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oF THE

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NEw Sousa VV ALES

FOR THE YEAR

1949

VOL. LXXIV.

WITH FOUR PLATES.
207 Text-figures.

SYDNEY:
PRINTED AND PUBLISHED FOR THE SOCIETY: BY

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

and
SOLD BY THE SOCIETY.
1949.

ii

CONTENTS OF PROCEEDINGS, 1949.

PARTS 1-2 (Nos. 341-342).
(Issued 15th June, 1949.)

CONTENTS.
Pages.
Presidential Address. Seventy-fourth Annual General Meeting: A Gummosis
Disease of Citrus in Relation to its Environment. By Lilian Fraser, D.Sc. i-xviii

Elections Bea, BOG ache Gee, AS yen At es aie Vib cr ool ca ern a CAS Rae aT eRe Re xix
Balance Sheets for the Year ending 28th February, 1949 itr a an Hien nce ee SRORS NCR T

Australian Formicidae. New Genera and Species. By J. J. McAreavey, S.J.
(Seventy Text-figures.) SO OR aoe Ip lectin) AC MUS Og, 2 (cic are Rea arta ee ae 1-25
The Distribution of Formic and Alcohol Dehydrogenases in the Higher Plants,
with particular Reference to their Variation in the Pea Plant during its
Life-cycle. By Daphne C. Davison, M.Sc. (Nine Text-figures.) .. .. .. 26-36

The Importance of Formic Dehydrogenase in the Oxidation Mechanisms of
Pisum sativum. By Daphne C. Davison, M.Sc. (Two Text-figures.) .. .. 37-56

On Australian Species of Creophilus (Coleoptera: Staphylinidae). By W. O.
Steel. (Communicated by J. W. T. Armstrong.) (Nine Text-figures) .. 57-61

Graptolites from Tallong and the Shoalhaven Gorge, New South Wales. By
Kathleen Sherrard, M.Sc. (Plates i, ii; thirty-three Text-figures.) .. .. 62-82

The Genus Dawsonia. By Alan Burges, M.Sc., Ph.D. (Twenty-six Text-figures.) 83-96

Revision of the Genus Brachycome Cass. Part II. New Zealand Species. By
Gwenda L. Davis, M.Se. (Twenty-four Text-figures. ) Becton teal diet eke 97-106

On Australian Dermestidae. Part V. Notes and the Description of Four New
Species. By J. W. T. Armstrong. (One Text-figure.) So EWn ae ere) eles eer OFS

CONTENTS.

PARTS 3-4 (Nos. 343-344).
(Issued 21st October, 1949.)

Australian Rust Studies. VII. Some Recent Observations on Wheat Stem Rust
in Australia. By I. A. Watson and W. L. Waterhouse. (Plates iii, iv)

Notes on Microspore-types in Tasmanian Permian Coals. By se A. and Roma
Dulhunty. (One Text-figure.)

A New Subspecies of Aédes (Stegomyia) scutellaris Walker (Diptera, Culicidae)
from Northern Australia. By A. R. Woodhill. (Four Text-figures.) ..

Revision of the Genus Brachycome Cass. Part III. Description of Three New
Australian Species and some New Locality Records. By Gwenda L. Davis.
(Seven Text-figures.)

Notes on the Morphology and Biology of a New Species of Tabanus (Diptera,
Tabanidae). By Kathleen M. I. English. (Fifteen Text-figures.)

Crania in the Macleay Museum. By N. W. G. Macintosh

The Hair Tracts in Marsupials. Part III. Description of Species, concluded.
By W. Boardman. (Two Text-figures.) ..

Studies on Australian Marine Algae. V. Observations on and Geographical
Records of Various Species, particularly those of the Gelidium Complex.
By Valerie May

iii

Pages.

113-131

132-139

140-144

145-152

153-160

161-191

192-195

196-202

1V CONTENTS.

PARTS 5-6 (Nos. 345-346).
(Issued 15th December, 1949.)

Pages.
The Stratigraphy of the Lower Marine Series of the Permian System in the

Hunter River Valley, New South Wales. By G. D. Osborne, D.Sc., Ph.D.

(Two Text-figures. ) 203-223
The Cotypes of Fordonia papuensis Macleay. By Arthur Loveridge. (Communi-

cated by S. J. Copland.) 223
A Note on the Experimental Crossing of Aédes (Stegomyia) scutellaris

scutellaris Walker and Aédes (Stegomyia) scutellaris katherinensis

Woodhill (Diptera, Culicidae). By A. R. Woodhill. (One Text-figure.) 224-226
A Detailed Study of the Field Distribution of Strains of Clover Nodule Bacteria.

By Hilary F. Purchase and J. M. Vincent. (One Text-figure. ) 227-236
Abstract of Proceedings Fp ae NCE areata ech Gis othe Cue e Ria LaDy eneim cL pie 8c XXili-xxvi
IS Or vNSTAD ERS te Aca eet et ote coastline ware eit seasaes Creuict tel UMERNT So ara Seba t)s raya amine kia opp TeRe XXV1i-Xxxi
List of Genera and Species XXXii
List of Plates XXxii

General Index ETN tie Sigs Ua ee uae NE AIRE el calle Pie Plea NN Ne eh at ann eR PAN Wank 5M: O.O-GUN k=O. 0,1

ANNUAL GENERAL MEETING.
WEDNESDAY, 30th Marcu, 1949.

The Seventy-fourth Annual General Meeting was held in the Society’s Rooms,
Science House, Gloucester Street, Sydney, on Wednesday, 30th March, 1949.

Dr. Lilian Fraser, President, occupied the Chair.

The Minutes of the preceding Annual General Meeting (31st March, 1948) were read
and confirmed.

PRESIDENTIAL ADDRESS.

During the past year the Society has made some advance from the inactivity neces-
sarily imposed on it by the war and members have shown increased interest in the
general meetings, the attendance at which has on several occasions strained the avail-
able seating accommodation of the meeting room. Some meetings were devoted entirely
to talks and exhibits and papers were read by title only. A symposium on “James Stuart
and his Paintings” was arranged by Messrs. A. Musgrave, G. P. Whitley, T. Iredale and
Dr. Frank Marshall for the June meeting, and members of the University Biology Society
presented Notes and Exhibits on an Excursion to the Great Barrier Reef to the July
meeting. In August, Miss Joyce Vickery gave a talk, illustrated by colour photographs,
on the Royal Botanic Gardens, Kew, and its association with the history of botany and
botanical exploration of Australia. The Secretary also gave a brief talk on several
Western Australian minerals of economic importance. At the November meeting Dr.
R. N. Robertson, who had recently been abroad, gave a talk on “Some Impressions of
Botanical Laboratories Abroad’’.

Exchanges received from scientific societies and institutions totalled 1,423 for the
year. There were many new requests for exchanges, those granted being to: Société
des Sciences Naturelles du Maroc; Miincher Entomologische Gesellschaft; Musée Heude,
Universite l’Aurore, Shanghai; Institute of Zoology, Academia Sinica, Shanghai;
Institute of Biological Research, Fukien Academy, Foochow; Institute of Zoology,
University of Florence. It is pleasing to note that many European exchanges which
lapsed during the war have been resumed. Some complete sets of Proceedings were
sent abroad to replenish libraries destroyed by enemy bombing and grateful thanks have
been received from the Société Linnéenne de Normandie, Caen, and the University of
Louvain. There has been a steady demand for reprints of papers and for subscriptions
to the Proceedings, the latter particularly from the United States. By exchange the
Society has received a very large number of reprints which, with those already in its
possession, will be card indexed for the use of members. The Library shelves have
been numbered and the numbers added to the card index of periodicals so that each
periodical may be readily located by members using the Library.

The printing of the Society’s Proceedings has continued fairly satisfactorily during
the year, Parts 5-6 of Volume 73 having been issued on 21st January, 1949. Volume 73
consists of 465 + lvi pages, 22 plates, and 579 text-figures. The Society is faced with a
further increase of approximately 50% in the cost of printing and the future will be
dificult. Financial assistance was obtained from the Commonwealth Publications Fund,
the School of Public Health and Tropical Medicine, and the N.S.W. Department of
Agriculture for the publication of certain papers. This further increase in costs will
seriously curtail the Society’s ability to accept lengthy systematic papers in the publica-
tion of which it has somewhat specialized.

A link with the Society’s past was severed last July when Macleay House in College
Street was sold to the Returned Nurses’ Association, who intend to build on the site
club-rooms and a hostel for nurses.

A

li PRESIDENTIAL ADDRESS.

The nett return from Science House was rather lower than in the previous year
because of the purchase of equipment for the main hall which, however, is credited to
the Society’s capital invested. The Science House Extensions Committee has been
revived and the Council will be discussing the proposed extension, particularly with
regard to the Library and Reading Room and the provision of amenities during the
coming year.

The numerical strength of the Society at the 28th February, 1949, was: Ordinary
Members, 201; Honorary Member, 1; Life Members, 14; Corresponding Members, 3;
total, 219. During the year 27 new members were elected and 5 members were lost by
death. Short obituary notices are given at the end of this address. New members from
the Science departments of the University took an active part in the proceedings of the
Society.

Linnean Macleay Fellowships.

The Council during the year presented a petition to the Suprenie Court in Equity
on 19th November to obtain a variation in Sir William Macleay’s Will so that it could
be more easily administered under the present economic conditions. As the result of
the Court’s decision the conditions upon which the Linnean Macleay Fellowships may
be awarded by the Linnean Society of New South Wales are (in addition to the applicant
being a member of the Society):

(a) The number of Fellowships to be awarded in any year shall be such number

as the Council of the Society shall determine, not exceeding four.

(0) Qualifications of candidates shall be as follow:

(i) Residence in New South Wales.
(ii) The expression Science degree in the said will shall include a degree
in the Faculty of Science in Agriculture.

(c) (i) The salary of each Fellow shall be such amount not being less than £400
nor more than £800 per annum as the Council of the Society shall
decide provided that the total amount of the salaries and allowances of
all Fellowships awarded in any one year shall not exceed £1,600.

(ii) The Council of the Society may upon the application of any Fellow
make grants to him out of the remaining income from the Linnean
Macleay Fellowship Fund received by the Society in any year provided
the total sum, including the amount of salaries paid to Fellows during
such year does not exceed £1,600, to defray field and other research
expenses incurred or to be incurred by him in carrying on his work and
investigations as such Fellow.

(d) The appointments to Fellowships shall be made from year to year provided

no Fellow shall hold a Fellowship for a total period exceeding five years.

The Court declared that persons residing in the Australian Capital Territory (who
are otherwise qualified as candidates) are not persons from whom the Council of the
Society may properly receive applications for appointment as Linnean Macleay Fellows.

The Society’s two Fellows appointed for 1948 made satisfactory progress in their
chosen investigations.

Miss Muriel Morris, Linnean Macleay Fellow in Zoology, had her first paper (on
work completed just prior to the award of the Fellowship), ‘‘Life-history of an Australian
Crustacean, Acetes australis (Decapoda, Tribe Penaeidae)”’, published in Parts 1-2 of
Volume 73. She was awarded a Fellowship to continue research on the planktonic
population of the Hawkesbury River Estuary, to determine the food of the commercial
oyster (Saxostrea commercialis) and to correlate this with the plankton hauls; and to
determine the mode of action of the foot of the sand-snail, Polinices. In the investiga-
tion on the plankton population samples were collected at regular intervals throughout
the year and the material worked through qualitatively, but no quantitative estimations
have yet been made. In collaboration with the Fisheries Division of the Council for
Scientific and Industrial Research a series of catches from four stations up the Hawkes-
bury Hstuary were examined, adding very greatly to the amount of material available

PRESIDENTIAL ADDRESS. lili

for estimating the seasonal variations of the plankton population. The investigation
of the mode of action of the foot of the sand-snail, Polinices, has been completed and a
paper for publication is being prepared. Miss Morris was reappointed to a Fellowship
for 1949,

Miss Mary Tindale worked throughout the year on a revision of the ferns of New
South Wales and completed a taxonomic revision of the south-east Australian species of
the following families: Azollaceae, Cyatheaceae, Gleicheniaceae, Angiopteridaceae,
Osmundaceae and Psilotaceae. In addition studies were carried out on some of the
members of the Schizaeaceae, Ophioglossaceae and Marsileaceae occurring in this region,
as well as on the Australian species of Cyathea and Rumohra. Detailed descriptions were
made of each species and subspecies. A representative list of specimens examined was
cited in each case. Artificial keys were provided to the genera and species. To supple-
ment the collections in Sydney, material was lent by the National Herbarium of
Melbourne, the Queensland Herbarium, the North Queensland Naturalists’ Club and the
Herbarium of the D.S. and I.R., Wellington, New Zealand. Field studies were carried
out on a number of species, particularly Gleichenia, Azolla and Marsilea. Miss Tindale
is desirous of continuing the work at the Kew Herbarium, and for this reason did not
apply for reappointment. She will leave shortly for England, where she proposes to work
for the next two years as Australian liaison officer.

The Council made three new appointments to Fellowships for 1949, Misses Judith
Balmain and Adele Millerd in Biochemistry and Miss Mary Hindmarsh in Botany. Miss
Balmain proposes to continue investigations of the biochemical basis of inhibitory
action of aromatic compounds related to phenol, benzoic acid, hydroxybenzoic acid in
bacteria and fungi. Miss Millerd will investigate the oxidative metabolism of Solanum
tuberosum from the point of view of the cytochrome oxidase system which is the
link between the cell and oxygen in animal tissue. Miss Hindmarsh will study the effect
of sulphanilamide in mitosis and it is proposed to investigate in detail the structural
changes in the chromosomes and the disturbances of the mitotic mechanism caused by
subjecting roots to various concentrations of sulphanilamide.

Macleay Bacteriologist—Following the resignation of Dr. H. L. Jensen in September,
1947, the Bacteriology Committee, consisting of Professors J. R. A. McMillan, H. K.
Ward, W. L. Waterhouse, N. A. Burges, Mr. J. M. Vincent, and the Executive, met on
several occasions to discuss the filling of this position. Financially the Bacteriology
Account has suffered through the general reduction of interest rates, and the Society
could only offer a little over £500 p.a., which would not attract a bacteriologist of the
standing required. On behalf of the Society a deputation waited on the Deputy-
Governor of the Commonwealth Bank to ask for financial assistance. Subsequently
appeals were made to the Commonwealth Bank, the Rural Bank of New South Wales,
the Commercial Banking Company of Sydney and the Bank of New South Wales,
with the result that the Rural Bank made a donation of £200 p.a. for five years to
assist the Society. Later the Commonwealth Bank appproved grants totalling £1,750
from the Rural Credits Development Fund towards the cost of the research programme
submitted by the Society, the grants to be in annual instalments after the appointment
of a suitable bacteriologist. The position is now being advertised.

Other Research.—The only other research sponsored by the Society was a
mineralogical investigation of sedimentary rocks which was done by the Secretary
largely in the Geology Department of the University through the courtesy of Professor
L. A. Cotton. A paper, “Mineralogy of the Cheltenhamian Beds at Beaumaris, Victoria’,
was accepted for publication in the Journal of Sedimentary Petrology.

Notes of Members.

Dr. Robert Broom was honoured on the occasion of his 80th birthday by a special
publication of the Royal Society of South Africa in recognition of his outstanding work
on fossil reptiles, fossil men and apes, etc. A bibliography of his published work includes
a list of 402 papers, beginning in 1885 and continuing to the present day. A tribute

AA

iv PRESIDENTIAL ADDRESS.

is paid to his work in Nature, Vol. 162, 20th November, 1948. He was awarded the
Wollaston Medal by the Geological Society of London.

Professor N. A. Burges was appointed to the Senate of the University of Sydney.

Professor J. B. Cleland received the C.B.H. in the New Year Honours List.

Mr. D. H. Colless was appointed Entomologist to the Malaria Research Station at
Labuan, British North Borneo.

Professor W. J. Dakin has been awarded the Mueller Medal by the Council of the
Australian and New Zealand Association for the Advancement of Science.

Miss Daphne Davison was awarded an 1851 Exhibition Scholarship and left to study
at Cambridge University in August.

Mr. E. H. Ealey left towards the end of the year to join the Australian Antarctic
Expedition. Mr. T. Manefield left in March for the same purpose.

Mr. D. J. Lee was appointed lecturer in Entomology at the School of Public Health
and Tropical Medicine, University of Sydney.

Dr. I. M. Mackerras was appointed Director of the Queensland Institute of Medical
Research.

The Rev. H. M. R. Rupp has been awarded the Royal Society’s Clarke Memorial
Medal for 1949 for his distinguished contributions to knowledge of Australian
Orchidaceae.

Dr. J. L. Still was in the United States on a Rockefeller Fellowship first at New
York and later at Wisconsin, where he became a foundation member of the Enzyme
Institute at the University of Wisconsin.

Mr. R. H. Wharton was appointed Entomologist at the Institute for Medical Research,
Kuala Lumpur, Malaya.

Dr. A. B. Walkom attended the Unesco Conference at Beirut, Lebanon, in November
as a member of the Australian Delegation, and subsequently visited museums in Great
Britain and the Continent, returning to Sydney shortly before the Hobart meeting of the
Australian and New Zealand Association for the Advancement of Science, at which he
delivered his Presidential Address entitled “Gondwanaland, a Problem in Palaeo-
geography”. He also attended the Seventh Pacific Science Congress in New Zealand in
February at the invitation of the President of the Royal Society of New Zealand.

Professor W. L. Waterhouse has been awarded the Medal of the Royal Society of
New South Wales and the Medal of the Federal Council of the Australian Institute of
Agriculture for his research on cereals and other aspects of agricultural science.

The Secretary was granted leave to attend the Pacific Science Congress in New
Zealand in February, where she represented the Australian National Research Council
and the Royal Society of New South Wales.

Obituaries.

It is recorded with regret that the following members died during the year: Messrs.
EK. C. Andrews, M. S. Barnett, Captain J. D. McComish, Rev. E. N. McKie and Mr.
Rowland EH. Turner.

ERNEST CLAYTON ANDREWS, B.A., F.R.S.N.Z., died at Bondi, New South Wales, on
Ist July, 1948, aged 77. He was a member of the Society since 1899 and a member of
Council from 1922 to 31st March, 1946, and President, 1937-38. He resigned from the
Public Service, December, 1931, after 45 years’ service. He was Government Geologist
from 1920 to 1931. Mr. Andrews received the David Syme prize for contributions to
Australian geology in 1915, and in 1928 he received the Clarke Memorial Medal of the
Royal Society of New South Wales for original contributions to Australian geology and
general natural science, and in 1931 the Lyell Medal of the Geological Society of London
for researches in economic geology in New South Wales, and physical geology. He led
the Australian delegation to the Pacific Science Congresses in 1929, 1933 and 1939. He
was President of the Australian and New Zealand Association for the Advancement of
Science in 1930 and General Secretary of this Association from 1922-1926. He was
awarded the Mueller Medal of this Association in 1946 in recognition of his work in
modern physiography and associated studies. Mr. Andrews was President of the Royal

PRESIDENTIAL ADDRESS. Vv

Society of New South Wales in 1921. He delivered the Silliman Lectures in April, 1927,
at Yale University, the subject being “The Geology of the Pacific Region’. He
contributed five papers to the Society between 1902 and 1913 and one with B. Sawyer
in 1901. (See also Obituary Notice in Australian Journal of Science, xi, 48.)

Marcus STANLEY Barnett died on 15th July, 1948. He had been a member of the
Society since 1919 but contributed no papers. He was formerly a chemist in the Colonial
Sugar Refining Company, but since his retirement he lived at Mt. Victoria, New South
Wales.

Captain JAMES Doran McComisu, F.R.G.S., died on 3rd June, 1948, aged 67 years,
shortly after his election as a member of the Society. He was Captain, N.Z. Military
Forces (Retired List), and for the sixteen years prior to 1948 had spent most of
his time in the capacity of an Honorary Botanical Collector (mainly in the South Sea
Is.) for the Royal Botanic Gardens, Kew, England; the National Herbarium, Sydney,
New South Wales; the Dominion Museum, Wellington, New Zealand; the Auckland
Institute and Museum, Auckland, New Zealand; the Bernice P. Bishop Museum,
Honolulu, Hawaii, and (latterly) was preparing his notes for publication. At the time
of his death he was engaged on identifying and classifying the plants and trees of
Lord Howe Island which he had collected over a long period and he was preparing
a paper which he wished to read to this Society. Captain McComish had spent many
years on various investigations in connexion with the Pitcairn Islanders and on early
missionary history of French Oceania and the trees and plants there. The manuscripts
have been given to the Mitchell Library and to Dr. Oliver, Dominion Museum, Wellington,
New Zealand, respectively.

Rev. Ernest Norman McKir, B.A., died on 19th May, 1948. He was a member of
the Society since 1927. The Rev. McKie was installed as Moderator of the General
Assembly of the Presbyterian Church on 10th May, 1938, and ministered for twenty-six
years prior to 1938 was minister of St. Columba’s Church, Guyra, New South Wales, the
centre of a large country parish, embracing also the Wandsworth district. He was born
at Guyra, but spent the whole of his earlier years at Glen Innes. Mr. McKie first
entered the Commercial Banking Company of Sydney, from which he resigned to enter
St. Andrew’s College within the University of Sydney, graduating Bachelor of Arts in
1906, and completing his theological training in 1908. He was licensed by the Presbytery
of New England and later was ordained and inducted by the same presbytery to his first
charge, Manilla—Bendemeer, in 1909. From Manilla he went to the charge at Guyra,
in 1912. In addition to his ministerial labours Mr. McKie found time to follow scientific
interests, of which botany has been the chief. He has worked on the flora of New
England and other districts for several years. He was a member of the Royal Australian
Historical Society and of the Royal Society of New South Wales. In his country parish
he assisted rural improvements comprised under the Junior Farmers’ Club and the
Agricultural Bureau organization.

RowLanp EH. Turner, F.R.E.S., F.Z.S., died on 29th November, 1945, at Mossel Bay,
South Africa. He had been a member of the Society since 1904. He contributed five
papers to the Society on Hymenoptera between 1907 and 1913 and one in 1904 in
collaboration with Dr. G. A. Waterhouse, on Australian Rhopalocera—Lycaenidae.

A GUMMOSIS DISEASE OF CITRUS IN RELATION TO ITS ENVIRONMENT.

Contents.
Page.
Introduction Leh chung aoe ae ed rae se See we & Saas bias oe ae Ave oe Pe ae ret Vv
Historical Survey SOMES iG cue Cased Ghiesec ra eceatlY Getcha? | LovAOus Ain haa oa eric ietoave ene Mare eLertsotitni ema Nia
IMATE OF Was lol G5 oo a 6 Oe an! 640. oe oo bol) oo! ba vob 965) op Xdhi/
Spread of Disease Tig CR eR Vora TAS ONO YAO Win rch PROD OU MCR Ste EGR. Rae RNa aire ane | tre -vip Aid
CONCLUSION ME eh ice ye, Gul gectth tien Scott Soto) Maem) yong colade “Weiict. aefawletp ute aie torieiet vera 60 oo BAA
INTRODUCTION.

The plant disease which is the subject of this address is the gummosis disease of
citrus caused by the fungus Phytophthora citrophthora (Sm. and Sm.) Leonian. The

vi PRESIDENTIAL ADDRESS.

title may seem unnecessarily obvious since all plant diseases are bound up more or less
closely with the environment and it is clear that the range of conditions under which
a disease can occur must be limited by the character of the fungus which causes it.
Those, for example, which attack the exposed parts of plants are more obviously at the
mercy of the environment than are the soil inhabiting species, which are subject to
changes usually less extreme and less sudden than occur in the atmosphere, but in
each series there are species which can exist under relatively dry conditions, which
require continuously high humidity or which flourish at high or low temperatures. Most
pathogenic fungi are limited in the parts of the plant which they can attack, for example,
to the leaves alone, to leaves and fruit, the young stem, special tissues or to special
parts of the root system. We have on the one hand fungi whose habitat is the soil, and
on the other those whose whole life is spent above the ground. P. citrophthora has this
very unusual ability that it can invade any part of its host. I propose to deal rather
generally with those environmental factors which govern the various phases of the
disease which it causes.

P. citrophthora belongs to the Phycomycetes, it produces a branching, non-septate
mycelium which is thin-walled and very vulnerable to desiccation. Its only recorded
spores are zoospores produced in water or in a saturated atmosphere, and requiring
water for their distribution. It is a simple species without specialized strains. It is
of microscopic size, and its presence can be diagnosed only by its production of
characteristic symptoms in the host and its recovery in culture from diseased plant
tissues. This has led to the confusion of the cause of the disease with the conditions
which favour the growth of the fungus, and to a widely held belief that the weather and
not a pathogen is the primary cause of the disorder; a belief which has persisted ina
way it could not have done if the fungus were more easy to see, produced masses of
spores like a powdery mildew, or rings of toadstools, like Armillaria mellea.

By modern standards the disease has a moderately long history. A disease which
attacks a staple food crop such as Irish Blight of potatoes or Stem Rust of wheat is
more likely to be referred to in historical records than a disease of a luxury crop such
as the orange, but some of the aspects of gummosis are sufficiently spectacular to have
aroused attention long before the days of plant pathologists, though the causal organism
was not isolated and described until comparatively recent times.

At the present time gummosis disease is co-extensive with its host, and it is without
doubt the most important disease affecting the citrus crop. It is the object of this
paper to show that P. citrophthora is the major factor determining the present distribu-
tion of the citrus tree.

The Causal Fungus.—The earliest attempt by a mycologist to discover the pathogen
was that of Briosi in 1878 (quoted by Fawcett, 1923). He described the fungus Fusarium
Limonis and was inclined to regard it as the cause of the disease. Later Comes, in 1891
(quoted by Fawcett, 1923), attributed the disease to a bacterium, but it is probable that
he had confused two separate diseases and had isolated the cause of the minor one.
The extensive literature dealing with the possible causes of gummosis is fully reviewed
by Swingle and Webber (1896) and by Fawcett (1923). Many early writers attributed
it to so-called physiological causes. There was no critical experimental approach to the
problem until after the description by R. HE. and E. H. Smith in 1906 of Pythiacystis
citrophthora as the cause of brown rot of lemon fruit. Fawcett (1913, 1923, 1936), in
a very exhaustive investigation, proved that this fungus, now known as Phytophthora
citrophthora, was the cause of collar and crown rot also. Two other related fungi,
P. parasitica and P. palmivora, have since been found to be associated with P. citrophthora
in the U.S.A., but in New South Wales only P. citrophthora has so far been implicated.

The Gummosis Disease.—P. citrophthora is, strictly speaking, a soil inhabitant, but
it can produce sporangia on the soil surface and these can be splashed up on to leaves
and fruit. It causes a brown rot of fruit, stem and shoot blight, leaf spotting, collar rot
(i.e., the rotting of the bark of the main trunk), foot rot, which is the rotting of the
base of the trunk and the upper parts of the crown roots and finally the rotting of both
fibrous and permanent roots. Considerable gum formation usually accompanies the

PRESIDENTIAL ADDRESS. vii

collar rot and to a less extent the foot rot phases. This gum formation is one of the
spectacular features of the disease and has earned for it its common name.

The severity of the root rot phase varies with the soil conditions. In the early
stages or in mild attacks when the environment is not altogether favourable for the
rapid growth of the fungus, the attack may be limited to the fibrous roots. The fibrous
roots normally die off as their period of activity ceases, but if the fungus is present in
the soil and the level of soil moisture is high, young active fibrous roots can be
destroyed in a few days and replacement may not be sufficiently rapid to maintain the
tree in a vigorous state. If the soil is wet for lengthy periods, the fungus extends its
attack to the permanent roots, invading the cortical and phloem tissues. Smaller
laterals are killed back and lesions are formed on the larger roots, first on the lower
roots and then on the lower sides of surface roots. The first roots to be affected are
the lowest, mainly because the lower soil levels are most likely to be wet continuously.
As the disease progresses the tree will come to exist with the aid of the surface roots
only. At this stage the tree is obviously far more sensitive to the drying out of the
surface soil than trees with intact lower roots. Such trees scattered through an other-
wise healthy block greatly increase irrigation difficulties, since they require lighter but
more frequent watering than healthy trees.

The parasite is very sensitive to fluctuations of moisture and when the soil dries
out its activity ceases, the lesions begin to heal around the edges and new fibrous roots
are formed. Renewal of wet conditions causes a renewal in fungus activity. The
surface roots feeding in a zone of soil most subject to drying out are the last to be
affected by the disease.

The first symptoms in the above ground parts of a tree affected with root rot are
the thinning out of the foliage and the failure to make vigorous new growth. As the
disease progresses the foliage becomes an unhealthy yellow and die-back of twigs occurs
which is not compensated by renewal. Depending on soil conditions and treatment, the
tree may die out quickly, or make periodic attempts at regrowth. It is quite usual for
a tree in which the disease is progressing rapidly to set a heavy crop of fruit before it
dies. Trees affected with collar rot show severe yellowing and the death of the whole
or part of the top is rapid.

The Principal Factors Influencing Disease Development.—By far the most important
factor is water. The fungus being without protective structures of its own development
ean grow only in an atmosphere, whether soil or surface, whose humidity approaches
saturation. This is very well illustrated by the history of the disease and the distribution
of its different phases in New South Wales. Other environmental factors which may be
of local importance are, directly, soil reaction and temperature, and indirectly through
its effect on tree vigour and the rate of replacement of roots, the general level of soil
fertility, and lastly, but from the point of view of control the most significant, there is
the differing susceptibility of the various species of citrus.

HISTORICAL SURVEY.

Distribution of the Species of Citrus—In order to provide a background for an
account of the history of the disease, let me outline very briefly the way in which the
economic variaties of citrus have reached the areas in which they now grow. The species
of the genus Citrus are native to the tropical and subtropical regions of Asia and the
Malay Archipelago (Webber and Batchelor, 1943). The sweet orange was grown in
China for centuries before its introduction to Hurope and the sour orange, mandarins,
citrons, kumquats and the trifoliate orange have also been in cultivation there for a
very long time. They have also been grown in India (to which some varieties are
thought to be indigenous) for centuries, the orange, no doubt, reaching there from
China along the trade routes which were well established in early times.

The first species to be introduced to the Western Hemisphere was the citron, which
preceded the now much more highly prized varieties by many centuries. It is thought
to have been brought to Egypt by one of the Egyptian kings following his wars in Asia
and it has been identified in the inscriptions in tombs of the 12th-15th centuries pB.c.

viii PRESIDENTIAL ADDRESS.

Citron seeds have been identified in southern Babylonian ruins of 4000 B.c., though there
is no means of finding out whence they came (Tolkowsky, 1938). The citron was well
known to the early Greeks and Romans; Theophrastus refers to its growth in Media and
Persia and says that its fruits were prized for their exquisite odour and its leaves used
as a protection against moths in clothing. It was well established in Italy and had
become wild in some southern parts by the middle of the first century A.D.

The next species to be introduced from the East was the sour orange, known to us
as the Seville. It appears to have been carried to Basrah from India, and spread by the
Arabs to Persia, Syria, Palestine and Egypt and later through the Mediterranean
countries. Its uses are referred to by the Arab writers of herbals of the eleventh and
twelfth centuries A.D.

There is evidence both geographical and etymological that the lemon is a native of
the eastern Himalayas or Upper Burma. It was established in India and introduced by
the Arabs into Spain and northern Africa by about a.p. 1150 and to Italy and other
Mediterranean and European countries from Palestine by the Crusaders in the thirteenth
century. It did not reach China until about a.p. 1175 and is said to have come from the
“southern barbarians”.

The rough lemon, which has become one of the most important citrus root stocks
in use today, is believed to have originated in India as a hybrid between the lemon and
possibly the citron. It is an immensely vigorous type and grows wild in parts of India.
It was obtained in India by the Portuguese, and has since become naturalized in Florida
and southern Rhodesia. The sweet orange is a native of China or Cochin China and had
reached India and become established there by the thirteenth century. It was introduced
to Europe in the fifteenth century and was well established in southern Europe by the
sixteenth century and an important item of trade. The introduction of a superior variety
by the Portuguese at the end of the fifteenth century gave a considerable impetus to the
industry.

Since their introduction, oranges and lemons have been grown very extensively in
the Mediterranean countries. They were introduced to the Americas in the early
sixteenth century by the Spaniards and Portuguese.

Citrus was brought to Australia with the first fleet in 1788 by Captain Hunter and
succeeded well enough in the Sydney district. By the middle of the 19th century quite
good types of oranges—the Parramatta (a local seedling), the St. Michael (from the
Azores), the Bahia (now known as the Washington Navel from Brazil), and mandarins
and lemons were commonly grown around Ryde and Parramatta and were a source of
great profit to the growers.

The Origin and Spread of the Disease——The early outbreaks of gummosis are well
summarized by Swingle and Webber (1896, quoting from Briosi), McAlpine (1899) and
Fawcett (1923). Fawcett referred to various minor types of gummosis which had been
described by early writers in the seventeenth century in various parts of the Mediter-
ranean, and early in the eighteenth century in South America, and Shiraz-el-din (1934)
quoted an early Arab source as mentioning gummosis in the tenth century in Spain. It
is open to doubt, however, that these were in fact due to Phytophthora attack. The
first record of a very serious attack, undoubtedly due to Phytophthora, is from the
Azores, where gummosis evidently gained a foothold about 1832. It is clear from the
accounts of this that it must have been newly introduced. The outbreak is vividly
described in a letter to Charles Moore, Director of the Sydney Botanic Gardens, by one
of the largest planters in the Azores. Unhappily none of Charles Moore’s correspondence
appears to have been preserved and we know of this letter only in a quotation given by
Alderton (1884). Part of it is worth requoting: “The disease was first observed in our
gardens about the year 1836, but it is probable that for a year or two previous it must
have been not inactive, inasmuch as the consequences we then observed were very general
and disastrous. At first we noticed that our orange trees were dying one after another,
and were wholly unable to divine the cause; ere long we found it was our best and
oldest trees that were thus disappearing. At that time we had trees producing from
6,000 to 20,000 oranges a year—trees that were as much as two hundred and three

PRESIDENTIAL ADDRESS. ix

hundred years old. By degrees we began to observe that all the trees affected with the
disease produced a very large crop exactly in the year that the disease manifested itself;
that the leaves became yellowish and fell off in large quantities; and that on the trunks
or stems near the ground (and sometimes beneath the ground) the bark opened and
drops of a kind of yellow gum exuded. ... At first many orangeries were quite destroyed
and had to be replanted, so that a remedy for so great a misfortune was earnestly sought
for. Opinion as to the cause of this disease was very much divided. Many thought it
must be that the orange tree had but a limited period for its existence, and that, this
being reached, the tree must naturally decay. As the only method which we then had
for propagating orange trees was by layers, we thought that the proposed explanation
was not an unreasonable one; but later it was found that new seedlings were attacked
in the same way, although not with so much violence. Then again, others thought that
it must be something in the atmosphere which prevented the free circulation of the sap
and other fluids, and supposed that this must be the origin of the disease. The great
point, of course, was to find an easy and practical mode for remedying the evil.
Gradually, after repeated trials, we found that superabundance of moisture in the soil
was one of the worst conditions for the disease, and every kind of drainage was the best
way of improving the health of the trees. Soon afterwards we found that the destruction
of all the diseased bark and wood in the stem of the tree affected was the best method
for us to adopt to save the trees.”

He goes on to describe the methods of treatment and maintenance of stocks for
replanting where trees had failed, and concludes: “although the disease continues to
exist, our gardens now look very prosperous for the remedy is known, and the disease
causes no such ravages as it did for the first seven or eight years after it made its first
appearance. Orange trees take very easily by transplantation and we always keep our
plantations fully supplied. The only and great difference which our orange gardens
present is that we have no longer old and large trees in our plantations; you can no
longer count on handing your orange trees over to your children.”

From the Azores the disease spread through the Mediterranean countries, causing
devastation, and a complete change in the system of citrus growing. It appeared in
California in 1875 and in Florida in 1876 and caused a discontinuance of the use of the
common lemon, lime and citron as stocks and the general adoption of sour and sweet
orange as the principal stocks. In the few cases where information is available, the
outbreaks of the disease in severe form have been related either to seasons of heavy
rain or associated with excessive irrigation.

At present the disease is the most destructive of the citrus crop in Japan and also is
of general occurrence in China. It is also well known in Egypt, India, Palestine, South
America, South Africa, etc.

By the year 1860 citrus growing was a very prosperous industry in the Parramatta,
Ryde and Baulkham Hills districts near Sydney. The trees were grown in loamy soil
derived from Wianamatta Shale or in pockets of sandy soil and were probably dependent
entirely on the natural rainfall without supplementary irrigation. In the year 1860 the
rainfall recorded at Parramatta was 72:9 inches and in Sydney 82-8 inches, and in that
year gummosis made its first appearance in Australia. In the year 1862 the rainfall
was only 24 inches in Sydney but in 1864 it was 72-1 inches. By 1867 many orchards in
the hills around Ryde were yellow and sickly, with gumming dying trees (McAlpine,
1899). The disease was known as Bark Disease, Sore Shin, Bark Rot, The Fatal, and
most invidiously, the Sydney Disease, as well as the Italian Mal di gomma and the
Portuguese Lagrima. Between 1860 and 1870 hundreds of acres of orange trees were
destroyed (Alderton, 1884). A commission was appointed which inquired into the loss
and attributed it to “unusual climatic influence, induced or accompanied by excessive
rainy seasons, the earth thereby becoming too cold and wet to support the orange tree
in perfection; and second to defective cultivation”.

The position was regarded as so serious that in 1867 Charles Moore, the Director
of the Sydney Botanic Gardens, was sent to Spain and Portugal to examine the state
of the citrus industry in those countries and to find whether something could be learned

x PRESIDENTIAL ADDRESS.

of the treatment of the disease which had recently devastated parts of the citrus growing
areas there which might be of value to New South Wales growers. His “‘Report on the
State and Management of the Orange Plantations in Spain and Portugal” is dated
18/3/1868, and I quote from it extensively because of the light it throws on horticultural
practice at that time as well as the relation of this to the progress of the disease. He
went first to Portugal, where “inquiries revealed that a disease of a most fatal character
had destroyed an immense number of trees, chiefly about Lisbon, Setubal and Evora,
and a commission had been appointed to inquire into the origin and nature of the disease,
but no satisfactory results had been obtained, not even a report. As Lisbon only a few
solitary trees remain .... The whole country (about Setubal) is occupied by orangeries,
the old trees being more or less affected, in a similar manner, in many respects as those
which perished in our colony. In every quarter the appearance of the orchards reminds
me of those about Ryde some few years ago, when they were fast going to destruction.
The foliage yellow, the tops of the branches dead; the bark dry, shrivelled and peeling
off, a small portion at the base of the stem on one side soft, and the wood underneath
rotten and discoloured, the roots on the same side being in a similar condition, emitting
an offensive smell. ...It was only when the roots were in a very putrid state that any
evidence of fungus was observed.’’

He then discussed whether this fungus might be the cause or the effect of the
disease. He went on: ‘No remedy which had been tried had proved effectual. That
generally adopted is, on its first appearance (indicated by a resinous gum exuding from
the bases of the stem) to carefully remove the earth on this side, cut out all the
unhealthy parts of the stem and roots, and to take these, together with the soil which
had been in contact with them, some distance off and burn them. Such parts of the
plant as are operated on are left for some time exposed to the air, and then fresh, richly
manured earth placed about them. Some cultivators, instead of exposing the roots to
the air, cover the wounds with pinepitch or coal tar and immediately cover the parts
with fresh earth. Both plans only retard—they do not cure the disease.”

He then summarized the position:

“1. Oranges have been cultivated in Portugal for upwards of a century.

2. There is no record of disease until the last 10 years.

3. Old trees have suffered more from disease than young ones.

4, Many young plantations are now quite healthy, although situated close to
those of older growth much affected by the disease.

5. It is not known in what part the disease commences. Some persons believe
that the young roots are first attacked, others that it begins at the base of
the stem.

6. The upper parts of the stem, the branches and leaves continue in apparent
health long after many of the roots are quite rotten.

7. Sometimes only half of the tree will die, and the other half bear good fruit
for 2-3 years and then perish....

10. In every case, constant irrigation during the dry season is considered to be
indispensable... .

12. The seedling orange is considered to be a better and far more permanent
stock than the lemon.

13. The tree attains here a considerable age and few were known to die until the
present disease appeared.”

In Andalusia in Spain, Moore found the plantations around Seville and Cordova in
beautiful condition: ‘At Seville, Palma, Mairena, Gibralean and Cordova, the soil is light
and loamy in comparatively flat situations, but the trees look equally well if not better
on the slopes of the Sierra Morena—a mountain range near Cordova. In every case
constant irrigation was resorted to, and to such an extent that in some places I observed
the water standing on the surface around the trees and retained there by means of basins
formed by ridges of earth, the whole extent of the ground being so thoroughly saturated
as to be quite unfit to walk upon....

PRESIDENTIAL ADDRESS. xi

“The great orange-producing parts of Spain, however, are further to the eastward of
this; much of the rich, low, cultivated land in the Province of Valencia and Murcia
towards and along the Mediterranean coast being occupied with this fruit. I therefore
proceeded to examine these districts, and entered the former Province via Almansa,
passing through the beautiful valley called Huerta de Manuel—one of the most extensive,
rich and best cultivated parts of the country. On every side oranges appeared and all
evidently affected by some disease; signs of decay being evident in the yellow foliage,
and in the great number of dead and dying branches. ... It or any other disease was
entirely unknown until about five or six years ago.”

Discussing the methods of control in vogue, he said: “When the stem at the base
is attacked and the disease extends in a lateral direction, or around the stem, the tree
seldom or never recovers. When, on the contrary, it proceeds upwards, affecting one
side of the stem only, it may be checked, and effectually eradicated by cutting out every
part that is diseased both in the stem and roots, covering the wounds with a liquid
preparation of lime and sulphur; removing all soil which has been in contact with the
diseased parts and replacing it with fresh soil enriched either by guano or good rotten
manure, and thoroughly incorporated by lime and sulphur in a powdered state... .”
The benefit of lime and sulphur “would naturally lead to the inference that the disease
in this part of Europe is attributable to a minute fungus whose insidious mycelium or
spawn penetrates the softer tissues of the roots, extending in the course of development
upwards to the stem and unless speedily arrested, causing the death of the tree... .
All the trees in this area, whether healthy or otherwise, were subjected to constant
irrigation during the summer months, every tree was kept thoroughly soaked. This
system had been practised from the time of the first introduction of oranges, but there
are now many intelligent growers who believe that it may be done to too great an
extent; as a proof of which I had pointed out to me fine orchards occupying the drier
situations in which the trees were altogether healthy. Some persons even attributed
the cause of the disease to excessive irrigation; but in Andalusia, where its fatal effects
are as yet unknown, the orange grounds are kept as thoroughly and as continuously
saturated with moisture as anything I had met with.”

The differences in susceptibility shown by different citrus stocks were clearly recog-
nized by Moore. “Formerly young trees from buds on stocks of citron or lemon were
preferred. These, at two years old, are now scarcely saleable, even at the low price
of two reals, or about sixpence each, whilst those of the same age, on stocks of the
bitter orange, are readily purchased at 12 reals, or about 2s. 6d. each. Although the
Spaniard is averse to change in any old custom, yet the fact that trees on stocks of the
bitter orange were seldom or never touched by the disease could not fail to fix his
attention, and the result has been to bring plants propagated in this manner into great
demand. One of the many remarkable instances of trees of this description resisting the
disease was pointed out to me in what had been a very fine plantation, near Alcira.
Here every tree was almost either dead or dying, with the exception of a group of about
a dozen all of the bitter orange, which were entirely unaffected, and in fine healthy
condition, while some of those surrounding them which had suffered so severely from
the disease were raised from seed, others budded both on the citron and lemon. .
Hitherto, in the Murcia and Valencia plantations, 90% of the worked orange trees were
upon citron stocks, about 8% on lemons, which were the first attacked by the disease,
as were the lemon trees themselves, and with such fatal results that this fruit had
almost ceased to be exported from these districts, at one time the most productive.”

It is obvious from this admirable report that Moore had a clear and accurate
conception of the problem. He recognized that a pathogen must be involved, that it
had been introduced and was spreading and that it was favoured by heavy irrigation,
and he pointed to the remedy, in the use of a resistant stock, the remedy which today
we have concluded is the only real one, although for the present we recommend a
different stock. If the report could have been followed up from this stage the problem
would have been solved, but the science of Plant Pathology was at that time in its
infancy. Investigation into the behaviour of plant pathogens was only beginning and

xii PRESIDENTIAL ADDRESS.

mycologists had barely started to work out techniques for the study of these fungi, and
it so happens that this pathogen, Phytophthora citrophthora is not altogether easy to
isolate except from very favourable material. It was not until nearly 25 years after
Moore’s report was written that a plant pathological laboratory was set up in the United
States Department of Agriculture and shortly after that pathologists were appointed
to the Departments of Agriculture in Victoria and New South Wales (1890).

Meanwhile after its first dramatic outbreak gummosis receded somewhat in import-
ance, and the New South Wales fruit grower, like the Azores orange grower before him,
became resigned to the continued presence of the disease. It became just an additional
burden to be borne. As the result of Moore’s recommendations the sour orange stock
was tried here, but it never gained favour in coastal New South Wales. Alderton (1884),
in his review of the citrus industry, states: “There is plenty of evidence to support Mr.
Moore’s theory that the seville is the hardiest of all trees, and is the least subject to
disease, but it is a very slow grower, and for that reason it finds but little favour. In
fact the stock which is known to be the most productive of disease, the lemon stock,
is the most popular in N.S.W., for the reason that it is a vigorous grower, makes a tree
the soonest, and produces the quickest return. A good grafted lemon stock will grow
into a tree twice as fast as a grafted orange stock, produce soon and heavily, and die
young (twenty years old and upwards)... .”

The recession in importance of the disease after the initial outbreak is to be ascribed
to a combination of improved cultural practice, extension of citrus growing to better
drained soils and the absence of such flooding rains as were experienced in 1860 and
1864. Previous to the original outbreak the trees were commonly planted in basin-shaped
depressions. The lodgement of water in these holes during wet weather became recog-
nized as conducive to gummosis and the planting system was improved by the trees
being set higher.

Gummosis, however, continued to be a source of loss, and frequent references to it,
citing various causes and treatments, occurred in the agricultural literature from that
time forward, and its association with heavy or ill-drained soils was confirmed by many
observations. However, for 75 years following Moore’s report there was no substantial
advance in Australia in treatment or recognition of the disease. Early plant pathologists
accepted the fact that a pathogenic fungus was responsible for gummosis, but the full
extent of the damage which could be done and the limitation of the productive life of
the tree caused thereby were not appreciated. The agricultural population generally
continued to blame the weather and the soil.

McAlpine (1899) in Victoria and Darnell-Smith and MacKinnon (1916) and Hamblin
(1920) in New South Wales followed Briosi in regarding Fusarium Limonis as the cause.
It is true that various soil-dwelling Fusaria are always close followers of the real
pathogen, which is itself fugaceous, and it is not in the least to be wondered that
Fusarium Limonis was blamed. Darnell-Smith and MacKinnon (1916) referred to
Fusarium Limonis as the cause of the disease known as collar rot, but did not link it
with gummosis and foot rot, which they associated with poorly drained or excessively
irrigated soils or heavy spring rain. They concluded that ‘‘a review of the whole case
leads to but one conclusion, viz., that the one great capital preventive measure for
gummosis to be applied when laying out an orchard is proper attention to drainage; and
the one supremely important remedial measure for gummosis to be applied to an orchard
already established is, again, proper attention to drainage”’.

The disease was not restricted to coastal orchards. In 1912 orange and lemon trees
at Hay, subjected to over irrigation, became sickly and died, the roots being stated to be
rotted.

The next outbreak of disease in New South Wales sufficiently serious to demand
attention occurred in the Murrumbidgee Irrigation Areas. This region had been
developed as a soldiers’ settlement in the years following the 1914-1918 war. In the
desire to get men started quickly on productive farms, large areas of land were laid out
for fruit growing, and were planted up before any soil survey had been made. A large
proportion of the settlers had little agricultural training, the irrigation layout was

x

PRESIDENTIAL ADDRESS. xiii

frequently inefficient, water was cheap and the tendency was therefore for the growing
trees to be given more water than they could utilize or which could be removed by
existing drainage systems. This did not matter in those soils where the natural drainage
was good, but much of it was rather heavy, or had a heavy phase below a permeable
top soil. The result was that artificial perched water tables were built up in many
sections. By 1928 this condition was well recognized and advisory services stressed the
need for minimum applications of water to citrus. The average rainfall received in this
area is of the order of 16 inches p.a. and the evaporation rate during the summer is
high. The rainfall is, however, not altogether reliable, and occasional seasons occur
when more than the average rainfall is received. The winter of 1931 was one such
season of high rainfall, and flooding occurred in some sections. Taken in conjunction
with the previous history of overwatering, this resulted in a considerable decline in tree
health and a survey was undertaken by Barnard at the request of the advisory committee
of the C.S.I.R. Citrus Research Station at Griffith, to inquire into the cause. In his report
Barnard (19386) cites some interesting figures regarding tree mortality. An analysis
of the planting figures showed that considerably more than 5,000 trees over six years
old had gone out of production between 1931 and 1935 and less than 28% of the new
trees planted in 1926-1929 reached the age of six years. The possibility that the
condition might be due to the action of a parasite was not raised, and waterlogging
due to faulty methods of irrigation accentuated by heavy rains and floods was given as
the main factor causing the decline. The possible value of the sour orange stock was
mentioned, but this was considered to be because it was resistant to waterlogging as
such.

The Murrumbidgee Irrigation Areas were again subjected to heavy rain and flooding
during the winter of 1939, and the winter of 1942 was continually wet though floods did
not occur. The decline in tree health continued. In a report of the Irrigation Research
Extension Committee of 1943 the following statement is made: ‘An orchard survey
carried out during the 1940 winter showed that of the 6,000 acres of citrus planting
only 53% of the trees were healthy, 25% were slightly unhealthy and 22% were very
unhealthy. Since then the decline in citrus tree health has become much worse. A
particularly heavy loss of trees followed the wet conditions during May and June, 1942.”
(1.R.E.C. Report, 1943.) In the report of a meeting of extension officers held in Leeton
in October, 1942, it is stated: “(a) By 1942 at least 50% of the citrus plantings in the
M.I.A. had gone out of commercial production; (0) no orchard in the whole of the Areas
is completely unaffected; (c) young as well as older orchards are being affected; and
(d) it is extremely difficult to establish new plantings.”

This story is quite similar to that of the early troubles of the citrus industry in
California and could be duplicated for the Australian settlements of Mildura, Tresco
and to a less extent Curlwaa, Coomealla and others.

Catastrophes like this meant that except in well-drained soils citrus growing could
not be regarded as a stable type of agriculture, and it was urgently necessary for the
cause to be established without doubt, and if possible a remedy found.

Investigations into the possible pathogenic nature of the condition of the trees in
the Murrumbidgee Irrigation Areas commenced in 1942, by good fortune a year of
unusual rain. The impetus which the weather gave to the disease during the winter of
that year greatly facilitated the isolation and identification of the organism, as well as
the experimental work necessitated by the discovery. It was possible to prove that
here was a pathological problem which had been treated as a cultural one and that the
pathogen was P. citrophthora. As by this time the fungus was well known and had
been thoroughly investigated overseas, this knowledge was at our disposal to apply to
our local problem. Here, however, the fungus was found to be playing a more insidious
role than had hitherto been ascribed to it. This is the root rot phase described above.
There was no foot rot associated, none of the gum production which was a feature of
the early outbreaks in New South Wales and elsewhere. The conspicuous feature of the
many root systems examined that year was the extremely poor development of fibrous
roots except in the upper soil levels (Fraser, 1942).

B

Xiv PRESIDENTIAL ADDRESS.

As well as trees showing clear evidence of infection there were other orchards in
which the trees were without obvious symptoms of decline. The colour of their foliage
was quite good, but a gradual decrease in fruit production took place after the fifteenth
year, soon after they had reached full bearing maturity. These trees might never have
been waterlogged, but it was always found that the soil in which they were growing
was retentive of moisture. The condition was found to be due to the continual rotting
of the fibrous roots before they had reached maturity, but without much damage to the
permanent roots. It was not due to nutritional exhaustion as suggested by Barnard
(1936). A precisely similar disease of avocados caused by P. cinnamomi has been
investigated in California and South Africa by Wager (1942).

INFLUENCE OF THE ENVIRONMENT.

The Present Distribution of the Major Disease Phases in New South Wales.—Iin the
Murrumbidgee and Murray irrigation settlements the disease is limited to the root
system. Though this phase of the disease had not been recognized previously, it is
probably in the aggregate the most important, since there is not any direct method of
dealing with it. With increasing age citrus trees on any but sandy, well-drained soils
gradually become unprofitable because of the continuous toll on the fibrous roots. The
process may take 20 years or more if excessive water is not applied, but it seems none
the less to be inevitable.

Though less general, this phase of the disease is also important in many coastal
districts. Along the Hawkesbury River and its tributaries citrus has been grown for
almost a hundred years. Parts of the river banks are fixed sandhills, and on these,
with their excellent under-drainage, citrus grows to perfection, but on the heavier soils
of the flats and in those areas subject to flooding the story is the same as in the Murrum-
bidgee Irrigation Areas. In parts of the Hills district where the soil is a clay loam
overlying heavy clay subsoil, this phase of the disease is also present. In special
localities in the Gosford district, too, root rot has been found to be a major problem.
Orchards where seepage collects from neighbouring hills, particularly if an impervious
sheet of rock underlies the soil, are likely to be affected. The importance of the removal
of surplus water from such soils is illustrated by the behaviour of two blocks of lemon
trees in an orchard at Calga. One of these was planted in the standard way, trees 20
feet by 20 feet on the square. The other block was double planted, the trees 10 feet by
20 feet apart. Both blocks of trees were well grown for their age (15 years) when they
were examined some years ago, but following on a season of heavy rain the double-planted
block appeared reasonably healthy whereas in the other block considerable root rot was
evident. Both blocks were subject to a comparable amount of seepage from adjoining
higher land.

The collar rot phase of the disease is quite common in coastal orchards, particu-
larly in the autumn following periods of rain and high humidity. The lemon is the
variety most usually affected, the bark of the butt above the bud union is attacked and
rotted, the lesion very often stopping at the bud union. The lemon is extremely
susceptible and can be attacked under conditions in which other more resistant species
are not. That this phase of the disease should be unknown in western irrigation districts
must be due to the fact that the air humidity is not sufficiently high for long enough
periods to allow this type of attack to take place.

In some coastal and Hills district orchards fruit and leaf attack is also not
uncommon under conditions of high humidity and discontinuous rain; this also is
unknown in the west. In coastal nurseries, where seedlings are grown in crowded beds
under moist conditions, outbreaks often occur in which leaf, shoot and roots are all
affected.

Soil Reaction.—In artificial culture P. citrophthora grows best in media where the
pH lies between 6:0 and 7-5. In media of pH below 5-0 it grows more slowly and below
pH 4:5 it makes only slight and abnormal growth. It is to be expected, therefore, that
a similar effect should occur in the field. Citrus trees are very tolerant of soil acidity
and grow well in soils of pH between 4-0 and 8-0. In soils whose reaction is naturally

PRESIDENTIAL ADDRESS. XV

around pH 4:5 or below, root rot is rarely found. In the Gosford district there is a
considerable acreage of citrus planted on flats along the banks of creeks, and these
are sometimes flooded. When this occurs leaf and fruit infection commonly results
but the roots remain almost unaffected. The pH of these soils is about 4:5. A striking
instance of the effect of soil reaction on disease intensity occurred in a nursery at
Wyong. One bed of seedlings on the river flat where the pH of the soil was 4:5 was
flooded for several days following heavy rain. The seedlings developed some shoot
blight, but the roots were little affected. Several other seed beds were situated not
25 yards away on sloping ground above the level of the flood. The soil, which was a
moderately heavy loam, was wet but not waterlogged for the duration of the rain. The
soil pH here was 5-6, and in these beds very extensive root rot developed which destroyed
all the seedlings within a month.

This relationship with soil acidity has been investigated under experimental
conditions, using year-old seedling rough lemon stocks grown in 6-inch pots in soil
which had received various admixtures of lime or sulphur. The greatest amount of
root rotting took place in soils of pH between 5:4 and 7:5; a moderate amount at 4-8 to
5-0 and very little between 4:3 and 4:5. This would appear to offer some prospects as a
means of field control but so far the results have not been altogether satisfactory,
perhaps because of the difficulty of obtaining the uniformity which is possible with
hand-mixed potting soil.

The effects of two different nitrogenous fertilizers are of interest in this connection.
Sulphate of ammonia has an acidifying effect when added to the soil, and this effect has
been demonstrated by a number of investigators to be greater in the immediate vicinity
of a root where absorption is actively proceeding. Nitrate of soda on the other hand
decreases the acidity. In pot experiments the addition of small quantities of sulphate
of ammonia gives very great protection to roots against the attack of P. citrophthora
even in soils where the average pH value may be in the danger range, and it seems
reasonable to assume that this is because of the development of a more acid shell around
the roots. In soils where the average pH value ranged from 3:8 to 5-1 as the result of
the addition of sulphate of ammonia to soils originally less acid, little or no damage
from root rot could be induced. Nitrate of soda has a precisely opposite effect. The
addition of small quantities was found to stimulate root rot greatly while raising the
general level of the pH value. Severe rotting occurred from 4:8 upwards, very much
more than in soils of the same degree of acidity where only sulphur had been added.
Probably in these cases the pH of the soil immediately around the roots was higher than
the general average. In soils between 5-8 and 7:0 practically the whole root systems
were destroyed three months after inoculation. The pots were not kept excessively
wet, but moist enough to ensure good growth in the control plants.

Some effects of the application of fertilizers on root rot development have been
observed in the field. In some orchards under high soil moisture conditions nitrate of
soda applications have resulted in rapid stimulation of root rot and lime or dolomite
applications have in few cases been known to have the same effect.

The effect of soil reaction on disease development has been established for a number
of pathogens. One of these, Phytophthora cinnamomi, is very similar in behaviour to
P. citrophthora. In Queensland this species causes a wilt of pineapples and there
Lewcock (1935) has found little disease in soils whose natural acidity is below pH 5:1.
White (1937) has found that the root rot of Rhodendrons caused by the same species is
very slight in soils of pH 4:0 and is much less serious at 4:7 than at 5-2 and 5-4.

Temperature.—The optimum temperature for the growth in culture of P. citrophthora
is 25-28 degrees C., no growth is made at temperatures over 32 degrees C. and only very
slight growth at 5 degrees C. This sensitivity to high temperatures no doubt is one of
the reasons for the absence of the pathogen from the surface soil layers of soil during
the summer, and the healthy nature of the fibrous roots developing there.

Variation in Susceptibility of Different Species of Citrus——The resistance of the sour
orange stock and the special susceptibility of lemon, lime and citron were well known
long before the cause of the disease was discovered. The relative susceptibility of a large

BB

Xvi PRESIDENTIAL ADDRESS.

number of different varieties and species of citrus to P. citrophthora was determined by
Klotz and Fawcett (1930) on the basis of the average size of the lesion produced following
inoculation of the trunk of mature trees. Their results showed the lemon to be very
much the most susceptible, followed by lime, sweet orange, certain varieties of mandarin,
rough lemon, Sampson tangelo, sour orange and kumquat. Certain tangelos were
resistant, others susceptible. Of three trees of Poncirus trifoliata inoculated two gave
no reaction but the third proved extremely susceptible. In Brazil Fawcett and Bitancourt
(1940) found much the same order of resistance to three species of Phytophthora, P.
parasitica, P. citrophthora and P. palmivora, lemon being the most susceptible followed
by sweet lime, grapefruit, valencia orange and tangerine.

With one important exception the general grouping has been confirmed by the inocu-
lation experiments carried out in New South Wales, firstly using potted seedlings in the
glasshouse and secondly using trees about 15 years old at Leeton. The method of
inoculation of the potted seedlings was (a) to make a small incision in the bark, insert
a fragment of mycelium and wrap the area in damped cotton wool. Wounding is not
necessary, but it is desirable to ensure a uniform start of infection in all plants inocu-
lated at the same time; and (0b) to inoculate the roots by inserting the inoculum into
the soil. In field inoculations all incisions were covered with vaseline. By this means
the following classification was arrived at:

Very susceptible: Lemon, sweet lime.

Moderately susceptible: Sweet orange, rough lemon, grapefruit.
Fairly resistant: Sampson tangelo, Thornton tangelo, sour orange.
Resistant: Citremon.

Immune: Carrezo citrange, Poncirus trifoliata.

The Cleopatra mandarin was intermediate in susceptibility between the sour orange
and the rough lemon. In all the experiments, which covered many hundreds of trees
and some dozens of different isolates of the fungus, the reaction of the two species chiefly
studied, the rough lemon and P. trifoliata, remained constant. The resistance of P.
trifoliata has never shown any signs of breaking down. For this reason the results
obtained by Klotz and Fawcett with one out of three trees appears to be open to doubt,
the most possible explanation being that their susceptible tree was of hybrid origin. The
immunity of the P. trifoliata stock has been confirmed by many field observations in
New South Wales and Victoria.

SPREAD OF THE DISEASE.

We are not likely to know where this fungus originated. The history of its ravages,
however, does very strongly suggest that it spread from a focal point rather than that
it is naturally present in soils. The description of the outbreak in the Azores and
Moore’s account of his observations in Spain leave little room for doubt on this point.
The widespread nature of the first bad outbreak in New South Wales suggests that it
had been introduced at an earlier date than 1860, and had become established through the
Ryde and Parramatta districts, causing only minor and unrecognizable injury, so that
when the wet conditions of the 1860’s were experienced, the collapse of trees was general.

Its distribution in New South Wales since that time is very easily understood.
Practically all the citrus trees grown in eastern Australia are raised in the Carlingford—
Ermington districts, close to, or even on the sites of old orchards. Field observations
jndicate that once introduced into the soil the fungus takes its place in the soil
assemblage of micro-organisms and may remain there indefinitely. In most seasons the
citrus trees grown in the metropolitan nurseries are healthy. If they carry infection, it
is of a minor type, but when a rainy autumn occurs considerable infection may be present
and this, of course, has a very serious effect on the young tree when it is planted out. For
instance, of the trees sent out by one Carlingford nurseryman to the Murrumbidgee
Irrigation Areas in the spring of 1945, over 60% died in the first year, though planted
in virgin soil, and many of the remainder produced only poor trees. On the coast losses
of up to 100% of trees from the same source occurred within three months of planting.

PRESIDENTIAL ADDRESS. Xvii

Young trees with partially affected root systems are particularly susceptible to frost
injury and are killed back badly at temperatures at which healthy trees are unaffected.

To illustrate the importance of nursery infection I quote the case of an orchard
planted at Narromine, close to the river. It consisted of valencia oranges and grape-
fruit, both from the same nursery and planted in adjacent blocks at the same time. The
ground was partially flooded during the first year of growth and several times subse-
quently. By the time the trees had reached the age of 12 years more than 50% of the
grapefruit were dead (some had died quite soon after planting) and most of the rest
showed a greater or less degree of infection. The valencias were all healthy and well
grown. In this case it was quite evident that the grapefruit had been infected in the
nursery, and the valencias, which would have been raised in a different bed, were free
of infection. The soil was a sandy loam of pH 6-0 and quite suitable for supporting the
healthy growth of citrus for considerably more than 12 years.

There is some evidence that the fungus can become established in soil in the absence
of the host plant. I shall quote one case from the Wyong district. Seedbeds were planted
on a flat quite close to Wyong Creek, and in these seedbeds three patches of seedlings
each about 4 feet across succumbed to root rot. The area had been under grass for a
few years previously, and before that under natural timber. No citrus had ever been
grown in or near it. But Wyong Creek flows through citrus country and floods have
been known to cover the seedbed site in past years. It seems most likely that the
infection was carried down in a flood, possibly as an infected fruit or leaf and that
the fungus was able to establish itself in the soil from this.

CONCLUSION.

The citrus tree is evergreen and its roots are active at all times of the year. There
are bursts of seasonal activity in the early spring and in the summer when new roots
are formed in quantity. The root system is a rather specialized type. The active absorp-
tion is carried out by what are called fibrous or feeding roots. They are rather coarse,
mycorrhizal, usually devoid of functional root hairs, closely branched but of limited
growth, often forming a mat close to the soil surface, and developing with great vigour
in organic matter. When actively growing they are white, becoming yellowish brown
with age, due to the suberization of the walls of the outermost cells, but active absorp-
tion of water and solutes goes on through this suberized layer (Kramer, 1945). The
length of time for which these roots are functional has not been determined, but it
probably exceeds one year. The health of the citrus tree depends on the condition of
the fibrous roots and on its ability to produce new roots in abundance. If Phytophthora
is present and periodically destroys a percentage of these, the tree will be reduced in
vigour, and if the process is unimpeded a stage will be reached when the number of
roots lost exceeds the new roots formed. When this balance is upset, the deterioration
in tree health will be evident and will proceed with increasing speed. It can be assumed
that the pathogen is now so widely distributed that it is always present in the soil in
which a citrus tree is growing, any factor, therefore, which favours the fungus will be
unfavourable for the tree, anything which promotes root growth will diminish the
importance of the fungus attack. The level of nutrition is of obvious importance since
good nutritional status encourages abundant root growth. Young trees whose roots are
spreading into new uninfected soil are more likely to survive an attack following
unusually high soil moisture than old trees whose new roots are made within the orbit
of its already formed root system. The importance of fluctuations of soil moisture
needs no emphasis.

A young tree planted in infested soil or with already infected roots is facing a very
heavy handicap, and only in soils quite unfavourable to the fungus—i.e., very freely
draining types—can it be expected to grow into a mature tree.

The advantage of even a slight degree of resistance is very obvious. The Seville
orange is an example of this; under glasshouse conditions, where everything favours
the pathogen, the Seville is readily attacked. P. citrophthora can invade the tissues at
a rate slightly less than half that at which it can invade those of the rough lemon. This

XVili PRESIDENTIAL ADDRESS.

means that in the field conditions favovring the fungus must be operative for much
longer for it to affect the Seville than for the rough lemon, thus giving the host much
more opportunity for throwing off the attack. In practice this gives the Seville a very
high degree of resistance, much more than would be expected purely from controlled
infection experiments. Similarly with the lemon and the rough lemon. Under conditions
in which the fungus can attack and rot the bark of the lemon, the rough lemon may
remain unaffected, so that the collar rots which commonly affect the lemon in coastal
orchards do not. often invade the tissue of the stock.

I have gone into the history of the disease in some detail because it shows clearly
the importance of the fungus as a determining factor in citrus culture. Particularly in
New South Wales, where the stock used is a susceptible one, it is responsible for the
distribution of the crop. Well drained sands and sandy loams are spoken of as “good”
citrus soils and it is clear that the reason for this is that P. citrophthora makes little
or no progress in such soils, and the small amount of damage which it occasionally does
is quickly repaired. Damp rather heavy soils are spoken of as unfit for citrus, but this
is so only because in these soils P. citrophthora is active and its continual ravages
gradually upset the balance of loss against gain of new roots. It is not because citrus
trees, even on rough lemon stock, will not grow in such soils, in fact they grow very well
until P. citrophthora overtakes them.

References.

ALDERTON, G. E., 1884.—Treatise and Handbook of Orange-Culture in Auckland, New Zealand.

BARNARD, C., 1936.—Citrus Decline in the Murrumbidgee Irrigation Areas. Comm. Aust. Jour.
Coun. Sci. Ind. Research, 9: 163-170.

DARNELL-SMITH, G. P., and MAcKINNON, E., 1916.—Fungus and Other Diseases of Citrus Trees.
Dept. Agric. N.S.W., Farmers’ Bulletin No. 90, Citrus Culture, 90-120.

Fawcett, H. S., 1913.—Two Fungi as Causal Agents in Gummosis of Lemon Trees in California.
Phytopath., 3: 194-195.

————, 1923.—Gummosis of Citrus.—Jour. Agric. Research, 24: 191-236.

————, 1936.—Citrus Diseases and their Control.

and Brrancourt, A. A., 1940.—Occurrence, Pathogenicity and Temperature Relations
of Phytophthora Species on Citrus in Brazil and Other South American Countries. Arquivos
do Instituto Biologico, 11: 107-118.

FRASER, L. R., 1942.—Citrus Decline in the Murrumbidgee Irrigation Area. Agric. Gazette N.S.W.,
53: 415-419.

HAMBLIN, C. O., 1920.—Collar Rot of Citrus Trees. Ibid., 31: 439-441.

IRRIGATION RESEARCH EXTENSION COMMITTEE, 1943.—Report on Citrus Decline in the Murrum-
bidgee Irrigation Areas.

Kuotz, L. J., and Fawcett, H. S., 1930.—The Relative Resistance of Varieties and Species of
Citrus to Pythiacystis Gummosis and Other Bark Diseases. Jour. Agric. Research, 41:
415-425.

KRAMER, F. J., 1945.—Absorption of Water by Plants. Bot. Review, 11: 310-355.

Lewcock, H. K., 1935.—Pineapple Wilt Disease and its Control. Queensland Agr. Jour., 43: 9-17.

McALPINgE, D., 1899.—Fungus Diseases of Citrus Trees in Australia and their Treatment.

Moore, C., 1868.—Report on the State and Management of the Orange Plantations in Spain and
Portugal. N.S.W. Parliamentary Papers, 1868.

SIRAG-EL-DIN, A., 1934.—Citrus Gummosis in Egypt. Ministry of Agric. Egypt, Technical and
Scientific Service, Bul. 131.

SmitH, R. E., and SmitrH, FE. H., 1906.—A New Fungus of Economic Importance. Bot. Gaz.,
42: 215-221.

SWINGLE, W. T., and WesBBER, H. J., 1896.—The Principal Diseases of Citrus Fruits in Florida.
U.S. Dept. Agr. Divn. Veg. Phys. and Path., Bull. 8.

TOLKOWSKY, S., 1938.—Hesperides: A History of the Culture and Use of Citrus Fruits.

WacerR, V. A., 1942.—Phytophthora cinnamomi and Wet Soil in Relation to the Dying Back of
Avocado Trees. Hilgardia, 14: 517-532.

WEBBER, H. J.. and BATCHELOR, L. D., 1943.—The Citrus Industry, Vol. 1, History, Botany and
Breeding.

WHITE, R. P., 1937.—Rhododendron Wilt and Root Rot. New Jersey Agr. Exp. Sta., Bull. 615.

PRESIDENTIAL ADDRESS. xix

The Honorary Treasurer, Dr. A. B. Walkom, presented the Balance Sheets for the
year ended 28th February, 1949, 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.

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

President: R. N. Robertson, B.Sc., Ph.D.

Members of Council: D. J. Lee, B.Sc., R. N. Robertson, B.Sc., Ph.D., E. Le G.
Troughton, C.M.Z.S., F.R.Z.S., H. S. H. Wardlaw, D.Sc., F.A.C.I., W. L. Waterhouse, M.C.,
D.Se.Agr., D.1.C., and A. R. Woodhill, B.Sc.Agr.

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

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

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AUSTRALIAN FORMICIDAE.
NEW GENERA AND SPECIES.
J. J. McAREAVEY, S.J.
(Seventy Text-figures. )

[Read 30th March, 1949.]

INTRODUCTION.

All but one of the new species of ants described in this paper have been collected
by Mr. J. W. T. Armstrong of Nyngan, N.S.W. During 1946, in the area around Nyngan,
over 160 different species have been collected, and it is interesting to note that the larger
Subfamilies, Cerapachyinae, Ponerinae, Myrmicinae, Dolichoderinae and Formicinae are
all well represented. There are about thirty new species, but unfortunately in the case
of several of the obviously polymorphic species, only one form has been located, so that
the descriptions of such species have to be delayed to avoid future confusion.

With some hesitation the new genus Nchizopelta is proposed for a very interesting
Myrmicine, collected by Mr. Armstrong in October, 1946. In certain notes this ant
resembles specimens of Vollenhovia, but the conformation of the clypeus is so peculiar
that it seems best to place the species in a new genus and await the decision of other
myrmecologists. Only one male has so far been collected, and consequently certain
details of the anatomy have been passed over rather than risk damaging this specimen
by more extensive examination. A considerable number of workers, however, have been
examined, and very little variation was found.

The species of the second new genus, Stenothorar, was collected on the top of a
sandy hill near Greensborough, Victoria, in 1938. A complete colony was taken, but
unfortunately during the years all specimens but three have passed through various
collections, and at the moment cannot be traced. In the paper the genus has not been
placed in a tribe, though it appears to be close to both Solenopsidini and Pheidolini.

Subfamily PoNERINAE Lepeletier 1836.
Tribe ODONTOMACHINI Mayr. 1862.
Genus ANOCHETUS Mayr 1861.
ANOCHETUS ARMSTRONGI sp. nov. (Text-figs. 1-6.)
Worker.

Length 5:5-6 mm. Head, thorax, node, legs, mandibles and antennae clear yellowish
red, with the vertex of the head darker brownish red; eyes, tips of mandibular teeth and
anterior border of the pronotum black; gaster deep reddish brown with the apex paler.

Head smooth and shining except for the longitudinal fine striae on the area between
the frontal carinae. Thorax shining and smooth except for the dorsum of the epinotum
which is very finely and transversely striate. Petiole, gaster and legs smooth and shining.
Scape with a few longitudinal ridges.

Hair confined to the inner border of the mandibles where it is long, erect and
yellowish. Pubescence yellow, very short, adpressed confined to the funiculus and tibiae.

Head, excluding the mandibles, as long as the broadest part, that is the line from
eye to eye, slightly broader in front than behind, sides straight except for the distinct
bulge at the region of the eyes, occipital border deeply concave; mandibles as long as
three-quarters of the length of the head, curved downwards abruptly at the tips, and
terminating in three teeth, of which the two outer ones are sharp and twice as long
as the centre one; clypeus short, narrow with the anterior border straight; frontal
carinae diverging behind and extending beyond the eyes while in front they form two

2 AUSTRALIAN FORMICIDAE,

flattened lobes; frontal groove distinct and terminating behind in a deep pit; eyes large,
placed just before the centre of the sides of the head; scapes extend beyond the occipital
lobes by twice their thickness; first and third segments of the funiculus equal, and
longer than the second, all segments longer than broad, the apical longer than the two
preceding together.

5
Text-figures 1-6, Anochetus armstrongi sp. nov.
1. Worker head. 2. Worker head in profile. 3. Worker, dorsal view. 4. Worker, lateral

view. 5. Female, dorsal view of thorax. 6. Female wings.

Thorax more than twice as long as the broadest part; pronotum slightly longer than
broad, with very convex sides and convex anterior border, which is slightly margined;
mesonotum broader than long, three-quarters as broad as the pronotum with feebly
convex sides; all thoracic sutures clearly impressed; epinotum more than twice as
long as broad, the dorsum is broader behind than in front. In profile the pronotum and
mesonotum form an even convexity with a slight dip at the promesonotal suture.
There is a marked depression before the epinotum. Dorsum of the epinotum straight,
lower behind than in front, one and a half times as long as the feebly concave declivity.
The epinotal corner is almost right angled. Petiole about three times as broad as long,
deeply concave on top. In profile higher than the epinotum, about twice as high as broad
at the base, and twice as broad at the base as at the top, the faces are almost straight.
Postpetiole slightly longer than broad, broader behind than in front, with convex sides
and anterior border. There is a deep constriction separating the postpetiole from the
first gastric segment. Sting long. Legs long and slender.

Female.

Length 6-6-8 mm. Colour as in the worker, though some examples have the legs,
mandibles and antennae more yellowish. :

Pilosity and sculpture of the worker, except that the striae on the epinotum are
confined to the posterior half.

Very similar to the worker, but the scape is shorter, not reaching the occipital
border by its thickness. Ocelli yellowish, small but distinct.

Thorax slightly more than twice as long as broad; pronotum almost twice as broad
as long, convex and margined in front, the sides feebly convex; mesonotum slightly
broader than long, the parapsidal furrows not impressed; all thoracic sutures deep;
scutellum large, twice as broad as long; postscutellum indicated; epinotum very slightly

co

BY J. J. MCAREAVEY.

longer than broad, with almost straight sides, and feebly concave posterior border.
Petiole as in the worker except that the dorsum is entire and almost straight. Rest as
in worker. Collected by J. W. T. Armstrong.

Type locality: Nyngan, N.S.W.

Material examined: Hight females and sixty workers. There are slight variations
in colour. The specimens are much larger than those of A. rectangularis Mayr, or of
Forel’s variety diabolus, and distinguished easily by the shape of the node.

Types: Holotype worker and female in C.S.I.R., Canberra.

Subfamily Myrmicinakr Lepeletier 1836.
Genus STENOTHORAX gen. NOV.
Worker.

Small. Mandibles large, triangular, and strongly dentate. Clypeus narrow with a
slight depression at the centre in front. In profile it is seen to project slightly over
the mandibles. The frontal carinae are broadly flattened and cover the insertions of the
scapes. Between the lobes, formed by the carinae, is a deep broad groove extending as
far back as the end of the carinae. Antennae eleven segmented, the scape reaching just
beyond the eyes. First segment of the funiculus as long as the four following which are
broader than long. There is a three-segmented club with the apical segment longer
than the preceding two segments. The eyes are very small, placed about the middle of
the sides of the head. The head is almost square. The thorax has no sutures on the
dorsum, but is greatly narrowed and depressed about the meso-epinotal area. The
pro-mesonotum is large in front and tapers away almost to a point. The epinotum is
roughly wedge-shaped without a distinct posterior border, since the dorsum is concave
and merges into the sloping declivity. There are no spines or teeth, but the lateral
border is sharp. The stalk of the first node is long and slender, but the nodes and gaster
are closely placed together. The nodes are cubic when seen from above with the first
node much smaller than the second. Gaster ovate. The anterior tibiae have pectinate
spurs, the middle and hind tibiae without spurs. Claws simple.

Worker major: Head very similar to that of worker. The clypeus is not notched
in the centre nor overhanging the mandibles. The frontal carinae are shorter. The
eyes, placed at the centre of the sides of the head, are very large, hemispherical. There
are two very distinct ocelli but no trace of the third. The rest of the body is exactly
the same as that of the worker though slightly larger. There are no traces of sutures
on the dorsum of the thorax. The legs are the same as in the worker.

Genotype: Stenothorax katerinae sp. nov.

STENOTHORAX KATERINAE Sp. nov. (Text-figs. 7-16.)
Worker minor.

