aust.
Geol. Surv. Bull., 88.

CHEYNEY, M. G., et al., 1945.—Classification of Mississippian and Pennsylvanian Rocks of
North America. Bull. Am. Assoc. Petroleum Geologists, 29 (2):125.

CROCKFORD, JOAN, 1941a.—Permian Bryozoa of Eastern Australia, Part I. A Revision of Some
Previously-Named Species of Fenestrellinidae (Fenestellidae). J. Roy. Soc. N.S.W., 74
(for 1940) : 397.

, 1941b.—Permian Bryozoa of Eastern Australia, Part II. New Species from the
Upper Marine Series of New South Wales. J. Roy. Soc. N.S.W., 74 (for 1940) : 502.

, 1948a.—Permian Bryozoa of Eastern Australia, Part III. Batostomellidae and
Fenestrellinidae from Queensland, New South Wales, and Tasmania. J. Roy. Soc. N.S.W.,
76 (for 1942) : 258.

, 1943b.—Bryozoa from the Port Keats Bore, Northern Territory. Proc. LINN. Soc.
N.S.W., 68: 145. ;

, 1944a.—A Revision of Some Previously Described Species of Bryozoa from the Upper
Palaeozoic of Western Australia. J. Roy. Soc. W.A., 28 (for 1941-2) : 187.

, 1944b.—Bryozoa from the Wandagee and Nooncanbah Series (Permian) of Western
Australia. Part I. J. Roy. Soc. W.A., 28 (for 1941-2) : 165.

, 1944¢.—Bryozoa from the Permian of Western Australia, Part I. Cyclostomata and
Cryptostomata from the North-West Basin and Kimberley District. Proc. LINN. Soc.
N.S.W., 69:139.

, 1945.—Stenoporids from the Permian of New South Wales and Tasmania. Proc.
LINN. Soc. N.S.W., 70: 9.

, 1946.—A Bryozoan Fauna from the Lake’s Creek Quarry, Rockhampton, Queensland.
Proc. Linn. Soc. N.S.W., 70 (for 1945) : 126.

, 1947.—Bryozoa from the Lower Carboniferous of New South Wales and Queensland.
Proc. LINN. Soc. N.S.W., 72:1.

, 1949.—Bryozoa from the Upper Carboniferous of Queensland and New South Wales.
Proc. LINN. Soc. N.S.W., 73: 419.

Dana, J. D., 1849.—In Wilke’s U.S. Exploring Expedition, 10, Geology, Appendix, and Atlas.

ELIAS, M. pis, 1937.—Stratigraphie Significance of Some Late Palaeozoic Fenestrate Bryozoa.
J. Paleontology, 11 (4): 306.

EXTHERIDGE, R., 1872.—Descriptions of Palaeozoic and Mesozoic Fossils from Queensland.
Q. J. Geol. Soc., 28: 317.

EXTHERIDGE, R., JNR., 1880.—On a Collection of Fossils from the Bowen Rivér Coalfield and
the Limestone of the Fanning R., North Queensland. Proc. Roy. Phys. Soc. Edinb., 5:1.

, 1891.—A Monograph of the Carboniferous and Permo-Carboniferous Invertebrata of
New South Wales, Part I, Coelenterata. N.S.W. Geol. Surv. Mem.. Pal. 3.

, 1892.—In Jack, R., and Etheridge, R., Geology and Palaeontology of Queensland and
New Guinea.

, 1903.—Palaeontological Contributions to the Geology of Western Australia, I.
Descriptions of Carboniferous Fossils from the Gascoyne District. W. A. Geol. Surv.
Bull. 10. ‘

, 1907.—Official Contributions to the Palaeontology of South Australia, No. 19.—
Fossils of Port Keats Bore, Thirty Miles North,of Fossil Head, Treachery Bay. Supple-
ment to Parliamentary Paper 55 of 1906.

—, 1915.—Palaeontological Contributions to the Geology of Western Australia, Series V,
No. 10. Western Australian Carboniferous Fossils, chiefly from Mount Marmion, Lennard
River, West Kimberley. W.A. Geol. Surv. Bull. 58.

, 1920.—Palaeontologia Novae Cambriae Meridionalis—Occasional Descriptions of New
South Wales Fossils, No. 8. Rec. Geol. Surv. N.S.W., 10 (1):1.

Hitt, DorotHy, 1934.—The Lower Carboniferous Corals of Australia. Proc. Roy. Soc. Queens-
land, 45: 62.

, 1943.—A Re-interpretation of the Australian Palaeozoic Record, based on a Study of
the Rugose Corals. Proc. Roy. Soc. Queensland, 54 (6): 53.

N

122 BRYOZOAN FAUNAS IN THE UPPER PALAEOZOIC OF AUSTRALIA.

HINDE, G. J., 1890.—Notes on the Palaeontology of Western Australia, Corals and Polyzoa.
Geol. Mag:, N.S., Dee. 3, 7: 194.

HUDLESTON, W. H., 1883.—Notes on a Collection of Fossils and Rock Specimens from West
Australia, North of the Gascoyne River. Q. J. Geol. Soc., 39: 582.

KOoNINCK, L. G. DE, 1877 (1898).—Ré€cherches sur les Fossiles Paléozoiques de la Nouvelle
Galles de Sud. Mem. Soc. Roy. des Sciences de Liege, 2: 8. (Reprinted as Descriptions of
the Palaeozoic Fossils of New South Wales. WN.S.W. Geol. Surv. Mem., Pal. 6.)

LASBERON, C. F., 1918.—Notes on Some Permo-Carboniferous Fenestellidae, with Descriptions of
New Species. J. Roy. Soc. N.S.W., 52: 81.

LONSDALE, W., 1844.—In Darwin, Geological Observations on Volcanic Islands.

, 1845.—In Strzelecki, Physical Description of New South Wales and Van Diemen’s
Land.

Moore, R. C., 1929.—A Bryozoan Faunule from the Upper Graham Formation, Pennsylvanian,
North-Central Texas. J. Pal., 3 (1):1 and 3 (2):121.

, and DUDLEY, RUTH, 1944.—Cheilotrypid Bryozoa from Pennsylvanian and Permian
Rocks of the Midcontinent Region. Kansas State Geological Survey Bull., 52 (6): 229.

NICHOLSON, H. A., and ETHERIDGE, R., JNR., 1879.—Descriptions of Palaeozoic Corals from
Northern Queensland, with Observations on the Genus Stenopora. Ann. Mag. Nat. Hist.,
(5) 4: 265.

, 1886.—On the Tasmanian and Australian Species of the Genus Stenopora Lonsdale.
Ann. Mag. Nat. Hist., (5) 17:173.

Nixkirorova, A. I., 1933.—Stratigraphic Range of the Carboniferous Bryozoa of the U.S.S.R.
Trans. Un. Geol. Prosp. Serv. U.S.S.R., Fasc. 268.

RaGceatTt, H. G., and FLeTcHerR, H. O., 1936.—A Contribution to the Permian-Upper Carboni-
ferous Problem and an Analysis of the Fauna of the Upper Palaeozoic (Permian) of
North-West Basin, W.A. Rec. Australian Musewm, 20 (2).

REED, F. R. C., 1933.—Anthracolithic Faunas of the Southern Shan States. Records Geol. Surv.
India, 67: 838.

Rep, J. H., 1930.—The Queensland Upper Palaeozoic Succession. Queensland Geol. Surv.
Publication 278.

TEICHERT, C., 1941.—Upper Palaeozoic of Western Australia: Correlation and Palaeogeography.
Bull. Am. Assoc. Petroleum Geologists, 25 (3) :371.

, 1947.—Stratigraphy of Western Australia. J. Roy. Soc. N.S.W., 80 (3): 81.

ULRICH, E. O., 1890.—Palaeozoic Bryozoa. Geol. Surv. Ill., Vol. 8.

WAAGEN, W., and WENTZEL, J., 1886.—Salt Range Fossils. Palaeontologica Indica, (13) 1.

WARTHIN, A., 1930.—Micropalaeontology of the Wetumka, Wewoka, and Holdenville Forma-
tions. Oklahoma Geol. Surv. Bull. 53.

123

SUCCINOXIDASE OF POTATO TUBER.

By ADELE MILLERD, M.Sc.
(From the Department of Biochemistry, University of Sydney.)

(Two Text-figures.)
[Read 29th August, 1951.]

Synopsis.

The participation of a tricarboxylic acid cycle, similar to that of animal tissues, in the
aerobic metabolism of plants would imply the participation of succinic dehydrogenase, an
enzyme which, in animal tissues, is coupled with the cytochrome system, the whole forming
the complex known as_ succinoxidase. Investigations carried out with potato (Solanum
tuberosum lL.) tuber have shown that this tissue contains an active succinoxidase system,
that the system is associated with particulate components of the cytoplasm and that succinic
dehydrogenase is coupled to the cytochrome system in the plant as in the succinoxidase of
animal tissues. Also associated with these particles, which resemble and may be identical
with mitochondria, are the enzymes, fumarase and malic dehydrogenase.

INTRODUCTION.

The participation of a tricarboxylic acid cycle, similar to that of animals tissues,
in the aerobic metabolism of plants would imply the participation of succinic dehydro-
genase, an enzyme, which, in animal tissues, is coupled with the cytochrome system,
the whole forming the complex known as succinoxidase. The in vitro demonstration of
succinic dehydrogenase activity in plant material has, however, been accomplished in
relatively few instances. Okunuki (1939) found succinoxidase activity in pollen of
Lilium auratum. Slight succinic dehydrogenase activity has been detected in extracts
of seedlings of some Leguminosae (Damodaran and Ramaswamy, 1940). Damodaran
and Venkatesen (1941) demonstrated the succinoxidase system in preparations of young
seedlings and pods of certain Leguminosae and examined in detail an active preparation
of the enzyme from seedlings of Phaseolus mungo. Goddard (1944) has also reported
a low succinic dehydrogenase activity associated with cytochrome oxidase preparations
from wheat germ.

From an examination of the effect of malonate on the accumulation of succinate,
Bonner (1948) concluded that succinic dehydrogenase must function in the metabolism
of pyruvate by segments of Avena coleoptile. Laties (1949) has also demonstrated the
accumulation of succinate in the presence of malonate in spinach leaves and in excised
barley roots.

The purpose of the present investigation was the study of the succinoxidase system
of the potato tuber both in relation to succinic dehydrogenase and in relation to the
cytochrome system. During the course of the investigation work bearing on this
problem has appeared from other laboratories. Cytochrome oxidase activity in prepara-
tions from potato tuber has been demonstrated by Levy and Schade (1948) and studied
in detail by Goddard and Holden (1950).

In general the results of the present work show, as do those of Levy and Schade
and of Goddard and Holden, that potato tubers contain an active succinoxidase system,
that the system is associated with particulate components of the cytoplasm, and that
succinic dehydrogenase is coupled to the cytochrome system in the plant as in the
succinoxidase of animal tissues.

PLANT MATERIAL.

Mature tubers from Solanum tuberosum L. (variety: Factor, grown at Blayney,
N.S.W.) were used throughout. Fifty pounds of freshly harvested tubers were stored
at approximately 15°C. and aliquots removed as needed. No systematic differences in

NN

124 SUCCINOXIDASE OF POTATO TUBER,

succinoxidase activity were noticed during the storage period of five months, nor were
differences found between potatoes from the two successive harvests used during the
investigation.

REAGENTS.

Cytochrome c was prepared from horse cardiac muscle according to the method of
Keilin and Hartree (1937), but was dialysed against distilled water rather than against
1% sodium chloride (Potter, 1941). The concentration was determined by measuring
the absorption at 550myu after oxidation with ferricyanide and reduction with
hydrosulphite.

Calcium cyanide was made according to Robbie and Leinfelder (1945) and estimated
gravimetrically by precipitation as silver cyanide.

Phosphate buffers (a mixture of Na,.HPO, and KH.PO,) were used throughout.
Hydrogen ion determinations were made by means of a Leeds and Northrop glass
electrode pH meter.

All solutions were made from A.R. chemicals.

Substrates and inhibitors were neutralized to litmus with sodium hydroxide where
necessary. 2 )

METHODS.

Succinoxidase activity was followed by measurement of oxygen uptake according
to the standard Warburg manometric technique. Potassium hydroxide was employed
in the centre well for absorption of carbon dioxide. In an examination of the effect of
cyanide on the aerobic system, calcium cyanide—calcium hydroxide mixtures (Robbie,
1946) were employed. Carbon dioxide output was measured by the direct method of
Warburg. Determination of dehydrogenase activity was done by the Thunberg technique
earried out in an atmosphere of nitrogen. Estimates of activity were based on’ the time
needed for complete decoloration of the redox dye used. All experiments were conducted
at 37°C. Succinic and fumaric acids were estimated quantitatively according to Krebs
et al. (1940). Chromatographic identification of organic acids followed the procedure of
Lugg and Overell (1948).

PREPARATION OF H}NZYME.

The tubers were first scrubbed in running tap-water, immersed for one minute in
a saturated, filtered solution of bleaching powder (CaClOC1), and thoroughly rinsed in
tap-water. The potato (300 gm.) was then diced into 100 ml. of 0:02M phosphate buffer
(pH 10:2). The diced material was next ground for one minute in a mechanical
macerator (overhead-drive Waring blendor). The pH of the mixture was maintained at
approximately 7:2 during this operation by the addition of 1N sodium hydroxide,
bromthymol blue being employed as an external indicator. The brei was now filtered
through muslin and allowed to stand in the refrigerator for ten minutes, after which
the supernatant was. decanted from the sedimented starch and centrifuged at approxi-
mately 1700g for ten minutes. The supernatant from the centrifugation was removed
by decantation and the residue, which contained the enzyme, suspended in 20ml. of
001M Na,HPO, for experiments on succinoxidase and in 20ml. of 0-01M KH.PO, for
experiments on succinic dehydrogenase activity. Two millilitres of this preparation,
containing 0-8—1:0mg. total nitrogen, was used in each experimental vessel.

All vessels and solutions employed were chilled before use and the entire preparation
of the enzyme was carried out at a temperature of 5°C. or below.

EXPERIMENTAL.

Demonstration of Succinic Dehydrogenase Activity.

Potato tubers contain an active succinic dehydrogenase as judged by ability to
catalyse the reduction of 2,6-dichlorphenolindophenol at the expense of succinate under
anaerobic conditions in the Thunberg technique. The enzyme is an insoluble one, bound
to particles which are sedimented in 10 min. in a field of 1700g, as noted above.
Microscopic observations show that the preparation as described above consists of
particles roughly ly in diameter.

BY ADELE MILLERD. 125

The standard method adopted for preparation of the particle-bound enzyme involved
sedimentation by centrifugation from the extract at pH 7:2. The pH of the extract
before and during centrifugation is critical, since it exerts an effect on the amount of

TABLE 1.

Effect of pH during Preparation on the Endogenous Substrate Level of
Succinic Dehydrogenase from Potato Tuber.
Main tube: 2-0 ml. enzyme, 0:5 ml. 0:2 M phosphate buffer (pH 7-3),
0-1 ml. 0-2 M succinate or 0-1 ml. water.
Sidearm: 0-3 ml. 0:05% 2,6-dichlorphenolindophenol.

Decoloration Time (Min.).
pH of
Extract. No Added Added
Substrate. Succinate.
Expt. 1 7-96 100 4:5
7:53 100 4-5
6:48 100 5
Expt. 2 6:48 55 3065)
6-00 20 4
Expt. 3 6-02 33 4
5:76 21 4
5-42 9 3
Expt. 4 5:35 3 1
5-02 1 1
4:46 1 al
3:98 1 1
3-44 1 il

endogenous substrate contained in the enzyme preparation. The data of Table 1 show
that when the extract is maintained at acidities greater than pH 5, the enzyme prepara-
tion obtained contains material which causes the rapid reduction of the redox dye
in the Thunberg technique even without the addition of further substrate. This effect
is minimized when the extract is maintained at pH values above 7.

Even preparations made by grinding potato tubers in water contain some succinic
dehydrogenase activity, as is shown in Table 2. The preparation for this experiment
was made by treating diced potato with one-third of its weight of distilled water for

TABLE 2.

Succinic Dehydrogenase Activity of Enzyme Prepared by
Maceration of Tissue in Distilled Water.

Main tube: 2-0 ml. enzyme, 0:5 ml. 0:2M phosphate
buffer (pH 7-3), 0-1 ml. 0-2 M succinate or 0-1 ml.
water.

Sidearm: 0:3 ml. 0:05% 2,6-dichlorphenolindophenol.

Decoloration Time
(Min.).
Control £5 4 A ye 35
+Succinate .. na Ke, Bt 8 =

two minutes in a Waring blendor. The enzyme was then separated by centrifugation
as described above.

pH.—The succinic dehydrogenase of potato was found to have a pH optimum of 6:5
in phosphate buffer, as shown in Text-figure 1. Succinic dehydrogenase from muscle
(Ohlsson, 1921) and from bacteria (Cook and Alcock, 1931) has heen shown to

126 SUCCINOXIDASE OF POTATO TUBER,

demonstrate maximal activity, as measured by the rate of decoloration of methylene
blue at pH 9.

20

150

oO
°
x
Z 15
=
=
: 100
= c
z =
2 10 S
= 4
fo} =z
=)
°o
re
2 50
yo ;
w
&
c
OG 7 8 oe 7 8
pH pH
1 2
Text-figure 1.—Effect. of pH on succinic dehydrogenase activity. Main tube: 2:0ml. enzyme,
0-5ml. 0:-2M phosphate buffer, 0-I1ml. 0:2M _ succinate. Sidearm : -3ml. 0:05% 2,6-dichlor-
phenolindophenol.
Text-figure 2.—Effect of pH on succinoxidase activity. Each flask contained 2-:0ml.

enzyme and 0-5ml. 0:2M phosphate buffer. The sidearm contained 0:2ml. cytochrome ec (final
concentration 2 x 10M), 0-1ml. 0:2M succinate or water to 0:3ml.

Redox Dyes.—The activity of potato succinic dehydrogenase as measured in the
Thunberg technique is markedly affected by the nature of the redox dye employed
(Table 3). Methylene blue and thionine were relatively ineffective as hydrogen acceptors,
while 2,6-dichlorphenolindophenol proved quite effective.

TABLE 3.
Effect of Nature of Redox Dye on Succinic Dehydrogenase Activity.
Main tube: 2:0 ml. enzyme, 0:5 ml. 0:2 M phosphate buffer (pH 6-5), 0-1 ml. water,
0-1 ml. 0-2M succinate or 0-1 ml. water.
Sidearm: 0-3 ml. dye.

Decoloration Time (Min.).
Cone.
Redox Dye. (M.). No Added Added
Substrate. Succinate.
2,6-dichlorphenolindophenol a5 5x10-* © | No apparent change in 15 hours.
10-3 120 33
10- 75 2-2
Ome 0:2 0:08
Thionine (Lauth’s violet) sa 10-3 No apparent change in 240 min.
10-* 240 50
10-5 10 9
Methylene blue At ate ao 10m No apparent change in 240 min.
Ome ” ” ”
Ome ” ” ”

On the basis of the data of Table 3, 2,6-dichlorphenolindophenol at a final con-
centration of 10-*‘M was chosen for all further Thunberg experiments. -
Succinate Concentration.

Data on the relation of potato succinic dehydrogenase activity to substrate con-
centration are presented in Table 4. The Michaelis-Menten constant, Km, derived from

BY ADELE MILLERD. 127

these data, is 1:9 x 107M. The Km for muscle succinic dehydrogenase has been found
to be 1 x 10°M (Lardy, 1949).

TABLE 4,
Effect of Concentration of Succinate on Succinic
Dehydrogenase Activity.

Main tube: 2-0 ml. enzyme, 0:5 ml. 0-2 M phosphate
(pH 6-5), 0-1 ml. water, 0-1 ml. succinate or
0-1 ml. water.

Sidearm: 0-3 ml. 10°? M 2,6-dichlorphenolindophenol.

Succinate Concentration Decoloration Time
(M.) (Min.).
75
6:67 x 107? 8308}
3-33 x 107? 2-5
1-67 x 107? Bye
6:67 x 1073 830[5)
3:33 x 1078 4:5
1-67 x10-3 6:2
6:67 x 1074 10-5
Soo Om 36

Inhibitors—Succinic dehydrogenase of animal tissues is known to be competitively
inhibited by malonate (Krebs and Eggleston, 1940) and to be sensitive to SH-reagents
(Potter and DuBois, 1948). Succinic dehydrogenase of potato tuber was likewise
inhibited by malonate as well as by phenylmercuric acetate, a potent SH-reagent. It
was not, however, inhibited by iodoacetate, a relatively weak SH-reagent (Table 5).

TABLE 5.
Effect of Varied Inhibitors on Potato Succinic Dehydrogenase.
Main tube: 2:0 ml. enzyme, 0-5 ml. 0:2 M phosphate buffer (pH 6-5), 0-1 ml.
0-2M succinate, 0-3 ml. inhibitor or 0-3 ml. water.
Sidearm: 0-3 ml. 10-°M 2,6-dichlorphenolindophenol.

Decoloration
Inhibitor. Conc. (M.). Time (Min.).

Phenylmercuric acetate A ine — 1:2
10-3 75
2x1074 75
1Om4 24
2x10-5
Ome
Ome
Malonate as we ob a —_—
10-?
10-8
10-4
10m—e
107°
Indoacetate a BG we as —
10-?
Ore
10-4
Ome
107§
Cyanide fee te ae ~ ——
10-2
Om
10-4
Ome
1@-e

<

-Q*

DION HON HH HHH HH HE pO HH Dp

* The slight increase in the rate of decoloration of the redox dye as noted
here, was shown by control experiments to be due to an effect of the inhibitor
in the absence of added substrate,

128 SUCCINOXIDASE OF POTATO TUBER,

Demonstration of Succinozidase Activity.—The potato enzyme preparation described
above is not only able to oxidize succinate anaerobically, but is also able to oxidize
succinate aerobically with the uptake of oxygen. This oxidation is markedly increased
by the addition of cytochrome ec to the reaction mixture, as is shown in Table 6. Succinic
dehydrogenase of potato tuber appears therefore to be, like the corresponding enzyme
from animal tissue, linked with the cytochrome oxidase system and may thus be
referred to as a succinoxidase.

TABLE 6.
Presence of Succinoxidase in Particulate Potato Succinic
Dehydrogenase.
Each flask contained 2:0 ml. enzyme and 0°5 ml. 0:2 M
phosphate buffer (pH 7-0).
The sidearm contained 0-2 ml. cytochrome c (final con-
centration 2x10~5M), 0-1 ml. 0-2M succinate or
water to 0:3 ml.

ul. O. Uptake
per Hour.
Control (no substrate, no cyto-
chrome) .. Ws ae oe 0
+cytochrome c os ae 4
+succinate at ah Be 11
-+cytochrome c+succinate .. 80

pH.—The succinoxidase of potato tuber shows a pH optimum in phosphate buffer
of 7-2 (Text-fig. 2). The system as a whole thus has a pH optimum different from that
of succinic dehydrogenase, which has been shown above (Text-fig. 1) to be ca. 6:5. The
pH optimum for muscle succinoxidase has been found to lie in the range pH 7:3-7:8
(Wieland and Frage, 1929).

Inhibitors —The succinoxidase system is sensitive to phenylmercuric acetate,
malonate, cyanide and iodoacetate (Table 7). Comparison of the results of Tables 5
and 7 shows that phenylmercuric acetate is an effective inhibitor of both succinic
dehydrogenase and of the complete succinoxidase system. The complete system is even
inhibited by concentrations of phenyl-mercuric acetate (10°M), whieh have little effect
on dehydrogenase activity. Malonate at high concentrations (10°M and 10°M) is an
effective inhibitor of both dehydrogenase and oxidase activity. Again, however, at a
concentration of 10-*M, which exerts no effect on the dehydrogenase, some inhibition
of the succinoxidase system is still observed. Cyanide has no inhibitory effect on
succinic dehydrogenase activity, but is an effective inhibitor of succinoxidase presumably
exerting its effect on cytochrome oxidase. Iodoacetate slightly inhibits the succinoxidase
system but has no effect on dehydrogenase activity.

Since phenylmercuric acetate is known to inhibit enzyme systems by its effect on
sulphydryl groups, the possibility of reactivating these groups, by glutathione, as
suggested by Barron and Singer (1945) was examined. The results of these attempts
were, however, entirely negative (Table 8).

Fate of Metabolized Succinate—The disappearance of succinate from a reaction
mixture containing potato succinoxidase may be shown to correspond quantitatively to
the reaction:

—2H + 1/20, +H,0

Succinic acid —————-—> Fumaric acid Fie Malic acid
—H,O
(1) (2)

The increased oxygen uptake displayed by the system in the presence of succinate
and cytochrome c takes place without concurrent increase in the small endogenous
carbon dioxide evolution (Table 9). It must be concluded, therefore, that although the
succinate is oxidized by the system it is not dismembered through further reactions of
the Kreb’s cycle,

BY ADELE MILLERD. 129

TABLE 7.
Effect of Inhibitors on Succinoxidase Activity.
Each flask contained 2-0 ml. enzyme, 0:5 ml. 0-2 M phosphate buffer (pH 7-2), and
0-3 ml. inhibitor or 0-3 ml. water. The sidearm contained 0-1 ml. cytochrome ec
(final concentration 10~>M) and 0-1 ml. 0-2M succinate.

Concentration ul. O2 Uptake %
Inhibitor. (M.). per Hour. Inhibition.
Phenylmercuric acetate an = 32
NO=9 0 100 °
10-4 0 100
10-> 17 47
10-8 28 13
Malonate Bs Ae on — 81
10-? 16 80
1078 44 46
10-4 49 40
10-5 62 24
10-° 66 19
Cyanide He ars as — 100
10-3 12 88
10-4 30 70
1075 54 46
Iodoacetate .. ou we — 81
2x10? 60 26
10-2 69 15
10-8 75 7
10-4 76 6
TABLE 8.

Effect of Glutathione on Inhibition of Succinoxidase by Phenylmercuric Acetate.
Hach flask contained 2:0 ml. enzyme and 0:5 ml. 0:2M phosphate buffer (pH 7-2).
The sidearm contained 0-1 ml. cytochrome c (final concentration 10-° M), 0-1 ml. 0-2 M succinate,
0-1 ml. phenylmercuric acetate (final concentration 10~° M), 0-1 ml. glutathione or water to 0-5 ml.

ul. O,
per Hour.
Control a6 3 is ae we a a ae or ae ae 0
+succinate+cytochrome c .. ale ~ at sik re Be ot 40
+succinate +cytochrome c+phenylmercuric acetate .. N56 eee ye 24
+succinate +cytochrome c+phenylmercuric acetate+glutathione (107? M) 28*
+succinate-+cytochrome c+phenylmercuric acetate+glutathione (1073 M) 15*
+succinate +cytochrome c+phenylmercuric acetate+glutathione (10~* M) s 16*

* These figures have been corrected for autoxidation of glutathione under the conditions of the
experiment.

TABLE 9.

Oaidation of Succinate: Examination of Oxygen Absorption and Carbon Dioxide
Evolution.
Each flask contained 2:0 ml. enzyme and 0:5 ml. 0:2 M phosphate buffer (pH 7-2).
The sidearm contained 0-2 ml. cytochrome c (final concentration 2 x 10-5 M},
0-1 ml. 0:2M succinate or water to 0:3 ml. ~

—ul. O» +ul. CO,

per Hour. per Hour.
Control .. Ae zis o% as Ae 0 5
+succinate Be a fs ee 0 7
+succinate+cytochrome c .. te 56 7

130 SUCCINOXIDASE OF POTATO TUBER,

Identification of Reaction Products.—Identification of the actual oxidation products
(Table 10) was carried out as follows. The aerobic oxidation of succinate was followed
manometrically in the usual fashion. At the end of the experimental period (one hour)
the manometer flasks were removed from the bath and to each cup was added l1ml.
5% metaphosphoric. acid and the contents mixed. The contents of each flask were
filtered through No. 1 Whatman filter paper, the flask washed repeatedly with small
quantities (2-3ml.) water and the washings collected. The filtrate and washings were
pooled and divided into two portions. On one portion succinic acid was estimated with
the aid of heart muscle succinoxidase. The other portion was reduced in such a way
as to convert any fumaric acid to succinic and the total succinic and fumaric acids
then’ determined as succinic acid by the muscle preparation (Krebs et al., 1940).

TABLE 10.
Quantitative Examination of the Fate of Added Succinate.
Eash flask contained 2:0 ml. enzyme (total nitrogen 0-8 mg.) and 0:5 ml. 0:2 M phosphate buffer (pH 7 -2). The
sidearm contained 0-1 ml. cytochrome c (final concentration 10~* M), 0-2 ml. 0-2 M succinate or water to 0-3 ml.

Succinate
Disappearance
ul. O» Corresponding Succinate
Succinate Absorbed in to Observed Disappearance
Added (Mg.). One Hour. Oxygen (Analytical)
Consumption (Mg.).
(Mg.).

Control ie ae 0. 5 0 0
+eytochrome c .. 0 7 0 0
+succinate 4-4 23* 0-19 0-2
+succinate +eytochrome c 4-4 132 1:39 1-4

* The enzyme preparation employed in the final section of this paper was prepared by centrifugation at —5° C.;
the lower temperature maintained during the preparation resulted in an enzyme with some succinoxidase activity
without added cytochrome c.

The validity of this procedure was established by recovery experiments carried out
under the conditions of the experiment with succinic and fumaric acids added in
amounts such as might be expected to be present. The method is accurate to +0:1mg.
succinic acid under the conditions of the experiment, and 80-90% recovery was obtained
with added fumarate.

TABLE 11.
Identification of Organic Acids formed during the Aerobic Oxidation of Succinate.

Rf Values of Plant Acids.
Acid Detected

Acid Tested Acid in in Reaction

Separately. Mixture. Mixture.
Succinic Ae Ae 0-701 0-694 0-687
Fumaric a 46 0-818 0-811 0-817
Malic .. ea oe 0-448 0-444 0-449
Solvent: n-butanol/formic acid/water. Chromatographic separation carried
out at 4°C.

If only reaction (1) were involved, fumaric acid should be present in amounts that
would be readily estimated by the method used. Fumaric acid did not, however, appear
as expected. On the contrary, only small increases in succinate concentration after

BY ADELE MILLERD, 131

reduction were obtained. It was, therefore, suspected that fumarase might be present
in the system and might then convert the fumarate formed to an equilibrium mixture
of fumarate and malate (Krebs ef al., 1940). That this is indeed the casé and that both
fumarate and malate are products of the oxidation of succinic acid by the potato enzyme
preparation was demonstrated with the aid of paper chromatography (Table 11).

The aerobic oxidation of succinate was followed by the Warburg technique. At the
end of the experimental period (one hour) the flask contents were acidified (pH 2) with
sulphuric acid and extracted with ether in a continuous extractor for thirty-six hours.
The ethereal extract was concentrated, 2-0ml. water added, the ether removed by heating
on a water bath, and the residue concentrated to approximately Iml. Ten ul. of this
extract was then applied to one end of a strip of filter paper and the chromatogram
developed with butanol saturated with formic acid. The control (no added substrate)
from the Warburg experiment was also extracted and examined in the same manner
but no organic materials were detected. Comparable Rf values for succinic, fumaric
and malic acids were determined by parallel experiments with the individual acids as
well as with a mixture of the three acids. Twenty ug. of each acid was used in each case.

DISCUSSION.
It has become increasingly evident that the particulate fraction of the cytoplasm .
is intimately concerned with the respiratory activity of animal tissues, since on these
particles, the mitochondria, are located inter alia, all the enzymes necessary for: the
metabolism of pyruvate to carbon dioxide and water via the Krebs cycle (Green e¢ al.,
1948). It is of interest, therefore, that in plants also it can be demonstrated that some
at least of the enzymes participating in the tricarboxylic acid cycle are located on
particles present in the cytoplasm—particles which would appear to correspond to the
mitochondria of animal tissues. It has been possible to prepare particles similar to
those described in this paper but which possess malic dehydrogenase activity in
addition to the enzymes succinic dehydrogenase, fumerase and cytochrome oxidase.
It will be of interest, therefore, to determine if further enzymes of the cycle are
located on these particles and whether or not particles may be prepared from plant
tissues which are capable of carrying out the complete oxidation of pyruvate by way
of the Krebs cycle.

- SUMMARY.

An enzyme complex has been prepared. from potato tuber which is capable of
converting succinic acid to a mixture of fumaric and malic acids. In this complex
the conversion of succinic acid to fumarie aid is coupled to the uptake of molecular
oxygen via the cytochrome system. The enzyme complex is associated with particles
which resemble and may be identical with the mitochondria.

ACKNOWLEDGEMENT.
This work has been supported by a grant from the Commonwealth Research

Committee.

References.

BarRRon, HE. S. G., and SINGER, T. P., 1945.—Studies on biological oxidations. 9. Sulfhydryl
enzymes in carbohydrate metabolism. J. Biol. Chem., 157: 221-240.

BONNER, J., 1948.—Biochemical mechanisms in the respiration of the Avena coleoptile. Arch.
Biochem., 17: 3811-326.

Cook, R. P., and Autcock, R. S., 1931.—The effect of hydrogen ion concentration on some
oxidations by B. coli communis. Biochem. J., 25: 523-533.

DAMODARAN, M., and RAMASWAMY, R., 1940.—Succinic dehydrogenase in germinating seeds.
Current Sci., 9: 319-320.

, and VENKATESEN, T. R., 1941.—Amide synthesis in plants. 1. The succinoxidase

system in plants. Proc. Indian Acad. Sci. B., 13: 345-359.

GODDARD, D. R., 1944.—Cytochrome ec and cytochrome oxidase from wheat germ. Am. Jd.
Botany, 31: 270-276.
, and HOoLpEN, C., 1950.—Cytochrome oxidase in the potato tuber. Arch. Biochem.,

27: 41-47.

132 SUCCINOXIDASE OF POTATO TUBER,

GREEN, D. E., Loomis, W. F., and AUERBACH, V. H., 1948.—Studies on the cyclophorase system.
1. The complete oxidation of pyruvic acid to carbon dioxide and water. J. Biol. Chem.,
172: 389-403.

KeEILIN, D., and HaArtrer, EH. F., 1937.—Preparation of pure cytochrome c from heart muscle
and some of its properties. Proc. Roy. Soc. (London) B., 122: 298-308.

KREBS, H. A., and HEGcuEston, L. V., 1940.—The oxidation of pyruvate in pigeon breast
muscle. Biochem. J., 34: 442-459. A

, SMYTH, D. H., and Evans, E. A., 1940.—Determination of fumarate and malate
in animal tissues. Biochem. J., 34: 1041-1045.

Larpy, H. A., 1949.—Respiratory enzymes. Revised edition. Burgess Publishing Company,
Minneapolis, Minn., U.S.A.

Latiges, G. G., 1949.—The oxidative formation of succinate in higher plants. Arch. Biochem.
22: 8-15.

Lrvy, H., and ScHADE, A. L., 1948.—Studies in the respiration of the white potato. 2. Ter-
minal oxidase system of potato tuber respiration. Arch. Biochem., 19: 273-286.

Luee, J. W. H., and OvERELL, B. T., 1948.—‘“‘One” and ‘‘two-dimensional” partition chromato-
graphic separations of organic acids on an inert sheet support. Aust. J. Sc. Res. A.,
1: 98-111.

OHLSSON, E., 1921.—Die Abhadngigkeit der Wirkung der Succinodehydrogenase von der
Wasserstoffionenkonzentration. Skand. Arch. Physiol., 41: 77-100.

OKUNUEI, K., 1939.—Uber den Gaswechsel der Pollen. 38. Weiters Untersuchungen tiber die
Déhydrasen aus den Pollenkérnern. Acta Phytochim., 11: 65-80.

PottTer, V. R., 1941.—Studies on the mechanism of hydrogen transport in animal tissues.
4. The succinoxidase system. J. Biol. Chem., 141: 775-787.

, and DuBois, K. P., 1943.—Studies on the mechanism of hydrogen transport in
animal tissues. 6. Inhibitor studies with succinic dehydrogenase. J. Gen. Physiol.,
26: 391-404. Fz

RosBig, W. A., 1946.—The quantitative control of cyanide in manometric experimentation.
J. Cellular Comp. Physiol., 27: 181-209.

, and LEINFELDER, P. J., 1945.—Calcium cyanide solutions as constant sources of
hydrogen cyanide gas for animal experiments. J. Ind. Hyg. Towxicol., 27: 269.

WIELAND, H., and FRAGE, K., 1929.—Uber den Mechanismus der Oxydations vorgange. 20.
Beitrage zur Kenntniss der Bernsteinsaure-dehydrase. Ann. Chem., 477: 1-32.

133

THE NOMENCLATURE OF HETERONYCHUS SANCTAE-HELENAE BLANCHARD
(COLEOPTHRA: SCARABAEIDAH; DYNASTINABE).

By EH. B. Britton,
Department of Entomology, British Museum (Natural History).

Synopsis.

Heteronychus sanctae-helenae Blanchard is an important pest of crops and grassland in
coastal New South Wales. The reasons for the change of the name of the species by Arrow,
in 1937, from arator F. to sanctae-helenae Blanchard, are examined. It is concluded that
' H. sanctae-helenae Blanchard is the valid name, while arator F., the type of which is in
Kiel, must be transferred to the genus Hybosorus.

Heteronychus sanctae-helenae Blanch. is an important pest of crops and grassland
along the New South Wales seaboard. Its distribution extends from Byron Bay in the
north to Moruya in the south (Wallace, 1946). The species is a native of South Africa
and was introduced into New South Wales about the year 1920, the first record of its
occurrence being that of, Gurney (1934). In Western Australia it was first observed
in 1938 at Albany; it is now widespread in the Perth area. In 1949 it appeared in
numbers in the neighbourhood of Adelaide.

The species (as an adult) is responsible for considerable damage to gramineous
crops, especially maize, and to vegetable and florists’ crops. It is troublesome in turfed
areas such as golf greens and grass tennis courts. The importance of the pest in pastures
is not known, but the larvae may well be an important factor in the decline of pasture
grasses after drought.

Until -1937 this species was known in the literature and in collections as
Heteronychus arator (F.), 1792. The later name, Heteronychus sanctae-helenae
Blanchard, 1853, was adopted by Arrow in his Catalogue of Dynastinae of 1937, with
arator Burmeister (nec Fabricius) as a synonym, implying that arator F. was a different
species and that Burmeister had misidentified it.

The reason for this change was not immediately clear, as the specimen labelled
Geotrupes arator in the Banks collection in the British Museum certainly belongs to
the species which we now know as sanctae-helenae. Because of the importance of the
species as a pest I have thought it advisable to investigate the matter and if possible
to stabilize the change of name. Fabricius described many species from the Banks
collection specimens, so that there was reason to believe that this specimen might be
regarded as the type. Burmeister’s redescription of H. arator F. is based on this
specimen, as he states that he saw a specimen named by Fabricius, standing as
Geotrupes arator in the Banks collection, and adds that this species was wrongly
referred to Hybosorus by Illiger and Schonherr.

Prell (1936) states that the Geotrupes arator (F.) in the Fabrician collection in
Kiel is a species of Hybosorus, and therefore is not even a Dynastid.

There are, therefore, two specimens of different species, one in the British Museum,
one in Kiel, either of which might be considered to be the type of arator KF. The choice
must depend on the Fabrician description.

It is noteworthy that in the descriptions of the species immediately before and
after that of arator, and elsewhere, Fabricius writes “Mus. Dom. Banks’, indicating
that the description is based on a specimen in the Banks collection. This is omitted in
the case of arator, the implication being that the type is not in the Banks coliection
but elsewhere. In addition, the description states “Clypeus integer, subscaber”. It is
more likely that Fabricius would have described the clypeus of H. sanctde-helenae as
“scaber”’.

(0)

134 NOMENCLATURE OF HETERONYCHUS SANCTAE-HELENAE.

The evidence therefore favours the acceptance of the specimen in Kiel as the type
of Scarabaeus arator F. The species represented in the Banks collection then becomes
Heteronychus arator Burmeister (nec Fabricius), 1847, but the continued use of the
name arator in Heteronychus is inadmissible, as it is a secondary homonym. The next
available name for the species, Heteronychus sanctae-helenae Blanchard, 1853, is
therefore valid.

I am indebted to Messrs. P. B. Carne, C. F. Jenkins and C. R. Wallace for the
information ov the status of the grass in New South Wales.

Bibliography.
FABRICIUS, 1792.—Entomologia Systematica, 1:33 (Scarabaeus arator).
Fapricius, 1801.—Systema Eleutheratorum, 1:21 (Geotrupes arator).
BURMBISTER, 1847.—Handbuch der Hntomologie, 5:94 (Heteronychus arator).
BLANCHARD, 1853.—Voyage au Poéle Sud., Zool., 4: 105, Pl. 7, f. 6.
WOLLASTON, 1869.—Ann. Mag. Nat. Hist., (4) 4: 312 (AH. arator = sanctae helenae).
PERINGUEY, 1901.—Trans. S. Afr. Phil. Soc., 12: 521, Pl. 40, f. 7.
JAcK, 1924.—Trans. Ent. Soc. Lond. (1923), 1924: 415, f. 20. (Monograph of Heteronychus.)
Paoutr, 1934.—Boll. Soc. Ent. Ital., 66:51.
PRELL, 1936.—Hnt. Blatt., 32:149 (arator F. transferred to Hybosorus).
ARROW, 1937.—Cat. Col. Dynastinae, pars 156: 31.
WALLACE, C. R., 1946.—Agric. Gaz. N.S.W., 57: 121.

135

CONTROLLED POLLINATION OF HUCALYPTUS.
By L. D. Pryor.
(Plate vi.)
[Read 29th August, 1951.]

Synopsis.
Successful cross-pollination in several pairs of Hucdlyptus species and the methods

employed, are recorded.
Self-fertility has been found in all species examined, but possibly some self-sterile indi-

viduals exist.
Attempts at wide crossing were unsuccessful.

INTRODUCTION.

At various times the possibility of hybridization between species in the genus
Eucalyptus has been discussed and in the taxonomic treatment of the genus some
described species have been considered as possible hybrids by both Maiden and Blakely.
There has been little if any record, however, of controlled hand-pollination within the
genus apart from some successful attempts which have been made by Brett (1949) and .
also Gilbert and Martin (1949) in Tasmania.

Recently abundant evidence has been produced by Brett (1949) and Pryor (1950) to
show that under field conditions hybrids between several well-established species can
be found quite easily. ‘The evidence for this has been principally morphological and
that secured from progeny tests. A rigorous demonstration of the contentions requires
that, in addition, the supposed hybrids shall be synthesized by controlled pollination.
A eritical examination of the methods of pollination, both artificial and natural, is also
necessary in many other ways. It is important to know the extent of self-sterility in
the genus, the amount of out-crossing in self-fertile species, and similar things before
we can hope to understand the complex genetic structure within the genus. This genetic
knowledge seems essential before Hucalyptus taxonomy can be placed on a scientific basis.
Barber (1950) has strongly emphasized this need for genetic knowledge and the experi-
mental method in taxonomy in general in Australia, while some knowledge of the
mechanism of reproduction is a prerequisite to breeding Hucalyptus for economic or
aesthetic needs.

POLLINATION METHODS.

The method of reproduction in the species examined is one which aids cross-
‘fertilization of each individual flower. Usually the pollen is shed soon after the
operculum falls, and from most anthers often within the first twenty-four hours.
However, to a varying degree some inner anthers retain pollen, which is not shed until
later, and then as they unfold more or less brush the stigma and no doubt encourage
self-pollination if there has not already been successful pollination. The stigma in most
cases is apparently not receptive until several days after the operculum falls, and then
it seems to remain receptive, often for four or five days, and sometimes up to ten days.
This means that the flowers are protandrous, which is common in many genera. In a
few flowers examined a little pollen has been shed before the operculum falls, but the
extent to which this occurs cannot be determined without more examination. Apparently
Mueller had this in mind when he expressed the opinion that cross-pollination is unlikely
in a genus where a calycine lid guards the stigma.

There seems to be little wind-pollination and undoubtedly the genus is principally
insect-pollinated. The arrangement of the stamens, stigma and nectaries, as well as the
time of maturation of pollen and stigma, favours cross-pollination in any one flower

136 CONTROLLED POLLINATION OF EUCALYPTUS,

with the help of insects. The flowers when open attract many kinds of insects, and in
some localities birds may be important.

Emasculation is a simple matter and may be effectively carried out, just before
the operculum falls, by cutting through the tissue just below the staminal ring but above
the top of the ovary. With most species the stamens and operculum come away freely
leaving the stigma uninjured, but in a few species examined the stigma is tightly
enclosed by the filaments and the stamens and operculum cannot be removed, if the
stigma is to remain undamaged, without a second cut longitudinally to free them.
If this has to be done the speed of emasculation is greatly reduced.

The method of preparing flowers for controlled pollination is quite easy; the
principal experimental difficulty is to prevent burning of the prepared flowers after
bagging. Where there is no free air circulation, such as by the use of brown paper
bags, or alternatively transparent cellulose bags, the flowers and young stems often die.
This is particularly the case with those species which flower in mid-summer or autumn.
Burning can generally be prevented by using muslin bags, but these possibly permit the
entry occasionally of stray pollen which, while of little importance in a general breeding
programme, may be troublesome if some critical examination is being made of repro-
ductive methods. While not easy with large trees, shading in some way is necessary
in these cases. Improved methods for this work should result with more investigation.

After pollination it is usually apparent in three months whether the fruit has set.
Unfertilized fruit does not as a rule develop and often falls. With most species examined
the fruit is ripe in ten to twelve months after pollination and may be harvested then.
_ The seed may be developed and viable a little earlier, but the fruit is often difficult
to open if not fully ripe and the seed may not be plump.

Very little fruit is set and almost no fertile seed obtained from fiowers that have
been emasculated and placed in muslin bags, showing that wind-pollination is prac-
tically absent and in fact the occasional pollination which occurs probably results from
a very small insect, such as a thrip, a chance dropping of pollen or perhaps some
pollen shed before the stamens were removed.

In most of the trials fresh pollen was used from flowers opening at the time the
prepared flowers of the seed parents were receptive. Two pollinations, seven days
apart, were made in most cases. There~is little doubt that pollen can be stored,
however, and successful germination was obtained with pollen that had been stored a
month in a desiccator at. 50% humidity and at about 35°F. Germination tests can be
made successfully with many species, using a 10% honey solution and placing in an
incubator at 70°F. The most striking feature from preliminary tests of this kind
was the great variability in germination percentage between pollen from flowers on
different branches of the same tree and even at times between different flowers on the
same branch. Germination percentage declines after three months’ storage in a
refrigerator... Dry storage at room temperature results in quicker loss of power to
germinate.

These preliminary tests indicate that separation of flowering times by as much as
three months, and probably more, is not likely to be an obstacle to successful cross-
pollination if the species are compatible and no barriers to successful fertilization exist.
Much more study of the possibilities in this direction is necessary.

INTERSPECIFIC HYBRIDIZATION.

After the basic method had been successfully determined as above, a series of
trials was made using principally E. bicostata and E. Maidenii as female parents. These
trials aimed to assess the possibilities of interspecific hybridization and to learn where
general barriers might exist within the genus. Because it was necessary, for the
reasons described, to use muslin bags the chance of some occasional stray pollen
fertilizing a flower could not be entirely removed. This means that some of the low
numbers of fertile seed obtained must be regarded with suspicion. "Where there have
been substantial quantities of seed obtained—about fifty or more from ten to twenty
flowers—there is no doubt that the pollination has been successful, and this is at once

BY L. D. PRYOR. UB57/

obvious from the seedlings because of the marked differences obtained in the lots of
seedlings from the same female parent but with different pollen parents.
Table 1 shows the results obtained with the species mentioned as female parents.
Keeping in mind the limitations of the data due to the failure to exclude all
possibility of chance pollination or perhaps occasional self-pollination before the
operculum falls, the trend is at once clear in compatibility in cross-pollination.

TABLE 1.
Number of
Flowers Number of Number of
Seed Parent. Pollen Parent. Treated Fruit. Plants Raised.
(Approx.).
E. Maidenii F. Muell. | #. bicostata Maiden, Blakely & 40 35 90
Simmonds.
E. rubida Deane & Maiden uy 12 10 35
E. Blakelyi Maiden 3 ar 10 7 33
FE. Macarthuri Deane & Maiden Ae 10 4 10
E. Maideni F. Muell. a3 Nee 6 4 16
E. cinerea F. Muell... an ae 10 3 16
FE. ficifolia F. Muell. ie ih; 10 2 1*
jE. Bridgesiana R. T. Bak. a 10 1 —_—

E. bicostata Maiden, | #. bicostata Maiden, Blakely & 15 12 Harvested too
Blakely & Simmonds Simmonds (another tree). early. Germina-
(Horse yards). E. Maidenii F. Muell. Sad ae 155 12 tion except for

E. Maidenii F. Muell. ae at 15 10 a few plants a
E. Maidenii F. Muell. Fe She 10 7 failure.

E. Blakelyi Maiden se BG 10 2

E. ficifolia F. Muell. i ae 10 1 '
E. cinerea F. Muell... ae +h 6 —

Self .. ae ae a Fa 8 —

HE. bicostata Maiden, | Self .. : a ae 15 15 Not yet raised.
Blakely &Simmonds | LE. Maidenii F. Muell. tf oe 10 a
(Acton Park). tE. Bridgesiana R. T. Bak. ae 10° 1

TH. delegatensis R.T. Bak. .. £% 6 —
E. Macarthuri Deane & Maiden .. 8 —
E. Blakelyi Maiden | LE. Blakelyi Maiden 6 4 12
(Capital Hill). E. rubida Deane & Maiden 5 2 3
E. cinerea F. Muell... Bie a 5 1 —
E. melliodora A. Cunn. ss =< 10 — —
Control (not poll.) # — —

E. Blakelyi F. Muell. | #. cinerea F. Muell.. . a He 12 8 ; 9
(Acton).

E. cinerea F. Muell. | EZ. Blakelyi F. Muell. 38 & 6 5 21
(Arboretum).

* Appears to be pure H. Maidenii.