Length 3 mm. Uniformly light reddish brown with the legs slightly paler.

Mandibles smooth and shining; clypeus and the space between the frontal carinae
faintly shagreened; head including the flattened frontal carinae longitudinally finely
rugose with the spaces between the rugae smooth and shining; pronotum finely rugose,
the rugae having a circular direction, the spaces smooth and shining; epinotum fainily
sculptured, microscopically reticulate-punctate; sides of pronotum transversely striate;
sides of rest of thorax and the whole of the stalk microscopically reticulate-punctate;
petiole, postpetiole and gaster very faintly shagreened; legs smooth.

Hair golden, long, erect on head and body, suberect on legs, abundant but never
hiding the sculpture. Pubescence paler and confined to funiculus, tibiae and tarsi.

Head without the mandibles square, with the sides and the occipital border almost
straight, the corners rounded; mandibles large triangular, with six strong teeth, the
apical three being stronger than the other three; clypeus narrow with anterior border
rounded. There is a faint depression at the centre of the anterior border. Frontal
carinae broadly flattened, forming two lobes that overhang the clypeus. Between the
lobes is a deep rectangular groove, twice as long as broad. The scapes extend to the
posterior quarter of the head, and are not thickened; first segment of the funiculus

Cc

4 AUSTRALIAN FORMICIDAE,

almost as long as the four following, second to sixth broader than long, seventh longer
than broad, the three following segments form a distinet club, of which the apical
segment is as long as the two preceding together; eyes very small, placed just behind
the middle of the sides of the head.

Thorax twice as long as broad and has no sutures; pro-mesonotum large, longer
than broad, in front slightly narrower than the head and tapering away to a point
behind, the anterior border rounded and without marked corners. The small area
between the pro-mesonotum and the epinotum strongly depressed, from this arises the
peculiar epinotum. It is wedge-shaped, and the dorsum is concave, so that it is difficult
to say where it emerges into the epinotal declivity. At its broadest part it is less than
half as broad as the pro-mesonotum. The lateral borders are sharp so that from
certain angles the posterior corners seem to be sharp. In profile the pro-mesonotum
is strongly convex, flattened slightly at the middle. There is a very small raised
metanotum which is not noticeable in the dorsal view. The epinotum is low, with the
dorsum straight and sloping into the slightly shorter and feebly concave declivity. The
stalk is long and narrow; petiole almost square, slightly narrower in front than behind,
the anterior angles rounded, the sides and posterior border straight. There is hardly
any stalk between the petiole and postpetiole. The postpetiole is larger than the petiole,
square, with the faces straight and the corners rounded. In profile the petiole is higher
than the postpetiole, as high as long, with the anterior face feebly convex, dorsum sharply
convex, and the posterior face straight. The postpetiole is dome-shaped. Gaster ovate.
Legs short, robust, with femora and tibiae slightly thickened.

Worker major.

Length 3-8 mm. Head reddish brown with the area around the ocelli very dark,
almost black; mandibles except for the reddish tips, antennae reddish brown, thorax light
reddish yellow with the legs pale yellow; petiole and postpetiole brownish yellow and the
gaster dull yellow. Hair golden, long, erect on body and suberect on legs, abundant
throughout, but not hiding the sculpture. Pubescence pale, confined to funiculus, tibiae
and tarsi.

Mandibles smooth with a few faint striae. Rest of the sculpture as in the worker
but the circular rugae on the pronotum are more pronounced, and the punctuation on
the epinotum very clear. The anterior coxae have a few longitudinal ridges, and the
sides of the petiole are densely and microscopically reticulate punctate.

Head almost square with almost straight occipital border and feebly convex sides;
clypeus not notched at the centre, and not overhanging the mandibles; mandibles as in
worker; frontal carinae as in worker, but not extending back quite so far; antennae as
in worker, but the scapes slightly thickened, extending as far as the posterior border of
the eyes; eyes very large, hemispherical, and placed at the centre of the sides of the head.
There are two very distinct ocelli but no trace of a third, nor even a pit for an ocellus.

Thorax is very similar to that of the worker, and has no sutures of any kind. In
profile the pro-mesonotum appears to be flatter, the small metanotum is hardly visible,
the dorsum of the epinotum is not so sloping. The petiole and postpetiole are very
similar to those of the worker. In profile they are almost the same size, the anterior
face and dorsum of the petiole form almost a single convexity. Rest as in worker.

Collected by J. J. McAreavey, S.J.

Type locality: Greensborough, Victoria.

Types: Holotype worker in collection at C.S.I.R., Canberra.

Tribe METAPONINI Forel 1911.
Genus METAPONE Forel 1911.
METAPONE TRICOLOR sp. nov. (Text-figs. 17-19.)
Female.

Length 7:2 mm. Head except for the mandibles and clypeus, which are reddish,
thorax and petiole dark chestnut (almost black in parts). The postpetiole is very dark
brown but lighter than the thorax. The first segment of the gaster is brownish red. The
rest of the gaster is yellowish red, The antennae and legs are yellowish.

BY J. J. MCAREAVEY. 5

Head smooth and shining except for the antennal grooves and the lateral portions
of the clypeus, which are evenly and strongly striate longitudinally. There are faint
traces of striae on the central portion of the clypeus. The thorax, petiole and post-
petiole are smooth and shining, with scattered piligerous microscopic punctures, which

22

23

Text-figures 7-25.

7-16, Stenothorax katerinae sp. nov. 7. Worker major, head. 8. Worker major, dorsal view
cf body. 9. Worker major, lateral view of head. 10. Worker major, funiculus. 11. Worker
major, lateral view of body. 12. Worker minor, head. 13. Worker minor, dorsal view of body.
14. Worker minor, funiculus. 15. Worker minor, lateral view of head. 16. Worker minor,
lateral view of body.

17-19. Metapone tricolor sp. nov. 17. Female, dorsal view. 18. Female, lateral view.
19. Female, wings.

20-25. Xiphomyrmex capitalis sp. nov. 20. Worker, head. 21. Worker, funiculus. 22.
Worker, lateral view. 23. Worker, dorsal view. 24. Female, dorsal view. 25. Female, wings.

cc

6 AUSTRALIAN FORMICIDAE,

are more numerous on the petiole and postpetiole. The sides of the thorax are striate.
The upper half of the sides of the petiole is striate, while the lower half is densely and
microscopically punctate. The sides of the postpetiole are densely reticulate punctate.
The gaster is smooth and shining with faint traces of reticulation.

Head, including the mandibles, nearly twice as long as broad, slightly narrower in
front than behind, with the sides and the posterior border straight; mandibles large, with
five strong sharp teeth; clypeus, flat, narrow, and projecting slightly over the mandibles.
The anterior border is feebly concave, and produced at each corner in a small tooth. The
sides are margined but there is no distinct posterior border. There is a deep groove
extending from the clypeal area to the anterior ocellus. Eyes are large, but placed low on
the sides of the head, so that they are not visible from above; ocelli yellow, hemispherical,
placed near the posterior border of the head; scapes stout, flattened and very short;
funiculus with an enlarged three-segmented club; the first segment nearly as long as
the three following, second to seventh very much broader than long, apical segment as
long as the two preceding together.

Thorax long and slender, two and a half times as long as broad, pronotum one and a
half times as broad as long, with the humeri bluntly angular, the anterior border and
sides almost straight; mesonotum. slightly longer than broad, slightly broader than the
pronotum, with the sides feebly convex; scutellum rather large, broader than long; wings
short, the veins extending to the apex of the wings; epinotum broader in front than
behind, almosi twice as long as the posterior border, the sides straight, and the
posterior border feebly concave. In profile the thorax is very low, at least three times
as long as high, with the dorsum almost straight. The epinotal declivity is slightly
concave, and one-third as long as the epinotal dorsum, into which it passes through an
abrupt curve.

Petiole twice as long as broad in front, broader behind than in front, with anterior
border and sides straight and the posterior border concave and terminating on each
side in a sharp tooth. In profile the anterior face is concave, the dorsum flat, and the
posterior face concave. The petiole is about one and a half times as long as high, and
on the ventral surface is a long sharp projection. Postpetiole is square with the sides
feebly convex and the corners rounded. In profile the anterior face is convex and merges
into the flat dorsum, while the posterior face is concave. On the ventral surface are
two massive, bluntly pointed projections. Gaster long and narrow, broader than the
postpetiole by only half the width of the gaster. First segment longer than broad.
Legs are short and the femora greatly enlarged. Collected by J. W. T. Armstrong.

Type locality: Nyngan, N.S.W.

Material examined: One female.

This species resembles M. leae Wheeler, but there are noticeable differences in colour
and sculpture. It differs also in the shape of the petiole, the width of the gaster,
the length of the first segment of the funiculus and the outline of the head.

Type: Holotype female in collection C.S.I.R., Canberra.

Tribe TETRAMORIINI Emery 1895.
Genus XIPHOMYRMEX Forel 1887.
XIPHOMYRMEX CAPITALIS sp. nov. (Text-figs. 20-25.)

Worker.

Length, 3-3-2 mm. Head almost black with the antennae and mandibles yellowish;
thorax, petiole and postpetiole brownish yellow; gaster and legs yellowish. There is
considerable variation in colour. Some examples have the thorax and nodes very dark,
almost as dark as the head, other examples have the head brownish with a darker area
on the vertex, and this dark area varies in size among such examples.

Mandibles finely longitudinally striate; head longitudinally striate rugose with the
spaces between the rugae feebly shagreened; thorax rugose longitudinally, the rugae
joined by transverse ridges to form large rectangular messes. The epinotum is more
irregularly rugose. The spaces between the rugae densely and microscopically punctate.

BY J. J. MCAREAVEY. 1

Petiole and postpetiole are densely microscopically punctate with a few longitudinal
ridges; gaster densely but faintly microscopically punctate.

Hair yellow, erect, long and abundant. Pubescence greyish, apparent only on the
funiculus.

Head very slightly longer than broad, with the sides and occipital border almost
straight, and the posterior angles rounded.

Mandibles large, triangular, with five or six sharp teeth; clypeus with the large
central portion raised and convex, the lateral portions narrow, with their posterior
borders raised to form strong ridges of the antennal sockets. The anterior border of
the whole clypeus is almost straight. Frontal area is not indicated; frontal carinae
rather long, flattened in front and widely separated, merging behind into the rugae of
the head. Scape does not quite reach the occipital border; first segment of the
funiculus as long as the three following, fourth to sixth as long as broad, apical segment
of the three segmented club as long as the two preceding together; eyes large, convex,
placed at the middle of the sides of head.

Thorax one and a quarter times as long as the anterior border of the pronotum,
which is the broadest part of the thorax. There are no thoracic sutures, though there is.
a marked depression at the region of the meso-epinotal suture. The pro-mesonotal area
is rounded in front, with sharp corners and the sides feebly convex, more than twice as
broad in front as behind. Epinotum is one-quarter longer than broad, with feebly
convex sides and concave posterior border. The posterior angles are produced as long
sharp spines directed backwards and slightly outwards, and almost as long as the
interval between them. There are two shorter spines at the base of the declivity. In
profile the dorsum is convex, more so on the pro-mesonotum than on the epinotum. The
epinotal declivity is short and almost vertical, though the upper and lower epinotal
spines give it the appearance of being concave. Petiole margined, slightly broader than
long, convex on the sides, but almost straight on the anterior and posterior borders. In
profile it is slightly higher than long, rectangular, with all faces straight and the angles
sharp. Postpetiole is ovate, almost twice as broad as long; in profile dome-shaped, as
long as high, with convex faces. Gaster ovate. Legs long but robust.

Female.

Length 3:2-4-2 mm. Head, thorax, petiole and postpetiole dark reddish brown, with
considerable variation in the shades of brown. Some have the pronotal angles lighter
brown, others the sides of the head lighter than the vertex and the front of the head,
others have the mesonotum lighter than the rest of the thorax. Mandibles, antennae and
legs brownish yellow. In some examples the legs are much lighter than the antennae.
Gaster dark orange.

Mandibles smooth; head longitudinally striate with the spaces between the rugae
shagreened; pronotum with a few transverse striae; mesonotum, scutellum and epinotum
regularly longitudinally striate, the spaces between microscopically punctate, especially
between the epinotal spines; petiole and postpetiole microscopically punctate and longi-
tudinally rugose; gaster finely reticulate.

Hair as in worker.

Head similar to that of worker, but slightly broader than long; the frontal area
clearly indicated; the frontal carinae slightly more diverging behind; the scapes slightly
shorter. The eyes are placed a little behind the middle of the sides; ocelli brownish,
large and distinct.

Thorax is more robust, nearly twice as long as broad. The pronotum from above is
narrow, three times as broad as long, rounded in front, the anterior corners sharp;
mesonotum slightly longer than broad; all thoracic sutures deeply marked; scutellum
elliptical, broader than long. The epinotum is slightly broader than long, the sides
straight and the posterior border feebly concave. The epinotal spines similar to those
of worker, though more robust. In profile the mesonotum and scutellum very feebly
convex, almost straight. The epinotum is lower and almost straight, as long as the
straight declivity. Petiole, postpetiole and gaster as in worker.

8 AUSTRALIAN FORMICIDAE,

Collected by J. W. T. Armstrong.

Type locality: Nyngan, N.S.W.

Material examined: Six females and ten workers. The workers show considerable
variation in colour.

This species differs from X. turneri Forel in colour and in the sculpture of the nodes
and gaster. The posterior border of the head is straight, not strongly concave. The
eyes are placed at the middle not before the middle of the sides of the head. The petiole
is broader in front than behind. The specimens are larger than those of X. turneri var.
fuscipes Viehmeyer, and the petiole and the postpetiole are broader, and the sculpture
is different.

Types: Holotype worker in collection at C.S.I.R., Canberra.

Genus pAcRYON Forel 1895.
DACRYON MARGINATUS sp. nov. (Text-figs. 26-29.)
Worker.

Length, 45-5 mm. Head, antennae, thorax, petiole and postpetiole rich red, gaster
‘darker, more reddish brown; legs brownish with the coxae and tarsal joints reddish;
eyes, margins of head, thorax and petiole black.

Mandibles finely longitudinally striate; head densely and microscopically punctate.
There are seven longitudinal ridges between the frontal carinae, and these ridges extend
to the occipital border. Antennae, legs, neck, thorax, petiole, postpetiole and gaster
microscopically punctate. There are a few longitudinal ridges on the thorax and nine
coarser ridges on the postpetiole. First segment of the gaster is finely longitudinally
striate.

Hair yellowish, confined to the clypeus and apex of the gaster. Pubescence greyish,
adpressed, abundant on the funiculus.

Head very slightly longer than broad, with the sides feebly convex, occipital border
feebly concave, the angles rounded. Mandibles triangular, with five sharp teeth; clypeus
almost straight, with a slight incision in the centre; frontal area hardly noticeable;
frontal carinae widely separated, raised and covering the insertions of the scapes, and
extending back almost to the occiput; scape short, extending beyond the eyes by their
width; first segment of the funiculus as long as the two following, second to seventh
as broad as long, the remaining three segments thickened, giving the appearance of a
club, the apical segment longer than the preceding but not as long as the two preceding
together; eyes large, convex, placed at the middle of the sides.

Thorax is almost twice as long as the broadest part, which is the anterior border of
the pronotum. Pronotum and mesonotum strongly margined, with anterior border convex,
the pronotal angles produced forwards and outwards as blunted teeth; the sides as
far as the mesonotal region almost straight. At that point on each side there is a short
triangular tooth, after which the dorsum narrows to half the width of the anterior
border of the pronotum. Meso-epinotal suture is deep. The strongly margined epinotum
is longer than broad, with convex sides and feebly concave posterior border. The
posterior corners are produced in two long-blunted spines, directed outwards and back-
wards, almost as long as their interval. In profile the pro-mesonotum is strongly convex,
the mesonotal tooth just noticeable. The epinotum is feebly convex, about twice as long
as the straight declivity. The epinotal spines almost as long as the declivity, directed
upwards and slightly backwards, the edge facing the dorsum straight, the other edge
convex.

Petiole broader than long, oval, the margined dome is convex and produced behind in
two spines which are directed slightly upwards as well as backwards. In profile longer
than high, wedge-shaped. The anterior face is straight, and sloping inwards to the apex
of the node and terminating above in a short, sharp spine. Dorsum almost missing,
the posterior face concave. Postpetiole broader than long (one and a half times), with
convex borders. In profile it is longer than the petiole, dome-shaped; the anterior face
almost straight, and parallel to the anterior face of the petiole; the dorsum and the

BY J. J. MCAREAVEY. 9

posterior face convex. Gaster ovate. Legs short and robust, the femora very strongly
thickened, the tibiae also thickened but not so much.

Collected by J. W. T. Armstrong.

Type locality: Nyngan, N.S.W.

Material examined: Three specimens of worker.

Text-figures 26-41.

26-29. Dacryon marginatus sp. nov. 26. Worker, dorsal view. 27. Worker, head. 28.

Worker, funiculus. 29. Worker, lateral view.
30-35. Monomorium (Holcomyrmex) armstrongi sp. nov. 30. Major worker, head. 31.

Minor worker, head. 32. Major worker, lateral view. 33. Minor worker, lateral view. 34. Male,

lateral view. 35. Female, dorsal view.
36-41. Monomorium (Holcomyrmex) niger sp. nov. 36. Major worker, head. 37. Major

worker, lateral view. 38. Minor worker, lateral view. 39. Minor worker, head. 40. Male, lateral
view. 41. Female, lateral view.

10 AUSTRALIAN FORMICIDAE,

Species nearest to D. ferruginea Clark but distinguished by the clear black margins
on head, thorax and petiole, the sculpture and the colour. The scape is shorter, the
petiole broader than long, the postpetiole broader, and in profile the nodes are quite
distinct.

Type: Holotype worker in collection at C.S.I.R., Canberra.

Tribe SoLENOPSIDINI Forel 1913.
Genus mMoNoMORIUM Mayr 1855.
Subgenus HOLCOMYRMEX Mayr 1878.
MONOMORIUM (HOLCOMYRMEX) ARMSTRONGI Sp. nov. (Text-figs. 30-35.)
Worker major.

Length, 6-8-7-2 mm. Black; mandibles and front of head more brownish black;
articulations of the legs, the tarsal segments, apex of gaster, tip of the apical segment of
the funiculus reddish.

Mandibles smooth and shining with a few small punctures; clypeus smooth with
scattered punctures; head smooth behind, but in front are fine longitudinal striae which
are coarser between the frontal carinae. Thorax reticulate-punctate, and also has
longitudinal fine striae which have a circular direction on the pronotum and a transverse
direction on the epinotum; petiole and postpetiole densely and finely punctulate; gaster
very finely and densely reticulate.

Hair yellowish, long, erect, sparsely scattered throughout, shorter and sub-erect on
the legs. Pubescence very short and fine, confined to the funiculus.

Head large, slightly longer than broad, almost as broad behind as in front, with
sides convex and the occipital border slightly concave, distinctly excised in the middle,
corners broadly rounded. Mandibles large, triangular, with five sharp teeth. Clypeus is
narrow on the sides, but the central portion is raised and bordered by two flattened
carinae which are produced forward in two sharp teeth. The interval between the
teeth is concave. Close to the outer side of these teeth is a much smaller tooth; frontal
carinae short and prominent, diverging at their posterior fourth, but parallel on anterior
three-quarters of their length; frontal area large, impressed with a narrow frontal
groove extending back as far as the level of the eyes; scape not thickened towards apex
and extending to the posterior quarter of the head; first segment of the funiculus
slightly longer than the second, all segments longer than broad. There is no club but
the last three segments are large, twice as long as broad, the apical slightly longer
than the second last; eyes large but very flat, placed about the centre on the sides of
the head.

Thorax broadest through the pronotum which is only half as broad as the head;
pronotum as broad as long, convex in all directions; promesonotal suture not distinct;
mesonotum short, broader than long, broader in front than behind with almost straight
sides; meso-epinotal suture impressed; epinotum about two-thirds as broad as the
pronotum, slightly longer than broad, with the sides straight and almost parallel. There
are two blunt tubercles directed upwards, towards the middle of the posterior border of
the epinotum, one each side of a concave depression. In profile the pro-mesonotum is
rather conical with the dorsal part of the mesonotum and the epinotum much lower
and sloping downwards towards the petiole. The dorsum of the epinotum is feebly
convex and slightly longer than the almost straight and vertical declivity.

Petiole pedunculate; the dorsum almost square, but the whole node is about twice
as long as broad. Postpetiole almost circular, very slightly broader than the petiole.
In profile the petiole is dome-shaped, with anterior face very feebly convex, the anterior
angle sharp, the dorsum, which is half as long as the base of the petiole, is feebly convex
and rounded into the posterior convex face. Postpetiole lower, dome-shaped, and convex
on all sides. Gaster large, elliptical. A small sting is exposed in some examples. Legs
long and robust.

Worker minor.
Length, 5-7-6-2 mm. Colour and pilosity as in major.

BY J. J. MCAREAVEY. 11

Sculpture as in major but much finer. This is especially the case with the striae
of the thorax, which are merely indicated. Head as in major though proportionately
smaller, and the scapes very slightly longer. Thorax as in major but the dorsum of the
epinotum slightly longer, and the tubercles on the posterior border are missing. Legs
long and slender. Rest as in major.

Female.

Length, 10 mm. Colour of major but the front of the head and the mandibles bright
red and the funiculus is wholly black.

Sculpture of the head as in major but coarser; pronotum, mesonotum and scutellum
strongly striate-rugose longitudinally; epinotum transversely striate-rugose; petiole
almost smooth, reticulate-punctate on the sides; postpetiole with fine circular striae and
a very finely and densely reticulate-punctate ground sculpture; gaster reticulate.

Hair reddish, longer and more abundant throughout.

Head broader than long, with feebly convex sides and almost straight occipital
border. Mandibles as in worker, but the apical tooth is extremely long. Scape extends
to the posterior fifth of head. Ocelli pearl white and large. Rest of head as in worker.

Thorax one and a half times as long as broadest part, which is a line through the
middle of the mesonotum; pronotum short, not noticeable from above; mesonotum
massive, broader behind than in front with convex borders. The parapsidal furrows are
deeply impressed. Scutellum is broader than long, broader in front than behind; all
thoracic sutures deeply impressed; epinotum as long as broad at anterior border, which
is twice as broad as the concave posterior border. There are no spines or tubercles. In
profile the pronotum is vertical; the mesonotum and scutellum almost flat or very feebly
convex; epinotum almost straight and sloping; the declivity as long as the dorsum.

Petiole is slightly broader than long, convex on all faces. There is a slight hollow
in the centre near the posterior border. In profile the anterior face is convex, the
dorsum very feebly convex and the posterior face feebly concave. Dorsum of the post-
petiole is reduced almost to a straight line, but it presents the anterior face which is
as long as broad and convex in all directions; in profile wedge-shaped, with a long,
sloping straight anterior face meeting the straight almost vertical posterior face in a
point. Gaster large, elongate. Legs long.

Male.

Length, 5:1 mm. Entirely black except for the tarsi and the apex of the gaster,
which are reddish.

Hair black, long, abundant throughout. Pubescence greyish, short, confined to the
funiculus and the tarsi.

Head and promesonotum irregularly, closely and finely shagreened; scutellum has in
addition some fine longitudinal striae; epinotum reticulate-punctate with a few fine
transverse striae; petiole, postpetiole and first segment of the gaster finely densely
reticulate-punctate; rest of gaster more transversely striate.

Head longer than broad, with the sides and occipital border strongly convex.
Mandibles rather large, furnished with denticles. Clypeus appears to be rounded in
front, but it is difficult to see the front of the head, which is concealed by long black
hair. Scape is short, reaching to the eyes, which are very large and occupy most of
the side of the head. All segments of the funiculus are much longer than broad, except
the first, which is short, and just slightly longer than broad. Ocelli are pearl white,
hemispherical, and large.

Pronotum short and not visible from above; mesonotum longer than broad, with the
sutures indistinct; scutellum as long as broad, broader in front than behind; epinotum
as long as broad in front, where it is twice as broad as the posterior border.

Petiole longer than broad, with feebly convex borders, the corners broadly rounded;
in profile rather bluntly conical, with sloping, feebly concave anterior border and feebly
convex posterior border; postpetiole slightly longer than the petiole, slightly longer than
broad, and slightly broader behind than in front, the lateral borders feebly convex; in

12 AUSTRALIAN FORMICIDAR,

profile very much lower than the petiole, forming merely a low convexity. Gaster
elliptical. Legs long and slender.

Collected by J. W. T. Armstrong.

Type locality: Nyngan, N.S.W.

Material examined: Large complete series of all forms. Wheeler described the major
worker of Holcomyrmex whitei, the first of the subgenus to be found in Australia. His
major worker is much smaller than the minor worker of H. armstrongi and the colour
is red and yellow. There is no possibility of confusing either of the following two
species with H. whitei Wheeler.

Type: Holotype worker major, and series of worker minor, male and female, in
collection at C.S.I.R., Canberra.

MONOMORIUM (HOLCOMYRMEX) NIGER Sp. nov. (Text-figs. 36—41.)
Worker major.

Length, 5-2 mm. Entirely black.

Except for the smooth frontal area, the anterior third of the head and the portion
between the frontal carinae are finely striate. The rest of the head is smooth with
scattered microscopic piligerous punctures. Thorax is shining, rather smooth with scat-
tered microscopic punctures and a few traces of striae, which are longitudinal on the
pronotum and transverse on the epinotum; petiole smooth; postpetiole very finely
transversely striate and with traces of reticulation; gaster feebly shagreened.

Hair reddish, short, erect on head; a few long hairs on the petiole and postpetiole;
longer hairs on the gaster but none on the thorax. Pubescence fine, reddish, confined
to the legs and antennae.

Head slightly longer than broad, as broad behind as in front, with sides feebly
convex and the occipital border concave with a marked incision at the middle of the
posterior border. Mandibles large, triangular with five strong teeth; clypeus narrow
on sides, the central portion raised and bordered by two carinae which are produced
forward as two stout blunted teeth. Flanking the outer side of each tooth is a sharper
slightly shorter tooth. Frontal carinae are short diverging outwards slightly; frontal
area large and distinct, the suture between it and the central portion of the clypeus
deep. Scape extends to the posterior fifth of the head, thickened very slightly towards
the apex; first segment of the funiculus longer than the second, the rest as long as broad,
except the three apical segments, which are thickened, and at least twice as long as
broad, the apical segment slightly longer than the second last; eyes moderately large
and very flat, placed at the centre of the sides of the head. :

Thorax twice as long as broad though the pronotum, with both sutures obsolete,
rounded in front, and constricted at the region of the mesonotum, where there is a
deep depression. The meso-epinotal area is reduced to half the width of the convex
pronotum and has the sides almost straight, and the posterior border coneave. There
is a small tubercle at each posterior angle of the epinotum. In profile the dorsum of the
pronotum convex; the dorsum of the meso-epinotum almost straight and sloping; the
short epinotal declivity feebly concave.

Petiole and postpetiole equal, as broad as long with feebly convex borders. In
profile the petiole is as high as long, with sloping, feebly concave anterior face, feebly
convex dorsum which is rounded into the almost vertical posterior face. The post-
petiole is much lower and forms a dome with the posterior face feebly concave. Gaster
ovate. Legs long and slender.

Worker minor.
Length, 4mm. Black, but the anterior border of the head, mandibles, joints of legs
and tarsi brownish. Hair similar to that of major.
Sculpture as in major but the sculpture of the pronotum is more reticulate and the
epinotal declivity transversely striate; petiole and postpetiole reticulate-punctate; gaster
smooth with scattered elongated punctures.

BY J. J. MCAREAVEY. 13

Head as in major but the scape is slightly longer, and the eyes a little more convex
and placed slightly nearer to the anterior border of the head. Thorax is similar to that
of major but proportionately shorter with the pronotum larger and more rounded. The
tubercles of the epinotum are not noticeable from above. In profile the dorsum is much
flatter, almost straight. The epinotal tubercles are visible, giving the declivity the
appearance of being concave, though actually the upper half is straight and sloping,
the lower half bulges outwards towards the petiole convexly.

Petiole twice as broad as long, the borders almost straight; postpetiole with the
dorsum reduced to a convex line, showing a full view of the almost square anterior face.
In profile the petiole is higher than long, with sloping almost straight anterior face,
convex dorsum, and feebly convex posterior border. Postpetiole as in major, low and
dome-shaped.

Female.

Length, 65-7 mm. Black with mandibles and front of head yellowish red. The
gaster varies from black with dull reddish bands at the borders of the gastric segments
to dull red with black borders to the gastric segments.

Front of head except for the smooth frontal area finely longitudinally striate-rugose.
The posterior half of the head and the thorax have scattered small elongated piligerous
punctures, which are denser on the head than on the thorax. The epinotum is finely
transversely striate. Petiole and postpetiole feebly shagreened on top, but the pedicel
is densely microscopically reticulate-punctate. Gaster smooth with sparse elongated
punctures. Pilosity as in worker but more abundant.

Head as in major but is as broad as long. The frontal carinae diverge further
behind and are wider apart. The carinae of the clypeus are more pronounced, further
apart in front and terminating in larger teeth. The eyes are comparatively smaller.
The ocelli brownish, small and convex.

Pronotum hardly visible from above; mesonotum massive, as broad as long, feebly
convex; the parapsidal furrows deeply impressed; scutellum circular, very slightly
broader than long; epinotum much lower and very sloping, so that at times it is difficult
to distinguish the dorsum from the declivity; dorsum as long as broad, broader behind
than in front. There is a slight margin between the dorsum and the declivity with a
small blunt tubercle at each corner. In profile the pronotum is vertical, the mesonotum
and scutellum very flat, the scutellum overhangs the epinotum slightly. Dorsum of the
epinotum straight, as long as the slightly concave declivity.

Petiole is twice as broad as long, excised on top in the middle; postpetiole twice as
broad as long, longer than the petiole, elliptical and slightly concave on top. In profile
the petiole is higher than long, the anterior face convex, rounded into the feebly convex
dorsum, posterior face straight and vertical; postpetiole lower, anterior face and dorsum
form one convexity while the posterior face is concave.

Male.

Length, 7 mm. Black; joints of legs and gaster, except for the black basal segment,
yellowish red. The reddish part of the gaster varies in extent among individuals of the
same nest. °

Head longitudinally striate on those parts where the sculpture is not concealed by
hair; dorsum and sides of thorax coarsely reticulate-punctate; petiole and postpetiole
densely and microscopically punctate. There are some transverse striae on the posterior
face of the petiole. Gaster is smooth with traces of shallow punctures on the first
segment, and very fine reticulation on the other segments.

Hair reddish and long on gaster. It is black on the head and thorax, copious,
especially on the head where it is longest.

Head as broad as long, with strongly convex sides and occipital border. Mandibles
are narrow, with three large strong teeth; clypeus rounded in front, apparently without
any teeth or spines; scape short, extending to the posterior border of the eyes; first
segment of the funiculus slightly longer than broad, second very long, fully three times

14 AUSTRALIAN FORMICIDAE,

as long as first, all the remaining segments at least twice as long as broad; eyes very
large, hemispherical, occupying most of the sides of the head; ocelli pale yellow,
hemispherical, rather large.

Pronotum from above small and rounded in front; mesonotum very large, as long
as broad, with convex sides; scutellum circular; epinotum very short, and from above
it is similar to that of the female, but the two tubercles on the posterior border are more
distinet. In profile the pronotum and mesonotum form a hump, with the top of the
hump about the pro-mesonotal suture. The epinotum is much lower than the scutellum
which slightly overhangs it. The dorsum of the epinotum is longer than the feebly
concave declivity, the faces separated by the small tubercles.

Petiole as broad as long, almost circular. In profile as high as long, dome-shaped,
with the faces forming an even convexity; postpetiole almost the same size, but slightly
broader behind than in front. In profile it consists of an evenly convex low hump on
the pedicel. Gaster long and slender. Legs long and slender.

Collected by J. W. T. Armstrong.

Type locality: Nyngan, N.S.W.

Material examined: Large series of all forms.

This species resembles H. armstrongi but is easily distinguished by the size and
sculpture. The scape of the major is longer and the petiole and postpetiole are alike,
whereas they differ greatly in H. armstrongi. The male and female are separated from
those of the other species in colour and sculpture, shape of the nodes. The male is,
moreover, relatively much larger.

Type: Holotype worker major, and series of worker minor, male and female, in
collection at C.S.I.R., Canberra.

Genus SCHIZOPELTA gen. Nov.
: Worker.

Monomorphie. Mandibles triangular, moderately large with few strong teeth.
Maxillary and labial palpi two segmented. Clypeus narrow, but produced forward in
two long prongs, which are slightly longer than broad at their base. The exterior
border of the projecting part is convex, and the inner border concave. The frontal
carinae are straight and almost parallel. Between these frontal carinae is a deep
furrow, bordered on each side by a carina, and this whole frontal area is produced
forward over the clypeus, and terminates in two small teeth. In profile the frontal
carinae are raised, and the clypeus extends over the mandibles. The antennae have
twelve segments. The first segment of the funiculus is long, the apical three segments
are also long and form an indistinct club. Eyes moderately large and convex, placed
about the middle of the sides. No ocelli. The head is rather square and broader than
the thorax. There are no thoracic sutures, and the pronotal area is strongly convex,
and twice as broad as the epinotal area. The pronotum is rounded in front, the epinotum
armed with moderately long teeth. The petiole is almost square, hardly pedunculate.
The postpetiole is also cubic but larger than the petiole, very close to the petiole and
gaster so that it is very difficult to see the stalk. Gaster ovate. Legs robust, the femora
and tibiae hardly thickened. The anterior tibiae have pectinate spurs, the middle and
hind tibiae without spurs. The claws simple.

Pseudogyne.

Slightly larger than the worker. Head similar to that of worker and there is no
trace of ocelli. Thorax has no wing sclerites but the other sutures are well developed.
The pronotum is more angular in front, and the mesonotum is large, and slightly higher
than the rest of the thorax. The epinotal spines are a little shorter than those of the
worker, and the dorsum of the epinotum is strongly concave. Rest as in worker.

Male.
Mandibles short and narrow. Clypeus raised and overhanging the mandibles.
Instead of the forked projections, found in the worker and female, the clypeus has a
deep notch in the middle and this is margined by a carina which extends to the posterior

BY J. J. MCAREAVEY. 15

border of the clypeus. The frontal carinae are raised and almost parallel. There is a
frontal groove and a small but deep depression just in front of the anterior ocellus.
Ocelli large and the eyes very large and convex. The articulations of the scapes are
exposed. The scape is slightly more than twice as long as broad. The funiculus is
twelve segmented with the first segment short. The parapsidal furrows are impressed.
The anterior wings have open radial cells, single closed cubital cells and a small
discoidal cell. The venation of the posterior wings is very restricted. The nodes are
low and dome-shaped. The fore tibiae have pectinate spurs, middle and hind tibiae
without spurs. Claws simple.
Genotype: Schizopelta falcata sp. nov.

SCHIZOPELTA FALCATA Sp. nov. (Text-figs. 42-47.)
Worker.

Length, 3-3-5 mm. Reddish brown with legs, antennae and gaster more yellowish;
eyes black and the front of the head very dark brown, almost black.

A yellowish short pilosity is confined to the clypeus, antennae, tarsi, tibiae and apex
of gaster.

The head is covered with small shallow punctures, slightly elongated. The spaces
between the punctures are smooth and shining. The dorsum of the pronotum has similar
sculpture. The rest of the thorax, the petiole and the postpetiole densely microscopically
punctate. Gaster smooth and shining.

Head slightly longer than broad, with convex sides and concave posterior border.
Mandibles, hidden by the clypeus, are moderately large, with four stout teeth. Clypeus
is narrow, but produced forward in two long prongs, which are slightly longer than
broad, with convex outer and feebly concave inner borders. Frontal carinae are straight
and parallel. Between these carinae is a deep furrow, bordered on each side by a
carina which terminates in a small tooth. Scapes extend to the middle of the eyes.
The first segment of the funiculus is almost as long as the two following together,
second to eighth almost as broad as long. The apical segment of an indistinctly three-
segmented club is longer than the two preceding segments together. Eyes moderately
large, rather flat, are placed at the middle of the sides.

Thorax not quite twice as long as broad; pronotum broader than long, strongly
convex in all directions. The pro-mesonotal suture is hardly present in most examples.
The meso-epinotum is halt as broad as the pronotum, with no trace of meso-epinotal
suture. The epinotum terminates in two long teeth, directed upwards and backwards,
the space between concave. In profile the dorsum is feebly convex and in most examples
there is no trace of sutures.

Petiole is hardly pedunculate, slightly longer than broad, and very slightly narrowed
in front, with feebly convex sides and straight anterior and posterior borders; post-
petiole more rectangular, longer than broad, the anterior border feebly convex, the
remaining borders straight and the corners bluntly rounded. In profile the petiole is
slightly higher than the postpetiole. The petiole is slightly higher than long with
feebly concave anterior face rounded into the feebly convex dorsum. The postpetiole is
cubic with the upper corners rounded into the feebly convex dorsum. The basal segment
of the gaster occupies most of the gaster. Legs robust.

Pseudogyne.

Length, 4:2 mm. Head except for a dull red spot on the vertex, deep black. Antennae
reddish yellow; thorax, petiole and postpetiole brownish red; the lower margin of the
sides of the thorax, the coxae and the upper half of the femora of the anterior pair of
legs black; remaining part of the front legs, the other four legs, and the gaster light
yellowish red.

The sculpture of the head as in the worker but the flattened frontal carinae are
finely longitudinally striate. The pronotum and mesonotum have scattered elongated
punctures. The rest of the thorax, petiole and postpetiole are densely and micro-
scopically punctate. Gaster smooth and shining.

D

16 AUSTRALIAN FORMICIDAR,

Head as in worker, but it is square with the sides and occipital border almost
straight, and the posterior corners rounded. Hyes flat, moderately large, are placed at
the centre of the sides; no trace of ocelli.

Thorax with very distinct sutures; pronotum three times as broad as long, with
convex anterior border and almost straight sides. The anterior angles are rather sharp.
Mesonotum is raised, large, twice as long as broad; metanotum depressed slightly, narrow
and indistinct; epinotum as broad as long, slightly broader behind than in front. The

Text-figures 42-56.

42-47. Schizopelta falcata sp. nov. 42. Worker, head. 43. Worker, funiculus. 44. Worker,
head in profile. 45. Worker, dorsal view. 46. Worker, lateral view. 47. Male, lateral view.

48, 49. Dolichoderus (Hypoclinea) armstrongi sp. nov. 48. Worker, lateral view. 49.
Worker, head.

50-52. Camponotus (Myrmogonia) sanguinea sp. nov. 50. Major worker, head. 51. Minor
worker, head. 52. Major worker, thorax, lateral view.

53-56. Camponotus (Myrmogonia) armstrongi sp. nov. 53. Major worker, head. 54. Minor
worker, head. 55. Minor worker, dorsal view. 56. Minor worker, lateral view.

BY J. J. MCAREAVEY. 17

epinotal spines are shorter than those of the worker, the dorsum of the epinotum
concave. In profile the pro-mesonotum evenly convex, a little flattened at the pro-
mesonotal suture. Dorsum of the epinotum is flat, nearly twice as long as the declivity;
epinotal spines stout, directed upwards and slightly backwards; petiole, postpetiole and
gaster as in worker.

Male.

Length, 3:2 mm. Entirely black, except for the tips of the mandibles and the tip of
- the gaster which are dull reddish brown.

Hair yellowish, long, and abundant throughout.

Densely reticulate punctate on head, petiole and postpetiole. The thorax is more
coarsely rugose and the gaster smooth.

Head very slightly longer than broad across the eyes, with the sides strongly
convex, and the occipital border short and almost straight. Mandibles short and narrow,
dentate; clypeus raised and overhanging the mandibles, deeply notched in the middle;
frontal carinae straight and raised above the articulations of the scapes. Frontal groove
is faint but there is a deep pit just before the anterior ocellus. Scape is short, about
three times as long as broad; first segment of funiculus almost as broad as long, all
other segments twice as long as broad; eyes large hemispherical; ocelli distinct.

Thorax slightly more than twice as long as broad. The pronotum-is hardly notice-
able from above; mesonotum very large. The parapsidal furrows are impressed, but
difficult to see because of the coarse sculpturation. Scutellum is broader than long,
broader in front than behind; epinotum much lower than the scutellum, as broad as
long, broader in front than behind, with almost straight sides and posterior border.
There are no spines on the posterior border of the dorsum but merely sharp corners.
The small spine at the base of the epinotal declivity is present, as in worker. Petiole
and postpetiole are much more elongated and more rounded than in worker. In profile
these are low humps. Legs rather long and slender.

Collected by J. W. T. Armstrong.

Type locality: Nyngan, N.S.W.

Material examined: One male, one pseudogyne, and a large series of workers. The
pseudogyne was taken with four workers from an incipient colony, some time after the
male and a large series of workers had been examined from the same area.

Type: Holotype worker and allotype male in collection at C.S.I.R., Canberra.

Subfamily DoLIcHODERINAE Forel 1878.
Genus DOLICHODERUS Lund 1831.
Subgenus HYPOCLINEA Mayr.

Hypoclinea Mayr, Verh. Zool. Bot. Ver..Wien., 5, p. 377, 1855.
Hypoclinea sens. str. Wheeler, Psyche, 42 (1), p. 68, 1935.

DOLICHODERUS (HYPOCLINEA) ARMSTRONGI sp. nov. (Text-figs. 48, 49.)
Worker.

Length, 3-5-4 mm. Head, thorax and node rich brownish red (the head just slightly
darker than the thorax and node); eyes and margins of the node black; margins of the
thorax brownish black; antennae and legs light brownish red. In some examples the
femora are light yellowish red. Gaster is deep orange or yellowish red.

Mandibles smooth; head shining, coarsely reticulate-punctate, the bottoms of the
punctures smooth and shining. (There is no fine reticulation between the punctures as
in H. reflexus.) Antennae are smooth; thorax and node reticulate-punctate but on the
node the sculpture is finer. The epinotal declivity is smooth (it is feebly transversely
striate in H. turneri). Gaster is very finely reticulate (reticulate punctate in H. reflerus
and H. turneri). Legs are smooth.

Hair greyish yellow, long, erect on head, mandibles, clypeus and antennae, on other
parts of the body there is very little hair. No pubescence. (Abundant pubescence
covering the sculpture of the gaster in H. turneri.)

DD

18 AUSTRALIAN FORMICIDAE,

Head one-quarter longer than broad, convex feebly on the occipital border and
strongly so on sides, widest just behind the eyes, narrowed greatly in front. Mandibles
large, armed with three strong teeth in front and about five or six denticles behind;
clypeus almost straight on anterior border with a deep longitudinal impression in front,
the posterior border is very indistinct; frontal°area not noticeable in most specimens;
frontal carinae raised, very slightly diverging behind, slightly longer than their distance
apart. There is a trace of a scrobe at the side of the carinae, and extending as far as
the end of these carinae. (This appears to be absent in both H. reflexus and H. turneri.)
Scape, as in H. reflexus, extends beyond the occiput by a quarter of its length. (Slightly
beyond occiput in A. turneri.) Eyes globular, placed slightly behind the middle of the
sides. First segment of funiculus twice as long as the second, which is shorter than the
third. (In H. reflexus the first segment is one-fifth longer than the second, which is
longer than the third.)

Thorax twice as long as broad; pronotum nearly twice as broad as the rest of the
thorax, one and two-thirds broader than long with the sides feebly convex and the
anterior angles bluntly produced; mesonotum as long as broad, broader in front than
behind, with the sides feebly convex and margined; epinotum one and three-quarters
longer than broad, with the sides and posterior border straight. In profile the dorsum
of the pronotum and of the mesonotum are convex, with the pronotum slightly longer
than the mesonotum. Dorsum of the epinotum is very high, convex, and raised behind
where with the very concave declivity it forms a sharp point. The dorsum of the
epinotum overhangs the declivity by half its length. At the base of the declivity,
just in front of the node, there is a small spine. Node massive, in profile more than
twice as high as long, with the faces straight almost to the top, where abruptly they
slope in to meet at a blunt point. From above the node is twice as broad as long.
Gaster oval. Legs robust.

Collected by J. W. T. Armstrong.

Type locality: Nyngan, N.S.W.

Material examined: About forty workers which show very little variation in colour
and size. :

This species is near both H. turneri Forel and H. reflexus Clark, but differs slightly
in the structure of the thorax and node, and clearly differs in the points indicated in
the description.

Type: Holotype worker in collection at C.S.I.R., Canberra.

Subfamily FormicinarE Lepeletier 1836.
Tribe CAMPONOTINI Forel 1893.
Genus CAMPONOTUS Mayr 1861.
Subgenus myrMogonta Forel (Mann emend. 1921).
CAMPONOTUS (MYRMOGONIA) SANGUINEA sp. nov. (Text-figs. 50-52.)
Worker major.

Length, 8-3 mm. Mandibles and anterior border of head black or brownish black;
clypeus dark reddish brown; frontal carinae black; insertions of scapes reddish yellow;
antennae dark brown with the tip of the funiculus lighter; head, thorax and node dull
blood red, the epinotum is darker, more brownish; legs reddish brown with tibiae and
tarsi darker; first segment of the gaster red, second brownish red, rest of gaster brown.

Mandibles with scattered elongated punctures; clypeus finely, densely reticulate-
punctate; head very finely and densely striate transversely. The striae are joined
together to form a wide meshed reticulation. Thorax is striate punctate, on pronotum
more transversely arched, and coarser on the meso-epinotum; node transversely finely
striate; gaster transversely striate.

Hair long, erect, reddish, abundant on mandibles, epinotum node and gaster.

Head slightly longer than broad, with the sides almost straight, and the occipital
border straight, corners abrupt. Mandibles thick triangular, with five strong teeth;
clypeus large, not carinate, the anterior border rounded and furnished with two large
blunt teeth in front; frontal area small, rhomboidal; frontal carinae elevated, very

BY J. J. MCAREAVEY. 19

distinct and diverging behind; frontal groove distinct. Scape extends to the posterior
border of the head. Eyes moderately large, flat, are placed at the posterior third. Ocelli
lacking but there is a pit in place of anterior ocellus.

Thorax nearly twice as long as the broadest part which is the pronotum; pronotum
nearly twice as broad as long, rounded in front and with feebly convex sides. The pro-
mesonotal suture is black and very distinct. Meso-epinotal suture very faint. Meso-
epinotum is nearly twice as long as broad in front, narrowed so that it is about a
quarter as broad behind as in front. In profile the dorsum of the thorax is flatly convex
with the epinotal declivity slightly concave. Node very transverse, from above it
presents a very narrow surface. In profile nearly three times as high as long, with
slightly convex anterior face, which meets the vertical, straight posterior face in a sharp
point. Gaster oval. Legs moderately long and strong.

Worker minor.

Length, 5-1 mm. Colour much more brownish and there is considerable variation.
Often the posterior part of the head is black, and the dark area varies in extent. The
node and legs in some species are very dark brown.

Sculpture similar to that of major.

The hair is much more noticeable on the thorax, more abundant on the whole body
than in case of major.

Head as broad as long, slightly broader behind than in front, with the sides and the
posterior border almost straight. There are no noticeable teeth on the anterior border
of clypeus and the teeth of the mandibles are longer and sharper. The eyes are larger,
more convex, placed at the posterior corners of head. The scape is longer, extending
beyond the occiput by half its length. The thorax like that of major, but the pronotum
is relatively broader, the meso-epinotum longer and more tapering, appearing to end in
a point. The node is thicker in profile, about twice as high as long. Rest as in major.

Collected by F. Heugel.

Material examined: Twenty minor workers and a major worker.

This species is nearest to M. sponsorum Forel, but is larger and differs in colour.
The node of the major is much thinner, the clypeus is more rounded in front and the
eyes placed further back.

Type locality: Broome, N.W. Australia.

Type: Holotype worker major in collection at C.S.I.R., Canberra.

CAMPONOTUS (MYRMOGONIA) ARMSTRONGI Sp. nov. (Text-figs. 53-56.)
Worker major.

Length, 6-8 mm. Head and gaster shining black, with the antennae slightly lighter
and the articulation of the scape yellowish; thorax, node and legs, except the tarsi and
the apex of the tibiae, bright brownish red; apex of tibiae and the tarsi more brownish.

Head, thorax, node and gaster very finely and densely striate transversely, finer
on the gaster which is shining and almost smooth in parts. The striae are joined
together to form a wide meshed reticulation. There are also a few scattered punctures
on the front of the head.

Hair yellowish, long, sparse on the head and thorax, more abundant on clypeus,
node and gaster.

Head rectangular, one-third longer than broad, with the sides almost straight, the
occipital border straight and the posterior angles abrupt. Mandibles furnished with four
strong, sharp, teeth; clypeus large, feebly carinated, the anterior border rounded and
faintly crenulated in centre; frontal area feebly marked; frontal carinae raised, very
distinct, and diverging behind; frontal groove distinct, reaching to the end of the
frontal carinae. Scape reaches just beyond the occiput; first segment of funiculus longer
than second, all segments twice as long as broad; eyes moderately large, rather flat,
placed at the posterior third of head; no ocelli but there is a slight pit in place of the
anterior ocellus.

20 AUSTRALIAN FORMICIDAE,

Thorax nearly twice as long as the broadest part, which is at the pronotum;
pronotum one and a half times as broad as long, rounded in front, with feebly convex
sides. On the front of the pronotum is a sharp semicircular ridge or margin which
extends some way back along the sides. Pro-mesonotal suture is distinct; mesonotum
broader than long; meso-epinotal suture feebly marked. The meso-epinotum is nearly
twice as long as broad in front, compressed laterally behind so that the dorsum at the
epinotum is reduced to a sharp ridge. In profile the whole dorsum of the thorax forms
a single strong convexity, with the pronotal region slightly flattened. The epinotal
declivity is feebly concave with the stigmata near the base very distinct. Node twice
as broad as long, elliptical. In profile thorn-like, twice as high as long, the anterior
face convex, merging into the dorsum. The apex is sharp and the posterior face is
straight and almost vertical. Gaster small and ovate. Legs robust.

Worker minor.

Length, 455 mm. Colour, sculpture and pilosity as in major.

Head as broad as long, broader behind than in front with convex sides and occipital
border. The scape extends beyond the occiput by a third of its length. Hyes large,
convex, are placed at the posterior corners of the head. The thorax as in major but the
meso-epinotal suture is missing. Ali the rest as in the major worker.

Collected by J. W. T. Armstrong.

Type locality: Nyngan, N.S.W.

Material examined: A large series of major and minor workers which shows very
little variation.

The species is smaller than most Australian Myrmogonia and the vivid red thorax
and shining black head and gaster readily distinguish it from all other species.

Type: Holotype worker major in collection at C.S.I.R., Canberra.

TRIBE MELOPHORINI Forel 1912.
Genus MELOPHORUS Lubbock, 1883.

Melophorus “Lubbock, Journ. Linn. Soc. Lond., Zool., 17, p. 51, 188.
Melophorus sens. str. Wheeler, Psyche, 42, p. 70, 1935.

MELOPHORUS (MELOPHORUS) BRUNEA Sp. nov. (Text-figs. 57-65.)
Worker maxima.

Length, 4 mm. Dark brown with the thorax slightly lighter than the head and
gaster; legs, antennae, mandibles, clypeus and cheeks more yellow.

Mandibles smooth; head almost smooth, very faintly reticulate on the sides of the
clypeus, the reticulation tends to be transverse; thorax reticulate but slightly coarser
than on the head, and the epinotum is almost reticulate-punctate; gaster smooth and
shining with faint reticulations similar to those of the head.

Hair yellowish, long, erect on clypeus, shorter on the rest of the body. Pubescence
yellowish, short, adpressed, confined to the antennae and legs.

Head, excluding the mandibles, as long as broad with very feebly convex sides and
occipital border, the posterior corners rounded. Mandibles large, triangular, with four
strong teeth; clypeus large, convex, rounded and entire on anterior border; frontal area
triangular and distinct; frontal carinae short and parallel. Scape fails to reach the
occipital border by twice its width; first segment of the funiculus as long as the two
following, second shorter than the third, as broad as long, apical segment just slightly
longer than the second last; eyes large, convex, placed at the middle of the sides: ocelli
very small but distinct.

Thorax half as broad as the head, and nearly twice as long as broad; pronotum as
broad as long, very convex in all directions; mesonotum almost circular; pro-mesonotal
suture distinct. There is a rather broad interval between the mesonotum and epinotum
and the stigmata of the dorsum are very distinct. Epinotum is less than half as broad
as the mesonotum, almost square, with the sides and the posterior border straight. In

BY J. J. MCAREAVEY. 21

profile the pronotum and mesonotum form a single strong convexity. The dorsum of
the epinotum is straight, slightly elevated behind. The epinotal declivity is abrupt and
straight or feebly concave, twice as long as the dorsum.~- Node scale-like, from above
it appears as a narrow transverse line; in profile three times as high as long with
almost straight faces and pointed apex. Gaster large. Legs slender.

Worker media.

Length, 3:2 mm. Colour, sculpture and pilosity as in maxima.

Head slightly broader than long, with convex sides and occipital border. The scape
extends just beyond the occipital border. In other respects the head is similar to that
of maxima. The thorax differs merely in not having any noticeable interval between
the mesonotum and epinotum. Rest as in maxima.

Worker minima.

Length, 2.1mm. Black with faint metallic sheen; legs brownish black with the tarsi
yellowish; mandibles, front of head and scape brownish; funiculus yellowish with the
apical segments slightly darker. Whole body densely microscopically punctate. Hair
silvery grey, most abundant on the gaster.

Head broader than long, the sides and occipital border convex. Mandibles triangular
with sharp teeth of which the apical is by far the longest; clypeus large with the
anterior border almost straight; frontal area triangular, very clearly indicated. There
is a well-marked frontal groove. Frontal carinae are short and almost parallel. Scapes
extend beyond the occiput by half their length; eyes extremely large, occupying most of
the side of the head; ocelli small but distinct.

Thorax slightly more than twice as long as broad at the pronotum; pronotum three-
quarters as wide as the head, slightly broader than long, convex in all directions; pro-
mesonotal suture indicated; mesonotum one-quarter longer than broad, broader in front
than behind, with the sides almost straight; meso-epinotal suture deeper than the pro-
mesonotal suture; epinotum as long as the mesonotum, one and a half times as long as
broad, very slightly broader behind than in front. In profile the pro-mesonotum forms
a single moderate convexity. Dorsum of epinotum is almost straight and rounded into
the straight declivity, which is almost twice as long as the dorsum. Node is elliptical,
twice as broad as long, slightly broader behind than in front; in profile twice as high
as long at the base, anterior face feebly convex, rounded into the convex dorsum,
posterior face straight and vertical. Gaster small. Legs long and slender.

Female.

Length, 45 mm. Head dark brown with mandibles, front of head, antennae reddish
brown; thorax dark brown with a patch of orange which varies greatly in extent in
different examples—in most the posterior half of mesonotum and all the scutellum are
orange; node and gaster dark brown; legs brownish yellow but much lighter than the
antennae.

Sculpture as in maxima but smooth except on gaster where it is coarser than in
worker and the reticulation has a transverse direction.

Pilosity as in maxima.

Head similar to that of maxima but slightly broader than long. Mandibles large,
triangular, with four strong sharp teeth, the apical tooth very long, and the third tooth
longer than second and fourth; clypeus produced to a blunt point in front; frontal
groove very clearly marked; scape extends to the occiput; eyes large, convex, placed
slightly behind the middle of the sides; ocelli brownish, large and convex.

Thorax nearly as broad as the head, almost twice as long as broad; pronotum from
above reduced to a narrow convex strip; mesonotum large, as broad as long; parapsidal
furrows deeply marked; scutellum very large, broader than long, slightly broader in
front than behind; epinotum twice as broad as long, slightly broader in front than
behind, sides and posterior border straight. In profile the pronotum is vertical,
_mesonotum and scutellum almost flat with a marked indentation between the mesonotum

22 AUSTRALIAN FORMICIDAE,

and scutellum. The dorsum of epinotum is much lower than the scutellum, straight, one-
third as long as the declivity, the posterior angle rounded. Node transverse, about six
times as broad as long. In profile thorn-like, anterior face convex, posterior face concave
and apex sharp. Gaster large. Legs slender.

Male.

Length, 2:1 mm. Black, legs brownish black with the joints and the tarsi lighter,
mandibles reddish brown.

68 70

65

Text-figures 57-70.

57-65. Melophorus (Melophorus) brunea sp. nov. 57. Maxima worker, head. 58. Maxima
worker, lateral view. 59. Maxima worker, dorsal view. 60. Media worker, head. 61. Minima
worker, head. 62. Minima worker, lateral view. 63. Male, lateral view. 64. Female, head.
65. Female, dorsal view.

66, 67. Plagiolepis nynganensis sp. nov. 66. Worker, lateral view. 67. Worker, head.

68-70. Stigmacros elegans sp. nov. 68. Worker, lateral view. 69. Worker, head. 70. Worker,
dorsal view.

BY J. J. MCAREAVEY. 23

Head and thorax densely microscopically punctate; node and gaster smooth.

Hair greyish, long, sparse, more abundant on gaster. Pubescence greyish, fine,
adpressed on antennae and legs.

Head as broad as long, slightly broader behind than in front, with the sides and
occipital border convex. Mandibles large, at each corner of the terminal border is a
small tooth, the edge between these teeth edentate; clypeus large, convex, with anterior
border rounded; frontal area small, merely indicated; frontal groove distinct; frontal
carinae small and parallel; scape long, extending beyond the occipital border by half
its length; funiculus twelve segmented, the first segment as long as the two following,
the second shorter than the third, the rest twice as long as broad, the apical slightly
longer than the preceding; eyes very large, convex, occupying most of the side; ocelli
large and convex.

Pronotum twice as broad as long, anterior border and sides almost straight;
mesonotum large, as broad as long, broader in front than behind; epinotum at least
twice as broad as long, broader in front than behind, sides almost straight, the posterior
border feebly convex. In profile the dorsum is convex, slightly flattened at the meso-
notum. The dorsum of epinotum is much lower than the scutellum, and about a quarter
as long as the convex declivity. Node is transverse; in profile scale-like, anterior face
straight and vertical, meeting the posterior face, which is convex, in a sharp point.
Gaster small. Legs long and slender.

Collected by J. W. T. Armstrong.

Type locality: Nyngan, N.S.W.

Material examined: A large series of all forms. There is very considerable variation
in colour in the females, worker maxima and worker media. The worker minima does
not seem to vary. This species is distinct from all known species, but in many cases
not all the forms of the other species are described, so that it is difficult to make
comparisons in this genus. The worker minima of this species resembles slightly
Melophorus (Erimelophorus) scipio Forel, but the dorsum is much shorter than the
declivity, the scape is longer, and the eyes larger, but the whole ant is much smaller.

Types: Holotype worker minima, allotype male, and a series of worker maxima,
worker media and the female in the collection at C.S.I.R., Canberra.

Tribe PLAGIOLEPIDINI Forel 1893.
Genus PLAGIOLEPIS Mayr 1861.
PLAGIOLEPIS NYNGANENSIS sp. nov. (Figs. 66, 67.)
Worker.

Length, 1-5-2 mm. Dull light brown to dark brown, with mandibles, clypeus, antennae
and legs paler.

Mandibles and clypeus smooth; rest of head densely microscopically reticulate-
punctate with the occipital region more longitudinally striate; thorax and gaster densely
microscopically punctate, the punctures on the gaster rather elongated.

Hair yellowish, long, erect on clypeus, more adpressed and thicker on the gaster.
Pubescence yellowish, fine adpressed, abundant but not hiding the sculpture.

Head, including the mandibles, almost one and a half times as long as broad, as
broad in front as behind, with feebly convex sides and occipital border and rounded
corners; mandibles large, triangular, with five small teeth; clypeus large, convex, not
carinated, the anterior border broadly rounded and entire; frontal area very distinct,
triangular; frontal carinae sinuous, very slightly approximating behind; eyes moderately
large, rather flat, placed just behind the middle of the sides; no ocelli; scapes extend
beyond the occiput by a quarter of their length. The funiculus is slightly enlarged
towards the apex; the first segment as long as the two following together, second and
third subequal, and as broad as long, the rest nearly twice as long as broad, the apical
as long as the two preceding together.

Thorax short and rather stout, broadest through the pronotum; pronotum, including
the neck, broader than long, very convex in front and on the sides; mesonotum three-
quarters as broad as the pronotum, as long as broad, with convex sides; both thoracic

24 AUSTRALIAN FORMICIDAE,

sutures deeply marked, and at the meso-epinotal suture are prominent stigmata.
Epinotum is almost as broad as the mesonotum, very slightly broader behind than in
front, twice as broad as long, with feebly convex sides and feebly concave posterior
border. In profile the pronotum and mesonotum form a single strong convexity. The
epinotum is much lower with the dorsum very short, passing gradually into the declivity
which is about five times as long as the dorsum, and is straight and sloping. Node is
hardly noticeable from above because of the overhanging anterior border of the gaster.
In profile it is low, rather thin on top, strongly inclined forward, the anterior face
straight, the posterior face convex and the apex sharp. Gaster broadly elliptical,
voluminous, and overhanging the node. Legs long and rather robust.

Collected by J. W. T. Armstrong.

Type locality: Nyngan, N.S.W.

Material examined: Seven workers which show only slight variation in colour.

Resembling P. lucidula Wheeler and P. squamulosa Wheeler, but differing from these
Western Australian species in colour and sculpture. The head differs from P. squamulosa
in length, and it is not broader behind than in front as in P. lucidula. The mandibles
and funiculus are distinct. The thorax is more like P. squamulosa but the dense pilosity
is lacking.

Type: Holotype worker in collection at C.S.I.R., Canberra.

Genus sTIGgMAcROS Forel 1905.
STIGMACROS ELEGANS Sp. nov. (Text-figs. 68-70.)
Worker.

Length, 1-8-2 mm. Head and thorax shining black; node very dark brown on top but
light brown towards the base; gaster clear brown or amber; legs, mandibles and antennae
brownish yellow with the funiculus slightly darker.

Head shining, shagreened and almost punctate; thorax, node and gaster smooth and
shining except for some very scattered elongated scratches.

Pilosity hardly noticeable on any part of the body.

Head, excluding mandibles, slightly longer than broad, with the sides feebly convex
and the occipital border straight, the corners rounded; mandibles triangular with at
least four strong teeth; clypeus convex above, with anterior border broadly rounded;
frontal area indistinct; frontal carinae short, and almost parallel; eyes large, convex,
and placed at the middle of the sides; scapes extend very slightly beyond the occipital
border; first segment of the funiculus as long as the two following, second to eighth as
broad as long, ninth twice as long as broad, apical nearly three times as long as broad,
and longer than the two preceding together.

Thorax one and three-quarters as long as the broadest part, which is at the pronotum.
Pronotum is twice as broad as long, strongly convex in all directions; pro-mesonotal
suture deeply impressed; mesonotum slightly broader than long, broader in front than
behind, with the sides convex; meso-epinotal suture deep and wide; epinotum nearly
twice as broad as long, broader behind than in front, the anterior corners rounded, the
sides almost straight, and the posterior border concave. In profile the pronotum and
mesonotum are flat, rounded in front and truncated behind. The dorsum of epinotum
is flat and shorter than the slightly concave declivity, which is almost at right angles
to the dorsum. Very near the top of the declivity is a short broad tooth directed back-
wards. Node from above transverse, three times as broad as long, feebly notched on top.
In profile it is thorn-like, twice as high as long, the anterior face feebly convex, and
rounded into the short dorsum, while the posterior face is almost straight and vertical.
Gaster longer than broad. Legs short and robust.

Female.
Length, 2-2-8 mm. Colour as in worker but the gaster is much darker.
Sculpture as in worker but the thorax is more shagreened, especially on the epinotum.
Pilosity as in worker but there is a fine greyish pubescence on the funiculus.

BY J. J. MCAREAVEY. 25

Head as in worker, except that there are three very smali and indistinct ocelli.
Pronotum is almost three times as broad as long, with anterior border and sides almost
straight, the anterior corners abrupt, almost right-angled, though not sharp; mesonotum
broader than long, with distinct parapsidal furrows; scutellum large, broader than long,
slightly broader in front than behind. The veins and pterostigma of forewings brown,
and there is a single cubital cell and no discoidal cell. Rest as in worker.

Collected by J. W. T. Armstrong.

Type locality: Nyngan, N.S.W.

Material examined: Six females and twelve workers.

This species resembles S. fossulata Viehmeyer and 8S. medioreticulata Viehmeyer,
but differs from both in colour and sculpture. The scape is much shorter than in either
of these species and segments of the funiculus differ in length. The thorax is nearer
to that of S. medioreticulata but the pronotal angles are not sharp. There are no lateral
teeth on the node.

Type: Holotype worker in collection at C.S.I.R., Canberra.

26

THE DISTRIBUTION OF FORMIC AND ALCOHOL DEHYDROGENASES IN THE
HIGHER PLANTS, WITH PARTICULAR REFERENCE TO THEIR VARIATION IN
THE PEA PLANT DURING ITS LIFE CYCLE.

By DAPHNE C. Davison, M.Sc.
(From the Department of Biochemistry, University of Sydney.)

(Nine Text-figures. )

[Read 30th March, 1949.]

INTRODUCTION.

In the study of any enzyme it becomes necessary to obtain some idea of its distribu-
tion in nature. So it was that the present investigation arose from a study of the formic
dehydrogenase system of pea seeds. This dehydrogenase has only been reported in the
seeds of pea and the French or Kidney bean so that the problem to be attacked was
whether it occurred in other plants and if so was there any generic significance in its
distribution. As a companion study the distribution of alcohol dehydrogenase has also
been investigated.

Thunberg (1921, 1936a) first discovered a formic dehydrogenase in the seeds of
Phaseolus vulgaris (French bean). The same enzyme, obtained from peas by Fodor and
Frankenthal (1930), was found to differ from the formic dehydrogenase of bacteria in
that it required a coenzyme which was present in boiled pea juice and, to a lesser
extent, in boiled yeast juice. Andersson (1934) and Lichtenstein (1936) were able to
identify this factor as coenzyme I. In 1937 Adler and Sreenivasaya, again using peas,
studied the formic dehydrogenase system in detail. They drew attention to the strong
inhibition obtained with small amounts of cyanide, a property differentiating this
enzyme from all hitherto known cozymase-requiring dehydrogenases.

The alcohol dehydrogenase of plants is much better known than is formic dehydro-
genase and although alcohol does not seem to be readily oxidized in some plants
(Thomas and Fidler, 1941), the presence of an active alcohol dehydrogenase has been
reported from many sources. Thunberg (1921, 1929, 1986b) found the enzyme in the
following species: Phaseolus vulgaris (French bean), Citrus aurantium (orange),
Corchorus capsularis (jute), Cucumis sativus (cucumber), Echinocystic lobata (a
climbing herb), Hvonymus europaeus (spindle-tree), Mucuna utilus, Pistachia vera,
Pisum sativum (green pea) and Poinciana regia. Andersson (1933) mentions this
enzyme as one of a number of cozymase-requiring dehydrogenases found in cucumber
seeds. Adler and Sreenivasaya (1937) studied the alcohol dehydrogenase of pea seeds
and determined its pH optimum and equilibrium constant. Some later work by Berger
and Avery (1943) includes a study of alcohol dehydrogenase as it occurs in the Avena
coleoptile.

These references are by no means exhaustive but point to the relatively widespread
interest in alcohol dehydrogenase as a distinct contrast to the lack of attention to formic
dehydrogenase by plant biochemists.

MATERIAL.

(a) Seeds.—Most of the seeds used in these studies were obtained from the National
Herbarium, Sydney. Pea seeds (unless fresh peas are speeified) were commercial
household dried peas bought in 1 lb. packets. The dehydrogenase content was indepen-
dent of the brand, several local brands being used.

(b) Pea Plants—These were grown in a normal garden plot between November and
February. They were always used within two hours of gathering.

bo
co |

BY DAPHNE C. DAVISON.

METHODS.

Preparation of Extract—For the preparation of materials used in testing for formic
and alcohol dehydrogenases a standard procedure was adopted. Seeds were soaked in
distilled water overnight. Often it was necessary to soften those with hard testas
either by putting in boiling water or by mechanical cracking before soaking. The
material was frozen before extraction but otherwise the preparations were made at room
temperature. All plant materials were homogenized by mincing and/or grinding in a
mortar with sand depending on the procedure required to produce a fine state of
subdivision. Water was added in small quantities, the amount depending on the
material—usually twice or three times the dry weight of the seeds; generally none was
needed with other plant material, the cell sap providing the necessary consistency.

Homogenates were squeezed through muslin and the extract centrifuged and/or
filtered through kieselguhr. The filtrate, which was examined immediately, was usually
moderately clear and colourless so that the decolorization of the redox dye was easy to
follow.

Thunberg Technique.—In studying the distribution of formic and alcohol dehydro-
genases in plants the Thunberg technique was invariably used. The contents of the main
part of the tube were: 1 ml. extract, 0-6 ml. 0:2 M phosphate buffer pH 7:0, 0:2 ml.
coenzyme I solution (60ug.), 0-3 ml. substrate or water. In the hollow stopper was
0-3 ml. 0:0025 M thionine. These concentrations of reactants are based on the series of
experiments outlined below which aimed to find the optimum conditions. The tubes
were evacuated three times by means of a water aspirator and refilled with nitrogen gas
freed of oxygen by passing over heated copper filings. The tubes were then closed and
incubated at 38°-C. for 10 minutes before the dye was tipped into the reaction mixture.
The times taken for decolorization of the dye were noted and the time of the blank
containing water was compared with those taken for the tubes containing alcohol and
formate.

Buffer —The buffer used had a final concentration of M/20, a concentration which
seems to be most suited for plant work, from frequent mention in the literature. Adler
and Sreenivasaya (1937) state that the optimum pH for formic dehydrogenase lies
between 5:5 and 6:0. Using both 0-5 M acetate and 0-2 M phosphate buffers the formic
dehydrogenase system has been tested in this laboratory at all hydrogen ion concentra-
tions from pH 4:4 to pH 8:0. The pH of each reaction mixture was checked at the
conclusion of the experiment and found to agree very closely with the pH value of the
buffer used. A Leeds and Northrup portable glass electrode was used for these deter-
minations. The optimum pH was found to be pH 6-0-6-1 (Text-fig. 1). Below pH 5:3
no reduction of methylene blue could be obtained and precipitation generally occurred.
On the acid side a sharp drop from the optimum is seen while the activity decreases
more slowly towards the neutral point and in alkaline reaction up to about pH 8-2 it
remains constant. It is noteworthy in connection with the type of buffer used that
this anaerobic system was just as active in borate, phthalate and acetate buffers as in
phosphate buffers of the same pH values. In these experiments as well as those in the
following sections an ammonium sulphate precipitate was used in order to reduce the
blank and obtain more clear-cut optima. The peas are soaked overnight in distilled
water. The soaked peas are homogenized under nitrogen in a Waring Blendor for two
minutes with twice their dry weight of distilled water. The homogenate is squeezed
through muslin and the extract centrifuged for ten minutes. The supernatant is half-
saturated with ammonium sulphate and the precipitate taken up in distilled water, equal
to half the amount used for extraction, and dialysed overnight in the refrigerator against
a large volume of distilled water. The dialysate is centrifuged for five minutes and the
supernatant is the enzyme preparation referred to in Text-figures 1, 2, 3 and 4.

The optimum pH of alcohol dehydrogenase is, between 7:8 and 8-3 (Berger and
Avery, 1943). For these experiments then pH 7:0 was chosen as it lies mid-way between
the optima of the two dehydrogenases, and both enzymes work well at that reaction.

Substrate—The concentration of formate used (0:3 ml. 0-2 M sodium formate) was
optimum as decided from experiments with pea extract purified by half-saturation with

105

28 FORMIC AND ALCOHOL DEHYDROGENASES IN THE HIGHER PLANTS,

ammonium sulphate as above (Text-fig. 2). Further increase of the formate concentration
from the optimum of 0-025 M to 0:18 M has no effect.

The rather high final concentration of alcohol was chosen to allow for loss of
substrate during the evacuation and gassing of tubes.

4

Ww

(mins)

IO/DECOLORISATION TIME

(mins)

TIME

50

DECOLORISATION

LS)
wn

oO 0:06 O12 O18
2 FORMATE CONCENTRATION (Molar)

Text-figures 1, 2.
1.—The effect of pH on the decolorization of methylene blue by the formic dehydrogenase
system of peas. Thunberg contents: 0:5 ml. enzyme preparation, 0:2 ml. coenzyme I (60xzg.),
0-3 ml. 0-2 M sodium formate, 0:5 ml. buffer of various pH values, 0:2 ml. 0:025% methylene blue.
2.—Formate concentration curve. Thunberg contents: 0:5 ml. pea enzyme preparation,
0-5 ml. 0-2 M phosphate buffer pH 6:0, 0-2 ml. coenzyme I (60ug.), 0-2 ml. 0:025% methylene
blue. Total volume 2:3 ml.

Coenzyme I.—This substance was prepared by the method of Williamson and Green —
(1940), and although purity was only about 6:0% as determined by spectrophotometric
measurements of the absorption of the dihydroderivative at 340my, nevertheless the
activity of the preparation in the formate and alcohol systems was very high. The
amount used (60ug.) was optimal as decided from Text-figure 3 where the amounts of
coenzyme stated are the amounts actually present, 6:0% purity being allowed for.

BY DAPHNE CG. DAVISON. 29

The reversibly oxidizable Dye—Theoretically any reversible indicator whose
potential is more positive than that of the formate-bicarbonate and alcohol-acetaldehyde
systems would be reducible by these two dehydrogenase systems. From a consideration
of H’, values it is probable that more negative dyes would have been more suitable for
the study of these systems but the choice was limited to methylene blue (H’, + 0:01 volt)
or thionine (Lauth’s violet; HE’, + 0-068 volt). That thionine is the more efficient of the

two is seen from Text-figure 4. This is particularly the case with alcohol dehydrogenase.

TABLE I.

Distribution of Formic and Alcohol Dehydrogenases in the Seeds of Gymnosperms.

Decolorization Times.
Order and Name of Genus
Family. and Species. Common Name.
Blank. Formate. Alcohol,
GINKGOALES—
Ginkgoaceae Ginkgo biloba. 29% m 12% mn. ++ 25 m +
CONIFERAER—
Pinaceae Callitris calcarata. Black Murray Pine. 20 hrs 20 hrs. = 20 hrs. _
Pinaceae Callitris cupressi- | Victorian Cypress. 22 hrs 21 hrs. _ 21 hrs. —
formis.
TABLE 2.
Distribution of Formic and Alcohol Dehydrogenases in the Seeds of Monocotyledons.
Decolorization Times.
Order and Name of Genus
Family. and Species. Common Name.
Blank. Formate, Alcohol.
GLUMIFLORAE
Graminae Triticum aegilopoides. | Wheat. 14 m. 18 m. _ 4m. dk ale
LILIFLORAE—
Amaryllidaceae Doranthes Palmeri. Palmer’s Spear Lily. 12 hrs. 103 hrs. ? 63 hrs. +
Liliaceae . | Smilax glycyphylla. N.S.W. Sarsaparilla. 10 brs. 103 hrs. — 94 hrs. =
SCITAMINEAE—
Cannaceae Canna spp. Canna. >4 hrs. 130 m. + 103 m Jt te
TABLE 3. ;
Distribution of Formic and Alcohol Dehydrogenuses in the Seeds of Dicotyledons.
Decolorization Times.
Order and Name of Genus
Family. and Species. Common Name.
Blank. Formate. Alcohol.
URTICIFLORAE
Moraceae Ficus Bellingeri. Bellinger River Fig 62 m. 57 m, + 15am: ++
Tree:
Urticaceae Urtica dioica. Stinging Nettle. 50 m. 43 m. =f 45 m. +
PROTEALES—
Proteaceae Grevillea Hilliana. White Silky Oak. 53 mM. 103 m. +4 34 m. +4
Proteaceae Macadamia terni- | Queensland Nut- 88 m 23 m. ++ 4 m. ++
folia. tree.
Proteaceae Stenocarpus sinuatus. | Queensland Tulip | 120 m 120 m. _ 62 m. zis
Tree.
Proteaceae Telopea speciossima. | Waratah. 31 omis 103 m. ab ae 2m. ++
SANTALALES—
Santalaceae Fusanus acuminatus. | Quandong. 40 m. ef Waal, qe P4005 ++

LE

30 FORMIC AND ALCOHOL DEHYDROGENASES IN THE HIGHER PLANTS,

TABLE 3.— Continued.