7 =H. Stuartiana F. Muell.

£=H. gigantea Hook. f., non Dehnh.

Highly successful cross-pollination is achieved by using pollen of the same species
and of more or less closely related species, for example, HH. Maidenwi with £. bicostata,
EH. Blakelyi, HE. Macarthuri and EH. cinerea, which are all in the anther group of
Blakely, Macrantherae; whereas H. ficifolia and H. Maidenii, E. bicostata and E. dele-
gatensis, EH. Blakelyi and E. melliodora are pairs in widely separated groups, and
practically no setting of fruit has been obtained. :

There is no doubt that successful crosses between #H. Maidenii and EH. Blakelyi;
H. \Maidenii and H. cinerea; H. Maidenii and H. rubida; FE. Maidenii and H. Macarthuri;
and HE. Blakelyi and EH. cinerea have been achieved, because of the distinct characters
already developed in these progenies,

138 CONTROLLED POLLINATION OF EUCALYPTUS,

In the main, the Fl hybrids show juvenile characters intermediate between the
parents.

Striking evidence of successful cross-pollination is furnished by the #. Maidenii x
E. Macarthuri hybrids due to the incorporation of the highly distinctive geraniol oil of
E. Macarthuri in the Fl hybrid after having been transmitted by the pollen of
E. Macarthuri.

The trend is therefore clear that the more widely separated the species within the
genus, the less likely that cross-fertilization can be achieved. To determine the absolute
limits and whether any progeny obtained is fertile will require more experiment.

This supports the field evidence that hybrids between species within the different
anther groups of Macrantherae, Renantherae and Terminales are rare or absent.

In one ease, in addition to H. Maidenii x FE. bicostata the reciprocal cross has also
been made between £. cinerea and H#. Blakelyi. The hybrid progeny with ZH. cinerea the
seed parent, and the other with #. Blakelyi the seed parent, is intermediate, at the ten
pairs of leaves stages, in morphology between the parents. Both progenies are closely
similar to each other but at the same time quite distinct from open-pollinated seedlings
of each parent.

SELF STERILITY.

Krug and Alves (1949) have stated that with the species of Hucalyptus examined
in Brazil they believe them self-sterile. This is contrary to the condition revealed in
the species examined here. The species critically examined are substantially different
from those used by Krug and Alves, the latter dealing with species common cn the
north coast of New South Wales, in the coastal climate. while the species which have
been subject to experiment here grow naturally on the cold Southern Tablelands. It is
clear that those on which the experiments have been made can successfully self-
pollinate because, if flowers are bagged before the operculum falls (and in one experi-
ment with #. Maidenii double bags were used consisting of an outer one of muslin and
an inner one of brown paper), good fruit is set, giving fertile seed. On the other
hand, if the flowers were emasculated and then bagged, fruit did not set and no fertile
seed was obtained. In a further experiment with the three species, H. Maidenii,
E. Blakelyi and EH. bicostata, bagging, emasculation and hand-pollination with pollen
of the same plant were followed by good setting of fruit and production of ample fertile
seed. This particular experiment on #H. bicostata was duplicated on separate trees. It
led to an interesting case of individual variation which must be examined in more
detail. In the second tree, unlike the first, flowers bagged before opening did not set
seed, therefore apparently not selfing. In addition, flowers emasculated and bagged
did not set when self-pollinated with other flowers of the same tree, but similarly
emasculated and bagged flowers pollinated with pollen from another tree of H..bicostata
set ample fruit.

This suggests that either the particular tree is self-sterile or produced defective
pollen in 1949, or perhaps does so always. Further examination,is necessary to find
the complete explanation, but in any one of the three cases there could be a factor
of considerable importance in the production of hybrid seed in quantity.

SUMMARY.

Simple trials show that many species of Eucalyptus are protandrous, entomophilous,
and with a mechanism to aid cross-fertilization of each flower but some capacity for
self-pollination if crossing fails.

Emasculation is easy and controlled pollination simple, provided the parents are
fairly closely related within the genus. Successful controlled pollinations have been
achieved with several pairs of species, and in two cases reciprocal crosses have been
made.

Absolute barriers and the limit of crossing, although indicated, have yet to be
accurately determined. Self-fertility exists in a number of species.

Burning of flowers and stems following bagging must be carefully guarded against.

BY L. D. PRYOR. 139

References. 4

BARBER, 1950.—Taxonomy. Aust. Journ. Sci., 12: 184.

Brett, 1949.—A.N.Z.A.A.S., Hobart, xxviii: 149.

GILBERT and MARTIN, 1949.—Unpublished communication.

Krue@ and Atves, 1949.—Eucalyptus Improvement. Journ. Hered., 40: 133-139, 143-150.
Pryor, 1950.—A Hybrid Eucalyptus. Proc. LINN. Soc. N.S.W., 75: 96-98.

EXPLANATION OF PLATE VI.

Fig. 1.—Progeny obtained from hand-pollinated capsules from the one tree of EH. Maidenii.
The four plants at the left are the result of cross-pollination with H. rubida. Central four
plants, cross-pollination with H. cinerea. The four plants on the right, pollination with
EF. Maidenii from another tree of the same species. The variation in population in form clearly
indicates the success of the controlled pollination.

Fig. 2.—Progeny obtained from controlled pollinations on a single seed parent individual
of HE. Maidenii. The four plants on the left are the result of pollination with FH. Blakelyi pollen;
the central four, H. bicostata, and the four plants on the right are from open-pollinated capsules
on the same tree. The marked variation in the form of the progeny as well as other
characteristics, such as colour, not apparent in the photograph, indicates the success of the
pollination.

Fig. 3.—Progeny from a seed parent of H. Maidenii. The four plants on the left are from
open-pollinated capsules on the parent tree. The four plants on the right are as the result
of hand-pollination with H. Macarthuri. Characters intermediate with H. Macarthuri are
apparent in the more tapering leaves, particularly noticeable in the first and third individuals
of the group. The unmistakable presence of geraniol in’ these individuals is striking confirmation
of the hybridization. ;

Fig. 4.—The four plants on the left were obtained from a EH. cinerea seed parent after
erossing with #. Blakelyi by hand. The four on the right were raised from seed from open-
pollinated flowers. The intermediate characters of the hybrid population are at once apparent.

Fig. 5.—The four plants on the left were obtained as the result of hand-pollination of a
tree of H. Blakelyi with H. cinerea. The progeny is intermediate. The four plants at the
right were the result of open-pollination on the same tree. This is a reciprocal cross of species
in Fig. 4, but not with the same individuals.

140

A GENETIC ANALYSIS OF SOME HUCALYPTUS SPECIES.
By L. D. Pryor.
(Plates vii—xi.)
[Read 29th August, 1951.]

Synopsis.
Genetic analysis by progeny testing based principally on morphological seedling characters,
has revealed probable F1 hybrids, and segregating swarms between several species. There is at
present no evidence of hybridization between certain major taxonomic groups of the genus.

INTRODUCTION AND METHODS.

The genus Hucalyptus is a complex one in which the separation of species by the
ordinary taxonomic method is difficult. As it has become better known the number of
described species has become more numerous. Even so, it is still easy to find
individuals and even populations which do not fit any described species as well as is
ordinarily the case within the majority of genera. One cause of trouble is that some
of the described species cover a latitude of variation which is much smaller than
that of others. In short, there has been “splitting” in some groups and not in others,
and it has been somewhat cynically observed that the density of Hucalyptus species
increases as one approaches centres of botanical study.

The past taxonomic approach in the genus has resulted in the naming, usually as
species, of divergent forms of varying biological status. If this process were continued
it would result in a multiplication of species which would contribute little to the
understanding of the genus or to its practical classification. A more reasonable
taxonomic approach involves the reclassification of many forms as geographical or
ecological subspecies, but the difficulty does not end there because some of the
variation is not of the geographic kind which would allow conveniently the erection
of subspecies.

Some variation encountered suggests that there is active hybrid development with
F2 and back-cross segregating swarms. Many of the Fl hybrids have been given
specific names, e.g. H. uwnialata (Brett, 1987 and unpub.). There is another type of
variation which again it is difficult to place in the category of subspecies. Hucalypt
forests show always a complex mosaic pattern, one species giving way to another
under very slight changes of such ecological conditions as aspect, edaphic qualities
or other slight habitat changes. The one species as it occurs in successive pieces of the
mosaic may show slight but: fairly constant differences. Brett (1937) has termed these
species polymorphic and suggests that the polymorphs have arisen by fixation under
varying conditions of selection of segregates of previously existing swarms. The
situation as he describes it shows many similarities to the introgressive hybridization
of Anderson (1949). These categories of variation, if they exist, present problems for
taxonomy which have so far not been satisfactorily resolved.

From accumulated knowledge of field variation of Hucalyptus on the Southern
Tablelands and Highlands of New South Wales an attempt has been made at a genetic
analysis of several of the species of this region and preliminary evidence which fully
supports many of Brett’s contentions has been obtained. The condition with Hucalyptus,
Brett suggests, seems to arise from the large number of species often growing side
by side, the general lack of sterility barriers between species, the stage of evolution
of the genus, the effect of sub-recent climatic changes and the still more recent effect
of settlement.

As luck would have it, one of the most consistent and distinctive sets of
characters possessed by different species is the morphology of the juvenile leaves.

BY L. D. PRYOR. x ‘ 141

This factor permits in many cases a preliminary assessment of the genetic make-up of
a plant within a few months by the simple means of growing seedlings from selected
individual trees. Apart from this source, which has been the principal means of
getting information, a series of preliminary experiments on controlled pollination has
been carried out which has indicated clearly that hybridization in many cases is a
simple matter (Pryor, 1951).

‘ SEGREGATION IN OPEN POLLINATED PROGENIES.
a. EH. pauciflora Hybrids. :

The situation in one of the hybrids included under “EH. vitrea’ has been discussed
previousiy (Pryor, 1950) where the individual concerned was a hybrid between
E. paucifiora and #. dives. Seed was collected from a somewhat similar tree with
morphological characters intermediate between #. pauciflora and EH. Robertsonii.
Seedlings were raised and fifty unselected plants in a block were taken from the flat
and grown on in tubes (PI. vii, fig. 2). At eight inches’ height they exhibit characters
showing complete segregation to parental types and intermediates (see Table 1). For
some specific characters an approximation to the Mendelian ratio for the independent
assortment of characters in the F2 generation is achieved. The juvenile foliage of the
putative parents is quite distinct. In the case of H. Robertsoni (Pl. vii, fig. 1) it is
narrow-lanceolate, sessile, thin, opposite for a large number of pairs, glandular hairy
on the stems. In #H. pauciflora, on the other hand, the leaves soon become petiolate
and become alternate after four or five pairs, they are thick and rather broadly oblong,
and the stems are almost smooth. The population obtained has some individuals
which are indistinguishable from seedlings of #. Robertsonii, and others which cannot
be separated from #. pauciflora. There is little doubt that the parent tree is an Fl
hybrid and that the generation raised, which is probably from seed that has been
produced by selfing, is an F2 generation. Similar individuals can be found almost
wherever there is a junction in the field between H. pauciflora and E. Robertsonii,
although they are not numerous and never form a pure stand or even a consistent
major component of a stand.

In the area in which this seed was collected there was a considerable number of
trees which would be considered H. Robertsonw. Nevertheless, though this is the only
possible determination amongst described species, many of the individuals are inclined
to be smooth-barked on limbs three or four. inches in diameter, somewhat heavy-leaved
and with fruits that are coarser than are often found on #. Robertsonii. From one of
these trees a small amount of seed was also collected and about twelve plants were
raised. While the number was small, three of the twelve plants corresponded with
individuals in the progeny from the Fl hybrid which were intermediate between the
two parental types. The deviation from an ordinary population of H. Robertsonii was
far too great to do other than lead to the conclusion that the parent tree in this
case could not have been “pure” H. Robertsonii (Pl. vii, fig. 3). Had there been no
morphological indications of this, one might have concluded that some of the seed in
-the capsules was of hybrid origin. Since most of the flowers are probably selfed on
-the one tree, if not individually self-fertilized, the variation’in the progeny in the
light of the morphological character of the parent is consistent with what one would
expect in an F3 or later generation, or from a back-cross. It is quite probable in the
regrowth area concerned that there is a population containing many segregates between
H. paucifiora and E. Robertsonii (Pl. viii, fig. 4) with a bias in survival towards those
individuals with a preponderance of H. Robertsonii characteristics.

Field examination of many stands suggests that mingling between various ecotypes
of #. pauciflora and ecotypes of Peppermints (#H. radiata, EH. Robertsonii and EH. dives)
is common, and this is perhaps one of the most frequent hybrid combinations found
on the Southern Tablelands.

EH. pauciflora enters into other combinations in addition. A stand was examined
at Barney’s Range, near Cooma, where there is an unusual occurrence side by side
of H. paucifiora and H. Rossii. After some search two trees were found which were

P

142 A GENETIC ANALYSIS OF SOME EUCALYPTUS SPECIES,

intermediate between these two species. Seed was collected from one and 50 plants
again raised without selection in pricking off from the flat. Complete segregation to
the putative parents was again attained although the juvenile characteristics of these
two species are not so clearly distinctive as in the previous case (Pl. viii, fig. 5).
Nevertheless the trend is unmistakable and the amplitude of variation in morphological
characters and the correspondence of the extreme forms closely with juvenile foliage
of the recognized putative parent species is such that there is no other readily acceptable
explanation than that we have an F2 population.

E. pauciflora has characteristics which are almost unique in the genus in its
practically parallel veins in the leaves, the very thick leaves and large pyriform fruit.
It appears these characters persist prominently in hybrid combination and the presence
ot E. pauciflora genes is therefore readily detected. Progeny tests are not yet complete
but it is highly probable that #. pauciflora x EH. delegatensis hybrids may be readily
found, and several trees are known which seem to be Fl hybrids of this combination.
One tree was also found on the dividing range near Badja which looked like a
hybrid between EF. pauciflora and E. fastigata. On the other hand, so far no individuals
have been found in the field which suggest that H. pauciflora enters into combination
with Gums such as #. Dalrympleana or E. Blakelyi, or with Boxes such as #. melliodora,
with all of which it can’ easily be found growing side by side.

b. E. Rossii Hybrids.

Eucalyptus Rossii, which, as has been mentioned, hybridizes with HE. pauciflora,
is commonly found in hybrid combination with other species. One of the commonest
of these is with #. macrorhyncha. Using the same procedure as in the examination otf
E. pauciflora and Peppermints, a tree on Mount Jerrabomberra was examined. It
has field characters intermediate between H#. macrorhyncha and H. Rossii. The bark is
smooth on limbs of six inches diameter or less, the fruit is rather small, with a flat-
domed disc compared with H. macrorhyncha; the leaves are also small and shiny for
‘normal’ H. macrorhyncha. The progeny obtained from seed of this tree provides a
most striking example of segregation and independent assortment of many characters
(Pl. viii, fig. 6). The juvenile characters of the two species are strikingly distinct.
E. macrorhyncha (Pl. viii, fig. ™)) has ovate-lanceolate or broadly elliptical leaves
covered densely with “stellate” hairs. EH. Rossii (Pl. ix, fig. 8), on the other hand, has
narrow-lanceolate, glabrous, grey-glaucous leaves. This affords an outstanding example
of the existence of an Fl hybrid and it is so distinctive that there is little need for
further demonstration.

in the same locality seed was also collected from another tree with similar morpho-
logical characteristics. The variation in the progeny was much more restricted than
in the first tree, although the trend is clearly in the same direction to the putative
parents, #. macrorhyncha and H. Rossii (Pl. ix, fig. 9). The conclusion, therefore, is
that though the morphological form of the parent is closely similar to that of the
undoubted F1 hybrid, this second tree must be a member of a later filial generation
or back-cross of the same hybrid combination in which the genetic variability has been
considerably reduced (see also Pl. ix, fig. 10).

The existence of Fl hybrids has been established on the basis of evidence of this
kind between several other pairs of species. #. Rossii x E. dives (Pl. ix, fig. 11) is
found quite frequently and progeny tests show marked segregation in some individuals.
whereas other individuals closely similar morphologically show less, as in the case of
E. macrorhyncha x E. Rossii combinations.

#. Rossii also hybridizes freely with EH. Robertsonii where these two species occur
side by side. Blakely (ined.) proposed a variety of H. Robertsonii on material from |
the Australian Capital Territory which is close to, if not identical with, the F1 hybrid
EB. Rossii x EH. Robertsonii. The juvenile foliage of these two species is so distinct
that segregation is at once apparent in seedlings (Pl. x, fig. 12).

BY L. D. PRYOR. 143

ec. Miscellaneous Hybrids.

A common hybrid combination is between H. Robertsonii and EH. fastigata, and F1
individuals have been located which segregate markedly when propagated (PI. x, fig. 14).
The segregation is easily discerned because the juvenile characters of each parent are
so distinct from those of the other.

Other hybrids have been found and tested between H. Blakelyi and E. elaeophora
(Pl. x, fig. 15) and between #. maculosa and H. elaeophora (PI. xi, fig. 16). Individuals
are occasionally found in which the morphological characters are intermediate between
EH. Blakelyi and E. elaeophora. Segregation of characters is again marked in the
seedling stage and the sessile, opposite, orbicular, glaucous juveniles of H. elaeophora
are so strikingly distinct from the stalked ovate-lanceolate, green, alternate, juvenile
leaves ot H#. Blakelyi that observation of segregation is quite simple. An Fl hybrid has
been progeny-tested and other similar individuals have been located on Black Mountain
(Pl. x, fig. 15). |

Likewise, extreme differences exist between the juvenile foliage of H. maculosa
and #H. elaeophora, and a progeny test of an anomalous individual has given a filial
population showing marked segregation to the putative parental types in the juvenile
foliage (Pl. xi, fig. 16).

Another interesting individual is an undoubted Fl hybrid between #H. viminalis
and H#. glaucescens. This individual was found on the Tinderry Ranges while examining
the vegetation in company with R. G. Brett. Segregation in the progeny is easily
recognized for the same reasons (Pl. xi, fig. 18).

Further combinations are being investigated but, since adequate diagnostic
characters are not displayed in the juvenile forms, results will not be available until
the progeny is more mature. Continued collection will produce more evidence as a
wider number of species can be: examined.

Table 1 shows the way in which characters have been observed to segregate in
the different progenies examined.

TAXONOMIC AND HVOLUTIONARY IMPLICATIONS.
a. Nomenclature.

The analysis given above is still incomplete, as it is based on juvenile characters
only, but problems of classification in Hucalyptus are profoundly affected by facts
established by genetic analysis of this kind. There is little doubt that segregating
swarms are common in the field and individuals from these have often been referred
to species while the types of some so-called species are probably members of such
swarms. The description of segregating forms as species is useless for taxonomy and
the classification of such individuals should be made differently. One of the first
steps in the revision of the genus Hucalyptus must be to determine which of the
' described ‘“‘species” are Fl hybrids or segregates.

The position with regard to hybrids known as “Hucalyptus vitrea’ has been
discussed previously and some additional similar or related matters have been disclosed
in the present study. For example, the Fl hybrid and the segregate between H. Rossii
and #. macrorhyncha have both in the past been referred to H. brevirostris. Comparison
of the material from the mature trees with the type of #H. brevirostris shows that
neither of these can be placed with that type, although they have some characters in
common, notably the buds and the general fruit size. However, the rim of the fruit
is quite different and the leaves are also different. There are some characters in
E. brevirostris which suggest affinity with HE. regnans on the one hand and a Stringybark
on the other. The record of distribution of this species also suggests that it might
well be a hybrid, as indeed Blakely himself supposed. It cannot, however, be a hybrid
between #H. macrorhyncha and H. Rossii. There is no type, therefore, with which the
Fl hybrid #. macrorhyncha x EH. Rossii corresponds. It is undescribed at present, and
in any case should not be described as a species.

Another hybrid combination of particular interest is that of #. fastigata and
E. Robertsonii. This illustrates one way in which the genetic approach has a marked

ECIES,

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UCALYPTUS

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ANALYSIS OF

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BY L. D. PRYOR. 145

bearing on taxonomic conclusions. H#. radiata and H. Robertsonii were regarded as |
distinct species by Blakely. One of the means of separation of these species is the
appearance of sub-parallel venation in H#. radiata as distinct from #. Robertsonii. The
twe species are obviously very closely related.

The presence of hybrids and segregates of H. fastigata and H. Robertsonii in the
field accounts for the sub-parallel venation which is sometimes found in areas where
the trees correspond generally with H. Robertsonii. One of the uncertainties which has
led to the suggestion that H. Robertsonui and H. radiata should not be separated as
distinct species is removed if it is recognized that this lack of consistency between
the two species is, in the case of H. Robertsonii, a result of hybrid influence by
H. fastigata, and the decision as to whether the two are “good” species rests on
whether the two populations are sufficiently distinct.

There is no doubt that the type of H#. Robertsonii which was collected by Blakely
and de Beuzeville on Talbingo Mountain represents the population of the western part
of the Southern Highlands, which has a number of characters different from H. radiata,
the type of which comes from the Blue Mountains. These characters, though not
marked, are nevertheless fairly distinctive. It seems, therefore, that H#. radiata and
E. Robertsonii, according to the definitions accepted in the concept, could be regarded
either as species or alternatively as subspecies within the #. radiata group.

On the other hand, the position of H. Westonii is similar to that of H. vitrea. The
hybrid between H. maculosa and H. elaeophora is identical with H. Westonii, the type
locality of which is Mt. Majura, A.C.T.

In the vicinity of the type tree there are populations of segregates which vary more
or less continuously between each of the supposed parents. There is little doubt that
E. Westonii is an F1 hybrid between EH. maculosa and E. elaeophora and that it is in
consequence genetically unstable. It therefore should not be regarded as a species.

b. “Phantoms” and Clines.

The detection of the Fl hybrid between H. Rossii and H. Robertsonii has already
been described where stands of these two species are side by side in the field; however,
in other areas somewhat remote from stands of H. Robertsonii individuals are found
which are intermediate between EH. Rossii and a Peppermint of the H. radiata type.
Usually there is no #. radiata or H. Robertsonii present, as at Black Mountain, KOE,
and Mt. Jerrabomberra, N.S.W., but the progeny from the tree shows a limited degree
of segregation which unmistakably reveals genetic diversity beyond that which is
normal in one of the accepted species (PI. x, fig. 13). The best explanation for such
occurrences is that given by Brett (1949). He supposes that the genes of the H. radiata
are preserved in hybrid form to some extent as a relict from former times when,
probably due to somewhat different climate, the H. radiata was much more widely
distributed. Brett has referred to such occurrences as a “phantom” of a species, and
this appears to be quite apt. The same condition has been found with ZH. fastigata
genes preserved in H. macrorhyncha on Mt. Macdonald, A.C.T. (Pl. ix, fig. 10). This
aspect must be studied more closely as information which relates critically to the
subject becomes available from the trials in progress.

H. Rossii also presents another phase of the same condition in that it appears to
contain, in many instances, genetic influence in varying degrees from Peppermints and
Stringybark species which could be expected as climatic changes and conditions for
survival of some species become more difficult. H. Rossii shows minor variations in
populations from place to place, apparently for this reason. Again this aspect must be
treated separately when more experimental data are available.

Another distinct condition resulting from gene movement is illustrated in the
clinal variation in #. fastigata and EH. radiata.

The hybrid #. radiata x H. fastigata is common on the dividing range in the vicinity
of Badja and in this locality exhibits an interesting feature. On the gentler grading
sheltered slope on the western fall of the range H. radiata is confined to a narrow strip
near the crest. At this point also hybrids of H. fastigata occur. As one then proceeds

146 A GENETIC ANALYSIS OF SOME EUCALYPTUS SPECIES,

down the slope, narrow-leaved, small-fruited forms of #. fastigata occur which gradually
become coarser and pass without discontinuity to typical H#. fastigata in the middle and
lower parts of the slope. This variation in a cline according to exposure grades from
a form intermediate between the two parent species to H. fastigata. A clinal sequence >
from the intermediate to HE. radiata is not to be found in this locality, presumably
because the site favours the combination on the #H. fastigata side of the mode of the
hybrid combination.

The same kind of clinal variation in individuals was observed from the area of the
occurrence of the Fl hybrid H. pauciflora x EH. Rossii, the gradation being from this form
to £. Rossii as the conditions become warmer.

c. Gene Invasion.

The evidence produced in the examination of H. glaucescens suggests that gene
invasion of this species by #. viminalis is a likely explanation of the facts.

E. glaucescens is an uncommon species which occurs in small communities on rocky
areas in sub-alpine regions. It has close affinity with two Tasmanian species, H. Gunnii
and #£. urnigera. This relationship and its scattered distribution suggest that it is a
relic of a group of species or a common precursor which flourished at some earlier
period in the south-east highlands and in Tasmania. Wherever it is in contact with
E. viminalis (and this is probably usual) it is being hybridized and invaded by the
more successful genes of that species. The evidence for this is that in the small
progeny test segregates intermediate in character between the apparent F1 hybrid and
the typical H. glaucescens are of low viability, whereas those segregates leaning to
E. viminalis are highly successful (Pl. xi, figs. 18 and 19). If this is so, it is an
illustration of gene invasion referred to by Brett (1950) and is one way in which
a species can be virtually extinguished. Specimens of #H. glaucescens collected by
A. B. Costin near the Victorian border have produced some plants tending towards
EH. viminalis (Pl. xi, fig. 19), thus clearly indicating the presence of the genes, though
the parent could not be taken for H. viminalis, and it could not be an F1 hybrid because
of the absence of segregation of the type expected in an F2 generation.

With #. Perriniana, by way of contrast, the position is quite the reverse. It is a
species of similar sub-alpine distribution, but, where examined, no trace of hybrids can
be found. While it is apparently a diminishing relic population, now occurring only
on hard sites, simple competition from more vigorous species appears to be the cause
of its near extinction.
ad. Sterility Barriers.

The evolution of the genus and the possible extent of the conditions described above
will be strictly limited by any barriers to breeding within the genus. These are not
yet fully known, but field occurrences suggest that hybridization is, at least under
natural conditions, confined within the major taxonomic divisions of the genus as given
by Blakely. Experimental controlled pollinations confirm this in oe terms but not
absolutely (Pryor, 1951).

On the other hand, Brett (1937) says there is some evidence of intergroup hybrids,
as, for example, #. ovata x EH. pauciflora, and EH. risdonii x E. viminalis.

While there is perhaps evidence of selection of similar characters, e.g., large leaves
or glaucousness, in unrelated species growing side by side in a particular habitat, a
careful search has so far failed to produce one of the wide crosses between the three
taxonomic groups, Macrantherae, Renantherae and Terminales, which often occur side
by side in the field on the Southern Highlands.

More critical work is necessary to establish the limits and such points as whether
such progeny is viable or, if so, whether it is fertile.

SUMMARY.
A genetic examination, principally by progeny testing, has been made of several
species common on the Southern Highlands of New South Wales and in Victoria.
Morphological seedling characters are of critical diagnostic value in the genus Hucalyptus

BY L. D. PRYOR. 147

and the segregation of some of these characters in progenies strongly suggests that a
number of the individuals examined are Fl hybrids. Other individuals similar in their
general morphology are revealed as probably hybrid but of a later filial or back-crossed
generation.

From the evidence obtained by progeny tests it is considered likely that segregating
swarms between well-established species occur in a number of cases. Under field
conditions hybridization between three of the major taxonomic groups of the genus
Eucalyptus, namely, Macrantherae, Renantherae, and Terminales, as given by Blakely,
has not been observed and it is considered that this, though perhaps possible, is unlikely.
Further work is in progress to examine the position with regard to other species -
combinations and to examine various aspects of genetic behaviour within the genus.

References.

ANDERSON, H., 1949.—Introgressive Hybridisation. New York.
Brett, R. G., 1937.—A Survey of Eucalyptus Species in Tasmania. Proc. Roy. Soc. Tas., 193
75-109.
, 1949.—A.N.Z.A.A.8., Hobart, xxviii: 149.
Pryor, L. D., 1950.—Proc. LINN. Soc. N.S.W., 75: 96-98.
——, 1951.—Proc. LINN. Soc. N.S.W., 76: 135-139.

7.
(

EXPLANATION OF PLATES VII-XI.
Plate vii.

Fig. 1.—#. Reberisonii. Progeny showing slight variation characteristic of a population
falling within the defined limits of the species without hybrid influence.

Fig. 2.—#H. paucifiora x E. Robertsonii, Cotter House, A.C.T. An F2 generation raised
from a probable Fl hybrid. Segregation is complete and there is independent assortment of
characters. The extreme left is identical with H. Robertsonii and the extreme right is
#H. paucifiora. The development of broad leaves, but practically sessile and opposite, in the
fifth from the left, is a mixture of E. pauciflora and E. Robertsonii characteristics, whereas the
petiolate leaves but relatively narrow, is a mixture of H. pauciflora and EH. Robertson charac-
teristics of a different degree. The development of alternate leaves is noticeable in the final
pair, especially of the fourth from the left and the seedling on the extreme right.

Fig. 3.—E. Robertsonii, Cotter House, A.C.T. There is unmistakable H. pauciflora in this
population, as is indicated by the seedling on the extreme left. The second from the left
shows thick and broad leaves but retains the Peppermint character of sessile, opposite leaves.
The seed parent of these seedlings appears to be Peppermint (H. Robertsonii) but was rather
clean-stemmed and with larger fruits and somewhat heavier leaves than is “normal’’. It
clearly contains EH. pauciflora genes.

Plate viii. :

Fig. 4.—EH. pauciflora, Cotter House, A.C.T. This shows a tendency to H. Robertsonii,
particularly in the individual at the extreme left. The leaves are narrower and inclined to
be sessile and opposite to an extent greater than that characteristic of EH. pawciflora, which is
typical in the two individuals on the right. The parent of this population, however, must have
been part of a hybrid swarm between EH. Robertsonii and E. pauciflora. ,

Fig. 5.—EH. paucifiora x E. Rossii. An F2 generation raised from an apparent F1 hybrid
from Barney’s Range. The thick broad-lanceolate leaves of the plants on the ‘left stand sharply
in contrast with the narrow-lanceolate, rather thin-stemmed and thin-petioled leaves on the
right. Characters which are more critically distinct may be expected to appear as the trees
become older, the juvenile characters being in this pair of species rather more similar than
in many others.

Fig. 6.—E. macrorhyncha x FE. Rossii. An F2 population showing complete segregation from
E. macrorhyncha type on the left to E. Rossii on the right. Note the change in leaf shape
and petiole length. The indumentum of “stellate” hairs characteristic of H. macrorhyncha
shows a similar gradation. There is a marked tendency for the intermediate hybrid type
(similar in form to the Fl hybrid), third from the left, to be the most vigorous.

Fig. 7.—H. macrorhyncha. A population characteristic of the species showing the typical
leaf shape and at times slightly crenate leaf margins.

Plate ix.
Fig. 8.—E. Rossii, Jerrabomberra. A population without hybrid influence showing relatively
uniform characteristics of the progeny.

Fig. 9.—H. macrorhyncha x HE. Rossii. Progeny from a tree\which was not an F1 hybrid,
showing a type tending to H. macrorhyncha in the plants at the left, and a type approaching
E. Rossii in the types at the right. Compare with Fig. 7, where the segregation runs from
one putative parent to the other, indicating a broader range of variation.

148 A GENETIC ANALYSIS OF SOME EUCALYPTUS SPECIES.

Fig. 10.—Progeny from an apparent H. macrorhyncha hybrid, Mt. McDonald, A.C.T. The
progeny suggests strong affinity with #. fastigata in the form on the right-hand side. The
two individuals at the left are closely similar with normal #. macrorhyncha. The fourth from
the left is roughly intermediate between H. macrorhyncha and HB. fastigata. This suggests the
preservation of B. fastigata genes in hybrid combination in a locality in which BP. fastigata
no longer exists.

Fig. 11.—#. dives x EH. Rossii. Note the broad, opposite, sessile leaves on the plant on
the left, grading to lanceolate, petiolate leaves approaching H. Rossii on the right. The fourth
from the left is becoming distinctly alternate. Probably not an F2, but a later generation
however.

Plate x.

Fig. 12.—Progeny from a parent belonging to the hybrid swarm EH. Rossii x EH. Robertsonii.
The plant on the left is close to EB. Robertsonii. Note the progressive development of stalked
leaves becoming alternate as one proceeds to the right, a generation later than an F2 generation.

Fig. 13.—Progeny from a tree on Mt. Jerrabomberra of a narrow-leaved Peppermint type.
Appears to be clearly a hybrid between a‘Peppermint of the #. Robertsonii or EB. radiata type
and #. Rossii, the characters of the putative parents being shown in the gradation from left
to right, the Peppermint being at the extreme left. The fifth from the right, however, indicates
a probable influence of #. dives, showing that this is not a simple hybrid. The occurrence of
this tree is remarkable, as there are no #. dives, EH. radiata or E. Robertsonii now on Mt.
Jerrabomberra. The trees from which the seed was taken to raise these plants are about
30 years old and appeared following disturbance in road making about 30 years ago.

Fig. 14.—Progeny from an F1 hybrid between £#. fastigata and EH. Robertsonii. Arranged
to show the sequence of characters from #2. fastigata on the left to #. Robertsonii on the right.
Note the change from alternate, petiolate broad-lanceolate leaves at the left, to opposite, sessile
narrow-lanceolate leaves on the right.

Fig 15.—Progeny from a hybrid between #H. elaeophora and HE. Blakelyi. Showing complete
segregation between the two putative parents, H. elaeophora on the left with opposite, orbicular,
sessile leaves, to #. Blakelyi with petiolate, broad ovate-lanceolate, alternate leaves at the right.
Note the gradations of characters between the two extremes and also the lack of vigour in
the intermediate types.

Plate xi.

Fig. 16.—Progeny from H. Westonii, Black Mountain, A.C.T. Indicating three main gene
types. At the extreme left, a Blue Gum resembling HE. goniocalyz, and the centre, types tending
to #. elaeophora or E. Bridgesiana, and at the right, EH. maculosa. A complex gene mixture that
needs further analysis. A variation in vigour of the various combinations is also interesting.

Fig. 17.—E. glaucescens x E. viminalis. Seedlings raised from a supposed F1 hybrid from
the Tinderry Mountains. Photographed in November, 1949. Compare with Fig. 18, in which the
order of arrangement is reversed, H. glawcescens being at the extreme right. The plant second
from the left, which is showing signs of weakness, is the second from the right in Fig. 18 and
is, in the six months’ interval, dead.

Fig. 18.—EH. glaucescens x E. viminalis—progeny raised from a supposed F1 hybrid collected

on the Tinderry Mountains. Note the segregation in the seedlings in the F2 generation between
E. vinminalis type at the extreme left and H. glawcescens at the right. Photograph taken in
May, 1950, indicating the difficulty in raising segregates which approach FH. glawcescens, the
fourth from the left being almost dead. Compare this plant with the same individual in
Fig. 17, which is the same population of five plants taken in November, 1949.

Fig. 19.—Progeny from a tree of H. glauwcescens showing the presence of H. viminalis
genes. The seed was collected by Mr. A. B. Costin from a tree, apparently H. glaucescens, at
Tingiringi. The unmistakable influence of EH. viminalis is apparent in the two individuals at
the right. a

149

INVESTIGATIONS OF THE PREFERENCES SHOWN BY AEDES (STEGOMYIA)
AHGYPTI LINN. AND CULEX (CULEX) FATIGANS WIED. FOR SPECIFIC
TYPES OF BREW®DING WATER.

By Tom MANEFIELD, B.Sc.,
Department of Zoology, University of Sydney.
[Read 29th August, 1951.]

Synepsis.

Comparative experiments are outlined showing the reaction of Aédes aegypti and Culex
fatigans to two types of water, representative of field habitats, these being tap water and a
“foul’ medium for which a manure infusion was used. Results are given showing the
oviposition responses of the adult females to both types of water and the survival percentages
of the larvae of the two species when reared in these media.

Each species showed preference for the foul medium for oviposition, that shown by
C. fatigans being completg. C. fatigans larvae were more able to survive than A. aegypti in
the foul medium at all stages of introduction into infusions of varying maturities. It was also
shown that development of a scum on the manure infusions was more lethal to the latter
species.

INTRODUCTION.

The reason for the differences shown by mosquitoes in the selection of larval
habitats is a problem which has received much attention from various workers. From
early observations in the field it was obvious that the occurrence of mosquito larvae
was not haphazard and that no one species is found in all types of water, marked
preferences being noticeable. Classifications of habitat have been given by Hopkins
(1936), who proposed a grouping according to the situation of the water (e.g. ground
water, rock pools), and Lee (1944) whose classification is based on the state of the water
itself (e.g. fresh, polluted). During earlier investigations masses of physico-chemical
data were obtained by such workers as MacGregor (1929), Senior-White (1934), Beattie
(1932). Few significant results were obtained, but Buxton (1934), examining Beattie’s
results, shows that her conclusion correlating larval prevalence with ammonia nitrogen
was significant. More recently work by Thompson (1940-41), Woodhill (1941) and
others concerning the selection by the adult female of the site of oviposition seems
to be of a more positive nature.

In this investigation A. aegypti and C. fatigans were selected as representing typical
eases of different breeding habits. A. aegypti is an entirely domesticated species, breeding
in and around human dwellings, usually in fresh water contained in artificial
receptacles. It is rarely recorded in foul water, whereas C. fatigans, on the other
hand, shows a distinct preference for such water. This investigation is of a preliminary
nature and was approached from two angles, namely, oviposition responses, comparing
the reaction of the two species to water which was believed to be typical of the location
in which the larvae have been recorded; and larval experimentation, to see if there
was any effect shown by the rearing of the species found in one habitat type, in the
alternate type. The two media used were a horse manure infusion and tap water (i.e.
the normal reticulated water supply of Sydney, which contains 0:0062% dissolved salts).

BREEDING TECHNIQUE.

The A. aegypti eggs were obtained from a stock which has been continuously bred
at the Zoology Department, University of Sydney, since 1938. Hgegs of C. fatigans,
which were collected from tubs outside the laboratory, were usually obtainable in
sufficient numbers all the year round, but in winter it was at times necessary to
maintain a laboratory culture. The technique used for breeding mosquitoes was
similar to that used by Woodhill (19386). A laboratory culture of A. aegypti was
continuously maintained, as fresh eggs were used during the larval experimentation.

150 PREFERENCES BY AEDES AEGYPTI AND CULEX FATIGANS FOR TYPES OF BREEDING WATER,

This was also done with C. fatigans when sufficient eggs were not obtainable outside
the laboratory.

The “foul” medium was prepared by mixing approximately equal volumes of dried
horse manure and water. These infusions were allowed to mature for three different
periods—7 days, 10-11 days, and 14-15 days. They were made up at varying times so
that the three states of maturity required would be available at the same time.

Pek EXPERIMENTATION.
Oviposition Responses. \
Males and females of the two species (about 100 of each) were released into
separate uniform cages, 12” x 10” x 9”, and given blood feeds and raisins. In each cage

TABLE 1.
Number of Eggs Deposited by C. fatigans and A. aegypti.
Culex fatigans. | Aédes aegypti.
| | Tap | Manure
Observation | Tap Water. | Manure Infusion. | Water. | Infusion.
Number. | | om ¥
| | |
Number of | Number of | Number of | Number of | Number of | Number of

Rafts. Eggs. Rafts. Bggs. Eggs. | Eggs.

1 Nil. Nil 4 299 299 489

2 a he 7 455 | 440 727

3 a Ke | 9 571 560 960

4 5 . 9 612 | 138 436

5 és e | 1 75 130 613

6 » i 9 635 / 39 741

7 he = 7 483 548 | 2,765

8 S es | 10 780 75 | 180

9 zs si 5 381 1,009 [eee s4i

10 i. a i | 351 545 1,521
11 A . 21 1 IS yay2 77 883
12 Za - 9 | 678 1053/9] 747
13 e ie 3 | 5 Ome 195 4,990
14 11 153
15 | | 166 | 805
16 | | | 495 | 8,472

| | }
Mean percentages ae 0 100 (Or) 82-3

were placed two oviposition dishes, one containing tap water and the other “foul”
water which had stood for at least ten days before use. They were placed at opposite
ends of the cage, the position reversed daily, and the cages placed parallel to the
window in order to eliminate any positional or phototactic response. The eggs were
collected and counted at regular intervals, and a record kept of the data and numbers
of females feeding. After collection of the eggs the contents of the oviposition dishes
were always replaced. This procedure was followed even when no eggs had been laid.
Results are given in Table 1.

Under statistical analysis the preference shown by both species for foul water
i
proved significant, giving the probability P = —.
232
Experiments with Larvae.
Fifty larvae at known stages were placed in uniform dishes containing 200 c.c. of
the manure infusions (at the required stages of maturity) plus food and incubated

ee

BY TOM MANEFIELD. i5aL

at 80°F. In each experiment controls were kept by placing larvae in tap water with
food in the normal manner. The amount of food used in each case was approximately
0:35 ¢m. The cultures were incubated and thespupae removed as they developed until
all the surviving larvae had pupated. The numbers of mature pupae resulting were
recorded.

The possibility of interference by scum formation was taken into account. For
comparison an effort was made in some cases to find the effect of the water alone,
without the interference of the scum. Removal of the scum daily with cotton wool
was satisfactory.

TABLE 2.
Percentage Aédes aegypti Surviving to Mature Pupal Stage.
Percentage Survival.
Scum
Larvae Stage of Condition Number of In Manure Infusions.
Introduction. y on Replications.| Control.
Surface. In Tap
Water. 7 Days 10 Days 14 Days
Maturing. Maturing. Maturing.
| |
sts. | |
| |
50 3rd stage larvae in each | |
replication .. as a Present. 6 99 | 70 | 88 98
50 2nd stage larvae in each | |
replication .. .. ..| Present. | 6 | 99 38 | 74 | 98
50 2nd stage larvae in each |
replication .. es Ae Removed. | 9 | 96 44 80
50 1st stage larvae in each | | | | |
replication .. #5 he Present. | 6 | 96 | 0 | 58 | * 94
50 1st stage larvae in each | | | |
replication .. 4 .. | Removed. | 9 | 96 | 3 oa
50 eggs introduced into each | | | |
replication .. ae OSE Present-—< | 12 | 99 | 4 44 94
50 eggs introduced into each | | | |
replication .. ae .. | Removed. | 6 | 99 | 9 | 90

| | | | |
| | | |

Table 2 gives the results obtained for A. aegypti at known stages -of introduction
into the “foul” medium cultures. These figures are compared with the controls taken
through at the same time in tap water.

Obviously from the results the lethal stage is during the first larval stage, probably
when newly emerged, so this stage was selected for a comparison of the effect on the
two species. Table 3 gives the results obtained when 50 newly emerged first-stage
larvae were introduced into each culture.

TABLE 3.
Percentage of A. aegypti and C. fatigans Surviving to Mature Pupal Stage.
Aédes aegypti. Culex fatigans.
| | |
|
Scum Number of | In Manure Infusion. In Manure Infusion.
Condition on | Replications. In Tap | In Tap
Surface. | | Water. | Water. |
; | Control. | 7-Day 11-Day Control. 7-Day 11-Day
| Infusion. Infusion. Infusion. | Infusion.
| | | |
| | |
| | |
Present | 6 99% 3% CUP mip 90% tI) )62% |) 92%
| | |
Removed Gumuener a gga, 24 b9R SCR oore |) | 7A 96%
| |

152 PREFERENCES BY AEDES AEGYPTI AND CULEX FATIGANS FOR TYPES OF BREEDING WATER,
The figures in Table 2 and Table 3 show the percentage survival of the two species.
For use in the summary these figures have been converted to percentage mortality by
subtraction from the 100% level.

DISCUSSION.
Oviposition Responses.

The potentialities of any water as a breeding ground depend primarily upon the
response of the female to that water. With the possibilities of phototactic and positionary
responses eliminated, results tor C. fatigans apparently ratified field observations. This
species showed a complete preference for foul water, laying 100% of the eggs on this
medium. In the field, C. fatigans is generally recorded in water with a high organic
content. The occurrence of this species in clean water would be due to the inability
of the female to find a more suitable breeding place.

A. aegypti did not show a complete preference for the “foul” water, but nevertheless
did have a very marked preference. In a total of 16 observations, when 29,905 eggs were
laid, 82% were deposited on the “foul” water. In the field, even when foul water
of the type used is available, A. aegypti larvae have rarely been recorded in it and almost
invariably occur in the water contained in rain water tanks and similar containers. In
all its usual breeding places the organic content is relatively low, yet in the laboratory,
under optimum conditions, the species preferred to lay its eggs on water with a high
organic content. These results seem to correspond with those obtained by Dunn (1927).
He tound that, by placing containers with various types of water in the open and
allowing A. aegypti to breed at will, the species showed a distinct preference for water
to which leaves had been added, rather than ordinary tap water. Apparently A. degypti
is attracted by some factors which are indicative of a higher food content.

From the evidence of these findings further oviposition experimentation is indicated,
giving a comparison of varying degrees of foulness or putrefaction rather than a
comparison with tap water, and also types of putrefaction, for example, using leaves.
It must also be noted that during these experiments the infusions used were all over
ten days’ maturity. Considering the larval findings with infusions of seven days’
maturity, it may be interesting to see the results of similar experiments using only the
seven-day infusion. A rejection point may be obtainable, giving a lead to the
actual limiting factor.

Larval Cultures.

The horse manure infusion proved a satisfactory medium and was easy to handle.
Initial observations were made on these infusions by exposing them to field conditions
so that the approximate time of entry of C. fatigans could be determined, giving the
time at which the infusion became a potential breeding ground. As the seventh day
after preparation was found to be the critical time for laying of eggs, it was decided
to see if there was any effect on A. aegypti by placing larvae in this medium. As
C. fatigans was then able to breed in these cultures it was considered unnecessary to
carry out comparable experiments between these species at first. The method was
altered variously to show the effect of infusions of different degrees of maturity on the
larval stages. Table 2 shows the results in a condensed form.

No statistical analysis was necessary for the verification of these results. The
major diminution of the population, such as in the introduction of first-stage larvae and
eggs into the seven-day infusion, was-obviously significant. The effect of the seven-day
infusions on all stages was the most noticeable. The earlier the larval stages at
introduction, the greater the mortality became; however, there was a slight rise in
survival with the introduction of eggs. In all cases the removal of the scum led to a
slight rise in survival rate. After ten days’ maturity the infusions became much less
toxic, but still showed this increase in mortality with the introduction of earlier
larval stages. The scum exerted a similar effect in these infusions, and with its
removal the percentage survival increased appreciably. This effect was greatest on
first stage larvae eggs. The 14-days infusion exerted no major influence on survival,

BY TOM MANEFIELD. 153

for in all cases it was almost normal. Scum formation in these experiments was only
slight when present, and it was never considered necessary to remove it.

“Having shown that the manure infusions had a definite toxic effect on the larvae
of A. aegypti, experiments were then carried out to compare the results with those
obtained from C. fatigans. Due to the difficulty encountered in setting up exact numbers
of C. fatigans eggs because of the raft form, it was decided to use newly hatched first-
stage larvae. This stage also presented the most critical stage in the development of
A. aegypti in the infusions. The results are shown in Table 3. For the seven-day
infusions the percentage survival of A. aegypti was 3%, compared with 62% for’ C.
fatigans. These increased to 5% and 74% when the scum was removed. For the
eleven-day period the percentages were: A. aegypti 60%, and C. fatigans 92%, with the
“scum left, and increasing to 84% and 96% when the scum was removed. The results
here for A. aegypti also closely correspond with those previously shown in Table 2.

A limiting factor is only evident for the seven-day infusions. As well as preferring
the manure infusion for egg laying, A. aegypti showed that they were quite able to breed
after the infusion was 14 days matured. At seven days the infusions were almost
completely toxic and could prevent breeding. At 10-11 days the larvae were able to
survive in sufficient numbers to set up a large community. ‘With this increasing ability
to survive in the maturing medium the lethal factor is lost. At this time some other
factor must come into existence in the field which deters the setting up of a community.
As well as showing a complete preference for oviposition, C. fatigans showed a greater
ability to survive in the manure infusion. The scum forming on the cultures was shown
to have more effect on A. aegypti than on C. fatigans. This is probably due to the
form of the respiratory siphon. The scum, however, was not the sole deterrent factor
on A. aegypti. It is regretted that a method for measuring the scum satisfactorily was
not devised. In view of the work carried out by Beattie (1932), it is suggested that
the lethal factor present is related to a change in the concentration of some nitrogen
product.

SUMMARY.

1. Aédes aegypti, when allowed a choice between clean tap water and foul water
(manure infusion) for oviposition, showed a distinct preference for foul water, the
percentages of eggs deposited on the two types of water being 17:7% and 82:3%
respectively.

2. Culex fatigans, when allowed a choice, showed complete preference for the foul
water, no eggs whatever being deposited on tap water.

3. In the seven-day manure infusions A. aegypti larvae showed a higher mortality
the younger the stage exposed. The percentage mortality in each stage was: larvae
introduced at the third stage gave 30% mortality; second stage, 62%: first stage,
983%; eges, 96%.

4. A similar result was obtained with A. aegypti in 10-11 day manure infusions;
but, even with early stages, there was sufficient survival to allow successful breeding.
The mortality percentages were: third-stage larvae introduced, 12%; second-stage
larvae, 26%; first-stage larvae, 41%; eggs, 56%.

5. A. aegypti larvae developed almost as well in 14-15 day infusions as in the
controls.

6. Scum formation was shown to have some effect on survival of A. aegypti in the
7-day and 10-11 day infusions. The percentage mortality decreased in all cases with
the removal of the scum. The effect of the scum was most noticeable with the
introduction of eggs and first-stage larvae in 10-11 day infusions, the percentage mor-
tality decreasing to 10% and 17% respectively when the scum was removed. (Cf. par. 4.)

7. C. fatigans was more able to survive in foul water in all experiments than
A. aegypti.’ The actual mortality percentages with first-stage larval introduction were
as follows: i

(a) In seven-day infusions with the scum untouched, A. aegypti showed 97% mor-
tality and C. fatigans 38%.

-154 PREFERENCES BY AEDES AEGYPTI AND CULEX FATIGANS FOR TYPES OF BREEDING WATER.

(bd) With the scum removed these decreased to 95% and 26% respectively.

(c) In 11-day infusions with the scum untouched, A. aegypti showed 40% mortality
and C. fatigans 8%.

(ad) With the scum removed these decreased to 16% and 4% respectively.

ACKNOWLEDGEMENTS.

The author’s thanks are due to Dr. A. R. Woodhill for his advice while carrying
out this work, and to him, Professor P. D. F. Murray, and Dr. W. A. McDougall,
Queensland Department of Agriculture and Stock, for many helpful criticisms during
the preparation of this paper.

References.

Brattip, M. V. F., 1932.—The physico-chémical factors of water in relation to mosquito breeding
in Trinidad. Bull. Ent. Res., 23: 477.

Buxton, P. A., 1934.—Further studies upon chemical factors affecting the breeding of
Anophelines in Trinidad. Bull. Ent. Res., 25: 491.

Dunn, L. H., 1927.—Mosquito breeding in ‘test’ water containers. Bull. Ent. Res., 18:17.

HopkKINsS, G. H. E., 1936.—Mosquitoes of the Ethiopian region. Part I.

LEE, D. J., 1944.—An atlas of the mosquito larvae of the Australasian region.

MacGrecor, M. E., 1929.—The significance of pH in the development of mosquito larvale.
Parasitology, 21:132.

SENIOR-WHITE, R., 1934.—Three years’ mosquito work in Calcutta. Bull. Ent. Res., 25: 551.

THomsSON. R. C. M., 1940.—Studies on the behaviour of Anopheles minimus. Parts I, Il and
Ill. J. Matar, Inst. Ind., 3: 265, 295 and 323:

, 1941.—/d., Part IV. The composition of water and the influence of organic pollution

on the selection of breeding place. Jbid., 4: 63.

WoOoDHILL, A. R., 1936.—Observations and experiments on Aédes concolor Tayl. Bull. Ent. Res.,
27:3 633.

, 1941a.—The oviposition responses of three species of mosquitoes (Aédes aegypti,

Aédes concolor and Culex fatigans) in relation to the salinity of the water. Proc. LINN.
Soc. N.S.W., 66: 287.

A MITE FROM A BEEHIVE ON SINGAPORE ISLAND (ACARINA:LAELAPTIDAB).

By Cart E. M. Guntruer, M.D., B.S., D.T.M. (Sydney), D.T.M.&H. (England),
Field Medical Officer, Bulolo Gold Dredging Limited, Bulolo, Territory of
Papua—New Guinea.

(Five Text-figures. )

[Read 25th July, 1951.]

Synopsis.

Four mites found in separate brood capsules in a hive of Apis indica are described as
Myrmozercon reidi, n. sp. It is thought likely that they belong to an undescribed genus; they
are quite close to Myrmozercon but it is felt that the erection of a new genus on nymphal
characters alone is not justified.

These mites were found in November, 1944, and given to me by Mr. John Reid,
Entomologist from the Institute for Medical Research at Kuala Lumpur (at the time a
member of the Federated Malay States Volunteer Forces, and a prisoner of war in
Changi Camp). While investigating mortality among bees (Apis indica) in one of the
hives kept for the camp hospital, he found four mites sealed in separate brood capsules.
The mites were alive and apparently feeding on the bee pupae; one mite actually had
its proboscis inserted into a pupa. There was no shed mite skin in any of the brood
capsules (i.e., there was no evidence that any stage of metamorphosis had taken place
while the mites had been enclosed in the capsules).

The cause of the trouble among the bees was wax moth (probably family Galleriidae).
Careful search revealed no more mites, in any stage, and it was finally assumed that
the ones found must have attached themselves to foraging bees and so been transported
into the hive, where they had managed to enter the brood capsules, being sealed up
with the pupae. That four were found among less than two hundred capsules opened
seems to indicate that this is a common occurrence, possibly the normal habit of
the mite. i

Three of these mites are smaller than the fourth and have the anal plate separate
from the ventral plate, whereas in the fourth the anal plate is closely attached to,
probably fused with, the ventral plate._ One of the smaller ones was dissected and was
found to have a testis overlying each second coxa. I can find no genital opening on
any of the specimens, and it seems, therefore, that these are all deutonymphs, the
smaller ones male, the larger one female. They are quite close to the genus Myrmozercon,
and while it is likely that they belong to some as yet undescribed genus, I do not feel
justified in erecting a hew one on nymphal characters alone, and so I include them
in Myrmozercon.

MyRMOZERCON REIDI, Nn. Sp.

Deutonymph: Body a transverse ellipse with a slight, smoothly-rounded projection
anteriorly; almost half as wide again as long; flattened, dorsum convex, venter concave.
Male: L. 1050u to 1090u, W. 1500u to 15404; female: L. 11254, W. 1600u. Colour brown.
A single dorsal shield covering the whole body, with a narrow thickened margin,
smooth, bearing setae as follows: general surface with sparsely-set fine setae up to 150
long, about 50u apart, those on the anterior half bearing minute setules, those on the
posterior half plain; from the inner edge of the thickened margin arise fine plain setae
75u to 150u long; at the sides, from two-fifths to four-fifths of the distance back around
the circumference, arising from the middle of the thickened margin, are stout, sharply-
tapering spines averaging 77“ in the male, 854 in the female, from 20 to 25 at each
side; all setae and spines pointing posteriorly or postero-laterally.

156 A MITE FROM A BEEHIVE ON SINGAPORE ISLAND,

Venter: Coxae close together, set in a semicircle well forward under the body,
coxae i at the anterior margin. The proboscis and palps completely retractable; when
retracted they lie between coxae i (Text-fig. 1), but when they are projected coxae i
move close together (Text-fig. 2), almost as if their squeezing action were forcing
and holding the mouthparts forward. An arch-shaped sternal plate overlies the tips
of the coxae; it bears five pairs of strong, stout, tapering spines as follows: one medial
to the tip of coxa i, one opposite the tip of coxa ii, and three overlying the tip of
coxa iii (there are neither metasternal plates nor epigynal shield, so, following Tragardh,
it would seem that this sternal plate represents the fusion of all these, since it bears

Myrmozercon reidi, n. sp.
1, Composite dorsal and ventral view; proboscis retracted between coxae i. 2, Ventral
view of appendages; proboscis projected and coxae i approximated. 3, Chelicerae and flagellum.
1, Hypostome and lingula. 5, Dorsal view of epistome. ;

9

five pairs of setae). Ventral plate pentagonal, occupying most of the posterior central
region, its lateral tips behind coxae iv, its anterior point smoothly rounded, its posterior
tip flattened; covered with sparsely-set short plain setae which extend forwards for
three rows on to the integument. Anal plate small, triangular, placed at the posterior
margin of the body; separated from the ventral plate by a narrow strip of integument
in the male, but in the female closely applied to, apparently fused with, the ventral
plate; anus with an anterior, backward-pointing triangular flap; a stout sharp seta at
each side of the anus and a shorter, more slender median seta at the posterior tip of
the anal plate; two stout setae on the strip of integument at each side of the anal plate.
Stigmata well behind the lateral ends of coxae iv; peritreme large, almost two and a
half times as long as wide, triangular, its apex curving back to alongside the anal

\

Proc. Linn. Soc. N.S.W., 1951. PLATE VI.

Controlled pollination of Bucalyptus.

Proc. Linn. Soc. N.S.W., 1951. PLATE VII.

1, 3. Eucalyptus Robertsonii. 2. BE. pauciflora x EH. Robertsonii.

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Proc. Linn. Soc. N.S.W., 1951. PLATE VIII.

Sed

i) a

4. H. pauciflora. 5. H. pauciflora x EH. Rossii.
6. EH. macrorhyncha x H. Rossii. 7. HE. macrorhyneha.

_
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Proc. Linn. Soc. N.S.W., 1951. PLATE 1X.

oe eee Soe

8. EH. Rossii. 9. EH. macrorhyncha x E. Rossii.
10. HE. macrorhyncha hybrid. 11. #. dives x E. Rossii.

Proc. Linn. Soc. N.S.W., 1951. PLATE X.

12. E. Rossii x E. Robertsonii. 13. Hybrid Peppermint x EH. Rossii.
14. Fl hybrid, #. fastigata x H. Robertsonii. 15. H. elaeophora x E. Blakelyi.

Proc. Linn. Soc. N.S.W., 1951. TACT ee xeTe

me,

mmm |

1 Se
if at Eee
Ye ies ey od

16. BH. Westonii. 17, 18. #. glawcescens B. viminadlis.
19. HB. glawcesceis.

BY CARL E. M. GUNTHER. 157

plate; covered with sparse short plain setae. Metapodal plates roughly triangular, their
apices behind coxae iv, their bases at the lateral margins of the body; with a few short
plain setae over the bases and the adjacent integument.

Coxa i bearing a stout tapering plain seta, 160” long, projecting straight forward
from a shoulder on the anteromedial angle; a shorter, stout seta projecting medially,
and a group of five stout setae on the lateral side. Coxa ii with one stout seta towards
its anterior margin and a group of six posteriorly. Coxa iii with one anterior and five
posterior setae. Coxa iv with four stout setae. Six segments in each leg, but the
last two segments in leg i fused, and partly fused in the other three legs. Legs short,
very stout, normally curved postero-ventrally; i, 5004; ii, 5254; iii and iv, 600 long.
Postero-ventral aspect of last five segments of each leg with a thickened plate. All
segments with several stout sharp setae; no spurs on any leg. Tarsus i longer than
the others, straighter and more slender; tarsi ii, iii and iv short, stout, curved. A large
caruncle, but no claws, on each tarsus (Text-fig. 2).

Epistome only visible from above when proboscis projected; a small, translucent
semicircle (Text-fig. 5). Mandibles with three segments, scissors-shaped, the medial fixed
blade with a row of minute teeth; a small flagellum at the lateral side of the base of
the blades (Text-fig. 3). Hypostome bifid, each half bearing three short stout forward-
pointing setae; lingula bifid, each point bearing three smail rounded projections on its
medial surface (Text-fig. 4). Palpi of five segments, with a few fine setae on segments
iii and iv; terminal segment rounded, with one short sharp stout seta on its disto-medial
aspect, and covered with several fine straight setae.

Type at the British Museum; paratype at the School of Public Health and Tropical
Medicine, University of Sydney.

158

A CRITICAL CONSIDERATION OF c-MITOSIS WITH REFERENCE TO THE
EFFECTS OF NITROPHENOLS.

By Mary M. HINDMARSH,
Linnean Macleay Fellow of the Society in Botany.
(Plate xiii.)
[Read 31st October, 1951.]

Synopsis.

Nitrophenols, as well as many other substances, suppress spindle formation and for this
reason have been described as c-mitotic. Many authors either assume without justification
that substances which suppress spindle formation have a common mode of action, or do not
recognize that, by their use of this term, they imply a common mode of action for all such
substances.

Although nitrophenol and colchicine cause spindle suppression, evidence is presented which
indicates that their modes of action are probably different. It is essential that the term
“c_mitotic’’ be confined to substances which have true colchicine action. If this distinction is
not observed, further confusion must result, which will tend to hamper research on mitotic
poisons. :

In 1938 Levan described the cytological action of colchicine on plant cells. He
showed that colchicine suppressed spindle formation and induced polyploidy without
apparently affecting any other cell process. The name “c-mitosis’” was given to this
phenomenon. Since that time the cytological action of a large number of chemicals
has been investigated and there are reports of numerous substitutes for colchicine.
Among substances reported to have an action like that of colchicine are phenols and
quinones (Levan and Tjio, 1948), veratrine sulphate (Witkus and Berger, 1944), erypto-
pleurine (Barnard, 1949), sulphonamides (Peters, 1946; Fuller, 1947) and various
inorganic salts (Levan, 1945; Galinsky, 1949). Although these compounds are chemically
unrelated, the fact that they are reported to cause a similar cytological effect suggests
the possibility of a common mode of action. It seems reasonable to suppose that
stages in cell division may be linked with metabolic processes and that cytological
aberrations may well result from artificial disturbances in the metabolism of the cell.
There are many substances which are known to interfere with physiological processes
in the cell, and the cytological action of some of these may be of interest.

Recent workers on the phosphorylation mechanisms have made great use ot
2,4-dinitrophenol, which seems to be a specific inhibitor of the phosphate transfer
mechanism in both plant and animal cells. The mononitrophenols do not have this
specific effect (Cross et al., 1949). The mononitrophenols are reported to cause c-mitosis
(Levan and Tjio, 1948a, 1948b). This would suggest that c-mitosis is not the result of
interference with the phosphate transfer mechanism. In view of the importance of
such a conclusion, it seemed desirable to reinvestigate the c-mitotic activity of
nitrophenols.

During the course of this investigation it became apparent that there is a marked
difference between the cytological abnormalities induced by colchicine and phenols. The
cytological action of phenols does not seem to be “c-mitosis’” and this may well apply
to many of the substances which have been described as ‘“‘c-mitotic’. If this is so,
it is essential to distinguish between true colchicine mitosis and other abnormal types
of mitoses before attempting either to look for a common mode of action or even to
evaluate the enormous literature on mitotic poisons.

Recovery experiments are an essential part of this type of investigation, as it is
necessary to know whether the roots are alive at the end of the experiment and also
whether induced cytological abnormalities are permanent or only temporary changes.

BY MARY M. HINDMARSH. 159

In the experiments to be described, onion bulbs with roots about 3 cm. long were
transferred from tap-water to the test solutions for periods of 1, 2, 4, 8, 12, 24 and 48
hours. Solutions of dinitrophenol, o- and p-nitrophenol at concentrations 3, 30 and 300
mg/l were made up in tap-water. After treatment some roots were fixed in acetic-
alcohol (3:1) and examined by standard aceto-orcein and Feulgen squash methods
(Darlington and La Cour, 1947). The remaining roots were transferred back to tap-
water for periods up to 48 hours, and recovery estimated by the ability of the roots
to continue or resume growth. At intervals during recovery roots were fixed for
examination. f

TABLE 1.
aie OF 2, 4-dinitrophenol. p-nitrophenol.
poe ee noi a) Wl a oumeay/ in les00lme/l. 3 mg/l. | 30 mg/l. | 300 mg/l.

1 hour. Metaphase Spindle | Spindle Normal. | Metaphase | Spindle
}chromosomes} Suppressed. |suppressed. chromosomes| Suppressed
short and Stickiness short and
thick. at anaphase. thick in
| Resting few cells.

nuclei |
granular.

2 hours. Metaphase Spindle Spindle Few meta- | Metaphase Spindle
chromosomes|Suppressed. |Suppressed. |phase with |chromosomes]suppressed.
short and Stickiness Stickiness short, thick |short and Stickiness
thick. | at anaphase.| at anaphase | chromo- thick. at anaphase.
Spindle | and somes.
suppressed | telophase.
ina few Resting
cells. nuclei

granular.

4 hours. Metaphase As 2 hours.|As 2 hours.|As 2 hours. | Metaphase Spindle
chromosomes| chromosomes]suppressed.
short and short and Stickiness
thick. thick. at anaphase
Spindle Spindle and telo-
suppressed suppressed phase. Rest-
in more |/in some ing nuclei
cells. cells. slightly

granular.

24 hours. | Spindle As 2 hours,|As 2 hours, | Metaphase Spindle As 4 hours,
suppressed. |but resting ;but all cells |chromosomes|suppressed. | but resting
Anaphase nuclei becoming short and Resting nuclei more
and telo- affected. diffuse. thick. nuclei granular.
phase Spindle slightly
rarely suppressed affected.
found. in few cells.

In water All Recovery No All All No

after recovered. after 2, 4, recovery. recovered. recovered. recovery
treatment. 8 hours. after 24
Not after ‘ hours.
24 hours. |

Cytological abnormalities induced by nitrophenols varied with the concentration,
length of time of treatment, and division stages of the cells at the beginning of
treatment. Cytological observations mentioned below are those obtained using dinitro-
phenol, but the effects with p- and o-nitrophenol were the same except that 2,4-dinitro-
phenol is effective at lower concentrations. This concentration difference is shown in
Table 1.

At the lowest concentration abnormalities were observed after two hours. Metaphase
and anaphase chromosomes were short and thick, but otherwise the mitotic figures
were normal. After four hours most of the cells in metaphase developed a normal
metaphase plate with short thick chromosomes, but a few cells had chromosomes
scattered at random in the cytoplasm. Twenty-four hours’ treatment allowed most cells

160 C-MITOSIS WITH REFERENCE TO THE EFFECTS OF NITROPHENOLS.

dividing during treatment to proceed to normal resting cells; some cells remained in
“blocked” metaphase, but unlike colchicine-treated cells, there was no evidence of
centromere division. These observations indicate that in the presence of nitrophenols
at low concentration spindle formation is completely suppressed, at least in some cells.

At higher concentrations the effects of dinitrophenol were noticeable in one hour.
In some cells a metaphase plate appeared in a normal manner; in others short thick
chromosomes were scattered in the cytoplasm or clumped together in the centre of the
cell. Anaphase stages were numerous; some were normal, some had lagging chromosomes,
and some had short thick chromosomes. This variability in the behaviour of different
nuclei could be due to irregular penetration across the root in a short period of time.
More than two hours’ treatment at 30 mg/l induced stickiness in anaphase chromosomes,
with sticky bridges in anaphase and telophase stages.

Changes in the resting nuclei were observed after 48-60 hours at 3 mg/l, after 24
hours at 30 mg/l and after 2 hours at 300 mg/l. Some resting nuclei lost their fine-
grained appearance, and the chromatin coagulated into coarsely grained masses, forming
nuclei about the same size as healthy resting nuclei. Others appeared as small deeply
stained “pycnotic’”’ masses with nucleoli standing out as unstained spots. With longer
treatment at the high concentrations resting nuclei and chromosomes became diffuse
and unstainable, and even cell walls were indistinct.

Length of treatment at any one concentration influenced the ability of the roots
to recover. When roots which had been in 3 mg/1 for 1—24 hours were transferred to
tap-water, all roots recovered, but after 24 hours in 30 mg/l] or 1 hour in 300 mg/l] no
recovery was observed. These roots became soft and flaccid at the tip.

The cells of roots which recovered in tav-water following treatment all commenced
normal: division after varying periods of time. Again there is a relationship between
length of treatment and concentration. Resumption of normal division is-rapid after
treatment at low concentrations, but much slower after high concentrations. When
transferred to water, all dividing cells pass into the resting state and there is a delay
depending on length of treatment at each concentration before division begins. As all
the cells which resume division are normal and diploid, it is possible that only cells
which did not divide during treatment are seen in mitotic stages after recovery.

If resting nuclei were affected, the roots did not recover when the nitrophenol was
washed out. Roots which had been in 300 mg/l even for one hour did not have normal
dividing cells after being in water for periods up to 48 hours. Abnormal division
stages exactly like those of treated roots were observed in these flaccid roots. These
results of recovery experiments, particularly those following treatment with 300 mg/I,
showed that nitrophenols possess a strong toxie action. At concentrations of 30 and
300 mg/l, nitrophenols appeared to behave as poor fixatives gradually killing and fixing
the cells in the stage in which they were when the substance penetrated the cells.

The cytological abnormalities induced by mono-nitrophenols on meristematic cells
has been described by Levan and Tjio (1948) as ‘‘e-mitosis’” over a limited concentration
range. This term “c-mitosis’” was originally used by Levan (1988) as follows. “The
effect of colchicine is entirely specific and the modification in mitotic behaviour will be
abbreviated ‘c-mitosis’. The c-mitosis can be referred to one single moment, viz. an
inactivation of the spindle apparatus connected with a delay of the division of the
centromere. The effect thus produced may be expressed as a completion of the
chromosome mitosis without nuclear or cellular mitosis.”

Levan then discusses the “course of c-mitosis”’, referring the term not to “one
single moment”, but to the whole abnormal cell division process which results in
telophase nuclei with twice the original number of chromosomes, and numerous
polyploid cells in the meristem. This explanation of the term “c-mitosis” is unfor-
tunately ambiguous. The first part of the explanation refers to the specific action of
colchicine on the spindle mechanism, whereas the second part, and the subsequent
discussion, refers to the whole cell division process. Since 1938 both the spindle

BY MARY M. HINDMARSH. 161

inhibition and the morphological picture of the colchicine mitosis have been described
as “c-mitosis” by Levan and other workers. It seems necessary to make a distinction
between these two aspects, since the work with nitrophenols indicates that spindle
suppression is not necessarily the same as colchicine action.

It seems generally accepted that colchicine has the single effect of upsetting the
spindle mechanism, thus altering metaphase and consequently anaphase separation and
cell wall formation. Other cell division processes, including the stages from resting
nucleus to metaphase and also centromere division, are allowed to proceed during
colchicine treatment and the result is polyploidy.

As shown earlier (pages 159-160), nitrophenols are not “entirely specific’? in their
action. They cause inactivation of the spindle mechanism, but they also alter other
vital and independent cell processes so that centromere division, that is “completion of
the chromosome mitosis”, rarely takes place in treated cells. A very small percentage
of cells became tetraploid during treatment, probably only those beginning metaphase
or anaphase at the time of treatment, but no tetraploid cells were seen in recovered
roots. Therefore, polyploidy, the permanent consequence of “c-mitosis”’, was not observed
as a result of nitrophenol treatment.

As nitrophenol-treated roots continue to grow by normal cell division when they
recover, it is not possible to determine whether the cells which show abnormalities
during treatment recommence mitosis on recovery. The normal diploid cells may come
from the large number of cells which remain in resting stage during treatment
and these may completely outgrow abnormal cells. Roots did not recover when the
resting nuclei were visibly affected, and this may indicate either that these are the
cells which continue growth or that granulation of the resting nuclei occurs only when
all the cells are killed.

Attempts to explain the action of colchicine on cell division, or to correlate the
physical and chemical properties of colchicine with those of other spindle suppressors
so far have been unsuccessful. The explanation of this would seem to be that colchicine
has a specific action on the spindle not possessed by other spindle suppressors. It has
been suggested (Levan and Ostergren, 1943; Ostergren and Levan, 1943; Ostergren,
1944) that there is a negative correlation between water solubility and the ability of
numerous substances to suppress the spindle. Colchicine, however, has a very high
water solubility and a strong activity, and does not fit this general theory. This
exception is too important to be disregarded and it must be assumed in contrast to the
assumption of Ostergren (1944) that colchicine activity is not the same as the spindle
suppression of other substances.

During division it is the spindle which is most sensitive to disturbance and there
are many observations to show that it is easily upset by changes in the environment
of the cell, e.g. temperature changes (Barber and Callan, 1943). Spindle formation in
plant and animal cells almost certainly depends on a number of reactions in the cell
and upset of one or more of these reactions may produce the same results at metaphase.
If the term “c-mitotic” is to be used for any spindle inhibiting substance, it can be
applied to a wide variety of substances which probably act on different parts of the
spindle mechanism. On the other hand, if it is to be used for true. colchicine-like
action, it applies only to a particular action on one part of the cell division process.
It would be better to retain the term “c-mitosis” for substances which are known to
have the same specific action on the spindle mechanism as has colchicine and no
other activity. This view agrees with that of Frahm-Leliveld (1949), who, after testing
many substances, concluded that only colchicine and acenaphthene act as polyploidogenic
agents on plant cells and are the only known “c-mitotic” substances. Y

Levan (1938) has shown that 2% colchicine for 72 hours produced cytological
abnormalities but was not toxic to the cells. With high concentrations the toxic action
of the nitrophenols is clearly demonstrated by recovery experiments. Treated cells
which did not recover, retained their abnormal cytological figures until all cell
contents became diffuse and unstainable, indicating that vital cell processes were stopped.

162 C-MITOSIS WITH REFERENCE TO THE EFFECTS OF NITROPHENOLS,

Concentrations of mononitrophenols reported by Levan and Tjio (1948) to induce
‘e-mitosis” were 5 x 10° to 2 x 10°M fon o-nitrophenol, 1 x 10° to 1 x 10?°M for
m-nitrophenol and 2 x 10° to 2 x 10*M for p-nitrophenol. All solutions were diluted
with distilled water. These concentrations are higher than those used in our experi-
ments where 2:16 « 10°M, the highest concentration, killed in less than 24 hours. The
toxic action of phenols is very much greater in Gistilled water than tap-water, probably
due to a different pH. As no recovery experiments were cited by Levan and Tjio, it
is likely that roots in which c-mitosis has been described were dead at the time of
sampling.

Results of experiments with low concentrations, where longer treatment was required
before the cells were killed, showed that spindle formation was ultimately suppressed.
The number of anaphase and telophase stages was reduced, indicating that the cells
go to apparently normal daughter cells, if the spindle is already initiated at the time
ot treatment. After 24 hours’ treatment there were no anaphase or telophase stages,
but some suppressed metaphases in which centromeres had not divided. The onset of
prophase was also affected as the number of cells dividing during treatment was
reduced. This suggests that there is a general slowing down of the metabolism of the
cell, which may be evidence for a toxic action even at the lowest concentration.

“Stickiness” of anaphase and telophase chromosomes occurred almost at once with
the high concentration, but more slowly at the low concentrations. It is induced by
many phenols (Levan and Tjio, 1948). Like spindle suppression, stickiness, particularly
of anaphase chromosomes, results from treatment with a wide variety of organic
substances and also by cold treatment (D’Amato, 1948). There is no evidence to suggest
that these treatments all have the same specific action on the chromosomes.

It seems possible that the cytological abnormalities induced by nitrophenols reflect
the general toxic action of these substances on the cells. Spindle suppression, stickiness
of chromosomes, and granulation and pycnosis of the resting nuclei could be a direct
result of the slow death of the cell. Phenols are known to precipitate proteins and the
results for nitrophenols can be explained as a slow fixation process, which first affects
spindle formation and then the onset of prophase, gradually slows the whole cell division
process and kills the resting nuclei:

On the other hand, the nitrophenols at low concentrations may have a specific
action on the spindle mechanism not at the same point as colchicine, as well as an
unrelated toxic action which kills the cells before the abnormal mitosis is complete.
As no abnormalities have been seen in recovered roots, it is not known whether
inhibited metaphase cells found in roots after 24 hours at low concentrations undergo
centromere division before reaching the resting condition. Further work is necessary
at low concentrations before this possibility can be excluded.

It seems essential to make a distinction between the specific action of a chemical
on one phase of the cell division mechanism and a general action which is apparent
only on cells which are dividing. Colchicine has a specific action on one stage of the
cell division process, whereas nitrophenols probably induce a general toxic effect by
some metabolic derangement which is visible at first only in dividing cells. Some
other substances described as ec-mitotic may well have this general action and the
literature should be re-examined from this point of view. The action later extends to
resting nuclei which are killed. It is difficult to make the distinction with this type
of experiment, as the course of an abnormal mitosis cannot be followed through a
complete division. It is necessary to reconstruct the process from a knowledge of the
normal mitosis and observations on a series of fixed roots.

Acknowledgements.

The writer wishes to thank Professor N. A. Burges, in whose department this work
was done, Mr. S. Smith-White, Dr. F. V. Mercer and Professor H. N. Barber for helpful
criticism of the manuscript, and the Department of Medical Illustration, University of
Sydney, for the plates,

BY MARY M. HINDMARSH. 163

References.
BarRBER, H. N., and CALLAN, H. G., 1943.—The effects of cold and colchicine on mitosis in the
newt. Proc. Roy. Soc., B, 131: 258-270.
BARNARD, C., 1949.—The c-mitotic activity of cryptopleurine. Aus. J. Sci., 12: 30-31.
BuLLouGH, W. S., 1950.—Completion of mitosis after death. Nature, 165: 493.
Cross, R. J., TAGGART, J. V., Covo, C. A., and GREEN, D. E., 1949.—Studies on the cyclophorase
system VI. The coupling of oxidation and phosphorylation. J. Biol. Chem., 177: 655-678.
D’AmatTo, F., 1948.—The Effect of colchicine and ethylene glycol on sticky chromosomes in
Allium cepa. Hereditas, 34: 83-103.
DARLINGTON, C. D., and La Cour, L. F., 1947.—The Handling of the Chromosomes. Allen,
Unwin, London.
DERMAN, H., 1940.—Colchicine, polyploidy and technique. Biol. Rev., 6: 599-635.
FULLER, T. C., 1947.—The effects of several sulpha compounds on nuclear and cell division.
Bot. Gaz., 109: 177-183.
FRAHM-LELIVELD, J. A., 1949.—Experiments with polyploidogenic and other agents in different
types of plants under tropical conditions. Ann. Roy. Bot. Gard. Buitenzorg, 51: 231-267.
GALINSKY, I., 1949.—The effect of certain phosphates on mitosis in Allium roots. Journ. Hered.,
40°: 289-295.
LEVAN, A., 1938.—-The effect of colchicine on root mitosis in Allium. Hereditas, 24: 471-486.
, 1940.—The effect of acenaphthene and colchicine on mitosis of Alliwm and Colchicum.
Hereditas, 26: 262-272.
, 1945.—Cytological reactions induced by inorganic salt solutions. Nature, 156: 750-751.
, and OSTERGREN, G., 1943.—The mechanism of c-mitotic action. Hereditas, 29: 381.
, and Ts1o, J. H., 1948a@.—Chromosome fragmentation induced by phenols. Hereditas,
34: 250.
, and ——, 1948b.—Induction of chromosome fragmentation by phenols. Hereditas,
34: 453-483.
LEVAN, A., 1949.—The influence on chromosomes and mitosis of chemicals, as studied by the
Allium test. Proc. Highth Int. Congress. Genet. (Hereditas, Suppl. Vol., 1949): 325-337.
LovELEsSS, A., and REVELL, S., 1949.—New evidence on the mode of action of ‘mitotic poisons’.
Nature, 164: 938-944.

OSTERGREN, G., and LEvAN, A., 1943.—The connection between c-mitotic activity and water
solubility in some monocyclic compounds. Hereditas, 29: 496-498.

OSTERGREN, G., 1944.—Colchicine mitosis, chromosome contraction, narcosis, and protein chain
folding. Hereditas, 30: 429-467.

PETERES, J., 1946.—Cytological effects of sulphanilamide on Alliwm cepa. Bot. Gaz., 107: 390-392.

RYLAND, A. G., 1948.—A cytological study of the effects of colchicine, indole-3-acetic acid,
potassium cyanide, and 2,4-D, on plant cells. Journ. Elisha Mitchell Sci. Soc., 64: 117-125.

WITKUS, HE. R., and BERGER, C. A., 1944.—Veratrine a new polyploidy inducing agent. Hereditas, ~
385: 1381-133. 4

EXPLANATION OF PLATE XIII.

Effect of 2,4-dinitrophenol on Onion root tip cells. (All photographs x ca 1000.)
1-3. Metaphase after treatment with 3, 30 and 300 mg/I1 respectively.
4-5. Anaphase showing sticky bridges after 30 and 300 mg/1 respectively.

6-9. Abnormal metaphase and anaphase stages after attempted recovery in water for
24 hours following treatment with 30 and 300 mg/l.

164

THE LIFE-HISTORY OF A PENAEID PRAWN (METAPENAEUS) BREEDING IN A
COASTAL LAKE (TUGGERAH, NEW SOUTH WALKS).

By Muriet C. Morris, M.Sc.,* and IsopeL BENNETT, Department of Zoology,
University of Sydney.

(Plate xii and 96 Text-figures. )
[Read 26th September, 1951.]

Synopsis.

This paper deals with the life-history of a species of Penaeid prawn (Metapenaeus, n. sp.,
Crustacea, Decapoda) and is based on a study of both living and preserved specimens. The
species is unique in that the whole life-cycle from egg to breeding adult is spent in a shallow
coastal lake (Tuggerah, New South Wales) ; in all other species whose life-histories are known,
the maturing adults migrate from estuarine waters to the sea, the iarvae returning to the
estuaries.

Stages from egg, through nauplius, protozoea, mysis, post-mysis and post-larval to adult
were obtained and the larval stages are described and figured hereunder. The life-history
follows the same general lines as that described by Heldt (1938) for a Mediterranean Penaeid

species.
INTRODUCTION.

This study was part of the long-range programme of plankton and general marine
investigations planned by the late Professor W. J. Dakin during his term of office as
Professor of Zoology at Sydney University. This particular study was begun by him in
1945 and the field work was carried out by one of us under his direction during the years
1945-47. Two short notes indicating the general findings have already been published
(Dakin, 1946, 1946).

Professor Dakin had completed a large number of drawings and had made rough
sketches and notes of all the stages in the early life-history before illness prevented his
finishing the work. It was felt that this effort should not be lost, and, as one of us
(1.B.) had been responsible for all the collecting, sorting and preserving of the stages,
it was a relatively easy matter to complete the work.

HISTORICAL.

The general story of the Penaeids, which are the Australian prawns of commerce,
together with full details of the species concerned in the fishery, has already been told
(Dakin, 1938, 1940, 1946).+

When prawn investigations were first begun in this Department, all the Penaeid
species which could be obtained along the New South Wales coast were collected and
sent, in 1937, to Mr. Martin D. Burkenroad, of the Bingham Oceanographic Foundation
at Yale, who at that time was engaged in a systematic revision of Indo-Pacific littoral
Penaeids. Burkenroad verified our identifications of Penweus esculentus (‘Tiger’ prawn),

* Formerly Linnean Macleay Fellow in Zoology.

7 In the year 1947 a chance discovery by fishermen led to commercial fishing for prawns
on a fairly large scale in open waters off the New South Wales coast. For the first time
‘School’ prawns, Metapenaeus macleayi (Haswell), with mature gonads, were found. This
seemed to prove Professor Dakin’s theory that this species, like Penaeus plebejus, also migrated
from the estuaries to the open ocean to breed.

A brief survey of the American Penaeid fishery (which is of very much greater economic
importance than the Australian fishery), together with details of the breeding migrations, is of
interest in that it indicates a very similar story (Idyll, 1950).

Just as this paper was going to press we were shown several specimens of females of
M. macleayi with spermatophores attached. These were taken in oceanic waters off the
northern New South Wales coast in the month of February, 1951, by Mr. J. H. Wharton,
Biologist, State Fisheries Department.

BY MURIEL C. MORRIS AND ISOBEL BENNETT. 165

P. plebejus (‘King’ prawn) and Metapenaeus macleayi (‘School’ prawn), but in a letter
dated 30/7/38, stated:

“Your two ‘Metapenaeus monoceros’ from Lake Illawarra may be of the ‘Dana’
species, or possibly of M. incisipes Bate, or of my (MS.) M. tweedii, or of some
other form. They are certainly not M. monoceros—since of the eight or ten species
of the group (unpublished studies) this last is the one which does not range east of
India. Determination of members of this group is difficult; it depends chiefly on
details of the genitalia.

“When you deal with hairy prawns with numerous but minute lateral telson
armature, you are on difficult ground. I am unable to say how many Metapenaeus,
which usually pass under the name of M. monoceros, may occur in the region of
Sydney, but I now have records of two species. One of these, the ‘Dana’ sp. n.,
must at some season be frequent in your commercial catch, since the sample, from
Sydney market, consists entirely of it. As the specimens are large and adult, they
may well have been taken at sea by the trawlers.” .

And later, on 18th January, 1940, Burkenroad wrote: “Your ‘Greasy Backs’ are the
same as the ‘Dana’ Metapenaeus sp. n.” :

This prawn species which, because of the very thick covering of minute hairs on
the carapace, was known locally to fishermen as the “Greasy Back’, had always been
regarded by Museum authorities and other workers in Australia as Metapenaeus
monoceros Fabr.

We have been unable to contact Burkenroad since the end of World War II, and
there is no mention of the genus Metapenaeus in the only published work on the ‘Dana’
collections we have seen (Burkenroad, 1940). In order to avoid possible duplication,
we have decided to refer to the species simply as quoted above from Burkenroad’s
letter, “Dana” Metapenaeus sp. n.

After Professor Dakin had first obtained our so-called Metapenaeus monoceros from
Lake Illawarra (see Dakin, 1938, p. 166), efforts were made over the years to procure
further specimens of this species. We were successful in obtaining only a few specimens
in the summer months of 1938-39 from Cook’s River, Sydney. These were all sent to
Burkenroad, who stated they were also the “same as my ‘Dana’ Metapenaeus sp. n.’.
On the outbreak of war in 1939, this work was discontinued, but in February, 1945, we
discovered that these same prawns were forming the bulk of the commercial catch sent
to the markets, and inquiry indicated that they were being caught in Tuggerah Lakes
(three closely-connected lakes, Tuggerah, Budgewoi and Munmorah), 70 miles north of
Sydney. Cooked prawns in which the gonads were fully mature were obtained from
the Sydney Fish Markets. The prawns_themselves were comparatively small, only 33
to 4 inches in length.

It is characteristic of the shallow coastal lakes along the New South Wales coast
to become silted up at the entrance, and, especially in dry periods, to remain closed
sometimes for as long as two years. It had been hoped to make a series of observations
on the various Penaeid species under these conditions, particularly in view of the
known habit of the group, in other parts of the world, of migrating to open ocean waters
to breed.

At this same time (early in 1945) it was learned that Tuggerah Lakes had been
closed for a period of approximately two years, and that the ‘King’ and ‘School’ prawns,
by the end of the second year (the summer season 1944-45) were scarce but very large.
None, however, was observed with any gonads. But the ‘Greasy Backs’ which, in that
particular season, were very numerous, forming the bulk of the catch, all had mature
gonads. This information seemed to fit in very well with our theories regarding the
three species concerned. By the time all these details were known, it was too late in
the season to carry out any field work to obtain larval stages, but plans were made for
an intensive effort for the next summer season (the months of November, December,
1945, and January—March, 1946).

Unfortunately heavy rains caused flooding in the lake backwaters during the
autumn, and in May, 1945, the entrance to Tuggerah Lakes was cut through and the

R

166 LIFE-HISTORY OF A PENAEID PRAWN BREEDING IN A COASTAL LAKE,

lakes have not been completely land-locked since that date. This entirely altered the
physical conditions within the lakes, but the work was carried out as planned.

Fishermen who tested the prawning grounds in the lakes stated that small prawns.
first began to appear in October, but these were well under regulation size (3% inches).
By November, 1945, however, intensive fishing was being carried on with three species
of prawns appearing in the catches—P. plebejus, Metapenaeus macleayi, and the so-
called “Greasy Back’. The two former were taken together, mainly in the largest lake,
Tuggerah itself, and always at night. The “Greasy Backs” were generally caught alone
in both Tuggerah and Budgewoi Lakes, and had a habit of shoaling during the day.

Many bulk catches were examined from different parts of the lakes and though a
very careful search was made throughout the whole period of the investigation (1945-47)
we were never successful in finding either P. plebejus or M. macleayi in the lakes with
any signs of developing gonads.

From November to March, however, the “Greasy Back” females were very
conspicuous in all catches because the gonads showed up as a dark green band along
the dorsal surface of the abdomen (Plate xii, fig. 2). Random samples of many
hundred specimens were taken, and the individuals sexed and measured. It was found
that the females were larger than the males—the largest female actually measured by
us being 5 inches, though the greatest number averaged between 34 and 4 inches. We
found only one male as long as 4 inches, the greater proportion being about 3 inches.
The females in all samples counted formed more than 50% of the total catch, but it
was felt that this was probably due to the size of the mesh of net used by the fishermen.

Close examination of specimens led us to suspect that, in common with many other:
marine animals, spawning took place at time of full moon. Our catches later substan-
tiated this. :

COLLECTION OF MATERIAL.

Since, to our knowledge, there had been no previous plankton collections in
Tuggerah Lakes, a great number of exploratory hauls had to be made, and these were:
carried out on all three lakes. Nets of varying degrees of coarseness were used, from
the finest mesh bolting silk to nets of coarse stramine. Since overseas work had
indicated (Pearson, 1939) that the eggs were probably demersal, various methods,
such as the attaching of heavy iron sledges to the nets, had to be devised for taking-
hauls just above the bottom layer of mud. This was extremely difficult since the floor
of the lakes, which was only about 8 feet deep in the deeper parts, was very uneven.
On more than one occasion in heavy southerly weather, the hauls had to be abandoned
and a fresh start made after the net had become clogged with mud from the bottom.

Another difficulty, accentuated by the opening of the lakes to the sea, was the
seasonal influx of Coelenterates. Finally two types of cones which had the same
diameter as the net and which fitted over the mouth of the net, were used. One was
made of flat galvanized sheeting with small holes punched all over it, and the other
of small gauge wire netting. The guy ropes of the net were threaded through a hole
in the peak of the cone, the base of which was firmly laced to the ring of the net. The
current set up by these cones as the nets were pulled through the water served, to a
certain extent, to force the jelly fish out of the course of the net, and the holes and wire
mesh permitted the net to catch smaller organisms. The method was quite successful
when dealing with the large medusa, Catostylus mosaicus, but when the Ctenophores.
came into the lakes, all our efforts were practically useless. The surface waters were
almost solid Ctenophora, which were so soft that the cones merely broke them and the
nets became coated with a thick gelatinous slime.

A large number of hauls using the different nets were taken at close intervals
throughout the summer seasons of 1945-46 and 1946-47. Throughout the winter months.
of 1946 regular monthly hauls were made so that some idea would be gained of the:
general plankton constituents throughout the whole year.

Since we are able to state with certainty that no other Penaeid prawn taken in
the lakes during this time had even immature gonads, and since the bulk of our
catches were taken in Budgewoi Lake, at least eight miles from the entrance, where-

BY MURIEL C. MORRIS AND ISOBEL BENNETT. 167

there is no tidal flow, we have no hesitation in regarding the eggs and nauplii as those
of the breeding Metapenaeus.

We were successful in obtaining eggs during February and March only, but
Protozoea and Mysis stages were found in all months from November to March inclusive.
We cannot be certain of the full number of nauplii but the most important stages in
development were verified with living material. No aquarium facilities of any kind
were available to us, but by means of a temporary laboratory set up in a tent on the
lake shore we were able to watch the egg developing through 2, 4, 8, and 16 cell stages,
and finally photographed several in which the nauplius could be seen actively moving
round within the egg membrane. The newly hatched nauplius was also photographed
(Plate xii, fig. 3). What we considered to be the last nauplius (Metanauplius) stage
moulted into Protozoea I. We also observed the moulting of Protozoea I into Protozoea
II, Protozoea II into Protozoea III, and Protozoea III into Mysis I.* It was only after
very considerable difficulty and a week’s concentrated effort that Mysis III stages were
obtained. It is interesting to note here that we had chosen a particular week in
February to make these catches, ten to eleven days after the full moon. In that week
practically no stages other than late Mysis were found in any of the catches. This
substantiated the assumption that spawning takes place at the time of full moon, as
the time taken for an egg to develop to a late Mysis stage is about ten days. No
Mysis were found in the surface hauls and it was only when the nets were weighted
and hauled just above the bottom that we were successful in finding them. Heldt
(1938) found that this was also the case with her Mediterranean Penaeid species: “Dans
les péches de plancton la quatriéme mysis et la premiére post-mysis sont trés rares.”

- We were able to keep most of the late Mysis alive until the first post-Mysis stage,
but without aquarium facilities were unable to go further. From the above, however,
we were able to place with certainty all our planktonic material as belonging to the one
species. A very long series of post-larval stages from 2 inch in length and leading
to the young adult, finally left us in no doubt about the identity of the species of which
they were the larval stages.

LARVAL AND PoST-LARVAL STAGES.

A series of 16 larval stages was taken in the plankton hauls. They consisted of
four Nauplius stages, three Protozoea stages, three Mysis stages and six post-Mysis.
stages. In addition several post-larval stages were taken, four of which have been
described.

The Nauplius stages taken did not constitute a complete series, and it would seem
that the full series, as obtained by Heldt,_numbers eight—the possible missing stages:
being two, three, four and six. The three Protozoea, of course, constitute the full
number of Protozoea stages, while three distinct Mysis stages have been obtained.
Heldt (1938) records four Mysis stages for Penaeus trisulcatus, while the American
workers record only two for their species.

There is quite a big gap between the first post-Mysis, which was obtained by direct
moulting, and the next stage taken in the plankton hauls (5-073 mm.). From the
5-073 mm. stage onwards a series of ten stages (to 23-572 mm. stage) were taken.
The first five of these are probably post-Mysis stages, while the other five are post-
larval. The distinction between post-Mysis and post-larval stages has been made on
the purely morphological basis that the first stage bearing biramous pleopods would be
the first post-larval stage. The important difference between post-Mysis and post-larval
stages is that propulsion is thoracic in post-Mysis and abdominal in post-larval. The
loss of feathered exopodites from the peraeopods (first post-Mysis stage) would mean a
loss of effective thoracic propulsion. However, the fact that the pleopods are still
ill-developed at this stage means that abdominal propulsion would be equally ineffective.
In fact, abdominal propulsion would not be effective until the appearance of biramous
pleopods (7-835 mm. stage), and for this reason it is felt that this stage should be

+ We have followed Heldt (1938) rather than Gurney (1927 and 1942) in the terminology
used for these stages.

168 LIFE-HISTORY OF A PENAEID PRAWN BREEDING IN A COASTAL LAKE,

considered as the first post-larval stage. It is realized, of course, that ever since the
appearance of a well-developed tail-fan at the moult to first Mysis, flexing of the abdomen
has no doubt constituted quite an effective means of propulsion. Madame Heldt (1938)
referred to the role of abdominal flexion in propulsion, but without the observation of
live specimens, which was not possible in this study, it would not be possible to say
just how effective a means of propulsion this would be in Metapenaeus. Heldt (1938)
made this division into post-Mysis and post-larval at an earlier stage—she named the
first post-larval as the stage preceding the one where biramous pleopods appeared.

NAUPLIUS STAGES.

First Nauplius (Text-fig. 1)—Several very early stages in the development of the
nauplius, all still enclosed in the egg membrane (Plate xii, fig. 3), were taken. They
later emerged as the first Nauplius larva.

The first Nauplius measures 0-23 mm. from the front end to the end of the body
(not including the spines). The body is somewhat pyriform in shape when viewed from
the dorsal or ventral aspect, but when viewed from a lateral aspect it has a ventral
flexure. At this stage the body shows no signs of segmentation.

A pair (1 +1) of spines extend out from the posterior margin of the body.

A Nauplius eye lies on the median ventral surface of the body close to the anterior
border. This eye persists through to the Protozoea stages and probably into the Mysis
stages. 2

The three pairs of appendages present at this stage, first and second antennae and
mandibles, are large unsegmented club-shaped structures with a purely natatory function.

The first antenna (Text-fig. 1) is a uniramous appendage bearing three terminal
setae and also three on the inner side and one on the outer side.

The second antenna (Text-fig. 1) is a biramous appendage bearing three terminal
setae on the endopodite and one short spine and two terminal setae and three lateral
setae on the exopodite.

The mandible (Text-fig. 1) bears no cutting surface at this stage, but consists of a
biramous appendage. The exopodite bears three terminal setae, whilst the slightly
longer endopodite bears two terminal setae and one on the inner side.