Decolorization Times.
Order and Name of Genus
Family. and Species. Common Name.
Blank. Formate. Alcohol,
CENTRO-
SPERMAE—
Chenopodiaceae Atriplex spongiosum. | Spongy Salt Bush. 10 hrs. 105 m. + 5 m. ++
RANALES—
Lauraceae Cryptocarya obovata. | Obovate - leaved 16 hrs. 18 hrs. — 18 hrs _
Australian Nut-
meg Laurel.
Lauraceae Cryptocarya tripli- | Three-nerved Aus- 16 hrs. 18 hrs. _— 18 hrs. _
nervis. tralian Nutmeg
‘Laurel.
Ranunculaceae Clematis aristata. Australian Virgin’s 20 hrs. 20 hrs. = 20 hrs. -
Bower.
RHOEADALES—
Capparidaceae + | Capparis nobilis. Caper Orange Tree. 43 hrs. 65 m. + + 3 hrs. ++
Cruciferae Brassica oleracea | Cabbage. 16 hrs. 16 hrs. _ 16 hrs. —
(var. capitata).
B. oleracea (var. | Brussels sprouts. 16 hrs. 16 hrs. — 16 hrs. -
gemnifera).
B. oleracea (var. | Broccoli. 16 hrs. 16 hrs — 16 hrs. —
botrytis). 3
B. oleracea (var. | Cauliflower. 17 hrs. 17 hrs. _ 17 hrs. -
boyrytis).
B. rapa. Turnip. 60 m. 50 m. + 25 m ++
Rasphanus sativus. Radish. 55 m. 66 m. — 20 m =
PARIETALES— ‘
Passifloraceae .. | Passiflora edulis. Passionfruit. >150 m. 120 m. ar 19 m aR Se
CUCURBITALES
Cucurbitaceae Cucurbito pepo. Pumpkin. 29 m. 22 m. + 43 sec. +
Cucurbitaceae Cucurbita pepo. Squash. 23 m. 20 m. — 1 m. +4
10 sec.
Cucurbitaceae Cucurbito melo. Honeydew Melon. 29 m. 35 m. — 34 m. ++
Cucurbitaceae Cucurbita melo. Rock Melon. 15 m. 14 m. - 1m. ++
10 see.
MALVALES—
Elaeocarpaceae Elaeocarpus cyaneus. | Pink-flowered Olive- 5 hrs. 5 hrs _ 4? hrs. _
Berry Tree.
Sterculiaceae Brachychiton discolor. 24 m. 60 sec. ++ 30 sec. +4
GERANIALES or
RUTALES—
Rutaceae Boronia Barkeriana. | Barker’s Boronia. 8 hrs 17 hrs ? 19 hrs. —
Simarubaceae .. | Guilfoylia monostylis. | Yellow - flowered 10 hrs 9 hrs _ 60 m ++
Pyramid Tree.
Meliaceae Synoum glandulosum. | Spurious Rosewood. 170 m 170 m. _ 70 m —
SAPINDALES—
Akaniaceae Akania Hillii. Hill’s Horse-Radish | 225 m 240 m. _ 10 m +4
Tree.
Sapindaceae Alectryon subcinereus. 20 hrs 21 hrs. - 20 hrs —
Sapindaceae Cupariopsis anacar- | Carrot Wood. 20 hrs 19 hrs. ? 20 hrs. -
diodes.
Sapindaceae Harpullia pendula. Moreton Bay Tulip 12 hrs 12 hrs. — 12 hrs =
Wood.
Sapindaceae Dodenaea viscosa. Victorian Lignum 60 m 11 m. ++ 3m. ++
vitae.
CELASTRALES—
Celastraceae Celastrus australis. Australian Staff 10 hrs. 150 m. ++ 22 m. +4
Vine.
Celastraceae Elaeodendron australe.| Australian Scarlet- | 225 m. 180 m. + 19 m. ++
fruited Olive Plum.
ROSALES—
Pittosporaceae Pittosporum phyl- | Butter-bush. 19 hrs. 20 hrs. = 20 hrs. -
lyraeoides.
Pittosporaceae Pittosporum revo- | Yellow -_ flowered 6 hrs. 6 hrs. - 4 hrs. 2+
lutum. Brisbane Laurel.
Rosaceae Pyrus malus. Apple. UO) W006 20 m. aR ar 14 m. ar SF

BY DAPHNE C. DAVISON.

TABLE 3.—Continued.

Decolorization Times.
Order and Name of Genus i
Family. and Species. Common Name.
Blank. Formate. Alcohol.
Leguminosae .. | Acacia elata. Cedar Wattle. 74 m. 30 mn. + + 83 m a IF
Leguminosae .. | Acacia hakeoides. Black Wattle. 66 m. 35 m 4 17 m a St
Leguminosae .. | Acacia mollissima. 50 m. 43 m + 84 m +4
Leguminosae .. | Cassia corymbosa. 145 m. 90 m + 25 m ++
Leguminosae .. | Cassia didymobotrya. 55 m. 20 m. let 24 m ar ar
Leguminosae .. | Cassia eremophila. Desert Cassia. 70 m. 40 m. + 10 m ++
Leguminosae .. | Cassia laevigata. Golden Glory Bush. | 120 m. 45 m. a Sr 154 m ++
Leguminosae .. | Chorizema cordatum. | blame Pea Bush. 3 hrs. 70 m. + 40 m +4
Leguminosae .. | Caesalpinia Gillesci. 60 m. + m. t+ 5 m Stet
Leguminosae .. | Dillwynia brunioides. 6 hrs. 64 m. ++ 34 m. ++
Leguminosae .. | Erythrina crista-galli. 50 m. 14 m. + + 4m. +4
Leguminosae .. | Genista alba. 3 hrs. 70 m a 23 m aoe
Leguminosae .. | Hardenbergia coc- > 330 m. 57 m. SF A 50 m aP 45
cinea.
Leguminosae .. | Hardenbergia ovata- 300 m. 120 m + 110 m +
rosa.
Leguminosae .. | Kennedya rubicunda. | Red Bean Flower. > 300 m. 63 m ++ 33 m aP oP
Leguminosae .. | Phaseolus caracalla. > 300 m. 95 m. ae Se 23 m coe
Leguminosae .. | Phaseolus vulgaris. French Bean. 4 m. 1 m. ap ar 55 sec ++
20 sec.
Leguminosae .. | Pultenaeae flezilis. Flexible Pultenaea. 2) hrs. 80 m. = 17 m. +4
Leguminosae .. | Leucena glauca. >300 m. 190 m. 3b 22 m. oP ae
Leguminosae .. | Spartium junceum. Spanish Broom. 195 m. 38 m. +--+ 84 m. +4
Leguminosae .. | Swainsona coronilli- > 280 m. 190 m. _ 220 m. ar
folia.
Leguminosae .. | Viminaria denudata. | Australian Bush |>280 m. 220 m + ca, 280 m. 2B
Broom.
Leguminosae .. | Sesbania tripellii. Sours: 65 m. 4p SP 13 m. qF ar
Leguminosae .. | Vicia faba. Broad Bean. 3m. 2m. + 1 m. ataesty
20 sec.
Leguminosae .. | Pisum sativum. Pea. 5 m. iL in; +--+ 1m. ++
10 see. 15 sec
Leguminosae .. | Glycine soja. Soya bean. 160 m 135 m + 145 m +
Leguminosae .. | Trifolium repens. White Dutch Clover. | 180 m 65 m. aa 70 m +
Leguminosae .. | Trifolium subter- | Subterranean Clover.| 55 m 30 m. + 32 m +
raneanum. |
MYRTIFLORAE
Thymelacaceae Phaleria Neumanni. iZehrss 9 hrs + 84 hrs. se
Myrtaceae Tristania laurina. Turpentine Box. 16 hrs. 16 hrs. _ 13 hrs. +
UMBELLI-
FLORAE—
Araliaceae Tieghemopanax 16 hrs 1534 hrs _ 153 hrs. _
elegans.
Umbelliferae Daucus carota. Carrot. 54 hrs. 5} brs — 53 hrs. —
Umbelliferae Pastinaca sativa. Parsnip. 5 hrs. 5 his — 44 hrs —
Umbelliferae Petioselinum hor- | Parsley. 43% hrs. 4% hrs — 4? hrs. =
tense.
EBENALES—
Ebenaceae Diospyros pentamera. | Black Myrtle Plum. 49 m. 27 m + 37 m +
CONTORTA E—
Apocynaceae .. | Carissa orata. Black Lime Bush. 20 hrs. 16 hrs ae 15 hrs +
TUBIFLORAE—
Verbenaceae Vitex acuminata. 2% hrs. 2¢ hrs = 23 hrs. =
Solanaceae Solanum lycoper- | Tomato. 100 m. 47 m. ar 23 m. aa
sicum.
PLANTAGINALES
Rubiaceae Hodgkinsonia ovati- 6 brs. 5 hrs = 6 hrs. =
flora.
CAMPANULALES
Goodeniaceae .. | Goodenia paniculata. | Panicle - flowered |>16 hrs. 6 hrs. ae 110 m. SP ar
Goodenia.
Compositae Cichorium endivia. Endive. 44 hrs. 44 hrs. — 30 m. ++
Compositae Lactuca sativa. Lettuce. 43 hrs. 4 hrs, = 25>m: SP Ar

32

FORMIC AND ALCOHOL DEHYDROGENASES IN THE HIGHER PLANTS,

TABLE 4,
Distribution of Formic and Alcohol Dehydrogenases in Pisum sativum during its Life Cycle.
Decolorization Times.
Age of
Seedling. Part of Seedling.
(Days.) Blank. Formate. Alcohol.
0) Mature seed .. 5 m. 1 malo s: aR 52 4n oP Abe rao, 155s) aR 45 4p SF
1 Residual seed 54+ m. an ap ar aR ar 50 s SP SF Sr ar
Testa >60 m. >60 m. = >60 m. =
Embryo 7 m. 14 wm. SE ae ae lems OSS SP SR SF =F
Whole seedling 53 m. I Sos 3) = stiataeate 55 s. Spar SP Oo
2 Whole seedling 6 m. Teme 5s: qe Se ae Se 1m. 50s. |}+++ (+)
3 Residual seed 64 m. 1 am: ar i 10, By Sy qe SP ae
Testa >150 m. >150 m. = >150 m. _
Embryo 6 m. 24 m. aR AP ar 3 m. qr Sr
Whole seedling 64+ m. 24 m. aR an ar 44 m. ah oe
4 Whole seedling 64 mn. 3} nak, Siaalceaia 44 m. af ar
5 Residual seed Tite OS 2m. 40 s. ap Arar 3m. 10's. ar oar ar
Radicle 5 m. 2m +++ 5 m =
Plumule 7 m. 2? m. iaiataala 6m aF
6 Residual seed 8m. 10s 34 m. SP AP ar 4m SE Se 5
Radicle 5S inal; YO) Se 3m. ap oF 5} m =
Plumule 10 m. 6 m. qe ar 8 m. Se
Whole seedling 74 mM. 4m. ++ 7 m. +
7 Residual seed 94 m. 4m. 10s. Satan 5m. 10 s. ap 6
Radicle 8 m. 8 m. = 8 m. =
Plumule 20 m. 17 m. ae 15 m. =F
14 Residual seed 52 m. 39 m. SPF 50 m. =
Root 5 m. 115) 190%, = 15 m. =
Shoot .. 31m 23 m. +b 27 m. ar
21 Residual seed >5 hrs. >5 hrs. = >5 hrs. =
Root 27 m™. 25 m. ar 24 m. +
Shoot .. 39 m. Lay 1001, sr 30 m. +
Nodules 40 m. 40 m. = 35 m. ae
28 Root 30 m. Pah 100s ae 24 m. oF
Shoot .. 45 m. 40 mn. ate 36g ae +
Nodules 8 m. 6 m. SF 45s SF SE ar
35 Root 38 m. 35 m. ae 21 m. ap ar
Shoot .. 50 m. 47 m. ate 42 m. aia
Nodules 5 m. By rel, ap ar Tain, 10) Ss a atiaealcicig
63 Root 80 m. 50 m. SE Se 2 nie ar SP ae
Nodules 7 m. 4m aP AF 14 m. siietatalpteste
Leaves... 70 m. 59 m. al 49 m. Spar
Stems ee 52 m. 44 m. = 38 m. or aR
Whole shoot 65 m. 52 m. ai 45 m. =F ae
Flowers and buds BY Adie 18 m. aP ae 25 m. ale
63 Pods 1:5 em. long .. 30 m. 18 m. aisecte 18 m. SP St
Pods 2:0-3°5 em. 35 m. 16 m. Shar ae 13 m. SP SP oe
Pods 3:5-6:0 cm. 50 m. 12 m. SP Se APs 8 m. SS ae op ae
Very young seeds... 17 m. 10 m. te 43 m. qc ar ae
77 Seeds from full pod.. 2m. 5) 20. ++4+ 14 m. Se i=teista te
RESULTS.

ile

DISTRIBUTION OF FORMIC

PLANTS.

AND ALCOHOL DEHYDROGENASES IN THE SEEDS OF
THE HIGHER

The preparations tested and the results found are contained in the following tables.

The system of classification adopted is that of Engler (taken from Rendle, 1930).

In

the tables actual decolorization times are given but in order to simplify their study signs
are also used. A negative result is denoted by the sign —, a positive result, indicating
the presence of the enzyme, by +, and if the enzyme is very active (decolorization time
less than half that of the control) the sign ++ is used.

On examination of the data presented in Tabies 1-3 no positive evolutionary trends

are appar

ent.

However, one important fact emerges and that is that in the seeds tested,

BY DAPIINE GC. DAVISON. 33

tormic dehydrogenase is relatively widespread, since the orders chosen cover an extensive
range of Dicotyledons as well as some Gymnosperms and Monocotyledons.

It remained to be seen whether this widespread occurrence of the formic enzyme was
purely a property of the seed. It was, theretore, decided to follow the activity of formic
dehydrogenase, together with alcohol dehydrogenase, throughout the life cycle of the
pea (Pisum sativum).

60

(rmins)

TIME

30

DECOLORISATION

fo) 2 3
3 COENZYME | (mg)

ifefe)

SO

DECOLORISATION TIME (mins)

4 5 10 ff 15
REDOX DYE CONCENTRATION (Molar x 10-)
Text-figures 3, 4.

3.—Coenzyme concentration curves for the formic and alcohol dehydrogenases of peas.
Thunberg contents: 0:5 ml. enzyme preparation (ammonium sulphate precipitate as in Fig. 1),
0-5 ml. buffer pH 7:0, 0:3 ml. 0:2 M sodium formate or 0:3 ml. 3:6 M ethyl alcohol, 0-2 ml.
0-025% methylene blue. Total volume 2:3 mls. x———x, formate as substrate; O—-——O,
alcohol as substrate.

4.—Methylene blue and thionine compared as redox indicators in the formate and alcohol
dehydrogenase systems. Thunberg contents: 0:5 ml. pea enzyme preparation (as in Fig. 1),
0-5 ml. 0-2 M phosphate buffer, 0:2 ml. coenzyme I solution (60u4g.), 0:3 ml. 0:2 M formate or
6-3 ml. 3:6 M aleohol. The dye, in various concentrations, was placed in the hollow stopper as
usual. Total volume 2:0 ml. x —x, formate as substrate ; O————O, alcohol as substrate;
e— @, blank.

Dotted lines denote the decolorization when methylene blue is used; the entire lines denote
the decolorization when thionine is used.

34 FORMIC AND ALCOHOL DEHYDROGENASES IN THE HIGHER PLANTS,

2. VARIATIONS IN FORMIC DEHYDROGENASE AND ALCOHOL DEHYDROGENASE CONTENT
OF THE PEA PLANT DURING ITS LIFE CYCLE.

Seedlings up to seven days old can be grown quite readily in a dissecting dish on
damp filter paper and covered by a sheet of glass. Specimens older than this were
obtained from a normal garden plot. Correlation of the activity of the enzymes at the
various times and from different parts of the plant were made from dry weight deter-

Ke) 8

WHOLE SEEDLING
RESIDUAL SEED

fe)
2) 4 6
6
60
40
20
fe)
7
40
NODULES
20
3 O ©
a 42 63
8 9

Text-figures 5-9.
5-9.—Variations in formic and alcohol dehydrogenase during the life cycle of the pea.
x—-———-x, formate as substrate; O————O, alcohol as substrate; e————e blank. Decoloriza-
tion time in minutes is plotted along the ordinate, the age of the plant in days along the
abscissa,

cto
ot

BY DAPHNE C. DAVISON.

minations of the water extracts used. The results are seen in Table 4. The decoloriza-
tion times quoted have been corrected for a dry weight of extract of 0:03 gm. Signs
have again been used to facilitate inspection of results. + indicates a decolorization
time >75% of the blank but still positive for the dehydrogenase; ++, 50-75% of the
control decolorization time; +++, 25-50% of the control, and +++ <25% of the control
time.

In Table 4 and Text-figures 5, 6, 7, 8 and 9 the general trends in the activity of these
two dehydrogenases are apparent. Alcohol dehydrogenase decreases in activity as
germination proceeds until after six days there is very little present. When the plant
becomes older, however, there is a gradual increase of this enzyme, especially in the
nodules and later in the root, shoot and young seed. Formic dehydrogenase similarly
diminishes on germination although not as much as the alcohol enzyme. It does not
reach high activity again until the seed pods begin to develop.

Finally it was necessary to discover whether these enzymes are dormant in the
mature dried seed, or whether they only appear upon soaking and germination.
Comparison between the dehydrogenase activity of seeds soaked overnight and the flour
of seeds ground up in a coffee grinder before extracting revealed no difference.

GENERAL CONCLUSIONS.

Once again the widespread occurrence of formic dehydrogenase must be emphasized.
Of the ninety-three species of plant seed examined it is of interest to note that fifty-four
contained formic dehydrogenase and sixty-nine alcohol dehydrogenase.

The striking generic trend in this survey is that all of the 28 species of Leguminosae
seeds examined contain the formic enzyme, mostly in a very active form. Some work
has been carried out on the possibility of a special role for this enzyme in nitrogen
fixation in the Leguminosae and it was shown (a) that not only is formate not
' metabolized by Rhizobia but that it actually inhibits the small endogenous respiration
of the latter and (b) that the presence of the formic dehydrogenase does not depend on
the nodule bacteria, since it is present whether the sterile plants are inoculated or
grown without nodules.

Finally, it appears from results in Table 4 that the formic enzyme is most active in
the seed. This activity is dormant in the mature dried seed, diminishes in activity with
germination of the seed and growth up to the adult stages and increases in activity
again in the germinating seed. Activity of the alcohol enzyme follows a similar course
except that it decreases much more rapidly on germination and it-is greater in the
mature plant than that of formic dehydrogenase. The extremely active alcohol dehydro-
genase of nodular material is worthy of special note.

SUMMARY.

1. Ninety-three species of plant seed have been examined for the presence of formic
dehydrogenase and alcohol dehydrogenase. Of these, fifty-four contained the former
enzyme and sixty-nine the latter. —

2. Variation in the activity of both enzymes throughout the life cycle of the pea are
reported.

ACKNOWLEDGEMENTS.

This work was supported by a grant from the Commonwealth Research Committee. The
writer wishes to express her thanks to Mr. Constable of the National Herbarium, Sydney, for his
assistance in obtaining most of the seeds used.

References.

ADLER, E., and SREENIVASAYA, M., 1937.—Uber die Komponenten der Dehydrasesystema. XVI.
Zur Kenntnis der Formicodehydrase und der Alkoholdehydrase in Pflanzensamen. JZ. physiol.
Chem., 249; 24-39.

ANDERSSON, B., 1933.—Uber Co-Zymaseaktivierung einiger Dehydrogenasen. Ibid., 217: 186-190.

—, 1934.—Die Co-Zymase als Co-Enzym bei enzymatischen Dehydrierungen. Ibid., 225:
57-68.

BerGER, J.. and AveERy, G., 1943.—Dehydrogenases of the Avena coleoptile. Am. J. Bot., 30:
290-297.

36 FORMIC AND ALCOHOL DEHYDROGENASES IN THE HIGHER PLANTS,

Fopor, A., and FRANKENTHAL, L., 1930.—The mode of action of dehydrogenases. 1. The dehydro-
genase of the pea and its so-called coenzyme. Fermentforsch., 11: 469-489. (Seen in abstract
only.)

LICHTENSTEIN, N., 1936.—The effect of enzymic dephosphorylation of cozymase on its action in
the dehydrogenase system of peas. Ibid., 15: 44-48. (Seen in abstract only.)

“RENDLE, A. B., 1930.—The Classification of Flowering Plants. Cambridge University Press.

THomas, M., and Fipuerr, J. C., 1941.—Studies in zymasis. VIII. The discovery and investiga-
tion of aerobic HCN zymasis in apples treated with HCN and comparison with other forms
of zymasis. New Phytologist, 40: 217-239.

THUNBERG, T., 1921.—The presence of certain oxidising enzymes in the seeds of Phaseolus
vulgaris. Arch. intern. physiol., 18: 601-606. (Seen in abstract only.)

————, 1929.—Uber das Vorkommen einer Hexose-diphosphorséure-dehydrogenase in Pflanzen-
samen. Acta Univ. Lund., 2 Avd., 25; 1-16.

——-.——, 1936a.—Uber das Vorkommen einer kraftigen Ameisensdure—Dehydrogenase in Samen
von Phaséolus-arten. Skand. Arch. Physiol., 74: 16-26.

————, 1936b.—Zur Kenntnis der Alkoholdehydrogenasen verschiedener Samen-arten. Ibid.,
75: 38-48. :

WILLIAMSON, S., and GREEN, D. E., 1940.—The preparation of coenzyme I from yeast.. J. Biol.
Chem., 135: 345-346.

THE IMPORTANCE OF FORMIC DEHYDROGENASE IN THE OXIDATION
MECHANISMS OF PISUM SATIVUM.

By DapPHNeE C. Davison, M.Sc.
(From the Department of Biochemistry, University of Sydney.)

(Two Text-figures. )

[Read 30th March, 1949.]

INTRODUCTION.

The widespread occurrence of the enzyme formic dehydrogenase in plant seeds has
been reported in a previous paper (Davison, 1949). Any attempt to understand the
position of this enzyme in the metabolism of the plant must include a study of the
respiratory mechanisms to be found. The oxidative chain reactions already postulated
for plants are summarized in Scheme I.

As ean be seen from this scheme, there are three main terminal oxidases and one
peroxidase and at least six paths by which biological oxidation could proceed. Attempts
have been made to find out which of these paths can be used in the breakdown of formic
acid to carbon dioxide and water in the pea seed. In order to make the results more
complete and perhaps discover some significant changes in activity, the enzymes of the
seven-day seedling were also studied.

METABOLITE

DEHYEROGENASB#B

es

PYRIDINE NUCLEOTIDE
- 285
- 2H
Fel ANV On PER OnTeko i Nid DEHYDROASCORBIC ACID \-24
ASCORBIC
24H ACID
(7) SUCCINATE- lM : @
OXIDASE
PUMARATE
+92
CYCLE ASCORBIC ACID__=28 »O0-QUINONE
-2 H Seog H +2 8 CATECHOL
CaY=T, 0 CoH ROM ES
- OXIDASE
QUINON POLYPHENOL
~ 2H PEROXIDASE
CYTOCHROME TH, 0%
OxIDASE PHENOL
OXYGEN

SCHEME I

MATERIALS AND METHODS.

The peas used were dried peas bought in 1 lb. packets and soaked overnight before
use when they took up their own weight of water. Various methods were used for the
preparation of the extracts depending on the nature of the enzymic relationships and
the conditions necessary for their demonstration. These methods are described in their
appropriate places. The seven-day-old seedlings were grown on damp filter paper in an

38 FORMIC DEHYDROGENASE IN THE OXIDATION MECHANISMS OF PISUM SATIVUM,

enamel tray under a glass cover. The whole seedling was used in the preparation of
extracts which were made in the same way as the corresponding seed extract.

The Thunberg technique was followed in anaerobic experiments on dehydrogenases
and a variation of this technique in work on ascorbic acid-formic dehydrogenase coupling.
Oxygen uptake was measured in Warburg manometers fitted with flasks carrying two
side arms; the volumes were between 20 and 25 ml. The manometers were shaken at
102 cycles per minute in a constant temperature water bath at 38°C. Usually the final
volume of each flask was 3:2 ml. To absorb CO, produced, 0:2 ml. 20% NaOH was placed
in the centre well, together with a KOH-paper made of Whatman No. 40 filter paper as
suggested by Dixon (1943). .

Coenzyme I was prepared by the method of Williamson and Green (1940) from yeast,
and its dihydro derivative estimated spectrophotometrically by absorption at 340mu.
Sodium pyruvate was prepared by the procedure of Robertson (1942) and estimated
manometrically by the production of CO, by enzymatic decarboxylation (Westerkamp,
1933). Other chemicals were of reagent grade purity and were neutralized or brought
to the appropriate pH before use. Buffers and the pH of flask contents were checked
frequently by means of a Leeds and Northrup portable glass electrode.

RESULTS.
DEHYDROGENASES AND CARRIER-LINKED DEHYDROGENASE REACTIONS.

In the first instance formic dehydrogenase must be regarded simply as one of the
many dehydrogenases to be found in the pea. In this connection the effect of a large
number of substrates has been tested anaerobically in Thunberg tubes and aerobically
in manometers.

In anaerobic studies peas were soaked overnight, homogenized for two minutes under
nitrogen in a Waring Blendor with an equal volume of water, squeezed through muslin
and the extract centrifuged for 5-10 minutes to remove starch. The green cloudy super-
natant fluid was 80% saturated with solid (NH,).SO,, the sediment taken up in water,
the solution centrifuged for 15 minutes and the supernatant used. Substrates which
were found to reduce the decolorization time include formate, alcohol, malate, fumarate,
glutamate and lactate (Table 1). As a rule succinate could only be included in this list
when an untreated, lightly centrifugd water extract was used. This is not surprising
since succinic dehydrogenase is not soluble in distilled water. Substances which either
made no difference to the decolorization time or else inhibited the reduction of the dye
were glucose, citrate and glycerophosphate.

TABLE 1.
Dehydrogenases Present in Pea Seeds and Seven-day Old Seedlings.
Tube contents : 1-0 ml. extract, 0-5 ml. 0-2 M phosphate buffer pH 6-4,
0-2 ml. coenzyme I (60 wg.), 0-3 ml. substrate (final concentration
0-02 M), 0-2 ml. 0-0025 M thionine.

Decolorization Time. (Minutes.)
Substrate.

Seed Extract. Seedling Extract.
Water +3 ae A 65 51
Formate te ee 7 16
Alcohol Bo as Si 4 45
Malate te a Ae 16 14
Fumarate .. re a 18 15
Glutamate .. ee Aas 30 32
Lactate BA Ne ae 40 42
Succinate oe ae as 34 35
Glucose oS xe ab 65 51
Citrate ats a 62 50

8-Glycerophosphate sis 67 ; 51

BY DAPHNE C. DAVISON. 39

In the seven-day seedlings there were only two marked changes in the dehydrogenase
content as measured by the Thunberg technique and correlated on the basis of dry weight.
The activity of both alcohol and formic dehydrogenases had fallen off, the former
considerably (Table 1).

In both preparations it was interesting to note that, when malate and fumarate were
substrates, the time of incubation before the dye was tipped was an important factor in
comparing the decolorization times. The longer the period of incubation (generally
15-20 minutes were enough) the more closely the times agreed. This was due no doubt
to the fact that the enzyme fumarase has first to add water to the fumarate to form
malate before it can be dehydrogenated by malic dehydrogenase.

Of the dehydrogenase found, formic, malic, alcohol, glutamic and lactie all required
the presence of coenzyme I before reduction of thionine would take place. Possibly the
negative result with citrate was due to lack of coenzyme II (necessary for the working
of isocitric dehydrogenase). However, citrate failed to reduce the decolorization time
even in the untreated water extract which, presumably, contains some coenzyme II
(van Herk, 1935).

For the aerobic experiments a water extract of peas was used after centrifuging
for 10 minutes to remove starch and cellular debris. Methylene blue was used as
hydrogen carrier in these manometric experiments and a pH of 6-4 was chosen since
that is the natural reaction of pea juice and of water extract of peas. The results of a
typical experiment using seed extract are seen in Table 2.

TABLE 2,
Effect of Added Substrates on Oxygen Uptake of Pea Seed Extract.
Manometer contents: 1:6 ml. extract, 0-8 ml. 0:2 M phosphate buffer pH 6-4, 0-2 ml, coenzyme I
(602g.), 0-3 ml. 0-2 M substrate, 0-1 ml. 0-5% methylene blue, 0-2 ml. 20% NaOH in centre

well.
Substances which have No
Substances which Increase - wl. O» in Effect or which Decrease ul. O, in
the Endogenous O, 60 Mins. the Endogenous O, 60 Mins.
Consumption. Consumption.
Water .. aS ae Na 85 Lactate an Re ue 79
Formate Ee ds 25 164 Oxalate tay ue ad 83
Alcohol ae #5 a 156 Glycerophosphate ay: 85
Malate is ae te 118 Formaldehyde ug, ae 75
Glucose ce ane Ae 127 Acetaldehyde ee
Fumarate on wis a 120 Acetate Ne ae re 85
Succinate a ae at 97 Pyruvate .. LS Ag 59
Glutamate .. a ae 138 Citrate ee ee ue 77
Glycine a: Hs ae 122 Sucrose bos ae xe 83
Tyrosine ne BS ay 80
Catechol ue os ue 69
m-Cresol He cae ee 77
Phenol

In aerobic experiments using seven-day seedlings the O, uptake of the blank
endogenous respiration was higher but the effect of the various substrates was much
the same as for seeds, except that, in the seedling, sucrose, lactate and citrate can be
included in the list denoting increases in oxygen uptake (Table 3).

Carrier-linked Reactions.—Eixperiments were next made to determine whether the
formic dehydrogenase could be linked with any of the other dehydrogenase systems found
in the pea seeds. Of these three were immediately eliminated because of the fate of their
reaction product. Both oxalacetate and pyruvate, the substances resulting from
dehydrogenation of malate and lactate, are immediately decarboxylated by pea extract.
In the case of glutamic dehydrogenase, the reversibility of the reaction, glutamic acid +
O——> a-keto-glutaric acid + NH,, is difficult. There remained the possibility of coupling

EF

40 FORMIC DEHYDROGENASE IN THE OXIDATION MECHANISMS OF PISUM SATIVUM,

TABLE 3.
Effect of Added Substrate on Oxygen Uptake of Seven-day Seedling Extract.
Flask contents as in Table 2.

Substances which Increase Substances which Decrease
O, Uptake. ul. O2/hr. O, Uptake. ul. O./hr.

Water .. es ap a 183 Oxalate mn ne ‘aA 180
Formate xe Be “4 239 2-Glycerophosphate om 183
Alcohol soe Ae ae 200 Formaldehyde e2 es 178
Malate a Ha ae 197 Acetaldehyde Bs a 185
Glucose ae at Se 199 Acetate aS LG oe 180
Fumarate a, ae ee 202 Tyrosine ae ee A 177
Succinate as me oe, 195 Pyruvate oh ah 3h 169
Glutamate a6 5G aie 214 Catechol Ses Be aA 179
Glycine ws ie os 212 Phenol ah Ls a 175
Sucrose ne Ae ne 190

Lactate ae a a 194

Citrate ae ae ae 192

with either succinic dehydrogenase and fumarate or alcohol dehydrogenase and acetalde-
hyde. Succinic dehydrogenase will dehydrogenate succinic acid to fumaric acid only if
an acceptor of hydrogen is present. If, however, activated hydrogen is supplied, for
example in the form of dihydrocozymase from formate dehydrogenation, and there is no
acceptor like methylene blue present, then one would expect the reaction to go in the
other direction. In the case of alcohol dehydrogenase in the absence of methylene blue
or other substances which oxidize the reduced coenzyme I, the equilibrium position of
the reaction, C.H;OH + coenzyme I — CH,CHO + reduced coenzyme, is far to the left
(Euler, Adler and Hellestrom, 1936).

For these coupling experiments an untreated water extract of pea seeds was used
after light centrifuging (5-7 minutes). The supernatant therefore contained formic,
alcohol and succinic dehydrogenases. Briefly, the principle of the method used was that
if one employs a bicarbonate buffer in an atmosphere of CO., CO. released by the enzymic
reactions taking place will escape into the atmosphere and can be measured mano-
metrically. If a coupling of dehydrogenases does occur this will be indicated by increased
formate oxidation, and hence increased CO, output, in the presence of either acetaldehyde
or fumarate. A wide range of pH values were used in these experiments. A typical
result at pH 6-4 is presented in Table 4.

It can be seen from this experiment that here there are two carrier-linked reactions
in each case, the carrier being reduced by the negative system, formate plus formic
dehydrogenase and reoxidized by the positive system, fumarate plus succinic dehydro-

TABLE 4,
Carrier-linked Dehydrogenases.

Flask contents: 1:9 ml. supernatant, 0-3 ml. 0-023 M NaHCO;, 0-2 ml. coenzyme I
(60 ug.), 0-3 ml. 0-2 M sodium formate or 0-3 ml. water in one side bulb. In the
other side arm was 0-3 ml. 0:05 M acetaldehyde, 0-3 ml. 0:2 M fumarate or 0-3 ml.
water. The gas phase was 5% carbon dioxide in nitrogen.

ul. CO, produced in $0 minutes.

CO, Due to
Water. Formate. Formate Oxidation.
Blank 90 102 12
Fumarate 115 155 40
Acetaldehyde 95 150 55

Note.—Acetaldehyde is extremely volatile and cn all occasions boiled extract controls

were done. These figures are corrected for the boiled extract.

BY DAPHNE C. DAVISON. 41

genase on the one hand, acetaldehyde plus alcohol dehydrogenase on the other. Coenzyme
I is the only carrier needed in the case of the alcohol enzyme. With the succinic dehydro-
genase flavoprotein may, and probably does, also function as an intermediate.

These coupled reactions are only two of the many possible, e.g., between the lactic
and malic enzymes as negative systems and the above two, alcohol and succinic dehydro-
genases, as the positive systems. Such coupled reactions must play a big role in the
economy of the pea seed and the fact that formic dehydrogenase can take part in them
increases the importance of this enzyme as a factor in the first step in biological
oxidation in the seed.

INTERMEDIATE HYDROGEN CARRIERS.
FLAVOPROTEIN AND THE DICARBOXYLIC AND TRICARBOXYLIC ACID CYCLES.

The next step in the study of formic dehydrogenase as related to oxidation
mechanisms was to examine pea extract for the presence of any intermediates or
terminal oxidases of Scheme I, and then to establish the relation of formic dehydrogenase
to any of the hydrogen carrier systems thus found. The enzyme preparation used for
these studies was generally a fine homogenate from which starch and cellular debris
had been removed by light centrifuging. Such a preparation contains much insoluble
material and particulate matter, including insoluble enzymes plus soluble enzymes and
a solution of the freely soluble diffusible components such as inorganic ions, coenzymes
and carriers. In addition the energy reservoir is maintained and the study of reaction
mechanisms which involve a number of co-ordinated enzymes is possible.

Flavoprotein.—A crude flavoprotein preparation was made from 12-day pea seedlings
by the method of Lockhart (1939), but this preparation contained too much formic
dehydrogenase to be used in experiments coupling formate oxidation with flavoprotein.
Attempts to purify it further by adsorption on C y alumina only led to its destruction,
because of the unstable nature of plant diaphorase (Lockhart, 1939). Soluble diaphorase
was therefore prepared from pig’s heart (Straub, 1939) and estimated by means of its
absorption at 451myu. The results of experiments with water extracts of pea seeds and
seedlings using this diaphorase preparation are seen in Table 5.

TABLE 5.
Effect of Flavopretein on Formate Oxidation.
Flask contents: 1 2 ml. enzyme, 0:8 ml. 0:2M phosphate buffer pH 6-4, 0-2 ml.
coenzyme I (60 g.), formate (0-3 ml. 0-2 M solution) and 0-5 ml. flavoprotein (5 Ug.
bound lactoflavin) as indicated, 0-2 ml. 20% NaOH in centre well, water to 3-2 ml.

|
ul. O, Uptake in 60 Minutes.
Increase.
Alone. Formate.
Seed Extract—
Blanks ee a or 48 58 21%
Flavoprotein A 52 75 44%
Seedling Extract—
Blank .. as See 110 109 —
Flavoprotein .. as 1a) 182 57%
|

Although flavoprotein does not markedly influence the blank O, consumption, it has
a stimulating effect on formate oxidation especially in the seedling. It can be concluded
then that, in the seedling particularly, the oxidation of any formate present can go
through this carrier.

Dicarbozylic and Tricarboxylic Acid Cycles.—Malic and citric acids are of common
and perhaps universal occurrence in plant tissues. Succinic, fumariec (Pucher and
Vickery, 1942) and isocitric acids (Krebs and Eggleston, 1944) have also been found.
The critical biochemical evidence for the occurrence of oxalacetic acid in plants rests

42 FORMIC DEHYDROGENASE IN THE OXIDATION MECHANISMS OF PISUM SATIVUM,

ultimately on Virtanen and Laine’s (1939) isolation of its oxime from legumes and
their nodules and the conversion of the oxime (oximinosuccinic acid) to aspartic acid
by reduction. However, no evidence has yet been obtained that these acids take part
in a Krebs or Szent-Gyorgyi cycle.

In peas, aS mentioned above, malate, fumarate and succinate increase the oxygen
uptake of pea extract but, so far from the acids acting catalytically, the extra oxygen
consumption was equivalent to only a fraction of the acid added. Citrate was found
to have no effect on seed extract. The effect of these acids on formate oxidation by pea
extract was measured manometrically at pH 6-4. The results are shown in Table 6,
from which it appears that citrate increases formate oxidation more than the other
added substrates. Malate and fumarate also have very marked effects but succinate
has no effect on the oxidation of formate as revealed by the oxygen consumption.

TABLE 6.
Effect of Acids on Formate Oxidation in Seeds.
Flask contents: 1-4 ml. water extract, 0-8 ml. 0-2 M phosphate buffer pH 6-4, 0-2 ml.
coenzyme | (60 Ug.). Various additions of C, acids, citrate, formate and water were
in the side arms, 0-2 ml. 20% NaOH in the centre well.

ul. O. Uptake in 60 Minutes.
Py ‘ Change.
No
Formate. Formate.
Blank a At ot 53 67 26% increase
Malate .. x oe 49 73 59% rh
Fumarate oe sa 47 72 52% »”
Succinate ae Ae 60 57 4% decrease
Citrate .. Se Sie 36 61 . 70% increase

3 6 ~9 9
' CYTOCHROME C (Molar x 10”)
Text-fig. 1.—The cytochrome saturation value for the formic dehydrogenase system of pH

7-0 buffer extract of pea seeds. Manometer flask contents as for Table 9. The cytochrome is
stated as final concentration. x...... x, formate as substrate; @...... @®, water (blank).

BY DAPHNE C. DAVISON. 43

In the seven-day seedling the effect on formate oxidation is not so great, possibly
because of the much lower activity of the formic dehydrogenase in seedlings (Table 7).
In the pea seed and seedling, then, although there is no evidence that the C, acids
and citrate act catalytically there is some indication that they hasten formate oxidation.

TABLE 7.
Effect of Acids on Formate Oxidation in Seedlings.

Flask contents as for Table 6.

wl. O, Uptake in 60 Minutes.
Change.
No
Formate. Formate.
Blank ie es ae 147 153 4% increase
Malate .. Me Ras 148 159 8% a
Fumarate oe oie 149 161 8% es
Succinate ss ea Ne 2 150 142 5% decrease
Citrate .. te a 131 142 9% increase
|

TERMINAL OXIDASES: CATECHOL OXIDASE, CYTOCHROME OXIDASE, ASCORBIC ACID
OXIDASE AND PEROXIDASE.

Catechol Oxidase.—Although this enzyme has been found in the nodules of peas
(Keilin & Smith, 1947), no evidence could be obtained in the present investigations that
it occurred in the seeds. No colour was obtained with catechol or a-naphthol, and none
of the substrates tested (Table 8) increased the oxygen uptake of an untreated buffer
extract. The extract was prepared by homogenizing 25 gm. peas (dry weight) in a
Waring Blendor under nitrogen in the usual way, 50 ml. 0:07 M phosphate buffer at
pH 7:0 being used instead of water. The supernatant was used after centrifuging for
seven minutes.

TABLE 8.
Catechol Oxidase in Pea Seeds.

Flask contents: 2-0 ml. supernatant, 0-3 ml. 0-5% gelatin, 0-3 ml.
substrate (2 mg.), 0-2 ml. 20% NaOH in the centre cup.

ul. O, Uptake

Substrate. in 45 Minutes.
Blank .. Sh Si is on Be eS 59
Catechol A ae ae ie A Be 49
Phenol a ao oa oe ar ae 53
m-Cresol af ae ate ae oe ans 55
Tyrosine ae Pe ae a wis Sis 57

Finally, injured seed tissue does not discolour on standing as is usual in plants
containing phenolases (and as was found in making extracts of nodules).

Similar experiments were made with seven-day-old pea seedlings again with negative
results so that it can be concluded that there is no catechol oxidase present, a decision
which eliminates two of the paths in Scheme I as applied to formate oxidation in peas.

Cytochrome System.—Hill and Bhagvat (1939) have found the four characteristic
absorption bands of cytochromes, a, b, and c in the embryos of legumes as well as other
plant tissues. Shaking with air caused the bands to disappear, succinate caused reap-
pearance, and reoxidation was prevented by the presence of cyanide and azide. Cyto-
chrome c from ox heart was rapidly reduced by the ground-up plant tissue poisoned with
cyanide.

44 FORMIC DEHYDROGENASE IN THE OXIDATION MECHANISMS OF PISUM SATIVUM,

In studying the effect of cytochrome c on formate oxidation the extract was made
in the same way as that used for catechol oxidase experiments, i.e., a pH 7-0 buffer
extract. The cytochrome c was prepared from ox hearts by the method of Keilin and
Hartree (1937) and estimated spectrophotometrically by absorption of reduced and
oxidized cytochrome c at 550myu. The results of experiments on pea seeds using this
preparation are seen in Table 9. The amount of cytochrome used is taken from Text-
figure 1 and is well over the saturation concentration for seed extract.

TABLE 9.
Effect of Cytochrome ce on Formate Oxidation in Seeds.
Flask contents: 1-5 ml. extract, 0-2 ml. coenzyme I (60ug.); 0-2 ml.
20% NaOH in centre well; in the side bulbs, 0-3 ml. 0-2 M formate,
0-3 ml. water or 0-5 ml. 2-7x10-7 M cytochrome as indicated
below. Total volume 3-2 ml.

ul. O. Uptake
in 50 Minutes.

Blank aes bys Me Be ae a 68
Formate Ag ons ae a, ae ae 70
Cytochrome ¢ ae Se sts oe oe 89
Formate and cytochrome ¢ AG ne 0 165

Results of this type show that cytochrome ec accelerates the endogenous O. uptake of
pea extract and that in the presence of formate this increase is much greater, indicating
that the cytochrome system is being used for formate oxidation.

Seedlings were prepared in the same way, i.e, a pH 7-0 buffer extract, and the
results are presented in Table 10.

TABLE 10.
Effect of Cytochrome ec on Formate Oxidation in Seedlings.

Flask contents as for Table 9.

ul. O. Uptake
in 50 Minutes.

Blank .. Be dys ae ee en at 210
Formate Se o0 50 ue SEER er 202
Cytochrome ¢ Mr ni ay BG ee 216
Formate and cytochrome ¢ to Le ae 234

These results show that cytochrome c has far less effect on the O. uptake of seedling
extract than on that of seed extract.

Ascorbic Acid Oxidase——Although peas before germination do not contain any
ascorbic acid, this substance begins to appear immediately upon soaking the dried seed.
The following table, taken from Harris and Ray (1933), gives the data on the synthesis
of ascorbic acid during germination.

The oxidation of ascorbic acid to dehydroascorbic acid is reversible and Szent-
Gyorgyi’s suggestion (1930) that it might behave as a redox body in plant respiration
has more recently been substantiated by the discovery of a mechanism for its reduction.
James and Cragg (1943) have shown that barley saps catalyse the oxidation of lactic to
pyruvic acid by means of the ascorbic system. Later James, Heard and James (1944)
established a similar link between the ascorbic acid system and an important inter-
mediate of glycolysis, hexosediphosphate, this time with the hydrogen transfer taking
place between coenzyme I and ascorbic acid. It was desirable to find out whether
coupling of this latter type could be demonstrated in pea seeds.

BY DAPHNE C, DAVISON. 4},

TABLE 11.
Synthesis of Ascorbic Acid During Germination.

Ascorbic Acid Content.
Material: Seed Feas. aL
u.g./Seed or
mg./gm. Seedling.
Before germination a 2 ae 0-00 0-00
Soaked 24 hours .. aye eh st 0:08 0-02
48 hours (germinated) os are ae 0-69 0-21
72 hours (germinated) ae ae ei 0-82 0-26
96 hours (germinated) ae ae & 0-86 0-27

Preliminary experiments designed to show the presence of ascorbic acid oxidase
in pea seeds were done on an ammonium sulphate precipitate (Text-fig. 2). The Waring
Blendor could not be used in this preparation because the oxidase was inactivated in the
process. Peas were ground up in a mortar with sand and twice their dry weight of
water. The enzyme was precipitated from the centrifuged extract by 83% saturation
with solid ammonium sulphate and the sediment was taken up in one volume of water,
dialysed overnight against distilled water and filtered through kieselguhr. This method
precluded the occurrence of any cytochrome oxidase in the preparation so that any
enzymic oxidation of ascorbic acid observed was due to ascorbic acid oxidase and not
the cytochrome—cytochrome oxidase system (see Stotz, Harrer, Schultze and King,
1937). Special precautions were taken to ensure the purity and freedom from copper
of all glassware and chemicals used in these experiments. Triple glass distilled water
was always used.

120

Bl O,

40

60 120 180
2 TIME (min)

Text-fig. 2.—The presence of ascorbic acid oxidase in pea seeds. Manometer flask contents:
1:2 ml. enzyme preparation, 1:5 ml. 0:07 M phosphate buffer pH 6-0, 0-3 ml. water or 0:3 ml.
ascorbic acid solution (0-01 mM=112 ul. O,), 0-2 ml. 20% NaOH in centre cup.

Curve 1: ascorbic acid. Curve 2: water. Curve 3: ascorbic acid plus boiled enzyme.

Identical experiments done on the pea seedling revealed a more active ascorbic acid
oxidase than in the seed. In the latter 0-01 in M ascorbic acid was oxidized in 130
minutes. In a seedling preparation, corrected for the same dry weight, the time taken
was 90 minutes. This increase in activity can be correlated with the increase in
substrate concentration in the seedling. As quoted from Harris and Ray (1933), there

46 FORMIC DEHYDROGENASE IN THE OXIDATION MECHANISMS OF PISUM SATIVUM,

is 0:02 mg. ascorbic acid per seed as used after soaking for 24 hours but 0-27 mg/seedling
after 96 hours’ germination.

From these preliminary experiments the presence of a specific ascorbic acid oxidase
in seeds and seven-day seedlings was confirmed. Further experiments, designed to
demonstrate the coupling of formic dehydrogenase and this ascorbic acid system, were
of two types, one aerobic the other anaerobic. In both cases the pea seeds or seedlings
were ground up in a mortar with twice their dry weight of Cu-free M/10 phosphate buffer
at pH 6:0. The extract was centrifuged for 15 minutes and the supernatant used.

Although formate by itself, with no carrier, caused little increase in the O, uptake
of the buffered extract of seeds, substantial increases were obtained on addition of
formate to ascorbic acid plus extract. To ensure that the increased oxygen uptake in
the presence of formate and ascorbic acid was not due to increased loss of ascorbic acid
rather than increased loss of formate, ascorbic acid was estimated at the end of each
experiment by titration with 2:6 dichlorphenol indophenol in acid solution (Harris and
Olliver, 1942). The results of a typical aerobic experiment are shown in Table 12.

TABLE 12.
Effect of the Ascorbic Acid System on Formate Oxidation.

Flask contents: 2:0 ml. enzyme preparation from pea seeds, 0-2 ml. coenzyme I
(120 wg.), 0-3 ml. water or 0-3 ml. ascorbic acid (0-01 mM) in one side arm,
0-3 ml. water or 0:3 ml. 0-2 M sodium formate in the other side arm, 0-2 ml.
20% NaOH in centre well.

Total Loss of
ul. O. Uptake Ascorbic Acid.
in 60 Minutes. (mg.)

Blank He ite af at oe 56 —
Formate is 2 as a oe 59 —
Ascorbic acid ay ox Be we 87 0-57
Ascorbie acid+formate .. Ree Pe 129 0-60

This experiment is typical of many and, in all cases, in the presence of ascorbic acid
and formate, there were increases in oxygen uptake over and above that caused by
ascorbic acid alone. In most experiments these were not accompanied by any increased
loss of ascorbic acid and even if some increased loss were observed (as in Table 12),
it was not equivalent to the extra oxygen consumed. For example, the increase of
0:03 mg. ascorbic acid lost here would only entail an increased oxygen uptake of 2ul., a
negligible quantity.

It seems clear then that the increased oxidation (hydrogen loss) is finally borne by
the formic acid and not by ascorbic acid. That is, the ascorbic acid system is acting as a
carrier in formate oxidation. The chain of reactions responsible for formate oxidation
here seems to be:

CH,OH CH,OH
. |
is(O—C——lal HO—C—Fi

| ascorbic |
H—C——_—— acid He

| -H,O+0 > |

C—OH oxidase HO—C—OH

| | | O

C=0H 0 HO—C—OH |

|

mae cca |

| ||

O
coms

reduced coenzyme I
Ascorbic Dehydroascorbie
acid acid

BY DAPHNE C. DAVISON.

47

The reduced coenzyme is produced during the formate oxidation and after being
reoxidized by the dehydroascorbie acid is again ready to be reduced by formate.

A buffer extract of seedlings gave evidence of a very active ascorbic acid oxidase
and, as with seeds, formate oxidation is increased by the ascorbic acid system although
the effect was not so marked probably because of the lower formic dehydrogenase activity

as this stage (Table 13).

TABLE 13.

Effect of the Ascorbic Acid System on Formate Oxidation in Seedlings.
Manometer flask contents as for Table 12.

ul. O. Uptake

Total Loss of
Ascorbic Acid.

in 40 Minutes. (mg.)
Blank 84 =
Formate 83 pas
Ascorbic acid é 179 1-50
Ascorbie acid +formate 186 1-56

The increased importance of ascorbic acid oxidase in the endogenous respiration of
the seedling as revealed by inhibitor studies is discussed below.
In the anaerobic experiments coupling between ascorbic acid oxidase and formic
dehydrogenase was demonstrated by observing the reduction of a redox indicator by the
negative system (formic dehydrogenase + formate + coenzyme I) and reoxidation of the

dye by the positive system (ascorbic acid oxidase + ascorbic acid).

The reducing system

together with the redox dye are placed in the main part of a Thunberg tube and the

components of the oxidizing system are placed in the hollow stopper.

In the control

tube water takes the place of ascorbic acid. After evacuation and filling with nitrogen
the tubes were incubated at 38°C. until the maximum reduction of dye is obtained.
Then the contents of the stopper are tipped in and if any reoxidation of the dye occurs,
the time taken for a second decolorization is noted.

It is evident that an indicator which is either more positive than the oxidizing
more negative than the reducing system cannot function as a carrier.
Theoretically any reversible indicator whose potential lies in the range between the two

system er

enzyme systems should be able to catalyse the interaction if one is to occur.

TABLE 14.

Methylene

Ascorbic Acid Oxidase-Formic Dehydrogenase Coupling in a pH 6-0 Buffer Extract of Pea Seeds.

Main tube—

Enzyme preparation

Coenzyme I (120 tg.)
0:2M sodium formate
0:05% redox dye

Stopper—

Enzyme preparation
Ascorbic acid (1-76 mg.)

Water

Time for first decolorization

Time for second decolorization

1 2 3 4
0-5 mil. 5 0-5 0-5
0:2 ml. as OR 2 0:2
0-5 ml. “5 0-5 0-5

Nile |blue Methyle|ne blue
0-5 ml. af) 0-5 0-5
0-5 ml. 5) 0-5 0-5

— -3 mil = 0-3
0:3 ml. — 0-3 =
25 mins. 25 mins. 17 mins. 17 mins.

No colour 80 mins. No colour |developed
developed

48 FORMIC DEHYDROGENASE IN THE OXIDATION MECHANISMS OF PISUM SATIVUM,

blue and thionine, whose potentials of + 0-011 and + 0-63 respectively are more positive
than the oxidizing system, were not suitable for these experiments. The carrier most
suited was Nile blue, whose potential of —0-116 lies in the range between the potentials
of the two enzyme systems. In all experiments with Nile blue the indicator was either
partly or completely reoxidized by the ascorbic acid system before being again reduced
to the leucoform. The extent of the reoxidation was found to be determined by the
relative activities of the oxidizing and reducing systems as shown by control manometric
experiments. The ascorbic acid and extract were mixed some time before gassing to
allow for formation of the dehydroascorbic acid necessary for the reoxidation of the
leuco-dye. Controls with enzyme + water in the stopper showed no trace of reoxidation
of the indicator.
From gas exchange measurements the equation for the oxidation of formate by
tormic dehydrogenase of pea seeds has been worked out by Adler and Sreenivasaya
(1937) and found to be: HCOOH + 4 0,——>H.0 + CO,. From the results in Table 14,
then it seems that the entire system in such anaerobic experiments is:
(a) Formate + coenzyme I ——> coenzyme I H, + CO.
(bo) Coenzyme I H, + dye —— coenzyme I + leuco-dye.

Possibly flavoprotein is involved as carrier between coenzyme and dye in reaction (bd).
(c) Ascorbic acid + H,O + % O,

IN
~ ascorbic
+ dye H, \ acid
(or leuco dye) \ oxidase

Dehydroascorbie acid.

The nature of the mechanism involved in the reduction of dehydroascorbic acid has
not been investigated. The close association of glutathione and ascorbic acid is seen in
the observation that in germinating pea seeds these two substances appear at the same
time and increase in concentration in a parallel manner (Hopkins and Morgan, 1943).
In any such plant tissue the possibility that the ascorbic acid—-glutathione relationships,
as outlined by Hopkins and Morgan (1936), have some physiological function cannot be
overlooked. That the reduction of ascorbic acid by glutathione in some plants is due
to a reducing catalyst, an enzyme to which the name dehydroascorbic acid reductase
has been given, is now an accepted fact and its properties, pH optimum, occurrence, etc.,
have been thoroughly worked out (Crook, 1941). Reducing catalysts of this type have
been demonstrated in the juice of peas and beans by Kohman and Sanborn (1937), who
showed that the juice of peas catalysed the reduction of dehydroascorbic acid by reduced
glutathione (G.S.H.). However, as shown by Crook and Morgan (1944), the activity of
the dehydroascorbic acid reductase is much lower than that of ascorbic acid oxidase so
that the presence of G.S.H. does not completely protect ascorbic acid from oxidation.
Some other system for the reduction of dehydroascorbic acid must therefore be operative
in peas. Possibly there is a simple transfer of hydrogen from coenzyme I to dehydro-
ascorbic acid. ;

Peroxidase.—In the question of ascorbic acid oxidation the presence of a specific
ascorbic acid oxidase as distinct from a cytochrome catalysed oxidation, and the absence
of any catechol action has left only the possibility of a peroxidase forming a terminal
oxidase system with ascorbic acid.

It was found that a fresh extract from either pea seeds or seven-day seedlings would
not produce any colour with 0-1 ml. of a 1% solution of guaiacol, benzidine (slight
coloration) or a-naphthol. However, on the addition of 1 drop of 6% hydrogen peroxide
to each tube, there was a marked positive reaction. Peroxidase then is present in both
stages.

An experiment carried out by James and Cragg (1943) on barley sap was repeated
with pea extract. The latter plus hydrogen peroxide will oxidize benzidine almost
immediately. If ascorbic acid is added there is a lag period until the oxidation of the
ascorbic acid by the quinone formed (p-quinone di-imide) is complete. Then the blue

BY DAPHNE C. DAVISON. 49

colour of the oxidized benzidine suddenly appears. As in James’ experiment, it was
found with pea extract that the length of the lag was proportional to the amount of
ascorbic acid added. From this it seems inevitable that, if hydrogen peroxide is formed
in the pea seed, the peroxidase system should co-operate in the oxidation of ascorbic
acid. However, the absence of H.O, in pea extract is shown in experiments quoted above
which showed peroxidase activity only when H.O. from an external source was added.
Also it is doubtful whether enough H.O. can occur in cells which, like those of the pea,
contain a strongly active catalase; and whether sufficient phenols are present is also
somewhat unlikely.

Further, 0:001 M cyanide almost completely inhibits ascorbic acid oxidation in a
pH 6:4 buffer extract of pea seeds. At this concentration the inhibition of peroxidase is
far from complete, so that peroxidase cannot be responsible for the oxidation of ascorbic
acid to the exclusion of a direct ascorbic acid oxidase.

What the value of peroxidase is to the plant is still an open question. On the whole
it seems extremely unlikely that peroxidase should combine with ascorbic acid in
forming a terminal oxidase system.

INHIBITOR STUDIES.

Some ideas on the relative importance of the various oxidation mechanisms in the
respiration of the pea seed and seedling can be obtained from studies using specific
inhibitors. The following experiments on the effect of inhibitors on endogenous respira-
tion were all carried out at the normal reaction of pea extract, pH 6-4. The enzyme
preparation is an untreated water homogenate centrifuged for 10 minutes. Manometer
flasks contained 1:9 ml. of this supernatant, 0-8 ml. 0-2 M phosphate buffer pH 6-4,
0-2 ml. 20% NaOH in the centre well. In the side bulb 0-3 ml. water or 0:3 ml. inhibitor
was placed. All inhibitors were neutralized before use and made up to their final
concentration in phosphate buffer pH 6-4. Oxygen uptake was measured over a period of
60 minutes and all results are given as percentage inhibition caused by the inhibitor.
Under the conditions set down above the oxygen uptake for seed preparation in 60
minutes is 42—55 ul., and for a seedling preparation, 50-67 wl.

Cyanide.—Both the cytochrome system and the ascorbic acid system are sensitive
to small concentrations of cyanide. The inhibitor is, therefore, by no means specific
but its effects are interesting in determining the activity of the respiratory system
resistant to poisoning with dilute cyanide. One cannot conclude at present that such a
fraction of the respiratory oxygen uptake is necessarily operated by flavoprotein systems.
However, coupled with the observation that the flavoprotein content of peas rises on
germination (after two days it increases to three times its original value; van Herk,
1935) the fact that cyanide inhibition is less in the seedling (Table 15) makes the flavo-
protein hypothesis extremely likely.

TABLE 15.
Inhibition Produced by Cyanide on Seeds and Seedlings.

Percentage Inhibition of Control.
Cyanide (Final
Concentration).
Seeds. Seedlings.

ea Ome Vi 58 25
5x10°3M 52 24
1x105°M 45 18
5 <x1074M 39 15
1x1074M 31 11
1x10°°>M 22 9

Another point to be considered here is that no serious inactivation of dehydro-
genases is obtained with cyanide concentrations of less than 0-002 M. The one exception

50 FORMIC DEHYDROGENASE IN THE OXIDATION MECHANISMS OF PISUM SATIVUM,

to this, however, is formic dehydrogenase, which is inhibited by concentrations of
1:3 x10 M. The fact that formic dehydrogenase activity decreases in the seedling is in
agreement with the decrease in cyanide inhibition in the seedling.

In experiments on cyanide loss of concentration of the inhibitor by distillation into
the alkali of the centre cup was prevented by using cyanide-alkali mixture (Krebs, 1935).

Azide.—This inhibitor is specific in that it poisons the cytochrome system but not
the ascorbic acid system (James, 1946). The results obtained with various concentra-
tions of azide are summarized in Table 16.

TABLE 16.
Inhibition Produced by Azide on Seeds and Seedlings.

Percentage Inhibition.

Azide Concentration.

Seeds. Seedlings.
10-2M aye site ei Ree 53 38
10-7 M Bd se bbe Aa 43 35
10°4*M se es a oe 37 26
107>M wee an aS as 21 12

Diethyldithiocarbamate.—Ascorbic acid oxidase is strongly inhibited by this copper
precipitant but cytochrome oxidase only slightly (James, 1946). Coupled with the
results obtained with azide then the figures in Table 17 can give some information on
the relative activity of these two terminal oxidases in seed and seedling. It appears
that azide inhibition is greater in the seedling whereas diethyldithiocarbamate is more
effective in inhibitng the oxygen uptake of the seed extract. These results can be
correlated with those reported earlier in this paper, namely, the increased ascorbic acid
oxidase activity on germination and the decreased effect of cytochrome on seedling
respiration. It seems very probable then that while these two terminal oxidases are of
more or equal value to the economy of the seed, the ascorbic acid system becomes the
more important in the seedling.

TABLE 17.
Inhibition Produced by Diethyldithiocarbamate on Seeds and Seedlings.
Percentage Inhibition.
Diethyldithiocarbamate
Concentration.
Seeds. Seedlings.
NO-SBIML 56 a0 a We oe 31 40
UOTE 5. ae oh Le Ble 25 36
TOS oe Ae oe ve a6 19 28

Iodoacetic Acid.—Triosephosphate accumulates in iodoacetate-poisoned barley digests
containing hexosediphosphate, and the acid is known to inhibit also the alcohol dehydro-
genase of oat seedlings (Berger and Avery, 1943). James (1946, p. 429) and Turner
(1937) agree in attributing the sensitivity of plant respiration to iodoacetate poisoning
to an inhibition of the triosis stage of glycolysis. Table 18 shows the inhibition produced
by iodoacetate on pea seeds and seedlings.

The increased inhibition on germination can be correlated with the fact that the
C, acids and citrate have been found to exert more effect in the seedling. Albaum and
Hichel (19438) found similar effects in the oat seedling where iodoacetate inhibition and
C, acid stimulation could be demonstrated only after 72 hours of growth. They were

BY DAPHNE C. DAVISON. 5]

TABLE 18.
Inhibition Produced by Todoacetate on Seeds and Seedlings.

Percentage Inhibition.

Todoacetate Concentration.
Seeds. Seedlings.
OWE 5. ae ee ae Aa 38 58
NOTE ME 5 - Aes abs By a3 21 3
UOTE 5, Ds ne ie aes 14 30
WOFSME os oe ae oe Ae 11 22

able to conclude from this and other evidence that fat is metabolized mainly in the early
stages and that sugar is the main metabolite after 72 hours. However, pea seeds have
very little fat, none of which is storage material but only phosphatides concerned in
the cell and nuclear structure. Also, in the seedling, pyruvate and the dicarboxylic acids
should produce a reversal of iodoacetate inhibition if this substance is poisoning the
triosis stage of glycolysis. Malate and glucose were found to overcome this effect slightly
but certainly not when used in catalytic amounts. Succinate and pyruvate were entirely
without effect. ;

It is impossible to conclude then that iodoacetate is inhibiting only the “triosis
stage of glycolysis” in peas or to make any suggestions as to the type of respiratory
substrate being used either before or after germination.

Malonate.—This acid is known to inhibit slightly succinic dehydrogenase and has
- been used to elucidate the relations between the C, acids making up what is known as
the Szent-Gyorgyi cycle. Neither in the pea seed nor seedling could any significant
inhibition be obtained with malonate, however, the results being given in Table 19.

TABLE 19.
Inhibition Produced by Malonate on Seeds and Seedlings.

Percentage Inhibition.

Malonate Concentration.
Seeds. Seedlings.
WOT 5.5 9 10
Om? Mie 8 5
1OmA Mae: 5 iG
Ope Vinee 4 3

The whole question of the significance of malonate inhibition in plants is at present
uncertain, but it seems clear from these results that no C, acid system in the sense of
the Szent-Gy6rgyi cycle is operating here.

Semicarbazide.—This substance has an interesting effect on the endogenous respira-
tion and alcohol oxidation of pea seeds and seedlings as shown in Table 20.

These results show that seedlings contain much less active alcohol dehydrogenase
than seeds and also the effect of semicarbazide is quite different in the two materials.
In the seed alcohol oxidation is greatly accelerated by semicarbazide presumably because
of its binding effect on the reaction product, acetaldehyde. Endogenous respiration in
the seed is also accelerated by semicarbazide. In the seven-day seedling the substance
was found to inhibit endogenous respiration and, to a lesser extent, alcohol oxidation.
These effects suggest that endogenous respiration in seeds is partly due to alcohol
dehydrogenase whereas in the seedling it is not.

52 FORMIC DEHYDROGENASE IN THE OXIDATION MECHANISMS OF PISUM SATIVUM,

TABLE 20.
Effect of Semicarbazide on the Endogenous Respiration and Alcohol Oxidation of
Pea Seeds and Seedlings.

Manometer flask contents: 1:0 ml. untreated water extract, 0:8 ml. 0-2M
phosphate buffer pH 7:9, 0-3 ml. coenzyme I (90 Ug.), 0-1 0:5% methylene
blue (one side bulb), 0-3 ml. 3-2 M ethyl] alcohol, and 0:5 ml. 0-25 M semi-
carbazide solution as indicated. Water to 3-2 ml.

ul. O, Uptake in 90 Minutes.
Addition.
Seed. Seedling.
Blank .. aN a ant ae 100 65
+Semicarbazide Sf ae Be 165 47
Alcohol .. a ae me 133 75
Alcohol +semicarbazide Bs Ne 337 62

In support of this hypothesis of an important role for alcohol dehydrogenase in the
endogenous respiration of the seed we find that pyruvic decarboxylase activity which is
extremely high in the seed falls off with germination (Tables 21, 22).

TABLE 21.
Carborylase Activity in the Seed.

Flask contents: 1:4 ml. pea extract, 0-8 ml. 0:2 M phosphate buffer pH 6-85,
0-1 ml. 0-5% methylene blue, 0-3 ml. 0-03 M pyruvate or 0-3 ml. water in
the side arm. R.Q. was determined by the direct method outlined in
Umbreit, Burris and Stauffer (1945, p. 16) over a period of 60 minutes.

O, Uptake. CO, Output. iRE@)>

Blank .. a 56 401. 661. 1-66

Pyruvate =e ate 25 wl. 275 wl. 10-83
TABLE 22.

Carboxylase Activity in Seven Day Old Pea Seedlings.
Manometer contents as in Table 21. These results are for an 80-minute period.

O, Uptake. CO, Output. R.Q.
Blanket eee ee 107 ul. 166 ul. 1°55
Pyruvate Bhs ahs 96 wl. 197 wl. 2-03

With the seeds 96% of the pyruvate added was decarboxylated in 60 minutes,
whereas in the seedling extract only 14% was decarboxylated in 80 minutes. This
decline of a reaction which seems the most likely source of alcohol, at a time when there
is a parallel decline of alcohol dehydrogenase, points to a probable use for the dehydro-
genase in the economy of the seed.

Finally, in discussing the place of alcohol dehydrogenase in the seed, it can be
recorded here that the endogenous respiration of the seed is accelerated by the addition
of small amounts of coenzyme I whilst in the seven-day seedling the effect is less marked
(Table 23). ,

Van Herk (1935) points out that the cozymase of germinating peas is initially large
and remains so for four days, but after seven days it has fallen to one-fifth or one-sixth
of its original value. These observations make it extremely likely that much of the

BY DAPHNE C. DAVISON. 53

TABLE 23.
Effect of Coenzyme I on Endogenous Respiration.
Flask contents: 1:6 ml. untreated water extract of peas, 0-8 ml. 0-2 M
phosphate buffer pH 6-4, coenzyme I added from side arm
(30ug.-180 zg.), 0-2 ml. 20% NaOH in centre well.

ul. O, Uptake in 120 Minutes.
Amount of Coenzyme I
Added.
Seeds. Seedlings.
= ae a He a 50 181
30ug. eee a 53 182
60 ug. fils as ul 58 185
120ug. i a! zt 65 186
180g. is Sey gies Ut 76 188

dehydrogenation in the seed depends on coenzyme I-requiring dehydrogenases and less
so in the seven-day seedling. The only plant dehydrogenases known to need coenzyme I
are alcohol, formic, lactic, malic, glutamic, $-hydroxybutyric and _ triose-phosphoric
dehydrogenases. The first five have been studied in peas and only alcohol and formic
dehydrogenase have been found to fall off in activity during germination. The difficuluty
in the way of proving formic dehydrogenase as a normal link in biological oxidation in
the pea is that there is no specific inhibitor of the enzyme which, by application to the
endogenoyis respiration, can prove a Similar inhibition. With alcohol dehydrogenase,
however, there is the supporting evidence of the semicarbazide and carboxylase experi-
ments quoted above.

In the seed, then, alcohol dehydrogenase and possibly formic dehydrogenase are the
main enzymes concerned in the first or dehydrogenating step of biological oxidation.

DISCUSSION.

On examining the results obtained with inhibitors in seeds and seedlings some
correlation can be obtained with the increase or decrease of certain enzymic activities
with growth. The results obtained have been tabulated below (Table 24).

TABLE 24,

Correlation of Results Obtained with Inhibitors in Seeds and Seedlings with the Increase or Decrease of Certain Enzymic
Activities with Growth.

Hydrogen Carriers. Inhibitors.
I. Seeds—
Ascorbie acid oxidase present. 10-3 M diethyldithiocarbamate gives 31% inhibition.
Cytochrome system operative. 10m2 M ‘cyanide—45%:
Alcohol and formic dehydrogenases active. 10°2°M azide—43%.
C, acid system—no activity. 10-2 M iodoacetate—21%.
High carboxylase activity. 10-2 M malonate gives no inhibition (2%).
Added coenzyme I has marked effect on endogenous Semicarbazide gives 63% acceleration of endogenous
oxygen uptake. respiration. :
II. Seedlings—
More ascorbic acid oxidase activity. Diethyldithiocarbamate inhibition rises to 40%.
Less cytochrome effect. Cyanide and azide inhibition fall to 18% and 35%
Alcohol dehydrogenase almost absent. No malonate inhibition.
Formic dehydrogenase less active. Todoacetate inhibition rises to 43%.
Flavoprotein effective in formate oxidation. Semicarbazide inhibits by 28%.
C, acids slightly more effective.
Low carboxylase activity.
Added coenzyme I has less effect on endogenous O,
uptake.

54 FORMIC DEHYDROGENASE IN THE OXIDATION MECHANISMS OF PISUM SATIVUM,

It is difficult to interpret the effects of poisons on such complex processes as oxida-
tion since so many consecutive reactions are involved. However, coupled with the
information obtained on the activity of the hydrogen carriers the following general
conclusions can be stated.

1. Based on the coenzyme activation of the respiration of seed extract and the
degree of activity of the various dehydrogenases in seed and seedling, it seems likely
that in the seed alcohol dehydrogenase and formic dehydrogenase are the main enzymes
concerned in the first or dehydrogenating step of biological oxidation. This is supported
in the case of alcohol dehydrogenase by the falling off of carboxylase activity in the
seedling, a reaction which seems the most likely source of alcohol and the acceleration
of seed respiration but not that of seedlings by semicarbazide.

2. Formic dehydrogenase is capable of coupling with other dehydrogenases. ;

3. Development of both iodoacetate inhibition and C, acid utilization with germina-
tion indicate the utilization of sugar as respiratory substrate only after about 72 hours.
However, this could not be confirmed by demonstrating a removal of iodoacetate inhibition
on the addition of C, acids and pyruvate.

SEED

FORMATE=@HycOENZYME I—Y FLAVOPROTEIN
COUPLED FCAYWALASE |
REACTIONS

Cc SYSTEM
y (?)
OTHER
3UBST

)
RATES SSRs
We “SS. cyTocHRomeE
AVSNOLOURUB TC! ea, GY S TELM
ACID

SYSTEM

SEEDLING

FORMAT pTE——2HycoOENZYME ean

COUPLED TCA LASE
REACTIONS

c eos
H
B

RATES Jeane

SYSTEM
a
AQID

SYSTEM

ie SLE
—— indicates main terminal oxidase.

SCHEME II

4. Lack of malonate inhibition and of noticeable effects on addition of catalytic
amounts of C, acids make the operation of the Szent-Gyérgyi cycle extremely improb-
able. However, formate oxidation is accelerated by two of these acids, malate and
fumarate and also citrate so that they may be of some importance in biological oxidation.

5. The main terminal oxidase in the seed is cytochrome oxidase. With germination
this activity falls off and the activity of ascorbic acid oxidase, already of importance in
the seed, increases. Finally, after seven days, most of the cyanide sensitive respiration
can be accounted for by the oxygen consumption of the ascorbic acid system. Formate
oxidation has been shown to be coupled with both cytochrome and ascorbic acid systems
in seed and seedling.

BY DAPHNE CG. DAVISON. 55

6. With regard to the cyanide insensitive respiration which seems to amount to a
fairly high percentage (41% in the seed, 75% in the seedling), this is in all probability
due to a flavoprotein system since there is a corresponding rise in flavoprotein content
on germination. Also formate oxidation is accelerated by flavoprotein, especially in the
seedling.

Some conclusions on the oxidation mechanisms of the seed and seven-day seedling,
and the place of formic dehydrogenase therein, can now be made. The paths used in
the breakdown of formate have been narrowed down and, in place of Scheme I, the
following diagrams give mechanisms for the oxidation of this substrate, based on the
results of experiments quoted above (Scheme IT).

SUMMARY.

1. Various oxidation mechanisms of the green pea seed and seedling and their
relation to formic dehydrogenase have been studied. These include other dehydro-
genases, intermediate hydrogen carriers such as flavoprotein and the dicarboxylic acids
and terminal oxidases.

2. Asa result of these studies a normal role in the first or dehydrogenation step of
biological oxidation can be assigned to formic dehydrogenase.

3. Inhibitor studies on seed and seedling extract have revealed some significant
changes in enzyme activity on germination.

ACKNOWLEDGEMENT.
This work has been supported by a grant from the Commonwealth Research Committee.

References.

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Bercer, J.. and Avery, G. S., 1943.—Action of synthetic auxins and inhibitors on dehydrogenases
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G

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WESTERKAMP, H., 1933.—Uber Ketoséuren im Blutserum. Biochem. Z., 263: 239-244. :

WILLIAMSON, S., and GREEN, D. H., 1940.—The preparation of coenzyme I from yeast. J. biol.
Chem., 135: 345-346.

57

ON THE AUSTRALIAN SPECIES OF CREOPHILUS (COLEOPTERA:
STAPHYLINIDAE).

By W. O. STEEL.
(Communicated by J. W. T. Armstrong.)

(Nine Text-figures. )

[Read 30th March, 1949.]

INTRODUCTION.

Three species of Creophilus have been recorded from Australia, erythrocephalus
{Fab.), lanio (Er.) and oculatus (Fab.).

Erythrocephalus is a very well known species and is common in carrion throughout
Australia. It has also been recorded from Tahiti and Chile.

Lanio is listed by Bernhauer and Schubert (Col. Cat. 57, 1914, 398) as a distinct
species, but is stated by Lea (Trans. Roy. Soc. S. Aust., 1925, 229) to be synonymous
with erythrocephalus. This latter view has apparently been accepted, as no records of
lanio can be traced in the literature.

Some years ago two specimens (4, 2) of a Creophilus collected at Lake Burrumbeet,
Vic., were received from Mr. M. F. Leask of Ballarat. These were obviously distinct
from erythrocephalus and agree well with Hrichson’s description of lanio. An examina-
tion of the series of erythrocephalus in my collection brought to light a further specimen
of lanio and the British Museum collection provided about a dozen more. In all, well
over 100 specimens of “erythrocephalus” were examined.

Oculatus is recorded by Fauvel (Ann. Mus. Civ. Gen., 1877, 250) as occurring in
Australia. This species is very common in carrion in New Zealand, and Lea (l.c.)
states that Fauvel’s Australian record requires confirmation, at the same time placing
the species as a variety of erythrocephalus. There is one specimen in my collection
from Rockhampton, Queensland (ex Pilcher coll.), and this is the only Australian
specimen seen up to the present. The insect would, however, be expected to establish
itself fairly quickly should it gain a footing in Australia.

The main purpose of this paper is to point out the differences between erythro-
cephalus and lanio, but it has been thought advisable to give, at the same time, full
descriptions and figures of the three Australian species of the genus, together with a
short key.

CREOPHILUS ERYTHROCEPHALUS (Fabricius). (Text-figs. 1, 4, 9.)
Staphylinus erythrocephalus Fabricius, Syst. Ent., 1775, 265.
Staphylinus unipunctatus Hope, in Gray, Zool. Misc., 1831, 32.

Head and pronotum shining, elytra (except humeral areas) and abdomen duller.
Head red, the dise with a rather small circular black spot, the edge of which is sharply
defined; narrowly black around the insertions of the antennae. Pronotum and abdomen
black, elytra blue-black, the humeral areas black. Antennae black, segments 7-11 with
close-set, short, greyish pubescence. Mandibles, palpi and legs black. Length: 17-22 mm.

Male.

Head as wide as or somewhat wider than the pronotum, about one and one-half to
one and three-fifths times as broad as long,* distinctly widened behind, the sides prac-
tically straight, the posterior angles very distinct, rather sharply rounded (Text-fig. 9),
the base almost straight. On the inner margin of each eye, towards the front, with a

* Wor the length, the head is measured from the “occipital suture’ to the front of the
clypeus.

58 AUSTRALIAN CREOPHILUS (COLEOPTERA: STAPHYLINIDAE),

well-marked setiferous puncture and a transverse row of some four similar punctures
near the base, the whole surface with rather diffuse, very fine punctules and between
these an indistinct ground-sculpture consisting of extremely fine, close-set wavy striae.
Antennae with the third segment slightly shorter than the second, the fourth subquadrate,
slightly broader than long, the fifth to tenth gradually becoming more transverse, the
eleventh about twice as long as the tenth, bluntly pointed.

Pronotum a little longer than the head, very slightly broader than long, widest in
front of middle, the sides slightly rounded, the anterior angles distinct, scarcely rounded,
the posterior angles normally very broadly rounded (sometimes much less so*), the sides
slightly emarginate just in front of these angles, the base rounded. Towards the anterior
angle on each side with a small patch of rather close punctures, each of which bears a
short seta; elsewhere impunctate and without visible ground-sculpture.

Scutellum rather closely punctured and with dark pubescence.

Elytra about one and one-sixth times as long as pronotum, very slightly broader
than long, a little widened behind. A small area around each humeral angle impunctate
and with, at most, traces of a rather coarse ground-sculpture; elsewhere closely punctured
and with dark pubescence, the surface between the punctures reticulate.

Tergites of abdomen finely and closely punctured and with dark pubescence, the
surface between the punctures with a ground sculpture consisting of extremely fine and
close, transverse, wavy striae. The sternites sculptured as the tergites. The apical
margin of the eighth sternite with a slight, median, triangular emargination.

Aedeagus as Text-figs. 1 and 4.

Female.

Head at most as wide as pronotum, often distinctly narrower, about one and two-
fifths to one and one-half times as broad as long, rather less widened behind than in
the male, with the posterior angles rather more broadly rounded. Sternite of the eighth
abdominal segment rounded apically.

In other respects, outwardly similar to male.

The differences between this species and lanio are discussed after the description
of the latter. -

CREOPHILUS LANIO (Hrichson). (Text-figs. 2, 5, 7, 8.)

Staphylinus lanio Hrichson, Gen. et Spec. Staph., 1840, 352.

Staphylinus erythrocephalus Fabricius, Syst. Ent. 1775, 265 ex parte.;

Staphylinus oculatus var. Gravenhorst, Mon. Micr., 1806, 127.

Head and pronotum shining, the elytra (except humeral areas) and abdomen some-
what duller. Head red, the disc with a rounded black spot which is wider than long,
with the edge not sharply defined; narrowly black around the insertions of the antennae.
Pronotum and abdomen black, elytra black, with a purplish reflection. Antennae black,
segments 7-11 with close, short greyish pubescence. Mandibles, palpi and legs black.
Length: 17=23) mm:

Male.