Fifth Nauplius (Text-fig. 2).—The next Nauplius stage which was taken in the
catches corresponds to Heldt’s fifth Nauplius for Penaeus trisulcatus. The only
differences between Heldt’s fourth and fifth nauplius stages are an increase in size;
the presence of seven segments in the exopodite of the second antenna of the Nauplius
four and eight in Nauplius five; and the growth of the terminal spine on the endopodite
of the second antenna of the fourth Nauplius into a long hair in the fifth Nauplius.

As the exopodite of the second antenna of this stage of Metapenaeus consists of
seven and not eight segments, it would seem that it is a fourth Nauplius stage. How-
ever, the presence of three well-developed hairs and a spine on the endopodite of the
second antenna identify it as a fifth Nauplius. Actually, Heldt gives the presence of
three plumose hairs and a spine on the endopodite of the second antenna as one of the
characteristics of the sixth Nauplius in Penaeus trisulcatus. However, the fact that
the telson bears four pairs (4 + 4) and not six pairs (6 + 6) of spines at this stage of
Metapenaeus, definitely classifies it as a fifth Nauplius.

The length of the body is now 0-27 mm.—0-29 mm., and the following significant
changes have taken place since the first Nauplius stage.

The posterior border of the body is now quite deeply indented to form a definite
fureca bearing four pairs of spines. The first antenna (Text-fig. 3) is now segmented and
the beginnings of a large number of segments are obvious at the base. It bears two
long plumose hairs and a short spine at the end; two small hairs well towards the
distal end on the outer side; two longer hairs on the inner margin—one in the middle
of the distal segment, and one further down.

The exopodite of the second antenna (Text-fig. 4) now consists of seven segments—
the division between the last two segments still being ill-defined. The most distal, or
seventh, segment bears two long plumose hairs and a small spine at its extremity, while

BY MURIEL C. MORRIS AND ISOBEL BENNETT. 169
the 3rd, 4th, 5th and 6th segments each bear a large plumose hair on their inner distal
margins. A small spine is present on the inner distal margin of the second segment.
The endopodite, which is still unsegmented, bears three long hairs and a spine at its

Text-figures 1-15.

1, First Nauplius, x 92.
2-5, Fifth Nauplius.
x 164.

6-14, Seventh Nauplius. 6 x 34; 7, First antenna, x 170; 8, Second antenna, x 171;

9, Mandible, x 284; 10, First maxilla; 11, Second maxilla; 12, First maxillipede; 13, Second
maxillipede; 14, Furca, x 462.

15, Eighth Nauplius, Fureca.

2, x 120; 3, First antenna, x 131; 4, Second antenna, x 66; 5, Mandible,

170 LIFE-HISTORY OF A PENAEID PRAWN BREEDING IN A COASTAL LAKE,

distal end; a shorter hair two-thirds of the way down and one one-third of the way
down on the inner margin.

There is now a very definite swelling at the base of the endopodite of the mandible
(Text-fig. 5) which will eventually give rise to the cutting surface of the future mandible.
The outlines of teeth can already be seen through the chitin on the inner side of this
swelling. The exopodite and endopodite still bear the same number of hairs as at the
first naupliar stage.

The maxillae and first and second maxillipedes are now visible under the cuticle on
the ventral side.

Seventh Nauplius (Text-fig. 6).—The next stage in the series which was taken
corresponds to Heldt’s stage seven for Penaeus trisulcatus. The presence of six pairs
(6 + 6) of spines on the furca marks it as either a sixth or seventh Nauplius stage, but
the facts that the posterior border of the carapace is beginning to appear, and that the
maxillae and first and second maxillipedes are now free, identify it as a typical stage
seven.

The body now measures 0-671 mm.

The number of segments in the first antenna (Text-fig. 7) has increased, an additional
small spine has developed on the distal outer corner of the most distal segment, and an
additional two have appeared on the inner side of the appendage.

Signs of further segmentation of the exopodite of the second antenna (Text-fig. 8)
are evident; a third hair has appeared, while the small spine, which was present at the
fifth Nauplius stage, has now developed into a small hair. The most distal segment of
the exopodite thus bears the full complement of four hairs—or what will be four hairs—.
found in the Protozoea stages. The endopodite remains unchanged.

The swelling on the base of the mandible (Text-fig. 9) has increased in size, and
the cutting surface is now clearly visible under the cuticle.

Highth Nauplius.—One later Nauplius stage, with seven pairs (7 + 7) of the spines
on the fureca, was taken. Only one specimen was available for study and only the telson
was drawn.

As the telson (Text-fig. 15) bears seven pairs of spines, it undoubtedly is a very late
Nauplius stage. The degree of development of the maxillae and first and second maxil-
lipedes at our seventh Nauplius makes it highly unlikely that there would be more than
one Nauplius stage between Nauplius seven and the first Protozoea stage. It seems
evident, then, that the telson figured belongs to the eighth Nauplius stage. Only two
specimens of very late nauplii were found by us in the plankton catches. One, of which
the telson is figured, was preserved, and the other which, so far as we could judge from
low-power magnification of a living specimen in the field, was identical with it, moulted
to the first Protozoea.

PROTOZOEA STAGES.

The last Nauplius stage is followed by the Protozoea stages. As Heldt (1938) has
dealt very fully with the change in habits that accompany the change from the last
Nauplius to the first Protozoea, it is not intended to cover this ground again. However,
two factors which influence the future development of certain appendages of the
Protozoea stages should be mentioned. These are, firstly that the larva now feeds and
there is a great development of the maxillae and the cutting surface of the mandible;
secondly, that certain of the thoracic appendages, the maxillipedes, now aid the first
and second antennae in propulsion. Up till the end of the sixth Nauplius stage the
maxillae and first and second maxillipedes are not free. By the end of the seventh
Nauplius stage they are free but still poorly developed. However, during the Protozoea
stages they increase greatly in size, acquire quite extensive feathered surfaces and are
thus able to aid the first and second antennae in propulsion.

First Protozoea (Text-fig. 16).—The larva now measures from 0-781 to 0-872 mm. in
length. All the larval stages show quite a considerable variation in length. (All
lengths are from the base of the rostrum to the end of the telson—not including the
rostrum or telson spines.)

BY MURIEL C. MORRIS AND ISOBEL BENNETT.

abgal

28

Text-figures 16-31.
16-22, First Protozoea. 16, x 54; 17, Second antenna, x 68; 18, Mandible, x 67; 19, First
maxilla, x 130; 20, Second maxilla, x 123; 21, Second maxillipede, x 91; 22, Third maxillipede,
x 148.

23-31, Second Protozoea. 23, x 31; 24, Lateral aspect of abdomen; 25, First antenna, x 115;
26, Mandible, x 167; 27, First maxilla, x 160; 28, Second maxilla, x 164; 29, First maxillipede,
x 126; 30, Second maxillipede, x 130; 31, Third maxillipede, x 135.

172 LIFE-HISTORY OF A PENAEID PRAWN BREEDING IN A COASTAL LAKE,

The shape of the body has changed considerably. It now consists of an enlarged
oval anterior section, making up a little less than one-half the total body-length, and a
long cylindrical hind portion, the abdomen. The abdomen, which is still unsegmented,
ends in the now widely-forked telson, which still bears seven pairs (7 + 7) of spines.
The fourth, or middle pair of spines is the largest, and remains so right up to the post-
larval stages.

The Nauplius eye is still present. The paired adult eyes are visible as two opaque
masses situated just beneath the carapace at the anterior margin of the body.

The first antenna (Text-fig. 16) consists of a basal five-ssegmented portion and two
long distal segments. There are two large and one small terminal setae; two near the
end on the outer side; and one on the inner side at the junction of the two large
segments; one half-way down the second segment; and one on the most distal of the
five basal segments.

The exopodite of the second antenna (Text-fig. 17) now consists of ten segments,
while the endopodite has two segments. The last or most distal exopodite segment now
bears four long terminal hairs—the small hair of the Nauplius seven stage having
developed into a long plumose hair. The 4th, 5th, 6th, 7th, 8th and 9th segments each bear
a hair on their internal borders, while the 4th and 6th segments each bear an external
hair. The endopodite now bears five terminal hairs; and on the inner margin two at
the junction of the two segments and two half-way down the first segment.

The mandible (Text-fig. 18) has now lost its biramous appearance, while the cutting
surface, consisting of two distinct regions, is well developed. The right and left
mandibles are quite similar at this stage, each having a long, pointed tooth and a short
thick-set tooth separating the lower molar region of the mandible from the upper
cutting region.

The first maxilla (Text-fig. 19) is a biramous appendage with a three-segmented
endopodite and an unsegmented button-like exopodite. The two protopodite segments
form two quite well-defined endites on their inner surface. The most proximal, or first,
endopodite segment bears two hairs, the second two, while the terminal segment carries
five. The exopodite carries four long plumose hairs, while the proximal endite carries
six and the distal endite four rather short hairs.

The second maxilla (Text-fig. 20) is also a biramous appendage, the endopodite
being five-segmented, while the exopodite again forms a button-like unsegmented
structure. The first or most proximal endopodite segment carries three setae, the
second two, the third two, the fourth two and the fifth three setae. The exopodite
carries five long plumose hairs. The protopodite forms four endites on its inner surface
carrying eight, four, four and three setae numbering from the most proximal endite.

The first maxillipede (Text-fig. 16) consists of a four-segmented endopodite and an
unsegmented blade-like exopodite. The endopodite carries three setae on the first
segment, one on the second, one on the third, and five terminal setae. The exopodite
carries two terminal setae and also one internal and four external. The protopodite
carries a large number of hairs on its inner surface.

The second maxillipede (Text-fig. 21) closely resembles the first maxillipede at this
stage, except that it is smaller. The endopodite is four-segmented, while the exopodite is
once again an unsegmented blade-like structure bearing two terminal setae and also one
internal and three external setae. The endopodite carries two setae on the first segment,
one on the second, one on the third, and five terminal setae. Once again, the protopodite
is thickly beset with hairs on its inner surface.

Both Heldt and Pearson found for their respective species of Penaeus that the third
maxillipede was not present in Protozoea I, but in our species it has definitely made its
appearance and is an unsegmented biramous appendage at this stage (Text-fig. 22). It
earries two terminal setae on the exopodite.

Behind the third maxillipede, there are five distinct thoracic segments already
formed, which, in the second Protozoea stage, will carry the buds of the peraeopods.

Second Protozoea (Text-fig. 23).—The larva now measures from 1-0 mm. to 1.36 mm..
in length.

BY MURIEL C. MORRIS AND ISOBEL BENNETT. 173

The body still consists of the large anterior portion covered by the carapace, and
a long cylindrical posterior portion. The carapace, which is lengthening rapidly and
is now pear-shaped, still reaches only to the end of the ninth (third maxillipede) body
segment. It is becoming more deeply indented posteriorly.

Anteriorly, the carapace now bears a rostrum and two supra-orbital spines, one on
each side of the rostrum. The eyes at this stage are large and stalked and project well
beyond the anterior corners of the carapace. The Nauplius eye still persists.

The six abdominal segments are now clearly visible, although the sixth is still not
separated from the telson. The seven pairs of spines on the telson have increased
considerably in size.

Apart from the acquisition of one seta half-way down on the outer side and a pair
of hairs on the outer distal corner of the large second segment, the first antenna has
undergone no change (Text-fig. 25). The second antenna has undergone no change
at all. ;

The cutting surfaces of the mandibles (Text-fig. 26) display a definite dissimilarity
between the right and left—the greater change having taken place in the left mandible.

The first maxilla (Text-fig. 27) carries an additional two to three setae on the distal
endite, while the first or proximal endite and the fourth endite of the protopodite of the
second maxilla (Text-fig. 28) each bear an additional seta.

The first maxillipede (Text-fig. 29) bears an additional seta on the inner surface
of the second and third endopodite segments and a few extra setae on the proximal
protopodite segment.

The second maxillipede (Text-fig. 30) which shows indications of a fifth endopodite
segment, the first being very indistinctly divided from the protopodites, bears an extra
seta on all the endopodite segments except the third.

The exopodites of the third maxillipede (Text-fig. 31) now bear three setae, two
terminal and one on the outer edge.

Behind the third maxillipede the five thoracic segments, formed in the first Protozoea
stage, now carry the buds of five pairs of peraeopods (Text-fig. 24).

Third Protozoea (Text-fig. 32).—The larva now measures from 1.53 mm. to 1.73 mm.
in length, the greatest increase having taken place in the hind thoracic and abdominal
regions.

The carapace, which still reaches only to the end of the ninth body segment, is
now quite deeply indented at the posterior end. The supra-orbital spines, which are
already decreasing in size, are now situated quite close together at the base of the
rostrum.

The Nauplius eye persists.

The sixth abdominal segment, which is now completely separate from the telson, is
by far the longest of the abdominal segments. The third, fourth and fifth abdominal
segments each bear a median dorsal spine on their posterior borders; the fifth carries
an additional dorso-lateral spine on each side. The sixth abdominal segment carries a
minute pair of dorso-lateral and also a pair of ventro-lateral spines (Text-fig. 33). There
are still no signs of abdominal pleopods.

The uropods are now completely free—the exopodite bearing six and the epappodie
three setae. The telson still bears seven pairs of spines.

The first antenna (Text-fig. 34) now consists of four segments instead of the seven
of the previous stage. This tendency towards a loss of flexibility of the first antenna
is very important as it serves as a pointer to the fact that in the Mysis stages the first
antennae will no longer be responsible for propulsion. The arrangement of setae has
undergone no change since the previous stage.

The second antenna remains unchanged.

The dissimilarity between the right and left mandibles (Text-fig. 35) is now more
evident. The right mandible bears two dentated spines between the molar and cutting
regions, while on the left mandible, this intermediate region which was showing signs
of forming spines in the previous stage, now bears seven spines, three of which are

174 LIFE-HISTORY OF A PENAEID PRAWN BREEDING IN A COASTAL LAKR,

dentated. The molar regions in both right and left mandibles have increased consider-
ably in size.

Apart from the fact that the distal endite of the first maxilla (Text-fig. 36) now
bears nine setae, the proximal endite eight, and the protopodite of the second maxilla
(Text-fig. 37) now bears more hairs, the first and second maxillae have undergone no
change since the previous stage.

Text-figures 32-54.

32-40, Third Protozoea. 32, x 24; 338, Lateral aspect of abdomen; 34, First antenna; 35,
Mandible, x 82; 36, First maxilla, x 86; 37, Second maxilla, x 86; 38, First maxillipede, x 65;
39, Second maxillipede, x 54; 40, Third maxillipede, x 59.

41-54, First Mysis. 41, Lateral aspect of abdomen; 42, Ventro-lateral aspect of 6th
abdominal segment; 43, Telson and uropods; 44, First antenna, x 88; 45, Lateral aspect of basal
segment of first antenna; 46, Second antenna, x 54; 47, Mandible, x 103; 48, First maxilla, x 83;
49, Second maxilla, x 103; 50, First maxillipede, x 62; 51, Second maxillipede, x 49; 52, Third
maxillipede, x 65; 53, First peraeopod, x 51; 54, Fourth peraeopod, x 76. ‘

BY MURIEL C. MORRIS AND ISOBEL BENNETT. : 175

The first maxillipede (Text-fig. 38) bears two extra setae on the exopodite—one on
the outer border and one on the inner border, and there are one or two additional hairs
at the base of the proximal protopodite segment.

The exopodite of the second maxillipede (Text-fig. 39) bears an additional seta
on the outer border and there is also one on the outer edge of the endopodite.

The endopodite of the third maxillipede (Text-fig. 40) now bears two terminal setae,
while the exopodite still bears three setae.

The five thoracic segments behind the third maxillipede now bear quite well-
developed biramous appendages which still show no signs of segmentation or setation
(Text-fig. 32).

MYSIS STAGES.

A. First Mysis (Text-figs. 41-54).

B. Second Mysis (Text-figs. 55-57).

C. Third Mysis (early stages, Text-figs. 58-62; late stages, Text-figs. 63-67).

With the moult from the third Protozoea to first Mysis stage, the larva undergoes
several quite radical changes.

Undoubtedly the most important development, and one which changes the whole
appearance of the larva, is the growth of the peraeopods.

At the end of the third Protozoea stage they are biramous, unsegmented buds, but
by the first Mysis stage they are quite distinct biramous appendages showing signs of
segmentation, and bearing several setae. Up till the end of the third Protozoea stage,
the antennae and maxillipedes are responsible for the propulsion of the larva through
the water. However, at the moult to the first Mysis stage, the first antennae lose their
feathery and flexible appearance, and propulsion is taken over by the peraeopods, aided
to a small degree by the rapid jerking of the abdomen and tail fan.

A. First Mysis (Text-figs. 41-54).

The larva now measures 2-07 mm. in length.

The carapace reaches to the end of the fourteenth body segment (i.e., to the end
of the fifth peraeopod segment). The supra-orbital spines have practically disappeared.

The third, fourth and fifth abdominal segments (Text-fig. 41) no longer bear dorsal
‘spines and the fifth segment has also lost its dorso-lateral spines. The sixth abdominal
segment now bears a median dorsal spine; one pair of ventro-lateral spines and a well-
developed ventral hook (Text-fig. 42).

The telson and uropods (Text-fig. 43) now form a well-developed swimming tail-
fan, and the prominent indentation of the posterior border of the telson, which was so
characteristic of the Protozoea stages, has completely gone. It is replaced by a very
small indentation, the two lobes diverging only slightly. Each lobe still bears seven
pairs (7 +7) of spines, the outer two pairs now being placed further up the sides of the
telson.

The exopodites of the uropods now bear a spine on the external border near the
extremity in addition to eleven feathered setae along the internal border and tip. The
endopodites bear ten setae.

The first antenna (Text-fig. 44-45) still consists of four segments, but now bears
‘an internal unsegmented branch attached to the extremity of the third segment. This
internal branch, which is about one-third of the length of the outer or fourth segment,
bears two terminal setae. At the base of the basal segment, on the outer side, is a
swelling with a squared distal corner. This swelling will eventually lodge the statocyst,
while the squaring of the corner will give rise to the stylocerite. There are two feathered
setae on top of the swelling. The basal segment also bears a well-developed spine on
the ventral face towards the inner border.

The second antenna (Text-fig. 46) has undergone quite a radical change. The
exopodite has lost all signs of segmentation, and now consists of a plate-like structure
with eleven hairs along its borders—six internal, three terminal, and two external. The
endopodite remains unsegmented and only bears five setae—two terminal and three on
the internal border.

176 ’ LIFE-HISTORY OF A PENAEID PRAWN BREEDING IN A COASTAL LAKE,

The mandibles (Text-fig. 47) which have undergone very little change since the
previous stage and still show a very marked dissimilarity, now bear a small bud which
will eventually give rise to the mandibular palp.

Apart from the possible loss of one seta from the proximal endite, the first maxilla
(Text-fig. 48) has undergone no change.

The exopodite of the second maxilla (Text-fig. 49) has increased greatly in size
and is now beginning to take on the shape of the scaphognathite which it will eventually
form. It bears nine setae along its border, whilst the number of setae on the internal
borders of the protopodite has also increased.

The first maxillipede (Text-fig. 50) has undergone little change except that the
number of setae on the exopodite has decreased to seven again, while the number on the
protopodite segments has increased.

Much the same type of development has taken place in the second maxillipede
(Text-fig. 51). The number of setae on the exopodite has decreased to five, while the
number on the protopodite segments has increased. ‘Three of the endopodite segments
now bear a seta on the outer edge.

The third maxillipede (Text-fig. 52) which was rudimentary at the last Protozoea
stage, has undergone a marked development. The endopodite is five-segmented and
carries a number of hairs on its internal and external borders and at the extremity,
while the exopodite, which is not segmented, bears six hairs at its extremity.

It is in the peraeopods (Text-figs. 538 and 54) that the most important change has
taken place. In the third Protozoea they were rudimentary, biramous buds, but they
now consist of a large unsegmented exopodite with eight terminal hairs, and a small
unsegmented endopodite. The endopodite, which bears three to four hairs at the end,
shows signs in the first three pairs, of bifurcation. This will eventually lead to the
formation of the typical chelate appendages seen in the adult.

Slight swellings on the ventral surface of the abdominal segments (Text-fig. 41).
indicate the future position of the five pairs of pleopods.

B. Second Mysis (Text-figs. 55-57).

The second Mysis measures from 2-14 mm. to 2-26 mm. in length.

The supra-orbital spines have disappeared from the carapace but the abdominal
spines remain unchanged.

The slit between the two lobes of the telson is becoming smaller, while the space
between the first and second and second and third spines is increasing (Text-fig. 56).

The exopodite of the uropods now bears twelve setae along the internal border and
at the tip in addition to the spine on the external border. The endopodite still bears.
ten setae (Text-fig. 56).

The stylocerite on the basal segment of the first antenna (Text-fig. 57) is becoming
more evident, while the ventral spine is still very obvious. The number of setae on
the various segments has increased, while the internal branch has increased quite
considerably in size and now makes up more than one-third of the length of the fourth
or external antennular segment. ;

A spine has appeared on the external border of the exopodite of the second antenna
a quarter of its leigth from the extremity. The endopodite -is still one-half of the
length of the exopodite.

The only change which has taken place in the mandible is an increase in relative
size of the palp.

The first maxilla has undergone no change, except that, throughout all the Mysis:
stages, the endopodite is gradually decreasing in size.

The scaphognathite of the second maxilla has increased in size and now carries:
twelve hairs along its borders.

The first, second and third maxillipedes have undergone no change.

The endopodites of the peraeopods have increased considerably in length and show
signs of segmentation into five segments.

The pleopod buds have increased in size and now are clearly visible on the ventral
side of the abdomen (Text-fig. 55).

BY MURIEL C. MORRIS AND ISOBEL BENNETT. 177

C. Third Mysis (Early Stages: Text-figs. 58-62).
The third Mysis measures from 2-41 mm. to 2-75 mm. in length.
The carapace remains practically unchanged, and there are indications of two
minute rostral teeth.

Text-figures 55-79.

55-57, Second Mysis. 55, Lateral aspect of abdomen; 56, Telson and uropods, x 44; 57,
Basal segment of first antenna.

58-67, Third Mysis. 58, Lateral aspect of abdomen (early stage); 59, Basal segment of
first antenna (early stage) ; 60, Second antenna, x 40; 61, Mandible; 62, Telson and uropods,
x 45 (early stage); 63, Lateral aspect of abdomen (late stage); 64, Basal segment of first
antenna (late stage); 65, First peraeopod, x 98; 66, Fourth peraeopod, x 98; 67, Telson and
uropods (late stage).

68-79, First post-Mysis. 68, First antenna, x 80; 69, Second antenna, x 44; 70, Mandible,
x 15; 71, First maxilla, x 23; 72, Second maxilla, x 28; 73, First maxillipede, x 24; 74, Second
maxillipede, x 60; 75, Third maxillipede, x 41; 76, First peraeopod; 77, Fourth peraeopod;
78, Telson, x 68; 79, Distal outer corner of exopodite of uropod.

178 LIFE-HISTORY OF A PENAEID PRAWN BREEDING IN A COASTAL LAKE,

The abdominal spines remain unchanged.

The indentation between the lobes of the telson is no longer conspicuous while the
same increase in the distance between the lateral telson spines continues (Text-fig. 62).

The spine on the distal outer corner of the exopodite of the uropods is becoming
larger, and while the exopodite still bears twelve setae, the endopodite now bears
fourteen.

The stylocerite of the first antenna (Text-fig. 59) is now very clearly formed, while
the basal swelling itself now lodges the statocyst. There are three hairs on the top of
the statocyst swelling and the ventral spine is still well developed. The internal branch
has increased quite considerably in size, is still smaller than the fourth antennular
segment, and is still unsegmented.

The number of setae on the exopodite of the second antenna (Text-fig. 60) has
increased to fifteen, while the endopodite is now completely devoid of setae. The latter
now consists of two segments, a long distal segment which will ultimately give rise to
the typical “feeler”’ of the adult, and a small basal segment. A spine has appeared on
the ventral aspect of the distal protopodite segment.

The mandibular palp is larger but still unsegmented and does not bear any setae
(Text-fig. 61).

While the exopodite has disappeared from the first maxilla, the exopodite of the
second maxilla has continued to increase in importance and now bears twelve to
thirteen hairs.

The first, second and third maxillipedes have undergone no change. The endopodites
of the peraeopods are now definitely longer than the exopodites and clearly four-
segmented. The bifidity of the endopodites of peraeopods one, two and three, which
was evident in Mysis I and II, has now given rise to definitely chelate appendages,
while the endopodites of peraeopods four and five are long, flexible, non-chelate struc-
tures. The exopodites on all five pairs of peraeopods still carry eight terminal setae.

The pleopods (Text-fig. 58) now show a definite division into a basal portion (the
protopodite) and an unsegmented distal portion (the exopodite). They are still
uniramous.

Late Stages of Third Mysis (Text-figs. 63-67).

In these later stages the two rostral teeth are now quite distinct.

The indentation between the lobes of the telson (Text-fig. 67) is now very small,
while the first spine is placed almost half-way up the side of the telson. The protopodite
of the uropods now bears a spine.

Another spine has appeared beside the spine described previously as being present
on the exopodite of the uropods. The exopodite now bears thirteen hairs, while the
endopodite still bears fourteen.

_ There are still three hairs on the top of the statocyst swelling of the first antenna
(Text-fig. 64) and the internal branch, which is slightly smaller than the fourth
segment, is not segmented. The fourth segment, however, is showing signs of division
into two segments.

The number of setae on the exopodite of the second antenna has increased, while
the endopodite has continued to increase rapidly in size, although it is still not as long
as the exopodite.

The mandibular palp is now quite large but bears no setae nor shows signs of
segmentation. The cutting surface of the mandible remains unchanged.

The only change in the first maxilla is an increase in the number of setae on the
proximal endite, while the exopodite of the second maxilla still bears twelve—thirteen
hairs.

The first, second and third maxillipedes remain unchanged except for the
acquisition of an additional seta on the distal protopodite segment of the second mawxilli-
pede at the point between articulation of the exopodite and the endopodite.

BY MURIEL C. MORRIS AND ISOBEL BENNETT. 179

The peraeopods have undergone no important changes. The endopodites have
continued to increase in importance, but the exopodites are still quite well developed and
still bear the eight terminal setae (Text-figs. 65 and 66).

The pleopods (Text-fig. 63) have increased considerably in size, but are still
uniramous and bear no setae.

Post-MysIs STAGES.

Heldt (1938) has dealt very fully with the development of the post-Mysis stages of
several Penaeid species, and has classified the post-Mysis appendage development into
four different types.

As there is a gap between the first and the five much later post-Mysis stages which
were taken by us in the course of this survey, it was not possible to follow in detail all
changes in the post-Mysis series of stages. However, reference will be made to these
trends throughout the description of the first post-Mysis stages. Since we obtained the
first post-Mysis larva by moult from the last Mysis, we had no doubts about this stage.

The larva reared from Mysis III measured 2-83 mm.

The rostrum still bears only two rostral teeth.

The posterior border of the telson is now completely straight, and the indentation
between the two lobes almost gone. The first and second spines are gradually moving
up the lateral border of the telson (Text-fig. 78).

The small spine which appeared on the external border of the exopodite of the
uropods at the previous stage, has increased in size, whilst the other exopodite spine
has also increased in size and is now almost as long as a very small seta (Text-
fig. 79).

There are now five hairs on the top of the statocyst swelling of the first antenna
(Text-fig. 68). The internal branch is longer than the fourth antennular segment, but
it is still unsegmented. The outer branch (or fourth segment) is now clearly two-
segmented and there has been quite a noticeable increase in the number of hairs on all
the antennular segments. The ventral spine is still present.

The exopodite of the second antenna (Text-fig. 69) bears twenty-three setae, while
the endopodite is now five- to six-segmented.

The mandibular palp (Text-fig. 70) is quite a large structure, bearing six hairs on
the distal segment, and four hairs on the proximal segment. The number of hairs goes
on increasing, and the distal segment continues to increase in relative importance in
succeeding stages. The cutting surface has now lost practically all its teeth.

In the first maxilla (Text-fig. 71) the endopodite is very much reduced, while the
endites have increased considerably in size and now bear a larger number of hairs, all
of which seem to be shorter and thicker than in the previous stages. The endopodite
never completely disappears and at a later stage grows into a long, thin, segmented
structure such as Heldt describes for her species.

The endopodite of the second maxilla (Text-fig. 72) is reduced, but grows again, and
in a 9 mm. post-larval stage is just as Heldt has indicated for Penaeus trisulcatus. The
most proximal protopodite endite has disappeared completely, while the second one
remains as a very small endite bearing a very few hairs. The two most distal endites
are quite well developed. The exopodite has expanded quite considerably, and is
gradually forming the scaphognathite. It now bears twenty-two setae along its border.

The endopodite of the first maxillipede (Text-fig. 73) has lost all signs of segmenta-
tion, while the hairs on this and the exopodite have gone also. The two protopodite
segments have increased considerably in size and have acquired several more hairs on
their inner borders. The endopodite, like that of the first maxilla, should regain its
hairs and segments at a later stage.

The exopodites of the second (Text-fig. 74) and third (Text-fig. 75) maxillipedes
remain as small withered chitinous structures, while the endopodites are quite powerful
five-segmented appendages. The endopodite of maxillipede two is flexed backwards
between its third and fourth segments. In the later stages the exopodites gradually
develop again like the endopodites of the maxilla.

180 LIFE-HISTORY OF A PENAEID PRAWN BREEDING IN A COASTAL LAKE,

The exopodites of the five pairs of peraeopods have now almost disappeared and
consist of small withered chitinous structures. They reappear at a later stage, but
never develop to the extent that the exopodites of the second and third maxillipedes do.

The endopodites of the first, second and third peraeopods have developed into quite
large and powerful chelate appendages, while the fourth and fifth still form rather
longer, thinner, five-segmented non-chelate appendages.

Text-figures 80-96.

80-88. Rostrum. 80, 5:07 mm. stage, x 52; 81, 4:95 mm. stage, x 52; 82, 5:95 mm. Stage,
x 52; 88, 6-29 mm. stage, x 52; 84, 7-42 mm. stage x 52; 85, 7-84 mm. stage, x 52; 86, 9-28 mm.
stage, x 52; 87, 11-34 mm. stage, x 52; 88, 16:67 mm. stage, x 52.

89-96. Telson. 89, 5-07 mm. stage, x 73; 90, 4:95 mm. stage, x 73; 91, 6:29 mm. stage, x 73;
92, 7-84 mm. stage, x 73; 93, 9:28 mm. stage, x 73; 94, 11:34 mm. stage, x 73; 95, 16-67 mm.
stage, x 73; 96, 23-57 mm. stage, x 73.

The pleopods have increased in size though they are still uniramous appendages,
but they now bear setae. é

As Heldt (1938) has dealt very fully with the subsequent post-Mysis and post-
larval development of several penaeid species, it was considered pointless to describe
the further development of the individual appendages in detail, and so it was decided
merely to follow the further development of the telson and see how this fitted in with
the acquisition of the adult number of rostral teeth.

As mentioned above, there is quite a big gap in the post-Mysis series between the
first post-Mysis and the next stage (5-07 mm.) obtained in the catches, but even so
the number of rostral teeth in the latter has only increased to three (Text-fig. 80) and
the posterior border of the telson is only slightly rounded. The fourth pair of telson
spines have reached their greatest length at this stage (Text-fig. 89).

The faint beginnings of a fourth rostral tooth can be seen in a specimen 4:95 mm.
in length (Text-fig. 81). The posterior border of the telson (Text-fig. 90) is showing
signs of becoming pointed, while the fourth pair of telson spines is definitely becoming
shorter. There are clearly four rostral teeth present at the 5-98 mm. stage (Text-fig. 82)

BY MURIEL C. MORRIS AND ISOBEL BENNETT 181

while by the time the 6-29 mm. length is reached, a trace of a fifth rostral tooth is
evident (Text-fig. 83); also, the telson can be seen to have become distinctly more
pointed, while the fourth pair of spines have shortened considerably (Text-fig. 91). At
the 7-42 mm. stage, the fifth rostral tooth is clearly formed (Text-fig. 84).

Post-LARVAL STAGES.

As previously mentioned, the next stage obtained (7-84 mm.) really constitutes the
first post-larval stage taken, as it is in this stage that the pleopods become biramous.
A sixth rostral tooth (Text-fig. 85) is just beginning at this stage, while the posterior
border of the telson has continued to lengthen and the fourth pair of spines is shorter
(Text-fig. 92).

By the time the 9-28 mm. length is reached, the sixth rostral tooth is well-formed
(Text-fig. 86) while the telson is showing definite signs of approaching the adult form
of a long, pointed structure with no spines. The posterior border is distinctly pointed,
while the fourth pair of spines only just projects beyond the pointed posterior border.
The last three pairs of spines have also shortened considerably (Text-fig. 93).

At the seven rostral teeth stage (11-34 mm.) the fourth pair of spines is shorter
than the pointed posterior border of the telson, and the three small spines on the
telson tip have gone altogether (Text-fig. 94).

The full adult number of eight rostral teeth are present at the 16-67 mm. length
(Text-fig. 88), while the remaining pairs of telson spines have shortened considerably
(Text-fig. 95). By the time the 23-57 mm. length is reached, only the most proximal
pair of spines (i.e., the first) remain on the telson (Text-fig. 96). This last pair of
spines disappears after the next one or two moults, thus giving rise to the adult telson
which bears no large lateral spines at all.

For purposes of clarity, no reference has been made to the increase in the number
of setae on the dorsal surface of the telson throughout the post-Mysis and post-larval
stages. There are one or two long setae present on the dorsal surface of the telson near
the posterior border in the first post-Mysis stage, and these increase in number, until
finally, at the 23-57 mm. stage, the telson is thickly beset with setae along its edges,
on the dorsal surface—from the tip to the base. In addition, several minute spines
appear on the dorsal surface, but their development is entirely separate from, and must
not be confused with the gradual loss of the seven pairs of spines which were present on
the telson borders right from the last Nauplius stage.

Acknowledgements.

The field work in connection with this study was carried out with the aid of a
Commonwealth Research Grant to the late Professor W. J. Dakin.

We are very deeply indebted to Mr. and Mrs. K. H. Hargraves of Toukley, Tuggerah
Lakes. Without their generosity with their boats and gear and their personal help at
all hours, day and night, in all weathers, we should never have achieved any success
in the field. They allowed us to turn their home into a temporary laboratory and made
us always welcome.

We should also like to thank Mr. Hilton Chalmers, who took plankton catches for
us periodically at the Entrance; Miss E. C. Pope, M.Sc., of the Australian Museum, for
help in the field; Miss H. Newton Turner, B.Arch., Biometrician of the C.S.1.R.O.
McMaster Laboratory, for checking our sampling results; Professor E. A. Briggs, D.Sc.,
and Mr. A. N. Colefax, B.Sc., of the Zoology Department of the University of Sydney, for
reading the manuscript, and finally Professor P. D. F. Murray, D.Se., Challis Professor of
Zoology in the University of Sydney, for his advice and forbearance during the writing-up

of this paper.
References.
ANDERSON, W. W., KING, J. E., and LINDNER, M. J., 1949.—Early Stages in the Life History of
the Common Marine Shrimp, Penaeus setiferus (Linn.). Biol. Bull., 96, No. 2.
BURKENROAD, M. D., 1934.—The Penaeidea of Louisiana—with a Discussion of their World
Relationships. Bull. Amer. Mus. Nat. Hist., |xviii.
, 1940.—Preliminary Description of Twenty-one New Species of Pelagic Penaeida from
the Danish Oceanographical Expeditions. Ann. Mag. Nat. Hist., (11) 6: 35-54.
, 1949.—Occurrence and Life Histories of Commercial Shrimps. Science, 110, No. 2869.

182 LIFE-HISTORY OF A PENAEID PRAWN BREEDING IN A COASTAL LAKE,

DAKIN, W. J., 1938.—The Habits and Life History of a Penaeid Prawn (Penaeus plebejus
Hesse). Proc. Zool. Soc. Lond., 108, A, Pt. 2.
, 1940.—Further Notes on the Life History of the King Prawn, Penaeus plebejus Hesse.
Rec. Aust. Mus., xx, No. 5.
, 1946.—A Further Step in the Elucidation of the Strange Story of the Penaeid Prawns.
Aust. Jour. Sci., viii, No. 6. 3 :
, 1946a.—Life History of a Species of Metapenaeus in Australian Coastal Lakes.
Nature, 158: 99.
GURNEY, R., 1927.—Report on the Larvae of the Crustacea Decapoda. Camb. Exped. Suez
Ganal. Trans. Zool. Soc: Lond., xxii, Pt. 2; No: 15.
, 1943.—The Larval Development of two Penaeid Prawns from Bermuda of the Genera
Sicyona and Penaeopsis. Proc. Zool. Soc. Lond., 113, B: 1-16.
HeELpT, J. H., 1938.—La Reproduction chez les Crustacés Décapodes de la Famille des Pénéides.
Ann. VInst. Oceanog. Monaco, xviii, 2: 31-206.
Ipyuu, C. P., 1950.—The Commercial Shrimp Industry of Florida. Florida Board of Conserva-
tion, Educ. Ser. No. 6.
PANIKKAR, N. K., and Alyar, R. G., 1939.—Observations on Breeding in Brackish-water Animals
of Madras. Proc. Ind. Acad. Sci., ix, No. 6, Sec. B: 343-364.
PEARSON, J. C., 19389.—The Harly Life Histories of Some American Penaeidae, chiefly the
Commercial Shrimp, Penaeus setiferus (Linn.). U.S. Dept. Commerce, Bur. of Fisheries,
Bull. 30.

EXPLANATION OF PLATE XII.

Fig. 1.—Head of a mature female ‘‘Greasy Back’’ showing rostrum.

Fig. 2.—Adult female dissected to show gonads.

Fig. 3.—Eggs with Nauplii showing through egg membrane and one hatched nauplius.
(Photos. Gwen Burns.)

183

THE USE OF EXCISED SHOOTS IN LINSEED INVESTIGATIONS.
By H. B. Kerr,
Thomas Lawrance Pawlett Scholar, Faculty of Agriculture, University of Sydney.
(Plate xiv.)
[Read 28th November, 1951.]

Synopsis.

Investigations dealing with physiologic specialization in Melampsora lini (Pers.) Lév.
and the genetics of resistance in crosses between varieties of Linum usitatissimum L. have
been in progress at Sydney University since 1940. Techniques used by other workers were
found to have certain disadvantages. The excised shoot technique has recently been developed
to overcome these problems. It involves the excision of shoots above the cotyledons, and
subsequent growth in tap water or nutrient solution. The advantages of the method are
discussed and further developments indicated.

INTRODUCTION.

During 1948 investigations were commenced at Sydney University with particular
reference to the inheritance of resistance to Melampsora lini (Pers.) Lév. The tech-
niques adopted were those used by previous investigators (Waterhouse and Watson,
1941, 1948; Baker, 1945; Charles, 1947) at Sydney University, and patterned on U.S.A.
work (Flor, 1947). Flor, of U.S.A., in his genetical investigations grew and tested his
material in four and a half inch pots. He used the tip inoculation method, removing
infected leaves aS soon as the reactions became distinct and inoculating the same plants
with new races of rust at 7—8-day intervals. Here this method was soon found to have
distinct limitations.

It was frequently necessary to allow infected leaves to remain on the plant for some
time after sporulation in order to distinguish between susceptible and certain meso-
thetic reactions. Spores liberated at this time were a serious source of contamination
because these plants usually had to be inoculated with other races of rust. The most
susceptible segregates usually had to be removed. Others, greatly weakened by the
rust attack, often succumbed to root rot or a later rust infection. Even when infected
leaves were removed, it was impossible to control the rust spread to the cotyledons and
stem. In order to reduce contamination the infected shoot was cut back and fresh
growth from the base inoculated with the second race. This control measure was only
moderately successful, since prolonged growth of the plants in the glass house and
successive rust inoculations made the plants particularly liable to root rot.

DEVELOPMENT OF THE EXCISED SHOOT TECHNIQUE.

During 1948 it was noticed that linseed seedling tips, cut or broken off and left
lying on a free water surface, continued to grow and soon developed a strong root
system. Shoots of several varieties placed in tap water gave normal reactions when
inoculated with different races of rust. Arnon’s nutrient solution (Arnon, 1938) was
subsequently used and a more vigorous growth obtained. The value of these observa-
tions was apparent, and after preliminary trials a new technique for linseed
investigations has been developed.

EQUIPMENT AND METHOD.

The equipment consists of a Pyrex dish approximately 12” x 8” x 2” of about two
litres capacity. Fitting into this is an aluminium grid made of two sheets clamped
about half an inch apart. The bottom sheet, of slightly smaller dimensions than the
dish, fits into it and is suspended in position by the top sheet, which slightly overlaps
the edge of the dish, One hundred and forty 4” holes arranged in 14 rows of 10 holes

184 THE USE OF EXCISED SHOOTS IN LINSEED INVESTIGATIONS,

each are bored through both sheets (Plate xiv, fig. A). Although suspension of the
erid in tap water or nutrient solution did not visibly affect the plant growth or rust
reaction, it was lacquered as a precautionary measure. Thanks are due to the Food
Preservation and Transport Division of the C.S.I.R.O. for this lacquering.

Seedlings to be tested with different races of rust are sown in four-inch pots. The
seedlings are identified by the pot number and by loops of bell wire of different shapes
and colour. When the seedling is about three inches high the growing shoot is excised
about half an inch above the cotyledons and placed in a hole in the grid corresponding
to its number in the series. The shoot may be inoculated with rust immediately, but
if it shows signs of wilting may be stood over for a short period to recover. As the
rust infection develops, the shoot forms a callus, often very large (Plate xiv, fig. D),
and if rust infection is not severe it roots within three weeks (Plate xiv, fig. E).
The rooted shoots may be transplanted and grown to maturity: quite sturdy plants
have been obtained from the weakest specimens. Alternatively the shoots may be left
to set seed in situ if it is desired to determine the flower colour or obtain only a
small quantity of seed.

While the first shoot is being tested, the parent plant, kept out of range of
infection, will throw out at least two basal shoots, which are available for further
use. When ten plants have been grown in a four-inch pot, five shoots per plant have
been obtained without difficulty, and this number may easily be increased by reducing
the number of plants per pot or by cutting back rooted shoots and using them as a
source of further shoots. A very large number may be obtained from plants growing
in the field. The tendency of the main stem of vigorously growing spaced plants in
the field to develop a heavy crop of buds from the leaf axils may be used to advantage.
One hundred such shoots have often been counted on the one stem.

MopIFIED TECHNIQUES.
The first has been developed for the race differentiation studies.

Several excised shoots of each of the 20 differential varieties are bound in bundles
with a single twist of bell wire to weight them, and then placed in 2” x 1” glass tubes
arranged in two rows of 10 in a wooden stand 34” x 14” x 16”. The shoots in position
are inoculated and incubated in the usual way. Clearly a great saving of space is
effected (Plate xiv, fig. C).

Quarter-pint cream bottles and other larger glass containers have also been used
(Plate xiv, fig. B). It is not necessary to water the shoots so frequently, since they
fall with the level of the water in the container. These containers may be capped and
used as incubation chambers by temporarily lowering the water surface till the shoots
fall below the level of the mouth. After incubation the water level may be raised
and the shoots floated back above the mouth.

Sydney tap water has been quite adequate to maintain growth and obtain normal
rust reactions. Growth is not so rapid and succulent as in nutrient solution, but this
is an advantage if the shoots have to be held back for any period prior to transplanting.

Root AND CALLUS FORMATION.

Preliminary studies indicate that the rate and vigour of rooting of the excised
shoot are not markedly determined by the size of the shoot nor by the meristematic
activity of the growing point. Shoots from which the growing points have been removed
appear to root as readily as normally excised shoots, so that a single stem may be
cut into several lengths and each length rooted. By the time such cuttings have
rooted, axillary buds have developed at the top and normal vegetative growth is
resumed. Excised shoots from which the growing points have been removed and left
lying horizontally on the surface of the water frequently take up water and develop
a distinct callus at the lower cut surface. A vigorous callus forms even when such a
shoot is inverted with the upper end under water and the cut surface exposed to the
air. A large branch of the inflorescence of one plant, half broken away from the main

BY H. B. KERR. 185

stem, was held in place with bell wire. It continued to grow normally and within a
fortnight the injured surface had developed a strong callus.

Associated with this vigorous callus-forming capacity is the tendency, observed in
a recent summer sowing of young seedlings, to develop small suppressed green hypo-
cotyledonary buds. The bulk of these buds was observed at soil level, but many were
noticed slightly higher up. Their development was probably stimulated by the light
friction of the hypocotyledonary epidermis against the sandy soil particles in windy
weather.

ADVANTAGES OF THE HE)XXCISED SHOOT TECHNIQUE.

Since excised shoots are used problems of rust contamination are eliminated. Bench
space requirements are reduced by more than 50%. The equipment is more easily
handled and is cleaner than where pots are used, and the nutrient level can be kept
constant from one test to the next. The use of young shoots eliminates any risk of rust
reactions varying with increasing age. Root rotting organisms like Pythium and
Fusarium have not yet affected growth of the shoots in solution.

The prolific shooting capacity of linseed makes it possible to test the one plant
with a great number of races simultaneously or consecutively. Uninfected shoots may
be kept in reserve against the loss of the parent plant or transplanted into the field
for seed increase.

It is useful.in plant breeding work, where it may sometimes be necessary to test
a plant with a particular pathogen without exposing the parent plant to the disease
or introducing the disease to the field. A rapid increase of valuable plants is possible.
Thus a single plant has been increased a hundredfold in the last four months. Since
each new plant can now be used for further increase, this number could be quadrupled
in a fortnight. This should help to offset the relatively low seed yield obtained from
fibre types. It may also be possible to maintain a particular plant or variety vegetatively
from one season to the next and then increase it clonally. This will overcome problems
of cross pollination.

PROBLEMS HNCOUNTERED.

A few set-backs have been experienced in the course of this work.

The shoots succumbed rapidly to a bacterial invasion in hot weather, when over-
crowded and grown under conditions of low light intensity. Chlorotic shoots suffered
more severely than normal shoots, and unrooted more than rooted shoots. No trouble
has been experienced recently, since every third row in the aluminium grid has been
left empty to allow for better light conditions and circulation of air. It is not yet
certain whether excised shoots will live through the summer without controlled-
temperature facilities.

Two or three weeks after they have been placed in Sydney tap water containing
Arnon’s nutrient solution minus the trace elements, excised shoots: pass through a
stage when chlorosis of the new growth occurs. This does not affect the rust reactions
shown by the leaves that were inoculated earlier. Later, with the addition of tap
water to make up for loss caused by transpiration and evaporation, the chlorotic
condition disappears. The chlorosis is probably due to a temporary trace element
deficiency accentuated during rapid growth, but remedied by subsequent addition of tap
water. Initial experiments suggest an iron deficiency. The addition of rusty nails to
the solution has given adequate control.

CONCLUSION.

The technique has now been developed beyond the experimental stage and has
become the basis of all glass-house studies at Sydney University. One hundred of the
aluminium grids and dishes are in use, together with the equipment for the modified
techniques.

Several interesting lines of work which are worthy of investigation are opening
up, as for example, nutritional studies, their effects on rust reaction prior to and after
rooting, studies on rooting physiology, and the relation of root development to general

186 THE USE OF EXCISED SHOOTS IN LINSEED INVESTIGATIONS.

plant growth. Heterosis studies based on terminal growth rates of the shoot are made
easy since lateral bud development is normally suppressed.

Other avenues are opening up as the work proceeds.

Acknowledgements.

Grateful acknowledgement is made to Meggitt Ltd., whose grant of financial aid
has made this work possible.

I also wish to express my special indebtedness to Professor W. L. Waterhouse for his
constant encouragement and advice.

References.

ARNON, D. I., 1938.—Microelements in culture solution experiments with higher plants. Amer.
Jour. Bot., 25: 322-325.

Baker, E. P., 1945.—Unpublished work.

CHARLES, A. W., 1947.—Unpublished Honours Thesis. Filed at Faculty of Agriculture, University
of Sydney.

Fior, H. H., 1947.—Inheritance of reaction to rust in flax. Jowr. Agric. Research, 74: 241-262.

WATERHOUSE, W. L., and I. A. Watson, 1941.—A note on the determination of physiological
specialisation in flax rust. Jour. and Proc. Roy. Soc. N.S.W., 75:115-117.

, 1943.—Further determinations of specialisation in flax rust caused by Melampsora lini
(Pers.) Lév. Jowr. and Proc. Roy. Soc. N.S.W., 77: 1388-144.

EXPLANATION OF PLATE XIV.

A: Grid with excised shoots in position and lifted out of pyrex dish to show heavy root
development after about four weeks. x 1/3.

B: Cream jar with shoots floated above mouth of jar after incubation. x 5/9.

C: A set of the rust differential varieties in which the excised shoots are used in short
specimen tubes placed in position in a frame. x 3/16.

D: Basal stems of excised shoots, after about five weeks, with roots removed to show
extreme cases of callus formation. x1.

E: Extensive root development shown by an excised shoot after about four weeks. Top

growth removed for photographic purposes. x1.