Head rather variable in size and shape. In large specimens (Text-fig. 7) somewhat
broader than the pronotum, rather more than one and three-fifths times as broad as long,
distinctly widened behind, the sides almost straight, the posterior angles rather broadly
rounded, the base almost straight. In small specimens (Text-fig. 8) narrower than the
pronotum, rather less than one and one-third times as broad as long, distinctly less

*JIn all the specimens seen, except two, the posterior angles of the pronotum, though
showing some variation, are broadly rounded and sometimes hardly discernible. The two
exceptions (two females collected by Mr. J. W. T. Armstrong, near Nyngan, N.S.W., and in
my collection) have these angles very well marked, just as marked as in lanio.

7 The type material of S. erythrocephalus Fab. is in the British Museum collection and
consists of two specimens. One of these is the species normally known as erythrocephalus
Fab., the other is lanio Er. To save confusion in nomenclature, the first mentioned specimen
is taken as the type of the species.

BY W. 0. STEEL. 59

widened behind, the sides rounded, the posterior angles very broadly rounded. Various
intermediates occur between the above-mentioned two forms. Sculpture similar to that
of erythrocephalus.

Pronotum similar to that of erythrocephalus, but with the posterior angles always
well marked.

Aedeagus as Text-figs. 2 and 5.

In other respects similar to erythrocephalus.

Female.

Head at most as wide as pronotum, usually narrower, about one and one-third to
one and one-half times as broad as long, widened behind, the posterior angles broadly
rounded.

In other respects, outwardly similar to male.

The head in the males of this species is very variable in size and shape, much more
so than is the case with erythrocephalus, but, even in large specimens, the posterior
angles are distinctly more rounded than in the latter species (cf. Text-figs. 7 and 9).
The black discal spot on the head is distinctly larger than in erythrocephalus, some-
times markedly so, and is broader than long, with the edge rather diffuse, instead of
being sharply defined and circular.

The posterior angles of the pronotum are always distinct and may normally be
used as a determinative feature, though, as mentioned above, specimens of erythro-
cephalus are occasionally encountered which have these angles just as marked.

The elytra are constantly black in colour, with a purplish reflection, in marked
contrast to the blue-black of erythrocephalus.

As will be seen from a comparison of Text-figs. 1 and 2, the aedeagi, particularly
the median lobes, of erythrocephalus and lanio are quite different and show without
doubt that the two species are distinct.

From material examined, lanio appears to be rather an uncommon species, though
it has most probably been mixed with erythrocephalus in collections. Having regard
to the fact that its habits would be expected to be the same as those of erythrocephalus,
which is common in carrion throughout Australia (as are other species of the genus in
the countries where they occur) it would be thought that lanio would be abundant where
it occurs. All the specimens seen have been collected in the south-eastern part of
Australia and the species may be restricted to that area. Specimens have been seen
from the following localities: Lake Burrumbeet, Vic.; M. F. Leask (Steel coll.);
Tasmania: M. F. Leask (Steel coll.); Victoria (British Museum, ex Sharp coll.);
Tasmania (British Museum, ex Sharp coll.).

CREOPHILUS OCULATUS (Fabricius). (Text-figs. 3 and 6.)

Staphylinus oculatus Fabricius, Syst. Ent., 1775, 265.

Head and pronotum shining, elytra (except humeral areas) and abdomen duller.
Head black, with a small red area on each side between the eye and the posterior angle,
pronotum black, elytra black, with a slight brownish reflection, abdomen black. Antennae
black, segments 7-11 with close set, short, greyish pubescence. Mandibles, palpi and legs
black. Length: 14-23 mm.

Male.

Head rather variable in size and shape, in large specimens slightly wider than the
pronotum, about one and three-fifths times as broad as long, scarcely widened behind,
the sides almost straight, the posterior angles broadly rounded, the base almost straight.
In small specimens, slightly narrower than the pronotum, a little less than one and one-
third times as broad as long, slightly widened behind, the sides straight, the posterior
angles broadly rounded. Various intermediates occur between the above-mentioned two
forms. Sculpture similar to that of the preceding two species.

Pronotum similar to that of erythrocephalus, the posterior angles broadly rounded
and not very distinct.

60 AUSTRALIAN CREOPHILUS (COLEOPTERA: STAPHYLINIDAE),

Aedeagus as Text-figs. 3 and 6.
In other respects similar to the preceding two species.

Female.

Head rather small, distinctly narrower than the pronotum, about one and one-third
times as broad as long, slightly widened behind, the sides straight, the posterior angles
broadly rounded.

In other respects, outwardly similar to male.

7

Text-figures 1-9.

1. Creophilus erythrocephalus (Fab.). Aedeagus, ventral view. 2. Creophilus lanio (Er.).
Aedeagus, ventral view. 3. Creophilus oculatus (Fab.). Aedeagus, ventral view. 4. Creophilus
erythrocephalus (Fab.). Apical portion of aedeagus, viewed from left. 5. Creophilus lanio (Er.).
Apical portion of aedeagus, viewed from left. 6. Creophilus oculatus (Fab.). Apical portion of
aedeagus, viewed from left. 7. Creophilus lanio (Er.). Portion of side of head of large male,
showing posterior angles. 8. Creophilus lanio (Er.). Portion of side of head of small male, to
same scale. 9. Creophilus erythrocephalus (Fab.). Portion of side of head of male, showing
posterior angles.

This species may easily be separated from the preceding two by colour alone. It
also differs in the shape of the head and in the form of the aedeagus in the male. It -
is definitely a distinct species and there seems to be no justification whatsoever for
regarding it as merely a variety of erythrocephalus, as was done by Lea,

BY W. 0. STEEL. 61

The males show much variation in size and also in the size and shape of the head,

though this latter is not as marked as in lanio. The shape of the head in the females
appears to be remarkably constant, even in very small specimens (14 mm.).

bo

Key to the Australian Species of Creophilus.

FeAdeReG a tiemCiscmwAth eam blacks Otemermkecccncnsstere «sks “ncieiorels sicbaioe: ciel ener etee (ie eh eusledeliee, «, seers 2
Headsplackaathessidestapehind themeyess rede... ss tidn etiehe «niet cle) soa elders = oculatus (Fab.)
Elytra blue-black, discal spot on head circular, basal angles of head less rounded (Text-fig. 9),

aecdeasusmqdistinete @Lext-fieshelmamd 4) cn . sais de seiae s+ oe ue. erythrocephalus (Fab.)

Elytra black, with a purplish reflection, discal spot on head transverse, basal angles of head
more rounded (Text-figs. 7 and 8), aedeagus distinct (Text-figs. 2 and 5) .. lanio (Er.)

62

GRAPTOLITES FROM TALLONG AND THE SHOALHAVEN GORGE,
NEW SOUTH WALES.

By KATHLEEN SHERRARD, M.Se.
(Plates i and ii; thirty-three Text-figures. )

[Read 27th April, 1949.]

INTRODUCTION.

The village and railway station of Tallong are situated on the Southern Tableland
of New South Wales 2,000 feet above sea-level. Into the tableland the Shoalhaven River
has cut a gorge 1,500 feet deep. For four-fifths of its length this river flows north, but
about four miles south of the village of Tallong it swings to the east, maintaining this
direction for about 40 miles until it enters the Pacific Ocean near Nowra. Steeply
folded Upper Ordovician slates and quartzites are exposed on the tableland (Woolnough,
1909) as well as in the gorge. Veins in the slate have been mined for tin and copper
(Carne, 1911) at the Tolwong mine in the gorge.

While the mine was being worked graptolites were obtained ‘‘a little north of
Tolwong Extended” (Carne, 1911) and were described by T. S. Hall (1909), first in a
paper “On a Collection of Graptolites from Tallong, N.S.W.”. Since these graptolites
were not obtained from Tallong, in the present meaning of that locality name, some
explanation of this title is called for. The syllable “Tal” or “Tol” is in wide use in the
district, for example in Tallong, Talwong, Tallawa, Tallawal and Tolwong and is of
‘aboriginal origin. It is variously interpreted as meaning in English either ‘‘water” or
“tongue of land”. The word ‘“Tallong”’ did not solely signify the immediate neighbour-
hood of the railway station until 1905, when the name of the railway station, known
until then as “Barber’s Creek”, was changed to Tallong. (Information from Public
Relations Office of N.S.W. Railways Department.) It seems probable that the graptolites
were collected before this change, because the metalliferous lode at Tolwong in the gorge
was discovered in 1904, when the word “Tallong” had not the restricted significance it
later acquired. The collection may have been sent to Hall at the Melbourne University
with the locality label ‘‘Tallong”’. Hence arose the title of his first paper. A second
paper by the same author dealing with a collection from the same locality was published
in 1920 (five years after Hall’s death). This was called “On a Further Collection of
Graptolites from Tolwong”, although the specimens described were from the same
locality as those dealt with in the 1909 paper. The name “Tolwong” here was perhaps
introduced by the Editor, since by 1920 Tallong would be well known as the name of the
railway station. However, in the body of this second paper Hall writes of “Tallong’’.
A further complication is introduced, since the specimens Hall described, now in the
Mining Museum in Sydney, are labelled as from ‘“Talwong”’.

A certain amount of confusion has resulted from these various names. It has, for
instance, been stated erroneously that graptolites from Ballanya Hill, Tallong (Naylor,
1935) were described by Hall. In neither of Hall’s papers were described any graptolites
from Tallong, using that word to mean the neighbourhood of the railway line on the
Tableland level.

In the present paper graptolites from eight localities on the Tableland level near
Tallong are described. Several of these localities lie on the Razorback, which Dr.
_Woolnough (1909) defines as the long spur extending south between the Shoalhaven
River and its tributary, Barber’s Creek. Graptolites collected recently at four localities

at the river level in the Shoalhaven Gorge are also described.

BY KATHLEEN SHERRARD. 63

As far as can be judged from information available about the locality from which
the specimens described by Hall (1909, 1920) were obtained, it is not the same as any
of the four places in the Gorge from which graptolites described here were collected.
Carne (1911) describes it as ‘‘a bed of carbonaceous shale a little north of Tolwong
Extended”. In the same report, Tolwong Extended is said to be on Mining Lease 6.
The Geological Survey Office in Sydney supplied the information that Mining Lease 6
was about one mile north of the main mine. Its position is shown on the map with this
paper. It is two and a half miles directly south of the nearest locality from which
graptolites described in this paper were obtained. That locality is at the mouth of
Bungonia Creek and separated from Mining Lease 6 by the tortuous windings of the
Shoalhaven River as well as by being on the opposite bank. The graptolites recorded
by Hall differ to a certain extent from those obtained from any locality described in
this paper. He records forms characteristic of more than one zone.

The site of the Tolwong Mine is now deserted and forgotten. The mine and the
remains of the smelting works are on the east side of the Shoalhaven River and the
trestles of the “flying fox’? which supplied the mine are 1,000 feet above those remains
on the opposite or western side of the gorge. The trestles are connected with Bungonia
village by a road built when the mine was being exploited, but now fallen into disrepair.

The site of Mining Lease 6, which is extremely difficult of access now, was visited
in the course of the present work. The river has to be forded and may be high after
rain. The ground rises precipitously from the water’s edge. A reasonable area north
of the lease was carefully examined but no graptolites were found.

SOURCES OF COLLECTION.

Graptolites have been collected from the following localities (see Text-figure 33).

Locality 1.—Portion 109, Parish of Bumballa, junction Old Bungonia road and the
timber track leaving it to the south, 0:8 mile west of the Caoura road. Slates
dip 60 degrees E.S.E.

Locality 2.—Ballanya Trigonometrical Station.

Locality 3.—North-west corner of Portion 96, Parish of Bumballa, about 200 yards
east of road to Long Point Lookout at point two miles south of railway line.
Slates dip S.E. at 50 degrees.

Locality 4.—Eastern edge, Portion 95, Parish of Bumballa near disused mining
shafts in dry, hanging tributary to Digger’s Creek. (Turn off from Long Point
Look-out road 3-1 miles south of railway line.) Slates dip W.N.W. at 65 degrees.

Locality 5.—Public Reserve, Long Point. On narrow track, which, about 30 yards
from signpost, leads downhill off the main track from the car park to the Long
Point Lookout. Slates dip W.N.W. at 70 degrees.

Locality 6—On scenic track, west of car park at Long Point overlooking Barber’s
Creek, about a quarter of a mile west of signpost, about Portion 128, Parish of
Bumballa. Slates dip W.N.W. at 60 degrees.

Locality 7.—On track from Long Point to Shoalhaven River, low cutting in slate
near junction of tracks, about 400 yards west of shelter shed at Long Point
Lookout. Slates dip W.N.W. at 50 degrees.

Locality 8—On track from Long Point to Shoalhaven River, cuttings in slate
between 1,475 and 1,500 feet above sea-level, that is 525 and 500 feet below
Lookout.

In the Shoalhaven Gorge graptolites have been collected at the following localities:

Locality 9—Junction of Barber’s Creeek and Shoalhaven River, on. left bank of
creek. Shales dip east at 75 degrees.

Locality 10—Bank of Shoalhaven River, a quarter of a mile downstream from
mouth of Bungonia Creek on left bank of river. In highly dipping micaceous
phyllites.

Locality 11—Bungonia Creek, from its mouth to a quarter of a mile upstream in
slates dipping E.S.E. at 60 degrees.

Locality 12—Two hundred yards upstream from Badgery’s Crossing of the Shoal-
haven River on left bank of river in quartzite dipping 80 degrees south of west.

iH

64 GRAPTOLITES FROM TALLONG AND THE SHOALHAVEN GORGE, N.S.W.,

NATURE OF ROCKS.

The Marulan igneous intrusion which outcrops to the west of Tallong has meta-
morphosed the Ordovician slates. The degree of metamorphism affects the ease of split-
ting the slabs and the condition of the graptolites contained in them. Intense folding
has also affected them.

The slates of Locality 4 are the least affected. Slabs down to almost wafer thinness
and up to eight inches across can be obtained, though some of the slates are considerably
leached. In Localities 2, 6 and 7 incipient chiastolite crystals powder the rock and
slabs cannot be obtained more than three inches across so that big graptolites are always
incomplete. At Localities 3 and 5 the slate is strongly silicified and it is almost impos-
sible to break off anything but cuboidal blocks with sides not plane and not more than
one or two inches across. However, comparatively good specimens were obtained from
slabs exposed in a small quarry made some years ago at Locality 5.

At Locality § intense folding and crumpling has affected the slate as much as has
the igneous intrusion. At fossil Localities 9 and 11 in the bottom of the gorge the
slates split well. It is remarkable that slates with admirably preserved graptolites are
obtained at these localities, but for long stretches elsewhere in the Gorge the slates are
completely metamorphosed to phyllites with few or no traces of fossils. However,
Messrs. Packham and Veevers were successful in finding graptolites in a phyllite at
Locality 10. At Locality 12 the rock is a coarse quartzite. Graptolites are accompanied
by phyllocarids at this place.

GRAPTOLITE ZONES REPRESENTED.

The assemblages of graptolites given in Table 1 show some characteristic differences.
Locality 11 is characterized by overwhelming numbers of Climacograptus bicornis and
its former varieties, by Corynoides and fragments of Mastigograptus. Orthograptus
truncatus var. pauperatus, Climacograptus minimus, a very small Dicellograptus
angulatus and the delicate Plegmatograptus nebula are found in Localities 4 and 9. At
Localities 4 and 8 Cl. tubuliferous and Cl. caudatus occur. However, at any one of these
localities big diplograptids, often fragmentary, are likely to be found. An Upper
Ordovician age is clearly indicated for all these localities.

Miss G. L. Elles, who has been good enough to examine some specimens from
Localities 3, 4, 9, 11 and 12, writes: “I think there is no doubt about the horizon. I
should refer all of them except those from Badgery’s Crossing (Locality 12) to the
zone of Dicranograptus clingani and near its base.”

Localities 2, 5, 6, 7 and 8, from which no specimens were sent to Miss Elles, seem
to belong to the same zone. It may be that localities 2, 3, 4, 7, 8 and 9 represent a
higher section of zone 12 (Dicranograptus clingani zone) than Localities 5, 6 and 11.

Locality 1 is indeterminate. Localities 10 and 12 are entirely distinct. At Locality 10
the rock is a phyllite and hence the contained graptolites must have been subjected to
considerable pressure. Dicellograptus cf. sextans and a Diplograptid, cf. Amplexo-
graptus have been identified. These suggest either zone 9 or 10 of the Upper Ordovician.

Graptolites obtained from Locality 12 at Badgery’s Crossing have been identified by
Miss Elles, who says they belong to one species only, a new species of Glyptograptus,
which is described later. Glyptograptidae are not recorded from Zones 12, 13 and 14 of
the Upper Ordovician, but are known from zones below and zones above these. Miss
Elles, referring to the Glyptograptus from Locality 12, has expressed the view: “I can
only say that it seems closer to the Silurian types, but it would not be safe to bank on it.”

The quartzite at Locality 12, from which this Glyptograptus was obtained, occurs on
the south-western limb of a steeply folded anticline, which has every appearance of
being conformable to the slates containing undoubted Upper Ordovician graptolites.
The Silurian rocks found in the south-west of the area have, however, been regarded
as unconformable to the Ordovician (Woolnough, 1909).

The implication from present information is, therefore, that Locality 12 belongs to
an uppermost Ordovician zone.

BY

KATHLEEN SHERRARD.

Icm.

Text-figure 1.—Dicellograptus caduceus Lapworth. Proximal portion, Loc. 9, No. S.1645.

(Drawings made of specimens examined through a camera lucida. Where
not otherwise stated, specimens are in the collection of the author.)

HH

65

66 GRAPTOLITES FROM TALLONG AND THE SHOALHAVEN GORGE, N.S.W.,

DESCRIPTIONS OF GRAPTOLITES.

Family DICRANOGRAPTIDAE Lapworth.
Genus DICELLOGRAPTUS Hopkinson.

DICELLOGRAPTUS CADUCEUS Lapworth (Text-fig. 1; Plate i, fig. 1).

Dicellograptus caduceus Lapworth, Elles and Wood, 1904, p. 161.

Stipes 9 ecm. long, enclosing an axial angle of 330 degrees and crossing about 1 cm.
from proximal end. Stipes slightly less than 1 mm. wide and frequently twisted. Sicula
with very long virgella (4 mm.). No lateral spines. Thecae 12 in 10 mm., nearly 2 mm.
long with curved ventral margin, aperture introverted and sac-like, overlap one-half.
Stipes show graceful double curvature after first crossing but do not cross again.

Associates: Plegmatograptus nebula, Diplograptus truncatus var. pauperatus,
Dicellograptus angulatus. .

DICELLOGRAPTUS cf. SEXTANS Hall (Text-fig. 2, a and D).

cf. Dicellograptus sextans Hall, Elles and Wood, 1904, p. 1538.

Stipes 4 mm. long and 0:6 mm. wide. Axial angle 300-325 degrees. Axil pointed.
Thecae 13-15 in 10 mm. Thecae 1:3-1:5 mm. long, overlap slight, ventral margins curved,
apertures sometimes introverted, through pressure, sometimes rolled out into long
mucros. Sicula faintly visible in axil. The stipes of some of these very small dicello-
graptids show what look like thecae on both sides. This appearance is too frequent to
be due to two minute diplograptids having come to rest in such a way as to make a
similar angle between them. Ruedemann (1908, p. 309) figures a similar braided appear-
ance in his Dicellograptus sextans var. tortus. Dicellograptus cf. sextans occurs in the
Shoalhaven Gorge only in a phyllite. Hence the graptolites have been subjected to much
pressure and the stipes are probably twisted until they are seen in scalariform view.

Associate: cf. Amplexograptus sp.

DICELLOGRAPTUS ANGULATUS Elles and Wood (Text-fig. 3, @ and b).

Dicetlograptus angulatus Elles and Wood, Elles and Wood, 1904, p. 149.

Stipes, greatest length seen 1:3 cm., shortest 3 mm. Generally straight. Diverging
at angles 260-340 degrees. Width 0-4-0:8 mm. Axil square, delicate lateral spines.
Thecae 11-10 in 10 mm., 1-5 mm. long, overlap one-quarter, sometimes introverted and
sometimes everted. Ventral margins slightly curved. Sometimes several of this species
occur together, but generally this is not the case.

Associates: Cryptograptus tricornis, Orthograptus truncatus var. pauperatus, Dicello-
graptus caduceus.

DICELLOGRAPTUS Cf. FORCHAMMERI Geinitz .(Text-fig. 4; Plate i, fig. 2).

cf. Dicellograptus forchammeri Geinitz, Elles and Wood, 1904, p. 150.

A very well preserved proximal portion of a Dicellograptus from Locality 4 is
difficult to place specifically. The axial angle is 305 degrees, the stipes are straight and
2 cm. of each is preserved. The stipes increase in width from 0-6 mm. to just over 1 mm.
Th. 1+ and th. 1° are inclined downward, their ventral margins making an angle of
90 degrees, and are produced in spines 3 mm. long. The virgella is very short. The
sicula is a large blunt knob. The second thecae on each side grow upward and not
horizontally. Spines are visible on all the proximal thecae and seem to-appear just
below the aperture, which is said to be characteristic of D. forchammeri (Hlles and
Wood, 1904, p. 137). The thecae are 10 in 10 mm. and seem not more than 1:3 mm.
long with scarcely one-third overlap. The curvature of the ventral margins is very
slight and the apertures almost everted. The excavation is triangular, occupying one-
half the stipe. The lateral spines‘on this form suggest D. elegans, but there is no double
curvature of the stipes nor are the ventral margins of the thecae sufficiently curved for
this species. The distal portion of the stipes is covered with a mass of stipes of frag-
ments of other graptolites.

Associates: Dicellograptus angulatus, Climacograptus tubuliferus, Glossograptus
hincksu.

BY KATHLEEN SHERRARD. 67

DICELLOGRAPTUS FORCHAMMERI Var. FLEXUOSUS Lapworth (Text-fig. 5, @ and 0).
Dicellograptus forchammeri var. flecuosus Lapworth, Elles and Wood, 1904, p. 152.

Stipes up to 5 cm. long. Width of stipes 0-4 mm., axial angle 260-305 degrees. Thecae
12-10 in 10 mm., each 1:5-1:0 mm. long, overlap one-third, aperture introverted, excava-
tion pocket-like, occupying half width of stipe proximally, apertural spines seen on many
thecae in some specimens. Fine lateral spines, 1 mm. long. Pronounced sicula.

This form suggests a Leptograptus, but lateral spines are not found there. The
thecae are not of the Leptograptid shape and they are more crowded than is the case in
that genus.

Associates: Orthograptus calcaratus var. basilicus, O. truncatus var. pauperatus,
Climacograptus bicornis.

Genus DICRANOGRAPTUS Hall.
DICRANOGRAPTUS NICHOLSONI Hopkinson (Text-figs. 6, @ and b; PI. i, figs. 3, 4).

Dicranograptus nicholsoni Hopkinson, Elles and Wood, 1904, p. 171.

Biserial section 5-7 mm. long, increasing in width from 1 to 2 mm. Spines on 4
proximal thecae on each side. Spines up to 1:3 mm. long. Slender virgella under 1 mm.
long. Axial angle 40-60 degrees. Biserial thecae slightly over 1 mm. long with one-
fourth overlap. Septum fairly well marked. Uniserial thecae 14-10 in 10 mm., 2 mm.
long, overlap about one-half, curved ventral margin with introverted and introtorted
apertures which practically fill excavations. No spines on uniserial thecae, thus differen-
tiating this form from Dicranograptus furcatus. The length of the uniserial stipes varies
considerably, it is often only 3 cm. but has been found up to 6 cm. Long stipes curve
back towards one another as recorded by Ruedemann (1908) of D. nicholsoni var. diapson.
In one specimen from Locality 12 the uniserial stipes meet at 6 cm. (Text-fig. 6, a;
Plate i, fig. 3).

Associates: Climacograptus bicornis, C. peltifer, Cryptograptus tricornis.

DICRANOGRAPTUS FURCATUS var. MINIMUS Lapworth (Text-fig. 7).

Dicranograptus furcatus var. minimus Lapworth, Elles and Wood, 1904, p. 179.

At Locality 5 the most common fossil is a Dicranograptus with biserial portion
uniformly 3 mm. long and 0-5—0:7 mm. wide proximally and up to 1-5 mm. wide at the
point of bifurcation. There are four thecae on each side of the biserial stipe and half
of these have spines up to 1 mm. long. The virgella is never distinct and the presence
of a septum is doubtful. The axial angle varies from 35-60 degrees. The uniserial stipes
are 1 mm. wide and never more than 2 cm. long and have only a few spines. After
diverging at the point of bifurcation, they curve back until they are more or less
parallel and resemble D. nicholsoni var. diapson Gurley, but the biserial portion in that
form is longer (up to 6 mm.).

DICRANOGRAPTUS cf. CoONTORTUS Ruedemann (Text-figs. 8 and 9; Plate i, fig. 5).

ef. Dicranograptus contortus Ruedemann, Ruedemann, 1908, p. 337.

ef. Dicranograptus contortus Ruedemann, Ruedemann, 1947, p. 389.

Biserial portion 2-4 mm. long and 1-2-1:8 mm. wide, sometimes with septum well
marked. Up to five thecae on each side, each nearly 2 mm. long, overlapping one-third
with curved ventral margin. Uniserial stipes diverge at angles up to 100 degrees from
sharp V, then either curve strongly concave upward, or one stipe may be concave and
the other convex. Uniserial stipes are 0-8-1 mm. wide and 1-3 cm. long. Thecae up to
12 in 10 mm., about 1-5 mm. long, with strongly curved ventral edge, slight overlap and
deep excavation. Apertures sometimes introverted. All thecae appear non-spinose.

Associate: Dicranograptus furcatus var. minimus.

Family DreLoGrRAprTipar Lapworth.
Genus CLIMACOGRAPTUS Hall.
CLIMACOGRAPTUS BICORNIS (Hall) (Text-fig. 10, a@ and Db).
Climacograptus bicornis (Hall), Elles and Wood, 1906, p. 193.

68

GRAPTOLITES FROM TALLONG AND THE SHOALHAVEN GORGE, N.S.W.,

2c \, s\

1 ty i
ns) P !

Db i
il

t

!

f

-——

Depa Neem ~ nw ew

]

Oe a nn oY al CO oe ne

O Icm.

Text-figures 2-9.
Dicellograptus cf. sextans Hall, Loc. 10, collected Messrs. G. Packham and J. Veevers.

Pay Oly (Oe
Dicellograptus angulatus Elles and Wood, Loc. 5, No. 8.1739.

ah Gh

3, b. Dicellograptus angulatus Elles and Wood, Loc. 4, No. 8.1771.

4. Dicellograptus cf. forchammeri Geinitz, Loc. 4, No. 8.1761.

5 Dicellograptus forchammeri var. flexwosus Lapworth, Loc. 4, No. 8.1748.

5 Dicellograptus forchammeri var. flexuosus Lapworth, Loc. 11, collected by Messrs.
Ingall, Joklik and Scott-Orr.

6, a. Dicranograptus nicholsoni Hopkinson, Loc. 11, No. 8.1657.

6, b. Dicranograptus nicholsoni Hopkinson, Loc. 11, No. S.1678.

> .

5 w

BY KATHLEEN SHERRARD. 69

C. bicornis and its former varieties now raised to specific rank, are found in profusion
at Loc. 11. The development of the appendages in the former varieties which Ruedemann
has studied and figured (1908, p. 80, Plate A) can be paralleled and surpassed at this
place, where the length of the virgella in Cl. tridentatus and the size of the wings on the
shield in Cl. peltifer greatly exceed anything shown in Ruedemann’s plate.

In C. bicornis itself, the rhabdosome is 2:0—2:5 ecm. long, 1 mm. wide proximally and
2 mm. wide distally. Thecae 12 in 10 mm., about 15 mm. long and overlapping about
one-third. Apertural excavations one-quarter width of rhabdosome. The spines on the
basal thecae vary from 3 to 6 mm. in length and may be thin downward curving threads
or stout structures protruding at an angle of 120 degrees.

Associates: Climacograptus tridentatus, Cl. peltifer, Cryptograptus tricornis.

CLIMACOGRAPTUS PELTIFER (Lapworth) (Text-fig. 11, @ and 0b; Plate ii, figs. 11 and 12).

Climacograptus bicornis var. peltifer Lapworth, Elles and Wood, 1906, p. 195.

Climacograptus peltifer (Lapworth), Elles, 1925, p. 340.

Rhabdosome 5 cm. long. Width proximally is not more than 0-7 mm., widens rapidly
to 2-5 mm. in 2 cm. Thecae 11 in 10 mm., about 2 mm. long, overlapping nearly one-
half, with small, square apertures occupying no more than one-sixth width of rhabdosome.
The basal spines are imbedded in a thick crescent shaped membrane nearly 2 mm. wide
normally. The virgula extends beyond the distal end of the rhabdosome. In some
specimens the crescent is surrounded and obliterated by an almost circular structure
(Text-fig. 11, b; Plate ii, fig. 12) measuring at its greatest extent 11 mm. long by 6 mm.
wide. The greater length is developed parallel to the length of the rhabdosome. It is not
unusual to see a small membrane surrounding the basal spines or bulging from portion
of them in this species, but the development now described far exceeds anything
previously seen. As mentioned above, it also exceeds anything figured by Ruedemann.
Cl. wilsoni is characterized by the development of a proximal vesicle of elliptical form.
However in that species the greater length is seen at right angles to the length of the
rhabdosome. In the specimens from the Shoalhaven Gorge the crescent shape is retained
to some extent in the vesicle, and since figure 10c, Plate xxvi, in Elles and Wood (1907)
shows a vesicle of the same shape though of a much smaller size, this supports
retaining these in Ol. peltifer. The virgella seems to protrude from below the membrane
in the Shoalhaven specimens.

Associates: as above.

CLIMACOGRAPTUS TRIDENTATUS (Lapworth): (Text-fig. 12, a and 0; Plate ii, fig. 13).
Climacograptus bicornis var. tridentatus Lapworth, Elles and Wood, 1906, p. 195.
Climacograptus tridentatus (Lapworth), Decker, 1935, p. 707.

Climacograptus tridentatus (Lapworth), Ruedemann, 1947, p. 439.

Rhabdosome not much more than 3 cm. long preserved, exclusive of the virgella,
which may add another centimetre. Width never more than 2:0 mm. There is a
prominent virgular tube commencing after the sixth theca and measuring 0-3 mm., where
the rhabdosome is broken distally. There are 11 thecae in 10 mm., each measuring
1-5 to 2 mm. long, and overlapping one-third. For half its length the ventral edge of
the theca is nearly parallel to the axis of the rhabdosome and then curves down towards
the proximal end. The aperture is generally concave upwards, sometimes showing a
denticle. The virgella is stout and may be slightly more than 1 cm. long. It is often
surrounded by a membrane for half its length. The spines on the basal thecae may be
5 mm. long and make with the virgella an angle of about 90 degrees. T. S. Hall (1902)
erected a species, Climacograptus hastata, to include forms with a long virgella. The
proximal extremity is also reminiscent of Diplograptus calcaratus. Miss Elles has kindly
examined examples from the Shoalhaven Gorge and identifies them as the species
tridentatus.

Associates: Orthograptus apiculatus, Climacograptus bicornis, Cl. peltifer.

7. Dicranograptus furcatus var. minimus Lapworth, Loc. 5, No. 8.1719.
8, 9. Dicranograptus cf. contortus Ruedemann, Loc. 11, collected Messrs. Ingall, Joklik,
Scott-Orr.

70 GRAPTOLITES FROM TALLONG AND THE SHOALHAVEN GORGE, N.S.W.,

4

=-—

O Icrn.

Text-figures 10-20.
10, a. Climacograptus bicornis (Hall), Loe. 11, No. 8.1683.
10, b. Climacograptus bicornis (Hall), Loe. 11, collected by Messrs. Ingall, Joklik, and Scott-Orr.
11, a. Climacograptus peltifer (Lapworth), Loc. 11, No. 8.1676.
11, b. Climacograptus peltifer (Lapworth), Loc. 11, collected by Messrs. Packham and Veevers.
12, a. Climacograptus tridentatus (Lapworth), Loc. 11, No. S$.1662.
12, b. Climacograptus tridentatus (Lapworth), Loc. 11, Coll. Messrs. Ingall, Joklik and Scott-Orr.
3. Climacograptus tubuliferus Lapworth, Loc. 4, No. 8.1754.
14. Climacograptus caudatus (Lapworth), Loe. 8, No. 8.1873.
15. Climacograptus minimus (Carruthers), Loc. 9, No. 8.1653.
16. Climacograptus brevis (Elles and Wood), Loe. 11, No. S.1657.

BY KATHLEEN STERRARD. 71

CLIMACOGRAPTUS TUBULIFERUS Lapworth (Text-fig. 13).

Climacograptus tubuliferus Lapworth, Elles and Wood, 1906, p. 203, Pl. xxvii,
fig. 8, a—d.

Rhabdosome 2 cm. long and 2 mm. wide attained gradually from 0-5 mm. proximally.
Thecae 12 in 10 mm., 2-5 mm. long, free ventral edge slightly inclined. Thecae closely
overlap so that excavations not more than one-sixth of width of rhabdosome. Virgella
1 mm. long, virgula 0:2 mm. wide and prolonged 2 cm. beyond rhabdosome and visible
in body. The specimens are short and narrow for Cl. tubuliferus, suggesting Cl.
minimus, but the virgula is strongly developed and the number of thecae in 10 mm.
agrees with Cl. tubuliferus.

Associates: Dicellograptus angulatus, Orthograptus calcaratus var. basilicus.

CLIMACOGRAPTUS MINIMUS (Carruthers) (Text-fig. 15).

Climacograptus minimus (Carruthers), Elles and Wood, 1906, p. 191, Pl. xxvii,
fig. 1, a-g.

Rhabdosome 7 mm. long, nearly 2 mm. wide distally, increasing rapidly from width
of 0-7 mm. proximally. Thecae alternate, 14 in 10 mm. with slightly curved free edges,
overlap slight, perhaps one-quarter. Apertural margin horizontal or everted, opening
within triangular excavation one-quarter width of rhabdosome. Virgella 0-7 mm. long,
virgula, thread-like, prolonged 1:5 cm.

Associates: Dicellograptus angulatus, Retiolites nebula.

CLIMACOGRAPTUS CAUDATUS Lapworth (Text-fig. 14).

Climacograptus caudatus Lapworth, Elles and Wood, 1906, p. 202.

Rhabdosome 1—2 cm. long, 2 mm. greatest width, gradually increasing from 0:7 mm.
Hair-like virgella prolonged 1:5 cm. surrounded by membrane for about 2 mm. near
rhabdosome. Thecae 11 in 10 mm., 1-5 mm. long, overlap one-third, ventral wall slightly
sloping, aperture opening within semi-circular excavation, taking up one-sixth of rhabdo-
some. Virgula prolonged up to 1 cm.

Associates: Orthograptus calcaratus var. basilicus, Climacograptus tubuliferus.

CLIMACOGRAPTUS BREVIS Elles and Wood (Text-fig. 16).

Climacograptus brevis Elles and Wood, Elles and Wood, 1906, p. 192.

Rhabdosome 5 mm. long, gradually increasing in width to maximum of 1 mm.
Thread-like virgula prolonged at least 3 mm. Thecae 15 in 10 mm., about 1 mm. long,
overlap one-third, with slightly curved ventral edge, short virgella, sicula indistinguish-
able. The differences between C. brevis and C. minimus are slight.

Associates: Lasiograptus harknessi.

Genus DrpLograptus McCoy.
Sub-Genus oRTHOGRAPTUS Lapworth.
DIPLOGRAPTUS (ORTHOGRAPTUS) Cf. QUADRIMUCRONATUS Var. SPINIGERUS Lapworth
(Text-fig. 17).

cf. Diplograptus (Orthograptus) quadrimucronatus var. spinigerus Lapw., Hlles
and Wood, 1907, p. 225.

Spined, biserial graptolite, the best specimen of which is unfortunately broken and
the distal end lost. A poorer specimen is not more than 2 cm. long in all. In the best
preserved form a length of 143 cm. can be seen with sicula and proximal thecae in
excellent preservation. Thecae are set 12 to 10 in 10 mm. They are 1:5 mm. long and
overlap one-half. Width of rhabdosome at sicular end is 0-6 mm. It increases rapidly
to 2-5 mm. and this width is retained. All thecae are furnished with apertural spines
and there is a short virgella. The apertural spines are rising in a distal direction as they
leave the thecae but soon become horizontal. They increase in length from 0:5 mm. on

17. Diplograptus cf. quadrimucronatus var. spinigerus (Lapworth), Loe. 11, coll. Messrs. Ingall,
Joklik, Scott-Orr.

18. Diplograptus truncatus var. pauperatus (Elles and Wood), Loc. 9, No. 8.1648.

19. Diplograptus calcaratus (Lapworth), Loc. 11, No. 8.1662.

20. Diplograptus calcaratus var. basilicus (Lapworth), Loc. 3, No. 8.1826.

72 GRAPTOLITES FROM TALLONG AND THE SHOALHAVEN GORGE, N.S.W.,

the first theca to 1 mm. on the 6th, while the 8th and 9th thecae on each side have spines
6 mm. in length, but spines on succeeding thecae are not more than 1 mm: long. The
spines on the 8th and 9th thecae are also broader than the others. The rhabdosome is
preserved as a clathria in the form of a hexagonal network with no exochitin remaining
between the crossbars. The dimensions are small for D. (0.) quadrimucronatus var.
spinigerus and the associated graptolites belong to a lower horizon than that which it
characterizes. Dr. T. S. Hall has described from near Mt. Haston, Victoria (Hall,
1906), a graptolite with four long spines on “about the 7th or 8th thecae” which he
referred to Diplograptus quadrimucronatus. D. quadrimucronatus var. spinigerus is
also recorded (Thomas and Keble, 1933) from a locality on Emu or Bolinda Creek,
Victoria, which was a collecting place of McCoy.

Dr. Ruedemann (1908) excludes all spinose forms from the genus Diplograptus
because he considers with the development of spinosity changes take place which are
of generic value. He points out further “that the spinose forms of ‘Diplograptus’ possess
as a rule a layer of retiolid meshes ...and that the development of this layer of meshes
and ledges is roughly proportional to that of the spines”. He places spinose forms
among the Glossograptidae. Elles and Wood (1907) state briefly the resemblance of
O. quadrimucronatus to the Glossograptidae but leave it among the Diplograptidae.

Dr. Ruedemann quotes Glossograptus quadrimucronatus in a discussion on spines,
copying a figure of Lapworth’s showing a graptolite from Ireland with two long spines
on the “fourth or fifth theca’. Ruedemann points out that spines in this position cannot
have been developed to protect the graptolite but “they express ... an intrinsic
tendency to a repetition of the lateral apertural spines’’.

O. ef. quadrimucronatus var. spinigerus is rare at Loc. 11 but found more commonly,
though in very bad preservation, at Loc. 5.

Associates: Olimacograptus peltifer, Cryptograptus tricornis, Dicranograptus
furcatus var. minimus.

DIPLOGRAPTUS (ORTHOGRAPTUS) TRUNCATUS Var. PAUPERATUS E. & W. (Text-fig. 18).

Diplograptus (Orthograptus) truncatus var. pauperatus EB. & W., Elles and Wood,
UNOS 1s BB.

Longest specimens nearly 4 cm., though most about 2 cm. Width 2 mm. Thecae
alternate, 13 in 10 mm., up to 2-5 mm. long, overlap one-half. Aperture horizontal to
everted. Ventral edge slightly curved. No virgula or virgular tube. The longest of the
forms included in this species suggest O. truncatus var. intermedius, but as 4 cm. is the
maximum length, probably all are O. truncatus var. pauperatus.

Associates: Dicellograptus caduceus, D. angulatus.

DIPLOGRAPTUS (ORTHOGRAPTUS) CALCARATUS Lapworth (Text-fig. 19; Plate ii, fig. 6).

Diplograptus (Orthograptus) calcaratus Lapw., Elles and Wood, 1907, p. 239.

Rhabdosome often in fragments, which may be 7 cm. long, though incomplete.
Complete examples 3 cm. long. Width 2-3 mm. proximally, increasing to 3-5 distally.
Thecae 11 in 16 mm., up to 3 mm. long and 0-4 mm. wide. Overlap one-half, inclined
30 degrees. Ventral walls generally convex outwards and apertures are horizontal,
everted or concave. Conspicuous virgular tube within rhabdosome throughout its
length, 0-6 mm. wide. Virgula continued more than 5 mm. beyond rhabdosome.
Proximally shows well-developed basal spines, 2-3 mm. long and virgella up to 8 mm.
Virgella sometimes appears distorted and split but this may be due to the covering
membrane breaking away.

Associates: Climacograptus tridentatus, Orthograptus apiculatus.

DIPLOGRAPTUS (ORTHOGRAPTUS) CALCARATUS Var. BASILICUS Lapworth (Text-fig. 20).

Diplograptus (Orthograptus) calcaratus var. basilicus Lapworth, Elles and Wood,
1907, p. 243.

Rhabdosome over 4 cm. long, 2-5 mm. wide, with short basal spines and virgella.
’Thecae 11 in 10 mm. Thecae 2 mm. long, overlap one-half or more. A suggestion of
spines on some thecae. Central virgular tube not always distinct.

BY KATHLEEN SHERRARD. 73

Associates: Dicellograptus angulatus, D. forchammeri var. flexuosus, Climacograptus
tubuliferus, Cl. minimus.

DIPLOGRAPTUS (ORTHOGRAPTUS) APICULATUS (EH. & W.) (Text-fig. 21, a, b, c; Plate ii, fig. 7).

Diplograptus (Orthograptus) rugosus var. apiculatus BE. & W., Elles and Wood, 1907,
p. 245.

Diplograptus (Orthograptus) apiculatus (Elles & Wood) Bulman, 1946, p. 51.

The rhabdosome is generally short, less than 2 cm. long, though broken fragments
of more than 4 cm. have been obtained. Its greatest width is slightly more than 3 mm.,
and is attained rapidly from an initial width of 1 mm. It narrows slightly distally.
The thecae number 11 in 10 mm. and are 2 mm. long overlapping one-half. The apertural
margin is usually everted and produced into a denticle. A short spine, 0-5 mm. long,
appears on the two basal thecae. A short, stout virgella is about the same length usually.
Excavations are pronounced. The thread-like virgula may be produced for at least 2 cm.
beyond the rhabdosome.

Associates: Climacograptus tridentatus, Orthograptus calcaratus.

Sub-Genus GLYPTOGRAPTUS Lapworth.
DIPLOGRAPTUS (GLYPTOGRAPTUS) ROSTRATUS sp. nov. (Text-fig. 22, a, b; Plate ii, figs. 9, 10).

Rhabdosome 4 cm. long, width proximally 1:5 mm., widening gradually to 3 mm.
Stout blunt sicula with no virgella preserved. Short curving basal spines. Thecae
alternate, 12-10 in 10 mm. Each theca about 2 mm. long and overlapping one-third to
one-quarter. Excavations wide and triangular. Ventral margins of thecae first convex
then concave. Apertural margin convex, produced to a denticle which forms a beak
when it meets the ventral edge, hence the specific name. Septal groove well marked.
Thecal margins strongly marked. Indistinct virgula. Specimens preserved in relief.
Moulds of thecae seen as comparatively deep pockets which deepen towards the apertural
denticle.

This species is found in one locality only, that is Locality 12, close to Badgery’s
Crossing of the Shoalhaven River (Joplin, 1945). Graptolites and phyllocarid carapaces
are preserved in black in low relief in a dark-grey coarse-grained quartzite. There is a
noticeable parallelism in the arrangement of the rhabdosomes. The majority are not
preserved in the bi-profile aspect but are seen in three-quarter face, half face and quarter
face. In some cases they almost look mono-serial. On one slab of 17 graptolite frag-
ments, only one when examined in the field with a hand lens appeared to show thecae
on both sides.

Miss G. L. Elles was good enough to examine some of these specimens which were
sent to her at the Sedgwick Museum, Cambridge, and she has stated:

“The specimens from Badgery’s Crossing are, I think, all different views of the
same graptolite which is certainly a Glyptograptus, but seems to be a new form, or rather
one unknown to me. The thecae are of the advanced type of Glyptograptus and recall
G. serratus but it is not serratus. All the Glyptograptids I know from early Ordovician
rocks have a simpler type of cell. I can only say that it seems closer to the Silurian
types but it would not be safe to bank on it. You have it in all sorts of aspects, half
face, quarter face, three-quarter face and scalariform views and those in (or nearly)
profile view are not quite like any known to me.”

Sub-Genus “mMESOGRAPTUS” Elles and Wood (Bulman, 1929).
Group II. AMPLEXOGRAPTUS.
cf. DIPLOGRAPTUS (AMPLEXOGRAPTUS) sp. indet. (Text-fig. 23).

Fragments up to 3 mm. long and 1 mm. wide. Well-marked septum, slightly
undulating. Thecae 16 in 10 mm., alternate, 1 mm. long, overlap one-third, with vertical
ventral edge and horizontal aperture. Excavation strongly marked, taking up one-third
width of rhabdosome.

Associate: Dicellograptus cf. sextans.

74

GRAPTOLITES FROM TALLONG AND THE SHOALHAVEN GORGE, N.S.W.,

2

| Z|

SPIT DUVIING SV QM
Cx

S-.
~My

TMA
ON
pvt tI
WAS

Text-figures 21-32.
a. Diplograptus apiculatus (Elles and Wood), Loc. 11, No. 8.1662.
b. Diplograptus apiculatus (EH. & W.), Loc. 11, coll. by Messrs. Ingall, Joklik, Scott-Orr.
c. Diplograptus apiculatus (EH. & W.), Loc. 4, No. S.1762.
a. Glyptograptus rostratus sp. nov., Loc. 12, No. 8.1926.
b. Glyptograptus rostratus sp. nov., Loc. 12, Sydney University Geological Department
Collection, Reg. No. 8289.
cf. Amplexograptus, sp. indet., Loc. 10, coll. Messrs. Packham and Veevers.

. a. Cryptograptus tricornis (Carruthers), Loc. 11, No. 8.1683.

BY KATHLEEN SILTERRARD. 75

Family CryproGRAPTIwDAr Hadding, restr. Bulman.
Genus CRYPTOGRAPTUS Lapworth.
CRYPTOGRAPTUS TRICORNIS (Carruthers). (Text-fig. 24, a and Db.)

Cryptograptus tricornis (Carruthers), Elles and Wood, 1908, p. 296.

Rhabdosome 0-5 mm. to 2 cm. long and slightly more than 1 mm. wide, sometimes
wider proximally than distally. Thecae 12 in 10 mm., of Climacograptus type and 1:5 mm.
long, overlapping for one-half their extent. Apertures sometimes inclined and sometimes
semicircular when they occupy one-fourth width of rhabdosome. Basal spines very
conspicuous up to 2 mm. long, may be drooping or stiff and at right angles to direction
of rhabdosome. Sicular spine wide and noticeable, up to 0:'5 mm. long. Faint thread-
like virgula prolonged beyond rhabdosome for 2 mm. and visible through test of rhabdo-
some from about third theca. Sicular ring bearing four spines very noticeable in some
specimens. (See Text-fig. 24, b.) As Ruedemann points out, the specific name is a
misnomer.

Associates: Climacograptus bicornis, C. peltifer, C. tridentatus.

Family GLossoGRAPTIDAE Lapworth.
Genus GLOSSOGRAPTUS Emmons.
GLOSSOGRAPTUS HINCKSII (Hopkinson). (Text-fig. 25; Plate ii, fig. 8.)

Glossograptus hincksii (Hopkinson), Elles and Wood, 1908, p. 309.

Well-preserved specimens of this graptolite are incomplete. Rhabdosome 2 cm. long
but broken at both ends. It maintains a uniform width of 1:8 em. Thecae of diplo-
graptid form and about 8 in 10 mm. Some thecae with aperture pressed into a denticle
only, but usually prolonged to a spine of 1 mm. length. Septal spines are very
conspicuous, and are stout and stiff and 1 mm. long. The impression is clearly that of
a form with a rounded or hexagonal cross-section. This is shown because of the
grouping of the spines which are more closely preserved in some places than others, on
account of the chance as to which side the spines behind the rhabdosome have been
bent over and preserved. For this reason the number of thecae per millimetre is difficult
to determine. The best preserved impressions are sub-scalariform and therefore the

width is not as great as in a bi-profile view. Septal threads are occasionally seen.
Associate: Cryptograptus tricornis.

Genus LASIOGRAPTUS Lapworth s. str.

LASIOGRAPTUS HARKNESSI (Nicholson). (Text-fig. 27 a, b, c; Plate ii, fig. 14.)
Lasiograptus (Thysanograptus) harknessi (Nicholson), Elles and Wood, 1908, p. 325.
Lasiograptus harknessi (Nicholson), Bulman, 1947, p. 71.

Rhabdosome up to 8 mm. long and 2 mm. wide, exclusive of spines. Width at
proximal end 0-5 mm. Three spines, possibly paired, seen at proximal end in some
specimens. In one example, what appear to be a pair of basal spines 4 mm. long and a
virgella (2-5 mm.) are preserved immediately below the rhabdosome, but with no
connecting test retained (see Text-fig. 27, c). Thecae 15-16 in 10 mm., from 1-2 mm.
long, overlapping one-half. They are diplograptid in shape with everted apertures.
Apertural spines up to 1 mm. long on all thecae which anastomose to form the marginal
network, which, however, is never entirely complete. The size of the rhabdosome allies
this with L. harknessi, though the number of thecae per millimetre, which can be
measured quite exactly in some cases, are those found in the variety costatus.

Associates: Orthograptus calcaratus, Corynoides calicularis.

24, b. Cryptograptus tricornis (Carruthers), Loc. 11, No. 8.1659.

25. Glossograptus hincksii (Hopkinson), Loc. 11, coll. Messrs. Ingall, Joklik, Scott-Orr.
26. Lasiograptus cf. mucronatus (Hall), Loc. 7, No. 8.1895.

27, a. Lasiograptus harknessi (Nicholson), Loc. 11, coll. Messrs. Ingall, Joklik, Scott-Orr.
27, b. Lasiograptus harknessi (Nich), Loc. 8, No. 8.1857.

27, c. Lasiograptus harknessi (Nich), Loc. 8, No. 8.1865.

28. cf. Neurograptus margaritatus (Lapworth), Loc. 2, No. S.1777.

29. Neurograptus cf. fibratus (Lapworth), Loc. 8, No. 8.1862.

30. Plegmatograptus nebula Elles and Wood, Loc. 9, No. 8.1645.

31, a and b. cf. Mastigograptus, sp. indet., Loc. 11, No. S.1667.

32. Corynoides calicularis Nicholson, Loc. 11, No. S.1670.

76 GRAPTOLITES FROM TALLONG AND THE SHOALHAVEN GORGE, N.S.W.,

Sub-Genus HALLOGRAPTUS Carruthers MS.
LASIOGRAPTUS cf. (HALLOGRAPTUS) MUCRONATUS (Hall). (Text-fig. 26.)

cf. Lasiograptus (Hallograptus) mucronatus (Hall), Elles and Wood, 1908, p. 321.

Rhabdosome 2:5 cm. long and 2 mm. wide, limp and lax. Thecae 12 in 10 mm., 2 mm.
long, overlapping half, mesial spines usually preserved on ventral edge. Virgella short.
What appears to be a scopulate process is shown on one theca. It is 4 mm. long and
0-$ mm. wide and has a two-pronged end.

Associates: Climacograptus bicornis, Lasiograptus harknessi.

Sub-Genus NEUROGRAPTUS Lapworth.
LASIOGRAPTUS (NEUROGRAPTUS) cf. FIBRATUS (Lapworth). (Text-fig. 29.)

cf. Lasiograptus (Neurograptus) fibratus (Lapworth), Elles and Wood, 1908, p. 381.

Incomplete rhabdosomes, 23 em. long and 24 mm. wide. Test attenuated, especially
distally. Clathria strongly outlined. Thecae 11 in 10 mm., Diplograptus-like, apertural
margins of adjoining thecae connected by vertical threads. Short spines on the outside
of these.

Associates: Climacograptus tubuliferous, Lasiograptus harknessi.

cf. LASIOGRAPTUS (NEUROGRAPTUS) MARGARITATUS (Lapworth). (Text-fig. 28.)

ef. Lasiograptus (Neurograptus) margaritatus (Lapworth), Elles and Wood, 1908,
Pp. 332.

Limp, lax rhabdosome, 2 cm. long, and 1:5 mm. wide. Thecae 15 in 10 mm., of
Lasiograptus type, 2 mm. long, overlap nearly one-half. Lacinia incomplete, but spines
visible.

Associates: Diplograptus cf. calcaratus, Climacograptus minimus.

Family ReEtTIioLiripaAr Lapworth (restricted).
Sub-Genus PLEGMATOGRAPTUS Elles and Wood.
RETIOLITES (PLEGMATOGRAPTUS) NEBULA Elles and Wood. (Text-fig. 30.)

Retiolites (Plegmatograptus) nebula Elles and Wood, Elles and Wood, 1908, p. 340.

This form is preserved as a delicately traced silvery network, without a continuous
membranous test. The rhabdosome consists of a clathria of horizontal threads joined
together by vertical strands of which the obverse and reverse members are almost super-
imposed by compression. Attached to notches on the threads of the clathria is the
reticula, made up of 5- or 6-sided meshes the sides of which are about 0-2 mm. long and
exceedingly fine, distinctly finer than the bars of the clathria. In some cases a network
representing a lacinia is fairly complete. The meshes of the lacinia are slightly larger
than those of the reticula. The largest form is 11 mm. long and 5 mm. wide. In the
absence of a continuous test the shape of the thecae is difficult to dissociate from the
reticula. Apparently there are 13 thecae in 10 mm., each theca being about 1 mm. long
and overlapping one-third. The ventral wall is strongly inclined. The virgula is
prolonged 14 cm. beyond the rhabdosome and a short virgella (0-5 mm.) can be seen.

Retiolites (P.) nebula is the only recorded Upper Ordovician Retiolites. Dr. T. S.
Hall found Retiolites to be very common in the New South Wales Geological Survey
Collection from Stockyard Flat Creek, County of Wellesley, which he described (1902),
and he erected a new species, Retiolites caudatus, with a conspicuous nema and virgula.
Specimens of the species were 22 mm.. long and 6 mm. wide. However, Miss G. L. Elles,
in the correlation table which she drew up for Sir Edgeworth David’s Explanatory
Notes (1932), indicates that Ret. caudatus T. S. Hall is equivalent to Plegma. nebula
EK. & W. The diagnosis of the form records a maximum width of 2 mm., but it is noted
that “a larger and wider form is referred to this species. (It) may eventually be
separated off as a distinct variety”.

Order DENDROIDEA Nicholson.
Family DENDROGRAPTIDAE Roemer.
Genus MASTIGOGRAPTUS Ruedemann. (Text-fig. 31, a, D.)

ct. Mastigograptus sp. indet. Ruedemann, 1908, p. 210.

STHLERRARD.

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Map of the Tallong and Shoalhaven Gorge

3.—Geological Sketch

Text-figure 3

Geological boundaries after Woolnough, Craft and Regional Map issued by Premier’s

Department, N.S.W.

78 GRAPTOLITES FROM TALLONG AND THE SHOALHAVEN GORGE, N.S.W.,

At Locality 11 a large number of minute graptolite-like forms are found on some slabs
of slate. They lie crowded in a random arrangement about 30 to the square inch. They
can be divided into two types. One type consists of a tangled mass of short, straight
or slightly curved forms, sometimes branched, all incomplete. None is longer than
1-5 cm., nor wider than 0-3 mm. They lie massed on top of one another at all angles.
Since there are no specimens with wide stipes such as Ruedemann (1908) describes in
Thamnograptus where the main stipe by secondary thickening is found up to 1-5 mm.
wide, these fossils seem best compared to Mastigograptus sp. indet. Only one branch
is seen in any one specimen, and no triangular appendages are found attached to stipes
or branches as described by Ruedemann (1908), p. 212. However, he states that these
must have been easily detached and loose fragments on the slabs from Locality 11 may
represent them. Ruedemann records Mastigograptus from the Trenton and Utica
horizons, which are equivalent to Zones 9-12 of the British succession of the Upper
Ordovician. In Victoria, Mastigograptus has been recorded from the Cambrian.
(Chapman and Thomas, 1936.)

Family Corynompipar Bulman.
Genus coryNnomeEs Nicholson.
CORYNOIDES CALICULARIS Nicholson. (Text-fig. 32.)

Corynoides calicularis Nicholson, Ruedemann, 1908, p. 234.

Corynoides ef. calicularis Nicholson, Bulman, 1945, p. 27.

The second type of graptolite-like form from Locality 11 is smaller than cf. Mastigo-
graptus, not being more than 5 mm. long (excluding nema) and slightly less than 0-5 mm.
wide. This has been identified as Corynoides calicularis Nicholson. The rhabdosome is
suspended from a very slender nema (2 mm. long) which sometimes, however, is not
preserved. At the base of the rhabdosome- can be detected the aperture of the sicula,
with that of a theca on each side of it.

The structure of Corynoides has been recently considerably elucidated by Bulman
(1945-47) from material preserved in limestone, which could be dissolved away leaving
Corynoides which could then be examined under a microscope. Bulman found that the
sicula extends the full length of the rhabdosome from the slender nema to the base
where is the aperture of the sicula with the virgella projecting. Its-aperture is seen
as a concave structure at the base of the specimen. Thecae up to three in number bud
off in turn from alternate sides of the sicula and near its apical end and grow down
along the sicula until their apertures, too, open on either side of the sicular aperture.
As well as the virgella of the sicula, each theca has a virgella also, which projects beyond
the base, making an angle with the sicular virgeila. These were taken previously for
spines. These features can be recognized in Corynoides calicularis from the Shoalhaven
Gorge, where it is, however, preserved as thin, chitinous films on slate.

The systematic position of Corynoides has been a question of debate and even now
Bulman says (1945, p. 23), “I am leaving it provisionally as a distinct family of the
Graptoloidea’’, which suggests he foresees later revision.

Corynoides has apparently not been recorded from any other locality in Australia,
until identified from the Shoalhaven Gorge (Sherrard, 1947). It occurs fairly commonly
in North America and Europe. The fact that it had not been identified from South
America caused Bulman to comment in 1931 “the absence of Corynoides is remarkable,
as this constitutes such an important member of the Canajoharie fauna and of that of the
Dicranograptus clingani Zone of Northern Europe”. Its discovery in Australia provides
further evidence, if any were needed, of the similarity of graptolite faunas of the same
age throughout the world.

SUMMARY.

Upper Ordovician graptolites from Tallong and the Shoalhaven Gorge are described
and figured, one of them a new species. Corynoides calicularis, characteristic of North
European and North American graptolite-bearing beds, is recorded from Australia for
the first time.

Graptolites Collected from Various Localities at Tallong and the Shoalhaven Gorge.

BY KATHLEEN SHERRARD.

TABLE 1.

79

Recorded from.

Extra-Australian Localities. Australian.
Zone 9. | Zone 10.| Zone 11.] Zone 12.} Zone 13.
Nema. CH Cl. Dicer. Pl. Gis- Hast- | Bolin-
gracilis. | peltifer. | wilsoni. | clingani.| linearis. | POTNian.| onian. dian.
LOCALITY 1 contains :
Indeterminate graptolite fragments.
LOCALITY 2 contains:
Climacograptus sp. indet.
Cl. cf. minimus (Carr.) = x Xe x
Diplograptus (Orthograptus) cf.
calcaratus Lapworth as Aa Xs X xx xx x
Cf. Lasiograptus (Neurograptus)
margaritatus (Lapworth) .. Xs x x
LOCALITY 3 contains :
Dicellograptus forchammert var.
flexuosus Lapw. é ee xe xX
Dicranograptus furcatus var.
minimus Lapworth x Bx ax x
D. nicholsoni Hopk. Xe x: X x x x
Climacograptus bicornis (Hall) x x Xs x aX: x
Diplograptus (Orthograptus) cal-
caratus var. basilicus Lapw. x x x: x
Cryptograptus tricornis (Carr.) x x Xe x x x
Retiolites (Plegmatograptus) nebula
E. and W. x x x
LOCALITY 4 contains:
Dicellograptus Gis forchammert
Geinitz ae ae ny axe xe x Bs x x x
D. forchammeri var. flexuosus
Lapworth xe x
D. angulatus E. and W. 3 x x
Climacograptus tubuliferus Lap-
worth xX x x x:
Cl. bicornis (Hall) xe x BX x x x
Cl. minimus (Carruthers) ; xX x x
Diplograptus (Orthograptus) trun-
catus var. pauperatus K. and W. axe x xe x
D. (O.) calearatus var. basilicus
Lapw. Ba ae x xX Xe x
D. (O.) apiculatus (E. and W.) x
Glossograptus hincksii (Hopk.) Xs x x Xe Dx x
Retiolites (Plegmatograptus) nebula
E. and W. x x x
LOCALITY 5 contains :
Dicellograptus angulatus Elles and
Wood a aD cas Xe x
Dicranograptus nicholsoni Hopk. aXe x He x Xs x
D. furcatus var. minimus Lapworth x x x x
D. cf. contortus Ruedemann Bx
Climacograptus sp. indet.
Diplograptus © (Orthograptus) cf.
calcaratus Lapworth ae ae aX xe aXe Xs x
D. (O.) cf. quadrimucronatus var.
spinigerus Lapworth x x x
Glossograptus sp. indet.
Lasiograptus harknessi (Nich.) x x x
LOCALITY 6 contains :
Dicranograptus nicholsoni Hopk. x Ke Ke x x x
D. furcatus var. minimus Lapworth x x x x
D. cf. contortus Ruedemann Xs
Climacograp‘us sp. indet.
Diplograptus (Orthograptus) cf.
calcaratus Lapw. . . a a: x x x x Xe

80 GRAPTOLITES FROM TALLONG AND THE SHOALHAVEN GORGE, N.S.W.,

LOCALITY 7 contains:

Climacograptus bicornis (Hall)

Cl. brevis E. and W. 55 Pie

Diplograptus (Orthograptus) cf.
calcaratus Lapw. . . ag

Lasiograptus (Hallapranene) cf.
mucronatus (Hall)

L. harknessi (Nich.) ..

TABLE 1.—Continued.

LOCALITY 8 contains:
Dicellograptus angulatus E. and W.
Dicellograptus sp. indet.
Climacograptus tubuliferus Lap-

worth t a
Cl. caudatus Lapw.
Cl. minimus (Carr) . ie
Diplograptus (Onnograpie) cal-
caratus var. basilicus Lapw.
Cryptograptus tricornis (Carr)
Lasiograptus harknessi (Nich.)
L. (Neurograptus) cf. fibratus (Lapw.)

LOCALITY 9 contains :

Dicellograptus angulatus Elles and
Wood 3

D. caduceus eprosn

Climacograptus bicornis (Hall)

Cl tridentatus (Lapw.)

Cl. minimus (Carr.) é ae

Diplograptus (Onnogranhe) trun-
catus var. pauperatus Elles and
Wood ae os x

D. (O.) calearatus var. basilicus
Lapw.

Cryptograptus TA CONIVILE (Came)

Retiolites (Plegmatograptus) nebula

Elles and Wood

LOCALITY 10 contains:
Dicellograptus cf. sextans Hall
Cf. Amplexograptus sp. indet.

LOCALITY 11 contains :
Dicellograptus forchammeri var.
flexuosus (Lapw.)

Cf. D. caduceus Lapworth
Dicranograptus nicholsoni Hopk. ..
D. cf. contortus Ruedemann
Climacograptus bicornis (Hall)

Cl. tridentatus (Lapw.)

Cl. peltifer (Lapw.) :

Cl. brevis Elles and Wood! é
Diplograptus (Orthograptus) cal-

caratus Lapw. oo
ID, (5) Pair Crone var.

spinigerus Lapw.

D. (9.) apiculatus (EB. and Ww.)
Cryptograptus tricornis (Carr.)

Glossograptus hincksii (Hopk.)
Lasiograptus harknessi (Nich.)

Corynoides calicularis Nich.

Cf. Mastigograptus Ruedemann

sp. indet.

7]

nw al

nw

nAnA WK

iv)

al

nnn wK

al

LOCALITY 12 contains:
Diplograptus (Glyptograptus) ros-
tratus sp. NOV.
Phy!locarid carapaces.

BY KATHLEEN SITERRARD. 81

ACKNOWLEDGEMENTS.

Professor L. A. Cotton, M.A., D.Sc., has been good enough to allow me facilities for work
in the Geological Department of the University of Sydney. Miss G. L. Elles has added
immeasurably to the value of this work by readily agreeing to examine some specimens sent
to her at the Sedgwick Museum. Dr. Ida Brown, of the University of Sydney, has generously
helped me with advice and has given liberally of her time and skill in photographing graptolite
specimens for me. Miss F. M. Quodling, B.Sc., of the University of Sydney, and my’ son,
J. M. Sherrard, have kindly accompanied me at various times to Tallong and the Shoalhaven
Gorge. Messrs. Ingall, Joklik, Packham, Scott-Orr and Veevers kindly lent for description
specimens collected by them.

References.

BuLMAN, O. M. B., 1929.—Genotypes of the Genera of Graptolites. Ann. Mag. Nat. Hist.,
10:4, 169-185.
————. 1931.—South American Graptolites. Arkiv. for Zoologi, 22A: 3, 111.
, 1945-47.—Caradoec (Balclatchie) Graptolites from MLimestones in Laggan Burn,
Ayrshire. Mon. Pal. Soc., xcviii-c, pp. i-xi, 1-78.
CARNE, J. E., 1911.—Tin Mining Industry in New South Wales. Geol. Surv. N.S.W. Min. Reés.,
14: 346.
CHAPMAN, F., and THomaAs, D. E., 1936.—-Cambrian Hydroidea of the Heathcote and Monegeeta
District. Proc. Roy. Soc. Vict. (n.s.), xlviii: 2, 193-212.
CraFrt, F. A., 1931.—Physiography of the Shoalhaven River Valley, i. Proc. LINN. Soc. N.S.W.,
Ivi: 99-132.
Davip, T. W. E., 1932.—Explanatory Notes to Accompany a New Geological Map of the
Commonwealth of Australia. S8vo. Sydney.
DeEcKER, C. E., 1935.—Graptolites of the Sylvan Shale of Oklahoma and Polk Creek Shale of
Arkansas. Journ. Pal., 9: 697-708.
Ewues, G. L., 1925.—Characteristic Assemblages of Graptolite Zones of the British Isles. Geol.
Mag., \xii: 337-347.
— and Woop, EK. M. R., 1903-12.—British Graptolites. Mon. Pal. Soc., lvii-lxiv.
Hau, T. S., 1902.—Graptolites of New South Wales. Rec. Geol. Surv. N.S.W., vii: 49-59.
, 1906.—Reports on Graptolites. Rec. Geol. Surv. Vict., i: 4, 266-278.
—_—— , 1909.—On a Collection of Graptolites from Tallong, New South Wales. Rec. Geol.
Surv. N.S.W., viii: 4, 339-341.
, 1920.—On a Further Collection of Graptolites from Tolwong, New South Wales. Ree.
Geol. Surv. N.S.W., ix: 2, 63-66.
Harris, W. J., and THomaAs, D. E., 1938.—A revised Classification of the Ordovician Graptolite
Beds of Victoria. Min. and Geol. Journ. Vict., 1:3, 62-72.
JOPLIN, G. A., 1945.—Petrological Studies in the Ordovician of New South Wales. iii. Proc.
LINN. Soc. N.S.W., Ixx, 158-172.
KEBLE, R. A., and BENSON, W. N., 1939.—Graptolites of Australia. Mem. Nat. Mus. Melb., 11:
11-99.
Naytor, G. F. K., 1935.—Note on the Geology of the Goulburn District. Journ. Roy. Soc. N.S.W.,
69: 75-85.
RUEDEMANN, R., 1908.—Graptolites of New York. New York State Musewm, Mem. 11: 2-488.
, 1947.—Graptolites of North America. Geol. Soc. Amer. Mem. 19: 1-652.
SHERRARD, K., 1947.—Exhibit to Geological Section, Royal Society. Journ. Roy. Soc. N.S.W..,
SxeReXCI Sy GREXTVe
THomaAs, D. E., and Kesey, R. A., 1933.—Ordovician and Silurian Rocks of the Bulla-Sunbury
Area. Proc. Roy. Soc. Vict (n.s.), xlv: 2, 33-84.
WooLNouGH, W. G., 1909.—General Geology of Marulan and Tallong. Proc. LINN. Soc. N.S.W.,
xxxiv: 782-808.

EXPLANATION OF PLATES.
(Photographs kindly taken by Dr. I. A. Brown.)

Puate I.

Dicellograptus caduceus Lapworth, Loc. 9, No. 8.1645. x 1.

Dicellograptus cf. forchammeri Geinitz, Loc. 4, No. S.1761. x 2:3.

Dicranograptus nicholsoni Hopkinson, Loc. 11, No. 8.1657. x 2-3.

Dicranograptus nicholsoni Hopkinson, Loc. 11, coll. by Messrs. Ingall, Joklik and Scott-
(Oper 3 74

5. Dicranograptus cf. contortws Ruedemann, Loc. 11, coll. by Messrs. Ingall, Joklik and

Scott-Orr. x 2.

BP wnw

PLATE If.

6. Diplograptus calcaratus Lapworth, Loc. 11, No. S.1662. x 2-3.
7. Diplograptus apiculatus (Elles and Wood), Loc. 11, No. 8.1662. x 4.

82

GRAPTOLITES FROM TALLONG AND THE SHOALHAVEN GORGE, N.S.W.,

Glossograptus hincksii (Hopkinson), Loc. 11, coll. by Messrs. Ingall, Joklik and Scott-
Orr oe2ro- ;
Glyptograptus rostratus sp. nov., Loc. 12, Sydney University Geological Collection Reg.
No. 8289, aspect one-quarter face. x 2:3.

Glyptograptus rostratus sp. nov., Loc. 12, No. 8.1926, aspect almost bi-profile. x 2:3.

Climacograptus peltifer (Lapworth), Loc. 11, coll. by Messrs. Ingall, Joklik and Scott-
Oyae; 9 3%

Climacograptus peltifer (Lapworth) with greatly enlarged vesicle, Loc. 11, coll. by
Messrs. Packham and Veevers. x 2:3.

Climacograptus tridentatus (Lapworth), Loc. 11, No. 8.1662. x 2:3.

Lasiograptus harknessi (Nicholson), Loc. 11, coll. by Messrs. Ingall, Joklik and Scott-
Orr °x 4:

THE GENUS DAWSONIA.
By ALan Burces, Botany School, University, Sydney.

(Twenty-six Text-figures. )

[Read 27th April, 1949.]

INTRODUCTION.

The species of the genus Dawsonia form a well-marked group of mosses, and
although closely resembling members of the Polytrichaceae in vegetative structure, the
dorsiventral capsule with a peristome composed of a large number of hairs clearly
distinguishes Dawsonia from any other genus. It was originally described by R. Brown
in 1811 for the species D. polytrichoides from Eastern Australia. Since then fourteen
other species have been recorded. The genus extends from Tasmania and New Zealand
through Australia and New Guinea to Borneo, the Celebes and the Philippines.

As in the related Polytrichaceae, there is a conspicuous uniformity of major
vegetative characters and considerable taxonomic interest has been centred on the
structure of the longitudinal lamellae which occur on the upper surface of the leaf.
Schleiphacke and Geheeb (1896) gave the outlines of a proposed monograph of the
genus but, so far as the writer has been able to discover, this monograph was never
completed. -In the preliminary report they divided the genus into two sections:

Section 1.—Polytrichoides, in which the apical cells of the lamellae, as seen in
cross section, were not differentiated from the lower cells but were only a
little longer; and

Section 2.—Superba, in which the apical cell of the lamellae was distinctly
different from the lower cells, usually the upper cell being considerably
larger and more transparent.

In the present treatment these divisions have not been followed, instead, two new
subdivisions are proposed: the Longifolia and the Brevifolia. The Longifolia section has
its main centre in New Guinea but extends southwards into Australia, Tasmania, and
New Zealand with D. superba, and northwards to the Philippines with the closely related
D. altissima. The Brevifolia section has a similar distribution with the northern
D. brevifolia in Borneo and D.. polytrichoides and D. longiseta forming the southern
limit in Eastern Australia and Tasmania.

DAWSONIA.
R. Brown: Trans. Linn. Soc., 10: 312, 1811.
Triplocoma La Pyl. in Desv. Jour. Bot., 1813, p. 7.

Robust plants, usually gregarious, covering considerable areas and forming distinct
communities. Stems simple, in the sterile and male plants, normally unbranched,
varying in height from a few cms. up to 70 or 80 ems. The lower part of the stem is
buried in the ground and covered with white rhizoids. Above ground the stems are
usually naked in their lower part, distinctly three-ribbed, shiny, dark-brown or black,
with scars indicating the positions of old leaf bases. The ribs on the stems twist
spirally, usually clockwise, when viewed from above. In most species the leaves are
uniform and show no marked differentiation into cauline and comal leaves, but a few
species have distinct scale-like cauline leaves with no lamina. The basic leaf structure
is very uniform throughout the genus. The sheathing base which encloses two sides
of the three-ribbed stem, the midrib corresponding to one of the stem ridges, is usually
colourless or of varying shades of orange-brown with a distinct red-brown nerve. The
cells of the wings of the leaf base are thin-walled, linear, about 10-124 broad by 80-120u
long, with either pointed or straight end walls. At the shoulders of the leaf base the
cell-walls are very conspicuously thickened and the cell-cavities become small and

I

84 THE GENUS DAWSONIA,

lenticular, about 8 x 3u with their axes at right angles to the long axis of the leaf. A
group of the most heavily thickened appear to form a group of motor cells which
determine the angle the leaf blade makes with the stem, according to their water
content. Above the sheathing base the leaf contracts abruptly to form a narrow lamina
which has a central nerve and is covered on most of its upper surface by vertical lamellae.
These lamellae never occupy the full width of the leaf and in all species there is a
distinct border of thickened, usually reddish brown cells. In some species, owing to the
lamellae being squashed somewhat flat during examination, it is often difficult to detect
the border, but in others the border is obvious and in D. longiseta may become consider-
ably infolded particularly at the apex and make the leaf almost cucullate. The margin
bears reddish brown spinose teeth in all species. These teeth are large, usually 60—-80u
long and are formed from a single cell with very heavily thickened walls. Occasionally
the apex of the teeth become hyaline. In older plants the teeth and margin become
eroded, particularly in the smaller species, and may disappear even before the capsule
is fully ripe. The lamellae are composed of a single layer of cells and vary in height
from 30-100u. There is a considerable variation in the structure of the lamellae in the
different species. The discoid male flowers are terminal, reddish in colour and usually
about 3-4 mm. in diameter. The perigonial bracts resemble very enlarged cordate leaf
bases with a short point in place of the leaf blade. After flowering the stems often
proliferate and plants may show evidence of many successive male flowers. Female
plants are similar to the sterile, the terminal female flowers being less conspicuous
than the male. At first the young capsules are erect and completely covered in a densely
hairy calyptra, which is usually bright orange to red. As the capsule matures it becomes
inclined and assumes its dorsiventral structure. The mature capsule is horizontal or
slight hanging with the operculum obliquely set and often almost vertical. The main
body of the capsule is broadly ovate, flattened or slightly concave above, convex below.
When old and empty it becomes almost V-shaped in cross section. The peristome is
composed of a large number of linear hairs forming a dirty white tuft which usually
is very slightly twisted and contorted. New shoots may occasionally arise below the
female flowers and form branched stems. Hach stem usually produces only a single
capsule but in most species two or more may sometimes be found arising from a single
female flower.

Key to Species.

ile j Leaf bladesmore-thant. 20mm VOM AI tens cussccern Geir bene vere noma aperote ales eect onal ceen on n rc aeenoe 2

| Leaf bladevless: thants2 Oram! Om Sar (Aer cata ale ee eule tae hay cae cie tee e ete TORE SDS ocean arene tl

2. if Mareintorlamellaexcrenace sin) SId emi Gwaun iercraciornometeucrsia ate noiene mectereee enelnee eens eee 3

) Margin Of lamellae rplanenycSi-ih.r iis as ey baad eas chats elhle, S Ebs te GROMER Ch Ia OSTEO une 4

3. { Lamellae 5-6 cells high’........... enn OIC OM eetins OMG hia asad clo SB SS bo D. Pullet

) eevencilene: 23s GSMS IMIR Sac aasecococoo abo Doce sono oon DCdoDOODOSOSCUOECH OOS D. papuana

4, iy Leaves not spirally twisted nor with faleate tips when dry .....................---- 5

| Leaves Spirally, twisted om with faleatel tips whenratryese sascenectee aciere ciokeloicinneieies 6

by § Apical cell of lamellae heavily thickened, lamellae of 3-5 rows of cells ...... D. gigantea

| Apical cells of lamellae not heavily thickened, lamellae of 5-9 rows of cells .. D. superba

6. Lowest row of cells of lamellae quadrate or hexagonal .................... D. altissima

LAt leastisomevot lowest row, of cellsrrhomboidalararcsrmieeie ie chore: D. grandis

fits j Stems Less) theames5) Crys Wa ey tes isan sas ere cep ees seiner ie pratie ees eee a koa ee mee NCH ie re ec 13

Sissy Tanvornsy Wingham, WAN Gos acdoccbsoodsuccndocsoslbansadsdcecdouoocUaoaCSLOE 8

8. j Leaves sSomewhatetal care wdistinctly crispedawhenednry: - se seis citer ieeen D. crispifolia

| Leaves not talcate;shardlyienispedenwhene dryer. a ace noncieie nee teie ee meats ie eee renee 9

@). { Leaves distinctlyz-appressed Swihen vary, Vy eA ins canine eiecenaialicl Sta eee ee ee Ae 10

) Leaves ONE Siaeosaked ye loi oneesierol yaveral \CbAY Go odosudGbocububbooUrboDDOO oo UbMObE AUD OS BOS ilal

10. j Leaves linear inanrow,, (9 hmm lone ene Miey gt. hiked Sivepe bcp eee Seat ater ene ae D. limbata
) Leaves INHARO\W? wokenorbilehe, CealGy incvan, WOK? SS on anocenoodonundcunogududeauocen D. Beccarii

ial, {f Stem with distinct scale-like leaves in the lower part ..................... D. brevifolia
) Stem without: distinct Scale leaviesn ru: fscrs ake eines ears he cure Ree Oe ee 12

N27 j Lamellae Of = Oy LOWS GOL COlISiicrah Sas pore oes toe caeemeter atone | eceee ua ilon sloth paren ore aT cra eet D. intermedia
| Lamellae OHENTC Hie MON ASuCOue MCX Binks og ctolao dole MIDS SG OU Ge Macao So oad ooo GOS D. polytrichoides

13, j Leaves IDG UES OLNWSS Or’ AMMO OHWSS Yocoodsssudussdiocochoedoooconbos D. longiseta

leaves linear wWanceolatetacnte lass np ee aie ieee D. polytrichoides var. minor

BY ALAN BURGES. 85

LONGIFOLIA.

Plants tall 20-60 cms., lower part of the stems naked and shiny, leaves long
20-35 mm., linear or narrow linear-lanceolate, very acute. Capsule robust, usually 1 cm.
or more long.

1. Dawsonia Papuana F.v.M., ex Schlie. et Geh.
Rev. Bryol., 23: 76, 1896. Schliephacke et Geheeb.

Stems tall, more than 20 ems. high, the lower part naked, the upper 15-20 cms.
clothed with leaves. Leaves, when moist, flat and standing almost at right angles to
the stem, when dry, twisted along their axis, flexuose, the tips usually somewhat falcate
and the margins only slightly inrolled except at the tip. Leaf base sheathing, reddish
brown 2-3 mm. long by 1:5-2 mm. broad, with a distinct nerve. Leaf blade narrow
linear lanceolate, tapering to an acute toothed apex, 25-30 mm. long. Margin toothed
almost to the base. Apex of the leaf thickly set both back and front with numerous
teeth 80u long. Lamellae about 80 in number, not quite touching each other, 30—-35u
high, usually of three rows of cells, occasionally only two, the apical cell only slightly
differentiated from the lower and a little larger. Margin slightly crenulate from the
curved upper walls of the apical cells. In side view apical cells usually elongated at
right angles to the long axis of the leaf, measuring 12u x 8u but occasionally square.
Cells of the central row fairly regularly hexagonal about 8u across. Male plants not
known. Seta long, 3:5 cm., yellowish below, reddish above, capsule 1 cm., horizontal to
almost pendant, light brown. Operculum somewhat attenuated 6 mm. long.

The above description is based on material from the type gathering now in the
Melbourne Herbarium and the Natural History Museum, South Kensington. Text-figures
1 and 14 were drawn from material now in the Natural History Museum Collection.

The original gathering was made by McGregor at 4,900 feet about 48 miles N.E. of
Kikori near the Hather Gorge on the Upper Purari River, Mount Musgrave, New
Guinea, on 25th June, 1889. Portions were distributed by Mueller as D. Papuana n. sp.,
Geheeb being one of the recipients. It has not been possible to find any trace of a
description having been published by Mueller and it would appear that the first published
description is that given by Schliephacke and Geheeb in 1896. The same species was
again gathered by McGregor in 1890 from Mt. Yule and in 1891 from Mt. Suckling,
New Guinea.

The species appears to be closely allied to D. superba and D. altissima. It is,
however, readily separated from the former by the lamellae, being no more than three
cells high, whereas in D. superba they are rarely less than five cells high and by the
difference in the marginal cells of the lamellae. From D. altissima it can be distin-
guished by the apical cells in side view being taller than they are wide, and in the
crenulate margin of the lamellae. In dried material the flexuose, twisted, flatter leaf
blades readily distinguish D. Papuana from the straight more inrolled and more
appressea leaves of D. superba.

2. Dawsonia superba Grev.
Greville, Ann. Nat. Hist., 19: 226, 1847.

D. longifolia Brown m:.s.

Stems tall 20-70 cms., usually about 380-40 cms., unbranched except the female
plants, which may sometimes produce one or more lateral shoots below an inflorescence.
In the field the stems are straight but often show a tendency to be flexuose when dried.
Lower part of stem naked, shiny, dark brown to black, in places light brown due to
persistent leaf bases. Upper 15 cms. of stem leafy, leaves when dry, straight and almost
vertical although not markedly appressed to the stem. Leaf margin inrolled from
immediately above the sheathing leaf base so that the leaf blade appears very narrow
and tapering and only very occasionally shows any tendency to be twisted or flexuose.
When moist, leaf blade flat, standing at right angles to the stem or slightly refiexed.
Leaf base sheathing, orange-red becoming light brown when old; up to 4 mm. long,
1:5 mm. wide at the base, 3 mm. wide at the shoulder with a distinct central nerve.

86 THE GENUS DAWSONIA,

Leaf base suddenly narrowed to the long linear blade. Leaf blade 20-24 mm. long,
slightly less than 1 mm. wide, remaining more or less constant in width for about
two-thirds of its length and then tapering to a fine point. Margin strongly toothed
throughout. Apex toothed at both back and front for about 5 mm., teeth at the back
of the nerve usually in pairs or in threes. Lamellae about 60 in number, 60-804 high
usually of about six rows of cells but the number is variable and may be as many as
nine or occasionally as few as four. Margin almost straight. Apical cells distinct from

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Text-figures 1-6.—Detail of lamellae shown in side view.
1. Dawsonia Papuana. 2. D. superba. 3. D. altissima. 4. D. grandis. 5. D. gigantea.
6. D. Pullei. AU x 220.

the lower cells of the lamellae, usually colourless and almost square, 14u across, very
variable in their degree of thickening. At times the apical cell wall hardly differs from
that of the lower cells; at others the apical cell walls are heavily thickened. A similar
variation is seen in sectional view; the apical may be much wider than the lower cells
and be almost obovate or may be very little wider and almost rectangular and subquad-
rate, about 10u in diameter. Female plants similar to the sterile plants, perichaetial
leaves hardly differing from the foliage leaves. Seta usually short and somewhat
flexuose, the capsule usually not exceeding the comal leaves in the dry condition. This,

BY ALAN BURGES. 87

however, appears to be a variable character and although average measurements for the
seta are 10-15 mm. by 0:5 mm., some New Zealand specimens have the setae up to
3-5 ems. long by 1 mm. thick. Calyptra large, up to 1 cm., densely covered with silky
red or orange hairs. Capsule flattened, borne at right angles to the seta, concave above,
convex below, 7 X 4 mm. Peristome a dense tuft of pale brown or dirty white hairs
3 mm. long. The orientation of the capsule is very variable owing to the manner in
which the seta curves when dry.

Type not seen. Text-figures 2 and 15 drawn from material collected from Dorrigo,
N.S.W.

This species is widespread and abundant throughout Eastern Australia, Tasmania
and New Zealand, where it is usually found in rain forests or wet sclerophyll forest.
Dixon (1922) includes D. superba for New Guinea. I know of no certain record from
New Guinea. The material collected by Brass and assigned by Bartram to D. superba is
D. Pullei (see p. 91). There is, however, a sheet in the Melbourne Herbarium simply
labelled “New Guinea” with no collector, locality, or date. This specimen is D. superba
but approaches much nearer to D. altissima than any Australian or New Zealand material
I have seen. Bartram (1939) refers the Philippine species to D. superba. The identity
of these Philippine plants is discussed on page 89 and reasons are given there for
regarding them as D. altissima Geh. For the present, the writer regards D. superba as
a plant confined to Eastern Australia, Tasmania and New Zealand, and considers the
closely related plant found in Borneo, the Celebes and the Philippines as D. altissima.

3. Dawsonia altissima Geh.
Geheeb, Flora, 69: 352, 1886.

Stems up to 50-60 cms., unbranched, the upper 20-30 cms. leafy, lower part denuded
of leaves, triangular, shiny dark brown, three ribbed. Leaves when moist straight and
flat, standing almost at right angles to the stem. When dry crisped and twisted often
markedly so. Leaf base sheathing orange brown, with distinct mid-rib 3 x 2-5 mm.
Leaf blade linear, tapering in the upper third to a sharply toothed apex, 30-35 mm.
long. Margin toothed to within a short distance of the base. Apex very strongly toothed
both back and front, teeth at the apex up to 250u long. Lamellae about 60u, not quite
touching each other, 35u to 45u high of 4-5 cells. Lower cells fairly regular, usually
subquadrate 6-8u across, apical row very distinct, usually without chlorophyll and much
wider than high, about 10u x 164. Margin of lamellae almost straight, each individual
apical cell often corresponding fairly distinctly with two cells in the layer below. Seta
variable in length 2-3-5 cms. by 0:5-1 mm., usually slender, slightly curved or very
slightly arcuate when dry. Capsule large, 9 x 5 mm. Dixon (1934) records that the
specimens from the Celebes were characterized by having a very short stout seta, but
otherwise did not seem to differ from the Borneo plant.

The material collected by Burbidge on Mt. Kinabulu, Borneo, 1877-78, part of
which is now in Kew Herbarium, would appear to be the type material. Text-figures 3
and 16 and the above description are based on this material.

Apparently not uncommon in the higher mountains of Borneo, the Philippines, and
the Celebes.

There appears to have been considerable confusion regarding this species. When
it was first received by Geheeb, he regarded it as merely a robust form of D. superba.
Subsequently, however, he considered it to be sufficiently distinct and named it
D. altissima. The differences which separate it are admittedly slight and somewhat
dificult to define, nevertheless they were regarded as sufficient to warrant is retention
as a separate species, by Gepp (1913) and Dixon (1922). Confusion is only likely to
arise with D. superba. In typical specimens the difficulty is not very great. D. altissima
is a taller plant with longer leaves which always show some twisting along their axis.
In dried material the leaf blade is always flatter than in D. superba and the leaves are
less erect. In D. superta the leaves are usually straight and, when dry, the leaf margin
becomes inrolled from immediately above the sheathing base making the leaf very
narrowly triangular. Differences also exist in the appearance of the apical cells of the

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BY ALAN BURGES. 89

lamellae when seen in side view. This difference is difficult to define but seems to be
due to the way in which the thickening is laid down. In D. superba the vertical walls
between the apical cells appear as if made of a single layer which may vary greatly in
thickness; in D. altissima the thickened wall seems to consist of two distinct layers. In
most specimens of D. superba the apical cells are approximately square in side view and
those of D. altissima are about twice as long as they are high, but this difference is of
less value than the others given above. In the collections examined there has been little
trouble in placing material from Borneo and the Celebes as D. altissima. The material
from the Philippines has, however, proved much more difficult and approached much
more closely to D. superba as it occurs in Australia than did the specimens from Borneo
or Celebes. The Philippine material has previously been regarded as D. superba, first
by Brotherus and subsequently by Bartram (1939). It is possible that Brotherus, when
he identified the Philippine material, did not fully understand D. altissima as when
Pflanzenfamilien, Ed. I was written he had not seen D. altissima. The Philippine
specimens examined by me were:

(1) Collected by Elmer, Mt. Apo, 1909, identified by Brotherus as D. superba.

(2) Collected by Hachisuko, Mt. Apo, 1921.

(3) Collected by Mearus and Hutchinson, Mindanoa, identified by Brotherus as

D. superba.

In this material the twisting of the leaves, while never so distinct as in material
from Borneo, was, however, quite apparent. Most of the leaves, when dry, lacked the
typical inrolled margins and straight triangular outline of typical D. superba. Examina-
tion of the lamellae showed them to be very variable both in shape of the apical cell
and in the height of the lamellae. Nevertheless all the preparations showed the charac-
teristic thickening of D. altissima. Taking all the characters together, I feel that the
Philippine plants are best placed as D. altissima.

Whether D. altissima is worthy of full specific rank must be reconsidered at a later
date. When a wider range of material is available it may be found that several of the
large species such as D. altissima and D. papuana should be regarded as geographical
subspecies of D. superba, but in view of the lack of material and scanty state of our
knowledge regarding the moss flora of the area between Australia and the Philippines,
I do not think it wise to lump at present and have therefore retained D. altissima as
distinct.

4. Dawsonia grandis Schliep. and Geh.
Schliephacke and Geheeb, Rev. Bryol., 23: 88, 1896.

Stems simple, tall, 40 or more cms. high, the upper 20-30 cms. leafy; the lower
part of the stem triangular, denuded of leaves or with the torn remains of the sheathing
leaf bases, shiny dark brown to black, with scars of the old leaves. Leaves, when moist,
straight or with their tips slightly curved, standing at right angles to the stem; when
dry, usually somewhat twisted, the leaf blade about 45° to the stem. In material which
has been heavily pressed, the leaves tend to be rather straight with the apices of the
leaves, particularly the upper ones very regularly and noticeably falecately hooked. In
material which has dried without being heavily pressed, the regular falecate curving of
the leaf apices is less noticeable owing to the general crisping of the leaves. Leaf base
4 x 2-5 mm. orange-brown; leaf lamina 3-4 cm. long, 1-1:5 mm. broad, linear, upper third
tapering to a point. Apex toothed back and front, margin toothed almost to the base.
Lamellae about 120, 20-304 high. Margin of the lamellae straight, apical cells not very
different from the lower cells, approximately 12 x 12u, sometimes a little wider than
high. Cells of the lowest layer somewhat variable with many cells distinctly rhomboidal.
Capsules usually borne singly but not infrequently two capsules occur on a single stem.
Seta stout, rigid, usually about 3 cms. long by 1-2 mm. thick, brownish, often showing
distinct ribs when dried. Capsule large, 12 x 8 mm. broadly oval, compressed, about
3 mm. thick, slightly concave above, convex below horizontal to somewhat downward
pointing; when old, often becoming V-shaped in cross section. Peristome dirty white
up to 5 mm. long.

90 THE GENUS DAWSONIA,

Type material was collected by W. Armit, Mt. Musgrave, New Guinea, June, 1894.
This was distributed by F. v. Mueller as D. papuana; Geheeb, when he received it,
recognized it as distinct, and described it as D. grandis. Figures 4 and 17 and the
description were made from part of the original material.

D. grandis is apparently restricted to New Guinea, where it is probably the
commonest species. The stout rigid seta and the rhomboidal cells of the lower row in
the lamellae readily distinguish this species from all others. In some lamellae the bulk
of the lower cells may appear regular, but groups of rhomboidal cells are always
present.

5. Dawsonia gigantea C.M.
C. Mueller, Hedw., 36: 336, 1897.

Stems simple up to 50 cms. high, upper 10-20 cms. leafy, the lower part denuded of
leaves or covered with the remains of tattered leaf bases. Leaf base 3-5 x 3 mm. orange
with distinct central nerve. Leaf blade about 25 mm. long, 1-1:25 mm. wide, very
narrowly lanceolate, slightly contracted immediately above the sheath. When dry,
leaves show a slight tendency to be crisped, the leaf blade, however, tends to remain
flatter and the margins are less incurved than in some of the other species. Margin
toothed to within about 3-4 mm. of the base, apex strongly toothed, nerve toothed at
the back near the apex. Lamellae about 90 in number, approximately 50u high usually
of four, occasionally of three or five rows of cells. Apical cell very distinct both in
side view and in section. In side view l16u high, 20-254 wide, walls considerably
thickened, outer wall 4-64 thick. Lower rows of cells, subquadrate, thin-walled about
10u across. Capsule on a relatively stout seta, 10-15 mm. long by 1-2 mm. thick. Capsule
large, 10 x 7 mm., horizontal; operculum 5 mm. long. Peristome dirty yellowish-white,
3 mm., slightly twisted.

Text-figures 14-19.—Leaf outline.

14. Dawsonia Papuana. 15. D. superba. 16. D. altissima. 17. D. grandis.
18. D. gigantea. 19. D. Pullei.

BY ALAN BURGES. 91

Type not seen. Collected by Beccari, July, 1875, from Mt. Arfak, New Guinea,
apparently as No. 160 in Geheeb’s collection. Material from which the above description
and Text-figures 5 and 18 were drawn was collected by Gibb (No. 5523), Mt. Arfak,
December, 1913.

A very distinct species readily distinguished by the thickened apical cells of the
lamellae. Apparently restricted to New Guinea.

6. Dawsonia Pullei Fleischer.

Stems tall, up to 50 ems., upper 15-30 cms. leafy, lower part naked or covered with
torn leaf bases. Leaves, when dry, strongly crisped, the upper third of the leaf often
tightly coiled, leaf margins not inflexed, except near the apex. Lower leaves 15 mm.
long, upper leaves up to 30 mm. Lamellae about 80 in number, 50-70u high usually of
4-5 rows of cells, margin crenulate, apical cells in side view 15u high by 9u wide,
occasionally almost square. Walls of the apical cells very heavily thickened, particularly
on the outer wall. Lower cells variable, quadrate, rectangular to rounded hexagonal.
Seta 3-5 cms. by 2 mm. wide, dull orange-red above, later becoming dark brown or black.
Capsule large, 12-13 mm. by 5 mm., horizontal when maturing, operculum up to 6 mm.;
calyptra covered with hairs, reddish orange. Peristome dirty white, 2-3 mm. long.

Type material collected by A. Pulle on Mont Hellwig, alt. 1,800 M., No. 727, 17th
December, 1912, on the Third Dutch Expedition to New Guinea. Material of this species
was distributed as D. Pullei n. sp. Fleischer. So far I have been unable to find any
published description of this species. What is clearly the same species was collected by
L. J. Brass (No. 10554) at Lake Habbema, New Guinea, in October, 1938. This material
was recorded by Bartram (1942) as a form of D. superba. The above description and
figures 6 and 19 have been drawn from the part:‘of the original collection now in Kew
Herbarium.

D. Pullei is by far the most distinct of the large Dawsonias; the tight spiral coiling
of the leaves and the characteristic lamellae readily separate it from the other species.
It is known only from the two New Guinea collections.

BREVIFOLIA.

Plants 10-25 ems. high, leaves usually under 15 mm. long, narrow linear or narrow
linear triangular, very acute, seta slender, capsule usually 4-8 mm. long.

The species of this section fall naturally into three groups. D. polytrichoides and
D. intermedia from Australia, which form the first group, suggest small specimens of
the longifolia. The New Guinea species D. Beccarii, D. crispifolia and D. limbata,
with their narrow, almost subulate leaf blade, are clearly closely allied to each other.
The Borneo and Celebes, D. brevifolia, as might be expected from its distribution, stands
somewhat apart from the others.

7. Dawsonia intermedia C.M.
C. Mueller, Hedw., 1897, 335.

Stems simple, variable in height from 10-30 cm., lower part of stems bare, triangular,
blackish, upper 6-10 cm. leafy. Leaves uniform, when moist, standing at right angles
to the stem; when dry, making an angle of about 45° with the stem. Sheathing leaf
base 3 x 2 mm. colourless or orange with an orange-brown central nerve. Lamina narrow
linear lanceolate, about 10-15 mm. long by 1 mm. broad, margin strongly toothed. Apex
with several closely set teeth, often reddish. Margin inrolled, when dry, making the leaf
almost terete. Lamellae 50-60 in number, about 100u high, of 5-9 rows of cells. Margin
of lamellae almost straight, marginal row of cells differentiated from the lower cells
variable in size and shape, usually rectangular, 16 x 244 with long axis either at right
angles or parallel to the leaf axis, thick outer wall usually with a granular appearance.
Lower cells quadrate or polygonal, 10u across. Perichaetial leaves grading into the
cauline, innermost with few lamellae, the lamina being reduced to a toothed subula at
the end of a large sheathing base. Seta 15 mm. x 0-75 mm., when dry somewhat flexuose
and twisted, ribbed with 5-8 ribs in the upper part. Capsule ovate 8 x 6 mm. Peristome
28 mm. long, dirty white, slightly twisted.

92 THE GENUS DAWSONIA,

Above description and Text-figures 7 and 20 are based on material collected by
Luehmann, 1881, Upper Yarra, Victoria, originally in Herb. Bescherelle, now in Natural
History Museum collection. Luehmann’s gathering apparently forms the type material.

Not uncommon in eastern New South Wales and Victoria, and probably also occurs
in Queensland. It appears to be a rain forest species and is fairly distinct. Although
resembling D. polytrichoides in the field, it can readily be separated on the size of the
lamellae.

8. Dawsonia polytrichoides R. Br.
R. Brown, Trans. Linn. Soc., 10: 316, 1811.

Stems simple, 5-20 cm., mostly about 10 cm., lower part of stem usually brownish
with the remains of the old leaf bases, triangular. Leaves, when dry, somewhat loosely
appressed but not markedly so. Sheathing base, 2 x 1-5 mm., pale or orange with orange-
brown central nerve. Lamina narrow linear lanceolate, 6-10 mm. x 0:75 mm., margin
and apex strongly toothed, nerve toothed at back above. When dry, leaf margin inrolled
making the leaves almost terete. Lamellae about 60 in number, 50u high, of 4 or 5 rows
of cells. Margin of lamellae distinctly crenate from the projecting margin cells.
Marginal cells distinct, thickened, outer wall often granular, about 10 x 12u elongated
in either direction. Lower cells 10 x 8u usually elongated parallel to the axis of the
leaf but sometimes almost quadrate or hexagonal. Perichaetial leaves consist of a spiny
flexuose subula and a greatly lengthened sheathing base. Seta about 2-5 cm. x 0-5 mm.,
orange-red when young, dark brown when old. Capsule ovate 5 x 3mm. Calyptra large,
1:55 cm. x 5 mm. densely set with crimson hairs. Peristome dirty white.

20
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23
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26

Text-figures 20-26.—Leaf outline.
20. Dawsonia intermedia. 21. D. polytrichoides. 22. D. brevifolia. 23. D. Beccarii.
24. D. limbata. 25. D. crispifolia. 26. D. longiseta.

Above description and Text-figures 8 and 21 are based on material collected from
National Park, N.S.W., by the writer. The material in the Herbarium, Kew, collected
by R. Brown, “Australia”, probably is the type. Examination of this material shows
it to agree well with numerous collections from New South Wales and Queensland.

Var. minor C.M., ? in lit. only.

BY ALAN BURGES. 93

Stems 2-4 cm., lower 1-2 cm. naked. Leaves when dry usually distinctly appressed
making the stems appear clavate, occasionally not very much appressed. Seta 1-5-2 cm.
Differing from the type in the short clavate stems. Superficially the plants resemble
D. longiseta in their short stems and by comparison long setas, but the structure of the
leaf is typical of D. polytrichoides.

Whitelegge’s specimen from Kangaroo Valley, 1885, would seem to be the type.

D. polytrichoides is by far the commonest species of the genus, it is widespread and
abundant in Eastern Australia where it and its var. minor most frequently occur on
freshly disturbed earth banks in sclerophyll forest. It is distributed from Tasmania to
Queensland.

9. Dawsonia brevifolia Gepp.
Gepp, Jour. Linn. Soc. Lond., 42: 209, 1914.

D. Kinabaluensis Broth. MS., in herb.

Stems mainly simple, female plants occasionally branched, 20-30 cm. tall, slender
lower part of stem base naked or covered with scale leaves, upper 3-10 cm. with normal
leaves. Leaves, when dry, appressed to the stems making the shoots almost cylindrical,
occasionally somewhat crisped, when moist, more or less patent. Sheathing leaf base
2-5 x 1:75 mm., orange to brown with a central dark brown nerve. Cells of the shoulder
uniformly thickened, irregular, oval 8 x 124 without any well-marked motor cells.
Leaf blade variable from 7-10 mm., occasionally up to 15 mm. in the comal tuft, narrow
linear acute. The cauline leaves reduced to a triangular leaf base with an apiculus
formed, from the excurrent nerve, devoid of lamellae; cells of the wings of the base
much laxer and more parenchymatous than of the comal leaves. Comal leaves with
strongly toothed margin, somewhat inflexed. Lamellae about 70 in number, 80-1004
high, usually of 6-7 rows of cells. Margin crenate with projecting cells. Marginal cells
16 x 104 without any marked thickening, elongated at right angles to the axis of the
leaf; lower cells very regularly quadrate, 10-124 across. Perichaetial leaves similar to
comal leaves, slightly more strongly toothed. Seta 1:5-2-5 em. x 0:75 mm., often slightly
curved. Capsule horizontal, ovate, 8 x 4 mm. Operculum narrow, pointed, 3 mm.

Description and Text-figures 9 and 22 are drawn from type material in Natural
History Museum, collected by Gibb, 1910, Kinabalu, Borneo.

This species has been collected several times from Borneo and also from the Celebes
(Dixon, 1934). Gibb collected it in 1910 and Gepp’s description was published in 1914.
The same species was collected by Clemens (her. No. 10549), also from Kinabalu in
1915 and provisionally named by Brotherus, D. Kinabaluensis n. sp. Apparently he later
became aware of Gepp’s description and dropped the name D. Kinabaluensis. The name,
however, does appear in Hertzog’s Geographie der Moose. An examination of material
collected by Clemens, a sheet of which is in Kew Herbarium, shows that Brotherus’
D. Kinabaluensis is clearly D. brevifolia, a very distinct species in the leaf structure.
The absence of typical motor cells in the leaf base gives the plant a very distinct habit
and the structure of the lamellae readily distinguishes it from any of the other species.
It occurs in both Borneo and the Celebes.

10. Dawsonia Beccarii Broth. et Geh.
Brotherus and Geheeb, Rev. Bryol., 23: 73, 1896.
Ibid., Biblioth. Bot. Hfte., 44: 14, 1898.

D, altissima C.M. MS., herb.

D, filicaulis Geh. MS., herb.

Stems simple, tall, up to 40 em., lower part of the stem bare, dark brown to blackish,
triangular. Leaves straight or occasionally somewhat crisped on the lower part of the
stem. When dry leaves closely appressed giving the stem a very slender appearance,
when moist standing away from the stem but still more or less erect. Leaf base
sheathing 3 x 1:5 mm., colourless to brownish, with an orange central nerve. Leaf blade
very narrowly triangular varying in length from 8-15 mm. Margin strongly spinose,
inflexed, apex very spinose, orange red, the lamellae ceasing some distance below the

/

94 THE GENUS DAWSONIA,

apex. Lamellae about 40 in number, 60u high, margin crenate due to projecting cells,
usually 5-6 rows high, margin cells distinct with thickened walls, square to rectangular,
12 x 12-20u, lower cells variable quadrate or hexagonal, about 10-12 across. Some of
the cells of the lowest row are often elongated parallel to the axis of the leaf and may
measure 20-30u x 8u. Seta about 2 cm. x 1mm. Capsule ovate 8 x 5 mm., horizontal.

Type not seen, originally collected by Beccari on Mt. Arfak, New Guinea, July, 1875.
The above description and Text-figures 10 and 23 are based on material collected by
Gibbs (No. 5521 and 6005) from Mt. Arfak. Subsequently recorded from Lake Habbema
and Mt. Wilhelmina, New Guinea.

The appressed leaves and the slender appearance make the species easily distinguish-
able from the others.

11. Dawsonia limbata Dix.
Dixon, Jour. Linn. Soc. Lond., 45: 486, 1922.

Stems simple, 10-15 cm., lower 3 ecm. bare, triangular dark brown. Leaves when
dry straight and appressed, when moist more or less spreading. Sheathing base of
leaves 3 X 2 mm. orange, with an orange brown central nerve. Leaf blade narrow linear
7-9 mm. x 0:5 mm. wide. Margin spinose, incurved, often somewhat whitish, apex
toothed. Lamellae about 40, 70u high, margin crenate from projecting cells, 6-9 rows
high, marginal row distinct thick-walled, 12 x 12u, lower cells quadrate or rectangular,
8 across, walls somewhat thickened. Capsule apparently unknown.

The above description and Text-figures 11 and 24 are based on material in Herb.
Dixon now in the Natural History Museum, South Kensington.

This and the two preceding species are readily separated from the remainder of the
genus when dry by their uniformly appressed leaves. The leaves of D. limbata are
shorter, more linear and less acute than those of D. Beccarii, which taper more or less
gradually throughout their length. The species is so far known only from New Guinea.

12. Dawsonia crispifolia Dix.
Dixon, Jour. Linn. Soc. Lond., 45: 486, 1922.

Stems simple, 15-25 cm. high, subfiexuose, lower part covered with the remains of
the old leaf bases, dirty brown. Leaves very characteristically crisped when dry, slightly
crisped to falcate when moist. Sheathing base almost triangular, orange brown with a
central reddish brown nerve. Leaf blade narrow, gradually tapering from 0-5 mm. to
an acute point, somewhat falcate, 8-10 mm. long. Margin with slender slightly curved
spinose teeth. Lamellae 40-50, about 80u high, margin crenate due to the projecting
cells. Lamellae mostly of 6 rows of cells, sometimes 7-8, marginal row conspicuously
thickened 14 x 6y elongated at right angles to the axis of the leaf, lower cells with
thickened walls, subquadrate or subrectilinear, mostly 6 x 10u. Seta about 2 cm. x
0-75 mm. slightly arcuate, capsule horizontal to hanging, broadly ovate, 8 x 6 mm.,
peristome large, 4 mm., dirty white.

The above description and Text-figures 12 and 25 are based on the type material
collected by Kloss, Mt. Carsters, New Guinea.

Readily distinguished from all other species of the genus by its narrow falcately
cerisped leaves and slender almost rigid habit. Brotherus in Pflanzenfamilien, Ed. II,
gives D. crispata Dix., apparently a slip, for this species. Fleischer evidently following
Brotherus returned D. crispata for specimens collected by Lam in New Quinea, and
this name thus appears in Lam’s Fragmenta Papuana (p. 119, English translation).

D. crispifolia is so far known only from New Guinea.

13. Dawsonia longiseta Hpe.
Hampe, Linn., 1860, 634.
Dawsonia appressa Hpe., Linn., 1860, 634.
Dawsonia Victoriae C.M., Hedw., 1897, 331.
Stems simple, 1-3 cm., lower part with small scale leaves, upper part densely
clothed with normal leaves. Leaves when moist set at right angles to the stem, usually
with a somewhat glaucous appearance; when dry appressed to the stem so that the sterile

BY ALAN BURGES. 95

shoots appear clavate. Leaves narrow lingulate, often slightly curved and asymmetrical,
incurved when dry. Margin wider than in the other species, strongly inflexed so that
at first sight the margin appears entire. At the apex the inflexed margin makes the
leaf almost cucullate. Leaf base sheathing 2 x 15 mm., usually colourless except for
the brown nerve; lamina 6-7 mm. long x 0:75 to 1 mm. wide. Lamellae numerous,
usually about 80, up to 65u high, of four to five rows of cells, margin crenulate due
to projecting cells. Marginal cells 20 x 15u, outer wall strongly thickened up to 5u
thick, usually elongated at right angles to the axis of the leaf. Lower cells very variable
from thickened quadrate or hexagonal to thin-walled rectangular, mainly 12 across.
Seta orange-red, long, up to 3:5 cm., by 0-5 mm. Calyptra conical, 8 x 2 mm., densely
covered with orange-crimson hairs. Perichaetial leaves with a very much reduced
lamina and usually without lamellae. Apex very variable, in some plants obtuse and
jagged toothed, in others finely acute. Capsule ovate 5 x 2 mm., horizontal, peristome
silky or dirty white, 1:5 mm.

The above description is a composite description based on a wide range of material
from Eastern Australia. Text-figures 13 and 26 are from the type specimen in Hampe’s
herbarium now at the Natural History Museum, South Kensington. D. longiseta is
common on recently disturbed soil in sclerophyll forest from Tasmania to Queensland.
Easily recognized by its small size and lingulate leaves. In this latter respect the
figures in Plate ix of Mueller’s Australian Mosses are somewhat misleading.

In reducing Hampe’s D. appressa and C. Mueller’s D. Victoriae to synonyms, it may
seem that I have lumped more than is justified. However, after examining almost a
hundred collections of plants attributed to these species I find no justification for
retaining them as separate species, as all intergradations exist between the three forms
which can be regarded as the original species. Furthermore, characters which might
perhaps be associated with one species may be found together with characters from
another and several combinations of characters will often occur in the one tuft.

Examination of the type specimens of D. longiseta and D. appressa, now at Natural
History Museum, South Kensington, and a comparison of these with part of the type
gathering of D. Victoriae from the Melbourne Herbarium shows them to be somewhat
distinct. D. longiseta with stems about 2 cm. long, seta about 3 cm. straight, the leaves
thick and inrolled appearing almost terete when dry; D. appressa with stems about
2 cm. long almost clavate from the appressed leaves, seta about 2 cm., usually with a
pronounced curve a few mm. behind the capsule; D. Victoriae a small plant usually
with the leaves flatter and more curved. There are also minor differences in the cells
of the lamellae. When a range of specimens is examined these distinctions break down
and it is impossible to assign the majority of specimens to one or other of the forms
just mentioned. C. Mueller separated his D. Victoriae from D. longiseta on its smaller
size, etc., and its obtuse perichaetial leaves. A dissection of a plant from the type
gathering, however, shows obtuse and acute perichaetial leaves on the same plant. He
does not compare his plant with D. appressa, which it much more closely resembles.
Hampe distinguishes D. appressa as follows: “D. longiseta proxima differt; caule
clavato longiore, foliis brevioribus, sicca appressis, seta breviore, theca minore, angulata
et peristomia sericeoniveo.” These characters, however, will not stand, and it seems
clear that both D. Victoriae and D. appressa are forms of a fairly variable D. longiseta.

Other workers seem to have met similar difficulties in separating these three
supposed species. Most Victorian material seems to have been labelled D. Victoriae,
apparently for no other reason than that it came from Victoria. All the New South
Wales and Queensland material has been called D. longiseta. In herb. Dixon there is a
good range of D. longiseta and alongside one packet from Fernshaw, Victoria, Dixon has
written “much of this might well be D. appressa’, yet some of the plants might equally
well have been the original for F.v.M.’s figure of D. longiseta in his Australian Mosses.

Invalid Names.

In addition to the species described above, the following names appear either in the
literature or in herbaria:

96 THE GENUS DAWSONIA,

D. altissima C.M., herbarium MS. name only = D. Beccarii Broth. and Geh.

D. crispata Dix. Appears in Pflanzenfamilien, Ed. II, and is apparently an error for
D. crispifolia. The same name appears in Lam’s Fragmenta Papuana.

D. insignis. In the Handbuch der Pflanzenanatomie, Bd. VII/I, Anatomie der Laubmoose.
Lorch refers (p. 65) to a D. insignis. I have been unable to find any other trace of
such a species.

D. Kinabaluensis Broth. Brotherus’ species was never published although distributed
with specimens. On the appearance of Gepp’s D. brevifolia, with which it is
obviously conspecific, D. Kinabaluensis was dropped.

D. longifolia R. Brown. Appears on herbarium sheets at Kew and South Kensington.
The specimens are clearly D. superba.

D. longisetacea appears as an error for D. longiseta in F. v. Mueller’s Australian Mosses.

D. Novae—Zeelandiae appears on a specimen at Kew, collected by Colenso near Auckland,
New Zealand. It had subsequently been labelled D. superba, which it clearly is.

ACKNOWLEDGEMENTS.

I would like to express my grateful thanks to the following for making available specimens
and records in their care and for their generous help in the present work: Dr. Ramsbottom and
Mr. Sherrin, of the Natural History Museum, South Kensington; the Director of the Royal
Botanic Gardens, Kew; Mr. Jessup and Mr. Willis of the Herbarium, Botanic Gardens,
Melbourne; Mr. Anderson of the National Herbarium, Sydney; and Mr. Bartram of Bushkill,
Pike County, U.S.A.

References.

Bartram, E., 1934.—The Mosses of the Philippines. Philippine Journ. Sci., 68:1.
—, 1942.—Third Archbold Expedition. Mosses from the Snow Mountains, Netherlands

New Guinea. Lloydia, 5: 245.

Dixon, H. N., 1922.—The Mosses of the Wollaston Expedition to Dutch New Guinea, 1912-13,
with some additional mosses from British New Guinea. Journ. Linn. Soc. Lond., 45: 477.

, 1934.—Mosses of the Celebes. Ann. Bryol., 7:19.

GeEpp, A., 1913, in Gibbs, L. S., 1913.—A contribution to the flora and plant formations of Mount
Kinabalu and the Highlands of North Borneo. Journ. Linn. Soc. Lond. (Bot.), 42: 1.

SCHLIEPHACKE, C., and GEHEEB, A., 1896.—Essai d’une monographie du genre Dawsonia. Rev.
Bryol., 23 2 (3-

97

REVISION OF THE GENUS BRACHYCOME CASS.
PART Il, NEW ZEALAND SPECIES.

By Gwenpa L. Davis, B.Sc., Lecturer in Biology, New England
University College.

(Twenty-four Text-figures. )

[Read 30th March, 1949.]

INTRODUCTION.

The genus Brachycome was first recorded from New Zealand by J. D. Hooker (1853)
with the description of B. radicata. Since that date extensive exploration has shown that
the genus is widely distributed in that country, particularly in the South Island.

In Part I of this paper (these ProcEeprines, Ixxiii, 142-241) the suggestion was put
forward that this genus had already colonized New Zealand prior to its connection being
severed with the Australian mainland. This hypothesis would account for the fact that
all species belong to the subgenus Hubrachycome, there being no representatives of the
subgenus Metabrachycome in New Zealand. If the present areas occupied by species be
any guide to their origin, it would appear that it was B. radicata, or its ancestral form,
which was the original colonist and that it spread widely before the land mass separated
into North and South Islands. B. Sinclairii, on the other hand, had it originated later
in the South Island, would have been prevented from migrating both to the North
Island and to Stewart Island by effective barriers of water. No specimens of
B. Sinciairii have been examined from either Stewart or North Islands, though the
possibility cannot be overlooked that it exists but has not been collected there. Should
this prove to be the case, it would go a long way towards explaining the presence on
Three Kings Islands of this species. Up to date no specimens of Brachycome have been
collected in the North Island north of the Kaimanawa Range.

The similarity in the flora and fauna between the islands of Subantarctica and New
Zealand has been the occasion for comment by various authors. Chiltern* (1909) points
out “the flora of the islands to the south of New Zealand is in its main characters and
alliances nothing more than a branch of the New Zealand flora, to some extent changed
and modified by long ages of isolation”. He also remarks that Campbell and Auckland
Islands ‘are not oceanic islands, but once formed part of a continental area connecting
them with the present mainland of New Zealand’’. It is therefore interesting to examine
specimens of B. radicata from Campbell Island. Unfortunately these are in flower only,
but the very young fruit present are densely glandular, and the habit agrees with that of
B. radicata from the mainland. It is more than likely that this species occurs also on
Auckland and Macquarie Islands. This record supports the view that B. radicata was
the original species of Brachycome in New Zealand, but it is remarkable that in isolation
on Campbell Island it has followed an identical line of development to that pertaining in
New Zealand. B. linearis has been collected only from the shores of Lake Te Anau. In
the glabrous and smooth nature of the fruit it appears to be most closely allied to
B. Sinclairii, but the affinity is not close.

NOMENCLATURE.

The general principles formulated in Part I of this paper are followed here. It is
regrettable that both the well-known specific epithets ‘odorata’ and “Thomsonii” must
be abandoned in favour of a little-known but earlier synonym.

In the past there was a strong tendency for botanists to regard each species and
variety as static. It is now coming to be generally recognized that each category is made

* The Subantarctic Islands of New Zealand. Vol. II, pp. 467 and 796.

98 REVISION OF THE GENUS BRACHYCOME CASS. PART II,

up of living individuals which, since they are living, have an inherent capacity for
variation, and once that capacity is lost the population is in danger of dying out in a
changing environment. The former practice of honouring each variation with a separate
name adds confusion rather than clarity to taxonomic work, and in this paper, separate
names are only applied to units of a discontinuous series. Within each unit limits of
variation are established within which the individuals of the particular population
oscillate about a hypothetical central point to which the term “typical” is applied.

SPECIMENS EXAMINED.

Several hundred specimens have been critically examined as a result of the courtesy
of the Directors of various public herbaria who lent their collections for this purpose.
These specimens are all listed under appropriate species, the source in each case being
indicated as follows:

Auckland Institute and Museum (A).
Canterbury Museum, Christchurch (C).
Dominion Museum, Wellington (W).
National Herbarium, Melbourne (MEL).
National Herbarium, Sydney (NSW).
Brisbane Herbarium (BRI).

Specimens in young flower only, are indicated by an asterisk, but microscopic
characters of the very young fruit are such that little doubt exists as to their correct
determination.

TAXONOMY.
Compositae, Tribe Asteroidea.

Brachycome Cass., Dict. Sci. Nat., 37 (1825), 491.
Subgenus Hubrachycome Davis.
Synonymy and definition of the genus as in Part I.

Key to the Species.
1. Glandular branching perennials up to 25:6 cm. high, with cauline leaves which are often
clustered basally.
2. Lower leaves oblanceolate or spathulate and toothed. Fruit oblong-cuneate,
slightly flattened, more or less densely covered with short glandular hairs.

Pa PPUSHCONSDICWOUS ase ez chaceasecio eee at acu ssecereaesianace = Ne Ree Eee EE Bode HOUT Rea 1. B. radicata
2*. Lower leaves spathulate and entire to pinnatifid or pinnatipartite. Fruit narrow-
obovate, flat, glabrous. Pappus conspicuous, deciduous ......... 2. B. Sinclairu

1*. Glabrous annuals not more than 3:4 em. high. Leaves radical, linear to narrow lanceo-
late, entire or with a single small tooth. Fruit narrow oblong, glabrous, turgid. Pappus
MICLOSCOpPIeFaNndAECIAUOUSM te ee cee IC ICO aon eae can 3. B. linearis

1. BRACHYCOME RADICATA Hook. f., Fl. N.Z., I (1853), 127.

(Text-figs. 1, 4, 7.)

Synonymy: B. radicata Hook. f., var. a, Fl. N.Z., I (1853), 127.

odorata Hook. f., Handb. N.Z. Fl. (1864), 138.

Thomsonii T. Kirk, Trans. N.Z. Inst., xvi (1884), 372.

Thomsonii var. membracifolia T. Kirk, St. Fl. N.Z. (1899), 261.

Thomsonii var. dubia T. Kirk, St. Fl. N.Z. (1899), 261.

polita T. Kirk, St. Fl. N.Z. (1899), 261.

Thomsonii var. polita Cheesem., Fl. N.Z. (1906), 277.

Haptotype: Patea, Wellington Province. Col. 1732. W. Colenso (A).

Weakly erect to erect stoloniferous perennials up to 25:6 em. high, branching from
the base, more or less densely glandular all over, rarely almost glabrous. Lower cauline
leaves oblanceolate or spathulate in gross outline, basally attenuate into a petiole, up to
9-5 cm. long, 2:3 cm. broad, with obtuse to acute teeth. Upper leaves sessile, often
becoming entire, and finally scale-like. Radical leaves present on young plants and
similar to lower cauline. Peduncles leafy and terminate branches, scape-like when
solitary. Inflorescences usually 1 or 2, but up to 6 may be present, 7-13 mm. diameter.
Involucral bracts 18-24, 3-5-4:-5 mm. long, 1-5-2 mm. broad, broadly oblanceolate, apex
obtuse to acute with torn-ciliate margins and usually bearing short glandular hairs on

by ty ty ty by by

vadicata.

1. B. radicata. Habit. x 3.

Fruit.

of B. radicata.

J

BY GWENDA L. DAVIS.

Text-figures 1-8.
2. B. Sinclairii. Habit. x 4. 3. B. linearis. Habit. x 1. 4. B.

x 22. 5. B. Sinclairii. Fruit. x 22. 6. B. linearis. Fruit x 45.
8. Distribution of B. Sinclairii (@) and B. linearis (x).

99

7. Distribution

100 REVISION OF THE GENUS BRACHYCOME CASS. PART II,

the outer surface. Ray florets 14-30, 3-5 mm. long, 1:1-1:5 mm. broad, apparently always
white. Receptacle 3-5-7 mm. broad, 1-7-5 mm. high, hemispherical. Frwit 2-3-5 mm.
long, 0:9-1:'4 mm. broad, dark brown to black, oblong-cuneate, slightly flattened, more or
less densely covered with short glandular hairs. Pappus conspicuous, straw-coloured,
bristles of irregular length and often spreading.

Range: North Island south of Kaimanawa Range, and widespread in both South Island and
Stewart Island; also Windlass Bay, Campbell Island.

Specimens examined: North Island: Tutira Run, Hawke Bay, H. Guthrie Smith (A);
Kaweka Mt., 2,500 ft., H. Tryon (W, BRI); Kaimanawa Range, 1.1914, B. C. Aston (A, W, C);
Kaimanawa Range, 1.1914, D. Petrie (W); Ruahine Range, 1.1914, B. C. Aston (W) ; Waimarino
Co., “in forest’, 1.1918, H. Carse (W); Matatoke Gorge, 1.1918, H. B. Matthews and H. Carse
(A); Wanganui (W); Patea, Wellington Province, W. Colenso n. 1732 (Haptotype of B.
radicata var. a, A); Mt. Holdsworth, Tararua Range, 3,000-4,000 ft., W. Townson (A).

South ~ Istand: ‘Cobb > Valley, Ne Wi -sNelson; pe Gs Gibbs EG) Me Arthur
plateau, Nelson, 4,000 ft., 1.1886, T. F. Cheeseman (lectotype and 2 lectoparatypes of B.
Thomsonii var. membracifolia, A; lectoparatype, W); Mt. Arthur, Nelson, J. Adams (A);
W. Nelson, W. Townson (C); Nelson, W. Townson (A); Mt. Murchison, W. Townson (W);
Mt. Murchison, 3,000 ft., W. Townson (A); Brunner Range, 3,000-4,000 ft., W. Townson (A);
Tarndale, Nothofagus forest, 3,800 ft., L. Cockayne (W); W. region of Amuri Co., W. G.
Morrison (W); Mt. Miro Miro, 19.1.1919, L. Cockayne (W); Mt. Miro Miro, 3,500 ft., D. Petrie
(W); Arthur’s Pass, 3,000 ft., T. Kirk (W); Arthur’s Pass, T. Kirk (lectotype of B. polita and
B. Thomsonii var. polita (A); Arthur’s Pass, “in moist ground near Lake Misery”, 26.1.1898,
L. Cockayne (W, A); Arthur’s Pass, T. F. Cheeseman (W); Arthur’s Pass, 2.1919, A. Wall
(C); margin of bush, Mingha Valley, 21.1.1932 (C); near Lyttleton, “‘rocky stations’, 1.1908,
D. Petrie (W); Oamaru Bluff, D. Petrie (W); Cape Whanbrow, T. Kirk (lectotype and lecto-
paratype of B. Thomsonii var. dubia, A; lectoparatype, W); Mt. Pisa Plateau, 5,500 ft., “soil
on rock’, 9.3.1921, lL. Cockayne (W); Mt. Pisa Plateau, 9.3.1921, W. R. Reid (A); *Upper
Routeburn, 4,000 ft., 7.5.1921, L. Cockayne (A); Otago Penin., 1.2.1920 (W); Dunedin, D. Petrie
(W); near Dunedin, between Otago Hbr. Isthmus and Green Is., 300 ft., D. Petrie (W); Green
Is., “sea cliffs’, D. Petrie (W); Black Head, St. Clair, Dunedin, 1.1920, A. Wall (C); Black
Head, Dunedin, A. Wall (A); Mt. Cleghearn, W. of Waiau R., Southland, 1.1914, I. Crosby
Smith (W); Takitimo Mts., ‘‘moist slopes in tussock’, 30.12.1912, D. Petrie (W); Bluff Hill,
Southland, 21.1.1913, D. Petrie (W); Fortrose, ‘‘near sea’’, 4.1.1913, D. Petrie (C); Fortrose,
“on clay near sea cliffs’, 4.1.1913, D. Petrie (W) ; Fortrose, 1.1913, D. Petrie (A).

Stewart Island: Mason Bay, 1.1907, LL. Cockayne (W); Mason Bay, ‘“‘open places amongst
scrub’’, 30.1.1907 (W); Mason Bay, 1910-1911, J. W. Murdoch (W); Mason Bay, 15-18, 1.1940,
J. E. Attwood (A); Paterson Inlet, Stewart Is., G. M. Thomson (A, C); near Pegasus Sound,
1.1877, D. Petrie (W); Sydney Cove, 1.1882, T. Kirk (W); Stewart Island, 1.1880, G M.
Thomson (BRI); Stewart Island, 12.1883, T. Kirk (W),; Stewart Is., J. W. Murdoch (W);
Stewart Is., T. Kirk (lectotype and three lectoparatypes of B. Thomsonii, A; two lectoparatypes,
MEL).

With the exception of var. minima Kirk (‘Dog Is.”’), syntype material of all
varieties of B. Thomsonii has been located in New Zealand and lectotypes nominated.

B. radicata var. a was described by Hooker from three specimens (“Northern Is.,
Cunningham, Colenso; Middle Is. Lyall’), and in 1864 he described B. odorata with the
comment “the B. radicata of Fl. N.Z. was founded partly on this, and partly on specimens
of a Brachycome in A. Cunningham’s herbarium, which I am now convinced were intro-
duced there by accident, and belong to an Australian species. The name of B. radicata
had therefore better be abandoned”.

B. odorata was accordingly described and the type locality given as “Patea Village,
Colonso”, a somewhat more detailed locality record than that quoted in connection with
B. radicata, but apparently based on the same specimen. Attempts to trace syntypes of
B. radicata have only been successful to the extent of locating a specimen collected by
Colenso bearing the type data in the Auckland Museum. Since, however, there must
remain some doubt as to whether Hooker actually handled this particular specimen, it
has been nominated haptotype both of B. radicata and B. odorata. It is unfortunate that
all Hooker’s syntypes are no longer extant, but in the event of one belonging to another
species, as Hooker suspected, the validly published name cannot be abandoned for that
reason alone, B. radicata is accordingly redescribed and the name B. odorata rejected as
a later synonym.

* Campbell Is.: Cliffs above Windlass Bay, 10.12.1944, K. L. Oliver (W) >; cliffs, Windlass
Bay, 12.1.1945, K. L. Oliver (W).

BY GWENDA L. DAVIS. 101

Specimens available show a continuous variation of vegetative features, and many have
been examined which occupy intermediate positions between varieties. Since the prac-
tice of naming variations in a continuous series is not upheld by the present author,
these varieties, having probably an ecological basis, are abandoned. In his description
of B. Thomsonii var. minima Kirk records “Flower heads smaller, ray florets wanting.
Habitat Dog Island’. Hither the ray florets were overlooked or this was a specimen of
Lagenophora, and since no syntype specimens have been traced this variety is not
listed in the synonymy of B. radicata. Syntype material of B. polita has been located and
type selection made. These specimens fall well within the new specific limits of
B. radicata except that the pappus recorded by Kirk as being absent is certainly present
but tends to be dehiscent with age.

Variation within this species is not great, the most striking feature being that of
the indumentum. The vast majority of specimens are more or less densely glandular,
certain of them being macroscopically so. However, sparsely glandular plants are not
uncommon, and a few were seen from the Otago district which were glabrous. The
degree of development of the indumentum cannot be correlated with either altitude or
geographical position and probably has a genetical basis. All specimens examined from
Stewart Is. were densely glandular, which tends to support this latter view. Variation
in size and shape of leaves is no more than would be expected in a species occupying a
variety of habitats in a relatively extensive range. In this connection it is interesting to
note Cheesman’s comments (1925) following his redescriptions of the following species:

B. odorata: “...1am inclined to think that this is not separable from B. Thomsonii.”

B. Thomsonii: “. .. A most variable plant, only separable from B. odorata by the
larger size, coarser habit, and larger heads.”

The strong fragrance of this species has been commented on by various authors but
no information is available as to the position of the glands responsible.

2. BRACHYCOME SINCLAIRII Hook. f. Handb. N.Z. Fl. (1864), 137.
(Text-figs. 2, 5, 8.)

. radicata var. B Hook. f., Fl. Nov. Zel., I (1853), 127.

Sinclairii var. a Hook. f., Handb. N.Z. Fl. (1864), 137

Sinclairii var. B Hook. f., Handb. N.Z. Fl. (1864), 137.

Sinclairii var. y Hook. f., Handb. N.Z. Fl. (1864), 137.

pinnata Hook. f. Handb. N.Z. Fl. (1864), 138.

Sinclairii Hook. f. var. montana T. Kirk, St. Fl. N.Z. (1899).

Weakly erect to erect stoloniferous perennials up to 26 cm. high with a glandular
hairy indumentum, rarely almost glabrous. Leaves clustered basally, up to 4:2 em. long,
1:1 cm. broad, very variable in shape, from spathulate and entire to pinnatifid or pinnati-
partite. Segments obtuse to acute, usually regular. Blade passes abruptly into the
petiole, or the leaf may appear sessile. Peduncles scape-like, naked or with a single
bract, usually densely glandular distally. Inflorescence 0-9-1 cm. diameter, usually 1 or 2
present, rarely 8. Involucral bracts 16-84, 3:5-5-5 mm. long, 1-2-1:-5 mm. broad, lanceolate,
glandular on outer surface, obtuse to acute, with torn ciliate margins. Ray florets 25-39,
5-9 mm. long, 1:2 mm. broad, apparently always white. Receptacle 2-5 mm. diameter,
1-4 mm. high, hemispherical. Frwit 1:7-2-3 mm. long, 0-:9-1:1 mm. broad, narrow-obovate,
glabrous, flat, with narrow smooth margins, dark brown. Pappus conspicuous, deciduous.

Synonymy :

my by by bY OD oy

Range: Widespread throughout the South Island, otherwise only recorded from Three Kings
Islands.

Specimens examined: *Three Kings Is. (W).

South Island: *Mt. Arthur, 3,000 ft., “in Beech forest’, 1.1.1934, E. M. Heine (W); *Mt.
Arthur Plateau, 1.1886, T. F. Cheeseman (A); Mt. Owen, 4,500 ft., 1.1882, T. F. Cheeseman
(A); Mt. Owen, “about 4,500 ft.”, W. Townson (W, C); Mt. Owen, W. Townson (A); *Wairaw
Valley, 4,000 ft., 1.1878, T. F. Cheeseman (A); Whites Bay, 18.1.1880, F. M. Reader (MEL) ;
Awatere Valley, 12.1926, I. M. McMahon (C); Ward, “limestone hills near coast’, 12.1915,
B. C. Aston (W); *Ward Hills (W); Camden Ra., Kiakoura Mts., 2.1930, A. Wall (C);
Clarence Valley, 3,800 ft., 1.1875, DT. Kirk (W); Clarence Valley, 3,000 ft., 1.1893, Tl m
Cheeseman (A); western region of Amuri Co., W. G. Morrison (C, W); Hanmer Plains,
Christenson (W); Mt. Percival, Amuri Co., about 4,000 ft., 10.2.1914, W. G. Morrison (CW);
Amuri. T. Kirk (W, MEL): *Mt. Halfmoon, Dillon R., about 5,000 ft., 12.1926, A. Wall (@)) 2

JS

102 i REVISION OF THE GENUS BRACHYCOME CASS. PART II,

*Upper Hurunui R., 10.1913, A. Wall (C); Bealey, Mingha, 1.1.1928 (C); Bealey Valley, near
Mingha Junction, “river bed’, 15.1.1928, W. R. B. Oliver (W); *Mingha Valley, “Beech Forest’,
15.1.1928, W. R. B. Oliver (W); slopes of Rolleston, 4,500 ft., 25.1.1929, LL. Cockayne (C);
Blimit Cirque, 16.2.1928 (C); *Blimit Cirque, 29.1.1928, W. R. B. Oliver (W); Rough Crk.,
Avalanche Ridge, 4,500 ft. (C); Graham’s Creek, 1.1928, M. Laing (C); mountain above
Avalanche Gully, 17.1.1928, W. R. B. Oliver (W); Broken River, D. Petrie (W); *dry banks of
Broken R., 11.1936, N. Lothian (MEL); Craigie Burn Mts., Canterbury Alps, 1.18938, D. Petrie
(W); *Canterbury Alps, N. T. Carrington (W); Canterbury Plains, T. Kirk (A,* MEL) ;
Canterbury, J. Haast (MEL); *Southern Alps, 2-5,000 ft. (W); Thomas River, Canterbury,
T. Kirk (C); Burnham (W); Ashburton Mts., T. H. Potts (W); Ashburton, 11.1921, H. H.
Allan (A); vicinity of Ashburton, H. H. Allan (A); *Mt. Potts, R. M. Laing (NSW) ; Woolshed
Creek, Mt. Somers, 4.1926, A. Wall (C); Little Spur Mt., Upper Rangitata, 5,000 ft., 1.1921,
A. Wall (C); *Mt. Peel, Canterbury, H. H. Allen (W); *Mt. Peel, “grassland. Abundant in
tussock, 1,500-3,000 ft.”, H. H. Allan (W); *slope of Mt. Peel, ‘‘about 600 metres, in tall tussock
land’, 2.1.1919, H. H. Allan (C); Ball Spur, above Tasman Glacier, 1.1924, A. Wall (C); Lake
Tekapo, 5,000 ft., 1.1853, T. F. Cheeseman (A); Tasman Valley, north of Lake Pukaki, 13.2.1911,
D. Petrie (W); *mountains near Hunter River, Lake Hawea, 1920, R. A. Wilson (W); Mt.
Arnould, head of Lake Hawea, 1920, R. A. Wilson (W); Lake Ohau (W); mountains on
Temple Crk., Lake Ohau, about 5,000 ft., 1.1931, A. Wall (C); Mt. Turner, 1.1926, A. Wall (C);
*Lake Wanaka (W); *Mt. Ernslaw, 5,000 ft., E. Phillips Turner (A); Mt. Pisa, about 4,500 ft.,
D. Petrie (W); Mt. St. Bathans, about 3,000 ft., 12.1892, D. Petrie (W); Mt. Cadrona, D.
Petrie (W); Mt. Cadrona, about 4,500 ft., D. Petrie (C, W); Ben Lomond, Lake Wakatipu,
1.1932, A. Wall (C) ; *Remarkables, east side of Lake Wakatipu, about 4,000 ft., 2.1886, D. Petrie
(W); Remarkables, 5,500 ft., D. Petrie (W); Remarkables, 5,000 ft., 1.1923, A. Wall (C);
*Cromwell, 12.1925, A. Wall (C); Matahanui, Manukerikia Plain, 11.1892, D. Petrie (W);
Kurow, Waitaki River, 4.1892; 10.1893, D. Petrie (W); *Mt. Ida, Maniototo Co., 10.1.1911,
D. Petrie (W); Mt. Ida, 12.1908, B. C. Aston (W); Ngapara, D. Petrie (W); Ngapara, L.
Cockayne (W); Earnsleugh flat, near Clyde, D. Petrie (W) ;.*Rock and Pillar Range, 12.1908,
B. C. Aston (W); Kyeburn Crossing, Maniototo Plain, about 1,400 ft., 11.1892, D. Petrie (W) ;
Maerea’s Flat, Petrie (W); Macrea’s Flat, about 1,800 ft., D. Petrie (W); near Green Island,
Dunedin, ‘‘sea cliffs’, D. Petrie (W); Old Man Range, about 5,000 ft., 2.1920, A. Wall (W, C);
*Old Man Range, 2.12.1923, J. Speden (W); *Old Man Range, 1.1924, J. Speden and H. L.
Darton (W); *Old Man Range, 2,000-4,000 ft., D. Petrie (W); *Strath Taieri, D. Petrie (W);
*Bluff (W).

The following localities have not been traced: Trelissick Basin, T. Kirk (W); Tinker’s
Diggings, 1,000 ft., 11.1892, D. Petrie (W); Mt. Princess, about 3,000 ft., 1.1927, A. Wall (C).

Since syntype specimens of all varieties are in the herbarium of the Royal Botanic
Gardens, Kew, they have not been examined by the author, and type selection has there-
fore not been made. There is no doubt, however, as to the identity of this species.

B. radicata var. B was described by Hooker from a single specimen (“Southern Is.,
Lyall’) with the note “var. 6 looks like a different species, but I have only one small
specimen and that in young flower only’. Later (1864), and apparently without having
obtained further specimens, Hooker raised this variety to specific rank as B. pinnata
with the comment “I have but one specimen of this pretty little plant’. The description
is, of necessity, limited to vegetative characters. The practice of describing any new
category on a single incomplete specimen needs no comment. Hooker’s specimen is now
at Kew, but several plants which agree perfectly with his description have been
examined, and the fruits are identical with those of B. Sinclairii.

The extremely variable nature of the leaves of this species is reflected in the long
synonymy listed above. This variation from entire to almost pinnate margins is shown
to be continuous (Text-figs. 9-24), and it is not uncommon for specimens to bear both
entire and divided leaves in varying proportions. Five specimens were examined from
Cromwell (12.1925, A. Wall), one of which bore exclusively the almost pinnate leaves
of Hooker’s var. pinnata, another bore the usual pinnatifid leaves, and the remaining
three specimens bore both pinnatifid and pinnate leaves. In view of this, to give
separate status to each peak of variation is a fruitless attempt to discover vegetative
stability where none exists, and this entire population is now grouped under one name.

In his original description of B. Sinclairii, Hooker describes the pappus as being
“minute, bristly or 0”. Examination of a long series of specimens shows that while in
mature fruit, the pappus is frequently lacking or poorly developed, in younger fruit on
the same plant it is invariably well developed. The pappus is therefore commonly
deciduous, seldom persisting into the mature condition.

BY GWENDA L. DAVIS. 103

9

3. BRACHYCOME LINEARIS (Petrie) Druce. Bot. Exc. Club, iv (1914-1916), 614.
(Text-figs. 3, 6, 8.)

Synonymy : Lagenophora linearis Petrie, Proc. N.Z. Inst., xxv (1892), 271.

; Brachycome lineata T. Kirk, St. Fl. N.Z. (1899), 259.

Lectotype: Lake Te Anau, 1.1886, D. Petrie (A).

Lectoparatypes: Three, Lake Te Anau, 1.1886, D. Petrie (A); *three (W).

Erect glabrous annuals, 1:7-3-4 cm. high. Leaves radical, linear to narrow oblanceo-
late, obtuse, up to 2:3 em. long, 1-2 mm. broad, entire or rarely with a single linear tooth.
Scapes 1-6, glabrous, naked, filiform. Inflorescences 2-5-3 mm. diameter. IJnvolucral
bracts 8-12, 1-1-6 mm. long, 0-6-0-9 mm. broad, ovate to oblong, obtuse to subacute with
minutely torn-ciliate margins. Ray florets about 12, the rays 1-1-2 mm. long, 0:2—0:3 mm.
broad, apparently white. Receptacle 1:5 mm. broad, 0:2 mm. high, very slightly convex.
Fruit 1-1-2 mm. long, 0-5-0-6 mm. broad, narrow oblong, glabrous, golden brown, turgid
at maturity. Pappus white, microscopic, deciduous.

Range: Apparently confined to the type locality.

Specimens examined: Lake Te Anau, 1.1886, D. Petrie (lectotype and three lectoparatypes,
A); three lectoparatypes (W); shores of Lake Te Anau, at foot of lake, 1.1892, D. Petrie (W) ;
Lake Te Anau, 1.1932, A. Wall (W). ?

B. linearis is known only from Lake Te Anau, though the possibility cannot be
ignored that owing to its small size its presence elsewhere has been overlooked. It shows
no close affinity with other described species, though in the glabrous and somewhat
flattened fruit it is nearest to B. Sinclairii. B. linearis may represent a relic of the
primitive stock persisting in isolation, but if so, similar forms should reasonably be
expected to occur in similar pockets elsewhere. If the distribution of this species is
restricted to its present known locality then it probably originated there directly from
the ancestral stock as a mutation and its lack of variation suggests that its chances of
survival are small.

The following three species were described in 1943, the types of which are, according
to the original descriptions, in the Herbarium of the Plant Research Bureau, Wellington,
but since it has not been possible to examine these specimens the original descriptions
are merely transcribed with comments.

Brachycome humile. Simpson and Thomson, Trans. Roy. Soc. N.Z., \xxiii, Pt. 3, 1943.

“A very small tufted plant. Leaves radical, 1-2 cm. long, linear spathulate, 1 mm.
broad at the tip, entire, or with 1-2 notches on one or both margins towards the tip,
rather fleshy, gland dotted or minutely pubescent. Scapes 1 or more from the base,
+ 3 cm. long, almost filiform, glabrous below, with glandular hairs at the tips; heads
solitary, + 6 mm. diam.; involucral bracts 9-10, lanceolate; margins purplish, scarious.
Ray florets about 12, twice as long as the involucral bracts, oblong obtuse, yellowish and
pilose at the base. Achenes obovate, slightly compressed, with a few stiff hairs; pappus of
small scales.

“Habitat: Snow hollows.

“Type specimen from the summit of Rock and Pillar Range, Otago, in the Herbarium,
Plant Research Bureau, Wellington. Smaller in size than B. linearis and differing by
larger flowers, longer ray florets, by the hairs on the corolla and achenes and the distinct
pappus scales.”

If this species is as close to B. linearis as the description suggests, it may well
represent another relic of the ancestral stock which, in isolation, has developed along
slightly different lines to the present B. linearis population. The presence of stiff hairs
on the fruit, however, is an unusual feature, unique among New Zealand species, and
what is meant by ‘distinct pappus scales” is, in the absence of a text-figure, obscure.

Brachycome longiscapa Simpson and Thomson, Trans. Roy. Soc. N.Z., \xxiii, Pt. 3
(1943).

“Rootstock short, stout, with many slender roots, branching closely to form small
dense clumps. Leaves radical, 4-6 cm. long, 8-12 mm. broad, spathulate or linear
spathulate, rather fleshy, darkish green, minutely or mealy pubescent; petiole stout,
flattened above, rounded beneath, widened at the base, as long as or longer than the
blade; blade obovate, rounded or blunt at the tip, narrowed to the petiole, entire or the

104 REVISION OF THE GENUS BRACHYCOME CASS. PART II,

margin very irregularly lobed or toothed or pinnatifid, the midrib sunken above,
prominent beneath. Scapes strict, long, 10-20 cm., slender, pale brown or purplish, naked,
glabrous at the base, closely covered towards the tip with short, stout, glandular hairs.
Heads 1:5 em. diam.; involucral bracts 4 mm. long, in two series, oblong, reddish,
glandular hairy, with the margins scarious, rough and blackened; ray florets 8 mm.

6 see eel?,

4 ee.
ee
aoe
ee.

20 20 208
ISA I9B
: 22 23 24
21A

Text-figures 9-24.
Leaf variation in B. Sinclairii. Leaves of the same ordinal number are taken from
the same plant. x &.

BY GWENDA L. DAVIS. 105

long, numerous, linear obovate, obtuse; dise florets green. Achenes obovate, compressed,
with minute pappus scales.

“Habitat: Grassland.

“Type specimen from the Upper Shag Valley, near Kyeburn, in the Herbarium,
Plant Research Bureau, Wellington.

“A common grassland plant in the valleys of the Shag and Waitaki Rivers and their
tributaries, Lindis Pass, L. Pukaki, etc. The irregularly margined, minutely pubescent,
darkish green leaves, and the long, slender scapes are unlike those of any other species.”

The vegetative characters as listed in the original description fall within the limits
of variation of B. Sinclairii, and although certain of these are referred to by the authors
as unlike those of any other species, in this revision vegetative characters alone are not
considered sufficient to warrant specific status. In the description of the fruit the pappus
is said to consist of minute scales whereas that of B. Sinclairii is conspicuous but
commonly deciduous at maturity. It is possible that in the fruit examined by the.
authors of B. longiscapa the pappus had already broken off leaving the persistent bases
of the original bristles which were then described as scales.

Brachycome montana Simpson, Trans. Roy. Soc. N.Z., xxv (1945), 197.

“Rootstock short, 2-3 mm. diam., with many slender roots, branching closely to form
small dense clumps. Leaves radical, 2-3 cm. long, 4-8 mm. broad, linear obovate-
spathulate, rather fleshly, greyish green, surfaces and margins closely dotted with stiff
glandular hairs; petiole as long as or twice as long as the blade, flat above and grooved,
rounded beneath; blade differing in shape and in the division of its margins, rounded or
narrowed to the petiole, sometimes almost entire, more usually with 2-7 irregularly
disposed, conspicuous, rounded lobes. Scapes 4-6 cm. long, rather stout, pale brown,
naked or with a single linear, acute bract, glandular hairy as the leaves. Heads 1 cm.
diam., involucral bracts 5 mm. long, in three series, linear oblong, acute, glandular
hairy, with scarious tips; ray florets 5 mm. long, in two series, linear obovate, rounded
at the tip, much recurved, disc florets numerous, yellowish. Achenes.

“Habitat: Type specimen from Mount Cardrona, Central Otago, collected by Mr. D. Leigh and
cultivated by Mr. W. B. Brockie at the Botanic Gardens, Christchurch, in the Herbarium, Plant
Research Bureau, Wellington.

“The diversiform, greyish green, glandular hairy and somewhat fleshy leaves
separate the species immediately from others of the genus.”

This description, being devoted exclusively to vegetative characters, is of little
diagnostic value, and could apply to certain variations of B. Sinclairii. Apart from the
word ‘‘Achenes’’, no mention is made of the fruit.

The following species are listed as nomena dubia, with transcriptions of the original
descriptions:

B. simplicifolia Armst., Trans. N.Z%. Inst., xiii (1881), 338.

“A small tufted branching perennial herb, 3-4 in. high, stout and leafy. Radical
leaves 2-3 in. long, linear-spathulate or linear, obscurely 3-nerved, obtuse, with broad
membranous sheathing petioles and revolute margins; quite entire, except the sheaths
which are somewhat shaggy, glandular-pubescent, or glabrous. Scapes 1-3 in. high,
pubescent, 1-2 flowered, striated or channelled. Cauline leaves or bracts few, 1 in. long,
linear-spathulate, obtuse, subamplexicaul, glandular-pubescent, more distinctly nerved
than the leaves. Heads half an inch in diameter. Involucral scales 12-16 in one series,
or with a second outer series of 2-3 scales only, linear-lanceolate or oblong, acute,
distinctly 3-nerved, coriaceous, appressed, with purplish membranous margins. Ray
florets short, oblong, obtuse, few or absent, white. Disc florets tubular, 3-5 toothed.
Pappus entirely absent. Achenes compressed, one-tenth inch long, glandular-pubescent,
thickened at the tips. Receptacle very narrow, convex, papillose.

“Habitat: Nelson Provincial District. Mr. C. W. Jennings, Marlborough Provincial District
(1869), J.B.A.”

Cheeseman (1925) notes: “B. simplicifolia J. B. Armstr. is quite unknown to me,
and there are no specimens in any public collection in the Dominion. It probably does
not belong to the genus.” Armstrong’s private herbarium is now housed in the Canter-
bury Museum, Christchurch, but contains no such specimens. It can only be assumed

106 REVISION OF THE GENUS BRACHYCOME CASS. PART II,

that all syntypes have been lost, and consequently there no longer exists any foundation
for the name.

B. alpina W. Colenso, Trans. N.Z. Inst., xxx (1898), 271.

“Plant small, slender, slight, hairy, simple (sometimes 2-branched); rhizome 3 in.
(and more) long, filiform. Leaves radical, scattered, suborbiculate, 4 lines diameter,
tapering, faintly crenate lobed, lobes few, their tips pointletted, hardened from vein
produced, dark-green above, paler below, membraneous, much veined; hairs sprinkled,
appressed, substrigillosa, white, flat, subulate, strangulated, thicker on upper surface.
Petioles very slender, 1 in. long, canaliculate and dilated at base with membranous
margins, and patent hairs, dark purple-brown. Scape erect, 3-34 in. long, filiform, with
2-3 small linear distinct bracts, glabrous but pubescent towards tip. Head small,
drooping, 2 lines diameter. Involucral bracts numerous, sub 20, linear dark-green with
a thick purple central nerve, margins membraneous, white; tips acute, jagged. Florets
few; ligulae white, revolute. Receptacle broad, naked, shining, alveolate. Pappus 0.
Achene sublanceolate, one-tenth inch long, slightly glandular, viscid.

Habitat: Ruahine Mountain Range, east side; Feb., 1898; Mr. H. Hill.”

This species is not mentioned by either Kirk or Cheeseman in their floras and no
specimens have been traced.

ACKNOWLEDGEMENTS.

This revision was made possible only by the co-operation of the Directors of various
herbaria, who generously made available their specimens for examination. In this connection
I would like to thank Dr. M. R. S. Oliver of Canterbury Museum, Christchurch; Dr. Archey
of Auckland Museum, Auckland; the Director of the Dominion Museum, Wellington; Mr. R. H.
Anderson of National Herbarium, Sydney; Mr. A. W. Jessep of National Herbarium, Melbourne;
and Mr. C. T. White of Brisbane Herbarium. I am also indebted to Miss B. E. Molesworth,
formerly of Auckland Museum, for transcriptions of several original descriptions and assistance
in tracing type specimens.

References.

ARMSTRONG, J. B., 1881.—Descriptions of new and rare New Zealand Plants. Trans. N.Z. Inst.,
xiii: 338.

CHEESEMAN, T. F., 1925.—Manual of New Zealand Flora, 2nd Hd., 907-910. Wellington.

CHILTERN, C. (Ed.), 1909.—Subantarctic Islands of New Zealand, ii. Wellington.

CoLENSO, W., 1898.—A description of a few more newly discovered indigenous plants; being a
further contribution towards making known the Botany of N.Z. Trans. N.Z. Inst., xxxi: 271.

Davis, G. L., 1948.—Revision of the genus Brachycome Cass., Part I. Australian Species. Proc.
LINN. Soc. N.S.W., Ixxiii, Pts. 3-4, 142-241.

DruseE, —, 1916.—Report of the Botanical Exchange Club, iv, 610.

Hooker, J. D., 1853.—Flora Novae-Zelandiae, Pt. I, 127. London.

, 1864.—Handbook of the New Zealand Flora, 137-138. London.

Kirk, T., 1884.—Descriptions of new plants collected on Stewart Island. Trans. N.Z. Inst., xvi:
372-373. ;

, 1899.—Student’s Flora of New Zealand: 259-261. Wellington.

PETRIE, D., 1892.—Descriptions of a new genus and new species of native plants. Proc. N.Z.
MGOMan, FO 273 11h

SIMPSON, G., and THOMSON, —, 1943.—Notes on Some N.Z. Plants. Trans. Roy. Soc. N.Z., xxiii,
Verbs G3 8° IHAILOe)e

, 1945.—Notes and Descriptions of New Species. Trans. Roy. Soc. N.Z., xxv, Pt. 2: 197.
WutuFr, EH. V., 1943.—Introduction to Plant Geography. Waltham.

CoRRIGENDA TO REVISION OF THE GENUS BRACHYCOME Cass. Part I.
(Volume 1xxiii, Parts 3-4.)

P. 171. The record of B. cardiocarpa from Queensland should be deleted. Southport, in
this case, refers to the Tasmanian locality.

223. Last line. For Nochnagar read Lochnagar.

228. Fourth line from the end of page. For obtuse to entire read obtuse and entire.

240. For map 24 B. longiscape read map 24 B. Readeri.

rd hd

107

ON AUSTRALIAN DERMESTIDAE. PART V.
NOTES AND THE DESCRIPTION OF FOUR NEW SPECIES.
By J. W. T. ARMSTRONG.

(One Text-figure. )

[Read 30th March, 1949.]

INTRODUCTION.
This paper contains the description of four new species of Australian Dermestidae
that have come under notice since Part IV* of this series was published.
Anthrenocerus stigmacrophilus n. sp. is a true inquiline, both the larvae and newly
emerged adults being found in the nests of the ant, Stigmacros foreli, which treats them
with complete toleration. Notes are also given on some previously described species.

TROGODERMA SILVICOLUM Nl. Sp.

Types: Holotype 6 exs. on card in coll. Armstrong, paratypes in British, Australian
and South Australian Museums and the collections of F. KE. Wilson and H. J. Davidson.
Type locality: Acacia Plateau, N.S.W.

DESCRIPTION.

Ovate, black, nitid, clothed with moderately short semi-depressed black setae,
antennae, mouth parts and tarsi fuscous.

Pronotum: Widest at base, sides evenly rounded to apex, medial lobe rather wide
with apex lightly emarginate, posterior angles acute, disc finely and not closely punctate,
with a wide, well-marked oblique depression on each side before the base.

Elytra: Three-quarters as wide as long, base as wide as pronotum, sides expanding
to shoulders, then lightly so for about half length, thence evenly rounded to apex, more
closely and strongly punctate than pronotum.

Antennae: ¢ long, extending well beyond base of presternum, first two joints rather
large moniliform, the remaining nine forming a long, loose, narrow club, of which the
first four become progressively larger, the next four are approximately similar and the
eleventh is ovate; 9? club four or five jointed, the seventh is somewhat wider than the
preceding one.

Size: 2-5-2 mm. x 1-:7-1:3 mm.

Discussion: Fifteen examples, taken on a flowering creeper in the edge of the rain-
forest in December, resemble the black variety of 7. reitteri Blackb., but that species has
only a three-jointed antennal club. They are more compact and shining than
T. maurulum Blackb. in which the club is eight-jointed.

Distribution: Acacia Plateau and Acacia Creek, N.S.W. (Armstrong).

TROGODERMA INCONSPICUUM Armst.

Armstrong, J. W. T., 1942.—Proc. Linn. Soc. N.S.W., Ixvii, 326.

Type: In South Australian Museum.

Type locality: Swan River, W.A.

Discussion: Hight specimens in the C.S.I.R. Collection evidently belong to this
species. Three have the elytra red except round the scutellum. In both sexes the
antennal club is three-segmented, but that of the ¢ is much stouter.

Distribution: Swan R. (Lea), Wurarga (A. Goerling), W.A.

* These PROCEEDINGS, Ixx, 1945, 47-52.

108 AUSTRALIAN DERMESTIDAE. PART V,

TROGODERMA LONGIUS Blackb.

Blackburn, T., 1903.—Trans. Roy. Soc. S.A., xxvii, 163.

Type: In British Museum.

Type locality: ? Glenelg R., Vic.

Discussion: Four specimens from Acacia Plateau were at first thought to be a new
species, but they agree with the description of this species, except that the antimedial
elytral fascia is absent and is represented only by a sparse pale pubescence. It is there-
fore considered better to treat them as a local variety, unless comparison with a typical
specimen, which is not now available to me, should reveal other differences.

Distribution: Glenelg R., Vic., Tas., Acacia Plateau, N.S.W. (Armstrong and
Davidson).

TROGODERMA CARTERI Armst.
Armstrong, J. W. T., 1942.—Proc. Linn. Soc. N.S.W., Ixvii, 328.
Types: In the author’s collection.
Type locality: Bogan R., N.S.W.

DESCRIPTION OF 6.

The ¢ differs from the 9 as follows:

Smaller.

Elytra: More or less darkly castanious.

Antennae: First two joints rather stout, club nine-segmented and stoutly pectinate.

Size: 2-75 x 1-25 mm.