187

REMARKS ON SOME AUSTRALIAN LAIUS GUER. (COL.: MALACHITIDAE).
By W. WITTMER, Buenos Aires.
(Communicated by J. W. T. Armstrong.)
(One Text-figure. )
[Read 31st October, 1951.]

Synopsis.

The Australian species hitherto placed in Laiws are distributed amongst five genera of
which two, Troglolaius and Flabellolaius, are described as new. Only one, possibly two, species
are retained in Laius; the majority are transferred to Dicranolaius Champ.; two to
Simoderus Ab.; the armicollis group is placed in the new genus Troglolaius and one new species
added, while microcerus is made the genotype of Flabellolaius. One synonym is noted L. ruguwli-
pennis Fairm. = nodicornis Blackb.

This note has been made possible through the kindness of Mr. J. W. T. Armstrong,
from whom I received a rich collection of Australian Malacoderms, and the comparison
of Fairmaire’s holotypes in the Paris Museum and paratypes of several Australian
species in the British Museum, London, which I had the opportunity to study in 1950.
This study showed that the genus Laius Guér. contains elements which are foreign to
Laius, and this to the extent that only one, or eventually two, species will remain in
this genus as far as can be seen from the material so far received. The different genera
can be easily recognized in the male through the key which follows. I take this
opportunity to express my sincere gratitude to Mr. Armstrong, of Nyngan, Dr. Balfour-
Browne (British Museum), and Prof. Jeannel and M. G. Colas (Paris Museum), who
helped me generously by lending me the material of the respective museums. As I am
continuing my studies on Laius and allied forms, I should welcome the opportunity of
examining additional material from Australia.

Key to Genera (males only).
1. Anterior tarsi simple in both sexes, second (third) joint of the antennae strongly dilated

TMT YD ctl een cicwerssteesres ay cslecs vase iey cxseinestaurawmetseiter oc ctcieey ci aieusts), cheers felcuici’s ye lei-eye einetieueisusnekegess euemereneys Laius Gueér.
Anterior tarsal joints 2 thickened, prolonged over 3 and nigropectinate at the tip in male,
second (third) joint of the antennae simple or strongly dilated ................... 2

Zee ccondss@third)santennaly joint stronely, dilatedias tye) leppelehs clsieloie ce ice) seksi) svete) aneiistelot el cllane=talne 3
Second (third) antennal joint simple~..=.......... ge Ne COTA REC OSM een Bear ata ars (res 4
SReEMainine, Joints) Of antennae Simple 3.5.2... .decee «cc 6 see ee Dicranolaius Champ.
ReEMAnMinewjOINts OL antennae Habellater yoy. -uerecie ere ieee sbsireneien ole Flabellolaius, nov.

4. Prothorax simple, without apical process, head simple, vertex not excavated .. Simoderus Ab.
Prothorax with an apical process, hornlike, more or less extended over the head, head
EXCAV ALE CMA beViCK LOR Wyte la eisrctetey strato cee ovata Star scie aa cares ete sreoa Vaan tice, Suata ara Maytetarie Troglolaius, nov.

LaIus Guér.

Among the material examined only L. filamentarius Lea was found to belong to
this genus, but according to Lea’s description L. minutus Lea might also belong to
this section.

DICRANOLAIUS Champ.

To this genus all those forms belong which have the general appearance of Laius
but the second anterior tarsal joint thickened, prolonged over the third and nigro-
pectinate at the tip in the male. The great majority of the Australian forms of Laius
belong in this group. Lea (‘Revision of the Australian and Tasmanian Malacodermidae’”’,
Trans. Ent. Soc. London, 1909, p. 151) says “. .. and the second joint of the front tarsi
in the males is always of peculiar shape and tipped with black”. It can therefore be
assumed that all the species mentioned in his revision, except the few referred to in
the next two genera, will have to be placed in Dicranolaius Champ. The following

188 REMARKS ON SOME AUSTRALIAN LAIUS,

were added by me on account of the character of this tarsal character, as described:
pallidus Lea, janthinipennis Lea, sordidus Lea, bellulus Guér., flavifrons Lea, cinctus
Redt., plagiaticollis Fairm., conicicornis Blackb., major Blackb., sinus Lea, villosus Lea,
carus Lea, nidicola Lea, alleni Lea, orthodoxvus Lea, tarsalis Lea, cavicornis Lea,
cyanocephalus Lea, rugiceps Lea, intermedius Lea, planiceps Lea, orcicornis Lea,
intricatus Lea, trifoveicornis Lea, albomaculatus Lea, inconstans Lea, semimaculatus
Lea, v-flavus Lea, maculiventris Lea, longus Lea, fimbriceps Lea, curvicornis Lea,
tetrasticus Lea, c-purpureus Lea, acervatus Lea, flavonotatus Lea, stenotarsus Lea,
concavifrons Lea, aulacophoroides Lea and rugulipennis Fairm. The type of L. ruguli-
pennis Fairm. (Pet. Nowv., 2, 1877, p. 174) is identical with specimens of L. nodicornis
Blackb. contained in the collection of the British Museum. L. nodicornis Blackb. (Trans.
R. Soc. 8. Austr., 10, 1886, p. 263) must therefore be considered as synonymous with
rugulipennis Fairm. (n. syn.).

Text-fig. 1.—Troglolaius armstrongi, n. sp.

SIMODERUS Ab.

This genus was erected by Abeille de Perrin in 1891 for one species from Siberia.
The Australian Laius flavopictus Lea (Trans. Ent, Soc. London, 1909, pp. 152 and 166,
figs. 148 and 149) and L. effeminatus Lea (Trans. R. Soc. S. Austr., xlv, 1921, p. 86) have
all the characters of Simoderus and must therefore be referred to it.

TROGLOLAIUS, nN. gen.

Several Australian species are characterized, in the male, by the prothorax provided
with a hornlike process extended more or less over the head and the head excavated at
the vertex with two or more foveae or impressions. Antennae, simple, 10-jointed,
sometimes joints 8 to 10 curved and atrophied (armicollis). Second tarsal joint of
anterior tarsi thickened, prolonged over third and nigropectinate at the tip. Last
abdominal tergite deeply excavated in some species. The following species can be
included in this genus: Laius armicollis (Lea), Trans. Ent. Soc. London, 1909, pp. 152 and
161, figs. 4 and 50; Laius apicicollis (Lea), Trans. R. Soc. 8. Austr., xli, 1917, p. 128;
Laius miraculus (Lea), l.c.; Laius sculptus (Lea), Trans. Ent. Soc. London, 1909,
pp. 152 and 162, figs. 61 and 144.

Genotype, Troglolaius armicollis (Lea).

TROGLOLAIUS ARMSTRONGI, N. SD.

6. Head black, sometimes with a faint bluish metallic shine, except genae, one very
small spot on the outer border of the two small interocular foveae and hind border of
the deep, transversal frontal excavation, reddish. Antennae black with the first and
second joints rufous underneath. Prothorax orange-red with a square black basal spot.

BY W. WITTMER. 189

Elytra bright metallic blue or purple with a transverse, complete antemedian orange-
red fascia, which is a little broader on the sides and suture than in the middle of each
elytron and a small orange-red spot on the suture at the apex. Legs and abdomen
black with a faint metallic shine.

Head with the eyes almost as broad as the prothorax, two small interocular foveae
and a quite deep transversal excavation on the front, less deep in the middle (Text-
fig. 1). Antennae short, serrate. Prothorax moderately transverse, hind angles rounded,
base feebly marginate, apex rounded and in the middle with a pointed process extended
over base of head, punctures small and sparse, surface clothed with a few erect black
hairs near the middle towards the pointed process, and several very long setae on the
side border. EHElytra almost parallel, strongly wrinkled, clothed with long blackish hair
intermixed with shorter whitish pubescence.

9. Head simple, slightly transversely depressed on the front.

Length, 4-5 mm.

Hab.— Bogan River, N.S.W., Australia, on Acacia pendula leg. J. W. T. Armstrong.
Holotype and allotype in Australian Museum collection (Nos. K.67692-3), paratypes in
my collection. I have much pleasure in dedicating this very interesting species to its
discoverer.

T. armstrongi is very closely allied to 7. sculptus (Lea) and can be easily separated
by the orange spotted apex of the elytra, which is metallic in sculptus.

FLABELLOLAIUS, nN. gen.
Genotype of this division is Laius microcerus (Lea) (Trans. R. Soc. 8. Austr., xii,
1917, p. 1380), characterized by having the antennae strongly flabellate from the fourth
joint onwards in male. Anterior tarsal joint 2 thickened, prolonged over 3 and
nigropectinate at the tip.

190

A REVISION OF AUSTRALIAN SPECIES PREVIOUSLY REFERRED TO THE
GENUS EMPOASCA (CICADELLIDAE, HOMOPTERA).

By Harotp F. Lower, Entomologist, Waite Agricultural Research Institute,
University of Adelaide.

(Plate xv, and seventy-five Text-figures. )
[Read 28th November, 1951.]

Synopsis.

This paper deals with the taxonomy of Australian species of the genus Empoasca
(Cicadellidae, Homoptera). As a means to this end, the external morphology of the genus
as a whole is discussed with special emphasis on the male genitalia. A brief account of the
distribution of the species together with their known economic importance is given and the
paper concludes with keys to the species, re-descriptions in detail of the already described
species, and the description of two new species.

INTRODUCTION.

Few, if any, complete studies of any Australian genus of leaf-hoppers have yet
appeared. From time to time, new species have been described, often without recourse
either to the types or the literature, in the transactions of various Australian and
overseas learned societies. For the genus Hmpoasca, this has resulted in the accumula- .
tion of unsorted species, misidentifications, synonymy, and even the inclusion of species
of other genera. Important economic species parade under synonyms, while others
which do not occur in Australia are recorded as doing damage to crops. The confusion
so established has been unwittingly perpetuated and added to by economic entomologists.

This study has therefore been undertaken, not with a view to making extensive
additions of new species, but to establish the validity or otherwise of those already
described, to give adequate re-descriptions of the accepted species, and to outline their
external morphology. The re-descriptions are given in more than usual detail since I
concur with Mayr (1942) that the fewer species of a genus there are known, the more
exact should the descriptions be. Unfortunately, all the original descriptions leave some-
thing to be desired in this respect. Six only of the previously described species are
shown to be valid, and to these, two new species are added.

The facts on which my conclusions are based are as follows: I have had, for study
purposes, the loan of every collection of leaf-hoppers in Australia; the only area not
represented is the Northern Territory. This material included the types of the six
species deposited in Australian collections as well as all specimens identified by Dr.
J. W. Evans. The authorities of the British Museum and the Hawaiian Sugar Planters’
Association have displayed endless patience in checking specimens with the three types
in, their possession, and in providing drawings of, and information about, these. To
reinforce this study, I have had material collected in the field for me by the officers
of the various State Departments of Agriculture. The morphological discussion has
been greatly assisted by the presence, in South Australia, of enormous numbers of
Empoasca viridigrisea Paoli in potato crops. I have read every original description and,
so far as I am aware, have missed no reference to the genus in Australia.

The limited number of known species gives no real picture of the actual position
of the genus in Australia, since six of the eight have been collected from cultivated crops
and are therefore of more or less economic importance. The two new species show that
others await collection, especially from native vegetation in the semi-arid parts of the
continent.

ECONOMIC STATUS OF THE GENUS.
The economic losses caused by leaf-hoppers have long been recognized in older
countries, but in Australia they have not yet received the detailed consideration which
they merit. If some relatively isolated examples be excepted, the damage done is not

BY HAROLD F. LOWER. 191

realized; it either passes unnoticed, or is ascribed to other agencies. Such damage may
be done in three possible ways.

Leach (1940) estimated that ninety per cent. of the known vectors of virus diseases
of plants occur in the Homoptera of which leaf-hoppers form a large part. As yet, no
species of Hmpoasca is so implicated, but this is possibly due to our lack of knowledge
of the biology of many species. Many closely allied forms are well-known vectors of
such diseases.

Others are toxicogenic feeders, that is, their saliva, which they inject while feeding,
has poisonous effects on plants. Such is H. fabae, the leaf-hopper responsible for the
“hopperburn” disease of potatoes in North America, a disease which, for many years,
was believed to be the result of adverse climatic conditions.

Many, when their rapid multiplication is favoured by their environment, can do
enormous direct harm by the withdrawal, during their feeding, of large quantities of sap
from growing plants. Such outbreaks may occur when a dry season, by restricting the
growth of their natural food plants, forces them to migrate to neighbouring cultivated
crops where they find ideal conditions.

Normally, most species are restricted in their breeding and feeding to particular
families of plants, and the introduction of new crops belonging to these families may
provide new sources of food for them. This has happened with both #H. terrae-reginae,
the cotton jassid, and H. viridigrisea, the vegetable jassid. The former originally bred
and fed on native Malvaceae, but it has now become a major factor in limiting cotton
production in Queensland (May, 1950). #. viridigrisea, which formerly lived on native
Solanaceae, has now become an established pest of potatoes and tomatoes in all parts of
Australia where these crops are grown. It has, however, adapted itself to a wide range
of plants of various families, having been recorded as attacking tobacco, cotton, beans,
lucerne, melons, celery, beet and many different weeds. (See Plate xv and Table 2.)

Apart from such obviously destructive species, others are annually responsible for
minor losses in various crops. Experimental work in the U.S.A. has shown that losses.
of up to twenty per cent. may occur yearly in lucerne crops as a result of the feeding
of species of Empoasca (Poos and Wheeler, 1943). To an unknown extent in South
Australia and elsewhere, H. viridigrisea, and in Queensland and New South Wales,.
H. alfalfae, may be so involved.

Furthermore, as cultivation in Australia is extended, with its consequent destruction.
of native vegetation, leaf-hoppers may be expected to become ever-increasingly important
crop pests, particularly in the warmer parts of the continent and in the artificial tropical.
environments provided in the Murray and other irrigated areas.

HIsTorIcAL RESUME. —

The first recorded species of the genus was described by Fabricius in 1794 as Cicada
flavescens. This was a European insect. Others were described under various genera
until Walsh (1865), in the U.S.A., erected the genus Hmpoasca for what we now know
to be three colour variations of H. fabae. These, he called viridescens, obtusa, and
consobrina. E. fabae Harris thus became the genotype.

Because colour was largely used as a distinguishing character, and this may vary
considerably in the one species, while two or more species may be almost identical in
colour, the taxonomy became obscure. Damage done by a single species was attributed
to several supposedly different species, while a number of species was grouped as one.
doing different kinds of damage in widely scattered areas.

The chaotic situation which had developed was made clear, and the mistakes
rectified only after the researches of De Long (1931) had revealed the highly charac-
teristic features of the male genitalia, and proven that distinct and constant differences
in these existed between the species. Using these differences as his criteria, he revised
the species in America, north of Mexico, and, in a lengthy series of subsequent papers,
first by himself, and later, with the aid of collaborators, added greatly to the numbers
and knowledge of the North American species.

The taxonomic history of Hmpoasca in Australia follows that common to the
taxonomy of Australian Insecta generally. The earliest descriptions were made by over-

192 AUSTRALIAN SPECIES PREVIOUSLY REFERRED TO EMPOASCA,

seas taxonomists working on material collected in Australia. As a consequence, the
older types came to be deposited in overseas collections.

The first Australian species recorded was described as Cicadula histrionicula by
Kirkaldy (1906), from material collected in Queensland by the Koebele and Perkins’
expedition in 1904. This species came from the Bundaberg district.

Two more Queensland species, Hmpoasca terrae-reginae and E. viridigrisea, were
added by Paoli (1936) from material in the British Museum. These came from Biloela
and Bowen respectively.

The following six species were all described from Queensland by J. W. Evans:
EH. bancrofti (EHvans, 1939), H#. athertoni and E. alfalfae (Evans, 1941), HE. maculata
(Evans, 1942a), and E. malvae and E. pulcherrima (Evans, 1942b).

As will be shown later, H. athertoni is not an Hmpoasca, while #. maculata and
E. pulcherrima are synonyms for #. terrae-reginae and EH. histrionicula respectively.

The genus therefore contains six valid, described species, and to these, I now add
merredinensis and bractigera bringing the number of species known to eight.

DISTRIBUTION OF THE GENUS (See map, Text-fig. A).

As stated above, six of the eight known species have been described from Queens-
land, to which State, so far as collections and records show, three of them, histrionicula,
terrae-reginae, and malvae, are confined. Two species occur in New South Wales as well
as in Queensland. These are alfalfae, apparently common in both States, and bancrofti.
The latter insect is rare in collections. The single specimen of bractigera came from
Mt. Keira in New South Wales, while all the specimens of merredinensis were collected
near the town of Merredin in Western Australia.

In very great contrast to the apparently limited habitats of these species, is the
exceptional distribution of H. viridigrisea, which ranges all over coastal Australia. It is
found all along such areas of Queensland as a common pest of tomatoes and lucerne. It
occurs in similar areas of New South Wales on potatoes, tomatoes, beans and lucerne.
In Victoria, it has been collected far inland on tobacco plants. i

I first identified specimens of it in South Australia in 1950. These came from the
irrigation districts of Renmark and Berri, where they were infesting crops of tomatoes
and potatoes. Shortly afterwards I collected it on cultivated Cucurbitaceae at Marion.
A very heavy infestation destroyed a large area of potatoes at Athelstone, and at the
same time it was common in lucerne at Mitcham.

In Western Australia the insects were first collected in 1942. These, and later
specimens, came from districts as far apart as Manjimup in the south and Wyndham
in the north. In these areas they were infesting solanaceous crops, other vegetables,
and home gardens.

The fact that the one species is common wherever its introduced hosts are cultivated
is very suggestive. Specimens from places as far apart as Wyndham in Western
Australia, Adelaide in South Australia, Nathalia in Victoria, and Charters Towers in
Queensland are identical. I have been unable to find either colour or morphological
differences in specimens from these or any other areas in Australia, and this statement
is based on the study of more than 2,400 specimens from all parts of the continent.
Were the species endemic in these areas, one would expect that evolution would have
resulted in at least a tendency towards sub-speciation. That such has not occurred
when the environmental conditions are so varied, and when it is remembered that
almost insuperable barriers to prevent natural dispersal exist, one is forced to the
conclusion that this insect has been distributed through the agency of cultivated
solanaceous crops since the coming of white settlement. This subject is of such interest
that I intend to deal with it in a future paper.

A careful search of literature and collections, as well as of the records of the
Tasmanian Department of Agriculture, has failed to reveal any species of Hmpoasca in
Tasmania, though I anticipate that #. viridigrisea will yet be found in those parts of
the island where potatoes are grown.

BY HAROLD F. LOWER. 193

THE GENUS AND ITS SYSTEMATIC POSITION.

Until 1931 the genus was regarded as one of world-wide distribution, having native
species in all the continents. In that year De Long (1931) demonstrated that the
“senus” was much more complex than had previously been thought, and since then it
has undergone subdivision.

De Long grouped the then known North American species into four subgenera.
These, in a recent letter to me, he states he has now raised to full generic status and
is erecting one or more genera for Mexican species.

His four North American genera are Empoasca s.str. Walsh (1865), Kybos Fieber,
1866, Hebata De Long (1931) and Jdona De Long (1931).

a viridigrisea

. terrae-reginaa

. alfalfae
. histrionicula — SCALE — Y
. malvae eed g

. merredinensis ;
; : no species known
. bancrofti

vouren-/

bDPOPrPLYryL

5 bractigera

Text-figure A.—Map showing known distribution of Auwstroasca in Australia.
The map has been made from actual records only. The shaded areas are those in which
A. viridigrisea is known to occur. The circles show where the other species have been collected.
1. A. terrae-reginae. 2. A. alfalfae. 3. A. histrionicula. 4. A. malvae. 5. A. merredinensis.
6. A banecrofti 7. A. bractigera.

Zakhvatkin (1946) has restored Fieber’s old generic name Chlorita for certain
Asiatic species, dividing it into the subgenera Chlorita s.str. and Hremochlorita.

To these I now add Austroasca, gen. nov., to include the Australian species formerly
included in Hmpoasca, and subdivide it into Austroasca s.str. and Paolia. The dis-
tinguishing characters of the new genus are given below.

As knowledge of the whole “group” increases, the number of genera must be
likewise increased. At present there is so little knowledge of the genera and species,
more especially in Asia, Africa, and Australia, that I am unable to make use of the
recognized taxonomic terms “tribe” or “group” for the assemblage of genera, since
possibly no one at present is in a position to circumscribe it; at the same time the
need is urgent for a term which will indicate the existence of such an unsorted and
unknown “group” of genera. For such a “group” of genera I propose the use of the
termination iti and refer to all the genera of “Empoasca” as the Empoasciti. No

194 AUSTRALIAN SPECIES PREVIOUSLY REFERRED TO EMPOASCA,

taxonomic significance is to be attached to the term. It is purely one of convenience
and signifies that such a “group” of genera exists and awaits the attention of a future
specialist, having at his command a representative collection from all parts of the
world so that the relationships between the genera can be defined and all these then
placed in their correct taxonomic category.

A viridigrisea Paoli

Text-figures 1-11.—A. viridigrisea Paoli.

1, complete insect, x 9; 1A, details of median white mark on scutellum; 2, face, x 9; 3,
antenna, x 109; 4, tegmen, x 9; 5, hind wing, x 9; 6, male genitalia (lateral view), x 27;
7, male genitalia (ventral view), x 27; 8, apical part of harpagone, x 109; 9, female genitalia
(lateral view), x 9; 10, blade of first valvula of ovipositor, x 109; 11, female genitalia (ventral
view), x 9.

The Empoasciti belong to the family Cicadellidae of the order Homoptera. They
constitute a well-defined natural group of leaf-hoppers, slender, and usually of small
size, few exceeding 5 mm. in length (Text-fig. 1).

BY HAROLD F. LOWER. 195

Austroasca resembles the other genera of the Empoasciti in all major morphological
structures except the dorsal hooks (see below). These have undergone such a reduction
as to be vestigial or no longer in existence.

EXTERNAL MorPHOLOGY OF AUSTROASCA.

In this section I deal with the external morphology of the Empoasciti, pointing
out, in the appropriate places, those features which are characteristic of Austroasca.

The Head. (See Text-figs. 12, 30, 38, 45, 53, 59, 68.)

When any species of leaf-hopper is viewed dorsally, the insect being so orientated
that the dorsal surfaces of the head and pronotum are in a horizontal plane, the visible
part of the head is the crown (Evans, 1946). This is seen to project medially to a
greater or lesser extent beyond the eyes, the extent of the projection (the crown
production) varying according as the horizontal is departed from. In order to be able
to make comparisons, a standard position must be adopted, and all the heads figured in
this paper have been so orientated before drawing. The term has no morphological
significance since it may comprise different morphological structures in different
genera; it is merely a convenient term for descriptive purposes. The crown may be
roundedly or angularly produced. In the Empoasciti it never includes any part of the
fronto-clypeus.

In Austroasca it is always broadly, more or less» roundedly produced, and the
sharp, pointed crown characteristic of the genus /dona De Long never occurs. On
either side, anteriorly, the dorsal half of each eye can be seen. The ocelli may or may
not be partly visible in the standard position. They are never completely visible. As
in leaf-hoppers generally, the crown is a very plastic feature and among the known
species of Awustroasca little correlation exists between crown shape and other
morphological characters.

For purposes of comparing the extent to which the crown projects in different
species I use what I term the Coronal Index (C.I.). This may be found as follows:
An exact drawing of the crown is first made and the distance between the eyes measured
at their anterior, visible, inner margins is the Eye Width (H.W.) (Text-figs. 20 and
21). The distance from the line joining these points to the anterior margin of the
crown, measured medially, is the Crown Production (C.P.) (Text-figs. 20 and 21). From

OJ2,
these measurements the Coronal Index is obtained by C.I. =

x 100 (Text-figs. 20

and 21) and is a number, varying within very small limits, characteristic of each species
irrespective of the considerable variations in size which occur among individuals of any
given species. In some species, for example, A. terrae-reginae, slight sexual differences
in the index occur, but these are never so large as to confuse the issue. The index for
each species is an average. In the case of A. viridigrise@, more than two thousand
specimens were measured. In all other species the index is an average of all the
specimens available, as shown in Table 1. In no instance was the variation between
minimum and maximum more than 1:5 per cent, an amount which, for this purpose,
is insignificant.

Crowns having a C.I. of 20 or more are obviously produced. Such are those of
A. terrae-reginae (C.I. = 27, Text-fig. 53), A. merredinensis (C.I. = 27, Text-fig. 59), and
A. alfalfae (C.I. = 23, Text-fig. 45).

Crowns having a C.I. of 15 or less tend to have their anterior and posterior margins
broadly rounded and parallel. This group includes A. bractigera (C.I. = 10, Text-fig. 68),
A. bancrofti (C.I. = 13, Text-fig. 12) and A. viridigrisea (C.I. = 13, Text-fig. 1).

In crowns of the intermediate group the production is not very obvious, though the
anterior and posterior margins are no longer parallel. Such are A. malvae (C.I. = 19,
Text-fig. 30) and A. histrionicula (C.I. = 17, Text-fig. 38).

The Face. (Text-fig. 2.)

When the head is viewed ventrally, three median sclerites are visible. These
collectively form what is referred to as the face, and comprise a small, triangular

196 AUSTRALIAN SPECIES PREVIOUSLY REFERRED TO EMPOASCA,

R A. bancrofti Evans R

Coronal Index, C.1=€% 2 4x100- \9 Coronal Index, C1.= Be = 3x19 =|6

Text-figures 12-19.—A. bancrofti Evans.
12, crown, pronotum and scutellum, x 9; 13, tegmen showing peculiarities of venation, x 9;
14, reconstruction of male genitalia (lateral view), x 27; 15, female genitalia (ventral view),
x 9; 16, harpagone, x 109; 17, brachone, x 109; 18, subgenital plate, x 22; 19, ‘dorsal hook,
x 109.
Text-figures 20 and 21.—Methods used in determining Coronal Index.

labrum, above which is the sub-rectangular ante-clypeus, while uppermost is a large,
long sclerite, the “‘clypeus” of Snodgrass (1935), or preferably the fronto-clypeus of
Evans (1946). In the Empoasciti the fronto-clypeus never transgresses on to the crown.
Underneath the labrum lies the long and rather stout labium. The three median sclerites
are long relative to their width, and this makes the face narrow and elongate. The
facial sutures are usually difficult to observe.

BY HAROLD F. LOWER. 197

On either side of the fronto-clypeus, and near its upper limit, is an ocellus. Hach
lies between the post-frontal sutures (Evans, 1946) and the eye. Sometimes they are
situated on the corono-facial angle. The two ocelli vary somewhat in size in different
species, but are never large and obvious.

On either side, near the termination of the frontal sutures (Hvans, 1946) and close
to. the eyes, are the antennae (Text-fig. 3), which are about equal in length to the face.
Bach consists of a large scape, pedicel and third segment. Then follows a number of
much smaller segments, the antenna terminating in a long flagellum. The antennal
ledges over the bases of the antennae are vestigial.

The Thorax. (Text-figs. 12, 30, 38, 45, 53, 59, 68.)

The thorax is well developed. It consists of a wide, rather long pronotum, concave
on its posterior margin, and a triangular scutellum.

The Tegmina. (Text-figs. 4, 13, 37, 44, 52, 58, 67, 74.)

The forewing or tegmen is long, narrow, slightly curved, and thickened on its basal
three-quarters. Its venation is highly characteristic. In dealing with this I follow, with
slight modifications, the notation of Evans (1946), who has given the most satisfactory
interpretation so far suggested.

In the Empoasciti the general plan is as follows (Text-fig. 4). The anterior margin
of the tegmen consists of C + Sec combined. Four long veins, R, M, Cu, and Cu.,, occur.
R divides into two branches, R, and R,», both of which tend towards the costal margin
of the tegmen. R,» always reaches the margin; Rij may or may not do so. Rs is always
absent. The second long vein is M, which terminates at the tegmen apex as M..,, the
anterior branch M,,, always being suppressed. M may or may not unite with R in the
‘apical part of the tegmen as R+ M. The third long vein is Cu,. This branches, at
about two-thirds of its length from the base of the tegmen, into Cu,,, which reaches
the margin at the tegmen apex and runs sub-parallei to M;,,. The second branch, Cup,
turns almost at a right angle and meets the posterior margin of the tegmen. The fourth
long vein, Cu., terminates in the posterior margin of the tegmen very close to the
termination of Cu,. A maximum of two cross-veins, r-m and m-cu, may be present. The
cross-vein m-cu is always present; r-m is present only when the veins R and M retain
their separate identities (Text-figs. 52, 58, 74). When R and M unite to form the
combined vein R + M (Text-figs. 4, 13, 37, 44, 67) it is part of M which connects R,» to
M;,,, The venation is distinct in the apical third only of the tegmen. The thickening
of its basal two-thirds and the thinning of the veins in this region make these very
difficult to see. Anal veins are never present.

All veins from R,» to Cu, reach the margin of the tegmen, and an appendix is
therefore never present. (An appendix is formed when the terminations of the veins,
instead of attaining the tegmen margin, are united by a sub-marginal vein (Text-fig. 5,
sub) which connects the ends of some or all of the veins, leaving a veinless area or
appendix between it and the margin.)

The principal variations in the tegmen venation of Awstroasca will be pointed out
as each species is dealt with later. One feature, however, may be mentioned here. In
all the specimens I have seen, the basal part of the tegmen is covered with a layer of
wax-like material which gives the tegmen a “mealy” appearance. Specimens which have
been stored in preserving fluids containing alcohol do not show this layer, since it
has been dissolved.

The Hind Wing. (Text-fig. 5.)

The venation is again characteristic. Near the base the combined vein C + Se
forms a thickened anterior margin to the wing, making a kind of “shoulder”. It
terminates suddenly near the end of this “shoulder”. The remainder of the wing
margin is very thin and difficult to observe. f

Six long veins occur, R, M, Cu,, Cu., 1A + 2A, and 3A. The tips of all these veins
are united by a sub-marginal vein (Text-fig. 5, sub) and hence none of them reaches
the wing margin. Between the sub-marginal vein and the wing margin is a large

198 AUSTRALIAN SPECIES PREVIOUSLY REFERRED TO EMPOASCA,

Text-figures 22-29.—A. terrae-reginae and A. viridigrisea (re-drawn from Paoli).

22-25, A. terrae-reginae.—22, subgenital plate; 23, brachone; 23A, tip of brachone showing
concave notching; 24, harpagone; 25, aedeagus.
26-29, A. viridigrisea.—26, subgenital plate; 27, brachone; 28, harpagone; 29, aedeagus.

veinless area. 1A + 2A, before joining the sub-marginal vein, forks into 1A and 2A.
A single, large, sub-rectangular apical cell is always present, closed by the sub-marginal
vein. Pre-apical closed cells never occur.

The Legs. (Text-fig. 1.)

The first two pairs of legs are of the generalized cicadellid type and need no
comment. The hind legs are long. Their femora bear a few strong spines distally.
The somewhat flattened tibiae bear strong spines in two or three series, more particularly
on the distal halves. The basal half of each tibia has a ventral comb of short bristles.

BY HAROLD F. LOWER. 199

A. malvae Evans

Text-figures 30-37.—A. malvae Evans.

30, Crown, pronotum and scutellum; 31, male genitalia (lateral view); 32, male genitalia
(ventral view) ; 33, aedeagus; 34, subgenital plate; 35, brachone; 36, harpagone; 37, tegmen.

The Abdomen.

This is typical of the Cicadellidae generally.

The Genitalia.

The female genitalia (Text-figs. 9, 10, 11, 15) follow the generalized homopterous
pattern and as this has been adequately discussed by Snodgrass (1935) little more need
be added. Only slight variations occur among the species of Austroasca. The female
genitalia are always large, prominent, and heavily bristled.

200 AUSTRALIAN SPECIES PREVIOUSLY REFERRED TO EMPOASCA,

The male genitalia are highly specialized and are characteristic. In discussing them
I use the terminology which Snodgrass (1935) adopted for his generalized description
of the male genitalia of the Homoptera, adding new terms for those structures only
which are peculiar to the Empoasciti, and giving these Greek names so as to be in
harmony with his terminology:

The aedeagus excepted, all elements of the male genitalia are paired, each member
of a pair being the mirror image of its fellow. The genitalia proper are so enclosed
as to be visible in prepared sections only.

From the lateral margins of the ninth abdominal dorsum there extends downwards
and posteriorly a pair of large lobes. The combined structure thus formed by the
dorsum and its lobes is called the pygophore (py, Text-figs. 31, 39, 46, 54, 60, 69), and
it encloses dorsally and laterally the genitalia proper.

From the ninth abdominal sternum there grow out two large, elongate, carinate
lobes which extend posteriorly to form the floor of the genital atrium. These are the
sub-genital plates (sgp, Text-figs. 6, 7, 14, 31, 39, 46, 54, 60, 69). In Auwstroasca their
tips are always upturned. They are always more or less heavily bristled, their
chaetotaxy being characteristic for each species. From two to four types of setae may
be present. Long, strong, thick, pointed bristles always occur. These are the ensiform
bristles. Along the apical half of the dorsal margin there is usually a row of very
short, stout bristles, the marginal setae. Similar setae may also.be scattered over the
outer surface of the apical parts of the sub-genital plates. Sometimes there occur very
long bristles, thick at the base and then rapidly tapering, ending in long flagella. These
are the flagellate bristles: they attain their greatest development in A. terrae-reginde
(Text-fig. 57). They occur to a lesser extent in A. alfalfae (Text-fig. 51). Finally, thin
seattered hairs of medium length may occur in various parts, but more especially
towards the base (Text-figs. 26, 34, 51, 57).

The genitalia proper consist of an aedeagus and two or three pairs of claspers.
The lower pair of claspers are the harpagones (h, Text-figs. 6, 14, 31, 39, 46, 54, 60, 69).
These are usually short and stout, and are always strongly sclerotized. Their bases
bear strong muscular attachments, the muscles controlling them being in the eighth
abdominal segment. Their muscular attachments readily distinguish them from other
members of the genitalia. They terminate in denticulations on the ventral side only.
Just before the denticulations begin, a small, variable number of strong bristles is
found. The position and number of the bristles, and the shape and number of the
denticulations are specific characters (see Text-figs. 24, 36, 41, etc.).

From the lateral walls of the pygophore there arises internally a pair of prominent
accessory structures whose purpose is to aid the harpagones during copulation. These
have no muscular attachments but are always strongly sclerotized, particularly towards
their apices. They are usually referred to as “lateral processes of the pygophore’”’. As
these must be repeatedly referred to in descriptions, and as the above expression is
somewhat lengthy and indefinite, I propose to call them brachones (that is, arms).
They are always much larger and more conspicuous than the harpagones, vary greatly,
in different species, in shape, length and curvature, and in themselves are often
sufficient for specific determination. They may be long and slender (Text-figs. 17, 23,
42), sail-like with broad bases and tapering tips (Text-figs. 27 and 63), almost straight
(Text-fig. 17) or strongly curved (Text-figs. 49 and 63). Their tips may be merely
tapered (Text-figs. 17 and 35) ‘or may be sculptured in various ways (Text-figs. 238A
and 49). In A. bractigera they are in the form of wide flat plates notched along the
apical margin (Text-fig. 72). They are one of the most plastic units of the anatomy of
the Empoasciti.

The aedeagus consists of a basal, more or less straight, attached portion and a
free dorsal portion. This latter consists of a pair of lobes whose size, shape and
curvature vary considerably in different species (Text-figs. 33, 56, 65).

The two terminal segments of the abdomen, segments X and XI,, together comprise
what is known as the anal tube. From its base, in most genera, there descends into

BY HAROLD F. LOWER. 201

A. histrionicula Kircaldy
Vie

42

VAN

dark brown
yellowish &reen

Text-figures 38-44.—A. histrionicula Wirkaldy.
38, crown, pronotum and scutellum; 38A, variation in crown markings; 39, male genitalia
(lateral view) ; 40, male genitalia (ventral view) ; 41, harpagone; 42, brachone; 43, subgenital
plate; 44, tegmen.

the genital atrium a pair of strong, usually curved, hooks. These are developed from
the tenth abdominal dorsum and serve, in those genera possessing them, as a third pair
of claspers. These are the dorsal hooks. They are greatly reduced in Awstroasca,
where they are either vestigial or entirely absent. When present, they are very small,
bluntly-pointed protuberances (ds, Text-fig. 46, and Text-figs. 19, 48, 62).

202 AUSTRALIAN SPECIES PREVIOUSLY REFERRED TO EMPOASCA,

DIAGNOSIS OF AUSTROASCA, gen. Nov.

Genotype, Austroasca viridigrisea Paoli (1936).

The general morphological characters of the Empoasciti are shared by all its genera.
Those characters which separate Austroasca from its allies are: The crown is always
wide relative to the distance between its anterior and posterior margins; it is broadly,
roundedly produced, the extent of the production never being great. In no known’
species is a C.I. of thirty or more found. Both male and female genitalia are heavily
bristled, ensiform bristles being always present and obvious. Dorsal hooks are either
vestigial or absent.

A.

alfalfae Evans

ZZ} thickened &reenish
area
Text-figures 45-52.—A. alfalfae Evans.

45, crown, pronotum and scutellum; 46, male genitalia (lateral view); 47, male genitalia
(ventral view); 48, dorsal hook; 49, brachone; 50, harpagone; 51, subgenital plate; 52, tegmen.
Note R and M distinct and presence of r-m.

In its affinities Austroasca most closely approaches the obtusa group of species of
Fieber’s genus, Kybos. In both, the crowns are broadly, roundedly produced and the
genitalia are heavily bristled. It differs greatly in the weakly developed or absent
dorsal hooks which, in Kybos, are prominent. Kybos is a North American genus, It is
probable that when the homopterous fauna of south-east Asia and Indonesia is better
known, more closely related genera will be found in these areas.

Characters Used in the Classification of Species.
With the small number at present known it is relatively easy to separate the species
of Austroasca, shortly after death, by colour differences. As the insects dry, however,

BY HAROLD F. LOWER. 203

great changes in the original colours occur. Blue and green (the latter a very common
colour in the Empoasciti) become a nondescript grey, and yellow tends to whiten.
White marks, originally present, may be lost, while a white pattern, not present in life,
may develop through various causes. The only colours permanent under all conditions
are black and brown. Specimens stored in liquids turn white except for such brown
or black marks as were originally present. As has been the experience in other parts
of the world, the discovery of new species will make even more difficult, and ultimately
impossible, specific separation on a colour basis alone.

A. terrae -reginae Paoli

Text-figs. 53-58.—A. terrae-reginae Paoli.

53, crown, pronotum and scutellum; 54, male genitalia (lateral view); 55, male genitalia
(ventral view); 56, aedeagus; 57, subgenital plate. Note great development of flagellate
bristles; 58, tegmen showing location of spot in cell Cu, and presence of r-m. R and M
separate veins.

For satisfactory determination of species, evanescent characters such as colour
(black and brown excepted) must therefore give way to characters which, irrespective
of the method or duration of preservation, will remain unchanged. Such stable
characters must be morphological, and are to be found in the venation of the tegmen,
the Coronal Index, and the male genitalia. Of these, the latter are of first importance.
When viewed laterally and ventrally, the general arrangement of the parts, and the
shape of each part, are characteristic for each species. Useful confirmatory evidence is
provided by the detailed structure of the brachone and harpagone, and the chaetotaxy
of the subgenital plate.

Adoption of these characters implies that new species shall not be described from
females alone. In the absence of good characters, identification of known female forms
is difficult; to set up new species without being able to separate them at any time from
the described forms is poor taxonomy. Where the species is of economic importance,

204 AUSTRALIAN SPECIES PREVIOUSLY REFERRED TO EMPOASCA,

both sexes will occur together; if the would-be identifier has no males in the material,
the remedy is obvious. To help the worker who may have female material only, I have
included separate keys for each sex, but the makeshift nature of the key for females

must be borne in mind.

A. merredinensis sp nov

C+Se Ria

67 Cu, Guib x?

Text-figures 59-67.—A. merredinensis, sp. nov.
59, crown, pronotum and scutellum; 60, male genitalia (lateral view); 61, male genitalia

(ventral view); 62, dorsal hook; 63, brachone; 64, harpagone; 65, aedeagus; 66, subgenital

plate; 67, tegmen.
A few words on the description of new species of the Empoasciti may not be out of
place here. To define the species clearly, figures are of much more value than verbal
descriptions. Apart from what else it contains, a description which does not figure the
male genitalia as a whole from both the lateral and ventral aspects, and separate details
of the structure of the parts of the genitalia, cannot be regarded as adequate. Without

BY HAROLD F. LOWER. 205

these figures, future identification is uncertain; with them, no question can ever arise
as to what species is referred to.

Methods of Study and Techniques Used.

The Coronal Index is first found as explained above. This limits the number of
possible species to which’ the specimen may belong, and this number is further limited
by the nature of the tegmen venation.

The next step consists in the preparation of the male genitalia so that the parts can
be clearly seen. After giving various methods a trial, I have finally standardized on the
following, which meets all my requirements:

If the insect has been freshly killed, or has been stored in a liquid medium, the
terminal five or six segments of the abdomen are removed. This is done with fine
needles, made by using the smallest entomological pins, with the heads cut off, in metal
holders. In dried specimens, the same segments may be removed with a small pair of
very sharp scissors.

A bractigera sp nov

Text-figures 68-74.—A. bractigera, sp. nov.

68, crown, pronotum and scutellum; 69, male genitalia (lateral view); 70, male genitalia
(ventral view); 71, subgenital plate; 72, brachone (note characteristic shape and denticula-
tions) ; 73, harpagone; 74, tegmen.

The sections are put in 20 per cent. caustic potash solution and left therein until
they have cleared as is shown by all the elements of the genitalia becoming plainly
visible. No heat should be applied, as this usually produces distortion. Dried insects
clear more rapidly than those which have been stored in preserving fluids. The latter
may take two or three days.

When cleared, the section is removed to distilled water for fifteen minutes, the
surplus segments providing a means of manipulating it with either needles or needle-
pointed forceps without damaging or causing distortion of, the parts of the genitalia.
At the end of this time, it is transferred first to 90 per cent. alcohol for ten minutes and
then to a one per cent. solution of basic fuchsin in 95 per cent. alcohol for ten minutes.
The staining is not essential but is desirable since, in pale-coloured specimens, some of
the structures may become invisible during subsequent treatment.

U

206 AUSTRALIAN SPECIES PREVIOUSLY REFERRED TO EMPOASCA,

From the stain, the section is put in absolute alcohol for ten minutes. From this it
is removed to a drop of cedar-wood oil in a hollow-ground slide where it is moved to a
lateral position so that the two harpagones and the two brachones are superimposed.
Fine needles are used to arrange the section in this position, and movement is prevented
by means of small fragments of glass placed round it. Using a x10 ocular and a x10
objective, a careful drawing of the genitalia is made. ;

The section is then turned so that its ventral surface is uppermost, kept in position
as before, and drawn.

These drawings having been completed, the section is washed in xylol, and then
placed in a drop of ‘“Sira’”’ or other permanent mountant on a microscope slide, where
it is carefully dissected under a binocular. This is best done by rotation the genitalia.
till the ventral side is uppermost and then dividing the section in two by inserting a
needle point at the base of the subgenital plates and carefully separating them with
another needle. The membranes will be found to tear much more easily than the
sclerotized parts of the genitalia. A cover slip is then applied.

From the slide so prepared, exact drawings can be made of all elements of the
genitalia. For this work, I find a x10 ocular and a x40 objective satisfactory, fine
details being checked by changing to a x25 ocular, thereby getting a magnification of
1,000.

All the figures in this paper have been made from my own drawings of the actual
insects and sections prepared as above, except Text-figures 14-19, which were drawn
from the prepared genitalia of the type, and Text-figures 22-28, which are redrawn from
Paoli (1936). For accuracy, these latter leave little to be desired. I have used a camera
lucida for the drawings. By doing them myself, I am able to guarantee accuracy of
detail.

Notes ON DISCARDED SPECIES, MISIDENTIFICATIONS, AND SYNONYMY.
Empoasca athertoni (Evans, 1941).
This is not a member of any genus of the Empoasciti. Lacking the necessary
knowledge, I am not qualified to place it in its correct genus. It appears, as the
following structural details show, to belong to the tribe Jassini of Evans (1947).

The face is very wide and short, its width being nearly twice its length.

The crown is extremely wide relative to the very short distance between its anterior
and posterior margins, and the ocelli are on it near the anterior margin.

The tegmen has a large appendix, and shows little reduction in venation. In addition
to the veins normally present in the empoascit tegmen, Rs, M,,., and a distinct Anal
vein occur. All veins are strong and clearly evident in the basal part of the tegmen.

The hindwing also has a much more complete venation. In addition to the veins
normally present in the empoascit hindwing, the following occur: Se is a separate vein;
Roa, Riss, Mis,, and M;,, are all present, as are both Cu, and Cu,. There are three apical
cells closed by the submarginal vein which terminates at the second Anal vein. These
cells are cell R.,;, cell M,,., and cell Msg,,.

The male genitalia are distinctive. The subgenital plates are very short and stout,
while the aedeagus is very similar to that of members of the genus Jassus, and quite
unlike that of any known genus of the Empoasciti.

“Empoasca terrae-reginae” for EH. viridigrisea Paoli.

In 1941, Evans identified as Empoasca terrae-reginae Paoli a green leaf-hopper
reported to be abundant in lucerne, on tomatoes and other vegetables, and on weeds, in
both Queensland and New South Wales. At the same time he gave a partial description
of the insect (Evans, 1941).

Amongst the material examined by me was a number of specimens identified by him
as E. terrae-reginae. I carefully compared these and his description with Paoli’s
original description and figures of H. viridigrisea (Paoli, 1936). As the specimens and
description agreed in every detail with Paoli’s account, I came to the conclusion that
Evans’ “EB. terrae-reginae Paoli” was, in fact, H. viridigrisea Paoli.

BY HAROLD F. LOWER. 207

A number of specimens of this insect, including two identified as “H. terrae-reginae”’,
together with a number of prepared genitalia, were submitted to the British Museum
authorities with the request that they be compared with Paoli’s type of HL. viridigrisea.
(All the specimens bore numbers only.)

They confirmed my opinion that the insects were H. viridigrisea Paoli and added,
“we consider it identical with a species referred to by Dr. J. W. Evans as Hmpoasca
terrae-reginae Paoli in Proc. Roy. Soc. Queensland, 52, p. 11, 1941”.

This misidentification has been perpetuated since that date in the literature of
economic entomology. The species must now revert to its correct name, A. viridigrisea
Paoli, 1936.

Empoasca maculata Evans (1942qa).

A study of Paoli’s original description of Hmpoasca terrae-reginae (Paoli, 1936)
leaves no doubt that H. maculata is a synonym for this species.

The subgenital plates, alone (see Text-figs. 22 and 57), are so highly characteristic
as to be sufficient in themselves, but all other parts of the genitalia of maculata are
identical with those of terrae-reginae. Both are yellow insects and each has a small
brown spot on the tegmen. I figure the drawings of Paoli of both this species and his
viridigrisea, and, in an appendix, include my translations of his descriptions so that the
facts can be checked by anyone interested.

It is peculiar that this synonymy should ever have arisen since, as late as 1939,
J. H. Smith (1939), in the Entomological Section of the Annual Report of the Queensland
Department of Agriculture and Stock, was well aware that the Queensland cotton jassid
was EH. terrae-reginae, for he writes: ‘The cotton jassid, H. terrae-reginae Paoli, proved
more serious than for some time past, and many areas were heavily infested late in the
season. Work on this pest at the Biloela Research Station is a joint project between the
plant breeder and the entomologist and strain resistance is a feature of present
investigations.”

It may be noted in passing that Paoli’s type of terrae-reginae came from Biloela,
where it was attacking cotton. The cotton jassid therefore reverts to its original
name, A. terrae-reginae Paoli.

Empoasca pulcherrima Evans (19246).

Having read Kirkaldy’s original description of his Cicadula histrionicula (Kirkaldy,.
1906), I was convinced that EL. pulcherrima was a synonym for this species.

Specimens of pulcherrima were therefore sent to Mr. C. EH. Pemberton, the
Entomologist of the Hawaiian Sugar Planters’ Association in Hawaii, asking him to
compare them with Kirkaldy’s type. His reply is as follows: “In comparing the
specimens [of H. pulcherrima] with _our type of EH. histrionicula Kirk., I find no differ-
ence. Your three specimens vary slightly in colour pattern on the dorsum. The specimen
in the center matches our specimen almost exactly. Though our specimen has lost its
tegmina, the wings are still present, and the venation seems to match exactly the
venation of your specimens. Your specimens are faintly larger; but I do not consider
the difference significant. .. . I am strongly inclined to believe that your specimens;
are actually E. histrionicula Kirk.”

Since pulcherrima corresponds exactly with Kirkaldy’s description, this name must
be discarded and the species revert to its original name of histrionicula Kirk.

Empoasca fabae Harris.

This insect has been recorded (Anon., 1944-1945) as attacking tomato plants in
Queensland. This reference has been carefully checked and it appears that no
identification of the insect was ever made. In view of the host plant, and the locality
from where it was reported, it is virtually certain that the insect was A. viridigrisea
Paoli, its general colour and pattern having much in common with those of the North
American species, though the different shapes of the crown in each would easily
differentiate them.

There is no evidence to suggest that H. fabae, or any other exotic species, occurs in
any part of Australia.

208 AUSTRALIAN SPECIES PREVIOUSLY REFERRED TO EMPOASCA,

THE GENUS AUSTROASCA, ITS SUBDIVISIONS, AND RELATIONS BETWEEN THE SPECIES.

The eight known species of Auwstroasca all share alike in the common heritage of
the Empoasciti. Seven of the species, however, resemble each other much more closely
than any one of them does the eighth.

The aberrant member of the genus is A. bancrofti, of which I have seen five specimens
only. This is the only species of which I have had no material for microscopic
preparations. Fortunately the tegmen and the genitalia of the type (male) have been
separately mounted. Of the remaining specimens, two are complete females and two
are females which have lost their abdomens.

For this species I erect the subgenus Paolia and distinguish it from the subgenus
Austroasca s.str. as follows:

SUBGENUS PAOLIA, NOV.

The average size is considerably greater than that of species of Austroasca s.str.

The sutures of the head are complete and easily seen.

The venation of the tegmen is unique. R, instead of occupying its normal position
(see R, Text-figs. 4, 37, 44, 52, 58, 67 and 74), runs parallel, and very close, to the
costa (see R, Text-fig. 13), until about half-way along the tegmen, where it unites with
the costa. It soon turns posteriorly and then apically uniting with M. The combined
vein R+M gives off the branch R, to the costal margin and then forks into R, and
M,,.,. In contrast to the venation of Austroasca s.str., where the cross-vein m-cu connects
M;,, only with Cu,, in this subgenus it connects R+M with Cu, so that the three veins
Ry», Ms.4, and m-cu are all in contact at one point. This has been brought about by
the basad movement of m-cu, its resulting increase in length making it as long as Cu,
though in Austroasca s.str. it is nearly always shorter, and usually very much shorter,
than Cup.

The genitalia apparently differ in certain respects from those of Austroasca s.str. A
normal pygophore either has not been developed or has been damaged in mounting.
A torn fragment (shown dotted in Text-fig. 14) may be part of a pygophore lobe.

The subgenital plates are much more weakly bristled than in other species of the
genus, while the harpagones appear to lack bristles.

Unfortunately the spatial relation between the parts cannot be known at present.
The genitalia of the type have been mounted after partial dissection, and Text-figure 14
is a reconstruction made from drawings, all on the same scale, of the separate parts.
The possibility of serious error is obvious when such a procedure is adopted, and
whether the position of the species in the subgenus is confirmed, whether it is shown
to be merely an aberrant form within the genus, or whether its complete removal
from Austroasca s.lat. is necessary, the future study of new material alone can decide.
In putting it in a separate subgenus, it can be later eliminated, if necessary, without
any alteration to Awstroasca s.str.

‘While the seven remaining species of Austroasca form a more or less homogeneous
group, three well-defined subgroups, based primarily on variation of the tegmen
venation, can be distinguished. The variation in the venation can be correlated with
the degree of specialization of the genitalia. When so grouped, genetic relationships
are made clear and, at the same time, there is some indication of the evolutionary
level attained by the various species.

I believe that the most generalized forms occur in the viridigrisea subgroup and
that these probably most closely approach the austroascan ancestor in structure.
In the tegmen of this group the combined vein R+M is present, r—m is therefore
lacking, and R,. meets the costal margin of the tegmen obliquely (Text-figs. 4 and 67).
The subgroup contains two species, viridigrisea and merredinensis. Both species exhibit
similar characters in the genitalia. The subgenital plates (Text-figs. 26 and 66) are
short, wide, and possess relatively few strongly-developed ensiform bristles. The
harpagones (Text-figs. 28 and 64) are relatively short and stout, and their denticulations
are of similar pattern. Both have wide sail-like brachones (Text-figs. 27 and 63) with

BY HAROLD F. LOWER. 209

broad two-pronged bases and tapering, curved, simple tips, while the free dorsal lobes
of the aedeagus are curved in an open arc (Text-figs. 29 and 65).

The histrionicula subgroup is equally distinctive. It contains the species
histrionicula and malvae. The tegmen venation is similar to that of the viridigrisea
subgroup except that R,. is vestigial, appearing as a mere thickening of R near where
R,. should fork. Rj, is not present as a distinct vein and does not reach the costal
margin (Text-figs. 37 and 44). It may be fortuitous that the tegmina of both species
show a similar colour pattern, and that the colours and pattern on the body generally
are not unlike. The genitalia show that this subgroup has attained a higher evolutionary
level than the first. The chaetotaxy of the subgenital plates (Text-figs. 34 and 43) is of
a similar pattern and is of a more complex nature than that of the viridigrisea subgroup.
A few flagellate bristles, though not well developed, are present. The harpagones are
of a similar structure (Text-figs. 36 and 41). but it is in the brachones (Text-figs. 35
and 42), the most plastic feature of the genitalia of the Empoasciti, that the greatest
changes have occurred. The broad two-pronged base has been lost, though the spur
in malvae may be a vestige of a prong. The brachones are now slender and cylindrical,
the terminal halves being straight with simple tapering tips. The aedeagus (Text-figs.
33 and 39) is very similar in each, the dorsal lobes assuming the form of short, thick,
crescents.

The highest evolutionary development is to be found in the terrae-reginae subgroup,
consisting of terrae-reginae, alfalfae and bractigera. In the tegmina, the cross-vein r-m
is present since R and M retain their separate identities (Text-figs. 52, 58 and 74).
R,a meets the costal margin perpendicularly, though it may be difficult to see, as is the
case in bractigera. The genitalia show the peak of evolution attained by known species
of the genus. The chaetotaxy of the subgenital plates is obviously of the same pattern
(Text-figs. 51, 57 and 71). All four types of bristles occur. There is a tendency towards
the development of an abundance of long flagellate bristles which are so strongly.
developed in terrae-reginae that it can be identified by this character alone. The
harpagones (Text-figs. 24, 50 and 73) have become short and stout, each terminates in
a Spine, and each bears a small number only of short stout bristles. The brachones
(Text-figs. 23, 23A, 49 and 72) have undergone further modification, since there is now
a tendency for their terminations to be sculptured. They are always strongly notched
in alfalfae, saw-like, apically, in bractigera, and very often concavely notched in terrae-
reginae. Text-figures 55, 47 and 70 show that the general arrangement of parts is
similar in the three species.

Based on the above facts, I arrange the species in the following evolutionary scale.
Of the known species I regard viridigrisea as the most generalized, followed in succes-
sion by merredinensis, malvae, histrionicula, bractigera, and alfalfae, with terrae-reginae
as the most specialized species.

In view of the small number of known species, these ideas must be tentative, but,
at the same time, the facts are suggestive.

CONCLUSION.

In this study I have conscientiously tried to apply those principles which I regard
as essential to the satisfactory revision of any group. I have done much painstaking
research, including a thorough study of types, and of all the literature bearing on the
subject, more especially the original descriptions. I have given complete re-descriptions
of all species needing it, together with, I hope, adequate illustrations. Finally, I have
tried to make keys which will enable any entomologist who is prepared to familiarize
himself with the characters used, and to make careful microscopic preparations, to
ascertain what species he is identifying (if known), or to determine definitely whether
it is an undescribed one. I believe I have laid a foundation on which those who follow
can build with confidence.

Note: In Table 1 is shown the number of specimens of each species examined and
the locations where these were collected; in Table 2 a list of the known food plants
of each species is given.

210

AUSTRALIAN SPECIES PREVIOUSLY REFERRED TO EMPOASCA,

TABLE 1.

Showing Numbers of Specimens Studied and Districts where These were Collected.

Queensland—
No district
Ayr..
Biloela
Bowen
Boyne River
Brisbane
Brookfield
Callide
Cambooya
Charters
Towers ..
Hidsvold
Gatton
Gayndah
Gracemere
Horne Hill
Humphrey
Indooroopilly
Jandowae ..
Kilmore
Kingaroy ..
Kongaroy ..
Mimgary
Mundubbero
Pentland
Rockhampton
Toowong
Yamala
Yeeroongpilly
New South
Wales—
Bellbrook
Bolwarra
Branxton ..
Camboyne
Cabramatta
Dundas
Dungog
Gunnedah
Maitland
Millthorpe
Mount Keira
Mudgee
Narellan
Nemingha
Paterson
Pitt Town..
Prospect
Raworth
Roslyn
Seaham
Sydney
Tamworth
Wauchope
Wilberforce
Windsor
Wingham

A. viridigrisea.

Oy
+0
Total.

—= il 1
1 3 4
1 1 2
A. 9
1 1 2
3 5 8

— ij 1
1 3 4
2 1 3

= ® 2
2 2 tl
——— 2

1 — il
2
1
1

[| paw rmwernmar

to e

_
PoPHoaRee pe | onman!] Haw nwmweePeno me
MAWOWHDAUMUNHOWHOWR SO

paeuicas|| (conemraal! oupores <a et oolmed

A. terrae-

| reginae 1.

Oy
+0
Total.

ry th
@ lil
10 15 25
3.3 6
2 4 6
2 ee Ss
iL By 1G
13 4

A. alfalfae

A. histri-
onicula 3.

Cy Ae
13 4

A. malvae

| A. merred-

inensis 5.

A. ban-
crofti 6.

bi 8
— 2 2

A. brac-
tigera 7.

1—

Total.

BY HAROLD F. LOWER. 211

TABLE 1.—Continued.

Showing Numbers of Specimens Studied and Districts where These were Collected.—Continued.

| A. terrae- | A. alfalfae| A. histri- | A. malvae | A.merred-| <A. ban- A. brac-
{

A. viridigrisea. | reginae 1. 2% onicula 3. 4. inensis 5. | crofti 6. tigera 7.
Core We, a4 SOS | gO Bg oS Se 2s Ores Ors
sal = sal ial cal | [= ial a)
Victoria—
Nathalia .. — 5 B ) a ee eee Soe pa yee del a
South Aus-
tralia—
Athelstone.. | 1000 1000 2000 | — — — | — — — | — — — |} — — — | —
Berri bee 2a 23, — = st
Marion a Tal. Ky. AD) =
Mitcham .. 18 25 43 — =
Renmark .. 6 eee ARS me eg [SS a ak =
Western Aus-
tralia— |
Bridgetown 2 2 4 a= —# | —i| ae
Kenwick .. 5 11 16 | — — — |] — — —] — — — | — — — | — — — | — — — | — — —
Manjimup 2 3 er rapa
Merredin .. — oS il 1) 19?
Perth 6 ea plain fie a RS a eS FS a, IP pe ts
Wyndham | 12 14 26 | — — —— — —._— — — | — — —
‘Tasmania— |
No species
known

Grand totals | 1174 1239 2413 | 27 40 67 | 20 32 52 o 8) 8 6 8 14 | 10 17 27 1 4 5 l= i

Key to genus Austroasca nov.

m-cu connecting M,,, with Cu,; R in normal position ...... —........ subgenus Austroasca
m-cu connecting R + M with Cu, (Text-fig. 13); R abnormally close to C .... subgenus Paolia

Subgenus Austroasca nov. Males.

i. Crown conspicuously marked with brownish-black ............... 00 cee eee eee ee eee Z
Crownenot marked) with) brownish=placks 2c - > secre ee dele cla cists e eiciel cle eemsleneu stele + s)le) = 3

2. (1) Four brown marks on crown in form of separate longitudinal bands touching anterior
MATL Ine CREXt= fis BO) Meee wusescuaveia sre chalebsescsue ea-o saysea suc isaee eal whee malvae Evans (1942b)

Two brown marks on crown either roughly U-shaped or subquadrangular, not touching
anterior margin (Text-figs. 38, 38A) ................ histrionicula Kirkaldy (1906)

SG) parachone swith) wide bases @Rext igs 2s Gopi stele cieirsiercciaselsiencbelclral el aliercrereie) -lieiehe “iekslcneaiel 4
Ga Chone: mwalthiom arr Owe. WW alSe i cise sseespesseeens sifei aio ok sire noueuteeneye: elietousireyieis ausiehs re selepellsiemens mel sive tacts eee ease 5

4. (3) Termination of brachone long, slender and much recurved (Text-fig. 63) ..............

2 brG2O0GHO G6 See IONE AID OR ERO SIG (CRORE OOHRS, HiesicDepeer career CnC erC arc eeh anaes merredinensis, sp. nov.

BRP sah cu ci sacs anes cyan ea RR SLR AUC iSua) OORT cal SLasasrar cece asaya rel ea iceatien ation eusiachaey Sas viridigrisea Paoli (1936)

5. (3) Brachones expanding to form broad plates, toothed on distal margins (Text-fig. 72)

70:10 -018 (0.650 Bud Ocp/ OPO BHOTD OG BiG TUIO Ort O NORGE Daca aa Seer A ncReS ae ZO SO RORD REE CREO NnaCecciLc Esean bractigera, sp. nov.
IS KAChHOnemiSlEMG Ore caicw wa tsy weptay ies hosed ste: <a salowra cae Pome ey wuedlen mas barral chertcumtara cust cmsiauah eos ere meee: amen erOreoret eas 6

6. (5) Brachone strongly angled, tip deeply notched (Text-fig. 49) ...... alfaifae Evans (1941)

Brachone almost straight (Text-fig. 23) ; tip sometimes concavely notched (Text-fig. 23A)
0 Ot BGO os CROTON DCL CACTI CRER ONSEN IG: OIC SEE EEL OEM CeO Canter nciaecs terrae-reginae Paoli (1936)

Subgenus Austroasca nov. Females.

1. Crown conspicuously marked with brownish-black ............... cc eee eee eee eee eee 2
Chroma maar manne! wrlel lomonwnMiM ele sobodacancdoc0d0b00d coon ODDO Oooo oO KUO OOOO 3

2. (1) Four brown marks on crown in form of separate longitudinal bands touching anterior
maeyeysinn (avers, BON soonsccogdcooo oud sD ago uDODOLOOUObCUE malvae Evans (1942b)

Two brown marks on crown either roughly U-shaped or subquadrangular, not touching
anterior margin (Text-fig. 38, 38A) ................. histrionicula Kirkaldy (1906)

2 Cl) ChesSAyEiin, “Tarn TOFOREME Fhe WEYSBONEN GoopoodceoudccoO dbo dOUOuKD OOOO COUODO Oooo ddDODOSD 4
Gross=veinerem abSentein teSMeMe. 95. eraie-ap slots. oh eotaptemetthods: ©. eoheic|s oops lyls okey ghen ous ecalle-e Scsnsteye « 6

4. (3) Tegmen with a small brown spot in cell Cu, near forking of Cu, (Text-fig. 58) ........

See eA STE TT TE eee nea ee ay ar EAs foo BG TE a Scale sal ehaoreiabiar Sehig) uae MARCO eoeeaTos terrae-reginae Paoli (1936)

212 AUSTRALIAN SPECIES PREVIOUSLY REFERRED TO EMPOASCA,

5. (4) Crown distinctly, obovately produced C.I. = 23 (Text-fig. 45)
Crown very little produced C.I. = 10 (Text-fig. 68)
6. (3) R,, directed towards base of tegmen (Text-fig. 67).

Subgenus Paolia nov.

. alfalfae Evans (1941)
bractigera, sp. nov.
merredinensis, sp. nov.
Greenish or greyish insects
viridigrisea Paoli (1936)

Head yellow with three oval black marks, two on crown, and a median one high on face

TABLE 2.

A.

Recorded Host Plants of Species of Austroasca.

(Paolia) bancrofti Evans (1939)

Species.

viridigrisea

merredinensis

malvae ..

histrionicula

Breeding and Feeding on.

Potato.

Tomato.

Lucerne.

French bean.

Rock melon.

Solanum nigrum L.
Chenopodium album L.
Malva parviflora L.

| Amaranthus viridis L.

Citrullus vulgaris Schrad.
Tobacco.

Silver beet.*

Cotton.*

Trianthema portulacastrum L.*
T. decandra L.*

Ricinis communis L.*

Atriplex sp.

Malva parviflora L.*
Abutilon sp.

Sida subspicata F.v. M.
S. rhombifolia L.*

alfalfae ..

terrae-reginae

bancrofti

bractigera

Feeding Only.

Cynodon dactylon Richard.
Celery.

Portulacca oleracea L.
Bidens pilosa L.*
Tribulus terrestris L.*

Cotton.*

Poona pea (Vigna sinensis L.).*
Pisum sativum L.*°

Lucerne.*

Cotton.*

Crotalaria sp.*

French bean.

Cotton.
Sida corrugata L.*
Abutilon sp.*

No hosts recorded.

No hosts known.

* Information from May, 1950.

Potato.*

Amaranthus viridis L.*
Chenopodium carinatum R.Br.*
Cleradendron tomentosum R.Br.*

No hosts recorded.

No hosts known.

AUSTROASCA (AUSTROASCA) VIRIDIGRISEA PAOLI (1936).
(Text-figures 1-11, 24-29.)

Empoasca viridigrisea Paoli, Mem. Soc. Ent. Ital., 15, 1936: 12-13.
Empoasca terrae-reginae Evans, Proc. Roy. Soc. Queensland, 52, 1941: 11.
Length.—Male: Average of 1,174
specimens, 3-8 mm.; max. 4:0 mm., min. 355 mm. Female: Average of 1,239 specimens,
3-9 mm.; max.4:2 mm., min. 3-8 mm. Colour: Green with white markings.

Suggested common name, the vegetable jassid.

Head.—Crown: C.l. = 13 (Text-fig. 1).

Anterior margin gently curved. Emerald

green with narrow, irregular median, white stripe, four lateral white spots, one on

BY HAROLD F. LOWER. 213

anterior margin close to each eye, and one posterior to each of these. Coronal suture
incomplete. Face: Shape typical (Text-fig. 2). Yellowish-green with white, irregular,
narrow, median white stripe from vertex to level of antenna bases. The pattern varies
considerably, some specimens showing more white than others. Two white bands, one
from below each ocellus, and directed outwards. Antennae typical, scape and pedicel
green, third segment and remainder of antenna dark green (Text-fig. 3). Ocelli two,
high on face, one between median and each eye, each in centre of yellowish-white,
circular spot. Hyes dark brown.

Thorax.—Pronotum (Text-fig. 1): Anteriorly yellowish-green becoming translucent
and paler posteriorly; an irregular median white spot, and four lateral white spots, two
on each side. All spots on anterior margin. Scutellum (Text-fig. 1): Emerald green,
yellowish laterally. A complex, characteristic, white median mark (Text-fig. 1A) and a
iong, white, irregular stripe along each lateral margin. The median mark varies little
in shape; in a few specimens it is not forked.

Wings.—Tegmen (Text-fig. 4): Basal four-fifths yellowish and opaque, almost con-
cealing venation, which is typical. Apical fifth brownish to colourless hyaline. Veins
' green and obvious in clear apical area. Hind Wing (Text-fig. 5) typical, colourless
hyaline, veins prominent and green. JLegs typical, green, tending to peacock blue
distally. Pretarsi dark brown. ;

Abdomen.—Typical of genus. Dorsal surface green, ventral surface yellow to
yellowish-green. Genitalia deep green (Text-figs. 6 and 7). Subgenital plate (Text-fig.
26) short and broad, with about fifteen stout ensiform bristles and about fourteen
long thin hairs on dorsal edge near base. Harpagone (Text-figs. 8 and 28): About eight
teeth and four stout bristles. Brachone (Text-fig. 27) subtriangular with broad base,
rapidly tapering to a blunt point; termination curved. Aedeagus (Text-fig. 29): Free
dorsal part curved in an open are.

Type in British Museum.

Type locality—Bowen, near Pomodori, Queensland, Australia, October 20, 1931
(D. O. Atherton).

Host plants.—See Table 2.

AUSTROASCA (AUSTROASCA) MERREDINENSIS, Sp. NOV.
(Text-figures 59-67.)
Length—Male: Average of 10 specimens, 3-9 mm.; max. 4:0 mm., min. 3:9 mm.
Female: Average of 17 specimens, 4-4 mm.; max. 4-5 mm., min. 4:3 mm. Colour: General
colour light brownish, showing no pattern. (Specimens have been preserved in alcohol.)

Head.— Crown: C.1. = 27. Anterior margin bluntly angularly produced, brownish
(Text-fig. 59). Face typical, brownish. Antennae typical, brownish. Ocelli high on face.
Eyes bleached to white by alcohol.

Thorax.—Pronotum: Anterior margin bluntly angled so as to be more or less parallel
with anterior margin of crown (Text-fig. 59). Scutellum (Text-fig. 59) brownish.

Wings.—Tegmen (Text-fig. 67) light brown to colourless hyaline. Venation typical.
Hind Wing colourless, venation typical. Legs brownish, typical.

Abdomen brownish, typical. Genitalia (Text-figs. 60 and 61). Swbgenital plate
(Text-fig. 66) short and broad. About 20 strong ensiform bristles; lower margin with
a fringe of about twelve thinnish bristles about half the length of the strongest central
ensiform bristle. Apical two-thirds of upper margin with marginal setae. Apex of
plate terminating in a concavity. Harpagone (Text-fig. 64) short and stout, apical part
curved with about eight or nine denticulations. Setae in two rows: an outer row of
three strong bristles and an inner row of three or four bristles not so long as outer
bristles. Brachone (Text-fig. 63) wide at base, rapidly tapering to a long, thin, very
recurved tip, terminating in a sharp point. Aedaegus (Text-fig. 65): Free dorsal part
curved in an open arc.

Type material.—Male holotype; tegmen and dissected genitalia mounted; remainder
of body in alcohol.

214 AUSTRALIAN SPECIES PREVIOUSLY REFERRED TO EMPOASCA,

Type material in Coll. Division of Entomology, C.S.1.R.O., Canberra, A.C.T.

Type locality—Merredin, Western Australia, April 3, 1940. “On Saltbush” (C. F. H.
Jenkins).

Host plant.—Atriplex sp.

AUSTROASCA (AUSTROASCA) MALYVAE EVANS.
(Text-figures 30-37.)
Empoasca malvae Evans, Proc. Roy. Soc. Queensland, 54, 1942: 49. (Note: The numbers

3 and 6 in Evans’ figures should be transposed.)

Length.—Male: Average of six specimens, 3 mm.; max. 3:1 mm., min. 2:9 mm.
Female: Average of eight specimens, 3-5 mm.; max. 3-7 mm., min. 3°33 mm. General
Colour greenish-yellow with brown markings.

Head.— Crown: C.1l. = 19 (Text-fig. 30). Pale green with four irregular longitudinal
brown stripes, one bordering each eye. Coronal suture incomplete. Face normal,
yellowish with a broad, median, irregular, longitudinal, white stripe. Antennae normal,
greenish. Ocelli close to eyes, one in each lateral pale green band on lower part of
corono-facial angle. Eyes brown.

Thorax.—Pronotum (Text-fig. 30) generally light brown with median greenish-white
mark immediately posterior to median pale green band of crown and a white mark near
each antero-lateral margin. Between each of these and the median mark is a sub-oval
white mark. Posterior median section translucent and greenish yellow; posterior lateral
lobes pale green. Scutellum (Text-fig. 30) yellowish-brown with a median white oval
spot continued posteriorly as a median greenish-white stripe to suture, where it expands
to a large sub-oval white blotch. On either side of median white blotch on anterior
of scutellum is a dark band. Tip of scutellum pale green.

Wings.—Tegmen (Text-fig. 37): Apical third brown hyaline, a wide greenish area
along costal margin between that margin and Cu, Area posterior to Cu, largely
brownish-yellow except the two whitish translucent areas shown. Area between Cu,
and Cu, chiefly whitish translucent. Veins greenish-white. Hind Wing normal. Legs
normal, green.

Abdomen normal greenish. Most of posterior of each tergite occupied by a large
brown area, the anterior being yellowish-green. Posterior of ninth tergite with an
almost black transverse band. Ventral surface yellowish-green. Genitalia pale green
(Text-figs. 31 and 32). Swbgenital plate (Text-fig. 34) long, broad at base and gently
tapering. About 25 strong ensiform bristles.. Near centre of plate are three flagellate
bristles, while a few shorter such bristles occur above these near the upper margin.
Terminal fifth of upper margin with marginal setae. Near base of plate and on upper
margin is a group of long hairs followed by six to seven short ensiform-like bristles.
Harpagone (Text-fig. 36) short and stout with between five and six denticulations and
four bristles. Brachone (Text-fig. 35) short, slender, curved and simple, with an upward-
directed spur near base. Aedeagus (Text-fig. 33): Free dorsal part in form of a thick
erescent.

Type.—Male mounted on point, in Coll. Queensland Museum. H0O5233.

Type locality —Gayndah, Queensland, April 7, 1942. “On Sida subspicata” (A. May).

AUSTROASCA (AUSTROASCA) HISTRIONICULA KIRKALDY (1906).
(Text-figures 38-44.)
Cicadula histrionicula Kirkaldy, Haw. 8.P. Exp. Sta. Entom. Bull., 1, part 9: 361.
Empoasca histrionicula Myers, Bull. Ent. Res., 18, 1928: 311.
Empoasca pulcherrima Evans, Proc. Roy. Soc. Queensland, 54, 1942: 49. (Note: The
numbers 3 and 6 in Evans’ figures should be transposed.)

Length.—Male: Average of three specimens, 2°7 mm.; max. 2°75 mm., min. 2:7 mm.
Female: Average of five specimens, 2-8 mm.; max. 2:9 mm., min. 2‘7 mm. Colour
yellowish-green with conspicuous brown markings.

Head.—Crown: C.1. = 17 (Text-fig. 88). Slightly produced, light green, most of
its area occupied by two large U-shaped almost contiguous dark brown marks. These

BY HAROLD F. LOWER. 215

marks vary considerably in shape. Frequently they are U-shaped, the arm of the U
nearer the median line being longer, but in many specimens the colour “overflows”
so as to make the marks roughly subquadrangular, as is the case in Kirkaldy’s type
(Text-fig. 38A). Face typical. Fronto-clypeus lemon yellow, ante-clypeus and labrum
light green. Antennae green. Ocelli high on face, each nearer the eye than median line.
Eyes dark brown.

Thorax.—Pronotum (Text-fig. 38) with a broad, median, velvety, brown band;
laterally, pale green. Scutellum (Text-fig. 38) wholly dark brown.

Wings—Tegmen normal, pale yellowish-green with large dark brown area (Text-
fig. 44). - Hind Wing normal. Legs normal, yellowish-green.

Abdomen.—Normal, yellowish in colour, a large.subtriangular brown area covering
most of the terga. Tergum of ninth segment brownish-black, a narrow yellow lateral
band on posterior of eighth tergite intervening between the two brown areas. Sternal
surface entirely yellow. Anal tube wholly dark brown. Genitalia (Text-figs. 39 and 40):
In both sexes the basal half is yellowish, the terminal half brown, intensifying to black
at the tips (Text-fig. 43). Swbgenital plate (Text-fig. 43) long, basal third wide, apical
two-thirds tapered with upper and lower margins subparallel; nine or ten ensiform
bristles present. Terminal half of upper margin with marginal setae. On upper margin
near base are three or four small hairs followed by four or five small ensiform-like
bristles and three or four flagellate type bristles. Harpagone (Text-fig. 41): Basal third
wide, apical two-thirds slender and almost straight. Nine to ten denticulations, four or
five small slender bristles in two series. Brachone (Text-fig. 42) strongly elbowed at
base, then straight, more or less cylindrical, tapering suddenly to a point, twisted.
Aedeagus (a, Text-fig. 39): Free dorsal part in form of a thick crescent.

Type—Female mounted on point. In Coll. Haw. Sug. Planters’ Assn., Honolulu,
Hawaii.

Type locality—Bundaberg, Queensland. Collected by Koebele and Perkins’
expedition in 1904.

AUSTROASCA (AUSTROASCA) BRACTIGERA, SD. NOV.
(Text-figures 68-74.)
Length—Holotype male, 3-6 mm. (only specimen Known). Colour yellow.

Head.—Crown (Text-fig. 68) extremely slightly produced; C.I. = 10. Bright yellow
with faint white stripe on coronal suture, which is incomplete. Face normal and
yellowish. Antennae normal, brownish-yellow, flagellum brown. Ocelli two, close to
eyes, and level with top of post-frontal suture. Hyes brownish-black.

Thoraz.—Pronotum (Text-fig. 68) robust; yellow possibly marked with white on
anterior margin. Scutellum yellow with apparently two white lateral bands and a
terminal white patch.

Wings—Tegmen (Text-fig. 74) normal, brownish-yellow and hyaline. Cross vein
r-m present, Ria weakly developed; near base in cell Se is a series of five circular
protuberances, and two elongated raised patches formed by the union of a number of
Similar protuberances. Hind Wing colourless; venation normal, veins white. Legs
normal, yellow.

Abdomen.—Yellow dorsally and ventrally, white laterally. Genitalia (Text-figs. 69,
70) yellow. Subgenital plate (Text-fig. 71) long, the distal half very narrow with dorsal
and ventral margins subparaliel. Ensiform bristles in two series; near apex are three
or four well-developed flagellate bristles; marginal setae also present. Harpagone
(Text-fig. 73) short and robust; about thirteen denticulations terminating in a curved
spine; four to five bristles; base rugged. Brachone (Text-fig. 72): In its shape the
brachone of this species is quite unique. It is short and from a narrow base opens
out into a broad flat plate with its apical margin notched like a saw. The brachone,
alone, will at once identify this species. Aedeagus (a, Text-fig. 69): The aedeagus is
relatively large. Its attached basal part is narrow but soon widens and gives off a
second strong point of attachment. The two free lobes are curved in an open arc.

216 AUSTRALIAN SPECIES PREVIOUSLY REFERRED TO EMPOASCA,

Type.—Holotype male, mounted on card; genitalia separately mounted. In Coll.
Department of Agriculture, Sydney, New South Wales.

Type locality—Only one specimen, the type, is known. This came from Mt.
Keira, New South Wales, and bears the label “Mt. Keira, New South Wales, at light
3.12.1949. C. E. Chadwick”.

AUSTROASCA (AUSTROASCA) ALFALFAE EVANS.
(Text-figures 45-52.)

Empoasca alfalfae Evans, Proc. Roy. Soc. Queensland, 52, 1941: 11.

Length.—Male: Average of 20 specimens, 2:75 mm.; max. 2°85 mm., min., 2:7 mm.
Female: Average of 32 specimens, 3:6 mm.; max. 3:8 mm., min. 3-4 mm. Colour yellowish-
green.

Head.—Crown: C.1. = 23. Obovately produced, yellowish-green (Text-fig. 45). Face
normal, yellowish-green. Antennae normal, green. Ocelli two, one much nearer each
eye than median line, near upper limit of face. Hyes dark brown.

Thoraz—Pronotum (Text-fig. 45) yellowish-green apparently unmarked with white.
Scutellum yellowish-green with indefinite white markings, green towards lateral margins.

Wings.—Tegmen (Text-fig. 52): Venation normal; the erect vein Ry is a charac-
teristic of this and the next species. Cross-vein r-m present. Basal two-thirds of tegmen
thickened, yellowish-green and translucent; apical third colourless hyaline. Hind Wing
normal, veins pale green. Legs normal, yellowish-green.

Abdomen.—Normal, yellowish-green darkening towards posterior end. Genitalia
(Text-figs. 46, 47) green. Suwbgenital plate (Text-fig. 51) relatively short, its upper and
lower margins subparallel. About fifteen ensiform bristles, the uppermost near the base
much longer and stronger than the others. Apical half of upper margin with marginal
setae. A series of six or seven hairs on upper margin near base. Scattered over the
“body” of the plate are long flagellate hairs which are very abundant towards its lower
apical region. Harpagone (Text-fig. 50) short and stout, terminating in a spine-like
point, with three or four denticulations, and with three short stout bristles. Brachone
(Text-fig. 49) slender and strongly angled upwards about half-way along its length,
where it is narrowest. It then widens for half the remainder of its length and is then
notched strongly, terminating in a long, stout-pointed portion. Aedeagus: Free dorsal
lobes somewhat ovate. Dorsal hook (Text-fig. 48 and ds, fig. 46): an extremely small
dorsal hock can be seen under a magnification of 1,000.

Type.—Male, mounted on point in Coll. Queensland Museum, Ho. 5225.

Type locality.—Lockyer, Queensland, Australia, November 6, 1939 (D. O. Atherton),
“On lucerne”.

AUSTROASCA (AUSTROASCA) TERRAE-REGINAE PAOLI.
(Text-figures 538-58 and 22-25.)

Empoasca terrae-reginae Paoli, Mem. Soc. Ent. Ital., 15, 1936: 13-14.
Empoasca maculata Evans, Pap. Proc. Roy. Soc. Tasmania, 1942: 27.

Suggested common name, the cotton jassid.

Length.—Male: Average of 27 specimens, 3:3 mm.; max. 3:6 mm., min. 3:0 mm.
Female: Average of 40 specimens, 4:0 mm.; max. 4:2 mm., min. 3:9 mm. Colour yellow
with white markings.

Head.—Crown (Text-fig. 53) distinctly produced in a blunt angle. C.I. = 27. Yellow
with median, longitudinal, narrow, irregular white stripe, and two sub-oval lateral
white marks, midway between eyes and coronal suture. Face normal, yellow with
broad, white, median, longitudinal stripe and two white stripes from middle of upper
part of face directed diagonally towards lower margin of each eye. In many specimens
the three stripes are united above to form a broad arrow. Frontal and epicranial
sutures obvious. Antennae normal, brownish-yellow. Ocelli two, one very close to each
eye on upper part of face. Eyes black.

Thorax.—Pronotum (Text-fig. 53) long, about twice crown length, yellow, with five
white marks close to anterior margin, one median and two laterally. These five marks

BY HAROLD F. LOWER. 217

vary greatly in size and shape but their number and position are constant. Scutellum
(Text-fig. 53) yellow with a large subrectangular median white blotch between suture
and anterior margin of scutellum, lateral margins white. Posterior to suture is a small
elongate median white blotch; the two anterior lateral parts of this area are white.

Wings.—Tegmen (Text-fig. 58) normal, colourless hyaline with a small brown
irregular spot in cell Cu, near the forking of Cu,». The colour of the spot varies from
dark brown to yellowish and its shape is very variable. Its area and position are
constant. In cell Sc and near the base of the tegmen is a line of small, circular,
‘colourless protuberances varying in number between twelve and sixteen. Posterior to
these is-a small raised area formed by the union of from seven to nine such protuber-
ances. A larger protuberance commonly occurs in cell M and a smaller one posterior to
it in cell Cu,. Cross-vein r-m present; R, perpendicular to costal margin. Hind Wing
normal, colourless, veins white. Legs normal, yello v.

Abdomen.—Normal, dorsally yellow, ventrally yellowish-white. Genitalia (Text-figs.
54, 55) yellow, highly characteristic. Subgenital plate (Text-fig. 57) extremely long and
narrow, its length being approximately ten times its greatest width. About twelve
well-developed ensiform bristles. Near its apex ventrally is a number of extremely long
flagellate bristles, while a row of similar but somewhat shorter bristles occur along the
terminal half of the upper margin, which bears marginal setae in the same region.
Towards the base and on and near the upper margin is a group of long thin hairs.
The flagellate bristles are so long as to give the plates a hairy appearance even to the
naked eye. Harpagone (Text-fig. 24) short and stout, terminating in a spine; seven to
eight denticulations; three short stout bristles. Brachone (Text-fig. 23) extremely long
and slender, slightly angled about one-quarter from its base, then straight with a
small upcurved tip. Many specimens show a concave notching at the tip (Text-fig. 23A).
Aedeagus (Text-fig. 56). Free dorsal part curved in an open arc.

Type.—Genitalia dissected and mounted separately, in Coll. British Museum.

Type locality.—Biloela, Queensland, Australia, January 4, 1932 (D. O. Atherton),
“From cotton”.

AUSTROASCA (PAOLIA) BANCROFTI EVANS.
(Text-figures 12-19.)

EHmpoasca bancrofti Evans, Pap. Proc. Roy. Soc. Tasmania, 1938: 40.

Length.—Male: One specimens (the type), 4 mm. (Evans’ measurement). Female:
Two specimens, 5:1 mm., 5-2 mm. (two specimens have the abdomens missing but are
adjudged as females on other evidence). Colour yellowish with brown markings.

Head.—Crown (Text-fig. 12): C.I. = 12-13. Bright yellow with two oval blackish
spots, one posterior to each ocellus. Coronal suture complete. Face normal in shape,
but differs from all other species in having the coronal, frontal and epicranial sutures
all complete and well defined; bright yellow with median, oval, black spot just above
level of antenna bases. Antennae normal; scape and pedicel yellow; third segment and
flagellum brown. Ocelli two, orange in colour, one between each eye and median line.
Eyes black.

Thorax.—Pronotum (Text-fig. 12) yellowish with large, median, transverse, crescentic
brown area, the whole finely white pollinose. Scutellum (Text-fig. 12) yellow medially
with two lateral brown areas and a median apical brown spot.

Wings.—Tegmen (Text-fig. 13): Venation unique; R runs parallel to and very close
to C + Sc, uniting with the latter about half-way along the costal margin. It then
turns sharply posteriorly and after a bend unites with M, the combined vein R+M
continuing on, giving off the branch R,: and eventually forking into R, and M3;,., thereby
eliminating the cross-vein r-m. The cross-vein m-cu, by moving basad, has lengthened,
and its anterior termination meets Ri» and Mz,, at the one point. Along the costal
margin is a blue-black band which fills the whole of the area between C and M from
the base of the tegmen to the point where R combines with M. A similar blue-black
area extends along the posterior margin to the union of Cu, with that margin. Scattered
over the basal half of the tegmen is a number of small circular protuberances, colourless

218 AUSTRALIAN SPECIES PREVIOUSLY REFERRED TO EMPOASCA,

in the colourless part and black when these occur in the blackened areas. The remainder
of the tegmen is milky white and translucent. Hind Wing not studied in detail for
reasons given above. Milky white and translucent, veins white. Legs normal, yellowish.

Abdomen.—Normal; dorsal surface yellow with median row of brown spots extending
along its whole length, two similar rows of lateral brown spots, one on each side of
each tergum; ninth tergum and anal tube entirely brown. Genitalia yellow (Text-figs.
14, 15). Female genitalia as in genus; male genitalia abnormal. Text-fig. 14 has been
reconstructed as explained above. Pygophore appears to be quite abnormal, there being
no true pygophore as generally understood. This is possibly due to defects in preparation
of the material. Subgenital plate (Text-fig. 18) short and tapering. Near lower basal
margin are five ensiform bristles; after a gap there follow twelve small ensiform
bristles and six small bristle-like hairs near the apex; marginal setae, flagellate bristles
and fine hairs appear to be missing. Harpagone (Text-fig. 16) large, stout, twisted, but
apparently of a simple type; the end is broadly rounded with five blunt teeth; there is
no sign of bristles. Brachone (Text-fig. 17) long, slender, curved at the tip. Dorsal
hook (Text-fig. 19), although much smaller than in other genera of the Empoasciti, is
larger than in any other species of Austroasca s.lat. Its tip bears a V-shaped ridge and it
is strongly sclerotized. Aedeagus has been torn in the type genitalia and its shape
cannot be made out.

Type material—Holotype male mounted on pin. Genitalia and tegmen mounted
separately. In Coll. Division of Entomology, C.S.I.R.O., Canberra, A.C.T.

Type locality—Hidsvold, Queensland, Australia (T. Bancroft). In view of the
paucity of material (I have not yet seen a complete male) the status of this species.
must be somewhat conjectural. The complete sutures of the head and the abnormal
tegmen venation are features common to the five specimens examined, but no certainty
can be arrived at until I have thoroughly examined and dissected a complete male. Only
in this way will it be possible to determine the exact nature of the parts of the genitalia
and their spatial relationship. When I have had the opportunity of so doing, certain
of the above statements may need modification.

APPENDIX.
Translations of Paoli’s descriptions of A. viridigrisea and A. terrae-reginae.
A. VIRIDIGRISEA.

“Colour of body (when dry) greyish green, with almost colourless tegmina; eyes
very pale.

Head with vertex rounded in the male (in the female?), its length not much more.
than half the width between the eyes (7:12), with its posterior margin somewhat
incurved.

Tenth abdominal segment rather short, the ventral surface with a sprinkling of
very short spines united into comb-like groups; processes of the anal tube [dorsal
hooks] very short in the form of small protuberances, weakly sclerotized.

Aedeagus strongly curved in the free terminal part.

Genital laminae [subgenital plates] short and broad with ne ensiform bristles
moderately dispersed; on the upper margin is a group of rather numerous (about 12).
long bristles, the remainder bearing short bristles; sternal surface with a fringe of
medium length bristles about half as long as the width of the lamina.

Upper styli [brachones] well protected, concealed, wide at the base, then rapidly
narrowing and curving, sharp at the apices.

The lower styli [harpagones] wide at the base, then gradually tapering and recurved
with 4-5 lateral bristles and 8-9 teeth at each apex.

The abdominal apodemes appear to be missing, since, in the one preparation that.
I was able to make, and that comprising almost the entire abdomen, these could not be
traced.

Body length of the male (when dry) mm. 1-9-2, including the wings 2-8-2-9.*

* These measurements are much smaller than in any specimen I have seen.—Author.

BY HAROLD F. LOWER. 215

Habitat: Bowen, Queensland (Australia) near Pomodori. (Sent by D. O. Atherton.)
I have seen three males in the British Museum Collection. These are labelled:
Bowen, Q. 20.10.31, D.O.A.
Queensland. R. Veitch.
Host: Tomato, D. O. Atherton.

Type in British Museum.

Observations: This species, by the reduction of the upper styli and the probable lack
of abdominal apodemes is most closely allied to H. decedens Paoli, but the aedeagus is
not sclerotized nor curved at an angle at the tip.”

A. TERRAE-REGINAE.

“Body wholly pale yellow (when dry) with colourless tegmina; eyes almost black,
ocelli brownish.

Head anteriorly rounded in the male, slightly more produced in the female; the
length of the vertex is a little more than half the width between the eyes. The long
pronotum is one and a half times the length of the head with its posterior margin a
little incurved.

The tenth abdominal segment has its ventral surface uniformly covered with spines
for the whole of its length; the processes of the anal tube [dorsal hooks] are short,
bent into straight teeth incurved at the apices. The free slender part of the aedeagus
is curved in an arc.

Laminae [sub-genital plates] long, straight, with the greatest width near the basal
part, and narrowing towards the apices; the ensiform bristles are of two kinds; those
near the apices are fine, about 4-5 times as long as the width of the lamina, and
terminate in flagella; towards the base they assume the normal form, but are always
fairly long; both kinds overlap in the medial region; the upper margin bears rather
slender bristles but these are shorter than those of the lower margin; in the basal
half, marginal setae are lacking on the upper margin. The sternal surface bears short
setae situated almost in the middle of the distal half; these become thin and very
long on the basal half.

The elongated upper genital styli [brachones] are almost straight with a slight
curve very near their apices; sometimes, at the curve the style shows a slight concavity.

The lower genital styli [harpagones] gradually taper with 2-3 big bristles at the
distal half and 7-8 teeth near the tip, which is prolonged into a sharp point.

Abdominal apodemes not seen through scarcity of material.

Length of body of male mm. 2:3; including the wings mm. 2°8; of the female 2:1—2:3;
including the wings 2-8—2:9.
Habitat: Biloela, Queensland (Australia), from D. O. Atherton. I have seen 2¢
and 29 collected from cotton, 4th January, 1932 (British Museum). These are labelled:
Biloela,
4.1.32,
From Cotton,
D. O. Atherton,
Queensland.

Type in British Museum.

Observations: In some characters, such as size and pale coloration, this species
shows affinities with H. facialis Jac., but more particularly through the long straight
laminae bearing abundant and long bristles. The scarcity of material has prevented
my examining other features such as the presence and shape of the abdominal apodemes
but the characters indicated above are so specialized that they are sufficient for the
precise determination of the species.” *

* The italics are mine.—Author.

220 AUSTRALIAN SPECIES PREVIOUSLY REFERRED TO EMPOASCA,

Acknowledgements.

The satisfactory completion of this piece of research has been made possible only
through the co-operation of a large number of scientific workers, and to the following,
I return my grateful thanks:

To Dr. Dwight M. De Long, Professor of Entomology, Ohio State University, U.S.A.,
for his sympathy, guidance, and encouragement, as well as his supplying me with
North American material for comparative morphological work. AM) IDyrs “bb tek, (Ch
Taylor, of the British Museum, and, through him, Dr. W. HE. China and Mr. N. C. E.
Miller, for comparison of specimens with types, for information, and for sketches of
genitalia of Paoli’s types of Hmpoasca viridigrisea and E. terrae-reginde. To Mr.
Cc. E. Pemberton, Entomologist of the Hawaiian Sugar Planters’ Association at Honolulu,
who was indefatigable in his checking of specimens with Kirkaldy’s type, and, even
more so, in his replying promptly and at length to my series of voluminous letters
requesting drawings, descriptions, etc.

To the following for the loan of material in their collections and general informa-
tion: Mr. G. Mack, Director of the Queensland Museum (especially for the loan of
Evans’ types of Hmpoasca athertoni, E. maculata, E. pulcherrima, HE. malvae and
E. alfalfae); Dr. A. B. Walkom, Director, and Mr. A. Musgrave, Curator of Entomology,
of the Australian Museum; Mr. T. G. Campbell, Research Officer, Division of Entomology,
C.S.1.R.0O., Canberra (especially for the loan of Evans’ type of Empoasca bancrofti) ;
Dr. Joseph Pearson, Director of the Tasmanian Museum; Messrs. H. Womersley,
Entomologist, and G. F. Gross, Assistant Entomologist, of the South Australian
Museum; Mr. L. Glauert, Curator of the Western Australian Museum; Dr. W. A.
McDougall, Senior Entomologist, Department of Agriculture and Stock, Queensland,
and, through him, Mr. W. A. Smith, Entomologist of the Ayr branch; Mr. T. McCarthy,
Chief Entomologist, Department of Agriculture, New South Wales; Mr. C. J. R. Johnston,
Entomologist, Department of Agriculture, Victoria; Mr. L. W. Miller, Senior Ento-
mologist, Department of Agriculture, Tasmania; Mr. C. F. H. Jenkins, Government
Entomologist, Department of Agriculture, Western Australia.

Finally, my thanks are due to the field-workers of the various Australian State
Departments of Agriculture, and to the collectors for the various museums, without
whose activities the large quantity of material investigated would never have been
assembled.

I wish to thank Miss Margaret EH. Morphett, who has done the lettering and
numbering of the figures. (In fairness to her, I must point out that I alone am
responsible for the headings of the figures of A. bancrofti and A. malvae and the
lettering on Text-fig. 37.)

References.
ANON., 1944-1945.—Entomological Investigations, 19th Ann. Rep. O.S.I.R., Canberra, A.C.T.,
p. 29. :
De Lone, D. M., 1931.—A Revision of the American Species of Empoasca known to occur
North of Mexico. U.S.D.A. Tech. Bull., No. 231. i
Evans, J. W., 1939.—Australian Leaf Hoppers (Homoptera, Jassoidea). Pap. Proc. Roy. Soc.
Tasmania, pp. 39-40.
, 1941.—Some Queensland Leaf Hoppers (Jassoidea, Homoptera) that Attack Lucerne.
Proc. Roy. Soc. Queensland, 52: 11.
, 1942a.—New Leaf Hoppers from Tasmania and Queensland. Pap. Proc. Roy. Soc.
Tasmania, p. 27.

, 1942b.—Some New Leaf Hoppers from Australia and Fiji. Proc. Roy. Soc. Queens-
land, 54: 49.
, 1946.—A Natural Classification of Leaf Hoppers (Jassoidea, Homoptera). Part I.
External Morphology and Systematic Position. Trans. Roy. Ent. Soc. Lond., 96: 47-60.
, 1947.—A Natural Classification of Leaf Hoppers. (Jassoidea, Homoptera). Part III.
Jassidae. Trans. Roy. Ent. Soc. Lond., 98: 105-271.
KiRKALDY, G. W., 1906.—Leaf Hoppers and their Natural Enemies. Haw. S.P. Exp. Sta. Entom.
Bull., 1 (€9) =: 361.
LeacH, J. G., 1940.—Insect Transmission of Plant Diseases. New York, p.° 285.
May, A. W. S., 1950.—The Cotton Jassid Problem in Queensland. Queens. Jnl. Ag. Sci., 7: 25.
Mayr, E., 1942.—Systematics and the Origin of Species. New York, p. 14.

BY HAROLD F. LOWER. 221

Myers, J. G., 1928.—A Note on Australian Typhlocybine Leafhoppers, with a Description of a
New Species. Bull. Hnt. Res., 18: 311.

Paoul, G., 1936.—Descrizione di Alcune Nuove Specie di Hmpoasca (Hemipt. Homopt.) e Osser-
vazioni su Specie Note. Mem. Soc. Hnt. Ital., 15: 12-14.

Poos, F. W., and WHEELER, NANcy M., 1943.—Studies on Host Plants of the Leafhoppers of the
Genus Empoasca. U.S.D.A. Tech. Bull., No. 850.