Discussion: The form of the ¢ antennae places this species close to T. macleayi
Blackb., but it is narrower, less shining, and the third antennal segment distinctly forms
part of the club, and is not minute as in Blackburn’s species. Larvae were found in the
oothecae of mantids, the adults emerging at the end of August. Larvae were also found
in spiders’ webs, under loose bark and infesting stored insects. They are remarkable
for the very strongly clubbed black hairs on their dorsal surface. They have short
hastisetae, rather evenly distributed, but not numerous, barbed setae at sides, but no
long caudal setae. Specimens have been submitted to Dr. Bryant E. Rees for description.
From late in June, the adult insect is usually found fully developed resting in the exuvia.
When disturbed they are quite lively.

Distribution: Only known from type locality.

TROGODERMA CALLUBRIENSE (Armst.).
Armstrong, J. W. T., 1945.—Proc. Linn. Soc. N.S.W., 1xx, 48 (Psacus).
Type: In the author’s coll.
Type locality: Bogan R., N.S.W.
Synonymy: Psacus callubriensis Armstrong, 1945, loc. cit. (n. syn.).
Discussion: Hinton (Mon. Beetles Assoc. Stored Products, 1945, 375) placed the

genus Psacus aS a synonym of Trogoderma. My species therefore also becomes a
Trogoderma.

MEGATOMA FOVEOLATUS Lepesme.

Lepesme, P., 1941.—Bull. Soc. ent. Fr., 142.

Type: Unique, ? either in coll. Lepesme or Paris Museum.

Type locality: Australia.

Discussion: This species is unknown to me in nature. The description, however, is
very suggestive of a 9 of Trogoderma attagenoides (Pasc.) = Psacus attagenoides Pasc.,
though my specimen of this has not the pronotal depressions mentioned. These, however,
may have been caused by damage to the specimen when immature.

Lepesme separates the two Australian species attributed to Megatoma as follows:

“Corps court et large, noir; une seule fascie transversale, étroite, subdroite, sur les
élytres) 222.555 tenuefasciata Rtt.”

BY J. W. T. ARMSTRONG. 109

“Corps allongé, brun rouge; trois fascies transversales, étroites, sinueuses, peu
marquées, sur les élytres ........ foveolatus, n. sp.”
Some specimens of 7. attagenoides are brown in colour.

ADELAIDIA RIGUA Blackb.

Blackburn, T., 1891.—Trans. Roy. Soc. 8S. Aust., xiv, 130.

Type: In British Museum.

Type locality: South Australia.

Discussion: A specimen taken in the Bogan River district is evidently the male of
this species. It is 3-25 mm. long and the combined length of the three segments of the
antennal club exceeds that of the remainder, each segment being more than twice as
long as wide, with the terminal the longest. This is the only specimen known to me in
Australian collections.

Distribution: S.A., Bogan R., N.S.W.

ANTHRENOCERUS ARROWI, Nl. SD.

Types: Holotype in the author’s collection; paratypes in the British Museum, etc.

Type locality: Bogan R., N.S.W.

Synonyms: Anthrenocerus bicolor Armstrong nec. Arrow, Armstrong, 1943.—Proc.
Linn. Soc. N.S.W., 1xvili, 61-(n. syn.).

DESCRIPTION.

Ovate, sub-convex, nigro-piceus, elytra dark brown, antennae and legs brownish,
clothed with not very short dark and light depressed setae, the latter white interspersed
with stramineous and disposed much as in A. australis (Hope), in four irregular elytral
fasciae and two apical spots, and at sides, front and on basal lobe of pronotum; clothing
oi ventral surface short, ashy white and very fine. (Hvans, “Insect Pests and their
Control’, Dept. Agr. Tas., 1943, 154, gives a good figure of Hope’s species.)

Pronotum: Transverse, widest at base, sides evenly rounded to apex, deeply, closely
and moderately punctate. ;

Elytra: As wide as prothorax at base, expanding to shoulders almost in the same
line as sides of pronotum, thence gradually narrowing for approximately half length,
then evenly rounded to apex, coarsely and closely punctate.

Size: 2:5-1:9 mm. x 1-6-1 mm.

Discussion: Arrow recently pointed out my mistake in identifying this species as his
A. bicolor, and kindly sent a cotype of the latter which comes very near some varieties
of the species, regarded by Blackburn and myself as A. variabilis Reitt., but the
pronotum is black. The species here described differs from it in having the elytra darker
and not so uniformly brown, the clothing longer and coarser and the light setae of two
colours. The brown elytra separate it from A. australis (Hope), signatus Armst., and
blackburni Armst. There are forty-one specimens under examination, and numerous
others have been identified by me as A. bicolor Arrow in the various museum and private
collections of this country. It is hoped these identifications will be altered. The species
is tabulated under Arrow’s name (Proc. Linn. Soc. N.S.W., 1943, 58). Specimens in the
British Museum from Townsville tend to be somewhat more robust and those from
Western Australia have finer clothing. The species is named in gratitude for the help
and co-operation received from Mr. G. J. Arrow of the British Museum.

Distribution: Bogan R., N.S.W. (Armstrong), Townsville, Q. (F. P. Dodd), Yallingup,
southern W. Aust. (R. E. Turner). Widespread in southern and eastern Australia.

ANTHRENOCERUS BLACKBURNI Armst.

Armstrong, J. W. T., 1943.—Proc. Linn. Soc. N.S.W., Ixviii, 60, fig. 4.

Type: In South Australian Museum.

Type locality: ? Victoria. .

Discussion: In view of the detailed description of A. australis (Hope) given by
Hinton (Mon. Beetles Assoc. Stored Products, 1945, 369), which is presumably based on
material collected in England, it seems likely that this form represents only an extreme

110 AUSTRALIAN DERMESTIDAE. PART V,

variety of that species. <A. arrowi, described above, would perhaps also be better treated

as a sub-species of the same species. It is thought possible that a study of the larvae of

these and nearly related species would throw considerable light on their relationship.
Distribution: Viec.; Bombala, Illawarra, Acacia Plateau, N.S.W.

ANTHRENOCERUS STIGMACROPHILUS, Nn. sp. Text-fig. 1.
Types: Holotype in the author’s collection, paratypes in the British, Australian,
National and South Australian Museums.

Type locality: N.S.W., Bogan R. (J. Armstrong), in nests of a small ant (Stigmacros
foreli Viehm.) during November.

DESCRIPTION.

Ovate, black, nitid, clothed with semi-erect black setae sparsely and obscurely
patterned with white, the white setae form two obscure fasciae on the elytra, one anti-
medial and one post-medial, and are also present at base and apex, and are present on
the pronotum, being almost confined to the basal lobe but sometimes faintly visible else-
where, ventral clothing dark, antennae and legs red.

Pronotum: Transverse, sides narrowing gradually, then more suddenly rounded to
apex, basal lobe wide, base with a depressed line in front of it, sides narrowly explanate,
dise strongly convex, finely and closely punctate.

Elytra: As wide as prothorax at base, expanding to shoulders, thence slightly
narrowing for approximately half length, then evenly rounded to apex, more coarsely and
closely punctate.

Size: 3-2-4 mm. x 1-9-1:4 mm.

Discussion: Fifteen specimens were taken in the galleries of two nests of the ant
Stigmacros foreli Viehm. (thanks are due to the Rev. J. McAreavey for the identification )
together with a number of larvae and pupae. The larvae are very different from those
of A. australis (Hope) as figured by Evans (loc. cit.). The species would fall beside
A. australis (Hope) in my table (loc. cit.), but the pronotum is different and the pattern
much fainter. At first glance it appears close to A. niger Armst., but in that species the

Text-figure 1—Anthrenocerus stigmacrophilus, n. sp. Larva.

pronotum is not margined, and, in the one specimen so far taken, the eyes are golden.
The larvae (exuvia Text-fig. 1) are rather dark stramineous and have very short bluntly
clubbed hairs on the dorsal surface, a lateral fringe of pointed barbed setae, but no
hastisetae. Specimens have been submitted to Dr. Rees for description. There can be
no doubt that the species is truly myrmicophilous as the larvae can be found in the nests
of its host all the year round, and the beetles also remain there until the weather is
suitable for them to emerge.

Distribution: In addition to the type locality, there are larval specimens in the
National Museum, Washington, from Hornsby, N.S.W., also from an ants’ nest, that
appear close to this species, but Dr. W. H. Anderson believes that they represent a
closely allied species.

ANTHRENUS VORAX Waterh.
Waterhouse, C. O., 1883.—Ann. Mag. Nat. Hist. (5), xi, 61; Hinton, H. E., 1945.—
Mon. Beetles Assoc. Stored Products, I, 334.
Locality record: N.S.W., Trangie (J. Armstrong).

BY J. W. T. ARMSTRONG. 111

Note: Advanced larvae from old saddlery in saddler’s shop taken 30th Sept., 1948.
Adults emerged about 1st Dec.

ORPHINUM NEALENSIS (Blackb.)

Blackburn, T., 1903.—Trans. Roy. Soc. 8S. Aust., xxvii, 170.
(Crypterhopalum.)

Armstrong, J. W. T., 1943.—Proc. Linn. Soc. N.S.W., Ixvili, 63.

Type: In British Museum.

Type locality: Oodnadatta, S.A.

Synonymy: Crypterhopalum nealense Blackb., loc. cit.

Distribution: Oodnadatta, S.A., Bogan R., N.S.W. (Armstrong), 60 specimens on
myall (Acacia pedula), February, 1945.

NEOANTHRENUS MACQUEENI, Nl. Sp.
Types: Holotype in the author’s coll., paratypes in British and Australian Museums.
Type locality: Milmerran, southern Queensland (J. Macqueen).

DESCRIPTION.

Elongate-ovate, black, legs and antennae ferrugineous, densely clothed with scales,
mostly ashy-yellow, but some brown forming a pattern much as in N. parallelus Armst.
(Proc. Linn. Soc. N.S.W., 1941, 390, fig. 2), those on the ventral surface ashen, closely
and evenly punctate.

Pronotum: Transverse, sub-rectangular, convex, transversely depressed just in front
of base, sides marginate, straight and gradually converging for about three-quarters of
length, then rounded to anterior angles, these obtuse, posterior angles slightly acute, base
strongly lobed.

Elytra: A little wider than prothorax, one and a quarter times as long as wide,
shoulders rounded, sides slightly curved almost parallel for approximately two-thirds
length then evenly rounded to apex.

Size: 2-1-8 mm. x 1-0-9 mm.

Discussion: Six specimens, named after their captor, are close to N. parallelus
Armst., but that is a proportionately longer species and the sides of its prothorax are
slightly curved in the first two-thirds. WN. ocellifer Blackb. has the sides of the prothorax
slightly incurved, and is a larger species, differently coloured. In the descriptions of
N. parallelus and N. niveosparsus (loc. cit.) the term “emarginate”’ was applied to the
sides of the pronotum, whereas ‘‘marginate” was intended.

Distribution: Only known from type locality.

NEOANTHRENUS PARALLELUS Armst.
Type: In the author’s coll.
Type locality: Lane Cove, N.S.W.
Distribution: Lane Cove, N.S.W., National Park, Q. (Hacker, Dec., 1919), eight
examples in Queensland Museum.

CORRIGENDUM.

Proc. Linn. Soc. N.S.W., Ixxii, 1947, 297, lines 26 and 38:
For ‘‘Xylabosca”’ read ‘“Xylobosca’’.

hseiosa a 08

; un Niue

EA Re She print ‘

Sew rasieet hha eis RES asc eee
Ag ay tees Sis y riae eont pnen shen PLY OD ata 28 ee > 4 ; ey ¥

a

Calf ah Se Seat

ts MWe Ae ais aneds avr strane: i
Cy OATAAT AL BY to ABE Mea
phe ae ay attend ; ce
TK aap ighoean Kits ty, i

eR Ae SET ES ae

Proc. Linn. Soc. N.S.W., 1949. PLATE I.

Graptolites from Tallong and Shoalhaven Gorge.

’ 113

AUSTRALIAN RUST STUDIES. VII.
SOME RECENT OBSERVATIONS ON WHEAT STEM RUST IN AUSTRALIA.

By I. A. WATSON AND W. L. WATERHOUSE,
University of Sydney.

(With Plates iii, iv.)

[Read 31st August, 1949.]

INTRODUCTION.

It is now generally recognized that the only satisfactory way of controlling wheat
stem rust is by the use of resistant varieties. In this country, as elsewhere, the progress
of wheat breeding has been such that the disease no longer constitutes a menace in rust
liable areas to those growers who cultivate resistant varieties. To a large extent the
success of these breeding programmes can be attributed to the fact that in the countries
concerned, annual surveys of the stem rust flora have been made. These surveys have
two essential objectives. They aim firstly to classify and describe in terms of Stakman’s
standard set of differential varieties the physiologic races present in a particular area,
and secondly, they aim to disclose whether new rusts are arising which are capable of
attacking either the resistant varieties under cultivation or those that are valuable as
parents.

There is at present considerable information available in the literature on the
results to be expected from physiologic race surveys. Most of the data have been
collected in the United States and Canada, but from the work of Waterhouse a good
deal is known of the cereal rusts present in Australia since 1920. Although the range
of variability of P. graminis tritici in Australia has been much lower than in North
America, rust surveys form an essential component of the local wheat-breeding
programmes, and population studies are reported from time to time. Although no report
has been published since 1939, in this paper certain facts are summarized concerning
recent major changes in the stem rust flora, leaving fuller details to be published later.

RESISTANT PARENTAL MATERIAL.

When the varieties Hope, Webster and Iumillo were imported into this country they
formed the main source of resistance to race 34 of stem rust which had first appeared
in December 1925 (Waterhouse, 1929). The first two of these varieties—Hope and
Webster—were crossed with Federation shortly after they were received in 1926, and
Hofed and Fedweb resulted as commercially cultivated stem rust resistant varieties.

About the time these three parents were received, the New South Wales Department
of Agriculture imported from Kenya Colony three varieties which have been used widely
in crosses. Macindoe (1931) and Waterhouse (1938) have reported on their value as
rust resistant parents. They have been carried in University of Sydney Accession Books
as Kenya 743, 744 and 745. The New South Wales Department of Agriculture designations
for these are C6040, C6041, and C6042 respectively. It has been shown (Watson, 1943) and
Watson and Waterhouse (1945) that these three varieties are very different both morpho-
logically and genetically. Each has a single major gene for resistance to race 34 of stem
rust in Australia, but each gene is inherited independently of the other two. Kenya 743
and 744 are only partially resistant to flag smut while 745 is practically immune.
Differences also occur in grain colour, awn production and several other characters.

K

114 AUSTRALIAN RUST STUDIES. VII,

Until 1942, the field reaction of all three Kenya wheats was highly satisfactory, and
no difficulty was experienced with high temperature effects as reported in Canada
(Johnson and Newton, 1941). They all had a further advantage in that their physio-
logical resistance made possible the use of glasshouse tests, since the reactions of seed-
lings and adult plants were correlated. Seedlings of these three varieties were not
identical in their reaction to the same race of rust, and there were also differences in
field reactions. Macindoe regarded Kenya 743 as having a greater field resistance,
and he concentrated on using this variety in breeding. At the University of Sydney,
where seedling reaction to several races was determined, it was found that Kenya 743
was only moderately resistant to race 45. For this reason Kenya 745 was preferred, and
of course it possessed flag smut resistance as well. Kenya 744, although it possesses
resistance to race 95 of leaf rust, was rejected mainly on account of its very weak straw
and lower seedling resistance to race 34. This variety has, however, been used to some
extent in Western Australia and Queensland.

Eureka was the first resistant commercial variety to be evolved from the Kenya
wheats. Actually Macindoe named two strains, Eureka and Hureka 2, from the cross
(Kenya 743 x Florence)F, x Dundee. Eureka 2 is distinguished only with difficulty
from Eureka, but in general the former strain is later, with a stronger straw and with
less tendency to shatter. When released to growers they each appeared to have the full
resistance of the Kenya parent (Macindoe 1937).

CULTIVATION OF RESISTANT VARIETIES.

In the north-western areas of New South Wales, the resistant varieties Hofed,
Fedweb and the two strains of Eureka made an immediate appeal to growers, since stem
rust is a potential serious menace every season in this area. The Hurekas rapidly
increased in popularity on account of their rust resistance plus attractive agronomic
characters, including ability to produce high yields. From 1938 onwards, increasing
acreages were sown to the Hurekas, mainly at the expense of Ford, Nabawa and similar
varieties.

Prior to 1941 the resistance of the two Hureka strains was undoubted, for repeated
close observations made throughout the wheat belt failed to reveal the presence of stem
rust on them. Kenya 743, the resistant parent of Hureka, remained virtually immune
in our field tests at Cowra, Glen Innes, Richmond, Tichborne and Gunnedah.

Rust Reactions of Eureka in 1941.

As a result of the strong resistance that Eureka had shown up to this time, it was
surprising to find rusted plants of this variety in 1941. In that year Mr. J. A. O’Reilly
forwarded rusted Eureka plants collected at Gunnedah and Tamworth. He reported
that the attack was not general, only isolated rusted plants being found in these fields
of Hureka. The rusted plants Mr. O’Reilly forwarded had well-formed grain, and were
accessioned in the University of Sydney varietal collection and have since been
carried on.

When the rusts from these collections were received they were used to inoculate
seedlings of Kanred, Arnautka and Hinkorn. In addition they were put on to
seedlings of EKureka from several different sources, as well as on to seedlings from the
actual plants that had been rusted in the field. S.U. Culture 334, which was regarded
as the standard Australian culture of race 34, was used as a check on seedlings from
the rusted plants and on Hureka seedlings from Gunnedah grain. Eureka was known
to be highly resistant to this culture both in the seedling and adult plant stage at
65-70°F.

The reactions are given in Table 1.

These tests were carried out during the latter part of November and the early part
of December 1941. The reactions of Kanred, Arnautka and Hinkorn, which up to this
time had been regularly sown to identify the Australian races in the survey work, failed
to differentiate between these three rusts. On account of the increased temperature at
this time of the year, Kureka from Gunnedah gave a fully susceptible reaction to

a at

BY I. A. WATSON AND W. L. WATERHOUSE. 115

TABLE 1.
Rust Reactions Produced by Three Cultures on Several Selected Vartieties at
Summer Temperatures.

|
Number and Locality of Rust Culture.
|
|

Variety. | 334
7150 7159 Sydney
Tamworth. Gunnedah. University.
| 2 SSS 2

Kanred 3 3
Arnautka 3) 3 |
Einkorn a Me e : !
Eureka (seed from Cowra) 3.4 3b

5 Ff ;, Gunnedah) 3 + 3 3
Eureka 2 Se ve ie 5) or 3. =
Eureka seed from rusted plants 7150 3 +E = 3

pees 5 a 3 7159 = 3 + 3

culture 334. From Table 1 it is clear that the two rusts 7150 and 7159 showed their
ability to attack Eureka from Gunnedah and each was capable of attacking seedlings
from the plants on which it had been collected. Culture 334 could not be distinguished
from them, since at these temperatures all Eureka seedlings tested were susceptible
to all three rusts.

Since these tests were unsatisfactory as a means of separating the two new cultures
from 334, it was decided to compare the three cultures on seedlings of Hureka at low
temperatures during the autumn of 1942. The average glasshouse temperatures had
fallen and the reactions given in Table 2 were obtained.

TABLE 2.
Reactions of Eureka to Three Cultures at Low Temperatures.

Rust Culture.
Variety.
7150 7159 334
Eureka an 56 30 eal sult Boal
Eureka 2 ae a ae ; 1 and 3 + ; l and 3 + ; 1 and 3 +

In this test Eureka showed the same resistance that had been characteristic of it
in previous years and produced mainly tiny flecks with all cultures. Hureka 2, on the
other hand, gave seedlings which proved to be a mixture of resistant and susceptible
types. The only conclusion possible from the tests thus far was that Hureka 2 showed
heterozygosity for reaction to stem rust. As this could be easily overcome by selection
no serious view was taken of it. The cultures 7150 and 7159 were, however, retained as
refrigerated stocks for further testing.

Eureka Rust Reactions in 1942.

During the spring of 1942, Mr. O’Reilly again submitted rusted stems of Hureka.
He reported that on the property of Mr. A. B. Foxe, “Lyndhurst”, Narrabri, Eureka was
fairly uniformly infected by rust. One culture, 7316, collected from this field was treated
in the usual way and was placed on to seedlings of Kanred, Mindum and Hinkorn as
well as of Eureka. At this time it was compared with culture 334 and with 7150 and
7159—the two latter being from rusted Hureka in 1941. These tests were carried out
in the summer of 1942: the glasshouses temperatures approximated to an average of

KI

116 AUSTRALIAN RUST STUDIES. VII,

80°F. Under these conditions Eureka was susceptible to all four cultures 334, 7150,
7159 and 7316. Johnson and Newton (1941) had already demonstrated that Hureka was
susceptible at these temperatures, and indeed it was known that Kenya varieties in
general did not retain their resistance when the temperatures were unduly high
(Newton et al 1940).

At this stage the four cultures appeared so much alike that it was considered that
any differences between them would not allow a reliable separation to be made at these
high temperatures and no further tests were made on seedlings during the summer
months.

Despite the fact that the glasshouse tests had failed to demonstrate clearly that a
new rust had arisen, it was obvious from at least two sets of field data collected late
in 1942 that a new rust capable of attacking Eureka was present. The first evidence
was from yield trials carried out by the Department of Agriculture on the property of
Messrs. White Bros., Boggabri, and already reported by Macindoe (1945). In these
tests the sowings had not been made until August and the rust developed on the
susceptible varieties to such an extent that they scarcely produced any grain. (See
Plate iii.) The yields are given in Table 3.

TABLE 3.
Yields of Grain in 1942 Harvest at Boggabri.
|
Yield,
Variety. Rust Reaction. Bushels/acre.

Charter .. Ae a Resistant. 34
Celebration as 25
Yalta si 20
Eureka 2 as a4 Susceptible. 6
Eureka ss 3
Ford $8 1

The second lot of data came from our own field experiments made on the property
of Mr. C. H. Beeson, Gunnedah, in the same year. It was found that an entirely new
situation had arisen, in that rows of Eureka and Eureka 2 were fully susceptible. It
was obviously not due to heterozygosity in the variety, as no plants showed any resist-
ance. It must have been due to the occurrence of a new type of rust.

In these field tests it was usual to sow seed of the three Kenya strains 748, 744 and 745
together with the varieties of Stakman’s differential set. For the first time in our
observations, Kenya 743 was completely susceptible. | Its reaction was indistinguishable
from that of Eureka, and it was apparent that the new rust could attack each with
equal virulence. In contrast to this, Kenya 744 and 745 were just as resistant as they
had been previously.

The occurrence of this new rust at Gunnedah in 1942 gave the first opportunity to
assess the value of the other potential stem rust resistant parents. The following were
quite resistant; Hope, Hofed, Webster, Fedweb, Gaza, (Bobin x Gaza) F, x Bobin [Gabo],
(Double Cross x Dundee)F, x Dundee [Celebration], (Kenya x Pusa 4)F, x Dundee
[Yalta], Kenya x Gular [Charter], Dundee x Kenya [Kendee], Hochzucht and Iumillo.
The susceptible varieties such as Bencubbin, Ford, Gular and others did not appear to be
more heavily rusted than normally, even though they were exposed to an epidemic caused
in the main by this new rust. The most striking feature of the field result was the
complete susceptibility of Eureka. (See Plates iii and iv.)

Glasshouse Tests since 1948.

During the winter of 1943 when average glasshouse temperatures had fallen to well
below 80°F., the refrigerated stock cultures collected on Eureka in 1941 and 1942 were
tested on the differential varieties, using culture 334 as a check. In the first instance,
cultures 7316 and 334 were each placed separately on to seedlings and adult plants of

BY I. A. WATSON AND W. L. WATERHOUSE. - 117

Eureka. After incubation the seedlings were kept inside the glasshouse but the adult
plants were grown outside.

As mentioned earlier, seedlings of this variety reacted similarly to both rusts
during the summer tests. Winter tests at temperatures approximating to 65°F. indicated
clearly that these two cultures, 334 and 7316, were very different rusts. Seedlings and
adult plants of Eureka when inoculated with culture 334 gave the characteristic fleck
and type 1 reactions, but 7316 on this same variety gave only 3 + reactions. These
winter tests on plants at two stages of development gave full confirmation of the field
results and demonstrated with certainty that a new race of stem rust had arisen
capable of attacking Eureka at low temperatures.

In order to determine the particular physiologic race involved in this new attack
on Eureka, seedlings of the standard differential set of varieties were inoculated with
cultures 334, 7150, 7159 and 7316. The seedlings were raised in an artificially heated
room at approximately 75°F. during July, 1943. On the standard set of varieties at,
those temperatures the four cultures could not be distinguished with certainty,
although the reactions on Kubanka and Acme were such as to indicate that possibly
separations could be made on these varieties. In the main, however, the reactions
were similar to those given by the standard 334 culture.

Although this general similarity existed, there appeared to be certain anomalies
in the reaction of some of the differential varieties and it was decided to make a
comprehensive analysis of collections taken from various localities. A series was
studied consisting of the four collections mentioned above as well as further collections
from Gunnedah, Tamworth, Narrabri and Richmond, N.S.W. To these were added
collections from Werribee and Longeronong, Victoria, and a collection of race 34
obtained from the United States (“U.S. 34”).

These comparisons were carried out over a period of three years, and for purposes
of differentiation, Kenya 743 was added to the Stakman list of differential varieties.
This was essential, as on the standard set, with the exception of the U.S. 34, the
cultures were very similar. The most important difference between them was their
ability or inability to attack Kenya 7438. As a result of this analysis, which was
carried out both during winter and summer, certain differences showed up on these
varieties. They can be summarized as follows:

(a) Kubanka. During the early stages of these critical comparisons it became
obvious that the Kubanka reactions were not similar to those previously recorded
for culture 334. A check on this rust reaction was made using Kubanka seed from
all possible sources. Since all seed lots gave the same result, fresh seed was requested
from Dr. BE. C. Stakman. However, on plants from the new sceed as well, culture 334
gave either flecks or x reactions or a combination of both instead of the usual 3 +
reactions. Repeated comparisons showed that culture 7316 also gave this reaction on
Kubanka and was indistinguishable from 334. This variety when tested with the
United States race 34 under the same conditions proved fully susceptible. All other
rust collections in the series gave Kubanka reactions which were not consistently
different from those of 7316 and 334.

(b) Acme. Cultures from Edgeroi, Narrabri, Richmond, Gunnedah, Tamworth .
(N.S.W.) and from Longeronong and Werribee (Victoria) were compared on the variety
Acme. There was considerable variation, and gradations between high and low reactions
on this variety were obtained. Cultures 334, 7150, 7159 were characterized by a mixed
reaction, the pustules being mainly of type 8 together with some of type 1. Repeated
separations showed that only one rust was producing this mixture. Culture 7316 and
several others showing Kenya 743 susceptibility also gave a mixture of type 3 and type 1
pustules, but the majority were of type 1, the overall reaction type being one of
greater resistance. Here also attempted separations demonstrated that only one rust.
was present.

These differences were consistent though slight. They could be decided upon with
some confidence when pure cultures were being compared, but mixed cultures could

118 AUSTRALIAN RUST STUDIES. VII,

not be determined with any degree oi certainty. In view of this, Acme was considered
of little use as a variety for differentiation.

(c) Arnautka, Mindum and Spelmar. It has already been pointed out that these
varieties are resistant to culture 334 at low but susceptible at high temperatures
(Waterhouse, 1929). All three varieties have reacted similarly in previous studies with
Australian collections of rust. During the late winter and early spring these varieties
appear to be mixed for rust reactions, giving the x type, but the “mixture” is in fact
due to the process of change from resistance to susceptibility. The cultures used in
this study, and others collected throughout the wheat belt, all showed this variable
reaction on Arnautka, Mindum and Spelmar, and confirmed the earlier work. As a
rule it was found that Arnautka showed less variation to these changes in temperature
and became susceptible at temperatures at which the other two varieties showed partial
resistance. Slight differences were found between certain collections in their ability
to attack these varieties, but it was impossible to use them as a safe basis for
differentiation. In contrast to this, the United States collection of race 34 gave the
characteristic 3 + and 4 reaction on these varieties at all temperatures.

(ad) Kenya 743. The local cultures of rust could be readily placed into two classes
on their ability or otherwise to attack Kenya 743. The U.S. race 34 was intermediate
in its reaction on this variety, and necrotic areas appeared around the large pustules
that developed. This reaction was quite unlike that of 7316. (See Plate iv.)

A typical set of reactions obtained on the varieties is shown in Table 4.

TABLE 4,
Reaction of Differential Varieties to Cultures 334, 7159 and 7316.

Typical Reaction to Rust Culture.
Variety. |
C.I. Number. 334 7159 7316
Little Club ae abe 4066 4 4 4
Marquis... Bye ie 3641 4 4 4
Kanred re as cs 5146 3) ae 3+ 38+
Kota ae ay a0 5878 3+ 3+ 3 +
Arnautka .. Se a 1493 x +, 4 x +, 4 x +, 4
Mindum .. pal a 5296 x, 4 x, 4 xan:
Spelmar .. Me ne 6236 x, 4 xa: 4:
Kubanka ats ah 2094 Bb SEX. 2X:
Acme AiG a A 5284 3 and 1 3 and 1 1 and 3
Kinkorn .. a ae 2433 : ; p 2
Emmer Ben Sis oo 3686 ; : :
Khapli Ne 5h i 4013 : : ;
Kenya 743 NR Be — : 5 3 +
Kureka a ate of = : ; 3 ar
|

These reactions reveal the tendency for Arnautka to be more severely rusted than
Mindum or Spelmar. The Acme reactions are shown as a mixture of pustule type with
all three cultures. With 7316 Acme had a predominance of the 1 type reaction while
with the other two cultures it showed a greater frequency of the 3 type reaction.

In view of the fact that Kenya 743 at low temperatures served to distinguish the
two groups of rusts as represented by 334 and 7316, tests were made to determine
whether other varieties would also serve this purpose. It was equally important to
find a variety that would separate the 334 and 7316 types at summer temperatures,
when the bulk of the rust survey work is done. Over 80 varieties selected from
earlier work for their seedling resistance to 334 were tested as seedlings at two
different temperatures with cultures 334, 7159 and 7316. The range of temperature
in one house was from approximately 60°-65°F. and in the other 70°-75°F. The
rust reactions in general were higher at the higher temperatures. It was found,

BY I. A. WATSON AND W. L. WATERHOUSE. 119

however, that the varieties could be classified according to their reaction type as
shown in Tables 5, 6, 7 and 8.

TABLE 5,
Varieties showing Fleck Reactions only when inoculated with Cultures 334, 7159 and 7316.

8.U. 5 Siu. S.U.
No. Name. No. Name. No. Name.
|

277 Gaza. 758 Beladi 129. 905 Portugal 24,
745 Kenya. 724 Marquillo. | 908 Portugal 60.
752 Beladi &5. 791 Poona. 911 Portugal 86.
753 Beladi 114. 806 Pinet. 913 Portugal 90.
754 Beladi 141. 880 Egypt 7. 916 Portugal 100.
755 Beladi 31. 893 Egypt 50. 945 Morocco 34,
756 Beladi 98. 896 Egypt 59. 1229 Rhodesian.
757 Beladi 132. 904 Portugal 3.

TABLE 6.

Varieties showing Flecks with some and X Reactions with others of the Three Cultures
334, 7159 and 7316.

$.U. 8.U.

No. Name. No. Name.
321 Palestine durum. 794 Poona 806.
668 Pinet. 795 Poona 809.
676 Russian. 797 Poona 808.
687 Trigo africano. 827 Russian 2995.
717 Vernal Emmer x Iumillo. 832 Russian 1309/2.
726 Nodak. 870 Greece 11.
728 Palestine. 882 Egypt 14.
741 Beladi. 889 Egypt 31.
751 Africano. 891 Egypt 45.
759 Beladi 26. 895 Egypt 56.
760 Bianrollo 20. 906 Portugal 24.
764 Covelle. 914 Portugal 98.
770 Greek 10. 939 Traq 11.

774 Hordeiforme. 1116 Margerito 11.
790 Nodak. 1227 Hochzucht.
792 Poona 804.

These varieties classified in Table 6 could probably be regarded as having physiologic
resistance to all three cultures. Under the conditions of fluctuating glasshouse tempera-
tures it was found that with any one variety there was a considerable range of

TABLE 7.
Varieties having Moderate Seedling Resistance to
the Three Cultures and Designated 3=C.

S.U. No. Name.
744 Kenya.
1228 Egypt NA965.
1231 Sabanero.
1238 Rhodesian.
1240 La Estanguela.
1313 Fronteira.
1314 Frondosa.
1315 Supreza.

120 AUSTRALIAN RUST STUDIES. VII,

reaction type. All varieties that appeared to have possibilities for the separation of
these cultures were tested during the summer of 1945 but none proved satisfactory as
a differential.

Another group of these varieties is shown in Table 7. Included in this list are
those lacking the physiologic resistance but nevertheless having some resistance to
the cultures. ;

In Table 8 are shown those varieties which at low temperatures served to
differentiate the culture 7316 from 334 and 7159. While these varieties served that
purpose, none of them was of any use at high summer temperatures: then they were
all completely susceptible.

TABLE 8.
Varieties which at Low Temperatures are Susceptible
to Culture 7316 but Resistant to Cultures 334 and 7159.

|
|

5.U. No. Name.
743 Kenya.
1232 Red Egyptian Special.
1233 Red Egyptian.
1234 Birdproof.
1235 Talberg.
1236 Rhodesian.
1237 ty
1239 5
1311 Eureka.

From these studies it is clear that no variety has been found which could be
used to separate the 334 and the 7316 rust types with any degree of certainty during
the summer months. Work has been in progress for some time with selections of
T. Timopheevi, and it is hoped that a satisfactory differential may be found among these.

DESIGNATION OF THE CULTURES.

It has been customary in the past to relate the reactions of any particular rust
to those that have been summarized and tabulated by Stakman and Levine (1944) for
their differential hosts. Such a procedure has not proved entirely satisfactory as far
as the Australian collections are concerned. The difficulty always arises of having
some relative standard with which to compare and contrast in detail rusts occurring in
different countries. Until some international clearing house is established in which
detailed comparisons can be made under identical conditions, we must continue to
assume that a good many of the large number of races already described are sufficiently
alike to be given the same race number. We must assume too that many of the rusts
designated as the same race by independent workers in different countries are
sufficiently dissimilar to be given the status of separate races. There is abundant
evidence for the existence of biotypes within the described races, and one must not™
assume that varieties resistant to race 34 in Canada will necessarily give that same
reaction to race 34 in Australia. For this reason, from the international viewpoint, the
present results from physiologic race surveys must be interpreted with extreme caution.

It is well known now that the breeding of varieties resistant to stem rust is not
nearly so complex as might be expected from the large number of races that have
been described. One of the reasons for this simplification is that races tend to fall
naturally into groups, and one major gene frequently governs resistance to all members
of a group. Stakman and Levine in an effort to draw attention to possible affinities
between the races, show in their key the races which may be related. For example,
they show race 34 to bear some relation to races 63, 77 and 126. Races 56 (which has
been recorded in Australia), 125 and 126 are related to race 127. Races 34 and 127
show affinity with race 126.

BY I. A. WATSON AND W. L. WATERHOUSE. 121

A consideration of the similarities between these races in the light of the patho-
genicity of cultures 334 and 7316 will probably help in the most appropriate designation
of these latter cultures. For this reason the respective pathogenicities recorded for all
nine are given in Table 9. The reactions of the described races are in this case taken
from the table of Stakman and Levine (1944).

: TABLE 9.
Reaction of the Differential Varieties to Seven Described Races and to Cultures 334 and 7316.
s :
= ; g ao g S :
Race or — | zr : = = S ‘S < 5 es ca

Culture No.| = 2 elas (tee SS & z = S Ss # = =

= a e S Es = eS = 5 = = r=

4 e 4 I < = Rn Na 4 ea es i

|
T 34 44 4 — re | 4 4= 4 = 4+ NSE i 0; 1+
r 56 4 3+ 3i = 3045 1 1 1 3 4 3 4 1 | 1 leeeeel
r 63 4 +4 Xo 4 3 ap 4+ Aaa 4+ 4+ 4n + i 1= OF
ar + qr |
me Eh 4 4 3 = 3 - 3e 3c BiG 4 — 3+ 1+ xs il
r 125 Be 44 4 4 4 0 0; 05 5X. 4 t= | 0 il =
r 126 | 4= 4 = 3 4 3+ xt ate x + x — Le | l= 1
| | |

Teles la AN |e A See a ft Py ie x 0; 0; 1 —
Culture 334 | 4 4 3 + 3 + |x +,4 oe, Lh x, 4 Gee (83: enael All) 8 laos $
Culture 7316| 4 4 | Bak Ise dh aie Al Se a aox \|fand 3": lee ;

When comparing these reactions with those observed during the course of this
study, race 63 can be eliminated from the list since at no time did the present cultures
show any tendency to produce an X reaction on Marquis. Similarly all cultures have
consistently shown the typical, or type 1, reactions on Emmer. Race 77 can therefore
be excluded. In all previous work done here, culture 334 showed the ability to attack
vigorously Kubanka, and with its other reactions, was appropriately designated race 34.
Much data, accumulated from 1926 to 1941, confirmed this reaction type many times.
The results reported here indicate that sometime during or just after 1942 a change
occurred in this rust so that it now gives a resistant or at high temperatures a meso-
thetic reaction on Kubanka. It is apparent that culture 334 can not now be designated as
either race 34 or 56. There remain to be considered races 125, 126 and 127.

As indicated earlier, it has been demonstrated that temperature is a very important
factor to be considered in the determination of rust reactions (Waterhouse 1929). For
this reason determinations have been made at a temperature as close to 70°F. as prac-
ticable. This is essential since the Australian culture 334 which up to 1941 has been
called race 34 is very sensitive to low temperatures. At temperatures below optimum,
Arnautka, Mindum and Spelmar are resistant to this culture, and the reaction type is
that of race 56 (Waterhouse 1929). This character of the Australian rust is absent
from the United States race 34 (Waterhouse and Watson, 1941).

In the above table showing the reactions of races 125 and 127, Arnautka, Mindum
and Spelmar show a resistant reaction. Cultures 334 and 7316 show these reactions only
at low temperatures, and as the temperature rises, X reactions change to X+ and finally
to the 4 type. As this susceptibility is not typical of either of the races 125 and 127
they can safely be eliminated from the list. When these eliminations have been made,
it appears that of the races described by Stakman and Levine, race 126 best fits the
reaction types of the cultures that have been examined in these studies.

There are certain anomalies to be explained before giving the Australian cultures
this designation. For example, race 126 is tabulated as showing X reactions on
Arnautka, Mindum and Spelmar whereas the conditions under which these cultures

1272 AUSTRALIAN RUST STUDIES. VII,

produced that reaction must be regarded as sub-optimal. The fact that they showed
the range from ; to 4 with changing temperatures, whereas the United States race 34
consistently gave 3+ to 4 reactions under the same conditions, seems to warrant their
designation as different races. To call these cultures race 126 would necessitate regarding
the x reaction as an average of those found with varying temperatures, and this seems
a satisfactory compromise.

Another difficulty is that Acme shows a variable reaction to these cultures, type 1
and type 3 pustules being mixed on the same leaf. This reaction is not typical of the
mesothetic reaction since the full range of pustule type does not occur. There is no
recognized reaction which adequately describes this condition. The X, however, probably
describes it as well as any, and this would be a satisfactory classification for all prac-
tical purposes. While admitting these two anomalies, it is considered that these cultures
can best be called race 126.

In order that some degree of international co-ordination could be arrived at in the
description of these cultures, Dr. Stakman kindly agreed to test certain material. In
1947 uredospores of culture 7316 were forwarded to him by air mail. This is identical
with 334 on the differential set, except for the Acme reaction. Although he carried out
two separate tests, the glasshouse conditions at the time were not good and the following
reactions were obtained:

Variety. Infection type
Marquis ae 34 4 — 4
Reliance .. aie 4 — 384+
Kota ee) ae “4 — 3+
Arnautka .. ait 4 —c¢ x+
Mindum ... ee x — xe
Spelmar .. At x x=
Kubanka .. ae x — x =
Acme thy Sn 3, 4-br 3¢br
Einkorn.. Ey, 0; bl 0; bl
Emmer ae ke 0; bl 0; bl]
Khapli ae ae Op, al, lol 2.

br indicates browning.

bl + blasting.

He remarked that on account of the browning observed on Acme and the tendency
to blast part of the leaf as on Hinkorn, Vernal Emmer and Khapli, the rust does not
appear to agree with their race 34. The reaction types he gives are very close to race
126 except for that on Acme: the reaction of this variety was complicated by browning.

From the evidence presented here and from the United States results, it appears
that culture 334 can now be listed as race 126. Since culture 7316 can only be differen-
tiated satisfactorily from 334 on varieties outside the standard set, it would be convenient
and satisfactory to regard the present Australian Eureka-attacking rust as race 126 B.

Comparison of 7316 with the United States Race 34.

Although independent workers engaged in rust survey working in different continents
may identify the same race from among their rust collections, it is now well known that
they may be dissimilar. Biotypes of existing races can be readily detected by the
inclusion of further differentials in the recognized set of 12 varieties. The authors
have found (Waterhouse and Watson, 1941) that an American collection of race 34
could be distinguished from an Australian 34 by incorporating into the differential set
the varieties Marouani, D5, Nodak, Pinet, Trigo africano, Egypt 75 and Greece 18.
Inbred lines of rye also separated these two rusts; and the temperature sensitivity of
the Australian 34 was a further important contrasting character.

When the detailed comparative tests were reported in 1941, the 7316 culture had
not been collected. About this time, however, Macindoe, working at St. Paul, Minnesota,
U.S.A., found that Eureka was susceptible both as seedlings and adult plants in the field
to the American culture of race 34 with which he worked. He showed that it was a
susceptibility which was not the result of high temperatures as reported by Newton
and Johnson (1941). Since this reaction of Eureka in the United States was so unlike

9

BY I. A. WATSON AND W. L. WATERHOUSE. 123

that of the New South Wales reaction, consideration was given at an early stage to a
possibility that American uredospores may have gained entry to this State and that
7316 was nothing more or less than a biotype of United States race 34. Repeated
comparisons of the three cultures 334, 7316 and U.S. 34 were made during the course of
the work. Reactions on certain of the differentials were compared more often than
others but Table 10 is an average of the reaction types:

TABLE 10.
Comparison of Reactions of Cultures 334, 7316 and U.S. 34.

Culture.
Variety. |
334 7316 | U.S. 34
|
| |
| |
Little Club | 4 | 4 4
Marquis | 4 4 3+
Kanred ul 3+ | 3+ 3+
Kota bi oh a) ae | 3 + 34+
Arnautka x-+, 4. | x +, 4 4
Mindum x 4 mg, us 4
Spelmar xan 4: | x 4 4
Kubanka .. -- | RaD:< 5x 34+
Acme oe ai ||| 3 and 1 | 1 and 3 | ict
Emmer we | : | ; | 3
Einkorn.. | : | ; ;
Kap ivees ees sees |
Bureka.. all ; | 3+ 3cn
Kenya 743 a | ; | 3+ gen

From these results it is clear that 334 and 7316 are separable only with difficulty
on varieties other than Eureka and Kenya 743. Both cultures can be readily distin-
guished from U.S. 34, however, by their reactions on Kubanka and Acme. Hureka gave
a characteristic reaction to all three rusts. It was resistant to 334, fully susceptible to
7316, and intermediate in its reaction to U.S. 34. The large pustules produced by this
rust at either low or high temperatures were surrounded by heavy chlorosis and
necrosis. (See Plate iv.) In this regard, U.S. 34 was distinct from any of the cultures
with which we had worked previously. Apart from the reactions on the differential
varieties, the Eureka reactions to these rusts made it unreasonable to suppose that
U.S. 34 had taken any part in the alteration of Eureka resistance.

Field Reactions of Eureka in 1946.

When new rust-resistant varieties are released and suffer the fate of Eureka, the
accusation may be made that the variety was never fully resistant to rust. The build-up
of the 7316 type of inoculum in north-western New South Wales has been such that
Eureka is probably the most susceptible variety grown in that area at the present time.
Indeed, it is not easy to realize that this variety was one of the most resistant grown in
these same areas prior to 1941.

Despite this change in N.S.W., Eureka has maintained its resistance in those areas
where only the old type of rust occurs. It has been reported that Eureka is still
resistant in Western Australia (Thomas, 1948) and except for one suspicious case in
1947, only the 334 type has been recovered from that State. Our own observations many
times confirmed the high degree of resistance which prior to 1941 had been shown by
Eureka, and this has been observed again in recent years. At Killara in 1946-47 observa-
tion plots of Pusa 4, Cailloux, Gabo, Eureka 2 and Charter were grown. A severe rust
attack developed on Pusa 4 and Cailloux and they suffered extensive damage. Despite
the widespread occurrence of the 7316 type in the County of Cumberland, Eureka 2,
Gabo and Charter remained completely free from rust. In the latter part of the summer
season identifications were made of the stem rust present in this plot. Only the 334

124 AUSTRALIAN RUST STUDIES. VII,

type was found. Under the conditions of this test where only the old type of inoculum
occurred, Hureka was quite as resistant as it had been when it was first released to
‘growers in 1988.

ORIGIN OF THE 7316 TYPE.

In the stem rust areas of North America it is not unusual to find new races of rust
arising from time to time. In many cases when such new races have been collected
their origin has been correlated with the presence of barberry bushes in the neighbour-
hood. Since it is well known that new races are produced by the crossing or selfing of
certain races on this alternate host, barberry infection is a very satisfactory explanation
for the occurrence of pathogenicity changes. It is difficult to advance this suggestion
as a means of explaining the origin of the 7316 type in N.S.W. Barberries are known
to occur on the tableland areas of this State, but on only one occasion have they been
reported as naturally infected by rust (Waterhouse, 1934). While this new 7316 rust
could have arisen in this way, it seems on circumstantial evidence most unlikely.

It must be conceded that as far as we know, new cereal rusts are continually arising
without the sexual stage on an alternate host. In the case of leaf rust (P. triticina),
for example, over 100 races have been recorded for this organism (Johnston et al, 1942),
but field-infected plants of the alternate host, Thalictrum spp., occur only very rarely,
and such have never been observed in Australia. For the present it must be assumed
that something in addition to hybridization is involved in the formation of new races.

We could conveniently postulate that these new races arise by mutation. However,
there is insufficient evidence to explain the process by which these mutations arise. It
is known that changes in the characters of fungi can be induced by incorporating
chemical substances into the substrate, but to date no such treatment has been possible
with the cereal rusts. Mutations both for pathogenicity and for spore colour have
been observed here as elsewhere, and their importance in explaining the variability of
rust fungi has been shown (Newton and Johnson, 1944). However, the causes responsible
are still unknown. Gene mutations may be involved in these changes, or they may be
associated with a rearrangement of the dicaryotic nuclei as suggested earlier
(Waterhouse, 1929).

As a result of the work by Rodenhiser and Hurd-Karrer (1947) it is clear that
hyphal fusions may take place between vegetative (dicaryotic) hyphae, but there is
no evidence yet that new races would result. Should contrasting types be brought
together by new associations of old nuclei in a dicaryotic mycelium, the “hybrid” may
exhibit enhanced pathogenic capabilities. It is probable that even if hyphal fusions
do occur within the host plant, a good many of the new combinations would not differ
pathogenically from the old. When one of us (I.A.W.) was engaged in extensive race
mixing experiments (Watson, 1942) a close watch was kept for the occurrence of
new races either by mutation or otherwise, but no changes in pathogenicity were
observed. Again, in earlier work by the other author, races 43 and 45 were kept in
association for 18 successive generations without any change being discernible. In
spite of the difficulty of explaining how a mutation has produced the culture 7316, it
seems more likely that it has arisen in this way than by hybridization on the barberry,
since this process is so rare in this country.

The mechanism concerned in the formation of these new types is important because
it is fundamental to the general problem of variability in fungi, and in this way it is
closely associated with breeding for disease resistance. The idea that certain species
may act as bridges, enabling pathogenicity to be built up, was initiated at a very early
stage of the work on specialization in fungi. However, it was dropped quickly when
physiologic races were sorted out and were regarded as stable entities fixed in their
pathogenic capabilities.

At the present time there are many happenings which lead to the belief that they
are not always very stable. It seems that the work of Reddick and Mills (1938) on
Phytophthora infestans has revealed certain hazards in the cultivation of varieties with
a moderate degree of resistance only. From this work they found that the virulence

BY I. A. WATSON AND W: L. WATERHLOUSE. 125

of the pathogen was increased by successive passages through certain varieties ranging
from somewhat resistant through considerably resistant to usually immune. In the
field, previously immune varieties became infected as a result of this progressive
increase in virulence.

If the result obtained with potatoes could be applied to wheat varieties infected
with stem rust, then it is easy to visualize certain varieties acting as satisfactory
bridging hosts, since many possess only a moderate field resistance. In 1944 a small
experiment was carried out to determine whether in fact seedlings of Hureka could
serve as a bridge to build up the virulence of culture 334 so that it could attack
Eureka. Four single pustules of this culture were taken, two were increased on
Federation and two on EHureka. Those on Federation. were kept at low temperatures
(60-65°F.) and those on Eureka at high temperatures (75—-80° F.) so that the seedlings
of this latter variety produced sufficient rust to enable sub-cultures to be made. After
the rust had produced each generation of uredospores, those on Federation were
transferred back to Federation, those on Hureka back to Hureka. These transfers
were continued on the duplicate pots of Eureka and Federation for six successive
generations. During the course of the experiment there was no sign of any increase
of virulence on Hureka. After the period of six generations, which occupied almost six
months, the cultures from Eureka and from Federation were compared at low
temperatures on Hureka. Those grown on Hureka for this period showed no greater
pathogenicity for this variety than those that had been grown on Federation. The
experiment proved of no value in helping to arrive at an explanation of the origin
of 7316.

With the limited amount of information available it is possible only to speculate as
to the origin of the 7316 culture. There is also no information to help in explaining
what has occurred in culture 334. The change in this latter rust is all the more
perplexing because not only has the glasshouse culture altered from race 34 to race 126,
but a similar change has been shown by all rusts that we have collected in the field
in recent years. It is most difficult to visualize any agency so far-reaching in its
effects that both glasshouse and field cultures would mutate at about the same time
from the old to the new rust. Nevertheless the fact remains that sampling widely in
the State gives in the main two rusts. One is of the 334 type, which cannot attack
Eureka and which we have called race 126. The other, which is of the 7316 type, occurs
on EKureka and we have designated it as race 126B.

THE OCCURRENCE OF NEw RUSTS IN RELATION TO THE BREEDING PROGRAMME.

When new physiologic races of stem rust arise, it is essential to know how potential
parents will react when tested with them. Observations since 1942 have indicated
that several of the well-known varieties, such as Hope, Webster, Iumillo, Gaza and
Khapli, are fully resistant to the 7316 type in the field. The chief undesirable agronomic
characters of Hope, Webster and Gaza have been removed and their resistance factors
are available in Hofed, Fedweb and Gabo respectively. In addition to these (Nabawa
<x Hope) [Warigo] and (Double Cross x Dundee)F, x Dundee [Celebration] have had
satisfactory field resistance to rust. The other varieties (Kenya x Gular) [Charter],
(Kenya x Pusa 4)F, x Dundee [Yalta] and (Dundee x Kenya 745) [Kendee] have all
shown resistance to all local races, both as seedlings and as adult plants. While the
seedlings of these latter varieties have reacted similarly to 7316, they do not all behave
alike in the field. On the basis of the 1-6 classification they could be ranked thus:

Charter 1

Kendee 1-2

Yalta 2-3
Eureka for comparison would be 5-6. Yalta is characterized by showing in the field
some plants typical of class 4, and this variety seems least resistant of those developed
in recent years.

The three varieties Charter, Kendee and Yalta have shown no field complications
resulting from the effects of high temperatures, and it is felt that considerable confidence

126 AUSTRALIAN RUST STUDIES. VII,

can be placed in the Kenya varieties as rust-resistant parents under Australian
conditions. It is, of course, to be expected that certain environments will be found
in which their resistance will break down. In 1942 a severe attack of downy mildew
(Sclerospora macrospora) in certain of the resistant varieties at Tichborne was
associated with complete susceptibility to the 334 type of rust. Heads and stems of
the varieties so affected are shown in Plate iv. The exact nature of the interaction
between host and pathogen is not known, but it is well established that infection by
one disease may lead to increased susceptibility to a second disease.

As mentioned earlier, the breeding work in this State has involved extensive use
of the Kenya varieties, and the three designated as 7438, 744 and 745, which were the
first available here, have been used most. Since the first introductions were made,
further samples from Kenya have been received. In addition, Dr. Margaret Newton
forwarded seed of a comprehensive list of the strains they had studied. Included
with this group was also seed of ‘“McMurachy’s Selection”’.

These latest Kenya strains have been tested extensively with cultures 334 and 7316
in the glasshouse: the reactions are given in Table 11.

Prior to the occurrence of the 7316 type in 1941-2, these Kenya varieties appeared
to provide a wealth of stem rust resistant material. Nothing was known of the genetic
relationships between them except that Kenya 743, 744 and 745 each had independent
factors for resistance. The arrival of the 7316 type has, as the results in Table 11 show,
completely changed that position, and the majority of the Kenya strains must now
be regarded as susceptible to rust in this country. They can be divided clearly into two
groups, those that are resistant to both 334 and 7316 types and those that are resistant
only to the 334. A good many of these Kenya strains were received without names,
so that they can only be identified by their Sydney University Accession Number. In
certain cases these numbers can be linked with a Rust Laboratory Accession number.
Among the varieties listed, “McMurachy’s Selection” appears to be susceptible to 7316
and it segregated for resistance to culture 334. Kenya 117A, which has been found
resistant to race 15B in the United States (Loegering, 1944), was resistant to culture 7316.

Culture 334 was used to inoculate F, plants of several crosses designed to get some
information on the relationship of the resistance genes present in these Kenya varieties.
Some of them were crossed among themselves and some were crossed with a Dundee x
Kenya 745 selection known to have inherited the resistant gene from Kenya 745. As
would be expected, certain crosses showed segregation for susceptibility in F., others
did not. If no segregation was obtained in several hundred plants, it was taken as
good evidence that the two parents had the same factor.

The following crosses did not show any segregation in F,:
Kenya 743 x Kenya 1037. E
Kenya 743
Kenya 743
Kenya 745
Kenya 745

x Kenya 1304 (RL1373) (Minn. 2693).
x Sweden 1230.
x Kenya 1035.
x Kenya 1049 (RL I/35-2).
Kenya 745 x Kenya 1053.
Kenya 1034 x Kenya 1037.
Kenya 1034 x Kenya 1304 (RL1373).
x
x
x
x
x
x

Kenya 1035 Kenya 745.
Kenya 1035 Kenya 1053.
Kenya 1037 Kenya 743.
Kenya 1049 Kenya 745.
Kenya 1049 Kenya 1053.

Kenya 1053 Kenya 745.

Kenya 1053 x Kenya 1035.

Rhodesian 1229 x Kenya 745.

Kenya 1304 (RL1373) x BHureka 1311.

(Kenya x Gular) [Charter] 1345 x Kenya 745.
(Kenya x Gular) [Charter] 1345 x Kenya 1053.
(Gullen x Gaza)F, x Gullen 1368 x Kenya 745.
Gabo x Kenya 745.

Gabo x Kendee.

Kendee x Gabo.

BY I. A. WATSON AND W. L. WATERHOUSE. 127

TABLE 11.
Seedling Reactions of a Selected Group of Kenya Varieties and ** MeMurachy’s Selection” to Cultures 334
: and 7316 at Temperatures of 65-70° F.

Reaction to

5.U. Name of Variety or Designation Given af
Accession by Other Workers.
NOD ls | | 334 7316
|

743 C6040 : 3 +
744 | C6041 i | 3)
745 C6042 5 | 5
1034 5 | 3+
1035 : | ;
1037 : 3)
1049 RB :
1053 5 :
1304 RL1373 : Ba
1347 117A :
14S 5 | 2010 Ti AVANT. :
1349 291 J iB 1 Seg 3
1350 290 Jets Taal : 3 +
1351 192 Q 2A (L) ; By
1452 RL 1/513 Seg. 3+
1453 — “RL 1/40-32 : 3+
1454 RL 1/40-33 3 = 3 =
1455 RL 1/40-34 Seg. Pp
1456 RL1387 C9906 3 3
1457 RL1388 C9965 ; 3
1458 RL1356 C9968 : 3
1459 RL 1/40-35 C9968 i 3
1460 RL1352 C10852 : :
1461 RL 1/40-36 C10854 5 p
1462 RL1379 C10857 g @ ar
1463 RL 1/40-37 C10857 ; 3+
1464 RL I/40-39 C10860 S 3
1465 RL 1/40-40 C10861 A 3
1466 RL 1/40-41 C10862 5 ueste
1467 RL1362 C10862 2 3+
1468 | RL1361 C10863 5 5
1469 RL 1/40-42 C10864 ¢
1470 RL1370 C10865 . 3
Ayre =) RL 1/40-43 10865 ; 3
1472 | RL1358 C10866 3 3+
1473 | RL 1/40-44 C10866 : 3
1474 | RL 1/40-45 3 3
1475 | RL 1/40-46 3. AL 3+
1476 RL 1/35-2 sea alt aul
1477 | 112A RL 1/35-4 ae il 3
1478 122.D.1.T.(L) Minn. 2693 RL1373 ; 3+
1479 | 117E.16.B.1 2694 RL1374 : 3+
1480 | 117B.5.B.2 Pr 2695 RL1375 Seal 38+
1481 117K.16A(L) A 2697 RL1376 ; 3 -—¢ 3 ot
1482 | 117 1.5¥(L) of 2696 RL1377 3; 3 — 3 +
1483 112E 19 J(L) RL1873 ; 1+
1484 60B12 B 16(L) RL 1/34-3 Di ;
1485 | 112B (15) L RL 1/34-5 sa! 8 ar
1486 | 131 C 5.E RL 1/34-2 : 3
1487 58F (L.1) RL 1/34-4 3 — 3+ | 3 4
1488 McMurachy’s Selection RL1313 Seg. 3+

Segregations were found in these crosses:
Kenya 743 x Kenya 1053.
Kenya 743 x Gabo.
Kenya 744 x Kenya 743
Kenya 744 x Kenya 745.
Kenya 744 x Kenya 1035.
Kenya 744 x Kenya 1049 (RL I/35-2).

128 AUSTRALIAN RUST STUDIES. VII,

Kenya 744 x Kenya 1053.

Kenya 744 x Rhodesian 1238.

Kenya 744 x Kenya 1349 291 J1B1.
Kenya 744 x Kenya 1351 192 Q2A(L).
(Federation x Kenya 745) x Kenya 743.
Kenya 745 x Rhodesian 1238.

Kenya 745 x Kenya 1349 291 J 1B 1.
Kenya 745 x Kenya 1351 192 Q@ 2A(L).

(Federation x Kenya 745) x Sweden 1230.
Kenya 1035 x Kenya 744.

Kenya 1037 x Kenya 745.

Kenya 1037 x Kenya 1035.

Kenya 1037 x Kenya 1049.

Kenya 1049 x Kenya 744.

Kenya 1049 x Rhodesian 1238.

Kenya 1053 x Kenya 744.

Kenya 1053 x Rhodesian 1238.

Rhodesian 1229 x Kenya 744.

Rhodesian 1238 x Kenya 744.

Rhodesian 1238 x Kenya 745.

Kenya 1034 x Kenya 745.

(Kenya x Gular) [Charter] 1345 x Kenya 744.

Kenya 117A 1347 x Kenya 744.

Kenya 117A 1347 x Kenya 745.

Kenya 1349 x Kenya 744.

Kenya 1349 x Kenya 745.

Kenya 1350 x Kenya 744.

Kenya 1350 x Kenya 745.

Kenya 1351 x Kenya 744.

Kenya 1351 x Kenya 745. :

(Guillen x Gaza)F, x Gullen 1368 x Kenya 743.

(Gullen x Gaza)F, x Gullen 1368 x Kenya 744.
In considering the group of crosses in which no segregation was observed, it
_ appears that two series of varieties are represented. The first series contains Kenya 743
and its related types. Kenya 1304 (RL1373, Minn. 2693), which, as shown in Table 11,
gave the same seedling reaction as Kenya 743, apparently has the same gene. Sweden
1230 behaves similarly and, as would be expected, Eureka 1311 belongs in a group
with 743. Two other strains, accessioned as 1034 and 1037, both have a single factor
for resistance to culture 334 and apparently in each this is the 743 gene. It is
obvious that no variety in this group is of any value in contributing to the problem
of rust resistance in this State.

The second series contains the variety Kenya 745 and those genetically similar
to it. Three strains, Kenya 1035, 1049 and 1053, behave in a way similar to 745. Hach
has a major gene for resistance, and crosses between these varieties and Kenya 745
give F, plants which are all resistant. Of the more recently developed varieties, we
have found that Charter and Yalta when crossed with Kenya 745 give F, plants which
are all persistent. It would appear from this that the two varieties were derived
from Kenya 745 as a common ancestor. Kenya 744, which gives segregation when
crossed with members of the above groups, is so far placed alone in a third class.

- It is of some interest to find that when Gabo and Kendee are crossed, no segregation
for rust reaction occurs. Crosses with Gabo and either Kenya 743 or 744 both give in F.
approximately one plant in 16 that is susceptible. As a result of observations on many
thousands of plants of resistant selections of- (Bobin x Gaza)F, x Bobin crossed with
Dundee x Kenya 745, it appears that the single major gene for resistance inherited from
Gaza (T. durum) is allelic to the one present in Kenya 745.

So far it has not been possible to put the other rust resistant varieties Hofed,
Fedweb, Warigo and Celebration into their respective groups, but at least it can be said
that Hofed x Eureka crosses show segregations in F, and later generations. This is
confirmed by the United States workers (Ausemus, 1944), as RL 1373 x Hope showed
segregation in F,. Ausemus also reports segregation in crosses between RL 1373 and
Thatcher. This probably means that the Australian variety Celebration would also give
this segregation for susceptibility in crosses with RL 1373, Kenya 743 or Eureka.

BY I. A. WATSON AND W. L. WATERHOUSE. 129

A considerable amount of work remains to be done on the available genes for
resistance to stem rust in Australia, particularly since the advent of culture 7316. All
the varieties belonging to the Kenya 745 group are still valuable parents. In addition,
Hofed, Fedweb, Celebration and certain (Timopheevi x Steinwedel) selections continue
to be very resistant in the field. Many of the varieties shown in Tables 5 and 6 will,
no doubt, be of some importance, too, but as yet insufficient data are available dealing
with their behaviour to rust in the field.

If we consider the problem of breeding for rust resistance here in the light of
American experience with race 15B, the number of varieties available as potential
parents is greatly reduced. This biotype of race 15 is differentiated by its ability to
attack Rival, but many other varieties are also susceptible to it. According to Ausemus
(1944), Hope, Thatcher and (7. Timopheevi x Steinwedel) S990 have been susceptible
to this biotype both in the seedling and mature plant stage under controlled experi-
mental conditions. He reports further that two Kenya spring wheats, Kenya 58 and
117A, are highly resistant to it, and Red Egyptian is moderately resistant. Da Silva
(1947), working at St. Paul, Minn., found Kenya 58 to range from susceptible to
moderately susceptible. Sydney University tests on seedlings of K58, 117A and Red
Egyptian with 7316 indicate that of the three, only 117A is resistant. For this reason
it appears to be a variety of considerable potential value as a source of resistance.
Nothing is known yet of the mode of inheritance of its resistance to 7316, but it is a
variety quite distinct from 743, 744 and 745, and as well gives a vigorous F, in crosses
with Khapli Emmer. Apparently its gene is not allelic with that of Kenya 744 or 745,
as in crosses with these varieties, susceptible plants have been obtained in the F,
generation.

The incidence of race 15B in the United States has necessitated an intensive search
for varietal resistance. In their present breeding programmes, the aim is to retain
resistances to stem rust now available, and wherever possible incorporate resistance to
new races. By this means it is hoped to solve new rust problems soon after they arise.
This line of approach is being followed to some extent here in that we have added to
Hureka the rust resistance of Gabo by back-crossing. Our present policy, however, is to
take certain of the very desirable commercial varieties and derive strains from them
which are at least morphologically indistinguishable but which have derived their rust
resistance from a different source. Hureka and Gabo may be used to illustrate this.

Eureka is a very desirable variety agronomically, and, as mentioned previously, its
popularity increased rapidly following its release in 1938. It now lacks resistance to
rust. Crosses such as the following have been made or will be made in the near future:
Eureka x Gabo, Eureka x Hofed, Hureka x 117A, Eureka x (T. Timopheevi x Steinwedel).
The resistant progeny from each of the crosses will be backcrossed to Eureka so that
most of its genotype will be recovered. In the end a number of different strains closely
resembling Hureka will be derived from these backcrossings. They should be prac-
tically indistinguishable except that their resistance in each case is due to a different
source. One strain, say the (Gabo x Eureka)‘ would, for the time being, be released to
growers since we have that in a fairly advanced stage. The others would be retained
jn the varietal collection and tested immediately a change in the rust flora occurs. If
1 new race of rust should arise, it is most unlikely that all derived strains will be
susceptible. One or other of those resistant both to the old and the new rust would
then be immediately increased for distribution. No extensive yield tests should be
necessary since the yielding ability will approximate that of Eureka.

Much the same policy is being followed with Gabo. This variety is also very
satisfactory commercially, and in the past few seasons has given some phenomenal
yields in the north-west of N.S.W. It is also highly resistant to stem rust. As shown
above, Gabo has the gene for resistance allelic to the one in Kendee, and this in turn is
allelic to that present in Charter and Yalta. It appears likely that should a new
strain of rust arise capable of attacking Charter, then Kendee, Gabo and Yalta will
also be susceptible. Since Gabo has proved such a high yielding variety, it is
considered that other strains, very similar to it but with a range of genic material

130 AUSTRALIAN RUST STUDIES. VII,

for rust resistance, should be available. Should a new rust arise against which the
Gabo gene is not effective, then one or other of the other genes that have been added
to the derived strains would contribute the necessary resistance.

In obtaining the new strains, where possible the technique for combining resist-
ances suggested by Macindoe (1948) will be used. If this is not possible—and it may
not be with varieties such as 117A—it is proposed to derive a susceptible strain of
Gabo and add the various resistances to it by crosses with the appropriate varieties
mentioned above. As with the Eureka strains, those derived from Gabo would be
retained until it was necessary to release them.

The essential difference between this approach and that suggested by Ausemus
is that the end products will possess in the main only one or two major factors for
resistance. It is probable that with the fewer genes they will individually not be
resistant to such a wide variety of rust types, but collectively their resistance should
be rather comprehensive. It is not possible to breed specifically against new races
until they arise, but it is expected that the procedure outlined above will give material
showing some resistance to new strains. It is fortunate that at the present time the
resistant material available to the Australian wheat breeder still offers a fairly wide
choice, and this will probably be the case until something like race 15B arrives. Until
then some progress could be made by having resistant hybrid lines tested in the
United States for their reaction to the more virulent races there. Preparations must
be made, however, for the possible occurrence of races here more pathogenic than any
yet recorded elsewhere.

SUMMARY.

During the period 1941-42 a new culture of stem rust arose in New South Wales
capable of attacking the variety Hureka. At about this time a change also occurred in
the race 34 which was so widespread throughout Australia. There are now present in
most parts of the country mainly two races of stem rust; one cannot attack Hureka and
is listed as race 126; the other is very similar to it, except that Eureka is susceptible.
This is regarded as a biotype of race 126 and listed as race 126B. Since 1942 the old
rust which for many years was regarded as race 34 has not been collected and it is
considered that race 126B represents a mutation from race 34. In the absence of race
126B, Eureka is still a highly resistant variety. Several varieties are very resistant to
races 126 and 126B. Most of the Kenya varieties are susceptible, but one, namely 117A,
is of special interest in that it is resistant to race 15B in the United States as well as to
races 126 and 126B here.

e ACKNOWLEDGEMENTS.

Our thanks are due to our colleague, Dr. E. P. Baker, who from time to time helped
in this investigation and made several field collections of rust. We acknowledge also
financial help given from the Commonwealth Science Research Grant.

References.

AUSEMUS, E. R., 1944.—Breeding for Stem Rust Resistance in Spring Wheat. Rep. Mill.
Baking Mtg. 7th Hard Spring Wheat Conf., Minneapolis, Minn., pp. 27. Mimeographed.

Da Sinva, A. R., 1947.—Estudos Preliminares Para a Producao de Variedades de Trigo
Resistentes as Ferrugens no Brasil. Bol. Serv. Nacion. Pesq. Agron., No. 1, pp. 53.

JOHNSON, T., and NEWTON, MARGARET, 1941.—The Effect of High Temperature on the Stem
Rust Resistance of Wheat Varieties. Canad. J. Res., 19, 438-445.
JOHNSTON, C. O., HUMPHREY, H. B., CALDWELL, R. M., and Compton, L. R. F., 1942.—Third
Revision of the International Register of Physiologic Races of Leaf Rust of Wheat.
LOEGERING, W. Q., 1944.—Stem and Leaf Rust in Relation to Wheat Breeding. Three Green-
house Inoculations with Stem Rust. Rep. Mill. Baking Mtg. 7th Hard Spring Wheat Conf.,
Minneapolis, Minn., pp. 27. Mimeographed.

MacInbDogE, S. L., 1931.—Stem Rust of Wheat. Observations at Glen Innes during 1930-31
Season. Agric. Gaz. N.S.W., 42, 475-484.

————,, 1987.—The New Era in Breeding Wheats Resistant to Stem Rust. Jour. Aust. Inst.
Aigric: (S¢:; 3, 20-01%

, 1945.—New Stem Rust Resistant Wheats to Replace Eureka. Agric. Gaz. N.S.W..
56, 530-531.

BY I. A. WATSON AND W. L. WATERHOUSE. 131

Macinpo#, 8. L., 1948.—The Nature and Inheritance of Resistance to Stem Rust of Wheat,
Puccinia graminis tritici, possessed by Several Parents. N.S.W. Dept. Agric. Science Bull.
No. 69. ;

NEWTON, MARGARET, JOHNSON, T., and PETURSON, B., 1940.—Seedling Reactions of Wheat
Varieties to Stem Rust and Leaf Rust and of Oat Varieties to Stem Rust and Crown Rust.
Canad. J. Res., 18, 489-506.

, 1946.—Physiologic Races of Puccinia graminis tritici in Canada, 1919 to 1944. Canad.
J. Res., 24, 26-38.

Reppick, D., and Miuus, W. D., 1938.—Building up Virulence in Phytophthora infestans. Amer.
Potato Jour., 15, 29-34.

RoDENHISER, H. A., and HurRD-KARRER, ANNIE M., 1947.—Evidence of Fusion Bodies from
Urediospore Germ Tubes of Cereal Rusts on Nutrient Solution Agar. Phytopath., 37
744-756.

STAKMAN, EH. C., Levine, M. N., and LOEGERING, W. Q., 1944.—Identification of Physiologic
Races of Puccinia graminis tritici. Paper 2148, Scientific Journal Series, Minn. Agr. Eap.
Station.

THOMAS, I., 1948.—Wheat Variety Trials. J. Agric. West. Aust., 25, 84-94.

WATERHOUSE, W. L., 1929.—Australian Rust Studies I. Proc. LINN. Soc. N.S.W., liv, 615-680.

, 1934.—Australian Rust Studies IV. Natural Infection of Barberries by Black Stem
Rust in Australia. Proc. LINN. Soc. N.S.W., lix, 16-18.

, 1938.—Some Aspects of Problems in Breeding for Rust Resistance in Cereals. J. Roy.
Soc. N.S.W., 72, 7-54.

, and Warson, I. A., 1941.—Australian Rust Studies VI. Comparative Studies of
Biotypes of Race 34 of Puccinia graminis tritict. Proc. LINN. Soc. N.S.W., lxvi, 269-275.

Watson, I. A., 1942.—The Development of Physiologic Races of Puccinia graminis tritici Singly
and in Association with Others. Proc. Linn. Soc. N.S.W., Ixvii, 294-312.

, 1943.—Inheritance Studies with Kenya Varieties of Triticum vulgare Vill. Proc.
LINN. Soc. N.S.W., Ixvili, 72-90.

, and WATERHOUSE, W. L., 1945.—A Third Factor for Resistance to Puccinia graminis
tritici. Nature, 155, 205.

EXPLANATION OF PLATES.

Puate III.

A. Typical straw of eight different varieties of wheat showing the effects of stem rust at
Gunnedah, 1942. 1. Ford. 2. Eureka. 3. Kenya 743. 4. Kenya 744. 5. Kenya 745. 6. Kendee.
7. Fedweb. 8. Gabo.

B. Typical grain showing the effects of stem rust at Gunnedah, 1942. The same number
of grains was photographed in each case. 1. Ford. 2. Eureka. 3. Fedweb. 4. Gabo.

PLATE IV.

A. Typical straw showing the effects of stem rust on three varieties at Badgery’s Creek,
1942. 1. Federation. 2. Hureka. 38. Kenya 745. Eureka, which was formerly resistant, is
now as susceptible as Federation, whilst Kenya 745 maintains its resistance.

B. Above are shown normal healthy heads to the right in each pair, and distorted heads
caused by Sclerospora macrospora to the left in each pair of 1. Kendee, and 2. Gabo. Below
are shown stems of these varieties showing how attack by S. macrospora now renders them
susceptible to stem rust. Grown at Tichborne, 1942.

C. A comparison between leaves of Eureka inoculated with the new race styled r. 126B(1.)
and similar leaves inoculated with U.S.A. r. 34(2.). In each pair of leaves the left shows
the upper and the right the lower surface. (Nat. size x 2.)

132

NOTES ON MICROSPORE-TYPES IN TASMANIAN PERMIAN COALS.
By J. A. and Roma DULHUNTY.
(One Text-figure. )

[Read 25th May, 1949]

INTRODUCTION.

A preliminary survey of microsporetypes in Tasmanian Permian coals was under-
taken with the object of recording the principal types present, obtaining data regarding
assemblages in different occurrences, comparing general assemblages in Tasmanian and
New South Wales Permian coal measures, and accumulating information which may
eventually be of use in palaeobotany and its application to stratigraphy.

Coal samples used in the investigation (Table I) were supplied by the Tasmanian
Department of Mines. It was not possible to obtain samples from all occurrences of
Permian coal in Tasmania, but the -material was sufficient to provide data for
preliminary purposes.

TABLE 1.
Permian Coal Samples Examined.
|

Syd. Uni.
Coal Serial Locality. Coalfield or District.
Number.

CS 157 Illamatha Mine. Mersey Coalfield.

CS 158 Aberdeen Mine. Mersey Coalfield.

CS 159 Tarleton Mine. Mersey Coalfield.

CS 190 Mole Creek. Mersey Coalfield.

CS 191 Mount Pelion. Western District.

CS 192 Cradoe. South-eastern District.

CS 213 Preolena. Western District.

CS 311 Latrobe. Northern District.

CS 324 Wynyard. North-western District.

Spore concentrates were prepared from the coal samples and mounted for
microscopical examination by a method already described (Dulhunty, 1945). The
Preolena material was cannel coal and no microspores were found in it. The Latrobe
sample was a type of oil shale, known as Tasmanite, consisting almost entirely of
macrospores. The concentrate prepared from this material contained an abundance of
well-preserved macrospores, but only a limited number of microspores could be found.
These were imperfectly preserved and insufficient in number to provide an assemblage
for comparison with other occurrences, but the types recognized are recorded in Table 4.
All the other coal samples contained large numbers of well-preserved microspores
suitable for the study of types and assemblages.

A thorough search was made in each concentrate for types known to occur in
New South Wales Permian coals; new types were studied; and spore counts were made
to determine the relative abundance of all types present. Each spore type was then
referred to one of five general orders of abundance depending on the percentage which
it represented of the total number of spores counted in the concentrate. The five
orders of abundance and the percentages to which they correspond are set out in

BY J. A. AND ROMA DULHUNTY. 133

Table 2. This procedure was adopted as it was considered that the limited number
of samples available for examination did not justify the consideration of actual
percentages in discussing the significance of results.

TABLE 2.
General Orders of Abundance.

Abundance General Order of Corresponding
Number. Abundance. Percentages.
|
1 Rare. 0:5% and less
2 Infrequent. | 0:°6% to 1:9%
3 Common. 2-:0% to 4:9%
4 Abundant. 5:0% to 7-9%
5 Very abundant. 8:0% and over

The system used for type-numbering of spores in this work was that adopted in
describing New South Wales Permian microspores (Dulhunty, 1945). For details of
the system, and descriptions and illustrations of spore-types, reference should be made
to the foregoing publication. For the convenience of readers, however, an abridged key
to the spore-types is given in Table 3. It is hoped that type-numbering systems may
eventually be abandoned in preference for the use of systematic binomial nomenclature.
However, type numbers were used for the purpose of presenting the present results
as a systematic classification of Australian microspores is not yet available.

TABLE 3.
Abridged Key to Spore-Types.
Examples of type-numbering: P2A, P29B, P34C, P40D.
Letter P preceding type-number indicates Permian type.
Letter A, B, C or D following type-number indicates variations in size or minor details belonging to the
same general type.
Type-numbers (2, 29, 34, 40 in above examples) refer to body-shape, tetrad scar and ornamentation, as

follows :
1. Angular tetrahedral; trilete ; psilate. 18. Ellipsoidal; monolete ; echinate.
2. Sub-angular tetrahedral; trilete ; psilate. | 19. Spheroidal; trilete ; echinate.
3. Ellipsoidal; monolete ; psilate. | 20. Spheroidal; monolete ; echinate.
4. Spheroidal; trilete; psilate. 21. Angular tetrahedral; trilete ; striate.
5. Spheroidal; monolete; psilate. 23. Ellipsoidal; monolete ;_ striate.
6. Angular tetrahedral; trilete; granulate. 26. Angular tetrahedral; trilete ; verrucate.
7. Sub-angular tetrahedral; trilete ; granulate. 28. Ellipsoidal; monolete ; verrucate.
8. Ellipsoidal; monolete; granulate. 29. Spheroidal; trilete ; verrucate.
9. Spheroidal; trilete; granulate. 30. Spheroidal; monolete ; verrucate.
10. Spheroidal; monolete; granulate. 32. Sub-angular tetrahedral; trilete ; monowinged.
13. Ellipsoidal; monolete ; reticulate. 33. Ellipsoidal; monolete ; monowinged.
14. Spheroidal; trilete ; reticulate. 34. Spheroidal; trilete ; monowinged.
15. Spheroidal; monolete ; reticulate. 35. Spheroidal; monolete ; monowinged.
16. Angular tetrahedral; trilete ; echinate. 38. Ellipsoidal; monolete ; biwinged.
17. Sub-angular tetrahedral; trilete ; echinate. 40. Spheroidal ; monolete ; biwinged.

Results of the examination of the microspore concentrates are summarized in
Table 4. In the case of each coal sample types present are shown by relative abundance
numbers and those not found are indicated by dashes. Similar data representing
average results for the Newcastle, Tomago, and Greta measures in New South Wales
are included for comparison. The table shows distribution of microspores with respect
to presence and absence of individual types, as well as variations-in relative abundance
and assemblages, in coals from the different Permian areas in Tasmania. In the case
of the Latrobe material only a small number of different types was found and
recorded owing to the rarity of microspores in the material.

L

134 MICROSPORE-TYPES IN TASMANIAN COALS,

Permian coal-measure stratigraphy in Tasmania was described by Johnston (1888) ;
Loftus Hills, Ried, Nye, and Keid (1922); and later by Voisey (1938). Voisey’s
publication deals with results of previous workers; gives a full list of references; and
reviews correlation of Permian stratigraphy in Tasmania and New South Wales. In
Voisey’s opinion there is difficulty in correlating the scattered occurrences of Permian
strata within Tasmania, and there is considerable uncertainty regarding specific
correlations (made by earlier workers) of various occurrences in Tasmania with the
different stages and measures in New South Wales. He concludes, however, that
Upper Palaeozoic rocks in Tasmania belong to the same system as those in type
sections in the Hunter River district of New South Wales and that the two can be
broadly correlated in a general way. In 1945 Dr. S. W. Carey kindly supplied notes
and correlations of Tasmanian and New South Wales Permian strata.

Based on conclusions and observation of the foregoing workers, tentative
stratigraphical positions of coals used in this investigation are as follows:

Upper Coal Measures: Mount Pelion; Cradoc; Preolena.

Lower Coal Measures: Mersey Coalfield, including Illamatha, Aberdeen, and Tarleton
samples; Mole Creek; Latrobe; Wynyard.
Marine beds occur at the base of the sequence and also between the Lower and
Upper Coal Measures which are generally believed to be equivalent to the Greta and
Upper Coal Measures, respectively, in New South Wales.

MICROSPORES IN TASMANIAN PERMIAN COALS.

The majority of types occurring in both Tasmanian and New South Wales coals
are almost identical and would appear to have come from similar plants. Some exhibit
slight differences which are insufficient to justify the adoption of new type numbers.
These may have been produced by plants representing variations of New South Wales
Permian species, but the spores have been grouped under similar type numbers, as
they possess the same essential features. Examples of such differences in spores of
the same general type are as follows, characteristics of the Tasmanian spores being
quoted. P1B, P3B, and P4B are somewhat smaller. P38A and P38C show less variation
in size. P5A and P5C are more variable in size. P10B seldom exhibits a well-defined
monolete opening. P14A is more variable in size and rarely shows the triangular
opening and short trilete sutures at the centre. P15A is smaller and the reticulum is
less distinct. The echinate spores, P16A to P20A, are smaller and possess shorter
spines. P26A is frequently less angular in outline. P28A is more variable in size but
tends to be larger. P40B exhibits slightly smaller wings. P40D is more variable in
shape and the wings are set farther apart. P41A appears to vary from psilate to
granulate and some examples were observed in which the exine was verrucate and
even slightly echinate.

Several spore-types, present in Tasmanian Permian coals, have not yet been
recognized in New South Wales and were not found in Queensland Permian coals
recently examined by de Jersey (1946). Three of these, illustrated in Text-fig. 1, were
found in sufficient numbers to establish types with definite characteristics. They are
as follows:

Type P5D (No. 1, Text-fig. 1).

Spheroidal; distinctly flattened; circular or slightly oval outline in axial view.
Eixine thick-walled, giving a well-defined marginal rim. A small inner body of variable
size and oval outline is situated within the spore near its centre or slightly to one
side. A longitudinal feature resembling a monolete suture frequently appears on the
inner body. No evidence of trilete sutures has been observed and the spore is
presumed to be monolete. Usually psilate but faint granulation appears in some
examples. Size is variable, diameter varying from one-third to two-thirds of the
over-all diameter.

BY J. A. AND ROMA DULHUNTY. 135

TABLE 4,
Distribution and Abundance of Microspores in Tasmanian Permian Coals.

Tasmanian Coals. N.S.W. Coal Measures.

Types. Tlla- Aber- Tar- Mole Mount Wyn-
matha deen leton Creek Pelion | Cradoc yard | Latrobe} New- | Tomago.| Greta.
CS 157. | CS 158. | CS 159. | CS 190. | CS 191. | CS 192. | CS 324. | CS 311. | castle.

| Oe eo Spe cone er ten ees Teese
emesis abe

eer oneres fo. || testes |) eo eos ean

[coed io ee at

| weste | epee | eee omean | am aes

ac}
or
Q
lH rmoeHPrmroneNnBPENanwoneHyReE wen
| Dee Tomer || = hem wre |

| ro | ow os |

tJ
He
© oO
>
hd
hd
I
|
bei esel Ss | oes | pees e ete as]
lrorvlwerwlrweenwen | wwmanwe |

cae) ine]
iS) 4
‘S) x
> ia

|
| |
| |

| = |

rm | = | ww |
|enomecveellbenoanouoncctcom arco lenomasecom un Omatecom om lmhsuicasteon Kon oMnonorl ono Komodns

[iS Soi |

Sos hea e eae enone pmol] Ssomeeswnw Vom ew Oe ewe mm

le lel el wel ol

|
|
|
wren | |
|
elel lilt]

He
mrewrprrmerweee |
rewrrunmrnuwl wl] |e |
eer! | eee |

Type P13B (No. 2, Text-fig. 1).

Ellipsoidal with oval to sub-oval outline. Monolete; frequently exhibiting well-
defined suture or opening running the full length of the body. Length 40 to 60;
width 30 to 40. Exine reticulate with a widely spaced system of anastomosing ridges

.

136 MICROSPORE-TYPES IN TASMANIAN COALS,

1 to 2 in width and 3 to 5 apart. Ridges appear at the margins of the spore producing
irregularities in its outline.

\

WAY : Si,

°
. \ \ ~
MAL :

Text-figure 1—Permian microspore types present in Tasmanian coals, but not
recognized in New South Wales Permian Measures.

Type P28B (No. 3, Text-fig. 1).

Ellipsoidal with oval to sub-oval outline. Monolete; at times showing a wide
longitudinal opening. Length 55 to 65; width 40 to 50. Exine verrucate with large
closely packed warts, 1 to 2 in width, arranged in a regular system of longitudinal
rows.

Distribution and variation in abundance of the foregoing types are indicated in
Table 4. P5D is widely distributed in Tasmanian coals, its abundance number varying
from 2 to 3. P13B is also widely distributed, varying in abundance from 1 at Mount
Pelion to 3 in the Mersey Coalfield. P28B was found in coal from the Illamatha Mine,
where its abundance was only 1.

In addition to the three types described above, a single example of an unusual
echinate spore (No. 4, Text-fig. 1) was found in Illamatha coal. A new type number
was not given to this spore, as one individual was not considered sufficient to establish
the occurrence of the type. It was spheroidal with approximately circular outline,
and exhibited dehiscence along trilete sutures extending more than half-way round
the body, which was 60 in diameter. The exine was echinate with very small spines,
0-75 to 1:0 in length, somewhat irregularly spaced at 1 to 2 apart. This spore is
distinct from any other spined spores so far observed in Permian coals, and it was
the only spined type found in samples from the Mersey Coalfield.

COMPARISON OF GENERAL ASSEMBLAGES IN TASMANIA AND NEW SOUTH WALES.

Of the 48 spore-types described from New South Wales Permian measures, 43 were
found in Tasmanian coals. The types not recognized were P21A, P29B, P32A, P34A, and
P34C. P21A is the unusual tetrahedral trilete type of special interest owing to the
presence of three dome-shaped protuberances, somewhat resembling pollen-tube pores,
situated at the centres of the three distal interfaces. It is rare in New South Wales
coals, occurring only in the Newcastle Measures with sporadic lateral distribution.
P29B is a large spheroidal verrucate trilete spore. It is widely distributed throughout
all three subdivisions of New South Wales Permian and attains maximum abundance
in the Greta Measures. P32A is a small tetrahedral trilete spore with a flangelike wing

BY J. A. AND ROMA DULHUNTY. Iasi

situated in one place and attached to the distal interfaces which appear as the triangular
outline of the body in proximal view. It is one of the two types (P32A and P15A) found
only in the Greta Measures in New South Wales. Types P34A and P34C are monowinged
spores with spheroidal trilete bodies. P384A occurs only in the Newcastle Measures,
but P34C is found in all New South Wales Permian coal measures.

Some important differences were found in the Tasmanian and New South Wales
microspore assemblages with respect to groups of closely related types. The mono-
winged spores (P33B, P34A, P34B, and P34C) which are relatively common in all
New South Wales coal measures are almost completely absent from Tasmanian coals.
Only three examples (one of P33B and two of P34B) of monowinged spores were found
in all the Tasmanian material examined. These were in coal from Mersey and Wynyard
areas, aS indicated in Table 4. Three individuals are not sufficient to establish the
presence of monowinged spores in the Tasmanian assemblage, as they may have been
transported by wind from the mainland. This apparent absence of monowinged spores
represents one of the principal differences in the two assemblages and it suggests
that certain plants bearing monowinged spores may have been absent in Tasmania,
although they flourished in New South Wales, during Permian time. Monowinged
spores, similar to the New South Wales types, occur in Permian tillite at Bacchus
Marsh in Victoria; in Lower Gondwana shales in India (Virkki, 1939); they have
been observed by the writer in Permian coal from Collie in Western Australia; but
they were not found in Queensland Permian coals examined by de Jersey (1946).

The echinate spores (P16A to P20A), absent from many of the Tasmanian coals,
are less abundant than in the New South Wales Permian, where they occur persistently
in coals from all measures. The biwinged spores, particularly P40A and P40B, are also
far less common in Tasmania than in New South Wales. Psilate tetrahedral trilete
spores (PIA, PIB, P2A) are represented in all Tasmanian coals but occur as minor
constituents, whereas in New South Wales they are among the most abundant types.
Ellipsoidal monolete types (P3A and P3C), and the small spheroidal monolete spore P5C,
occur persistently and abundantly in both Tasmanian and New South Wales Permian
measures.

The above results do not appear to bear any special stratigraphical significance.
Of the types not found in Tasmanian coals, some are confined to the Upper Coal
Measures and some to the Greta Measures in New South Wales. It would seem,
however, that the results are significant in relation to the distribution of Permian
plants in Australia. This subject, which is beyond the scope of the present work,
requires further knowledge about relations between Permian spores and plants, but
the general absence in Tasmania of certain spores relatively abundant in New South
Wales, and vice versa, suggests that important results regarding distribution of Permian
floras may be obtained from the study of microspores when the plants to which they
belong are known.

VARIATIONS IN TASMANIAN SPORE ASSEMBLAGES.

Spore assemblages vary considerably in coals from different Permian localities
in Tasmania, as illustrated in Table 4. The spined spores (P16A to P20A) appear to
be completely absent from Illamatha, Aberdeen, and Tarleton coals in the Mersey
Coalfield and also the Wynyard sample, although they are present in the other coals,
including Mole Creek. Types P40A and P40B were found only in Mount Pelion and
Cradoc materials, being absent from all the Mersey coals and Wynyard sample. The
only examples of spheroidal monowinged spores, whether transported or indigenous,
were found in Illamatha and Wynyard samples, and these were also the only two coals
in which P38B and P4A were found. Mole Creek and Mount Pelion were the only
coals in which P38A was present and from which P5A was absent.

These results indicate a general similarity in spore assemblages in coals from
Iliamatha, Aberdeen, Tarleton, and Wynyard, supporting the conclusions of previous
workers that the Mersey and Wynyard coal measures are stratigraphically equivalent.
Assemblages in. Mount Pelion and Cradoe coals are similar, but differ somewhat from

138 MICROSPORE—-TYPES IN TASMANIAN COALS,

those of the Mersey and Wynyard materials. This also supports existing views that
Mount Pelion and Cradoec measures are equivalent but occur on a different horizon to
the Mersey and Wynyard coals. The Mole Creek sample, however, contains an
assemblage similar to that of Mount Pelion and Cradoc coal, and distinctly different
from assemblages in samples from the Mersey field. This suggests that Mole Creek
coal may be stratigraphically equivalent to that at Mount Pelion and Cradoc rather
than seams mined at Illamatha, Aberdeen, and Tarleton collieries, as previously
suggested.

The foregoing results suggest that it may be possible to use the absence of spined
spores in correlating Tasmanian Permian areas equivalent to the Mersey and Wynyard
coals, which are regarded as lower Permian in age.

Insufficient spore-types were recognized in the Latrobe material to indicate an
assemblage, and the types identified bear no special stratigraphical significance, as
they were found in practically all the Tasmanian coals examined.

APPLICATION OF RESULTS TO CORRELATION WITH NEW SOUTH WALES PERMIAN.

Attempts to apply preliminary results, recorded in this paper, to correlation of
various Tasmanian occurrences with subdivisions of New South Wales Permian are
not promising. The two assemblage groups, into which Tasmanian coals fall, cannot
be matched with any assemblages in New South Wales coal measures. Certain spore-
types appear to have restricted stratigraphical ranges in New South Wales (Dulhunty,
1946). Some of these occur in Tasmanian coals, but results based on their presence,
or absence, are conflicting. P15A (confined to New South Wales Greta coal) is present
in two of the Mersey coals and absent from Mole Creek, Mount Pelion, and Cradoc
samples. Also, P40A and P40B (confined to Upper Coal Measures in New South Wales)
were found only in Mount Pelion and Cradoc coals. While this suggests that Mersey
coal is of Greta age and that Mole Creek, Mount Pelion, and Cradoc occurrences are
of Upper Coal Measure age, it is contradicted by the fact that P8B and P13A (confined
to Upper Coal Measures in New South Wales) occur in the Mersey and Wynyard coals
but not in the Mount Pelion and Cradoc samples. Furthermore, P9A (absent from
New South Wales Greta) was found in two of the Mersey coals and the Cradoc sample.

It follows that no conclusions can be reached at this stage by studying the
presence and absence of types which appear to have restricted ranges. This is not
surprising in view of the distance between Tasmania and New South Wales and the
palaeogeographical variations in environment which probably existed between the two
areas during Permian time. Conditions favouring certain plants in New South Wales
during Greta or Upper Coal Measure deposition may not have existed during deposition
of equivalent strata in Tasmania, some 700 miles away.

SUMMARY.

Permian coal samples were examined from nine localities in Tasmania and microspores
were found in all except one from Preolena. Of the forty-eight spore-types described
from New South Wales Permian coals, forty-three were found in Tasmania. Four
new types, not recognized in New South Wales, were observed in Tasmanian coals.
Monowinged spores are almost completely absent in Tasmanian coals, although they
are present in the Permian of New South Wales, Victoria, Western Australia and
India. Biwinged spores are far less common in Tasmania than in New South Wales.
Coals from Wynyard and the Mersey field (Illamatha, Aberdeen, and Tarleton) are
characterized by the absence of echinate spores which are relatively abundant in Cradoc,
Mount Pelion and Mole Creek coals. Evidence based on assemblages supports existing
views that Mersey and Wynyard coals are equivalent and occur on a different horizon
from Mount Pelion and Cradoc coals. The assemblage in Mole Creek coal resembles
that of Mount Pelion and.Cradoc materials rather than the Mersey coals, with which
it was previously believed to be equivalent. The study of presence and absence of
types with limited ranges in New South Wales produced conflicting results when

BY J. A. AND ROMA DULHUNTY. 139

applied to correlation of Tasmanian occurrences with subdivision of New South Wales
Permian.

ACKNOWLEDGEMENTS.

This investigation was part of a programme of coal research supported by funds from
the Commonwealth Research Grant to the University of Sydney. The authors wish to
acknowledge the generous co-operation of the Tasmanian Department of Mines in providing
samples of coal and stratigraphical information, and the assistance of Miss N. Hinder in the
preparation of microspore concentrates.

References.

DE JERSEY, N. J., 1946.—Microspore Types in Some Queensland Permian Coals. Univ. Q’land
Papers Dept. Geol. I11(5), 12 pp.

DuLHUNTY, J. A., 1945.—Principal Microspore-Types in the Permian Coals of New South
Wales. Proc. LINN. Soc. N.S.W., Ilxx: 147.

JOHNSTON, R. M., 1888.—A Systematic Account of the Geology of Tasmania.

Lortrus HILus, RIED, A. M., Ny#, P. B., Kemp, H. G. W., 1922.—The Coal Resources of Tasmania.
Tas. Dept. Mines., Geol. Sur., Min. Res. No. 7.

VirKKI, D., 1939.—On the Occurrence of Similar Spores in a Lower Gondwana Glacial Tillite
from Australia and in Lower Gondwana Shales in India. Proc. Indian Acad. Sci., Vol. ix,
No. 1, Sect. B.

VoisEyY, A. H., 1938.—The Upper Palaeozoic Rocks of Tasmania. Proc. LINN. Soc. N.S.W.,
G8 8 309.

140

A NEW SUBSPECIES OF AEDES (STEGOMYIA) SCUTELLARIS WALKER
(DIPTERA, CULICIDAE) FROM NORTHERN AUSTRALIA.

By A. R. WooDHILt,
Department of Zoology, University of Sydney.

(With Four Text-figures. )

[Read 31st August, 1949.]

INTRODUCTION.

A considerable amount of work has been carried out in recent years on the
scutellaris group by Farner and Bohart (1945), Stone and Farner (1945), and by
Stone (1947). The last-named author has shown that Aédes variegatus var. hebrideus
Edwards is synonymous with Aédes scutellaris Walker. Prior to 1944 no members of
the scutellaris group had been recorded from Australia, but in December, 1944, and
January, 1945, R. H. Wharton collected three females and one male at Batchelor in
the Northern Territory of Australia. These were forwarded to Stone, who pointed out
that they differed in the femoral markings from A. scutellaris, but the material was
considered inadequate for description. In January, 1948, Mr. A. EH. Wynn forwarded to
the author a batch of eggs from Katherine, Northern Territory of Australia, and these,
when developed, proved to be similar to the specimens collected by Wharton. They
have been retained as a continuous laboratory culture for a year and several hundred
specimens have been examined. It has been found that they show a constant variation
in the marking of the mid femora from A. scutellaris Walker, but are identical as
regards the male genitalia and in all other respects, including the eggs, larvae and
pupae. In addition, males of this North Australian form, when crossed with females
of A. scutellaris from New Guinea, gave fertile progeny to the F, generation, although
the reciprocal cross was sterile. In view of these facts it was considered advisable to
describe the new form as a subspecies of Aédes scutellaris Walker.

AKDES SCUTELLARIS SCUTELLARIS Walker.*
Synonymy.

Culex scutellaris Walker, 1859. Proc. Linn. Soe. Lond., Zoology, 3, pp. 77-131.

Culex zonatipes Walker, 1861. Proc. Linn. Soc. Lond., Zool. 5, p. 229.

Aédes variegatus var. hebridews Edwards, 1926. Bull. Ent. Res., 17, p. 102.

Aédes scutellaris var. hebrideus Edwards, 1932. Gen. Insect., fase. 194, p. 163.

Aédes scutellaris hebrideus. Edwards. Knight, Bohart and Bohart, 1944, Keys to the
Mosquitoes of the Australasian Region, Nat. Res. Council, Washington, D.C., p. 55.

Aédes hebrideus Edwards. Farner and Bohart, 1945, U.S. Naval Medical Bulletin, 44, p. 30.

Type Locality: Aroe Islands.

Distribution: Farner and Bohart (1945) and Stone and Farner (1945) give the
distribution as eastern New Guinea (including the D’Entrecasteaux and Trobriand
Islands), New Hebrides, Palau Islands, Ceram and the Philippine Islands. Additional
records supplied to the author by D. J. Lee are as follows:

New Britain: Rabaul (Taylor), 1933.

New Guinea: Gona (O’Connor), 1948; Lae (Clinton), 1943; Lalapipi (Atherton),
1943; Finschaffen (Berril), 1944; Dobadura (Ratcliffe), 1943; Salamaua
(Woodhill), 1944; Merauke (Roberts), 1943; Port Moresby (Wharton), 1947;
Kanusia (Lee), 1947.

Morota (Clarke), 1945.

* This combination has previously been used by Knight, Bohart and Bohart (1944), and
Farner and Bohart (1945) give it as a synonym of Aédes quasiscutellaris F. and B.

BY A. R. WOODHILL. 141

AEDES SCUTELLARIS KATHERINENSIS, nN. subsp.

Type Locality: Katherine, Northern Territory of Australia.

Types: Holotype female, allotype male, ten female and ten male paratypes and a
series of mounted male genitalia deposited in Macleay Museum, University of Sydney.

Distinctive Characters: This subspecies can be distinguished by the presence of a
broad distinct line of white scales on the anterior surface of the mid femur; this line
is entirely lacking in A. scutellaris scutellaris (Text-figs. 1 and 2).

iD

Text-figures 1 and 2.
1. Anterior surface of mid femur of Aédes scutellaris katherinensis, n. subsp. x 50.
2. Anterior surface of mid femur of Aédes scutellaris scutellaris Walker. x 50.

DESCRIPTION OF FEMALE.

Head.

The head is clothed with broad flat black scales and with flat white scales forming
a median longitudinal band and a lateral and ventro-lateral longitudinal white band
on each side. The median white band includes the inter-ocular vertex and extends as a
few white scales along the dorsal margins of the eyes; the ventro-lateral white bands
also extend along the ventral margins of the eyes. The occipital region carries a
small patch of black upright forked scales and a row of long black post-ocular bristles
is also present on each side. The clypeus and antennae are black with a band of flat
white scales round the anterior margins only of the pedicels. The palpi are clothed
with black scales except for a conspicuous patch of white scales covering the dorsal
aspect of the third segment. The proboscis is entirely black scaled. ;

Thorax.

The thorax is dark brown to black in colour, conspicuously marked with bands and
patches of white scales. The scutum is covered with narrow dark brown scales, except
for a small median bare area at the posterior margin and two small bare areas on the
postero-lateral angles, and carries scattered dark bristles which are particularly abundant
above the wing bases. A conspicuous white longitudinal stripe which narrows
posteriorly extends from the anterior margin of the scutum to the bare patch on the
posterior margin, and a band of broad flat white scales occurs on the postero-lateral
margin above the wing base and extends to the bare area on each side, i.e., almost to
the scutellum. In a few specimens out of some hundreds examined there is also an
indistinct line consisting of a few yellowish scales on each side of the posterior bare
area. The scutellum is dark with a broad band of flat white scales extending across
its whole width, and each lobe carries four to six long dark bristles. The metanotum
is bare and dark brown in colour. The anterior and posterior pronotum carry a band
of broad white scales continuous with the white scales on the lateral region of the
head and with the lateral white band on the scutum, so that a continuous white bang

M

142 A NEW SUBSPECIES OF AEDES (STEGOMYIA) SCUTELLARIS WALKER,

runs from the head over the wing bases to the scutellum. Below this and parallel to
it is another continuous band of white scales extending from the propleuron to the
posterior margin of the mesepimeron. In addition, there are two separate patches of
white scales near the lower margins of the sternopleuron and mesepimeron. The
ornamentation of the dorsal and lateral aspects of the thorax is identical with most
other species of the scutellaris group and has been admirably illustrated by Farner
and Bohart (1945).

4

Text-figures 3 and 4.
3. Basal lobe of coxite of Aédes scutellaris katherinensis, n. subsp., dorsal aspect. x 650.
4. Basal lobe of same from the lateral aspect. x 650.

3)

Pleural Chaetotazxy.

The anterior pronotum carries a tuft of ten to twelve strong dark bristles and
the posterior pronotum has a single strong dark bristle near its posterior margin. The
propleuron has a group of three to five bristles, there are no spiraculars, and the post-
spiracular bristles are usually two in number, but may vary from one to three, including
a small weak bristle. The pre-alar bristles vary from five to nine, the upper sterno-
pleurals from one to three, and there is a row of lower sternopleurals varying from
three to five. The upper mesepimerals occur as a tuft of weak pale bristles varying
in number from two to eight, and lower mesepimerals are absent.

Halteres.
These are pale basally, with the apical knob covered with flat black scales.

Wings.

The veins are clothed with both broad and elongated dark scales, with particularly
dense scaling on the costa, subcosta and R,, these scales being mainly broad. At the
extreme base of the costa is a small patch of white scales. The upper fork cell is
slightly longer than the lower fork cell. The wing fringe is of the usual type and
the alula bears a row of small flattened scales on its margin.

Legs.

The coxa of each leg bears a conspicuous patch of white scales anteriorly and
also a row of strong bristles. The fore femur is black with an anterior apical patch of

BY A. R. WOODHILL. 143

white scales and a narrow line of somewhat scattered white scales on the anterior
surface extending from the base to one-half to one-quarter the length of the femur.
This line is very variable and may be absent or represented only by a few scattered
scales. The posterior surface of the fore femur is pale scaled on the basal half. The
mid femur is black with a white anterior apical patch and a broad distinct line of
white scales on the anterior surface extending from the base almost to the apical white
patch; this is a constant character in all specimens examined. The posterior surface
of the mid femur is black with a few scattered pale scales on the basal half. The
hind femur is black with an apical anterior white spot and practically the whole of
the anterior surface is covered by a broad longitudinal band of white scales which
tapers apically; the posterior surface also has a broad white longitudinal band which
tapers apically and extends about half the length of the femur.

The fore, mid and hind tibiae are completely black.

The fore and mid tarsi are black with the exception of incomplete narrow basal
rings on tarsi I and II only. Hind tarsi I to IV are black with complete wide white
basal rings, occupying approximately the following proportions of the segments:
tarsus I, one-third; tarsus II, two-fifths; tarsus III, one-half; tarsus IV, three-quarters.
Tarsus V is completely white.

Abdomen.

The abdomen is clothed with flat black scales and has conspicuous transverse bands
and patches of flat white scales. Tergite I is clothed with long fine hairs, sternite I
lacks both hairs and scales, and the remaining tergites and sternites each carry a
row of short fine hairs on their posterior margins. The white markings are arranged
as follows: Tergite I has a lateral white patch on each side and tergites II to VII
have transverse bands which commence at the antero-lateral corners and run backwards
and upwards to cross the tergites transversely slightly closer to the anterior than to
the posterior margins. These bands are complete on tergites IV to VII but may be
interrupted dorsally to a greater or lesser extent on tergites II and III. Sternites II
to VII have similar bands which commence at the antero-lateral corners and run
backwards and downwards to cross the sternites slightly closer to the anterior than
the posterior margins. These are complete on all sternites and may be expanded on
sternites II to V only to form mid-ventral patches which may extend from the anterior
to the posterior margins.

DESCRIPTION OF MALE.

The markings and chaetotaxy are similar to those of the female. The palps are
approximately equal in length to the proboscis and are black scaled with the exception
of a basal white patch dorsally on the second segment, a white ring covering the basal
two-fifths of the third segment, and ventral basal white patches on the fourth and
fifth segments. The genitalia are indistinguishable from those of Aédes scutellaris
scutellaris, the basal lobe of the coxite carrying a series of hairs at the apex with
several longer hairs joined to form a spine. The degree of development of this spine
varies in both sub-species and it is frequently very transparent and difficult to detect.
The shape of the basal lobe also varies in accordance with the exact angle from which
it is viewed and also with the degree of flattening in the individual preparation. Text-
fig. 8 shows the basal lobe in the allotype male from the dorsal aspect and Text-fig. 4
shows the lateral aspect from another specimen.

EGGS, LARVAE AND PUPAE.
These are also indistinguishable from those of Aédes scutellaris scutellaris.

Material Examined.

Several hundred specimens of males, females, eggs, larvae and pupae of laboratory
cultures ex Katherine, three females and one male from Batchelor and two females
from Kummunya Mission.

MM

144 A NEW SUBSPECIES OF AEDES (STEGOMYIA) SCUTELLARIS WALKER.

Distribution.

Batchelor (R. H. Wharton, 27/12/44, 5/1/45, 15/1/45) and Katherine (A. G. Wynn,
17/1/48), Northern Territory of Australia; Kummunya Mission, Port George IV (EH. J.
Davies, 1/2/44), Kimberley Division, Western Australia.

References.

KnicHT, K. L., BoHART, R. M., and BoHarT, G. E., 1944.—Keys to the Mosquitoes of the
Australasian Region. Nat. Res. Council, Washington, D.C., 55.

FARNER, D. S., and BoHArRT, R. M., 1945.—A Preliminary Revision of the Scutellaris Group.
U.S. Naval Medical Bulletin, 44 (1), 21-37.

SToNE, A., and Farner, D. S., 1945.—Further Notes on the Aédes Scutellaris Group. Proce.
Biol. Soc. Washington, 58, 155-162.

SToNE, A., 1947.—A Topotypic Male of Aédes scutellavis Walker. Proc. Ent. Soc. Washington,
49) 13), (815:

145

REVISION OF THE GENUS BRACHYCOME CASS.

PART III. DESCRIPTION OF THREE NEW AUSTRALIAN SPECIES AND SOME NEW
LOCALITY RECORDS.

By Gwenpa L. Davis, B.Sc., Lecturer in Biology, New England
University College, Armidale.

(Seven Text-figures. )

[Read 25th May, 1949.]

INTRODUCTION.

Recent intensive collection of this genus by Victorian botanists has brought to hand
several specimens which are quite distinct from any described species. Similar collecting
in other States will probably increase the number of known species considerably and
at the same time lead to a more accurate knowledge of the distribution of those
already described.

TAXONOMY.
COMPOSITAR, tribe ASTEROIDEA.
Brachycome Cass., Dict. Sci. Nat., xxxvii (1825), 471.
Subgenus EUBRACHYCOME G. L. Davis.
Superspecies TENUISCAPA.

BRACHYCOME STOLONIFERA, Sp. nov. (Text-figures 1-3.)
Holotype: Summits of Kosciusko plateau, N.S.W., ca. 7,000 ft., 3, 1947, A. Costin (MEL).
Paratypes: Eleven, loc. cit. (MEL).

Herba perennis, erecta, glabra, stolonifera, ad 6-7 cm. alta; folia ad 3:2 cm. longa,
1:2 mm. lata, radicalia, lato linearia, integra, obtusa, basibus involutis; omnis herba
1-4 stolonibus quorum folia, brevia sed typica, ultima parti radicata, moriuntur.
Capitula sola, 6-7 mm. transverse lata; pedunculi uno phyllo robusti; involucri phylla
18—24, circiter 4-5 mm. longa, 1:4 mm. lata, lanceolata, acuminata, marginibus fimbriato-
ciliatis; flores radii circiter 38, ligulis 6 mm. longis, 2:1 mm. latis, albis; receptaculum
2 mm. latum, 2 mm. altum, praecipiter conicum, punctum; achaenia 1:8 mm. longa,
0-9 mm. lata, fusca, cuneata, in septato-piloso disco turgida, margine angusto et tenui
et in labro septato-piloso; pappus setis, sericis, albis, conspicuis.

Erect glabrous stoloniferous perennials up to 6-7 em. high. Leaves up to 3:2 cm.
long, 1:2 mm. broad, radical, broad-linear, entire, obtuse with sheathing bases. Hach
plant bears 1-4 stolons on which the small but typical leaves die off when the tip
becomes rooted. Inflorescences solitary, 6-7 mm. diameter. Scapes robust and provided
with a single bract. IJInvolucral bracts 18-24, about 4:5 mm. long, 1-4 mm. broad,
lanceolate, acute, with torn-ciliate margins. Ray florets about 38, rays 6 mm. long,
2:1 mm. broad, white. Receptacle 2 mm. broad, 2 mm. high, steeply conical, pitted.
Fruit 1:8 mm. long, 0-9 mm. broad, dark brown, cuneate, thick in central region, with
a narrow thin margin. Septate hairs are scattered over the central area and along
the edge of each margin. Pappus of silky white conspicuous bristles.

Habitat: Not recorded.
Range: Only known from the type locality.
Specimens examined: Type series only.

This species is vegetatively very similar to certain specimens of B. nivalis var.
alpina (F. Muell. ex. Benth.) G. L. Davis, and B. radicans Steetz, which is possibly the
reason it has been overlooked for so long. The fruit structure is closest to B. Stuartii
Benth., but the margin is thinner and wider and the longitudinal folds are absent.

146 REVISION OF THE GENUS BRACHYCOME CASS. PART III,

Text-figures 1-7.
1-2. B. stolonifera. Holotype and paratype. x i. 3. B. stoloniferas Hruit. << 920%
x

4. B. obovata. Holotype. x 3. 5. B. obovata. Fruit. 20. 6. B. petrophila. Holotype. x #.
7. B. petrophila. Fruit. x 20.
BRACHYCOME OBOVATA, Sp. nov. (Text-figures 4-5.)
Holotype: Echo Flat, Lake Mountain, about 4,700 ft., margin of alpine sphagnum bogs,
25.1.1948, J. H. Willis (MEL).
Paratypes: Two, loc. cit. (MEL).

Herba perennis, glabra, ad 19 cm. alta, radicalibus foliis in basi. Folia ad 10-5 cm.
longa, 1-1-3 mm. lata, linearia—lineari-obovata, integra, quorum latiora in basi petiolum
habent. Capitula 1-2; 8-9 mm. transverse lata; pedunculi 4—6, foliosis phyllis; involucri
phylla 22-26, 4:3-4-5 mm. longa, 1:1-1:-5 mm. lata, subacuminata-acuminata, serrata,
glabra aut in parti exteriora mic. glandulosa; flores radii circa 24, 5 mm. longi, 1:4 mm.
lati, simulate albi-caeruiei; achaenia 2:3-2-5 mm. longa, 1—1:3 mm. lata, fusca, obovata,
levia et planata; pappus setis brevibus fulvis.

Erect glabrous perennials up to 19 cm. high with a basal cluster of radical leaves.
Leaves up to 10-5 em. long, 1:1-3 mm. broad, linear to narrow-obovate, entire, the broader
leaves tapering proximally into a petiole. Inflorescences 8-9 mm. diameter, 1 or 2 present
on each plant. Scapes provided with 4-6 leaf-like bracts. Involucral bracts 22-26,
4:3-4-5 mm. long, 1:1-1:5 mm. broad, subacute to acute with serrulate margins and
either glabrous or microscopically glandular on the outer surfaces. Ray florets about
24, rays 5 mm. long, 1-4 mm. broad, apparently white to bluish. Fruit 2:3-2:5 mm. long,
1-1:3 mm. broad, light brown, obovate, smooth and flattened. Pappus of short straw-
coloured bristles.

Habitat: Apparently swampy conditions at high elevations.

Range: Eastern Victoria.
Specimens examined:

Victoria: Echo Flat, Lake Mountain, about 4,700 ft., margin of alpine sphagnum bogs,

25.1.1948, J. H. Willis (MEL. holotype and paratypes) ; Snowy Mountains, 6-7,000 ft.,
2.1890, W. Bauerlen (MEL); Genoa district, 3.1885, W. Bauerlen (MEL).

BY GWENDA L. DAVIS. 147

The above series, although small, shows a certain amount of variation in the shape
of the leaves. Those of specimens from Lake Mountain and Genoa agree in that they
are linear and grasslike and the plants resemble certain specimens of B. radicans Steetz
ex Lehmann. On the other hand, the specimens from Snowy Mts. bear narrow-obovate
leaves and are very similar to B. nivalis var. alpina (F. Muell. ex Benth.) G. L. Davis.
It is anticipated that more critical collecting in the highlands of Victoria will produce
specimens intermediate between these two forms of B. obovata.

The fruits are very distinctive but are superficially similar to those of B. graminea
(Labill.) F. Muell. They can, however, be readily distinguished by the small but
distinct pappus. Dissection of the fruit shows that the whole of the interior is
occupied by the embryo and cotyledons, whereas in B. graminea these are completely
enclosed by the spongy margins. For this reason the affinities of B. obovata are to be
found within the superspecies tenuiscapa.

BRACHYCOME PETROPHILA, Sp. nov. (Text-figures 6-7.)

Holotype: Little River Falls, about five miles N.E. of Wulgulmerang, E. Vic., “on dripping
cliff faces’, 16.1.1948, J. H. Willis (MEL).
Paratypes: Four, loe. cit (MEL).

Herba perennis, inferme erecta aut adscendens, ramosa, glanduloso-pubescens, ad
33°5 em. alta; folia caulina, ad 4:5 cm. longa, 1:7 cm. tata, grosse orbicularia, superiora
plerumque sessilia, dentata, 5 utrimque dentibus, inferiora petiolo brevi et interdum
fere crenata; capitula 1-3, 6-7 mm. transverse lata; pedunculi graciles, axillares,
6-5-10:-7 cm. longi, uno folioso phyllo; involucri phylla circiter 24, ad 4 mm. longa,
1 mm. lata, angusto-lanceolata, acuminata, in exteriora parti glandulosa, marginibus
fimbriato-ciliatis; flores radii circiter 28, ligulis 6 mm. longis, 1:3 mm. latis, albis;
receptaculum 3 mm. latum, 2:1 mm. altum, hemisphaericum, punctum; achaenia 1-9 mm.
longa, 0:9 mm. lata, fusca, cuneata, compressa, margine angusto glabro 2 longis sinibus
utrimque separato; discum tubercula pauca, parva, sparsa, in ultima parti mic.
glanduloso pilo, habet; pappus clarus, fulvus, setis in fasciculis inaequibus dispositis.

An apparently weakly erect or ascending branching ? perennial, shortly glandular
pubescent all over and up to 33:5 cm. high. Leaves cauline, up to 4-5 cm. long, 1-7 cm.
broad, orbicular in gross outline, mainly sessile but the lower ones tapering proximally
into a short petiole; upper leaves dentate with usually five teeth on either side,
lower leaves sometimes almost crenate. Inflorescences 1-3 on each plant, 6-7 mm.
diameter, and borne on slender axillary peduncles 6:5-10:7 cm. long, each of which
is provided with a small leaf-like bract. IJnvolucral bracts about 24, up to 4 mm. long,
1 mm. broad, narrow-lanceolate, acute, glandular on the outer surface and with torn
ciliate margins. Ray florets about 28, rays 6 mm. long, 1:3 mm. broad, white.
Receptacle 3 mm. broad, 2:1 mm. high, hemispherical, pitted. Frwit 1:9 mm. long,
0-9 mm. broad, dark brown, cuneate, flat, with two longitudinal folds down each
side, cutting off a narrow smooth margin. Central region of fruit bears a few small
scattered tubercles, each of which is tipped with a microscopic glandular hair. Pappus
conspicuous, straw-coloured, the bristles grouped in bundles of irregular length.

Habitat: “In moist shaded places in soil amongst the rocks, on ledges or in crevices” (N. A.

Wakefield).

Range: Eastern Victoria.
Specimens examined:

Victoria: Little River’ Falls, about five miles N.E. of Wulgulmerang, HE. Vic., “on dripping
cliff faces’, 16.1.1948, J. H. Willis (MEL, holotype and paratypes); Little River Falls,
Wulgulmerang, ‘on ledges and in crevices amongst moist porphyry rocks, 2,500 ft.,
17.1.1948, N. A. Wakefield (NAW); Murrundal River, “in shaded crevice of porphyry
rock, about 500 ft.’, 19.10.1947, N. A. Wakefield (NAW).

Vegetatively this species is very similar to B. Nova-Anglica G. L. Davis, and the
leaves, though larger, are of the same type. Both species appear to occupy the same
type of habitat.

In details of the fruit (ie. shape, presence of longitudinal folds and a narrow
margin) a definite affinity is indicated with B. Stuartii Benth,, but a pappus of the

type present in B. petrophila has not been previously described in the superspecies.

148 REVISION OF TIE GENUS BRACHYCOME CASS. PART ITI,

Superspecies tenwiscapa.
Amended Key to the Species.

(1). Herbs with a glandular indumentum. Fruit cuneate and flat.
(2). Leaves radical, oblanceolate to ovate-cuneate, distally dentate, with sheathing bases.
Fruit 1-2 mm. long, 0-9 mm. broad, smooth, with slightly thickened margins.
IZM oP OOK STONE EOC” WAIL). © idaho oo o-p-d.o-0 bd oo. opD Strom Old odo blO.0 B. tenuwiscapa.
(2).* Leaves cauline, orbicular in gross outline, dentate to almost crenate, usually sessile.
Fruit 1:9 mm. long, 0:9 mm. broad, with a longitudinal fold down each side
and smooth margins. A few tubercles are present on the central area. Pappus
conspicuous and straw-coloured, bristles grouped in bundles of irregular length
AEE Rae TARA Laon Ry Bethe TA APIS, Ge LCR elas Ruacianitc-s eae Dta cy. 0.-C “O -aeeROec Non eG 8 Unc. B. petrophila.

(1).* Glabrous herbs with radical leaves.

(3). Leaves entire, linear to oblanceolate or narrow-obovate.
(4). Base of the plant surrounded by the dead remains of former leaves. Fruit
2-3-3 mm. long, 0:8-1:2 mm. broad, glabrous, the margins slightly thickened
Bliovol Sranoyoidoy - JERVIS SINOIAE SoaucoosousoocudubddooooesuououS B. scapigera.

(4).* Bases of dead outer leaves not persistent.

(5). One to four leafy stolons present on each plant. Fruit 1:8 mm. long,
0-9 mm. broad, cuneate, thick in central region, with a narrow margin.
RappuswCONSPICUOUS! we: cas eueteeesk el Cheese Dksdonee sale ere nate there ele B. stolonifera.
(5).* No development of stolons. Fruit 2:-3-2-5 mm. long, 1-1-3 mm. broad,
obovate, smooth and flattened. Pappus short ............ B. obovata.
(3).* Leaves pinnatisect, crenate or acutely toothed. ;
(6). Leaves oblong-cuneate to elliptical, crenate to acutely toothed distally. Fruit
3 mm. long, 1-4 mm. broad, with relatively long straight hairs on each flat
surface and a shallow longitudinal groove down the outer edge of each
slightly thickened margin. Pappus short .........-......... B. decipiens.
(6).* Leaves pinnatisect, the segments entire or again divided. Fruit 0-9-1-7 mm.
long, 0:5-0:9 mm. broad, with two longitudinal folds on each face. Pappus
Shores bristles motesimne quae emeiuhmeeeresrcsrry eae sierenen iemeam ese n B. Stuartii.

Since the publication of Part I (Proc. LInn. Soc. N.S.W., Ixxiii, 1948, 142-241) of
this series further specimens have been examined from the herberia of two private
collectors. In addition, a large amount of unworked material collected by Mueller
and his contemporaries has recently been found in the National Herbarium, Melbourne.
Although the majority of these specimens. are from well-collected localities, and therefore
of no great interest, a number of them establish new locality records and extend the
known range of certain species, in some cases, considerably. These fresh records are
recorded under the appropriate species, the present location of each specimen being
indicated as follows:

National Herbarium, Melbourne (MEL).
F. A. Rodway (FAR).
N. A. Wakefield (NAW).

Superspecies TENUISCAPA.
BRACHYCOME TENUISCAPA Hook. f., var. a TENUISCAPA (Hook. f.) G. L. Davis.

Proc. Linn. Soc. N.S.W., Ixxiii, 1948, 153.
New South Wales: Moona R., Walopa (MEL).

Tasmania: Cradle Mt., 12.1915, F. A. Rodway (FAR); Cradle Valley, 12.1915, F. A. Rodway
(FAR).

This species has not previously been recorded from New South Wales, and
unfortunately the locality has not been traced. Since ‘N.S.W.” is recorded on the
label it is assumed that Walopa is either too small to be indicated on the maps
consulted or is now known by another name.

BRACHYCOME TENUISCAPA Hook. f., var. 8 PUBESCENS (Benth.) G. L. Davis.

Proc. Linn. Soc. N.S.W., Ixxiii, 1948, 153.
New South Wales: Swamp near Tenterfield, C. Stuart, n. 925 (MEL).

BRACHYCOME SCAPIGERA (Sieb. ex Spreng) DC.,
Prod. vii, (1838), 277.
New South Wales; Braidwood district, 3,000 ft., 10,1886, W. Bauerlen (MEL).

BY GWENDA L. DAVIS. 149

BRACHYCOME DECIPIENS Hook. f.,
London Journ. Bot., vi, (1847), 114.

New South Wales: *Near Dubbo, 1882, J. M. Curran (MEL); Upper Macquarie R., 10.1882,
J. M. Curran (MEL).

Victoria: Hume R., 1883. Jephcott (MEL); *Upper Livingstone, Omeo, 1882, Howitt (MEL) ;
Slopes around Omeo, metamorphic schist for 2,200 ft. above sea level, 26.9.1882 (MEL) ;
Gippsland, 1882, Howitt (MEL); Daylesford, 1880, Wallace (MEL); Ballarat, 9.1884,
D. M. Spence (MEL); near Geelong, 1884, Wilson (MEL).

Tasmania: *Victoria Valley, past R. Ouse crossing on road to L. St. Clair, 12.1917, F. A.
Rodway (FAR).

This-species was previously recorded in New South Wales only from the southern

highlands.

BRACHYCOME STUARTII Benth.,
Fl. Aust., iii, (1866), 5138.
New South Wales: Mudgee, Woolls (MEL). ’
These specimens depart from the typical condition in that they are glandular hairy
and the pappus is microscopic. Hitherto this species was thought to be confined in
New South Wales to the New England Tableland.

Superspecies LEPTOCARPA.
BRACHYCOME DEBILIS Sond.,
Linnaea, xxv, (1852), 477.
New South Wales: Lower Edward’s R., Mein (MEL).
Victoria: Little R., Fullager (MEL); Werribee, Fullager (MEL); Wimmera, 1890, J. Eckert

(MEL).

Specimens from the above Victorian localities were intermixed with vegetatively
identical ones of B. leptocarpa F. Muell., an association previously noted (Davis, 1948).
The range of this species is now extended to south-western New South Wales, west of
Port Phillip and western Victoria.

BRACHYCOME ANGUSTIFOLIA A. Cunn. ex DC. var. a ANGUSTIFOLIA (A. Cunn. ex DC)
G. L. Davis,
Proc. Linn. Soc. N.S.W., 1xxiii, 1948, 161.
New South Wales: Foot of the Pulpit Rock, 54 miles south-west of Nowra, Sandstone,

22.3.1947, F. A. Rodway (FAR); Nowra, roadside, 1.1921, F. A. Rodway (FAR); Grassy
Gully, Shoalhaven R., 14 miles west of Nowra, moist slope, 3.5.1941, F. A. Rodway (FAR).

BRACHYCOME ANGUSTIFOLIA A. Cunn. ex DC. var. 8 HETEROPHYLLA (Benth.)
G. L. Davis,
Proc. LINN. Soc. N.S.W., Ixxiii, 1948, 162.
New South Wales: Joadga (Southern Tablelands), 10.1919, F. A. Rodway (FAR).
Victoria: Prince’s Highway at Karlo Crk., Mt. Drummer and Wingan River, E. Vic., 25.1.1947,

J. H. Willis (MEL).

Previously this species was only recorded from New South Wales, where the
southern limit was Newcastle, so the range is extended considerably. These specimens
differ from the type series in that the leaves are orbicular and regularly dentate, and
Bentham’s type specimens are to be regarded as representing one extreme of a
variable series.

BRACHYCOME DISSECTIFOLIA G. L. Davis,
Proc. Linn. Soc. N.S.W., Ixxiii, 1948, 163.
Victoria: Moyston, 10.1881, D. Sullivan (MEL); Mt. William Creek, 10.1879, D. Sullivan
(MEL).
This species has not been previously recorded from Victoria.

BRACHYCOME PROCUMBENS G. L. Davis,
Proc. Linn. Soc. N.S.W., Ixxiii, 1948, 164.

Victoria: Gorge country of the Upper Snowy River, near Deddick, 21.1.1948, J. M. Béchervaise
(MEL).

The fruits of these specimens show a small variation from those of the type series
in that the wing is approximately equal to the breadth of the body.

150 REVISION OF THE GENUS BRACHYCOME CASS. PART IL,

This is the first record of this species from Victoria, where it appears to occupy
a similar habitat to that of specimens in New England.

BRACHYCOME LINEARILOBA (DC) Druce,
Rep. Bot. Hach. Cl. Brit. Isles, iv (1917), 610.
New South Wales: Trundle, 8.1916, H. M. R. Rupp (FAR).

South Australia: Charlotte Waters, 1889, W. Schwartz (MEL).
Western Australia: Eucla, 1877, Richards (MEL); Eucla, 1882, J. Oliver (MEL).

BRACHYCOME GRAMINEA (Labill.) F. Muell.,
Frag. Phytog., i, (1858), 49.
New South Wales: Moonie Creek, Jervis Bay, 4.1916, 2.1921, F. A. Rodway (FAR); Tabourie
Island, 15.5.1938, F. A. Rodway (FAR).
This species is not confined to the tablelands of New South Wales, as was previously

stated.

Superspecies BASALTICA.
BRACHYCOME BASALTICA F. Muell. var. 6 GRACILIS Benth.,
THE, AMSiop. sa (AUS) 5 SLs
New South Wales: Balranald, 1878, Lucas (MEL); Murrumbidgee R., 9.1878, EF. Mueller
(MEL).
Victoria: Swan Hill district, 10.1888, C. French (MEL); Albury, 1890, J. Wilson (MEL) ;
Western border of Victoria, C. Walter (MEL).
These records are interesting in that they link up those already recorded and

support the suggestion advanced previously (Davis, 1948) that this variety migrated
to Queensland along the basin of the Darling River.

BRACHYCOME MULTIFIDA DC, var. a MULTIFIDA (DC) G. L. Davis,
Proc. Linn. Soc. N.S.W., Ixxiii, 1948, 181.

Victoria: *Bolwarra, near Ballarat, 1882, W. H. Wooster (MEL); Bendigo, 12.10.1892 (MEL).
These specimens extend the range of this species into central Victoria.

Superspecies ACULEATA.
BRACHYCOME PAPILLOSA G. L. Davis,
Proc. Linn. Soc. N.S.W., lxxili, 1948, 191.
New South Wales: Murrumbidgee R., 9.1878, F. Mueller (MEL); Murrumbidgee, 1885, Bruckner
(MEL). -
BRACHYCOME MUELLERI Sond.,
Linnaea, xxv (1852), 475.
South Australia: Yorke’s Penin, 1888, Beythieu (MEL). é
In this specimen the fruits are smaller than previously recorded, being 1:3 mm.
long, 0-9 mm. broad.

BRACHYCOME MUELLEROIDES G. L. Davis,
Proc. LINN. Soc.. N.S.W., Ixxiii, 1948, 194.
Victoria: Granite summits of the mountains on the lower Snowy R., 1.1874, F. Mueller (MEL).
This species was known previously only from two localities, Wagga and Nathalia,
consequently further records are of considerable interest in determining the limits of
variation. The specimens from the Snowy River bear fruit 1 mm. long and 1 mm.
broad, so that they are slightly larger than those of the type series. A further
difference is apparent in that the body of the fruit is dark brown and the wings golden
brown. Vegetative variation is shown in the leaves, which bear 3-6 acute linear lobes.

BRACHYCOME CARDIOCARPA F. Muell. ex. Benth.,
Fl. Aust., iii (1866), 517.
New South Wales: Upper Darling R., near Queensland border, 1884, L. Henry (MEL).
Victoria: Lower Glenelg R., 1891, Eckert (MEL).

Tasmania: *Cape Portland, 1884, Bandinet (MEL); Lake Crescent, 9.12.1938, E. P. Rodway
(FAR).

Hitherto this species has been recorded in New South Wales only from the Southern
Highlands.

BY GWENDA L. DAVIS. 151

BRACHYCOME DIVERSIFOLIA (Grah. in Hooker) Fisch and Mey, var. y DISSECTA
G. L. Davis,
Proc. Linn. Soc. N.S.W., Ixxili, 1948, 202.
Victoria: Treasure’s Homestead, Dargo High Plains, E. Vic., 4,500 ft., 28.1.1946, J. H. Willis

(MEL).
This variety has not previously been recorded from Victoria.

SuperspecieS TESQUORUM.
BRACHYCOME BLACKII G. L. Davis,
Proc. Linn. Soc. N.S.W., xxiii, 1948, 206.
Central Australia: James Range, 3.1883, Kempe (MEL).

Subgenus MrETABRACHYCOME G. L. Davis.
Superspecies IBERIDIFOLIA.
BRACHYCOME IBERIDIFOLIA Benth.,
Enum. Pl. Hueg. (1837), 59.
South Australia: Mt. Eba, 1880, E. Giles (MEL); between the Alberga R. and Mt. Olga, 1873,
Giles (MEL).
BRACHYCOME TATEI J. M. Black,
Proc. Roy. Soc. S8.A., lii (1928), 227.
Western Australia: Eucla, 1889, J. D. Batt (MEL).
Based on these specimens the following notes can be added to the redescription of
this species (Davis, 1948).
“Receptacle 2-5 mm. broad, 1 mm. high, conical, shallowly pitted.”

BRACHYCOME PARVULA Hook. f. var. a PARVULA (Hook. f.) G. L. Davis,
Proc. Linn. Soc. N.S.W., Ixxili, 1948, 212.
Victoria: Lakes Entrance, 12.1878, D. Sullivan (MEL); Gippsland, 1882, Howitt (MEL); Nhill,
St. Eloy D’Alton (MEL); Lower Glenelg R., 1891, Eckert (MEL).
Tasmania: Cape Portland, 2.1885, Baudinet (MEL).
This variety is now known to extend along the whole coast of Victoria and to occur

also in the western district. In Tasmania the only definite record hitherto was from
the Gordon River on the west coast.

BRACHYCOME PARVULA Hook. f. var. 6 LissocarRPA (J. M. Black) G. L. Davis,
Proc. LINN. Soc. N.S.W., lxxiii, 3-4, 1948, 213.
Victoria: Port Phillip, 1891, J. G. Luchmann (MEL); Ballarat, 1887, G. Day (MEL).
A specimen was examined from the Wimmera district (1893, W. E. Matthews,
MEL), which in size and general appearance very closely resembled this variety
except that the leaves were entire.

BRACHYCOME PUSILLA Steetz,
Pl. Preiss, i, (1845), 426.

Western Australia: Between Dundas Hills and Lake Lefroy, 1893, J. D. Batt (MEL).
This constitutes the most easterly record of this species.

BRACHYCOME EXILIS Sond.,

Linnaea, xxv (1852), 449.
New South Wales: Between Bogan and Darling Rivers, 1877, L. Morton (MEL).
Western Australia: Greenough R., 11.1877, F. Mueller (MEL).

It would now appear that this species occurs in western New South Wales, and
since there are now records from two widely separated localities in Western Australia
(Israelite Bay and Greenough River) it is probably to be found also in intermediate
areas.

Superspecies TRACHYCARPA.
BRACHYCOME TRACHYCARPA F. Muell.,
Linnaea, xxv, (1852), 339.

Victoria: Werribee, Fullagher (MEL).
South Australia: Great Bight, 1875, E. Giles (MEL); Eucla, 1889, J. D. Batt (MEL).

152 REVISION OF THE GENUS BRACHYCOME CASS. PART III.

Hitherto this species was only recorded from western Victoria, and in South
Australia no farther west than Denial Bay.

BRACHYCOME CILIARIS (Labill.) Less. var. a cILIARIS (Labill.) G. L. Davis,
Proc. Linn. Soc. N.S.W., 1xxiii, 1948, 221.
Victoria: Snowy R., 2.1890, W. Bauerlen (MEL).
Tasmania: Near Swansea, 1882, A. Simson (MEL); Port Arthur, 1892, J. Bufton (MEL).
From previous records this variety appeared to be confined in Victoria to the
western districts and no specimens had been examined from Tasmania..

Superspecies CILIOCARPA.
BRACHYCOME CILIOCARPA W. V. Fitzgerald,
SOW WA NiGigeHist: |SOGe iim 905) uae.
New South Wales: Yandarlo via Wilcannia, 1886, B. Kenney (MEL); Cobar, 1883, H. Andrae
(MEL); Parkes, 22.9.1947, E. F. Constable (NSW).
The known range of this species is now extended to western and central New

South Wales.

Superspecies SILPHIOSPERMA.
BRACHYCOME GLANDULOSA (Steetz in Lehmann) Benth.,
FY. Aust. iil (1866;) 521:
Western Australia: Stirling Range, F. Mueller (MEL).

No previous records exist of this species occurring in the coastal area, and although
vegetatively similar to the inland specimens, the growth of some specimens is more
luxuriant. The largest specimen reaches a height of 26-5 cm. and bears 44 inflorescences.
The largest fruit examined was 4 mm. long and 3 mm. broad, but although exceeding
the previously defined upper limits of size shows no structural variation.

BRACHYCOME PERPUSILLA (Steetz) Benth. var. TENELLA (Turez) G. L. Davis,
Proc. Linn. Soc. N.S.W., Ixxiii, 3-4, 1948, 231.

Victoria: Werribee, Fullager (MEL). :
Western Australia: Hast of York, 8.1889, A. Eaton (MEL); Stirling Range, F. Mueller (MEL).

These specimens extend the range of this species to the Port Phillip district of
Victoria and to the south-western area of Western Australia, where it was only
previously recorded from far inland. ©

An atypical specimen was examined from Yorke Peninsula (1879, Tepper, MEL) in
which a height of 5-5 cm. was attained and whose leaves were up to 2 em. long and
entire. In all other specimens examined the leaves were pinnatisect, but in view of
the robust branching habit its affinities seem closest to this variety.

ACKNOWLEDGEMENTS.

My thanks are due to Mr. A. W. Jessep, Director of the National Herbarium,
Melbourne, for forwarding a number of specimens for examination, to Dr. F. A. Rodway,
Nowra, and Mr. N. A. Wakefield, Cann River, Victoria, both of whom placed their
private herbaria at my disposal.

I am also indebted to Miss Greta Baddams, of New England University College,
Armidale, for writing the Latin descriptions of new species.

All examinations and drawings of specimens were carried out with equipment
provided by a Commonwealth Research grant.

Reference.

Davis, G. L. 1948.—Revision of the Genus Brachycome Cass. Part I. Australian Species.
Proc. LINN. Soc. N.S.W., Ixxiii, 142-241.

153

NOTES ON THE MORPHOLOGY AND BIOLOGY OF A NEW SPECIES OF
TABANUS (DIPTERA, TABANIDAE).

By KATHLEEN M. I. ENGLISH, B.Sc.
(Fifteen Text-figures. )

[Read 25th May, 1949.]

INTRODUCTION.

This new species was found on sandy ocean beaches near Narooma on the south
coast of New South Wales. Adult flies were first taken on Mystery Bay beach in
January, 1937; others were taken on the same beach in January, 1938, when also a
pupa was found emerging from damp sand at about 3.15 p.m., from which the fly
emerged about fifteen minutes later. The part of the beach from which the pupa
emerged had been covered by water at high tide between 6 and 8 o’clock that morning.
A week later another fly was found emerging from its pupa in the damp sand.
Altogether five pupae were found, and of these, two died and three adults emerged. A
search was made for empty pupal cases and many were found on this beach and on
others near Narooma.

In November, 1938, during a search for the immature stages of an Apiocerid (Proc.
Linn. Soc. N.S.W., Ixxi, p. 296), ten large Tabanid larvae were found, one large larva
was found dead on the surface of the sand, and six larval exuviae were found in the
sand. Of these larvae five pupated, but only two adults emerged, both females. One
pupated on 24th November and emerged on 15th December, 1938 (21 days); the other
pupated on 19th or 20th December and emerged on 2nd January, 1939 (13 or 14 days).
Fourteen pupae were found in the sand, and of these, eight died and six emerged.
Another pupa found was deformed and from it many small nematodes emerged. Larvae
and pupae were more numerous about the level of high-water mark than farther back
on the beach;. no search was made in the sand near low-water mark because of the
difficulty of sifting the wetter sand. The depth at which larvae occurred was not
determined, as none were found in position, but all sand sifted. was taken from
between two and’ about ten inches from the surface, the top two inches of sand being
cleared away before sifting was commenced.

Adult flies were humerous on the beach in January, many were to be seen resting
on the sand in the sun, mating pairs were seen and one pair was caught. No egg-laying
was observed nor were eggs or young larvae obtained. All larvae found were large,
probably last instar. The flies were not troublesome on the beach, but men who had been
fishing on headlands near by said they had been bitten by similar flies; however, this
is inconclusive evidence, as no biting specimens were procured.

The adult was identified tentatively by Mr. G. H. Hardy as being near to
Tabanus rubricallosus Ric. (1914) from New Caledonia, so specimens were sent to Mr.
H. Oldroyd at the British Museum for comparison with the type and paratype, and
. to Dr. Bequaert at Harvard, who has specimens from New Caledonia. Mr. Oldroyd
instanced a number of small differences between the flies from Narooma and the type
series of 7. rubricallosus, but he also considered that the close resemblance between
these two littoral forms should be emphasized. Dr. Bequaert instanced a number of
differences, sufficient, he considered, to constitute a separate species. The most striking
differences are:

1. The presence of erect hairs on the subcallus; no trace of these is to be detected
in his two females of 7. rubricallosus.

N

154 A NEW SPECIES OF TABANUS (DIPTERA, TABANIDAE),

2. The eyes in both sexes are decidedly pilose all over, the hairs being readily seen
with a hand lens. In 7. rubricallosus females the hairs are extremely sparse and short
and not to be seen with a hand lens.

3. The basal (or anterior) bare area and callus of the frons is very extensive,
occupying slightly over half the length, whereas in 7. rubricallosus this bare area
extends only over about one-third of the frons. Also the lower margin of the callus
is only slightly convex, much less so than in 7. rubricadllosus.

4. The antennae are more thickset in the New South Wales species.

5. The New South Wales species is more hairy than 7. rubricallosus; for instance,
the upper part of the frons has many long black hairs; in 7. rubricallosus such hairs
are few and very short.

There are no males of 7. rubricallosus in the other collections for comparison.

No precise locality was given by Miss Ricardo for 7. rubricallosus, but Dr. Bequaert
had been told by the collector that one of his specimens was caught on a beach at Lebris.

TABANUS ORARIUS, Nl. SD.

In Hardy’s key to species of Tabanus with hairy eyes (1939) this new species
runs down to the first part of couplet three and falls into the regisgeorgii group;
group characters are: frons diverging towards the antennae, and thorax with well-
defined dark stripes, four anteriorly and three posteriorly. Included in the group are
the species 7. regisgeorgii Macquart with synonyms 7. spadix Taylor and T. brisbanensis
Taylor, and the species 7. diemeniensis Ferg. These are of a general brownish colour.
The new species can be very easily distinguished from them by the general grey
colour and the more extensive bare area and callus of the frons.

DESCRIPTION.
ADULT.

A medium-sized, grey, hairy species with a large frontal callus.

Female.

Length 13 mm., width across head 5 mm., length of wing 12 mm.

Head: Eyes sparsely covered with fine short hairs of an indeterminate colour
and which can be seen with a hand lens. Behind the eyes is a fringe of erect silvery
hairs. Frons with sides almost parallel, very slightly wider anteriorly, twice as long
as wide, posterior half clothed with white tomentum and dark brown hairs which are
longer on the vertex. Bare area and callus large (Text-fig. 1), extending back half
the length of the frons and reaching the eyes for almost its whole length, lower margin
very slightly convex and posterior margin irregular, shining chestnut brown in colour,
lower centre bare, the rest with sparse hairs, mostly brown. Subcallus with white
tomentum except on narrow median groove, and with a small patch of erect fine
white hairs on each side (Text-fig. 2). Antennae (Text-fig. 2), length 1-5 mm., first
segment with light olive-grey tomentum and clothed with short hairs, mainly black on
dorsal half and mainly white on ventral half; second segment with some grey tomentum
and short hairs mostly black, a few white on ventral surface; third segment broad,
clove brown in colour, with some very short recumbent metallic pubescence. Face
not sunken, covered with white tomentum and fine silvery-white hairs, beard long
and white. Palpi long (Text-fig. 2), extending past the middle of the proboscis, first
segment short and bulbous, second long and tapering to a point, both segments
ochraceous-buff in colour, clothed with fine silvery white hairs, and on the second
segment a very few black hairs also.

Thorax: Dorsum with tomentum in longitudinal stripes of iron-grey or dark olive-
grey alternating with light olive-grey (Text-fig. 3), four main anterior and three main
posterior dark stripes; median stripe very fine anteriorly, short lateral posterior
stripes partly obscured by a fold in the dorsum on each side. The whole dorsum
clothed with erect fine silvery hairs and coarse black hairs, together with recumbent
bronze, and some recumbent black, hairs. Long silvery white hairs form a fringe

BY KATHLEEN M. I. ENGLISH. 155

above the base of the wings and on the edge of the scutellum. The notapleura have
cinnamon drab tomentum and very long hairs, fine silvery white and coarse black.
The ventral surface is covered with tomentum, mostly olive-grey, and densely clothed
with long silvery white hairs. .

Subcallys
with hairs.
i]

OF
Vy,
My, \\

ly,

Ny

Text-figs. 1-3. Tabanus orarius, n. sp.
1. Head of female, front view, x 8. 2. Head of female, side view, x 10 approx.
3. Dorsum of thorax, x 8.

Legs: All coxae and femora covered with olive-grey tomentum and fine silvery
white hairs, except on the inner edge of the fore femora, where the hairs are short
and dark; and on the distal ends of all femora which are bare of tomentum, tawny
in colour, with a few long black hairs. Fore tibiae russet, mid tibiae tawny, on
proximal three-fourths, with many silvery hairs and some black hairs, the distal
fourth and the tarsi mummy brown in colour with short hairs, mainly black. Hind
tibiae russet with black hairs in a line along mid-dorsal surface and covering apex; and
with a very noticeable fringe of long hairs along edges, on inner edge silver hairs
only, except at distal end, where they are short and nearly all black, on outer edge
silver with one or two black hairs on the proximal half, then black and silver mixed
on the distal half. Hind tarsi mummy brown with short hairs mainly black.

Wings: Clear, veins hazel brown, appendix short, stigma inconspicuous.

Abdomen: Dorsal surface covered with tomentum, mostly light olive-grey toning
into light cinnamon drab on posterior and lateral edges of segments 3 to 6. Silvery
white hairs cover most of the first segment and the lateral edges of all segments;
they occur on the posterior edges of segments 2, 3 and 4; and also form a small
triangular patch in the centre of segments 2 to 5; these patches form a median
longitudinal stripe; elsewhere the surface is clothed with short black hairs. The
ventral surface is covered with tomentum, pinkish buff on lateral and posterior edges
of all segments except the first, elsewhere light olive-grey, with short silvery hairs
on all segments and many black hairs also on the seventh segment.

In the female paratypes the colour of the callus varies from hazel brown to
almost black, and the colour of the legs varies correspondingly. The labial palpi are
deformed in one specimen. The cinnamon tinge in the tomentum is more evident in
some specimens than in others, and the dark transverse bars on the abdomen, as
described in the male type, are very evident in some of the female paratypes. The
appendix in the wing varies in length from 0:3 mm. to little more than a sharp angle
on the vein. The wings are damaged in most of the bred specimens. Neave (1915)
states “it is really more difficult to obtain perfect specimens of Tabanidae from bred

156 A NEW SPECIES OF TABANUS (DIPTERA, TABANIDAE),

individuals than from collected ones”, for “Flight seems invariably to take place before
the wings are completely hard and dry”.

Male.

Length, 15-5 mm., width across head, 6 mm., length of wing, 11 mm.

Head: Eyes with large facets hazel brown in pinned specimens and densely covered
with bronze hairs; small facets fuscous in colour and sparsely covered with shorter
hairs. Subcallus with white tomentum except on median groove and in angle between
eyes, where it is bare of tomentum and is fawn in colour, there is a small patch of
hairs on each side, the hairs are shorter and finer than in the female. Antennae not
as broad as in female. Face sunken, same covering as in female. Palpi with first
segment about the same width as the second and more than half as long; second
segment has rounded end; the first segment is light olive-grey at base, the rest of the
segment and the second light ochraceous buff in colour and clothed as in the female.

Thorax: As in female except that recumbent hairs are very sparse and the erect
hairs are longer.

Abdomen: On dorsal surface the anterior border of segments 3 to 5 is bare of
tomentum, and the dark surface with black hairs forms almost black transverse bars.
The hairs are longer except on the median triangular patches, where the silvery hairs
are shorter and sparser, so the central stripe is a little less evident than in the female.

In one male paratype there are no hairs on the subcallus; in the other the dark
transverse bars on the abdomen are less evident than in the type.

The colours in this description have been identified as accurately as possible with
the aid of Ridgeway’s colour chart (1912), a binocular microscope with artificial light
being used to examine the colours of the specimens.

LARVA (Text-figs. 4-10).

The larva is white in colour, the skin striated, shining, and transparent enough
for the internal organs to be seen. A live larva when quiescent measured 22 mm. in
length and 4 mm. in breadth; when active the larva extended up to about 28 mm. At
the anterior end (Text-fig. 4) the larva tapers gradually to the small head; at the
posterior end it tapers only slightly and the last segment (Text-fig. 5) is abruptly
truncated. ;

Head: The head capsule measures up to 5 mm. in length and about 1 mm. in
width, it is mainly cream in colour with some light brown portions and very dark
brown mandibles. The eye-spots are small. Various parts of the head of Tabanid
larvae have been described in detail by Boving (Webb and Wells) 1924, Stammer 1924,
Cameron 1934, and Fuller 1937, and these authors differ somewhat in naming parts.

The antennae (Text-fig. 6) are two segmented, the first segment is long and
cylindrical in shape, the apical segment is much shorter, it is slender and tapers to a
rounded point, and at its base is a very short and slender branch. Hach antenna
arises from near the end of a structure called by Cameron (1934) “a flattened cephalic
sclerite’, and by Fuller (1937) “an elongated plate of the lateralia’’, this structure is
considered by Boving (Webb and Wells 1934) to be the basal segment of the antenna,
which he describes, therefore, as three-segmented. Philip (1931) devotes a paragraph
of “Special Comment” to this question. Situated just behind the antenna on the
lateralia are two minute structures, probably sense organs.

Mouth-parts (Text-fig. 7): The heavily chitinized mandibles agree in general with
the descriptions of other writers. The maxillae differ in shape from that figured by
Fuller (1937), but resemble the one figured by Boving (Webb and Wells 1934) and
called by him “the proximal part of the mandible’. The maxillary palpi are three
segmented, the basal segment is long, thick at the base and tapering distally; towards
its anterior end on the ventral surface there is a side growth bearing a long hair.
The middle segment is short and thick, the distal segment is about the same length
but more slender, and it bears sensory papillae at the tip.

157

BY IXATHLEEN M. I. ENGLISH.

2 External pore of

Graber’s orga.

il ip Sf. Be zi
Seracular SH.

|

AGU AMM

| B

(GRIND

Text-figs. 4-15. Tabanus orarius, n. sp.
4. Anterior end of larva, lateral view, x 16 approx. 5. Posterior end of larva, lateral view,
x 16 approx. 6. Antenna of larva, x 130 approx. 7. Mandible and maxilla of larva, x 50 approx.
8. Posterior spiracle of larva (with fringe shown on one side only, and filament branches

omitted), x 50 approx. 9. Branched filament of spiracular fringe, x 135. 10. Anterior portion of

Graber’s organ, x 150 approx. 11. Pupa, lateral view, x 5. 12. Anterior end of pupa, ventral

view, x 5. 13. Orbital seta of pupa, x 25. 14. Thoracic spiracle of pupa, x 50 approx. 15.
Posterior segment of male pupa, end view, x 20 approx.

158 A NEW SPECIES OF TABANUS (DIPTERA, TABANIDAE),

Thorax: The prothorax is encircled anteriorly by pale brown unstriated skin
densely covered with minute hairs or spines; this hirsute skin covers the anterior
retractile portion of the prothorax and forms a broad collar on the non-retractile portion
(Text-fig. 4); running back from it for more than two-thirds the length of the segment
are five projections, paired laterals and a single ventral one. At the anterior edge
of both meso- and meta-thorax is a band of hirsute skin, much lighter in colour than
on the prothorax, and on these are small backwardly directed projections situated at
the lateral furrows, along which the hirsute skin is continued in broken lines. On
the ventral surface of each thoracic segment are two groups of fine hairs, one group
on each side of the middle line, five or six hairs in each group. Spiracular openings
occur on the prothorax near the posterior edge, and on the meta-thorax in the hirsute
band at the anterior edge of the segment. The actual evagination of the anterior
spiracles during the prepupal stage described by Philip (1931) and Cameron (1934)
was not observed.

Abdomen: At the anterior border of segments one to seven are pseudopodial
cireclets, each composed of four pairs of retractile prolegs; the ventral, latero-ventral
and lateral pairs are more or less rounded and prominent, the dorsal pair is only
slightly raised and is more or less elongated. In conjunction with these locomotor
swellings are more or less complete bands of hirsute skin. On the posterior border of
segment seven there is a wide complete band of hirsute skin, which is hidden when
the larva contracts. On segment eight brown hirsute skin forms a wide collar round
the spiracular prominence; it covers the ridges round the anus and forms two
irregularly shaped patches on each side of the segment. Between the ventral pseudopods
on segments one to seven is a pair of fine hairs, one on each side of the middle line.
Isolated single fine hairs occur also on the thorax and abdomen. There are very small
spiracular openings on segments one to seven, posterior to the lower edge of the
lateral pseudopods. Very slender tracheae were found connected to some of these
openings in the last larval exuviae, the only stage secured. Philip (1931) states “close
observation during the act of moulting reveals that tracheal filaments are cast loose
laterally on every segment except the prothoracic and anal’. The posterior spiracle
(Text-fig. 8) is situated on the last segment on an oval area surrounded by a thickened
collar. It has the felt chamber and antechamber depicted by Stammer (1924). The
visible part of the spiracle is in the form of a pair of vertical chitinous ridges each
crossed by a series of bars of thicker chitin, as depicted by Stammer (1924) and
described by Fuller (1937); this external part and the felt chambers are orange-
coloured. The lips of the spiracular slit can be drawn over the chitinous ridges of the
spiracle and so close the antechambers leading to the trachea. A little back from the
edge of the spiracular slit is a fringe of branched filaments (Text-figs. 8 and 9); these
are long enough to extend across the oval area and on to the hirsute skin of the
surrounding collar, and in some preserved specimens this fringe can be seen standing
out just beyond the end of the segment with magnification as low as ten. A group of
four setae is situated at the dorsal end of the chitinous ridges, and other groups
occur on the surrounding area.

Graber’s organ (Text-fig. 10). This organ was not observed in living specimens, as
the larvae had been killed and preserved before close examination was possible. In
the preserved specimens, all large, probably last instar, two black bodies only were
found; and similarly two black bodies were found in the last larval exuviae, in the
posterior section of the tube cast off from Graber’s organ. The external opening of
this tube can be seen in mounted exuviae (Text-fig. 8); it lies above the spiracle
in the fold between the oval area and the hirsute collar.

PuPA (Text-figs. 11-15).
Pupae vary in length from 18 to 20 mm., and the largest was 4 mm. in width
across the thorax.

The head and thorax (Text-figs. 11 and 12) are armed with long slender setae
(Text-fig. 13), the ends of which are expanded, and on the distal surface have a

BY KATHLEEN M. I. ENGLISH. 159

minutely branched coral-like structure. A central canal runs the length of each seta,
and all setae are borne on prominent tubercles. There is a pair of frontal setae, a
pair of anterior and a pair of posterior orbital setae, and on each side there is a
large lateral orbital tubercle bearing two setae. On the dorsal surface there is a pair
of vertical setae and a pair each of anterior and of posterior meso-notal setae. Laterally
there is a pair of basal alar setae. The nomenclature of the setae is that used by
Cameron (1934).

There are no setae on the meta-thorax. The thoracic spiracle is prominent (Text-
fig. 14), and the mesal opening is large, with a pair of minute openings on the
dorsal edge.

The wing sheaths extend to the second abdominal segment.

There are eight abdominal segments. On the first there are two tergal and
three pleural setae on each side; they are long and slender and the ends are only
slightly expanded; they are each borne on a very small tubercle. On pupal exuviae
the expanded ends have usually been broken off the setae on the first abdominal
segment, and sometimes off the thoracic setae also. Segments 1 to 7 each bear
laterally a pair of small spiracles on low backwardly directed elevations. Segments 2
to 7 are divided by longitudinal lines into dorsal, ventral, and paired lateral regions.
These segments each bear a girdle of spines on the posterior half, the girdle is formed
by a posterior series of mostly long spines, and an irregular anterior series of mostly
short spines, but there is considerable variation in the lengths of the spines in both
series. The spines become more numerous and slightly longer on each segment as they
progress backwards. The last segment terminates in an aster (Text-fig. 15) of paired
tubercles each bearing a long slender spine. In male pupae there is a large anal
tubercle with a continuous row of spines beneath it, and in the dorso-lateral comb the
number of spines varies from three to five, usually there are three long and one or
two short spines, and in the dorsal comb there are only two short spines. In the
female pupae the anal tubercle is small and there is a wide median gap in the row
of spines beneath it, the spines in the dorso-lateral comb are usually longer and more
numerous than in the male (as many as seven occurred in one specimen), and the
spines in the dorsal comb are longer and may be four or five in number.

TYPES AND DISTRIBUTION.

Types: Holotype female, allotype male with pupal exuvia, morphotype pupae (male
and female) and larva; six female paratypes (one with larval and pupal exuviae, three
with pupal exuviae) and one male paratype with pupal exuvia. These, together with
slides used in the prepration of this paper have been placed in the Macleay Museum
at the University of Sydney. Paratypes male and female have been placed in the
School of Public Health and Tropical Medicine, Sydney, and in the C.S.I.R. Museum
at Canberra. Specimens were sent to the British Museum and to the Museum of
Comparative Zoology at Harvard University.

Type locality: Narooma, N.S.W.

Distribution: In the collection at the Macleay Museum there are two females from
Sydney, N.S.W., and three from Rockhampton, Queensland.

CONCLUSION.