Smi1TtTH, J. H., 1939.—Rep. Ent. Sect. Ann. Rep. Queensland Dept. Agric. and Stock, p. 34.

SNopGRASS, R. E., 1935.—Principles of Insect Morphology New York.

WALSH, B. D., 1865.—On Certain Remarkable or Exceptional Larvae, Coleopterous, Lepidop-
terous, and Dipterous, with Descriptions of Several New Genera and Species, and of Several
Species Injurious to Vegetation which have already been Published in Agricultural Journals.
Boston Soc. Nat. Hist. Proc., 9: 286-320.

ZAKHVATKIN, A. A., 1946.—Studies on the Homoptera of Turkey I-VII. Tvrains. Roy. Ent. Soc.
Lond., 97: 149.

EXPLANATION OF PLATE XV.

Left.—Portion of stem of black nightshade (Solanum aigzwm), x 0-8. Plant has been
killed by feeding of A. viridigrisea.

Right.—Typical damage done by feeding of A. viridigrisea. 1. Leaf of Amaranthus
viridis. The black specks are the execreta of the insects. 2. Leaf of Malva parviflora (small-
flowered mallow). 3. Leaf of Medicago sativa (lucerne). 4. Leaf of Apiun graveolens
(celery). 5. Leaf of Citrullus vulgaris (pie-melon).

MISCELLANEOUS NOTES ON AUSTRALIAN DIPTERA. XV.
TABANUS, HETEROPSILOPUS.
By G. H. Harpy.

[Read 28th November, 1951.]

Synopsis.

In family Tabanidae two new synonyms are given for Tabainus imperfectus Walker, and
the characters are discussed. In Dolichopodidae, a key is given for the genus Heteropsilopus
and new synonymy is recorded.

Genus TABANUS, Subgen. DoLICHAPHA Enderlein.
Hardy, 1948, Proc. Roy. Soc. Queensland, 59: 169-178.

The subgenus is defined in the above reference, and it covers the Vabanus
circumdatus-complex and the closely allied imperfectws-complex, both of which have
become so involved in literature that no author seems to have a clear understanding
of the components. The typical circumdatus form is light brown, and the typical
imperfectus form is blackish, but varying in the amount of lighter brown which is more
or less confined to the base of the abdomen. Specimens in collections show a wide
range between these colours and, as shown below, the shape of the frontal callus cannot
be considered fully reliable for specific identification.

There seems to be no hope of disentangling the confusion brought about in these
species without field experience, and the collection from each type locality of sufficient
specimens to cover all variations that may occur there, for study whilst in a fresh
condition. Especially is the male required, from which sex has to be established the
specific status in each locality. I anticipate that the species passing under the name
imperfectus in Tasmania, which has no enlargement of facets in the eye of the male,
will be found to differ from the mainland form which apparently has enlarged facets,
but this character may be subspecific in status.

TABANUS IMPERFECTUS Walker.
T. imperfectus Walker, 1848, List Dipt. Brit. Mus., 1: 179.—Ricardo, 1915, Ann. Mag.
Nat. Hist., (8) 16: 278.—? White, 1915, Proc. Roy. Soc. Tasmania, 11.
T. dubiosus Ricardo, 1915, Ann. Mag. Nat. Hist., (8) 16: 284 (dubiosa in error).
T. indefinitus Taylor, 1918, Rec. Austr. Mus., 12: 68.

Synonymy.—Walker’s type of imperfectus was described by Ricardo as having an
enlarged callus taking up “the whole of the anterior half of the forehead” and no
extension of the callus was seen by her. She records it from New South Wales.

T. dubiosus Ricardo, based on Queensland specimens, at the same time recorded
from Katoomba, New South Wales, is said to have the callus almost reaching the eyes,
oblong, the linear extension ending in a point.

T. indefinitus Taylor, also from the Blue Mountain area, is said to have its callus
as wide as the frons and with a short extension. The type shows the callus agreeing
with the description, and yet corresponding with two specimens from Katoomba the
callus of which is open to varied interpretations explained below. In Australian
collections specimens standing under the three names are certainly one species.

These two Katoomba specimens were caught early in November, 1949, and, although
not long emerged, already had their frons abraded, being bare of vestiture over a wide
area. The apparent callus is therefore large, somewhat gothic-arched in shape, ending
in a point two-thirds towards the summit. The lower half of the frons is almost
completely occupied by this pseudocallus and the arching extension is in no way
divided from it. When judged to be abraded, and seeking the true outline, one can

BY G. H. HARDY. 223

imagine a slender process arising from a practically eye-to-eye oblong callus, or even
an oblong without a process. Even this may be reduced to a squarish or roundish
object, well separated from the eyes, a view justified by the evident but very minute
hair-pits scattered over the surrounding area.

After collecting various extra specimens at Katoomba, and after re-examining the
material used by Taylor, I considered that both Ricardo and Taylor were deceived by
appearances, due to the condition of the specimens they studied, all specimens from the
Blue Mountain area being but one species.

T. imperfectus White, described from Mangalore, Tasmania, shows a wide variety
of pseudocalli on specimens available for study, but it is doubtful if it be truly con-
specific with the mainland form.

Characters.—A dark, hairy-eyed species, with the apical markings of each segment
ashy-white, faintly yellowish at times, the colour merging into a central white spot on
a blackish-brown abdomen; the sides of the first two segments are often lighter brown.
The true callus is probably small, well separated from the eyes, but the frons becomes so
readily denuded that the callus becomes ill-defined and it is not known if a true
extension occurs. Minute hair-pits scattered over the surface will denote that abrasion
has taken place, forming a pseudocallus. The vestiture of the frons is mainly dark
greyish with some brown staining over the central area. Apparently on specimens that
are old when caught, the frons slightly collapses, becoming parallel sided, but normally
the frons widens slightly towards the antennae.

Remarks.—Taylor refers to golden hairs in the abdominal spots and margins. These
are slightly yellowish on his type, so I presume Taylor examined them under artificial
light to see them as golden.

Hab.—In literature the species is recorded from Queensland to Tasmania. It occurs
at Katoomba, New South Wales, rather sparingly, specimens being taken from
November, 1949, through the summer months, but only 14 were captured over the period.
In the summer 1950-51 only three were seen. The collecting area is around a swamp
on Katoomba Creek, which supports Scaptia auriflua Don., S. patula Walk. and S. brevi-
cornis Maeq., all in small numbers, but no male of a Tabanus was seen there. Another
Tabanus occurs and seems to be a rare undescribed species of subgenus Cydistomyia.

Genus HETEROPSILOPUS Bigot, 1859.

The limits of the genus are not certainly known. Chrysosoma interruptum Becker
and C. caelicum Parent, differing only in having the third antennal segment at least
one and a half times longer than broad and with a terminally placed arista, probably
belong here, but the following key contains only those with typical antennae.

Key to species of genus Heteropsilopus. "
1. Anterior tarsi combined are 2% times longer than the tibia .......... jacquelinei Parent.

Anterior tarsi combined are less than twice the length of the tibiae ................ 2
2. First posterior tarsus as long as the four following segments combined .............. 3
IMDS DORUWSPIOR TAURI) SINGH) SoeucooducosudnaeduuoncsouuocODOUbaogdomOooOEo Do eDOOE 4

Abdomen rarely with yellow above, then limited to the anterior margins of the first two
segments. Male with intermediate tarsi fringed with hook-shaped cilia ............
Teast etre is sete ae eres nrss. ete ico te, oii aciavaaeets aaecaastnrniicnape ue ener teense Pees cingulipes Walker.

Abdomen with conspicuous yellow parts at least basally. Male with a cuticular apical
spur on the first segment of the intermediate tarsus ........... brevicornis Macquart.

4. First posterior tarsus as long as the three following segments combined. Anterior tarsi
combined only 1% the length of tibia. Male intermediate tarsi with the two apical
Sesmentse peculiarliye ORME! 0c cits auspsessersashepeusy cont oes) ceneiolel san ean nee aa ingenwus Hrichson.

First posterior tarsus as long as the two following segments combined. Anterior tarsi
combined 14 length of tibia. Male with a broadened fifth segment on the intermediate
ENT SMe ee ya Te ARO R LES yt oie. My E i cndutancuha amie Si Gb prop aed Chae MPEP OME GRR iene trifasciatus Macq.

In this key the proportional lengths of tarsal segments are those given in
descriptions, but slight variations may be found on specimens from different regions. It
has yet to be shown that the two names in couplet 4 are not based on the one species,

(Jt)

224 MISCELLANEOUS NOTES ON AUSTRALIAN DIPTERA. XY,

HETEROPSILOPUS CINGULIPES Walker.
Psilopus cingulipes Walker, Ent. Mag., 2, 1835: 472.—Sciapus plumifera Becker, Cap.

Zool., 1, 1922: 206.

Synonymy.—tThat Becker’s species should prove conspecific with that of Walker is
quite unexpected, but the evidence is based on field observations, leaving no doubt
concerning this synonymy. Collecting around my laboratory at Katoomba, I found not
only were the two forms present, but also during the latter half of December all grades
between them in colour variations were present. The species occurred in enormous
numbers, persisting through the summer in diminishing quantities.

In various collections, where specimens come from mainly low-land areas, the
dominant type is the clear winged cingulipes, but the dark winged form also occurs
there. From the Blue Mountains come (dad) specimens with wings clear and faintly
marked; (b) two bands formed but isolated from the costa, and these bands when
interrupted centrally may become four spots on the wing; (c) the two bands, though
reaching the costa, do not join together, varying to join together as in Becker’s
illustration. The third band, which is normally small and interrupted, also varies in
density of colour and in the amount of area covered.

Other parts are similarly variable, the anterior coxae varying from pale yellow,
almost white, to intensely yellow, and usually they are fuscous coloured on the anterior
side. The abdomen may be entirely metallic, varying to mainly yellow below, this
yellow spreading above along the anterior border of the first two segments.

In December, 1950, five specimens with lightly marked wings were kept, each in a
separate glass tube; one died, the other four had developed full and dense colour
markings five days afterwards.

On this evidence it would seem that the synonymy given by me in 1935 is mainly
correct, the exception being that doubtfully placed Chrysosoma metallicum Parent, 1932,
a female now removed. Some mainland specimens have been referred to as trifasciatus
Macq. and most of these are H. ingenuus EHrich., but probably some belong to the present
species, the two being very alike in characters.

Characters.—On the male the intermediate tarsi contain easily overlooked cilia,
forming a row of bent-over, hook-shaped hairs, about 20 of these hairs to the millimetre.
The row forms a fringe reaching the tibia, on which it peters out by narrowing down
to very small hairs. Other characters are given in the key.

Notes.—On the above evidence it may be assumed that other species are similarly
variable in coloration to a greater or less extent and that these variations are found
mainly during the early part of the season in which they occur. There are apparent
trends to regional variations which also must be taken into account.

HETEROPSILOPUS TRIFASCIATUS Macquart.

Psilopus trifasciatus Macquart, 1849.—Chrysosoma metallicum Parent, 1932.

Synonynvy.—Macquart’s type, redescribed by Parent as having the first segment of
the posterior tarsi equal to the length of the two following segments combined, conforms
to C. metallicum, which feature suggests the above synonymy. The type of the latter
is a female, whereas Macquart’s type is a male, of which Parent gives the tarsal
drawing that does not conform with H. ingenuwus Erich., and is perhaps due to slight
damage.

Hab.—Macquart gives Tasmania as type locality in his fourth supplement, but
Sydney is likely to be the true locality.

HETEROPSILOPUS INGENUUS Erichson.
Psilopus ingenuus Hrichson, 1842.—Scapius chalceus White, 1916.

Synonymy.—The type of chalceus, a female specimen, may be the present species
with duller colour and less wing marking than is normally found. It was captured
early in the season.

Characters.—On the male the apical segments of the intermediate tarsi are
peculiarly formed, the shape being illustrated by Parent under the name gloriosus,

BY G. H. HARDY. 225)

which is a synonym. The smaller size and the extension of the apical band along
the costa to the wing tip (as illustrated for trifasciatus by Macquart) will serve to
distinguish the form from cingulipes.

Hab.—The species is definitely known from Tasmania, Victoria, and the Blue
Mountains of New South Wales. It was found in moderate number at Katoomba in
December, 1950, but became scarce in the following month. One specimen had as
prey a species of Simulium.

HETEROPSILOPUS BREVICORNIS Macquart.

Synonymy.—Parent’s remarks about the type state that the middle tarsi are not
ornamented, and this was supplemented later in a letter which stated that he could
not dissociate the name from the present form. There is need for more clarity in
references, but it seems that Scapius chalceus White, 1916, must be removed from
the synonymy.

HETEROPSILOPUS JACQUELINEI Parent.

Sciopus jacquelinei Parent, Ann. Soc. Sci. Bruxelles, (B) 52, 1932: 169.

Characters.—Presumably the intermediate tarsi are simple on the male, the first
segment being described as one and a half times the length of the following segments
combined. The description was based on a unique male from Canberra and has not
been recognized in collections available to me.

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Xxili

ABSTRACT OF PROCEEDINGS.

ORDINARY MONTHLY MEETING.
28th Marcu, 1951.
Mr. A. N. Colefax, President, in the Chair.

Library accessions amounting to 28 volumes, 246 parts or numbers, 32 bulletins,
12 reports and 58 pamphlets, total 376, had been received since the last meeting.

PAPERS READ (by title only).

1. A Septoria Disease of Huphorbia peplus L. By Dorothy E. Shaw.

2. Serological Studies of the Root-nodule Bacteria. IV. Further Analysis of
Isolates from Trifolium and Medicago. By Hilary F. Purchase, J. M. Vincent and
Lawrie M. Ward.

3. Preservation Techniques for Scarabaeid and other Insect Larvae. By P. B. Carne.

ORDINARY MONTHLY MEETING.
26th Apri, 1951.
Mr. A. N. Colefax, President, occupied the Chair.

The President announced that the Council had elected the following office-bearers
for the 1951-52 session: Vice-Presidents, Dr. G. D. Osborne, Dr. Lilian Fraser, Dr. R. N.
Robertson and Mr. D. J. Lee; Honorary Treasurer, Dr. A. B. Walkom; Honorary
Secretary, Dr. W. R. Browne.

Dr. T. Clive Backhouse, D.P.H., D.T.M. & H., F.R.A.C.P., Sydney University, was
elected an Ordinary Member of the Society.

Library accessions amounting to 22 volumes, 139 parts or numbers, 26 bulletins and
19 pamphlets, total 206, had been received since the last meeting.

PAPERS READ.

1. The Anatomy and Morphology of the Operculum in the Genus Hucalyptus.
Part I. The Occurrence of Petals in Hucalyptus gummifera (Gaertn.) Hochr. By J. L.
Willis.

2. Some Notes on Athrotavis. By Charles G. Elliott. (Communicated by Dr.
Patrick Brough.)

3. The Paramphistomes (Trematoda) of Australian Ruminants. Pants

Systematics. By P. H. Durie.

Short addresses were given by Professor Kmmons (Professor of Veterinary
Physiology, University of Sydney) on Biology and Chemistry of Sex Hormones; and
Dr. Margaret Hardy (McMaster Laboratory, University of Sydney) on Tissue-culture
of Hair Follicles.

ORDINARY MONTHLY MEETING.
27th Junp, 1951.
Mr. A. N. Colefax. President. occupied the Chair.

Mr. A. S. Fraser, M.Sc., Sydney University, Mr. H. B. Kerr, B.Sc.Agr., Croydon,
Dr. B. J. F. Ralph, B.Sc., A.A.C.I., University of Technology, Sydney, and Mrs. C. E.
Seccombe, B.A., Bellevue Hill, were elected Ordinary Members of the Society.

The President offered congratulations to the following members: Dr. Joan Beattie,
on the award of the D.Sc. degree; Miss Judith Fraser, on obtaining her M.Se. degree;
Dr. R. J. Noble, on receiving the Outstanding Achievement Award of the University of

W

XX1V ABSTRACT OF PROCEEDINGS.

Minnesota, U.S.A.; Miss Hilary Purchase, on the award of the Thomas Lawrance Pawlett
Research Scholarship of the University of Sydney, being the first occasion on which the
scholarship has been awarded to a woman; and Dr. A. R. Woodhill, on obtaining the
D.Sc. degree of the University of Sydney.

Library accessions amounting to 23 volumes, 144 parts or numbers, 33 bulletins,
13 reports and 14 pamphlets, total 227, had been received since the last meeting.

PAPERS READ.

1. A Review of the Australian Species of Sarcochilus (Orchidaceae). By H. M. R.
Rupp.

2. Australian Rust Studies. VIII. Puccinia graminis lolii, an Undescribed Rust of
Lolium spp., and Other Grasses in Australia. By Professor W. L. Waterhouse.

3. On Trombicula minor Berlese, 1905. By Carl E. M. Gunther.

4. Studies on Australian Marine Algae. VI. New Geographical Records of Certain
Species. By Valerie May.

5. The Marine Algae of Brampton Island, Great Barrier Reef, off Mackay,
Queensland. By Valerie May.

6. The Evolution of the Radio-medial Area in the Wings of the Muscoidea Acalyptrata
(Diptera). By H. F. Lower.

An address was given by Dr. Baas-Becking, a distinguished Netherlands botanist,
entitled “A Biologist Looks at the Problem of Waste’.

ORDINARY MONTHLY MEETING.
25TH JULY, 1951.

Mr. A. N. Colefax, President, occupied the Chair.

Mr. B. R. Hewitt, Kempsey, N.S.W., was elected an Ordinary Member of the Society.

The President announced that Dr. F. V. Mercer had been elected a member of
Council in place of Dr. R. N. Robertson, who had resigned.

The President referred to the death of Dr. Robert Broom, who had been a Corres-
ponding Member of the Society since 1902, on 6th April, 1951, aged 84.

The President offered congratulations to Dr. Janet Harker, F.R.E.S., on obtaining the
Ph.D. degree of the University of Manchester, and to Dr. G. F. Humphrey, on the award
of the Ph.D. degree of the University of Sydney.

Library accessions amounting to 17 volumes, 116 parts or numbers, 4 bulletins,
2 reports and 13 pamphlets, total 152, had been received since the last meeting.

PAPERS READ.
_ 1. A Mite from a Beehive on Singapore Island (Acarina: Laelaptidae). By Carl

BE. M. Gunther. 5

2. Celaenopsoides Gunther, 1942: A Synonym of Ophiomegistus Banks, 1914
(Acarina: Parasitidae). By Carl E. M. Gunther.

3. The Development of Bryozoan Faunas in the Upper Palaeozoic of Australia. By
Joan Crockford (Mrs. Beattie).

Lecturettes, illustrated by lantern slides, were given by members of the recent
expedition to the Kosciusko area, in regard to the Glaciology, Soils, Vegetation, and
Land Usage.

ORDINARY MONTHLY MEETING.
29th AuaustT, 1951.
Mr. A. N. Colefax, President, occupied the Chair.

Messrs. E. 8S. Lowery, M.A., Northmead, N.S.W., and D. K. McAlpine, Bronte, N.S.W.,
were elected Ordinary Members of the Society,

ABSTRACT OF PROCEEDINGS. xXxXV

The President announced that Dr. A. R. Woodhill had been elected a Vice-President
for the remainder of the current session in place of Dr. R. N. Robertson, who had
resigned from the Council.

Library accessions amounting to 7 volumes, 73 parts or numbers, 44 bulletins and
3 reports, total 127, had been received since the last meeting.

PAPERS READ.
Succinoxidase of Potato Tuber. By Adele Millerd.
Controlled Pollination of Hucalyptus. By L. D. Pryor.
A Genetic Analysis of some Hucalyptus Species. By L. D. Pryor.

im elcioe es hoa te

Investigations on the Preferences shown by Aédes (Stegomyia) aegypti Linn.,
and Ciilen (Culex) fatigans Wied., for Specific Types of Breeding Water. By Tom
Manefield.

5. The Nomenclature of Heteronychus sanctae-helenae Blanchard (Coleoptera:
Scarabaeidae: Dynastinae). By E. B. Britton. (Communicated by P. B. Carne.)

NOTES AND EXHIBITS.

Miss Kathleen M. I. English exhibited an introduced slug, TVestacella haliotidea.
Testacella, a carnivorous snail slug, was discovered in considerable numbers in Adelaide
Botanic Gardens in 1931. In 1949, they were recorded from a garden in Geelong,
Victoria, where they had been noticed for some years. Distribution: Hurope and
Canary Is. References: Wild Life, xi, pp. 506 and 516 (Nov., 1949); xii, p. 84 (Feb.,
1950).

Mr. A. Musgrave exhibited specimens of the Giant Orb-weaver, Nephila maculata
(Fabricius, 1793), a member of the Family Eperidae (= Argiopidae) from the North
Coast of New South Wales. Three females of this species have been forwarded to the
Australian Museum, Sydney, during the past decade. Two of these were sent by Dr.
G. H. Hewitt of Bellingen, Bellinger River, and were collected about 12th January,
1941, and the 2nd February, 1951 (the last from Repton near the mouth of the river).
The third specimen came from Glenreagh, a little north of Bellingen, and was collected
by Mr. D. Thompson, 1ith June, 1950. The spider has a wide range from Africa,
through India to China and the Pacific Islands. In Australia it has only been
recorded from Queensland, so that these records from the Bellingen district extend the
range of the spider further south.

On behalf of Professor P. D. F. Murray, Mr. A. N. Colefax exhibited specimens of
young turtles from the Great Barrier Reef. These had been stained with alizarin, and
cleared, in order to show the development of the bony skeleton. All of the bony
elements were bright red, and were in strong contrast to the remaining tissues which
remained unstained, though transparent. Fixed, but otherwise untreated specimens,
were also shown by way of comparison.

ORDINARY MONTHLY MEETING.
26TH SEPTEMBER, 1951.

Mr. A. N. Colefax, President, occupied the Chair.

Miss Helen P. Laneaster, B.Se., University of Sydney, was elected an Ordinary
Member of the Society.

The President referred to the death on 19th September, 1951, of Mr. Edwin Cheel,
who had been a member of the Society since 1899, a member of Council from 1925 to
1940, and President, 1930-31; also to the death, on 30th August, 1951, of Professor T.
Harvey Johnston, a member of the Society since 1907.

The President announced that the Council is prepared to receive applications for
Linnean Macleay Fellowships tenable for one year from Ist January, 1952, from qualified

Www

xxvi ABSTRACT OF PROCEEDINGS.

candidates. Applications should be lodged with the Hon. Secretary not later than
Wednesday, 7th November, 1951.

Library accessions amounting to 9 volumes, 155 parts or numbers, 36 bulletins,
3 reports and 13 pamphlets, total 216, had been received since the last meeting.

PAPER READ.

The Life-history of a Penaeid Prawn (Metapenaeus) breeding in a Coastal Lake
(Tuggerah, New South Wales). By Muriel C. Morris and Isobel Bennett.

Addresses were given by Dr. F. V. Mercer on “Biology, the Poor Relation”, and by
Dr. R. Catala, a visiting French marine biologist, on his observations in the Gilbert
Islands, illustrated by lantern slides and photographs.

NOTES AND EXHIBITS. -

Mr. A. J. Bearup exhibited a blood-film of an Australian gecko, Phyllurus platurus
(Shaw), collected at Glen Davis by Mr. R. Palmer on 6th August, 1951, showing
chromatin bodies and vacuoles similar to those described by Chatton and Blane as
Pirhaemocyton tarentolae. The film also showed haemogregarines and microfilariae.
The adult filarial worms were found in the mesentery supporting the liver.

Dr. F. V. Mercer exhibited a lantern slide of a caterpillar, which had been collected
by Miss K. English, on a dried branch of Hucalyptus ecugenioides, exhibiting mimicry
of its surroundings to a remarkable degree. The most striking feature of the mimicry
was the similarity between the top of the head and the broken end of the twig.

ORDINARY MONTHLY MEETING.
31st OcToBER, 1951.
Mr. A. N. Colefax, President, occupied the chair.

Miss Isobel I. Bennett, Crow’s Nest, N.S.W., and Mr. G. H. Packham, Earlwood,
N.S.W., were elected Ordinary Members of the Society.

The President announced that the Council is prepared to receive applications for
Linnean Macleay Fellowships, tenable for one year from ist January, 1952, from
qualified candidates. Applications should be lodged with the Hon. Secretary not later
than Wednesday, 7th November, 1951.

Library accessions amounting to 22 volumes, 116 parts or numbers, 5 reports and
5 pamphlets, total 148, had been received since the last meeting.

PAPERS READ.

1. Remarks on some Australian Laius Guér. (Col. Malachiidae). By Walter
Wittmer. (Communicated by J. W. T. Armstrong.)

2. A Critical Consideration of c-mitosis with Reference to the Effects of Nitrophenols.
By Mary M. Hindmarsh, Linnean Macleay Fellow in Botany.

NOTES AND EXHIBITS.

Miss M. Hindmarsh exhibited a film of mitosis in the reproductive cells of the
Grasshopper. The development of the phase contrast microscope has made possible
observations on living cells and this technique has been employed in making this
film.

Dr. R. N. Robertson exhibited a preparation of plant mitochondria which had been
extracted from tissue of red beet and an electron micrograph of mitochondria taken
by Dr. E. H. Mercer.

Mr. A. N. Colefax exhibited a device for the rapid moulding of metal foil containers
for paraffin embedding. Attached to a wooden base is a rectangle of 18g. flat brass the
size of the blank, and a small block of brass the size and shape of the desired container.

ABSTRACT OF PROCEEDINGS. XXV1i

The lead foil is first rubbed on the flat plate to imprint the size of the latter
on it, cut to size with a pair of scissors, and then moulded, by the fingers, around the
brass: block. This gives a one-piece container with the usual triangular flaps at the
ends.

Speed of production is high, and it is recommended that several sizes of blanking
plate and block be mounted on the one base, in order to cover a range of containers
commonly used in embedding.

Mr. D. J. Lee gave a short summary of recent observations and the importance
of entomological work in connection with outbreaks of myxomatosis and encephalitis
in Eastern Australia.

ORDINARY MONTHLY MEETING.
28th NoveMBER, 1951.
Mr. A. N. Colefax, President, occupied the Chair.

Professor L. G. M. Baas-Becking, Ph.D., D.Se., Botany Department, Sydney
University, and Mr. Colin L. Macdonald, Lewisham, N.S.W., were elected Ordinary
Members of the Society.

The President announced that Miss Mary Hindmarsh and Mr. T. G. Vallance had
been reappointed to Linnean Macleay Fellowships in Botany and Geology respectively
for the year 1952.

The President also announced that a series of eight broadcast lectures on “The
Nature of the Universe’, by Fred Hoyle, will be broadcast by the National Station on
the first eight Friday nights in 1952, ie, 4th January to 22nd February, 1952, at
7.30 p.m.

Library accessions amounting to 20 volumes, 168 parts or numbers, 24 bulletins and
3 reports, total 215, had been received since the last meeting.

PAPERS READ.

1. Miscellaneous Notes on Australian Diptera. XV. Tabanus, Heteropsilopus. By
G. H. Hardy.

2. A Revision of Australian Species previously referred to the Genus Empoasca
(Cicadellidae, Homoptera). By Harold F. Lower.

3. The Use of Excised Shoots in Linseed Investigations. By H. B. Kerr.

The President referred to the 60th anniversary (on 7th December, 1951) of the
death of the Society’s benefactor, Sir William Macleay, and historic personal relics.
were exhibited.

A welcome was extended to a party of scientists from the Danish research frigate
“Galathea”; and a lecture was given by Dr. Anton Bruun, leader of the expedition, on
the aims and achievements of the scientific expedition, concluding with a film produced
by the party’s information officer, which showed the early work in fitting and equipping
the ship, its trials and departure from Copenhagen.

XXV1il

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1227
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1$22
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1912

LIST OF MEMBERS.
(15th December, 1951.)

ORDINARY MEMBERS.

(An asterisk (*) denotes Life Member.)

Abbie, Professor Andrew Arthur, M.D., B.S., B.Sec., Ph.D., c.o. University of Adelaide,
Adelaide, South Australia.

*Albert, Michel Francois, ‘““Boomerang”’’, 42 Billyard Avenue, Elizabeth Bay, Sydney.

*Allman, Stuart Leo, B.Sc.Agr., M.Sc., Entomological Branch, Department of Agriculture,
Farrer Place, Sydney.

Anderson, Robert Henry, B.Sc.Agr., Botanic Gardens, Sydney.

*Armstrong, Jack Walter Trench, “Callubri’, Nyngan, N.S.W.

Aurousseau, Marcel, B.Se., c.o. Mr. G. H. Aurousseau, 16 Woodland Street, Balgowlah,
N.S. W.

Backhouse, Thomas Clive, M.B., B.S., D.P.H., D.T.M. & H., F.R.A.C.P., School of Public
Health and Tropical Medicine, Sydney University.

Baddams, Miss Greta, B.A., B.Sc., New England University College, Armidale, N.S.W.

*Barber, Professor Horace Newton, M.A., Ph.D., Department of Botany, University of
Tasmania, Hobart, Tasmania.

Barrett, Mrs. Judith Hope, M.Se. (née Balmain), Dairy Research Institute, Shinfield,
near Reading, Berks, England.

*Beadle, Noel Charles William, D.Se., Botany School, Sydney University.

Bearup, Arthur Joseph, 66 Pacific Avenue, Penshurst, N.S.W.

Beattie, Joan Marion, D.Se. (née Crockford), Bradley Street, Cobar, N.S.W.

Benson, Professor William Noel, B.A., D.Se., F.G.S., University of Otago, Dunedin, New
Zealand.

Besly, Miss Mary Ann Catherine, B.A., 7 Myra Street, Wahroonga, N.S.W.

Birch, Louis Charles, B.Ag.Se., M.Se., Department of Zoology, Sydney University.

Blake, Stanley Thatcher, M.Se., Botanic Gardens, Brisbane, Queensland.

Boardman, William, M.Se., Zoology Department, University of Melbourne, Carlton, N.3,
Victoria.

Bradhurst, Miss Peggy Joan, B.Sc., 25 Belgium Avenue, Roseville, N.S.W.

Brett, Robert Gordon Lindsay, B.Sc., 7 Petty Street, West Hobart, Tasmania.

Brown, Kenneth George, 6 Dolphin Street, Randwick, N.S.W.

Browne, Ida Alison, D.Se. (née Brown), Department of Geology, Sydney University.

Browne, Lindsay Blakeston Barton, 34 Kent Road, Rose Bay, N.S.W.

Browne, William Rowan, D.Se., Department of Geology, Sydney University.

Bryan, Clement, B.A., Intermediate High School, Corowa, N.S.W.

Burden, John Henry, 1 Havilah Street, Chatswood, N.S.W.

*Burges, Professor Norman Alan, M.Sc., Ph.D., Botany School, Sydney University.

Burkitt, Professor Arthur Neville St. George Handcock, M.B., B.Se., Medical School,
Sydney University.

Campbell, Mrs. Emily Mary, 22 Madeline Street, Hunter’s Hill, N.S.W.

Campbell, Thomas Graham, Division of Economic Entomology, C.S.L.R‘O., P.O. Box 109,
City, Canberra, A.C.T.

*Carey, Professor Samuel Warren, D.Sc., Geology Department, University of Tasmania,
Hobart, Tasmania.

Carne, Phillip Broughton, B.Agr.Se. (Melb.), Department of Entomology, Imperial
College of Science and Technology, Prince Consort Road, London, S.W.7, England.

Carne, Walter Mervyn, 7 De Villiers Avenue, Chatswood, N.S.W.

*Chadwick, Clarence Harl, B.Sc., Entomological Branch, Department of Agriculture, Farrer
Place, Sydney.

Christian, Stanley Hinton, Malaria Survey, Banz, Central Highlands, via Lae, New
Guinea.

*Churchward, John Gordon, B.Se.Agr., Ph.D., 1 Hunter Street, Woolwich, N.S.W.

Clark, Laurance Ross, M.Sc., c.o. C.S.I.R.O., P.O. Box 13, Bright, Victoria.

Cleland, Professor John Burton, M.D., Ch.M., 1 Dashwood Road, Beaumont, Adelaide,
South Australia.

Cleland, Kenneth Wollaston, M.B., Department of Anatomy, Sydney University.

Colefax, Allen Neville, B.Sc., Department of Zoology, Sydney University.

Colless, Donald Henry, Borneo Malaria Research, Labuan, British North Borneo.

1942
U9 7,
1908
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1945
1948
1950
1936
1934
1925
1937
1927
1948

1926
1946

1948
WG)eei

1949

1947
1930

1947
1948
1948
1930

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1935
1944

1946
1936
1939

1950
1928
1917
TAT

1932
1947
1951
1930

1938

1943
1930

1943

1932
1942

1917
1938

AUG YeST
1945
1937

1930
1948
1933

LIST OF MEMBERS. XXix

Copland, Stephen John, B.Sc., Chilton Parade, Warrawee, N.S.W.

Costin, Alec Baillie, 12 Barambah Road, Roseville, N.S.W.

Cotton, Professor Leo Arthur, M.A., D.Sc., Department of Geology, Sydney University.
Crawford, Lindsay Dinham, B.Sc., 4 Dalton Avenue, West Hobart, Tasmania.

Davis, Mrs. Gwenda Louise, B.Sc., New England University College, Armidale, N.S.W.

Davison, Miss Daphne Claire, M.Sc., 9 Brissenden Avenue, Collaroy, N.S.W.

Day, Alan Arthur, 13 Besborough Avenue, Bexley, N.S.W.

Day, Maxwell Frank, Ph.D., B.Sc., C.S.1.R.O., Box 109, Canberra, A.C.T.

Day, William Eric, 23 Gelling Avenue, Strathfield, N.S.W.

de Beuzeville, Wilfred Alexander Watt, J.P., ““Melamere’’, Welham Street, Beecroft, N.S.W-

Deuquet, Camille, B.Com., 126 Hurstville Road, Oatley, N.S.W.

*Dixson, Sir William, ‘‘Merridong”’’, 586 Gordon Road, Killara, N.S.W.

Drover, Donald P., Institute of Agriculture, University of Western Australia, Nedlands,
W.A.

Dumigan, Edward Jarrett, 10 High Street, Toowoomba, Queensland.

Durie, Peter Harold, B.Se., C.S.I.R.O., Veterinary Parasitology Laboratory, Yeerongpilly,
Brisbane, Queensland.

Haley, Eric H. M., 18 Ray Road, Epping, N.S.W.

Edwards, Eric Thomas, Ph.D., M.Sc.Agr., National Press Pty. Ltd., 126-130 Phillip Street,
Sydney.

Blliott, John Henry, 8 Shellecove Road, Neutral Bay, N.S.W.

Endean, Robert, M.Sc., 15 Milton Avenue, Eastwood, N.S.W.

English, Miss Kathleen Mary Isabel, B.Sc., 1 Mt. Morris Street, Woolwich, N.S.W.

Fenton, Miss Enid Grace, 45 Cecil Street, Gordon, N.S.W.

Fraser, Ian McLennan, 131 Fox Valley Road, Wahroonga, N.S.W.

Fraser, Miss Judith A., M.Sc., 14 Milray Avenue, Wollstonecraft, N.S.W.

Fraser, Miss Lilian Ross, D.Se., ‘““Hopetoun’’, 25 Bellamy Street, Pennant Hills, N.S.W.

Garden, Miss Joy Gardiner, B.Sc.Agr., Botanic Gardens, Sydney.

*Garretty, Michael Duhan, D.Sc., “‘Surrey Lodge’, Mitcham Road, Mitcham, Victoria.

Greenwood, William Frederick Neville, c.o. Colonial Sugar Refining Co. Ltd., Lautoka,

Fiji.

*Griffiths, Mrs. Mabel, B.Se. (née Crust), 2 Carden Avenue, Wahroonga, N.S.W.

Grifiths, Mervyn Edward, M.Sc.,:Australian Institute of Anatomy, Canberra, A.C.T.

*Gunther, Carl Ernest Mitchelmore, M.B., B.S., D.T.M., D.T.M. & H. (England), Bulolo
Gold Dredging Limited, Bulolo, Territory of Papua—New Guinea.

Hallinan, John Michael, ‘‘Quirindi’, 65 Dunmore Street, Bexley, N.S.W.

Hamilton, Edgar Alexander, 16 Hercules Street, Chatswood, N.S.W.

Hardy, George Huddleston Hurlstone, “The Gardens’, Letitia Street, Katoomba, N.S.W.

Harker, Janet Elspeth, M.Sc., Ph.D., F.R.E.S., c.o. Zoological Department, The University,
Manchester 13, England.

Harris, Miss Thistle Yolette, B.Sc., 14 Pacific Street, Watson’s Bay, N.S.W.

Henderson, David Leonard Wylie, L.S.M.I.S., ‘‘Berida’”’, R.M.B. 420, Bourke, N.S.W.

Hewitt, Bernard Robert, c.o. Box 177, Kempsey, 2C, N.S.W.

Heydon, George Aloysius Makinson, M.B., Ch.M., Flat 5, 79 O’Sullivan Road, Rose Bay,
N.S.W.

Hill, Miss Dorothy, M.Se., Ph.D., Department of Geology, University of Queensland,
Brisbane, Queensland.

Hindmarsh, Miss Mary Maclean, B.Sc., Botany School, University of Sydney, N.S.W.

Holmes, Professor James Macdonald, Ph.D., B.Sc., F.R.G.S., F.R.S.G.S., Department of
Geography, Sydney University.

Horowitz, Benzoin, Hng.Agr.S., Dr.Agr.Se. (Cracow, Poland), C.S.I.R.O., c.o. Waite
Institute, Private Mail Bag, Adelaide, South Australia.

Hossfeld, Paul Samuel, M.Sc., 132 Fisher Street, Fullarton, South Australia.

Humphrey, George Frederick, M.Se., Ph.D., Department of Biochemistry, Sydney
University.

Jacobs, Ernest Godfried, ‘““Cambria’’, 106 Bland Street, Ashfield, N.S.W.

Jacobs, Maxwell Ralph, D.Ing., M.Se., Dip.For., Commonwealth Forestry Bureau, Canberra,
JN AC AN

Johnson, Lawrence Alexander Sidney, B.Sc., 178 Beecroft Road, Cheltenham, N.S.W.

Johnston, Arthur Nelson, B.Sc. Agr., Hawkesbury Agricultural College, Richmond, N.S.W.

Jones, Mrs. Valerie Margaret Beresford, M.Se. (née May), Mooloolabel Esplanade,
Narrabeen, N.S.W.

Joplin, Germaine Anne, B.Se., Ph.D., Bureau of Mineral Resources, Canberra, F.4, A.C.T.

Jopling, Alan Victor, B.Sc., B.E., 86 Alfred Road, Brookvale, N.S.W.

Judge, Leslie Arthur, 87 Eastern Road, Turramurra, N.S.W.

XXX

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1938
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1939

1946
1946
1932
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1936

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1943
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1951

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1922

1931
1948
1948

1949

1932
1905

1933
1951
1932

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1949
1944

1947 °

1949
1938
1948
1947
1946
1949
1947

1944
1939
1926
1949

1920
1949

LIST OF MEMBERS.

Keast, James Allen, B.Sc., Australian Museum, College Street, Sydney.

Kerr, Harland Benson, B.Sc.Agr., 41 Badminton Road, Croydon, N.S.W.

Kesteven, Geoffrey Leighton, D.Sc., c.o. F.A.O. of United Nations, Milawa Mansions, Para
atit Road, Bangkok, Siam.

Kesteven, Hereward Leighton, D.Se., M.D., Palmwoods, Queensland.

Kiely, Temple Baylis, D.Se.Agr., Caroline Street, East Gosford, N.S.W.

Kinghorn, James Roy, C.M.Z.S., Australian Museum, College Street, Sydney.

Kooptzoff, Miss Olga, 322 Moore Park Road, Paddington, N.S.W.

Krishnamurthy, Kolipakkam Venkatesa, M.A., Ph.D., Central Tobacco Research Institute,
Rajamundry, South India.

Langford-Smith, Trevor, M.Sc., Department of National Development, Acton, Canberra,
AL Cure ;

Larcombe, Miss Pauline Gladys, B.Sc., 17 Ethel Street, Burwood, N.S.W.

*Lawrence, James Joscelyn, B.Sc., 91 Boundary Street, Clovelly, N.S.W.

Lawson, Albert Augustus, 9 Wilmot Street, Sydney.

Lazer, Mrs. Ruth, M.Se. (née Wolf), 25 Sir Thomas Mitchell Road, Bondi, N.S.W.

Lee, Mrs. Alma Theodora, M.Se. (née Melvaine), Manor Road, Hornsby, N.S.W.

Lee, David Joseph, B.Sc., School of Public Health and Tropical Medicine, Sydney
University.

Liddell, Miss Jean Joyce, M.Se., Zoology Department, University of Melbourne, Carlton,
N.3, Victoria.

Lothian, Thomas Robert Noel, Botanic Gardens, Adelaide, South Australia.

Lower, Harold Farnham, Waite Agricultural Research Institute, University of Adelaide,
Private Mail Bag, G.P.O., Adelaide, South Australia.

Lowery, Edwin Sanders, M.A., 13 Francis Street, Northmead, N.S.W.

Macintosh, Neil William George, M.B., B.S., 32 Benelong Crescent, Bellevue Hill, N.S.W.

Mackerras, David, 48 Edgecliff Road, Bondi Junction, N.S.W.

Mackerras, Ian Murray, M.B., Ch.M., B.Sc., Queensland Institute of Medical Research,
Herston Road, Valley, Brisbane, Queensland.

*Mair, Herbert Knowles Charles, B.Sc., 5 Collaroy Street, Collaroy Beach, N.S.W.
Manefield, Tom, Department of Agriculture and Stock, P.O. Box 65, Nambour, Queensland.
Marks, Miss Elizabeth Nesta, M.Sc., Biology Department, University of Queensland,

Brisbane, Queensland.
Marshall, Professor Charles Edward, B.Sc., Ph.D., D.Se., F.G.S., M.I.Min.E., Department
of Geology, University of Sydney, N.S.W.

Martin, Donald, B.Se., C.S.1.R.O., Stawell Avenue, Hobart, Tasmania.

Mawson, Sir Douglas, D.Sc., B.E., F.R.S., University of Adelaide, Adelaide, South
Australia.

Maze, Wilson Harold, M.Sce., University of Sydney.

McAlpine, David Kendray, 12 St. Thomas Street, Bronte, N.S.W.

McCulloch, Robert Nicholson, B.Se.Agr., B.Se., Roseworthy Agricultural College, Rose-
worthy, South Australia.

McKee, Hugh Shaw, B.A., D.Phil. (Oxon.), c.o. C.S.I.R.O., Private Bag, Homebush P.O.,

N.S.W

McMillan, Bruce, 171 Lawson Street, Hamilton, Newcastle, N.S.W.

McPhail, Miss Isabel Jean, B.Se., 3 Margaret Street, Canterbury, E.7, Melbourne, Victoria.
Mercer, Frank Verdun, B.Sc., Ph.D. (Cambridge), Botany School, Sydney University.
Messmer, Mrs. Pearl Ray, 64 Treatts Road, Lindfield, N.S.W.

*Miller, Allen Horace, B.Se., Dip.Ed., 1281 Canterbury Road, Punchbowl, N.S.W.

Miller, David, Ph.D., M.Se., F.R.S.N.Z., F.R.E.S., Cawthron Institute, Nelson, New Zealand.
Millerd, Miss Alison Adele, M.Sce., 51 Park Avenue, Roseville.

Millett, Mervyn Richard Oke, B.A., 19 Avoca Street, South Yarra, Vic.
Millington, Richard James, New England University College, Armidale, N.S.W.

‘Minter, Philip Clayton, 49 Cotswold Road, Strathfield, N.S.W.

Morris, Miss Muriel Catherine, M.Sc., c.o. Commercial Banking Company of Sydney Ltd.,
18 Birchin Lane, Lombard Street, London, E.C.3, England.

Moye, Daniel George, B.Sc., Dip. Ed., S.M.H.H.A., 6 First Avenue, Cooma, N.S.W.

Moye, Mrs. Joan, B.Sc. (née Johnston), S.M.H.E.A., 6 First Avenue, Cooma, N.S.W.

Mungomery, Reginald William, c.o. Bureau of Sugar Experiment Stations, Department
of Agriculture and Stock, Brisbane, B.7, Queensland.

Murray, Professor Patrick Desmond Fitzgerald, M.A., D.Se., Department of Zoology,
University of Sydney, N.S.W.

Musgrave, Anthony, F.R.E.S., Australian Museum, College Street, Sydney.

Myers, Kenneth, C.S.1.R.O., Wildlife Survey Section, Field Station, 598 Affleck Street,
Albury, N.S.W.

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1927

1921
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1245
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1932
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1948

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1898
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1948
1911

LIST OF MEMBERS. XXXi

Newman, Professor Ivor Vickery, M.Sc., Ph.D., F.R.M.S., F.L.S., Department of Botany,
University of Ceylon, Colombo 3, Ceylon.

Nicholson, Alexander John, D.Sc., F.R.E.S., C.S.I.R.O., Box 109, Canberra, A.C.T.

*Noble, Norman Scott, D.Se.Agr., M.Se., D.I.C., C.S.1.R.O., 314 Albert Street, Hast
Melbourne, C.2, Victoria.

Noble, Robert Jackson, B.Sc.Agr., Ph.D., 324 Middle Harbour Road, Lindfield, N.S.W.

North, David Sutherland, 42 Chelmsford Avenue, Lindfield, N.S.W.

O’Brien, Brian Robert Alexander, Department of Histology, Sydney University.

O’Farrell, Antony Frederick Louis, A.R.C.Sc., B.Sc., F.R.E.S., New England University
College, Armidale, N.S.W.

O’Gower, Alan Kenneth, B.Sc., 59 Brighton Street, Harbord, N.S.W.

Oke, Charles George, 34 Bourke Street, Melbourne, C.1, Victoria.

Osborn, Professor Theodore George eee, D.Sc., F.L.S., Professor of Botany, University
of Oxford, Oxford, England.

Osborne, George Davenport, D.Sc., Ph.D., Department of Geology, Sydney University.

Oxenford, Reginald Augustus, B.Sec., 9 Cambridge Street, Singleton, N.S.W.

*Pasfield, Gordon, B.Sc.Agr., 20 Cooper Street, Strathfield, N.S.W.

Perkins, Frederick Athol, B.Sc.Agr., Biology Department, University of Queensland,
Brisbane, Queensland.

Phillips, Miss Marie Elizabeth, M.Sc., 4 Morella Road, Clifton Gardens, N.S.W.

Plomley, Kenneth Francis, 50 Domain Street, South Yarra, Melbourne, Victoria.

Pope, Miss Elizabeth Carington, M.Sc., Australian Museum, College Street, Sydney.

Pryor, Lindsay Dixon, M.Sc., Dip.For., Parks and Gardens Section, Department of the
Interior, Canberra, A.C.T.

Purchase, Miss Hilary Frances, B.Sc.Agr., 19 Hampden Avenue, Cremorne, N.S.W.

Raggatt, Harold George, D.Sc., 60 Arthur Circle, Forrest, Canberra, A.C.T.

Ralph, Bernhard John Frederick, B.Sc., Ph.D, (Liverpool), A.A.C.I., N.S.W. University
of Technology, Broadway, Sydney.

Rickwood, Frank Kenneth, B.Sc., Department of Geology, Sydney University.

Riek, Edgar Frederick, B.Sc., C.S.I.R.O., Division of Heconomie Entomology, P.O. Box
109, City, Canberra, A.C.T.

Roberts, Noel Lee, 43 Hannah Street, Beecroft, N.S.W.

Robertson, Rutherford Ness, B.Sc., Ph.D., Food Preservation Research Laboratory,
C.S.1.R.0O., Private Mail Bag, Homebush, N.S.W.

Ross, Donald Ford, c.o. Ross Bros. Pty. Ltd., 545-547 Kent Street, Sydney.

Roughley, Theodore Cleveland, B.Sc., F.R.Z.S., Chief Secretary's Department, Box 30a,
_ G.P.O., Sydney. :

Salter, Keith Eric Wellesley, B.Se., ‘““Hawthorn’’, 48 Abbotsford Road, Homebush, N.S.W.

*Scammell, George Vance, B.Se., 7 David Street, Clifton Gardens, N.S.W.

Scott, Miss Beryl, B.Sc., Department of Geology, University of Tasmania, Hobart, Tas.

Seecombe, Mrs. Lorraine, B.A., 28 Fairweather Street, Bellevue Hill, N.S.W.

Sellgren, Miss Elise Evelyn, B.Sc., 76 Bream Street, Coogee, N.S.W.

*Sharp, Kenneth Raeburn, B.Sc., Eng. Geology, S.M.H.H.A., Cooma, 4S, N.S.W.

Shaw, Miss Dorothy Edith, B.Sc.Agr., Faculty of Agriculture, Sydney University.

Sherrard, Mrs. Kathleen Margaret, M.Sc., 43 Robertson Road, Centennial Park, Sydney.

Shipp, Hrik, 59 William Edward Street, Longueville, N.S.W.

Smith, Robert, B.Sc. (Agric.), Institute of Agriculture, University of Western Australia,
Nedlands, Western Australia.

Smith, Miss Vera Irwin, B.Sc., F.L.S., “Loana’’, Mt. Morris Street, Woolwich, N.S.W.

Smith-White, Spencer, B.Se.Agr., 7 Merriwa Street, Gordon, N.S.W.

Southcott, Ronald Vernon, M.B., B.S., 12 Avenue Road, Unley Park, Adelaide, South
Australia.

Spencer, Mrs. Dora Margaret, M.Sc. (née Cumpston), Moorabinda, Tenterfield, N.S.W.

Stanley, George Arthur Vickers, B.Sc., c.o. Messrs. Robison, Maxwell and Allen, 19 Bligh
Street, Sydney.

Stead, David G., ‘“Boongarre’’, 14 Pacific Street, Watson’s Bay, N.S.W.

Stevens, Neville Cecil, B.Sc., 12 Salisbury Street, Hurstville, N.S.W.

Still, Professor Jack Leslie, B.Sc., Ph.D., Department of Biochemistry, Sydney University,
N.S. W.

Stokes, Miss Anne, B.Sc., 45 Walkerville Terrace, Gilberton, S.A.

*Sulman, Miss Florence, 11 Ramsay Street, Collaroy, N.S.W.

XXXii

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3926
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1942

LIST OF MEMBERS.

Taylor, Keith Lind, B.Sc.Agr., ¢.o. Division of Entomology, C.S.I.R.O., Box 109, City,
Canberra, A.C.T.

Tchan, Yao-tseng, Dr., és Sciences (Paris), Botany School, Sydney University.

Thorp, Mrs. Dorothy Aubourne, B.Se. (Lond.), ‘‘Carinya’, Tara Street, Kangaroo Point,
Sylvania, N.S.W.

Thorpe, Ellis William Ray, B.Se., New England University College, Armidale, N.S.W.

Tindale, Miss Mary Douglas, M.Sc., 60 Spruson Street, Neutral Bay, N.S.W.

Tipper, John Duncan, A.M.I.E.Aust., Box 2770, G.P.O., Sydney.

*Troughton, Ellis Le Geyt, C.M.Z.S., F.R.Z.S., Australian Museum, College Street, Sydney.

Vallance, Thomas George, B.Sc., Department of Geology, Sydney University.

Veitch, Robert, B.Se., F.R.E.S., Department of Agriculture and Stock, William Street,
Brisbane, Queensland.

Vickery, Miss Joyce Winifred, M.Se., Botanic Gardens, Sydney.

Vincent, James Matthew, B.Sce.Agr., Dip.Bact., Faculty of Agriculture, Sydney University.

*Voisey, Alan Heywood, D.Sc., New England University College, Armidale, N.S.W.

Walkom, Arthur Bache, D.Sc., Australian Museum, College Street, Sydney.

Wallace, Murray McCadam Hay, B.Sc., Institute of Agriculture, University of Western
Australia, Nedlands, Western Australia.

Ward, Mrs. Judith, B.Se., 24 St. Alban’s Road, London, N.W.5, England.

Ward, Melbourne, Gallery of Natural History and Native Art, Medlow Bath, N.S.W.

Wardlaw, Henry Sloane Halcro, D.Se., F.A.C.1., 71 McIntosh Street, Gordon, N.S.W.

Waterhouse, Douglas Frew, M.Sc., C.S.I.R.O., Box 109, Canberra, A.C.T.

Waterhouse, John Teast, B.Se., ‘““Invermay”’’, Collarenebri, N.S.W.

Waterhouse, Professor Walter Lawry, D.Sc.Agr., M.C., D.I.C., Faculty of Agriculture,
Sydney University.

Watson, Irvine Armstrong, Ph.D., B.Se.Agr., Faculty of Agriculture, Sydney University.

Wearne, Walter Loutit, c.o. James Gillberg & Co., Suite 102, 243 Elizabeth Street, Sydney.

Whaite, Mrs. Joy Lilian, c.o. Mrs. D. Bailey, Tower Street, Panania, N.S.W.

Wharton, Ronald Harry, B.Sc., Institute of Medical Research, Kuala Lumpur, Malaya.

*Whitley, Gilbert Percy, Australian Museum, College Street, Sydney.

Wilkins, Miss Marjorie Jessie, M.Sc., 33 Muston Street, Mosman, N.S.W.

Willings, Mrs. Lucy May, B.A., C.S.I.R.O., Division of Fisheries, P.O. Box 21, Cronulla,
N.S.W.

Willis, Jack Lehane, B.Sc., 15 Muttama Road, Artarmon, N.S.W.

Wilson, Ralph Dudingston, M.Se., M.Se.Agr., Biological Branch, Department of Agricul-
ture, Box 36A, G.P.O., Sydney.

Winkworth, Robert Ernest, Botany Department, University of Melbourne, Carlton, N.3,
Victoria.

Womersley, Herbert, F.R.E.S., A.L.S., South Australian Museum, Adelaide, South
Australia.

Wood, Edward James Ferguson, C.S.I.R.O., Marine Biological Laboratory, P.O. Box 21,
Cronulla, N.S. W.

Woodhill, Anthony Reeve, D.Sc.Agr., Department of Zoology, Sydney University.

Zeck, Emil Herman, 694 Victoria Road, Ryde, N.S.W.
Zeck, Mrs. Nance (Anne), 694 Victoria Road, Ryde, N.S.W.

HONORARY MEMBER.

Hill, Professor James Peter, D.Sc., F.R.S.Z., F.Z.S., F.R.S., ‘Kanimbla’”’, Dollis Avenue,
Finchley, London, N.3, England.

CORRESPONDING MEMBERS.
Jensen, Hans Laurits, D.Se.Agr. (Copenhagen), State Laboratory of Plant Culture,
Department of Bacteriology, Lyngby, Denmark.
Rupp, Rey. Herman Montague Rucker, B.A., 24 Kameruka Road, Northbridge, N.S.W.

XXXiii

LIST OF NEW GENERA, SUBGENERA, AND SPECIES.
VoL. 76.

Genera and Subgenus.

Page.

Austroasca (Cicadellidae) .. .. .. 202

Flabellolaius (Malachiidae) Bio tiaecicaen si)

IEOONG (AMSUTOUSGD)) 55" o5-. 06. -n0 7Ale

Troglolaius (Malachiidae).. .. .. 188
Species.

Page. Page.
armstrongi (Troglolaius) .. .. .. 188 aL (GNGMEOTUO)) 56 Sse a6) bo ee 28S
bractigera (Austroasca) ae Mean eesti 4 ta) reidin @CUyrmozercon) <2 Baie. ecne. 155
lolii (Puccinia graminis) .. .. .. 63 tricalliatus (Sarcochilus) .. .. .. 54
merredinensis (Auwstroasca) Mma oho.

LIST OF PLATES.
PROCEEDINGS, 1951.

imWSeptoria pepli on leaves of Huphorbia peplus.
F iv.—Ceylonocotyle streptocoelium.

v._Ceylonocotyle streptocoeium, Paramphistomum ichikawai, Calicophoron
calicophorum.

vi.—Controlled Pollination of Hucalyptus.

vii-xii—Seedlings of Hucalyptus species and hybrids.

xiii Penaeid Prawn (Metapenaewus, n- sp.).

xiii—Effect of 2,4-dinitrophenol on Onion root tip cells.

xiv.—Use of Excised Shoots in Linseed investigations.

xv.—Stem of black nightshade (Solanum nigrum) .—Typical damage done by
feeding of Austroasca viridigrisea.

CORRIGENDUM.

The following lines were omitted under the heading “Notes and Exhibits” at the
meeting of the Society on 26th July, 1950, and should be inserted in the PROCEEDINGS,
Vol. lxxv, 1950, p. xxvii, following line 22:

Mr. A. K. O’Gower exhibited the eggs, larvae and pupae of cat fleas and pointed out
the possibility of high infestations in domestic situations from the presence of cats.
Two collections were made at a Dulwich Hill joinery works during the last week of
June and early July. The situations examined were the sleeping sites of two cats.
Four such sites were found, each being less than one square yard in area. From these
collections a total of some 500 eggs together with more than 8,000 larvae resulted.

XXXiV

INDEX.

(1951. )

Abstract of Proceedings .......... Xxili

Aédes (Stegomyia) aegypti Linn.,
and Culex (Culex) fatigans
Wied., Investigations on _ the
Preference shown by, for Specific
Types of Breeding Water ...... 149

Algae, Marine, of Brampton Island,
Great Barrier Reef, off Mackay,

Queensland nano sero eee 88
Algae, Australian Marine, Studies on,
VL ey dy baciceeae Cee eae. eos oie heck ie 83

Anatomy and Morphology of the
Operculum in the genus Huca-

LY DEUS. Lee oes hohe Stee ove Re ata Le 31
Athrotaxis, Some Notes on ........ 30
Australian Diptera, Miscellaneous

INOLES SONEGERV we vues ono e Gaoe 222
Australian Laius Guér., Remarks on

SOMVC: 5.12 eesick eters ane RES Ge 187
Australian Marine Algae, Studies on,

SVilig ch retsnemtene trent cyanea rates creases eis dercue 83
Australian Ruminants, The _ Par-

aM phiStoniesmotasl mere ener 41
Australian Rust Studies, viii ....... 57

Australian Species previously re-
ferred to the Genus Hmpoasca,
AMRVE VASTONNIO hacer cio tee 190

Baas-Becking, Prof. L. G. M—
Address given by
Hlectedwamivicmber essence ener XXVii

Backhouse, T. C., elected a Member xxiii

Balance Sheets for the Year ending

28th February, 1951 ........ XXi-xxii
Bearup, Av) J.” see) Hxhibits) vat
meetings.

Beattie, Joan, congratulations to .. xxiii

Bennett, Isobel I.—

Hlected ae Members. essences XXvi
See under Morris, Muriel C., and
Bennett, Isobel.

Brampton Island, Great Barrier
Reef, off Mackay, Queensland,
the Marimg Alzacyot tse nk 88

Britton, E. B., The Nomenclature of
Heteronychus sanctae-helenae

Blan chands etsacrcci ee ae 133
Broadcast lectures by Fred Hoyle
AND OUNCCHRI iene: ee ae ee XXV1i

Broom, Dr. R., reference to death of xxiv
Browne, Ida A., resignation as a

MembereotL Council vane ee li
Browne, W. R., elected Honorary
SEChOtany oa neeyeiican ke eee XXili

Bryozoan Faunas, The Development
of, in the Upper Palaeozoic of
AUSTTALTA ey Aree eer oe ee ae 105

Page.
Carne, P. B., Preservation Tech-
niques for Scarabaeid and other
Insect Warvaere sonic c serene 26
Catala, Dr. R., address given by ... xxvi
Celaenopsoides Gunther, 1942: A

Synonym of Ophiomegistus
Banks 14 ye lesseske arr ain ie ioe 65
Cheel, E., reference to death of .... xxv
Colefax, A. N.—
Hlected President ................ Sap

See Exhibits at Meetings.
Controlled Pollination of Hucalyptus 135
Critical Consideration of c-mitosis
with Reference to the Effects of
INMARODNATNOIS soo dacndcooceaccoe 158
Crockford, Joan (Mrs. Beattie), The
Development of Bryozoan Faunas
in the Upper Palaeozoic of
AUISUT ATT ARG ohcaics hci ani ee ee 105
Culex (Culex) fatigans Wied., Inves-
tigation on the Preference shown
by Aédes (Stegomyia) aegypti
Linn., and, for Specific Types of
IBGE Cdn eg Wiatela weir ie reer 149

Development of Bryozoan Faunas in
the Upper Palaeozoic of Aus-

Cray a hacer mete aoe 105
Diptera, Australian, Miscellaneous

NOteS Om; AxVics tenant eee eis 222
Durie, P. H., The Paramphistomes

(Trematoda) of Australian

EU UIITNITEDTICS SIU eye eene ene ete 41
Hlections Wigs aan eee Xix, XXili
Elliott, C. G@, Some Notes on

AER TOUOLIS. Crea aa ee eee 36
Emmons, Prof., short address given

BDO AEE i Re te DARE ae XXili

Empoasca, A Revision’ of the Aus-
tralian Species previously re-
ferred to the Genus ............ 190

Endean, R., congratulations to ...... ili

English, Kathleen M. I., see Exhibits
at Meetings.

Eucalyptus, Controlled Pollination of 135

Eucalyptus Species, A Genetic
ATIALY SIS OLSON GC marie teeter 140

Eucalyptus, The Anatomy and Mor-
phology of the Operculum in the

GenNists fli ends coroner rion tren anes 31
Euphorbia peplus L., A Septoria
Disease® of vile Ag RA ee 7

Evolution of the Radio-medial area

in the wings of the Muscoidea

A CALY DULALA. | 5.0 Sapte eieiays coene ernier noes til
xchange: relavions” a... cece i

INDEX.

Page.
Exhibits at Meetings:
Bearup, A. J.—a blood-film of an

-Australian gecko, Phyllurus
platurus (Shaw) collected at
Glen Davis, N.S.W. ..........- XXvi

Colefax, A. N.—a device for the
rapid moulding of metal foil
containers for paraffin embed-
GIT Spares pip oita,scitats sary 8 eaelawermerarnena sharos XXvVi

English, Kathleen M. I.—an intro-
duced slug, Testacella haliotidea xxv

Hindmarsh, Mary M.—a film of
mitosis in the reproductive cells
of the Grasshopper ............ XXvi

Mercer, F. V.—a lantern slide of a
caterpillar, exhibiting mimicry
of its surroundings to a remark-
DCG SS GEOC. Lois hy wshetsnegsciosve aecreuaye XXvi

Murray, Prof. P. D. F.—Specimens
of young turtles from the Great
Barrier Reef, stained with ali-
zarin and cleared to show
development of the bony skele-
ton; also fixed but. otherwise
untreated specimens for com-
[ORNPIS OTR ee aoa eae heater char ieeere mi portiesa or XXV

Musgrave, A.—specimens of the
Giant Orb-weaver, Nephila macu-
lata (Fabricius, 1793) from the
North Coast of N.S.W. ........ XXV

Robertson, R. N.—a preparation of
plant mitochondria extracted
from tissue of red beet and an
electron micrograph of mito-
GINCIACHPIE, =" Seats cameo cree es Oe ere XXvi

Fraser, A. S., elected a Member .... xxiii
‘Fraser, Judith, congratulations to .. xxiii

Fraser, Lilian, elected a Vice-
IERESIMEM be he vera eal Aes eee Xxili

“Galathea”’, Danish research frigate,
welcome to party of scientists
from, and lecture by Dr. A.
ESSIGUUUIN Ng yeic ehei sence ake ense sGacckelng Deen XXVii

Genera and Species, New, List of, in
NOEs Olen sever ncp gona slo elewoti ony eis f XXxili

Genetic analysis of some Hucalyptus
SPECIOSA ca cwct eh eet eo sat ee Tas 140

Gunther, C. H. M.—

A Mite from a Beehive on Singa-
DOG MUSTAM CP eckee hia arse ae 155
Celaenopsoides Gunther, 1942: A

Synonym of Ophiomegistus

Banikss Fe MGW A cy ys spel ee bes ho share et 65
On Trombicula minor Berlese,

TUSK SF aiero eid orcleltesrs cre oicttac ior it baaeenene 66

Hardy, G. H., Miscellaneous Notes on
Australian Diptera. xv. Tabanus,

HReterOpSilOpuUs *.......26+-++--- 222
Hardy, Dr. Margaret, short address
VETO VE iontns ciercuararsreio tens ce anus Xxili

Harker, Janet, congratulations to iii, xxiv

Heteronychus sanctae-helenae Blan-
chard, The Nomenclature of .... 133
Hewitt, B. R., elected a Member ... xxiv

XXXV

Page.
Hindmarsh, Mary M., Linnean Mac-
leay Fellow in Botany—
A Critical Consideration of c-
mitosis with Reference to the

Effects of Nitrophenols ........ 158
Reappormbedseal9 a0 aac cioeierscen ili
WORK Ob ei Ca Pate TER nr ae iii
Reappointed, 1951 ............... iv
Reappointedy e952) asec ane xxvii

See Exhibits at Meetings.
Humphrey, G. F., congratulations to xxiv

Insect Quarantine, The Problems of v

Investigations on the Preference
shown by Aédes (Stegomyia)
aegypti Linn., and Culex (Culex)
fatigans Wied., for Specific
Types of Breeding Water ...... 149

Johnston, Prof. T. H., reference to

Cleat nOty sae tace ai: oe coaeerra nu ctoa cies XXV
Jubilee Celebrations, co-operation
with other scientific bodies re .. iii
Kerr, H. B.—
Elected a Member .............. Xxili
The Use of Excised Shoots in
Linseed Investigations ......... 183
Kiely, T. B., congratulations to .... iii

Kosciusko area, lecturettes given by
members of expedition to the .. xxiv

Laius, Guér., Remarks on some Aus-

Creal mays eye Mee eee eet 187
Lancaster, Helen P., elected a
Miennlb@iys 2.5 50) ers Pe RS XXV
Lee, D. J.—
Elected a Vice-President ........ XXili

Short summary of recent observa-
tions and the importance of
entomological work in connection
with outbreaks of myxomatosis
and encephalitis in Hastern Aus-
trralliiayiey ts er ee! a Pe Su 2 XXvVii

The Problems of Insect Quaran-
tine (Presidential Address) .... Vv

Library accessions ...:.... i, Xxili—xxvil

Liddell, Jean, congratulations to .... iii

Life-history of a Penaeid Prawn
(Metapenaeus) breeding in a

Coastalivlake) v5 jews hey orn poe erate oy. 164
Linnean Macleay Fellowships—
Reappointments for 1950 ......... ili
Reappointments and appointments
LOTS Hae Ae ee Mik raen el tae een A eee iv
Applications for 1952 invited xxv, xxvi
Reappointments for 1952 ....... XXVii
Linseed Investigations, The Use of
JDPCONSECL SIMOOUS Th Socccascouo 183
Lower, H. F.—

A Revision of the Australian
Species previously referred to
the Genus Hmpoasca ........... 190
The Evolution of the Radio-medial
Area in the Wings of the Mus-
coidearAcaliy pinata. a. cele roel 71
Lowery, E. S., elected a Member ... xxiv

XXXV1 INDEX.

Page.
Macdonald, C. L., elected a Member xxvii
Mackerras, I. M., congratulations to iii
Macleay Bacteriologist, see under
Tchan, Yao-tseng.
Macleay, Sir William, reference to
60th anniversary of death and
exhibition of personal relics .. Xxvii

Manefield, T., Investigations on the
Preference shown by Aédes
(Stegomyia) aegypti Linn., and
Culex (Culex) fatigans Wied.,
for Specific Types of Breeding
Wee Trea tary tres eae ohana ea saekewaite 149

Marine Algae of Brampton Island .. .88

May, Valerie— ;

Studies on Australian Marine
Nev aeen fl Oe Seen Gack rrimig Gab or ee Gra oA 83

The Marine Algae of Brampton
Island, Great Barrier Reef, off

Mackay, Queensland ........... 88
McAlpine, D. K., elected a Member xxiv
MembersrnlaiSGrOl caer cine cto ely XekoValilil
Mercer, F. V.—

AQGEeSS T2iIVeN ID) \ceslee eaaletee cae. XXVi
Elected a Member of Council .... xxiv

See Exhibits at Meetings.
Millerd, Adele, Linnean Macleay
Fellow in Bio-chemistry—

Consranulationsy stones rei. cai. otek iii
Reappoiniedawl9 50s telcrrnpiicr rae iii
IRYASIATEVINGIN OE “Soba oac boc eoabooos ili
AWiO Tt O fa ieee yr ence reese oe een ier,
Succinoxidase of Potato Tuber ... 123

Miscellaneous Notes on Australian
Diptera. xv. Tabanus, Heterop-

SULOD US tats See Rhee Ae Ot epee 222
Mite from a Beehive on Singapore
Ts Ven Gee pe cea ee nro sce anata re 155

Morris, Muriel C., Linnean Macleay
Fellow in Zoology—

Coneratnlarionshitomae. eee ili
Rieappomtedh s195 Oiee chris cenit lii
Resiematilom FOL eae. wore taere ore iv
Wonk Ob ein ahr is SR eet iv

Morris, Muriel C., and Bennett,
Isobel, The Life-history of a
Penaeid Prawn (Metapenaeus)
breeding in a Coastal Lake
_(Tuggerah, New South Wales) 164

Muogamarra Sanctuary ............ ili
Murray, Prof. P. D. F.—
Elected a Member of Council .... ii

See Exhibits at Meetings.

Muscoidea Acalyptrata, The Evolu-
tion of the Radio-medial Area in

ae Ay beets Giese an po necoe caus Tak
Musgrave, <A., see Exhibits at
Meetings. :
Natural Areas, Preservation of ..... ii

Nitrophenols, A Critical Considera-
tion of c-mitosis with Reference

LOMEME MELE CES MOL \si.,6 cect a mawumeele one 158
Noble, R. J., congratulations to ... xxiii
Nomenclature of Heteronychus

SONCUMETUCLET MC ree iiie.6 eye ae 133

Page
Obituaries:
ProfessOrmwe tdi DWakink ewe eter iv

DrwbywiamViiddletoneen. ties eer Vv

Osborne, G. D., elected a _ Vice-
IPTeSidenitae tA fee. ace XXili

Packham, G. H., elected a Member .. xxvi
Paramphistomes (Trematoda), The,

of Australian Ruminants. i.... 41
Plates ee TStaOL see ce ee ee ReSeRA IT
Potato Tuber, Succinoxidase of .... 123

Prawn, Penaeid (Metapenaeus), The
Life-History of a, breeding in a
Coastal Lake (Tuggerah, New

SOUtH*AWalles! ieee) on onhy come come ee 164
Preservation Techniques for Scara-
baeid and other Insect Larvae .. 26
Presid enrialarAd dress see eee i
Problems of Insect Quarantine ..... Vv
Pryor ae)
A Genetic Analysis of some
Hucalyptus Species ............ 140
Controlled Pollination of Huca-
LUDIELUS Merch gts DURA Cena 27. 1k neat 135
Purchase, Hilary F., congratulations
LOPATA Aaa a satay Sete meus here SEC EN aera XXiV

Purchase, Hilary F., Vincent, J. M.,
and Ward, Lawrie M., Sero-
logical Studies of the Root-
nodule Bacteria. iv. Further
Analysis of Isolates from Tri-
folium and Medicago

Ralph, B. J. F., elected a Member .. xxiii
Review of the Australian Species of
SQECOCKILUSH ee a ee 49
Revision of Australian Species pre-
viously referred to the genus
IBGGNOUSGOS “Soa boes ddo6ds soa Gococ 190
Robertson, R. N.—
Elected a Vice-President

5 the aks ofr Xxili
See Exhibits at Meetings.

Root-nodule Bacteria, Serological
SUOMI OWASP Soccccccnescce
Rule svar ad cditionmonen ean ii

Ruminants, Australian, The Param-
phistomes (Trematoda) of, i .. 41

Rupp, H. M. R., A Review of the
Australian Species of Sarco-
chilus, (Orchidaceae)... 23.025. 49

Rust Studies, Australian, viii ...... ST
Sarcochilus (Orchidaceae), A Review

of the Australian Species of .... 49
Searabaeid and other Insect Larvae,

Preservation Techniques for ... 26
Science House, re net return, annual

meetings and survey of libraries

Ofmowmenr-bodicSmeaceMal ee ee ee ii
Seccombe, Mrs. C. E., elected a

avaMiember (tvs. 2a Gu oerareiee xxiii
Secretary, resignation of .......... li
Septoria disease of Huphorbia peplus

JRE p Sieen SPE Maca aoe anes aire Pie tps 1

Serological Studies of the Root-nodule
Bacteria. iv. Further Analysis
of Isolates from Trifolium and
MI COUCHG OLA ioh rere tate ote hss cetietol ae 1

INDEX.

Page.

Shaw, Dorothy E., A Septoria Disease
of Huphorbia peplus lL. ........
Special General Meetings
Stevens, N. C., appointed Linnean
Macleay Fellow in Geology, 1951
Studies on Australian Marine Algae.
Ns eR seas Fe GI en vitars Pec US BUR
Succinoxidase of Potato Tuber .....
Synopses of papers to be published
below titles commencing 1951

Tchan, Yao-tseng, Macleay Bac-
teriologist—

Arrival in Sydney and commence-

TNE THE OTWiOIvKey nea ees eee
NOC TONE SS aie ie ena omni Aiea ome
Tindale, Mary 1) appointed
Society’s representative at 7th
International Botanical Con-

gress, 1950

Upper Palaeozoic of Australia, The
Development of Bryozoan Faunas
in the

Use of excised
investigation

shoots in linseed

iV

ii

105

XXXViil

Page.
Vallance, T. G.—
Appointed Linnean Macleay Fellow
iin “(Greoloray; WGIL, goto ooancoeonde iv
IRGEVO OUMNH, IUOE~A Sseoobesoobuoo XXVli
Vincent, J. M., see Purchase, Hilary

He Vincent ards Vie, lands Ward:
Lawrie M.
Walkom, <A. B., elected Honorary
AIRE AS URC Te eiy sh ta tense ui a ames Xxili
Ward, Lawrie M., see Purchase,
EtilanvachsVancent.Js VME vanid

Ward, Lawrie M.
Waterhouse, W. L., Australian Rust

SLUTS Sevilla eae eters ora. 57
Wild flowers and _ native plants,
renewal of proclamation for pro-
Fechlions Of —“SUpPpPOGLEds sess. ere ili
Willis, J. L., The Anatomy and Mor-
phology of the Operculum in the
Genus Hucalyptus. 1 ..5....... 31

Wittmer, W., Remarks on some Aus-
tralian! saws) Gueiws. a. one see: 187

Woodhill, A. R.—
Congratulations to
Elected a Vice-President

ba ‘ - i ’ v
2 d ‘ 4 ie: i ee 3 ©.
= i ee { js .
_ ae - 4 iy
. Pht LY + tain ay fl eat
; ik eee a? ayers : i Pee andsayol: vi
sist Lon aatoT a gee pnieanas ath
; ys a PR teoy yt ve > 5 wat: path Heiss fat)

i‘ AP Waste >, wr atu in 8 oy ta. ‘ ye Gis Fees’ ae « “seer

: hs : : : ae ae ee ‘ ¢ : heat tate eh Tere oper
eat? . ' : yn Leet RT ROLE Aidatt By a A

; a eteers Ws ain SP 7 Pa : Whe oe ar Oa uf wile peter ay ee

Fes pet me nea

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tai os eae ‘i
an e ¥

ren rope

44 av o he 3 tet aif

' ey Paley The We’ a

: bee hie cL ted ar its pee
syria igs | Pd rwckested

tis “Lagi Lagi
ng os Aye, ca

an Bae a ys ae

ny pa ee) od 2

“aed ae
wd 1 Bas oes

eee

axa & Sas >

Oa Ra

Beppe - oes
cer vad

Proc. Linn. Soc. N.S.W., 1951. PLATE XII.

Penaeid Prawn (Metapenaeus, n. sp.).

uf

‘ jl a 7A ‘ 2s sf 7 af

bts 4 i

8 ; : i |

'
Leh Cee e
re Sots eo i” Oe; v ne ;
as ‘ i , —_ 7 A e ‘i a
g° - 733? Se F
‘= e ty

= Fj .
=
eg j ;
\ * ie ul :
i . ‘! ' ’
.
hacks
' ‘. i ; 7
i:
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; .
. “
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‘ # = = 7 a
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Te) 4

er RM ae me

eT
.
rs
x
ms x

ee
ar

Proc, Linn, Soc, N.S.W., 1951, PLATE XIIT

Uo a2

Effect of 2,4-dinitrophenol on Onion root tip cells,

.

ar at

Te ul ee, Ni
siamese

PLATE XIv.

Proc. Linn. Soc. N.S.W., 1951.

igations,

seed Invest

in

ised Shoots in Li

Hxe

Use of

*\,
,

Tt

. ¥ Ki ak ef
OHTA tae

PLATE xv.

Proc. Linn. Soc. N.S.W., 1951.

DISLIB VPA

DISHOLISNF JO

suIpesy Aq suop asvUep [BoId AT,

“CWUNnsHYU WNUWYDIOS )
epeys JYUSIU Yyoulq Jo ways jo uolIy1Og

_ (Issued 81st May, 1951.) ;

foc= Jo

Vol, LXXVI. 55: ene l
Parts 1-2. 1 Marine Biological Labom
LIBRA RR

THE MAY 5 - 1959I|
: PROC 5 E DI N S WOODS HOLE, a
LINNEAN SOCIETY

New SoutH WALES

FOR THE YEAR

1951.

Parts 1-2 (pp. i-«x; 1-48).
CONTAINING THE PROCEEDINGS OF THE ANNUAL GENERAL
MEETING AND PAPERS READ IN MARCH-APRIL.
With five plates.
[Plates i-v.]

SYDNEY:
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by post as a periodical. :

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David Nutt, 212 Shaftesbury Avenue, London, W.C.2.

fie nia Society of New South Wales

LIST OF OFFICERS AND COUNCIL, 1951-52.

President: :

A. N. Colefax, B.Sc.
Vice-Presidents: ‘
G. D. Osborne, D.Sc., Ph.D. _ R. N. Robertson, B.Sc., Ph.D.
Lilian Fraser, D.Sc. D. J. Lee, B.Sc.

Hon. Treasurer: A. B. Walkom, D.Sc.

Hon. Secretary: W. R. Browne, D.Sc. :

Council: .

R. H. Anderson, B.Sc.Agr. G. D. Osborne, D.Sc., Ph.D. ¢
W. R. Browne, D.Sc. R. N. Robertson, B.Se., Ph.D.
Professor N. A. Burges, M.Sc., Ph.D. T. C. Roughley, B.Sc., F.R.Z.S.
A. N. Colefax, B.Sc. E. Le G. Troughton, C.M.Z.S., F.R.Z.S.
Ss. J. Copland, B.Sc. J. M. Vincent, B.Se.Agr., Dip.Bact.
Lilian Fraser, D.Sc. A. B. Walkom, D.Sc.
Professor J. Macdonald Holmes, B.Sc., H. S. H. Wardlaw, D.Se, F.A.C.I.

Ph.D., F.R.G.S., F.R.S.G.S. : Professor W. L. Waterhouse, M.C.,
D. J. Lee, B.Sc. D.Sec.Agr., D.I.C.
Professor P. D. F. Murray, M.A., D.Sc. A. R. Woodhill, B.Sec.Agr.

Auditor: S. J. Rayment, F.C.A. (Aust.).
NOTICE.

Complete sets of the Proceedings of the Linnean Society of New South Wales (the
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Separate Parts may be obtained also from David Nutt, 212 Shasta ee Avenue, London,
W.C.2. x

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. Authors are requested to assist in facilitating the Society’s publishing work by observing
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Manuscript. —Manuscripts should be double-space typed with a margin of at least one inch on
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BuLuoueH, W. S., 1939.—A Study of the Reproductive Cycle of the Minnow in Relation
to the Environment. Proc. Zool. Soc. Lond., 109, A, Pt. 1: 79-108.

Abbreviations.—Standard abbreviations should be used in tabulations and after numerals in the
text. The abbreviations of names-of periodicals should conform to those in the World List
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. ‘Tabulations. —Tables should be maapeced consecutively and referred to specifically in the text
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Proofs should be corrected and returned as soon as possible together with an order for any
reprints required in addition to the 50 supplied gratis by the Society.

=

PROCEEDINGS, LXXVI, PARTS 1-2, 1951.

CONTENTS.
Pages
Presidential Address, delivered at the Seventy-sixth Annual General Meeting,
28th March, 1951, by D. J. Lee, B.Sc. The Problems of Insect Quarantine .. i-xix
LE CCLOMS HCO iy et casi Ie GEL SI fea a ees ieee a te luke te BURG eti rd MCR IESE ANN AVN MDE LO eR Rs ae xix
Balance Sheet for the Year ending 28th February, 1951 .. .. .. .. .. .. xx

Serological Studies of the Root-nodule Bacteria. IV. Further Analysis of —
Isolates from Trifolium and Medicago. By Hilary F. Purchase, J. M. -
Vincent and: Lawrie (Me Ward. ite ys oe end ease ee Ee Cet gene Molt aNtoaee ade a 1-6

A Septoria Disease of Huphorbia peplus L. By Dorothy E. Shaw. ‘(Plate i and
OWE DEKE ME UTES) ye aye shes eee Ne ates Sida e alae ah Seer Ear Wea ec icaaM Sr tiny He feo ae Jo a 7-25

Preservation Techniques for Scarabaeid and other Insect Larvae. BY SPy ea:
Carne Pacis eA MA I CAIN Nn tag LGU ANI a es a eS eR 26-30

The Anatomy and Morphology of the Operculum in the Genus Hucdlyptus.
Part I. The Occurrence of Petals in Hucalyptus gummifera (Gaertn.)
Hochr. By. J. L. Willis. (Plates ii-iii and two Text- ‘figures. ) Sipe es Ae ete 31-35

Some Notes on Athrotaxis. By Charles G. Elliott. (Communicated by Dr.
Pairick Brough.) (Sixteen: Téext-feures:) 205) cee ieee ate cee vee ele ee 36-40

The Paramphistomes (Trematoda) of Australian Ruminants. Part I.
systematics, | By P: A. Durie: OCPIAtEs VV) i ivatak ih i) fe tin wae ake Nien ae 41-48

“

Corrigendum.

The following lines were omitted under the heading “Notes and Exhibits” at the
meeting of the Society on 26th July, 1950, and should be inserted in the PROCEEDINGS,
Vol. Ixxv, 1950, p. xxvii, following line 22:

Mr. A. K. O’Gower exhibited the eggs, larvae and pupae of cat fleas and pointed out
the possibility of high infestations in domestic situations from the presence of cats.

Two collections were made at a Dulwich Hill joinery works during the last week of

June and early July. The situations examined were the sleeping sites of two cats.
Pour such sites were found, each being less than one square yard in area. From these
collections a total of some 500 eggs together with more than 8,000 larvae resulted.

| Nos. 355-386.
| Po

INNEAN SOCIETY

New SoOuTH

FOR THE YEAR

1951.

Parts 3-4 (pp. 49-157).
CONTAINING PAPERS READ IN JUNE-AUGUST.

With six plates.
a ¢ [Plates vi-xi.]

- SYDNEY:

PRINTED AND PUBLISHED FOR THE SOCIETY BY
AUSTRALASIAN MEDICAL PUBLISHING CO., LTD.,
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and
“SOLD BY THE SOCIETY,

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x

Registered at the General Post Office, Sydney, for transmission
by post as a enos ed:

Agent in Europe:
David Nutt, 212 Shaftesbury Avenue, London, W.C.2.

WOODS HOLE, MASS.

The Laaneah Society of New South Wales

LIST OF OFFICERS AND COUNCIL, 1951-52.

President:

A. N. Colefax, B.Sc.
Vice-Presidents:
A. R. Woodhill, D.Sc.Agr. Lilian Fraser, D.Sc.
G. D. Osborne, D.Sce., Ph.D. D. J. Lee, B.Sc.

Hon. Treasurer: A. B. Walkom, D.Sc.
Hon. Secretary: W. R. Browne, D.Sc.

Council: :

R. H. Anderson, B.Sc.Agr. Professor P. D. F. Murray, M.A., D.Sc.
W. R. Browne, D.Sc. G. D. Osborne, D.Sc., Ph.D.
Professor N. A. Burges, M.Sc., Ph.D. T. C. Roughley, B.Sc., F.R.Z.S.
A. N. Colefax, B.Sc. ; E. Le G. Troughton, C.M.Z.S., F.R.Z.S.
S. J. Copland, B.Sc. J. M. Vincent, B.Sc.Agr., Dip.Bact.
Lilian Fraser, D.Sc. A. B. Walkom, D.Sc.
Professor J. Macdonald Holmes, B.Sc., H. 8S. H. Wardlaw, D.Sc., F.A.C.I.

Ph.D., F.R.G.S., EF.R.S.G.S. Professor W. L. Waterhouse, M.C.,
D. J. Lee, B.Sc. D.Se.Agr., D.1.C.
*H, V. Mercer, B.Sc., Ph.D. A. R. Woodhill, D.Sc.Agr.

Auditor: S. J. Rayment, F.C.A. (Aust.).
* Elected 18th July, 1951, in place of Dr. R. N. Robertson, resigned.

NOTICE. ee

Complete séts of the Proceedings of the Linnean Society of New South Wales (the
stock of the First Series, Volumes I-VI, 1875-1881, being limited), and separate Parts, with
the exception of Volume II, Part 4, Volume V, Part.2, and Volume VI, Part 4, of the First
Series, may be purchased from the Society, Science House, 157 Gloucester Street, Sydney.
Separate Parts may be obtained also from David Nutt, 212 Shaftesbury Avenue, London,

W.C.2.

Year. Volume. Part 1. | Part 2. | Part 3. | Part 4. Year.| Volume. | Part 1. | Part 2. | Part 8. | Part 4,

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1879 IV 6 0 6 0 8 0 6 6. 1902 XXVII|} 7 0 7/6 7 6° |} 15 O
1880 Vi 6 6 — 7 6 7 6 1903 XXVIiI 9.0 | 1246 14 0 15 0
1881 VI 6 0 10 0 10 0 _ 1904 XXIX | 10 O 7 6 9 0 10 0
1882 VII 7 6 10 0 a) 10 0 1905 xXX 6 04 /}10 0 12 6 8 0O5
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with portrait, and forty-two plates. Price £2 2s.
DEscRIPTIVH CATALOGUD OF AUSTRALIAN FISHES. By William Macleay, F.L.S. [1881]. A few

eopies only. Two volumes and supplement. Price £2 2s.
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u
Bh
a

BY CARL E. M. GUNTHER. 157

plate; covered with sparse short plain setae. Metapodal plates roughly triangular, their
apices behind coxae iv, their bases at the lateral margins of the body; with a few short

plain setae over the bases and the adjacent integument.

Coxa i bearing a stout tapering plain seta, 160u long, projecting straight forward

from, a shoulder on the anteromedial angle; a shorter, stout seta projecting medially,

and a group of five stout setae on the lateral side. Coxa ii with one stout seta towards
its anterior margin and a group of six posteriorly. €oxa iii with one anterior and five
posterior setae. Coxa iv with four stout setae. Six segments in each leg, but the
last two segments in leg i fused, and partly fused in the other three legs. Legs short,

‘very stout, normally curved postero-ventrally; i, 500u; ii, 5254; iii and iv, 600 long.

Postero-ventral aspect of last five segments of each leg with a thickened plate. All
segments with several stout sharp setae; no spurs on any leg. Tarsus i longer than
the others, straighter and more slender; tarsi ii, iii and iv short, stout, curved. A large
earuncle, but no claws, on each tarsus (Text-fig. 2). -

Hpistome only visible from above when proboscis projected; a small, translucent

_ semicircle (Text-fig. 5). Mandibles with three segments, scissors-shaped, the medial fixed

blade with a row of minute teeth; a small flagellum at the lateral side of the base of
the blades (Text-fig. 3). Hypostome bifid, each half bearing three short stout forward-
pointing setae; lingula bifid, each point bearing three small rounded projections on its
medial surface (Text-fig. 4). Palpi of five segments, with a few fine setae on segments
iii and iv; terminal segment rounded, with one short sharp stout seta on its disto-medial
aspect, and covered with several fine straight setae.

Type at the British Museum; paratype at the School of Public Health and Tropical

‘Medicine, University of Sydney.

PROCEEDINGS, LXXVI, PARTS 3-4, 1951.

CONTENTS.

A Review of the Australian Species of Sarcochilus (Orchidaceae). By H. M. R.
Rupp. (One Text-figure.) .. Sa MCT tae Li
Australian Rust Studies. VIII. Puecinia graminis lolii, an Undescribed Rust
of Lolium spp., and Other Grasses in Australia. By W. L. Waterhouse ..

Celaenopsoides Gunther, 1942: A Synonym of Ophiomegistus Banks, 1914
(Acarina: Parasitidae). By Carl E. M. Gunther :

On Trombicula minor Berlese, 1905. By Carl E. M. Gunther. (Three Text-
figures. )_ Pi enema inch oe Ga ner DINAH SIA PA.

The Evolution of the Radio-medial Area in the Wings of the Muscoidea.

Acalyptrata (Diptera). By H. F. Lower. (Thirty Text-figures.)

Studies on Australian Marine Algae. VI.

Certain Species. By Valerie May ..

New Geographical Records of

The Marine Algae of Brampton Island, Great Barrier Reef, off Mackay,
Queensland. By Valerie May .. ERP obesity Mts, oo ; rete

The’ Development of Bryozoan Faunas in the Upper Palaeozoic of Australia.
By Joan Crockford (Mrs. Beattie.) (Four Text-figures.)

Succinoxidase of Potato Tuber. By Adele Millerd. (Two Text-figures. ) ts

The Nomenclature of Heteronychus sanctae-helenae Blanchard (Coleoptera:
Scarabaeidae: Dynastinae). By E. B. Britton. (Communicated by P. B.
Carne.) Rhea ee Rests races ani le tele race Ua aR Vo atom Rp a Mea

Controlled Pollination of Hucalyptus. By L. D. Pryor. (Plate vi.) ..

A Genetic Analysis of some Hucalyptus Species. By L. D. Pryor. -(Plates
vii-xi.)

Investigations on the Preference shown by Aédes (Stegomyia) aegypti Linn.,
and Culex (Culex) fatigans Wied., for Specific Types of Breeding Water.
By Tom Manefield By i Ag te LRM ee kt a RCAC T Eas ae it Loe ea sires ;

@

A Mite from a Beehive on Singapore Island (Acarina: Laelaptidae). By Carl
E. M. Gunther. (Five Text-figures.) Pia mea aN age bea pt

. 105-122

Pages.

49-56

57-64

co
66-70
71-82 4
83-87

88-104.

123-132

133-134

135-139

140-148

149-154

155-157

tle allt —
s

(Issued 15th January, 1952.) , ad

Vol. LXXVL | y Nos. 357-358. |!
PROCEEDINGS
LINNEAN SOCIETY

NEW SOUTH

Marine Biological Laboratory!

FOR THE YEAR a ESR A ER x,

1951.

MAY 2 6 1°53
WOODS HOLE, MASS.

Parts 5-6 (pp. 157-225; xxi-

CONTAINING PAPERS READ IN SEPTEMBER-NOVEMBER,
ABSTRACT OF PROCEEDINGS, LIST OF MEMBERS
AND INDEX.

With four plates.
[Plates xii-xv.]

SYDNEY:

PRINTED AND PUBLISHED FOR THE SOCIETY BY
AUSTRALASIAN MEDICAL PUBLISHING CO., —
Seamer Street, Glebe, eyduey:
and
SOLD BY THE SOCIETY.

1952.

Registered at the General Post Office, Sydney, for transmission
- by post as a periodical.

Agent in Europe:
David Nutt, 212 Shaftesbury Avenue, London, W.C.2.

The Linnean Society of New South Wales

LIST OF OFFICERS AND COUNCIL, 1951-52.

President:
A. N. Colefax, B.Sc.

Vice-Presidents:
A. R. Woodhill, D.Se.Agr. Lilian Fraser, D.Sc.
G. D. Osborne, D.Sc., Ph.D. D. J. Lee, B.Se.

Hon. Treasurer: A. B. Walkom, D.Sc.
Hon. Secretary: W. R. Browne, D.Sc.

Council:

R. H. Anderson, B.Sc.Agr. Professor P. D. F. Murray, M.A., D.Sc.
W. R. Browne, D.Sc. G. D. Osborne, D.Se., Ph.D.
Professor N. A. Burges, M.Sc., Ph.D. T. C. Roughley, B.Sc., F.R.Z.S.
A. N. Colefax, B.Sc. E. Le G. Troughton, C.M.Z.S., F.R.Z.S.
S. J. Copland, B.Sc. J. M. Vincent, B.Sc.Agr., Dip.Bact.
Lilian Fraser, D.Sc. A. B. Walkom, D.Sc.
Professor J. Macdonald Holmes, B.Sc., H. S. EY Wardlaw, D:Se., F:A.C.L

PhD.) HRiG.S. RS: G:s: Professor W. L. Waterhouse, M.C.,
D. J. Lee, B.Sc. DISGAS DEG
*K. V. Mercer, B.Sc., Ph.D. A. R. Wocdhill, D.Sc.Agr.

Auditor: S. J. Rayment, F.C.A. (Aust.). ;
* Blected 18th July, 1951, in place of Dr. R. N. Robertson, resigned.

NOTICE.

Complete sets of the Proceedings of the Linnean Society of New South Wales (the
stock of the First Series, Volumes I-VI, 1875-1881, being limited), and separate Parts, with
the exception of Volume II, Part 4, Volume V, Part 2, and Volume VI, Part 4, of the First
Series, may be purchased from the Society, Science House, 157 Gloucester Street, Sydney.
Separate Parts may be obtained also from David Nutt, 212 Shaftesbury Avenue, London,

W.C.2.

Year.| Volume. Part 1. | Part 2. | Part 3. | Part 4. Year.| Volume. Part 1. | Part 2. | Part 3. | Part 4.
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1878 Ill By) 5 0 6 0 " 1901 XXVI/}10 O 9 0 5 0 LNG
1879 IV 6 0 6 0 8 0 6 6 1902 XX VII Ge © LE f-6*) Th aK0
1880 V 6 6 — 0 (nn! 1903 XXVIII 9 0 12-6 14 0 15 0
1881 VI 6 0 10 0 10 0 _ 1904 XXIX | 10 O if AD 9 0 10 0
1882 VII Tf Gs 10 0 by) 10 0 1905 xXxXX 6 0* 10) 0 12 6 8 05
1883 Vili | 10 O 5 0 TanO) 8 0 1906 XXXI/} 12 6 1G 12) 6 15 0
1884 Ix 8 0 12 0 25 0 25 0 1907 XXXII 8 0 8 6 15 0 18 0
1885 X12 0 fo @ 150 6 1908 XXXII nO 9 0 14 0 12 6
1886 If| 10 6 120 13 0 12 6 1909 XXXIV} 12 O Li 0 14 0 16 6
1887 II rhe 3 @ 12 0 PAW) 1910 10 il) 0 12 6
1888 MAL |), Thy 24 0 20 0 18 0 1911 XXXVI 9 6 9 6 9 6 10 0
1889 inte eat 5a) 16 0 19 0 LO) 1912 XXXVII 8 6 2ban0 L276 15 0
1890 WaieliiierO 9 0 9 0 9 0 1913 XXXVIII | 14 0 iG 6 0 13 0
1891 WAP a (0) 9 6 ile) ane 1914 XXXIX | 13 0 7-0 Zo 19 0
1892 VII 6 6 4 6 8 0 8 0 1915 XL /|17 0 12796 10 0 168
1893 VIII Dia it’ @) 6 0 9 0 1916 XLI } 10 0 12 0 15 0 19 0
1894 EX | 12 0 12 0 on0 8 0 1917 XLII | 14 0 90 12 6 16 6
1895 ; >] alla Ay) 8 6 10-0? | 12 0 1918 XLITI | 20 0 14.0 21 0 19 O
1896 XXI oo? 6 6 TaG 27-16 1919 XLIV | 12 6 inh Livia 6. 13 0
1897 XXII | 10 O 8 6 9 0 12 6 1920 XLV /}10 9 7°6 9 O spent)
1898 XXIII 3 0 6 0 12 0 14 0 1921 XLVI 9 0 8 0 nO 9 6

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Notice To AUTHORS. P
Authors are requested to assist in facilitating the Society’s publishing work by observing
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BuLLouGH, W. S., 1939.—A Study of the Reproductive Cycle of the Minnow in Relation
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PROCEEDINGS, LXXVI, PARTS 5-6, 1951.

CONTENTS.

A Critical Consideration of c-mitosis with Reference to the Effects of Nitro-

phenols. By Mary M. Hindmarsh, Linnean Macleay Fellow in Botany.
(Plate xiii.)

The Life-history of a Penaeid Prawn (Metapenaeus) breeding in a Coastal
Lake (Tuggerah, New South Wales). By Muriel C. Morris and Isobel
Bennett. (Plate xii and ninety-six Text-figures. )

The Use of Excised Shoots in Linseed Investigations. By H. B. Kerr.
(Plate xiv.)

Remarks on some Australian Laius Guér. (Col.: Malachiidae). By W.
Wittmer. (Communicated by J. W. T. Armstrong.) (One Text-figure.) ..

A Revision of Australian Species previously referred to the Genus Hmpoasca

(Cicadellidae, Homoptera). By Harold F. Lower. (Plate xv and seventy-
five Text-figures. ) 4

Miscellaneous Notes on Australian Diptera. XV. Tabanus, Heteropsilopus.
By G. H. Hardy

Balance Sheets
Abstract of Proceedings

List of Members

Pages.

158-163

164-182

183-186

187-189

190-221

222-225

XXi-XxXli

XXili-xXxvli

XXVili-xxxii

List of New Genera and Species ..
List of Plates

Index

XXXiii

XXXili

. XXXiv—-xxxvii

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