Three records have been found of Tabanid larvae in beach sand. Surcouf, in
Wytsman’s Genera Insectorum (1921), refers to the finding in sand on the coast of
Brittany of a larva which developed into an adult identified as 7. nigrifacies Gobert,
later included by Szilady in the sub-genus Ochrops. Again in 1922 Surcouf records the
finding of a hundred Taon larvae in damp sand on the shore in Tunisia, and some
more in Algiers; adults obtained from these larvae were described as a new species,
Ochrops seurati. Spencer (1942) records the finding of a larva on the sea shore in
British Columbia from which was obtained a Tabanid identified as a new species of
Hybonitra. Unfortunately no description of any of these larvae has been found.

160 A NEW SPECIES OF TABANUS (DIPTERA, TABANIDAE).

In the larva of 7. orarius are two features, the striated dorsum of the thorax and
the abruptly truncated posterior segment, which occur also in 7. froggati, T. gentilis
and JT. neobasilis described by Fuller (1937), who states “all other species described
have the thoracic segments unstriated on the dorsum” and “all other Tabanus larvae
described are pointed posteriorly, most having long siphons”.

Also in this larva the fringe round the posterior spiracle, and in the pupa the
length, and expanded ends, of the cephalothoracic setae are characters that have not
been found in descriptions of other Tabanidae.

ACKNOWLEDGEMENTS.

The writer is indebted to Dr. E. A. Briggs, Department of Zoology, University of Sydney,
for the use of laboratory accommodation at the Department; to Mr. G. H. Hardy, University
of Queensland; Mr. H. Oldroyd, Department of Entomology, British Museum; and Dr. J. C.
Bequaert, Museum of Comparative Zoology, Harvard, for their interest and assistance in
identifying the adults; to Mr. A. R. Woodhill, Department of Zoology, University of Sydney,
and Mr. D. J. Lee, School of Public Health and Tropical Medicine, Sydney, for their helpful
criticism of the paper.

References.

CAMERON, A. EH., 1934.—The Life-History and Structure of Haematopota pluvialis Linné
(Tabanidae). Trans. Roy. Soc. Hdinburgh, lviii, pp. 211-250.

FERGUSON, EH. W., 1921.—A List of the Tabanidae (Diptera) in the South Australian Museum
with Descriptions of New Species. Rec. S.A. Mus., i, No. 4, p. 374.

Fuuuer, M. E., 1937.—Notes on the Biology of Tabanus froggati, T. genitilis and T. neobasilis
(Diptera). Proc. Linn. Soc. N.S.W., Ixii, pp. 217-229.

Harpy, G. H., 1939.—Miscellaneous Notes on Australian Diptera. V. Proc. LINN. Soc. N.S.W.,
Ixiv, p. 43.

NEAvE, S. A., 1915.—The Tabanidae of Southern Nyasaland, with Notes on their Life Histories.
IONE, JH LIS Wo Iies 4h, To, 74Bile

PuHtuip, C. B., 19381.—The Tabanidae (Horseflies) of Minnesota with Special Reference to their
Biologies and Taxonomy. Univ. Minnesota Agric. Exp. Sta., Tech. Bull. 80, 132 pp.

Ricarpbo, G., 1914.—Species of Tabanus from Polynesia in the British Museum and in the late
Mr. Verrall’s Collection. Ann. Mag. Nat. Hist., 8, xiii, pp. 476-479.

, 1915.—Notes on the Tabanidae of the Australian Region. Ann. Mag. Nat. Hist.,
8, xvi, pp. 278 and 284.
, 1917—New Species of Tabanidae from Australia and the Fiji Islands. Ann. Mag.

INGE. VASES Shexixeyep. i228

RIDGEWAY, R., 1912.—Color Standards and Color Nomenclature. Washington.

STAMMeER, H. J., 1924.—Die Larven der Tabinaden. Zeitch. fiir Morph. und Okcologie der Tiere,
i Band, Berlin, pp. 121-170.

Surcour, J., 1921.—Wytsman’s Genera Insectorum. Fasc. 175, Diptera Fam. Tabanidae, p. 22;
and Supplement, p. 193.

———., 1922.—Note sur un Diptére 4 Vie Larvaire Littoral. Bull. Soc. ent. France, No. 19,
pp. 297-299.

WALKER, F.., 1848.—List of Diptera of British Museum, Pt. i, p. 178.

WesB, J. L., and WELLS, R. W., 1924.—Horse-flies: Biologies and Relation to Western
Agriculture. U.S. Dept. Agric. Bull., 1218, 35 pp.

WHItTsm, A., 1915.—The Diptera-Brachycera of Tasmania. Ite Th Families Tabanidae and
Therevidae. Pap. and Proc. Roy. Soc. Tas., 1915, p. 11.

161

CRANIA IN THE MACLEAY MUSEUM.

By N. W. G. MAcINTOSH,
Anatomy Department, University of Sydney.

[Read 31st August, 1949.]

Contents.
At aie Page.
Introduction EET ROME Te Nat RECs AP ree reo oid oh car eshte ete Makan LUNecdive | BuSW Uncek@ hh duster fetes 6) thee. a Gd:
Analysis Raa ey eae a ee aU tg Skirt GMa tn Ae wh aay es osu d. w.st Patek Loe kay rah “Epo mos Loma, wae OD
History PMR Spt Ue Ra hal SOG out ode DEE rete ihe peer) Say) a ey ists 2 = euelp canes Cuiener daa eal O8
IEresent Status.) a... cae eerie ager Cunt Meares bos tedMeat er Nitec son wera Ro eeuuen |G
Historical aspects meleinen to ine eotlection®
NDOSULACESEEelauine to thes pastestatusmor the collectionman) Gell se slate eee) so e8) 1 64
Expeditions possibly related to the collection .. .. Beeh we Sorehom pense) cegee s ee LOT:
References in literature to Crania now located in the MEKIAG Meeunes
Exhibitions of Crania MPR TIE Ae: AeA EAS my Peal Ardsley eh Yea Mus meet, eee eT BS
Wilhelm Krause ‘ove fic TCR seri a cae art Ne el see PICaMe CAM, We ip LEE LP eRe eS 8 emma Besa
N. N. GeeMiilounio Macias ee ENCE Se Leh PMR ce Me USAT YS de Bull Meee EPs eels s ek Gt cel)
Geographical Catalogue.
MOLLeSe Stuaite Groups OLraniaveNOS l= S\cand) SUMMarye ase el eeole es) se eect | en Te:
Admiralty and Ninigo Group, Crania Nos. 19-22, and summary .. .. Sie eerie Wepre LIfe)
North East Coast of New Guinea Group, Crania Nos. 23-38, and summary Sa eat ee eS,
Philippines and Hast Indonesia Group, Crania Nos. 39-45, and summary .. .. .. .. 180
USI nine GrOUD as CraniagieNOS4 6-40 sande SUmMIManya iy, le ucie Fie een ini a) eee ell Sell
Various Regions, Crania Nos. 71-81 .. .. Seudtes ae: Pee Ree AL Saat ech ital pices orci a beer il Lteie
IP Ghrowio, (Ciena INOS: Sob), Ennel Gibkeaban po Ge Nao ego oo. pe ee © Sol pac ea al3hs
Viti Levu Group, Crania Nos. 96-99, and summary Dey Lb ese! Um eneceey cota aueO eee eM tte ae be woe D6eh59
Various Melanesian Islands, Crania Nos. 100-103, and summary .... .. .. .. .. 186
Mallicolo Group, Crania Nos. 104-129, and summary Sooay Merole Pach eee sent cre ah Oe iirc’ UM ier Wepre sibel ieoi(e)
Vivant CAvGl Origen as pete sete Meese rae a) EAS SE ie et el ee eet. SA Our FREE Sos
VEEN GH CS ane ROT ER ie ear, rn er ame nay de eats ire tet cee (as. Moet ite okt EG LE TE QIG
Miscellaneous ob ie Lee Bima Rope RES Ge DG IE Co eRe iry patie hry 2 MOON ot GMS EPL SE SAR MD AL dentine [°C
Acknowledgments =e oi ae Bi een Me ai aie More aie BA tg tes oes EP ie ea ee ee LSD)
Bibliography POL a ge ye bees Rete) eae Bae Race RP eaewer SRI Baie se a wae BNR ao ney ea ciel ects)
INTRODUCTION.

A year ago Messrs. Lee, Henry and Woodhill asked the writer if he would examine
the craniological collection in the Macleay Museum and offer some suggestions or
assistance.

The skulls were found scattered in different parts of the Museum, some wrapped in
brown paper, some in canvas bags, some lying loose among a variety of other specimens
(shells, grass skirts, etc.). The majority of the mandibles were found piled here
and there in loose heaps. No catalogue could be found. A heavy coating of dust and
other accumulations concealed any identification such as labels or markings on the
skulls themselves. Some paper labels which had originally been attached to the skulls
were lying on the floor of one cabinet.

The material was first segregated into groups, on the basis of morphological
appearance. Then the mandibles were restored to their rightful crania, by comparing
texture of bone, relationship of condylar process to mandibular fossa, age similarities,
general morphological congruities, and character, colour and fit of teeth.

The crania were cleaned, care being taken to preserve gummed paper labels, ink,
or pencil inscriptions or numbers as they emerged from the coating of grime. Highteen
of the crania had no attached documentation of any kind.

The literature was searched for data relevant to the labels or inscriptions remaining.
The inscriptions in some cases required translation from Russian to English, and the
geographical location of these inscriptions (village names) had then to be discovered.

O

162 CRANIA IN THE MACLEAY MUSEUM,

The frontal regions of the skulls were then marked with consecutive numbers in
Indian ink, a typed card index was provided, and the collection set up in two large
glass display cabinets, where comparison and contrast of the geographical groups can
be seen at a glance.

A catalogue of the crania is appended, recording the original data—i.e., serial
numbers, paper labels, ink and pencil inscriptions, symbols—that accompanied each
skull. This is followed by comments for further identification.

The collection now consists of one hundred and twenty-nine crania (including
fifty which are complete with mandible), three full skeletons, two mummified bodies,
and thirteen mandibles.

Among the best preserved skulls are twenty of ethnological interest in which over-
modelling with clay or fibre, or integumentary mummification has been attempted.

Five of the crania must remain in the category of “unclassified”. Four have the
base missing and gaps near ‘the bregma and suggest head-hunting trophies. Two have
gaps in the temporal bones and a missing base suggesting artificial procedure of a
cannibal or ritual nature. Two show pre-mortem trauma with healing. Twenty-seven
are artificially moulded (deformed) skulls.

In order that the documentation of these crania may be as complete as possible,
considerable research into their past history has been undertaken. Reviews of the
history of the collection as a whole and of any later references to them in the literature
are first presented as part of the evidence on which is based their individuai
documentation. Also an attempt has been made to establish the story of these skulls
particularly in relation to their collectors. In this latter aspect it is necessary to
indicate that Maclay (Baron Nicolai Nicolaevitch Miklouho-Maclay) and Macleay (Sir
William Macleay) are two separate and distinct identities. Although contemporary
and sometimes working together in the same place and on the same material, they
were not related, and a careful distinction between Maclay and Macleay must be kept
in mind.

This paper purports to be no more than a superficial record of cranial material.
Nevertheless its preparation has consumed nine months of spare time and involved
many searches through literature in various languages, often without any relevant
information resulting.

The writer thinks the paper has some value in putting on record material which
has never been fully catalogued and which, for the past fifty years, has been almost
completely overlooked. The material would undoubtedly have been used by anatomists
and anthropologists during that time had its existence been known and data relevant
to it available. ;

ANALYSIS.

Apart from entries in Sir William Macleay’s private diaries or journals of additions
received, the only effort in the past to preserve the identity of the crania has been
(1) the writing of a place name on the skull, which may have been done by the original
collector; (2) the affixing of labels Sie XIII, 1644-1669, probably between the years
1875 and 1888.

It is very obvious that some selection was made and that twenty-six skulls were
serially labelled and placed on display; tit» remainder were consigned to obscurity and
ignored. Labels in a more or less semi-legible and tattered state have remained on 26 of
the selected crania, Sie XIII 1660 being one not found, although two appear to have the
label Sie XIII, No. 1659.

Eleven skulls have pasted above the glabella a 1:0 sq. cm. label with a legible or
partly legible number. Another eighteen have in the same situation a 1:0 sq. cm. area
bearing traces of gum or a shred of paper where a similar label was formerly attached.
Twenty-six of them bear Miklouho-Maclay’s handwriting. There are a further ten skulls
carrying Miklouho-Maclay’s handwriting. Crania Nos. 73 (label 49) and 74 (label 50)
are two heads identified as Maclay’s embalmed Chinese and Malay who were hanged

BY N. W. G. MACINTOSH. 163

in 1880. Cranium No. 3 (label 37) is a curious specimen in which squamous occipital,
parietals and temporals have been removed by disarticulation at the sutures, and the
facial skeleton has been decorated with clay ochre and fibre. The technique differs
rather from that of Darnley and could possibly be from New Ireland. Cranium No. 72
has its inscription printed in English script and there is nothing to identify its
collector or donator.

Twenty-four of these skulls carry a year alongside the place name which coincides
with the date of Maclay’s expeditions to the regions named, but none of them has a
Sie XIII No. .. . label attached to it.

It appears fairly certain, then, that the fifty Maclay skulls presented by Lady
Maclay in 1889 were numbered 1-50 by Miklouho-Maclay himself. After being accepted
by the Macleay Museum they were probably neither catalogued nor exhibited, but stored
out of sight in cupboards, where their numbered paper labels fell off, and their inscrip-
tions gradually became obscured by dust. Forty of these can now be definitely identified
and one more probably.

Twice Miklouho-Maclay lost specimens. In January, 1880, when he transferred
from the Sadie F. Caller to the Hllangowan he left his collection in the former ship
and half of it was lost, apparently consisting of skulls from Noumea, Loyalty, New
Hebrides, Solomons, New Ireland and the Admiralty Group.

There is no record of what cranial specimens Miklouho-Maclay had in the Garden
Palace, but whatever they were, all were lost in its destruction by fire in 1882.

Maclay often found himself unpacking collections about which he had forgotten—
for example, the three skulls obtained ix December, 1872, from HEnglam and Bongu
(Maclay, 1874, p. 246). In 1885 he unpacked a lot of accumulated material which
included the fifty skulls presented by Lady Maclay. It seems probable that the Luzon,
Cebu, Sulu, Timor, Solor crania (Nos. 39-45) belong to his 1873 expedition to the
Philippines and subsequently via Timor to the south-west New Guinea coast.

The Englam, Sangdinbi, Sambul, Bongu, Gorendu, Bili Bili skulls belong to the
1876-77 second sojourn on the Maclay coast of New Guinea. The Ninigo, Andra, Erub,
Mabiak skulls belong to his 1879-80 voyage through the Melanesian islands, Torres
Strait, and south-east coast of New Guinea. The Balonna skulls and the embalmed
Chinese and Malay heads belong to the 1880-81 Australian period of research.

HIstTory.
It would appear that historically the existing material falls into four sections:

1. An original few skulls represented by the massive rugged skull No. 66,
marked “Old Collection”, which is undoubtedly an Australian aboriginal
cranium. No labels, numbers, documentation or record accompanied these.

2. About seventy crania, the product of expeditions or the donations of
collectors. Examples are skulls from Darnley, Derby, Darwin, Cape York, Fiji
and Mallicolo.

3. Occasional crania donated singly or in pairs. Examples are two
Australian skulls, one collected at Bondi by Mr. O’Brien and another obtained
via the Nicholson Museum; also skull No. 62 from Bega, exhibited by Masters,
and skull No. 59, dated 1924.

4. Fifty skulls which had been collected by Miklouho-Maclay.

The interest of the curators in the Macleay Museum has always been, and is still,
predominantly entomological. The least emphasis has been on human skeletal material.
This makes the lack of any deliberate recording. or cataloguing of these crania easier
to understand.

Absence of such documentation as precise locality, tribal affinities, genealogy, time
period, age at death, mode of death, etc., takes from a skull almost all its value.
Without the data that have been collected, correlated and interpreted in this paper
the Macleay crania have unidentified curio value only.

164 CRANIA IN THE MACLEAY MUSEUM,

PRESENT STATUS.

The collection now represents value in the following terms.

The Miklouho-Maclay skulls are unique historical relics of an extraordinary man.
It is doubtful whether an equal number of any of his specimens in any of his fields of
enquiry exists elsewhere in the world. It is also a link with early Australiana and
with the early days of the Linnean Society of New South Wales.

The entire collection is unique in its historical associations in similar, though
perhaps less spectacular terms. A collection of 129 crania, of which 124 are reasonably
documented and classified, though small, is by no means to be despised.

It is worth observing that the Anthropological Section of the Anatomy Department,
University of Sydney, contains only 447 skulls, of which approximately 330 have
reasonable documentation (Macintosh, 1948). The huge collection of the Royal College
of Surgeons, London, was destroyed by bombing during the 1939-45 war. The
Tasmanians became extinct as recently as 1876, yet no more than 200 authentic
Tasmanian crania exist in the world’s total collections (Wunderly, 1939, p. 306).

During the nineteenth century crania of many native races were obtained by
trifling purchase and traded as curios—tattooed Maori heads for example. Today
customs export and import barriers have become rigid; international conventions
prescribe burial for the fallen in battle; legislation aims at the prevention of most
aspects of native exploitation; Western Australia has declared her native regions out
of bounds; New South Wales has at least partial prohibition of the excavation of
native burial grounds. Native peoples in most parts of the world have become
somewhat emancipated, have learned something of White monetary standards, and
either refuse to provide, or ask high prices for, their crania.

The scarcity of cranial material is indicated by the fact that the individual work
of anatomists and physical anthropologists has been for some years performed on
known collections which have already been the repeated subject of scrutiny by a
succession of workers. Krause toured the museums of several countries to describe 200
Australian skulls. Hrdlicka in 1925 visited the museums of the world to place on
record the precise measurements of as large a number of crania as he could find,
recognizing that hew additions would be few in number. That Hrdli¢ka, when visiting
Australia, missed the Macleay crania indicates the obscurity to which they had been
relegated (Hrdliéka, 1928).

HISTORICAL ASPECTS RELATING TO THE COLLECTION.
ABSTRACTS RELATING TO THE PAST STATUS OF THE COLLECTION.

Sir William Macleay kept his collections in.a single building at Elizabeth Bay
until 1876, and subsequently in two buildings in the same grounds. His collections did
not go to the Garden Palace, but following its destruction in 1882, he presented Linnean
Hall, Elizabeth Bay, to the Linnean Society in 1885. The collections were removed to
the University building in 1889-90.

University of Sydney. Chancellor’s address, 13th April, 1889. Hon. William Macleay.
New benefactions:

ce

. consist of Mr. Macleay’s payment to the Senate of the sum of £6000 for a
Curatorship in Natural History, and his handing over to the University of his large
collection. Both had been promised as far back as 1876... . This promise is now
carried out, and the Senate has appointed Mr. Macleay’s own Curator, at a salary of
£300 a year, to remove and take charge of the Collection.”

Calendar of the University of Sydney for the year 1890: “In the year 1874, the
Hon. W. Macleay, M.L.C., undertook to present to the University of Sydney his
collection . . . the collections were, with the exception of the department of Entomology,
removed to the University in the beginning of the year 1889.”

In the Donations Ledger in the Macleay Museum there are only two entries relating
to crania:

BY N. W. G. MACINTOSH. 165

“1889. Oct. 17. Presented by Lady Miklouho-Maclay, 50 Human skulls, 1 skeleton
of child, several miscellaneous bones (human), 5 Human Tongues, 1 ear and portion
of scalp.”

“Web. 15. 1876. Two Fijian skulls—Archibald Boyd.”

In Sir William Macleay’s 1876 diary volume is an entry referring to ‘Mr. J. A. Boyd,
collecting at Herbert River’. In his 1879 diary he refers to Mr. J. A. Boyd of Ripple
Creek, Ingham, making collections from the vicinity of Herbert River. Also in this
1879 volume he writes of Mr. A. Boyd of Fiji sending seven consignments from Fiji
and the New Hebrides: ‘‘Native weapons and implements, and osteological specimens
(human) from Fiji, Mallicolo, and other islands.” There is mention of the material
being too dear, but shortly afterwards he records that purchases were made from
Mr. A. Boyd, Fiji—received March 15th.

Fletcher (1929, pp. 267-269) quotes these two entries and makes the additional
comment that no further particulars can be discovered.

Dr. Cox in a Presidential Address (25/1/1882) refers to purchases from Mr. A. Boyd,
Fiji, on 14th March, of specimens from Espiritu Santo, Solomons and New Hebrides.

It would appear, then, that Mr. J. A. Boyd and Mr. A. Boyd are two separate
individuals, as they would hardly be in Queensland and Fiji simultaneously in both
1876 and 1879.

It is reasonably certain that cranium No. 55, marked in pencil as Herbert River,
was obtained from Mr. J. A. Boyd, of Ingham; and that crania Nos. 82-94 (Fiji)
and Nos. 104-129 (Mallicolo) were purchased from Mr. A. Boyd of Fiji, between 1876
and 1881; while the crania from Viti Levu, Pentecost, Lakoni and Solomons (Nos.
96-103) are probably from the same source.

There are also references in 1879 to material obtained from Mr. Cockerell
(Solomons), Mr. A. Goldie (Port Moresby), and Mr. Goodman (Richmond River).
Cranium No. 60 (Sie XIII No. 1659), Richmond River, seems a probable link with
Mr. Goodman.

The Visitors’ Book in the Macleay Museum contains no entry relevant to crania.

Several drawers in the Macleay Museum contain a mass of loose papers and
correspondence; a single sheet of manuscript was found, dated 19th January, 1892,
unsigned, listing the material in the Macleay Museum. The list mentions the
following: “Six human skeletons, 1 Darnley Island and 1 Peruvian Mummy. Two
hundred and six Human skulls from Australia, New Guinea, South Sea Is. ete.
Ethnological. A fairly representative collection numbering hundreds of specimens
from Australia, New Guinea, and South Sea Islands.” The handwriting is that of
George Masters, then Curator.

The Macleay Memorial Volume, in enumerating the contents of the Macleay
Museum, contains the following paragraph:

“The anthropological and ethnological collections include over 200 crania of
aboriginal Australians and natives of New Guinea and the South Sea Islands, six
entire skeletons of natives of Torres Straits, and many hundreds of specimens of
native weapons, implements, and utensils from Australia, New Guinea, Melanesia ete.

~ (®letcher, 1893, p. xlix.)

A letter from J. J. Fletcher (11/10/1920) to Mr. Shewan, Curator, requests
to see Sir William Macleay’s notebook containing his record of the additions to the
collections between 1874 and 1876. Fletcher thinks he gave the notebook back to
Mr. Masters. He wants to compare the notebook with “Sir William’s private Journals
for 1874-82 (with some blanks) now in my possession” and not available when the
Macleay Memorial Volume was produced. The notebook has not been traced in the
present search, but the private journals for “1873-81”? have been perused.

Letter from Thomas Steel (26/7/20) to the Curator, Mr. J. Shewan: “Could

you inform if circa 1877-79 N. de Miklucho-Maclay deposited specimens illustrative of
‘Macrodontism’ in the Museum .. .?”

166 CRANIA IN THE MACLEAY MUSEUM,

Answer (29/7/20): “I am unable to find any records of donations to Maclay
collection such as you state in your letter. There were no records kept as far as I
know until the collection came to the University ten years later... .”

A letter (25/9/30) from Dr. W. Arndt, Keeper of the Zoological Museum, University
of Berlin, asking for information re specimens of sponges collected by Miklouho-Maclay
from Kamaka-Vallar, S.W. New Guinea, produced the Curator, Mr. Shewan’s reply
(10/11/30): “I have had a careful search through our Museum material and records,
but found no trace....I have also consulted with Dr. Anderson, Chief of the Australian
Museum in Sydney, who confirms my opinion that Baron M. Maclay did not give any
material to Sydney Museums. He is believed to have sent all his material to
St. Petersburg some time before his death, and the boxes are reputed to have been
there unopened some years after his death, regretting that I have no further information
on the subject.”

“Veuillez corriger cette 6épreuve’, dated 1934, from M. de Montessus de Ballore,
Directeur, under the proof paragraph headed “Macleay Museum, 1933, University,
Sydney (founded 1874—W. Macleay)”, gives no mention to crania.

A copy was found of the reply of the Curator, Mr. K. HE. W. Salter (30/11/34). The
text of this reply appears in the 1934 edition, under the Hthnological subheading:
“Large colection of New Guinea native weapons, aboriginal implements and a series of
120 skulls of South Sea Island Natives.” (Index Generalis, Sorbonne.)

Undated. Cyclostyle Notice. Practical Geography II. Ethnology. Technology.
A. Each student must study the examples in the Macleay Museum, from Australia,
New Guinea, Solomon Islands, New Hebrides, Fiji, New Zealand and Samoa. B. With
the permission of the Trustees, Mr. Shewan has kindly agreed to allow students to
handle specimens. These must not be removed from the Museum, etc.

This series of abstracts suggests that Miklouho-Maclay’s cranial collection com-
pletely lost its identity from the time of his death up to the present year, a period
of sixty years, also that all the crania in the Macleay Museum were relegated to
obscurity for about thirty years (1902-1932) and that possibly the majority of the
crania in the collection were never individually recorded.

Among loose papers in the Macleay Museum are two University of Sydney Examina-
tion Books. One is labelled “Contents of Case C, Macleay Museum, 1913”. In it, under
the heading Drawer 39, is written ‘Skulls taken from downstairs, Balance in M.22’. The
other is labelled “Contents of Case M, Macleay Museum, 1913”. In it, under the heading
Drawer 22, is written ‘Skulls, too large for Drawer 29. (Taken from downstairs.) The
handwriting is Mr. Shewan’s.

The deduction could be that the “Balance in M22”, presumably an odd 77 skulls,
represents the difference between the 206 of Masters (1892) or over 200 of Fletcher
(1893), and the 120 of Salter (1934) or the 129 of Macintosh (1948); the six full
skeletons (1893) are reduced to three (1948), and the two mummies are still present.
In other words, almost one-third of the cranial collection disappeared between 1893 and
1934. Yet every single skull of the eleven described by Krause in 1897 is accounted for
in the present collection.

On the other hand, Dr. Cox, in his Presidential Address (25/1/82), when listing
exchanges, mentions skulls, birds, crustacea and three mammals in spirit, sent to Dr.
Bidie, Madras, on 22nd February, 1881.

Could the assessment of Mr. Masters (19/1/1892) have overlooked the skulls sent to
Madras in 1881? If his estimate was made from a notebook of specimens received,
rather than from an actual stocktaking of the material, it might be that the 200 or so
skulls mentioned in the Memorial Volume, should in fact have been recorded as 130
approximately.

Two photographs of the interior of the Macleay Museum were found, both dated
1918. In these the building has no second floor. Rearrangement of the collections
occurred following the building of the second floor, and also when the ground floor was
given over completely to geological material. Finally, following the June 1942 submarine
alarm in Sydney, much of the collection was bundled into cases for evacuation, which

BY N. W. G. MACINTOSH. 167

did not eventuate. During the 1939-45 war a considerable quantity of correspondence,
loose paper documents, calendars, etc., relating to the early 1900’s, or earlier, was given
over to pulp salvage.

EXPEDITIONS POSSIBLY RELATED TO THE COLLECTION.

The original log of the Chevert, 1875, kept in the Macleay Museum, records the
taking of fishing and hunting parties to and from the barque, but no reference is made
to any anthropological matters. The Chevert Expedition had as its object the collection
of “all branches of Natural History”, in the north-east coast of Australia, New Guinea,
and Torres Strait. Places visited included Brooke Is., Coconut Islands, Darnley Is.
(2/52), Dungeness Is., Fitzroy Is., Flinders Is., Palm Is., Percy Is., Somerset (Cape
York), Three Sisters (Sue, Bet, Poll), Turtle Reef, Warrior Is., York Islands and Yule
Islands. .

Sir William kept an independent journal (Diary) for the cruise, the first entry
being on the 29th May, 1875, and the last on the 28th August, 1875.

The following are the only references to human skeletal material. Sat. 19th June

(p. 26). Somerset. Cape York. “Natives dying out ...are lying unburied....’ He
intends to get the doctor to conduct investigations into the cause.
Mriday, 25th June (p. 32): “Mr. Brazier . . . brought back, inter alia, four

aboriginal skulls.” These must almost certainly be the Cape York crania Nos. 50, 51,
52, 53; 50 having the label Sie XIII, No. 1648, 51 having the label Sie XIII, No. 1650,
and 53 the label Sie XIII, No. 1651.

Saturday, 3rd July (p. 38): “Katow River, Mokatta village ... all around the
houses ... remains of animals, human skulls... .” It is probable that crania Nos. 78,
79 and 80, with the word “Papuan” pencilled on them, can be ascribed to this occasion.
Sir William uses the word ‘‘Papuans” (p. 44) for these people, and the pencil writing
of Papuan on the skulls is very like Sir William’s handwriting. The newspaper dated
14th Oct., 1878, wrapped round three teeth and tucked inside one skull would seem to
indicate a scrutiny of the skulls at leisure at a later date.

Monday, 2nd August (p. 49). At Treacherous Bay, Darnley Island. “I went ashore
with Brazier and in the evening I purchased ...a mummified human head.”

Friday, 16th August (p. 50). “I have succeeded in getting several mummified heads
and the mate has gone round in the boat to the village today to get me a complete
mummy which has been promised him.”

Wednesday, 21st August (p. 51). ‘Mr. Williams got the mummy on Friday and it
is now securely packed away with sundry heads.”

In a published preliminary review of the results of the Chevert Expedition, he said
he had not neglected the Hthnological and Anthropological branches, but made no further
comment on these subjects (Macleay, 1875, p. 36).

In his 1876 diary volume, under 4th July, Sir William writes: ‘‘two tables erected
... the other with Darnley Island Mummy and a number of human skulls.’”’ This extract
is quoted also by Fletcher, 1929, p. 259.

Crania Nos. 4 (Sie XIII, No. 1644), 5 (Sie XIII, No. 1645), 6 (Sie XIII, No. 1646),
11 (Sie XIII, No. 1647) are beyond any doubt identified; Nos. 2, 7, 8, 9 and 10 obviously
represent the members of the series which were not selected for display purposes and
were not given a serial number. The selection was apparently based on a desire to
display a variety, i.e, 1 full mummy, 1 integumentary preservation—the male cranium
No. 4, 2 clay restorations—the female crania Nos. 5 and 6, and 1 bare skull—cranium
No. 11. The choice of No. 11 rather than No. 10 was probably influenced by the fact
that No. 11 was complete with mandible.

Cranium No. 1 cannot be identified by deduction. It represents, in company with
cranium No. 3, a departure from Darnley Island technique and may be a Miklouho-—
Maclay specimen.

Walter Wilson Froggatt “In 1886 collected for the Macleay Museum at Cairns,
Russell River, Daintree and Mossman Rivers, North Queensland. In 1887-88 collected

168 CRANIA IN THE MACLEAY MUSEUM,

for the Macleay Museum at North West Kimberley, West Australia, and Derby, Barrier
Range, Leonard and Fitzroy Rivers’. (Musgrave, 1932, p. 101).

The Froggatt Collections received much attention. Macleay exhibited snakes from
Port Darwin, Fletcher plants from Derby, Ramsay marsupials from Derby. Macleay
published notes on the collections from Derby in 1887, and Froggatt gave an anthro-
pological account of the natives of West Kimberley. (Froggatt, 1887, pp. 556, 557,
550, 1017; 1888, p. 651).

Reticence is always more pronounced in the collection and/or donation of human
remains, than in other fields; such reticence may partly account for the absence of
precise information about the origin of many of the Macleay Crania.

It appears quite probable that the Derby and Port Darwin skulls were obtained by
Froggatt.

REFERENCES IN LITERATURE TO CRANIA NOW LOCATED IN THE MAcLEAY MUSEUM.
EXHIBITION OF CRANIA.

“Mr. Masters exhibited the skull of an aboriginal female remarkable on account of
a large oval aperture about 1” x 4” at the junction of the two parietal bones. The
exhibit was the skull of the ‘gin’ of ‘King Charley’ of Bega. The woman had been
accustomed for many years to wear a cap of wet clay upon her head.” (Masters, 1877,
p. 30.) This skull carries the label SIE XIII, No. 1649, and is numbered “62” in the
present arrangement of the collection. Inscribed in ink across the frontal bone is
Coolangubre, N.S.W. (Map of the County of Auckland Hastern Division N.S.W., 1929,
29/11/29, gives a parish of Coolangubra, 7 miles S.W. of Bega.)

This same skull was described by Krause, 1897 (vide infra) as having a parietal
hole 24 x 16 mm., the margins showing exostoses, in a gutter 10 x 5 cm., which begins at
the anterior end of the sagittal suture. Krause calls it a male skull; apparently he was
not aware of its history.

This skull is dolichocephalic, has characteristic Australian aboriginal features and
is probably female. The lesion looks traumatic rather than congenital, suggesting
successful marginal healing, the posterior margin of the aperture being situated at the
mid-point between bregma and lambda, on the sagittal suture line. The measurements
of the lesion are as quoted by Krause, except that the width of the aperture is 18-5 mm.
A slicing blow from a fighting boomerang or a furrow from a bullet might be consistent
with the condition. The wet clay cap mentioned by Masters suggests a protective
mechanism for her traumatic cranial decompression.*

Mr. Masters exhibited “A new species of Serranus and a new Percis from the Fiji
Islands, and two heads of Malicolo Chiefs, showing the skull lengthened out as is done
during infancy. The profiles had been reproduced with prepared clay”. (Masters,
1879, p. 244.)

Sir William Macleay’s 1879 diary volume under Wed., 26th March, refers to the
meeting and comments on the exhibits in these terms: “... 2 skulls of Chiefs of Island
of Mallicolo by Mr. Masters from my museum.”

(Vide supra—the purchase from Mr. A. Boyd in March, 1879, of human osteological
specimens from Mallicolo, Fiji, etc.)

“Mr. Harper exhibited also deformed Malekulan crania to illustrate photographs
sent to him by Mr. R. Parkinson of New Britain, who had recently found among the
inhabitants of South Cape, New Britain, exactly the same deformation as that practised
by the people of South Malekula and the Maskelyne Islets.” (Harper, 1901, p. 311.)

There are 26 deformed Mallicolo crania in the collection, consisting of 19 bare skulls
and 7 with the face modelled in the “Rambaramp” (Deacon, 1934, p. 544) fashion. Of
these, 2 built up skulls and 7 bare skulls have labels Sie XIII, Nos. 1661-1669. The two

*T am indebted to Professor W. K. Inglis for the following comment relative to the same
skull: “On the whole, the appearances suggest that the abnormality is due to a pathological
condition, inflammatory in nature, though it is possible that trauma predisposed to the inflam-
matory reaction.” This opinion was obtained on ist July, 1949, after the manuscript had
been submitted.

BY N. W. G. MACINTOSH. 169

heads labelled 1661, 1662, would be those exhibited by Masters, the 7 bare skulls 1663-
1669 those exhibited by Harper.

HERR WILHELM KRAUSE.

Krause in 1897 says: “W&ahrend meines Aufenthalts in Australien in Sommer 1897
. Die Schidel ... Nr. 34-36 und 38-45 befinden sich im Macleay Museum in Sydney.”

Krause described about 200 Australian skulls distributed in various museums, and
his conclusion was that the mean indices of Australian skulls from various regions
differed very little (p. 519).

He was apparently directed to the cranial collections in the Macleay Museum, the
Anatomy Department, University of Sydney, and the Australian Museum, Sydney, by
J. T. Wilson, Professor of Anatomy in Sydney. In the reprint of Krause’s paper in the
Library of the Anatomy Department, University of Sydney, a marginal note in Professor
Wilson’s handwriting occurs against each description of a Macleay Museum skull.

The eleven skulls in the Macleay Museum described by Krause are:

*

Krause’s No. Present Catalogue No. Other Identification on Skull.
34 60 Sie XIII, No. 1659, Richmond River.
35 46 Derby, N.W. Aust.

36 53 Sie XIII, No. 1651, Cape York.

38 54 Sie XIII, 1652, Cape York.

39 50 Sie XIII, 1648, Port Darwin.

40 62 Sie XIII, 1649, N.S.W.

41 51 ; Sie XIII, 1650, Cape York.

42 48 Derby, N.W. Aust.

43 56 Hinchinbrook Is., N.E. Australia.
44 52 Cape York.

45 58 [35| ¢ H. Austral., Balona R., 1880.

(Krause, 1879, pp. 519, 534, 535, 536.)

BARON N. N. DE MIKLOUHO—-MACLAY.

The classification of Maclay’s skulls presents a threefold problem. 1. The inter-
pretation of the Russian script for the Villages and Islands and/or people written on
the skulls. 2. The geographical placing of these names, which existed over 75 years ago,
but which in the custom of native villages have since changed or disappeared. 3. The
correlation of the named and geographically placed skulls with Maclay’s published
observations and deductions.

The Maclay Coast is defined by Miklouho-Maclay as extending for 150 miles between
Cape Croisilles and Cape King William, and 30-50 miles inland to the mountains of
Mana-Boro-Boro or Finisterre Mountains (Maclay, 1885, p. 713).

The village of Aralu is on the coast between the Gabeneu and Koli Rivers, Gumbu
village is slightly further inland, and Sangdinbimana is near the River Koli, not less
than 1,200 feet above sea-level (Maclay, 1885, pp. 964, 967).

On his last visit to the Maclay Coast in March, 1883, Miklouho-Maclay distributed
seeds from Macassar and Amboina to villages of Bongu, Bogati and Bili-Bili. (Maclay,
1886, p. 354.) His last visit was of one day duration; these can therefore only be coastal.

The geographical situation of some villages is indicated in the following:

Maclay explained to the people of Bongu that he wanted guides to take him to
Kuliku; “. . . ging ik Bongoe en deed den inwoners verstaan, dat ik eenige mannen
noodig had om mij naar Koelikoemana te geleiden’. Footnotes: “Bongoe: Hen dorp
aan de kust (p. 117). Koelikoe: Hen dorp in de bergen—Mana beteenkent berg”
(p. 118). “o.a. had ik medegewerkt tot de herstelling van een Papoea van Gorindoe,
wiens schedel zwaar beschadigd was door den val van een boom” (p. 119). “Van daar,
dat ik aan dien eilandengroep den naam gegeven heb van ‘Archipel der tevreden
menschen’” (p. 121). “de eilanden Bili-Bili en Kar-Kar (Vitias eiland en Dampier
eiland)” (p. 123). “Zoo noem ik de kust, gelegen om de Astrolabe golf en de baai,

170 CRANIA IN THE MACLEAY MUSEUM,

waarin de ‘Archipel der tevreden menschen’ ligt...” (p. 125). “Gabinau—Inlandsche
naam van de kleine baai bij mijn woning” (p. 126).

After a general comment about the Astrolabe Gulf, the neighbouring islands, the
6-S000 feet mountain ridge, and the many scattered villages (p. 277), Miklouho-Maclay
speaks of ‘Die Hinwohner von Sambul- und Seguana-Mana (Bergodrfer) binden an
einzelnen Locken verschiedene kleine Gegenstande ...” (p. 236).

As very few ‘Papuaschadel” are to be found in Huropean museums, he tried very
hard, but could only get 10: ‘mir nur 10 Stuck zu erhalten” (p. 238). The custom is
exhumation after a year, the mandible is cleaned and stored and is not willingly
surrendered; the rest of the skull is thrown anywhere in the village or bush, and can
be taken. With great difficulty he got two full skulls only: “‘“Mit vieler Muhe habe ich
mir zwei vollstandige Schadel verschaffen konnen” (p. 239). He once got five skulls
in less than 10 minutes, brought from different corners of huts of the village, by boys,
for tobacco and calico: “Ich habe ein Mal in einem Dorfe (Gumbu) in weniger als
10 Minuten 5 Schadel erhalten...” (p. 239, footnote). (Maclay, 1873, pp. 225-250.)

Miklouho-Maclay expressed his intention of working out the Papuan skull
thoroughly at a later date; in the meantime he presented a general picture:

Maclay-Kuste: Dolichocephalic, Breadth Index 77, Height 72, lateral view suggests
considerable height, posterior view shows roof-shaped vault; forehead is laterally
compressed, malars very prominent, supraorbitals often strongly developed, occiput
wide but flat; maxillae strongly prognathic with protruding upper teeth; mandible
narrow with angles projecting laterally, and lower teeth receding; chin also receding;
contrast between prominent malars and narrow, low, sloping forehead is marked;
nose is wide and flat. This is the dominant type, but variants exist (pp. 239, 240).
(Maclay, 1873, pp. 225-250.)

Among general observations he says circumcision does not occur in the mountain
villages of Englam, Tengum, Maragum, or in Tiara Island, though it is general and
skilful on the Maclay coast: “In den Bergdorfern Englam-Mana, Tengum-Mana,
Maragum-Mana (Mana heisst Berg), auch der Insel Tiara (eine der Insel des Archipels
der zufriedenen Menschen) fand ich diesen Gebrauch nicht” (p. 244).

Papuans generally die between 50 and 60, because of accumulated ill health; “die
altesten schatzte ich zwischen 50-60 Jahre” (p. 249).

His conclusion is that Papuans are members of one race which falls into not
clearly distinguishable types: “Hin Schluss aber drangt sich schon jetzt, nachdem ich
die Papuas Neu-Guinea’s (der Maclay-Kuste und von Doreh), Neu-Ireland’s, die
Negritos von Luzon (die zweifelsohne echte Papuas sind), die Bewohner der Neu-
Hebriden und der Salomon-Insel gesehen habe, entschieden auf, namlich der, dass der
Papua-Stamm in mehrere von einander distincte Varietaten zerfallt, die aber nicht
schroff von einander geschieden sind” (p. 250). (Maclay, 1873, pp. 225-250.)

In deseribing three skulls obtained in the last days of December, 1872, Miklouho-
Maclay says two are from the mountain village of Englam-Mana and one from the
coastal village of Gumbu, on the Maclay-Kuste (Maclay, 1874, p. 346). t

Other references to locations given by Miklouho-Maclay are: ‘Insel Bili-Bili im
Archipel., Zufriedener Leute; Bili-Bili Jam-Bomba Griger Tiara; Englam-Mana und
Tiengum-Mana; Bergdorfer Tiengum-Mana, Englam-Mana, Maragum-Mana; Bili-Bili,
Kar-Kar (Dampirs Insel), Wag-Wag (Insel Rish). (Maclay, 1875, pp. 77, 78, 83, 87, 88.)

On the Maclay Coast again, the following are indicated: “des Dorfes Bogati;
Bongu; Billi-Billi, Jambomba und den Inseln des Archipel’s der Zufriedenen Menschen;
Insel Tiara; In Englam-Mana (einen Bergdorf); des Dorfes Tiengum-Mana; ‘Bogge’
(Mann von Englam-Mana), ‘Buggay’ kam nach Bongu ... um bei (mein gutter Freund);
‘Saoul’ von Bongu; In Gorendu; In Billi-Billi; ‘Tui’ (Bongu); des Kindes von Gagu
und dem Bau; Mareolan und Lalaulan waren Higennamen Gorima ist der Name eines
Kastendorfes.” (Maclay, 1876, pp. 300, 312, 315, 316, 318, 319, 320, 325.)

Galton’s “reviews” in Nature are abbreviated semi-literal translations of some of
Miklouho-Maclay’s publications in Natuurkundig Tijdschrift, and are not as helpful as

BY N. W. G. MACINTOSH. zl

the originals; for example, 1876, p. 108, briefly refers to “the inhabitants of Bili-Bili
and the Archipelago of Contentment” (Galton, 1876, 1880).

Maclay’s intended regions of investigation, after leaving the Maclay Coast, were
Cebu, Luzon, Sunda Island, and another part of the coast of New Guinea: “Ik maak
den tocht mede en ben van plan onder anderen Cebu aan te doen en de Negrito’s
op Lucon te bezoeken; dan ga ik naar de Soenda-eilanden en van daar naar Nieuw
Guinea, maar aan een ander gedeelte van de kust” (Petermann, 1873, p. 113).

In notes to Petermann, abstracted from a letter to Baer, Miklouho-Maclay says he
arrived in Manila on the 21st March, 1873, went next day by perahu to the coast
village of Limai, the following day to the mountains of Limai, and stayed two and a
half days. The people were negritos, and a typical skull was exhumed for him. “Der
Negrito-Schadel, der fur mich in den Bergen beim Dorfe Pilar (auf Luzon) ausgegraben
wiirde, hatte ebenfalls eine Breite von 89:5” (Maclay, 1874, p. 22).

In a footnote to a letter read by Virchow in 1878, Miklouho-Maclay, referring back
to the same 1873 visit to Cebu, Manila and the Limai mountain, says that after
measuring 21 living Negritos he subsequently, through the kindness of an official in
Balanga, obtained a true negrito skull of which the C.I. of 89-5 was in harmony with
the 21 he measured. “Ich erhielt bald darauf durch die Freundlichkeit eines Beamten
in Balanga einen sicher achten Negrito-Schadel, dessen Breitenindex (89-5) mit dem
durch die Kopfmessung gefundenen Extreme (87-5-90-0) gut stimmte’ (Maclay, 1878,
p. 101).

Maclay supports his opinions by referring to measurements he made on 148
individuals and 23 skulls of the Maclay Coast: “so dass, gestutzt auf meine, an 148
Individuen (beiderlei Geschlechts) gemachten Kopfmessungen und auf die Unter-
suchung von 23 zweifellos achten Schadeln, ich ein ziemlich sicheres Urtheil uber die
Schadel der Papuas der Maclay-Kuste werde fallen konnen” (Maclay, 1878, p. 101).

Maclay regarded the Ninigo people as similar to Micronesians, who were very
different from the Melanesians of Agomes and Kanies (Maclay, 1878, p. 100). The
two Ninigo skulls (Nos. 21, 22 in the present arrangement) are almost perfect in
condition, are of massive appearance, and look very different from any other skull
in the collection.

Carl Baer wanted a perfect brachycephalic skull from Papua and from the
Philippines. Maclay soon concluded that dolichocephaly was by no means as common
in Melanesians as was supposed. He estimated the Cephalic Index of New Guinea
natives as varying from 62—84:3; the 62 was a Namatote Islander of the Kowiay Coast.
He also observed that short heads were more frequent in the women. In New Guinea,
Maclay’s interest in skulls was directed towards the frequency of brachycephaly
(Maclay, 1876, p. 167); while in Australia he was more interested in extremes of
dolichocephaly (Maclay, 1884a, pp. 395-396; 1884b, 401-403).

The difficulty of geographically locating native named villages was recognized by
Miklouho-Maclay himself: “Es ist sehr wahrscheinlich, dass dei Namen ofters veloren
gehen, und statt derselben, neue eingefuhrt werden, denen es wiederum ahnlich gehen
kann.” Apart from that, very few place names existed on the Kowiay-Kuste; he heard
some natives calling part of Triton’s bay “Uru-Lenguru” (Maclay, 1876, p. 160).

Phonetic spelling of native names presents difficulty in geographical diagnosis.
Maclay made his notes in Russian, then translated them into German. He also wrote
in HWnglish, French and Dutch. His spelling varied (Greenop, 1944, p. 265). Note
Bili-Bili and Billi-Billi; Belonne, Balonne, Balonna, Balona River. Amboeang might
indicate Zamboeanga Harbour in Mindanao or a mixed Russian-Dutch phonetic for
Amboina. In either case, why does it occur on a skull also marked Timor? (Skull
No. 44.) Similarly, Skull No. 45 is marked Solor, Kidang, Lomblen; Solor and Lomblen
are separate islands near Timor. The bibliography indicates some of the variations
in the spelling of his own name in his different publications. Some of his comments
written on the cranial bones are initialled M—M, others MM, others B—M, and B—Mkl.

172 CRANIA IN THE MACLEAY MUSEUM,

A single sheet of blue foolscap in the Mitchell Library purports to represent a list
of fifty place names in New Guinea as employed in Maclay. Of names already placed ~
from Maclay’s own publications abstracted above, the list omits Englam, Sangdinbi, and
Gorendu, but confirms Bili-Bili, Bongu and Sambul.

In the villages of the Mariveles Mountains of Luzon he found Negritos who had
customs and facial features similar to Papuans. While admitting that, in general,
the Papuans of the Maclay Coast are dolichocephalic with some variation to brachy-
cephaly, and the Negritos of Luzon are brachycephalic with some variation to
dolichocephaly, he considered that their origin was probably the same (Maclay, 1874,
p. 23). The primitive mountain dwellers on the borders of Pahang, Trengenau and
Kelantan had the heavy brows, thick lips, receding foreheads, kinky hair and skin of
Melanesians; nomadic, unmixed, few in number, retiring before the Malay, they were
to Maclay the faint echo of a Melanesian past. Dark people of the Malay Peninsula,
Negritos, Andamanese, Melanesians, were linked by his observations, whereas previously
the link had been merely presumed. Maclay was looking for similarities. Hence the
Luchon (Sbu) skulls might have come from Luzon or Cebu in the Philippines, from
south-west New Guinea, or even from the Admiralties, or the Maclay Coast.

The map accompanying Maclay’s address to the Imperial Russian Geographical
Society about his March 1879—January 1881 expeditions (Iswestija, 1882—Mitchell
Library) gives no additional solution to the problem.

Luzon in Maclay’s time would receive in Spanish a pronunciation of “Luthon”’.
The Russian language does not provide for “th”, and a Russian phonetic representation
might be ‘“Luchon’’, the “ch” being a soft modification of the pronunciation ‘‘ch” as in
the Scottish “loch”. In Dutch, Luzon is expressed as Lucon in Maclay’s publication;
and in German he writes it as Luzon. A large Russian commercial atlas in the
Geography Department, University of Sydney, shows Luzon as Luson; the Russian
characters are identical, save that Maclay’s Russian X is shown in the map as a
Russian C, that is, an English S. The same problem of the Timor Amboeang and
Solor-Kidang-Lomblen skulls occurs again with the Luchon (Sbu) skull (No. 40). The
English pronunciation of the S is “see’. Why are two separate islands, Luzon and
Cebu, written on the same skull?

The next question is whether the negrito skull exhumed for Maclay in the Limai
Mountains and the negrito skull obtained by the kindness of the official in Balanga
are to be regarded as two references to the same skull or whether they are two
separate skulls. After several discussions with several linguists, and after reconsidera-
tion of Maclay’s story in his own words, and of the appearances of the skulls, the
two skulls marked Luchon and Luchon (Sbu) (Nos. 39 and 40) have been accepted
as the two mentioned in his Luzon excursion. ;

There is one fact against this conclusion: Maclay gives the Breadth Index as 89-5
for one or both Negrito skulls. The Breadth Indices of Skulls 39 and 40 are, by
present-day methods, 74:7 and 74:0. On the other hand, Maclay says he had a makeshift
instrument; he does not define his bony points for measurement, and the indices of
87-5-90:0 which he quotes for 21 living Negritos, are higher than the highest in the
world as quoted by Martin (Martin, 1928, pp. 775-778).

The Russian script in pencil on Skull No. 41, like that on Skulls 39 and 40, is
partially rubbed. Having received preliminary assistance from others (acknowledged),
the writer accepts responsibility for the translation as Sulu (Island or Sea). In
having no base, and in demonstrating large post-mortem temporal gaps, the skull is
very similar to skull No. 40. The bony gaps are similar to those in the classical
descriptions of many writers, of ritual cannibalistic and other procedures, particularly
in the Philippines regions.

Miklouho-Maclay lists places visited by him between March, 1879, and January,
1881, as Lifu in New Caledonia, Tanna, Vate, Tongoa, Mai, Epi, Ambrim, Malo, Vanua
Levu, in the New Hebrides; Lub, Ninigo, Trobriant, Solomons, islands of south-east
end of New Guinea, and the islands of Torres Strait; the terminal period was spent
in Brisbane and on the Balonne River. Also, “In order to eliminate any doubts as

BY N. W. G. MACINTOSH. 173

to the correctness of the cranial measurements on living individuals, I have not
neglected to collect a considerable number of undoubtedly authentic skulls from New
Caledonia, New Guinea, the Admiraltys, Ninigo, and Solomon Islands” (Maclay, 1881,
pp. 171-5).

The Murray Islands consist of three islands, Mer, Uga and HErub; Hrub is now
known as Darnley Island. They lie at the eastern end of Torres Strait. Mabuiag is a
populous island half-way between Cape York and New Guinea, and its Anglicized
spelling is Mabiak. Referring to his observations in 1880 in Mabiak Island, Torres
Strait, and in the south-east coast of New Guinea, Miklouho-Maclay says: “I possess a
skull from one of these villages of the south coast of New Guinea, on which the above
described saddle-shaped ridge is well marked .. .” (Maclay, 1881, pp. 627-630).
Miklouho-Maclay states that he visited the hill villages on Basilaki Island (Moresby
Island) in 1880 (Maclay, 1886, p. 353).

Skulls Nos. 12, 13 and 14 marked Erub (translated from Russian script) and Nos.
15, 16, and 17 marked Mabiak, are obviously Maclay’s, the names being in his own
handwriting. Skull No. 18 is marked in Russian script, which translates possibly to
Mabnyly, 1880. It therefore has to be from Torres Strait or the south-east coast of
the eastern tip of New Guinea. Maclay speaks of obtaining from this southern coast
one skull with a saddle depression 4 mm. deep across the anterior margins of the
parietal bones. Skull No. 18 is the only one in the collection which has such a
transverse depression behind the coronal suture. The precise geographical position
of the name has not been traced.

Maclay said he received brains and skulls, including a range of interesting brains
of Melanesians, from Sydney Hospital in 1881, Sydney being one of the centres of
trade with the Pacific Islands (Greenop, 1944, p. 188, quoting from Maclay to the
Russian Geographical Society). It is worth observing that between 1868 and 1876 over
11,000 New Hebrideans were brought to Australia and approximately 4,000 only of these
were returned. In 1886 there were 10,000 recruits in Queensland (Harrisson, 1937,
pp. 197-8). These included deformed Malekulans, for trade was always concentrated
in the southern part of the group rather than the north. However, Maclay probably
had no relation to the Mallicolo skull collection, because in his fight against the trade
me says: “New Ireland, New Britain, and the Solomon Islands are now the chief
places of kidnapping” (4/3/1884, to Admiral Wilson; quoted by Greenop, p. 223).

Gulnarber Station, near St. George, on the Balonne River, Queensland, was visited
by Miklouho-Maclay in 1880, to examine “hereditary atrichia universalis” in Australian
aborigines (Maclay, 1881, p. 175). Subsequently, ‘Baron Maclay exhibited photographs
and dissections of the brains of a Malay, Chinaman, and Australian Aboriginal;
photographs of the hairless family, Balonne River’ (Maclay, 1881, p. 196).

Greenop went to the Macleay Museum in search of any surviving Maclay specimens.
The Curator, Mr. Salter, showed him two heads; the Sheriff of the Supreme Court,
Brisbane, furnished extracts of official records identifying these as Jimmy Ah Sue,
convicted on 20th April, 1880, at Rockhampton and hanged at Brisbane on 31st May,
1880; and posibly Maximus Gomez, alias Pedro, convicted at Cooktown on 30th April
and hanged on 21st June. Greenop says: “I was able to examine and photograph
these—the only two specimens in existence now, I believe, upon which Miklouho-Maclay
actually worked in pursuit of his studies” (Greenop, 1944, pp. 174, 175; photograph of
Jimmy Ah Sue, p. 160).

Of all the evidence collected, the last quotation from Greenop is the one most
indicative of the almost total eclipse of Maclay’s craniological material for a period
of sixty years.

A Russian film, “Without Prejudice’, as reviewed in The Sydney Morning Herald
(12/10/48), allegedly states: “Sidney’s Anglo-German conspirators callously destroy
Maclay’s all-important anthropological specimens.”

At the present moment 40 skulls arranged on the shelves of the Macleay Museum
can be conclusively identified as Maclay’s and linked with his published observations.

174 CGRANIA IN THE MACLEAY MUSEUM,

Of the total 206 crania (19/1/1892) there are now 129. These 206 included 50 from
Lady Miklouho-Maclay, presented in October, 1889, of which there are now 40, or
perhaps 41, that can be identified. Yet of the 11 Australian skulls examined by
Krause in 1897, six having a Sie XIII No... . label, four having a place name, and
one being a Miklouho-Maclay skull, all have survived.

GEOGRAPHICAL CATALOGUE.

A new series of numbers has been instituted and the skulls arranged on a
geographical basis. The notes include any previous numbers, or distinguishing marks
on the skull; also brief comments of ethnological, historical or anatomical interest.
The date and circumstances under which the skull was obtained are added wherever
possible.

TORRES STRAIT.
1. Skull and mandible. No documentation.

The exterior of the facial skeleton is built up into a clay mask and painted brown
and white. The clay has been pushed into the nasal cavity as far back as the posterior
nares; also into the orbital cavities, and from the lateral aspect into the temporal
and infratemporal fossae. The modelling of the face presents a thin straight nose,
straight lips of medium thickness and slightly protruded. The clay appears to have
been fired after application. The teeth have been retained by the clay and can be
seen from the base of the skull. The skull is intact, the basilar suture ununited.
Possibly female, aged 18. Location possibly Torres Strait Islands; the technique is a
little different from that of Darnley or Mabiak. Possibly a Miklouho-Maclay specimen.

2. Skull and mandible. No documentation.

The tissues of the face and scalp have been mummified, possibly by smoke curing.
Deficiencies have been replaced by rolls of fibre thrust into the infratemporal fossae.
A very thin layer of clay closely follows the outline of the facial bones. The nose is
narrow and straight. A dise of pearl shell, with a central daub of clay for pupil,
occludes each orbit. The integuments over the vault are not covered with clay. The
entire skull is coloured red. Fibre cords hold the mandible to the skull by passing
round the symphysis and through each nasal aperture, and by binding the condylar
neck to the zygomatic arches. Other fibre cords pass coronally round the skull in
front of the mastoid processes, are gathered into a suspensory loop above the skull,
and end in a further decorative set of strings suspending eight nuts and four carved
sticks. The teeth are large and show considerable attrition. Muscle markings are
pronounced in the bare occipital region. Almost certainly male, probably over 40.
The technique is identical with that in specimens 4 and 9. Darnley Island technique.
Chevert Expedition, 1875.

3. Facial skeleton and mandible. Paper label one cm. square with the number 37
pasted on the frontal bone.

A most unusual preparation in which the parietals, the temporals, and the squamous
occipital bones have been removed by disarticulation at the sutures. The remainder of
the skull is intact, including sphenoid, basi-occiput, and condylar occiput. The outer
and inner tables are well displayed at the coronal margin, and the diploe has been in
process of resorption. Clay modelling has overbuilt the face, eyeballs, floor of the
mouth and tongue, the lower jaw being wide open. The teeth show considerable wear
by attrition. A fringe beard of fibre is attached round the mandibular border up to
the zygoma. The clay face is painted black, white and red, the eyeballs being black,
the nostrils red and directed forwards. The appearance is that of a venerable old man.
Almost certainly male, possibly 50. The technique is that of New Ireland. The label
37 places it as one of Miklouho-Maclay’s 1880 collection.

4. Skull and mandible. Paper label SIE XIII, No. 1644, pasted on right parietal.
The tissues of the face and scalp have been mummified, the tissues of the base
removed. A thin layer of clay has been modelled over the tissues of the face, nose

BY N. W. G. MACINTOSH. 175

and orbits. Pearl shell discs, with a central daub of clay occlude the orbits. The
face is coloured red, the vault black. Fibre cords passing through the nares and
round the symphysis are slack, following shrinkage of the tissues. The teeth are
perfect save that three lower incisors have fallen out post mortem; horizontal wear
on the upper incisors almost suggests artificial filing rather than attrition. The
exposed base of the skull shows fire blackening. Almost certainly male, possibly
35. Technique is Darnley, identical with that in specimens 2 and 9. Chevert Expedition,
1875.

5. Skull and mandible. Paper label SIE XIII, No. 1645, pasted on frontal; Darnley
Island, Macleay Museum, printed in Indian ink on frontal region.

The integuments remain on the mid-face only; the upper face is over-modelled
with a thick layer of clay, which probably covered the skull more extensively, but has
since broken away. The moulding of the nose is thin and straight; nostrils and septum
are so arranged as to present a side-to-side pathway for a nose stick. Pearl shell discs
with a central clay daub occlude the orbits. The clay is a dull red colour. Fibre cords
attaching the mandible pass from the symphysis through the nares, and from the
condyles to the zygomatic arches. The remaining teeth show little attrition. One
molar shows advanced caries. Epipteric bones are present. Muscle markings are not
prominent. The line of the suture between squamous temporal and parietal is unusual.
Probably female, possibly over 35. The technique is identical with specimens 6, 7 and 8,
but differs from that in specimens 2, 4 and 9. Chevert Expedition, 1875.

6. Skull and mandible. Paper label Sie XIII, No. 1646, pasted on frontal; Darnley
Island, Macleay Museum, printed in ink on frontal.

The face has been modelled with a thick layer of clay, which follows the contours
of the facial bones. Rolled leaves are inserted in the infratemporal fossae and nasal
cavities and clay superimposed. Pearl shell discs with a central clay daub occlude the
orbits. The face is red, the vault black. The nose is thin and straight, with a septal
aperture for a nose stick. Fibre cords attach the mandible by passing from symphysis
through nares and from condyles to zygomatic arches. The teeth show attrition, but
no caries. Probably female, probably adult. Chevert Expedition, 1875.

7. Skull and mandible. No documentation.

The face is modelled with a thick layer of red clay. The bones of the vault are
blackened. Pearl shell discs with a central clay daub occlude the orbits. Fibre cords
attach the mandible by passing over the symphysis and nares only. The nose is
modelled as in specimens 2, 4, 5, and 6. The majority of the teeth have fallen out
post mortem. Probably female, possibly over 55. The technique is identical with
specimens 5, 6, and 8. Chevert Expedition, 1875.

8. Skull and mandible. Letter M or W incised on right parietal.

The technique is very similar to specimens 5, 6, and 7, but the clay modelling is
more extensive, the entire mandible being included in the clay which extends on to the
basi-sphenoid. The fibre cords passing from the symphysis through the nares are
incorporated in the clay. Fibre and rolled leaves are used as packing for the nasal
cavities. Pearl shell discs with a central daub of clay occlude the orbits, the nose
has a septal perforation, the face is red and the vault and eyebrows blackened.
Probably female, adult. Chevert Expedition, 1875.

9. Skull and mandible. No documentation.

The integuments are mummified, the shrivelled ears well defined, and a very thin
clay reinforcement is present in the upper facial region. The pearl shell discs and
septal perforation are similar to the other specimens, the face is red and the tissues
over the vault are blackened. There are no fibre supports to the mandible. A fragment
of decorative cloth, portion of a head band, still adheres to the left frontal region.

The teeth show no caries and the third molars are unerupted. Probably male, possibly
14. Chevert Expedition, 1875.

176 CRANIA IN THE MACLEAY MUSEUM,

10. Skull. Darnley Is. 1. printed in ink on frontal bone. In pencil on right parietal
appears 725 grms., 1578 cc... .

The teeth show no caries, very little attrition and the incisors are shovel-shaped.
Probably male, adult. Chevert Expedition, 1875.

11. Skull and mandible. Paper label Sie XIII, No. 1647, pasted on frontal; Darnley
Island, Macleay Museum, printed in ink on frontal.

Fibre cords attach the mandible to the skull in precisely the same type of knotting
as occurs in the integumentary and clay heads described above. Some teeth have
fallen out post mortem; the second left lower premolar was lost pre-mortem. Epipteric
bones are present. Chevert Expedition, 1875.

12. Skull and mandible. EHrub Is. 9 in ink on left parietal. Russian script. A one cm.
square area on the frontal indicates the site of a former label.

Corded fibres attach the mandible to the skull in precisely the same manner as
described in the built-up heads, passing from symphysis through the nares, and from
the condyles to the zygomatic arches. A small, delicate, well-preserved skull.
Undoubtedly one of Miklouho-Maclay’s collection, probably 1880.

13. Skull and mandible. Erub Isl. in ink on left parietal. Russian script. Also
possibly a translation of Sangdo. ¢ on the left temporal. Corded fibres attach
the mandible as in No. 12.

A skull very similar in appearance to specimen 12. Undoubtedly one of Miklouho-
Maclay’s collection, probably 1880.

14. Skull. Erub Isl. in ink on left parietal. Russian script. A one cm. square area on
the frontal indicates the site of a former label.
A skull very similar in appearance to specimens 12 and 13. A double loop of cord
fibre braided in similar fashion to the fibre cords already described is loosely wound
round the skull. Undoubtedly one of Miklouho-Maclay’s collection, probably 1880.

15. Skull and mandible. Mabiak in ink on left temporal. Russian script.
Undoubtedly one of Miklouho-Maclay’s collection of 1880.

16. Skull and mandible. Mabiak in ink on left temporal and in blue pencil on left
parietal. Russian script. Paper label one cm. square with the number 38 pasted
on the frontal bone.

Undoubtedly one of Miklouho-Maclay’s collection of 1880.

17. Skull and mandible. Mabiak Isl. M. 1880 in ink on left parietal. Russian script.
The entire skull and mandible are coloured light red. One of Miklouho-Maclay’s.

18. Skull and mandible. Mabnyly ° M. 1880 in ink on left parietal. Russian script.
A one cm. square area on the frontal indicates site of former label.

This skull carries flecks of red colouring matter. There is a slight depression of
the parietals, one cm. behind and parallel to the coronal suture. It is possibly the skull
obtained from a village on the south-east coast of New Guinea and referred to by
Miklouho-Maclay (1881, pp. 627-630).

Summary.

These eighteen Torres Strait crania offer an excellent study of range of genetic
expression within the series; also an opportunity for measurement, detailed description,
and contrast with other Torres Strait series already in the literature. The details of
tissue preservation, remodelling and decoration offer a study of random behaviour super-
imposed on a basic pattern.

Geography.—Erub Island is the former name for Darnley Island and is one of the
Murray Group at the eastern end of the Strait. Mabiak Island is further west, half-way

BY N. W. G. MACINTOSH. 747

between Cape York and New Guinea. Mabnyly may be a village on the northern
shores of the Straits.

Specimens.—Erub (Darnley) is represented by crania Nos. 2, 4, 5, 6, 7, 8, 9, 10, 11,
12, 13, 14; Mabiak by crania Nos. 15, 16, and 17. Cranium No. 1 can only be hazarded
as from somewhere in the Torres Strait area. No. 3 is certainly from New Ireland.

Anatomy.—The eighteen Torres Strait crania in the Macleay series offer a range from
males with heavy supraorbital and occipital ridges to almost delicate female crania.
The males with narrow temporals, low sloping frontals, wide zygomatics, low orbits
and prominent brows, marked alveolar prognathism, and teeth worn by attrition have
distinct Australoid traits. It is doubtful whether the noses are as narrow and straight
as the clay modelling would suggest. The average cranial capacity of the males looks
rather higher than Australian averages. Thomas speaks of the “low Australoid natives
of the Torres Straits Islands” ... “ prognathism and extreme length of palate” .
in one skull ‘‘all the marks of degradism .. . in an exaggerated degree ... the lowest
and most simian human cranium likely to occur at the present day” (pp. 335-6). The
measurements and descriptions he gives are typical of the extreme type of Australian,
Cull, GileY ues

Material—A modelled or mummified head ensures a much greater likelihood of
obtaining a perfect skull. The teeth are less apt to be lost, bones less likely to be
broken, the mandible is practically never missing. Thomas, describing his series of
49 Torres Strait crania, complains that nearly all have lost the greater part of their
teeth. The Rev. S. MacFarlane, who obtained these 49, observed that they were “painted
a deep vermilion which came off as a fine red powder; ... the skulls are rubbed
with earth from Laibi and New Guinea fresh-water swamps; it is a light colour until
burnt, when it becomes a deep red”. His 49 were Banks, Mulgrave and Dauan skulls,
taken by skull-hunting Jervis Islanders, and ornamented with artificial wooden noses
(Thomas, 1885, p. 328). Barnard Davis had no Torres Strait crania in his collection
and could only refer to Pls. 33 and 34 in Dumoutier’s Atlas, showing Tond Island skulls
(Davis, 1867, p. 305).

Social Anthropology.—Haddon (1932, pp. 58-59), however, says that in Erub and
Murray Islands the wizened corpse ... was lashed to a bamboo framework, ... was
painted red, and pieces of mother-of-pearl from a nautilus shell were inserted in the
orbits, a round spot of black beeswax serving for a pupil. After it fell to pieces an
artist who was “skilled in making portrait faces in beeswax on skulls” made a wax
model on the deceased’s face. Haddon says the length of the nose was accurate, as
immediately after death the length of the nose was measured with a piece of wood
which was kept to secure the right proportion of the imitation nose. The modelled
skulls of relatives were kept partly for sentimental reasons, but mainly for divinatory
purposes.

He says (1932, p. 83) that on the sacred islet of Pulu near Mabuiag, skulls were
painted red and provided with noses made of beeswax and eyes of mother-of-pearl, but
this was probably done in only a few cases.

This would infer that Haddon considered that male and female skulls were treated
alike; Haddon apparently did not see the actual procedure. In view of MacFarlane’s
information and of the findings on this Macleay Series, the following interpretation is
put forward.

Crania 2, 4 and 9 show a technique which is essentially integumentary mummifica-
tion with a minimum of clay restoration; they are light red in colour and are probably
all males. Their preparation is identical with that of the Darnley Island mummy in
the Macleay Museum, which is also male. Crania 5, 6, 7, and 8 display a technique
which is essentially clay restoration with practically no attempt at tissue preservation.
Their colour is dark red, and they are probably all female.

Some elements are common to both sets—the pearl shell occlusion of the orbits and
the septal nose-stick perforation.

1

178 CRANIX IN THE MACLEAY MUSEUM,

Within each group the basic procedure has minor variants of decoration; for
example, in the male group, additional sticks, cords, nuts occur in one, a head band
occurs in another.

Cranium No. 1 is probably female and basically belongs to the clay restoration
technique; minor variation lies in the varied colour pattern of brown and white.
Cranium No. 3 is probably male and essentially belongs to the mummification technique,
but variants of the standard lie in a non-perforated septum, forwardly directed red
nostrils, rounded black clay modelling of the eyeballs, absence of pearl shell, bizarre
decoration, and the curious removal of the vault bones to leave a skeletal mask.

This series reveals a general principle for the Torres region; while perpetuation
of the heads of both sexes is attempted, the attitude demands mummification of the
actual tissues of male heads, but merely a clay modelling of the females.

The neat and careful binding of the mandible to the skull in the bare crania Nos.
11 and 12 and the red coloration of both mandible and skull in No. 17, indicate that
the mandible is given equal ritual value with the skull. This is in marked contrast
with the north-east coast of New Guinea, where the mandible is treasured and the
skull thrown away; and with some parts of Indonesia, where the skull is preserved
and the mandible thrown away.

The writer has not had the modelling material analysed and has used the word
clay as a general term. Haddon specifically mentions beeswax, while MacFarlane
speaks of earth from fresh-water swamps.

ADMIRALTY AND NINIGO GROUPS.
19. Skull and mandible. Andra Is. (Mt. Munrach) MM 1879 in ink on left parietal.
Russian script. A one cm. square label with a partially obliterated number
ending in 6 (? 46) pasted on the frontal.
Undoubtedly one of Miklouho-Maclay’s specimens.

20. Skull and mandible. Andra Isl. (Mt. Munrach). MM 1879 in ink on left parietal.
Russian script. A one cm. square label with a partially obliterated number
ending in 5 (? 45) pasted on the frontal.

Undoubtedly a Miklouho-Maclay specimen.

21. Skull and mandible. Ninigo in pencil on the left temporal and on the left mandible.
Also M.. English script. A one cm. square label with a partially obliterated
number, either 6, 4 or 8.

22. Skull and mandible. Ninigo in pencil on a thin cardboard label, tied to a leaf
wrapped round a broken fragment of the left zygomatic arch and inserted in
the left orbit. English script. A one cm. square area on the frontal indicates
the site of a former label.

Summary.

These four crania are undoubtedly of the Miklouho-Maclay collection of 1879, and
represent the only survivors of an apparently large number of skulls which he left
in the American vessel Sadie F. Caller and did not recover. Andra is an island off
the north-east coast of the Admiralty Group; Ninigo is the name of a small group of
islands, equidistant west of the Admiralty Group and north of New Guinea.

Maclay considered the Andra crania to be typically Melanesian; the Ninigo skulls
he thought were very similar to Micronesian crania.

The two Ninigo skulls, Nos. 21 and 22, are large, handsome, rugged; apart from
the small fragment broken from the left zygomatic arch of No. 22, they are in perfect
condition and are possibly the most striking skulls in the collection. They are
probably reasonably rare as well.

NORTH-EAST COAST OF NEW GUINEA.

23. Skull. Mau ¢ Englam 1876. B—M in ink on right parieto-temporal. A one cm. square
area on the frontal indicates site of former label. Russian script.

BY N. W. G. MACINTOSH. 179

24. Skull. ¢ Englam M. 1876. B- in ink on right parieto-temporal. Russian script.
A one em. square area on the frontal indicates site of a former label.

25. Skull. Englam 1876 B—M in ink on right parieto-temporal. Russian script. A one
em. square area indicates site of former label.

26. Skull. ¢ Englam. M. M—M 1876 in ink on right parietal. Russian script.
27. Skull. Sangdinbi. 1876. Russian script in ink on right parietal.
28. Skull. Sangdinbi. M. 1876. Russian script in ink on right parietal.

29. Skull. Sangdinbi. M. 1876. Russian script in ink on right parietal. A one cm.
square area on the frontal indicates site of former label. Paper fragment with
Dutch printing wrapped round bone inside skull.

30. Skull. Sangdinbi. M. 1876. Russian script in ink on right parietal.

31. Skull. 2 Sangdinbi. M. 1876. A one cm. square area on the frontal indicates site
of former label. Russian script in ink on right parietal.

32. Skull. Sangdinbi. M. 1876. Russian script in ink on right parietal. A one cm.
square label with the number 15 pasted on the frontal.

33. Skull. Sambul. M. B-M. 6 1876. Russian script in ink on right parietal and
temporal.

34. Skull. Andan. Bongu. @2 Bongu. B-M. 1877. Russian script in ink on right
parietal and temporal.

35. Skull. Ibor or Ebor. ¢ Bongu. B-M. 1877. Russian script in ink on right parietal
and temporal. A one cm. square area on frontal indicates the site of a former
label.

36. Skull. Keik. 92 Bongu. B-M. 1877. Russian script in ink on right parietal and
temporal. A one cm. square area on frontal indicates the site of a former label.

37. Skull. Panake. ¢ Gorendu. B. Mkl. 1877. Russian script in ink on right parietal.

38. Skull. Bili Bili Isl B-M. 1877. Russian script in ink on right parietal. No base
or facial skeleton. The loss has been by weathering.

Summary.

Geography.—The Maclay Coast extends 150 miles from Cape Croiselles to Cape
King William and 50 miles inland to the 6,000-feet Finisterre Range. Bongu and
Gorendu were villages one mile apart on the sea coast. Sangdinbi, Sambul and Englam
were mountain villages near the Koli River, 30 miles inland, 1,200 feet above sea
level. Bili-Bili or Vitias Island is near Madang.

Material—In his first (1872) visit Maclay pointed out the difficulty of obtaining
skulls, and particularly mandibles. He got 10 skulls without mandibles and two with
mandibles from Gumbu and Englam. These he either left in Java or sent to Virchow
or Car! Baer.

In his second (1876-7) visit Maclay obtained 23 skulls, and crania Nos. 23-38 are
16 of these 23.

Anatomy.—Maclay gave the following data: Breadth Index 77. Height Index 72.
Nose wide and flat. MRoof-like vault; narrowed frontals, wide malars, strong supra-
orbitals, flat occiput, alveolar prognathism with protruding upper teeth; narrow mandible
with laterally deviated angles.

Physical anthropologists and anatomists, particularly Germans, for example Poch
(Bondy-Horowitz, 1930), have been over the New Guinea coast extensively, so the
main value of these 16 skulls lies in their historical significance.

They are all without mandibles. Six are marked male, three female, and seven are
not sex labelled by Maclay.

The fact that the five skulls (Nos. 23, 34, 35, 36, and 37) carrying an additional
name in front of the village name are all sex labelled would appear to indicate that

180 CRANIA IN THE MACLEAY MUSEUM,

Mau, Andan, Ibor, Keik and Panake are the personal names of natives known to
Maclay and who had died between his 1872 and 1876-7 visits.

These 16 crania are all on the small side, lack a robust appearance, and have less
sex differentiation than, for example, Australian crania. Similar observations can be
made in over 100 New Guinea crania in the Anatomy Department, University of Sydney.

PHILIPPINES AND EAST INDONESIA.

89. Skull. Luzon derived from Luchon in Russian script in pencil on left temporo-
parietal. A one cm. square area indicates site of former label.

The skull is smoke-blackened, remnants of caked soil adhere to its interior. Its
features are compatible with negrito affinities, but it is mesocephalic. A 1-0 mm. thick
encrustation of reddish clay coats the skuil deep to the smoke blackening.

40. Skull. Luzon (Cebu) derived from Luchon (Sbu) in Russian script in ink on left
temporal, and in pencil on right parietal. A one cm. square label with the number
16 is pasted on the frontal.

The nose is rebuilt with a hard reddish black clay. A 1:0 mm. thick incrustation
of reddish clay coats the skull, which is fire blackened in the parietal and occipital
regions. There is an elusive similarity between skull 39 and skull 40 in anatomical
features, and between skull 40 and skull 41 in treatment.

41. Skull and mandible. Sulu in Russian script in pencil on right temporo-parietal.
A one cm. square area indicates site of former label.
The base is missing, there is a large ragged gap in the right temporal, and a
smaller circular gap in the left temporal. The mandible is attached by ratten cane
bindings from neck to zygomatic arches.

42. Skull. Timor on right temporal in ink, in English script, in Miklouho-Maclay’s
handwriting. A one cm. square area on the frontal indicates site of a former
label.

The base is missing, there is a ragged gap one inch in diameter at the bregma.

43: Skull. Details the same as No. 42.

44, Skull. Amboeang in pencil and Timor in ink, in Maclay’s handwriting on right
temporal. Other details the same as Nos. 42 and 43.

45. Skull and mandible. Solor. Kidang-Lomblen in pencil on the frontal, in ink on
the right parietal. Homme in blue pencil on the right parietal. English script.
Maclay’s handwriting. A one cm. square site of former label on frontal.

The base is missing and a gap in the vault is situated one inch behind the bregma.

Summary.

Crania Nos. 39-45 are undoubtedly Miklouho-Maclay specimens, and probably belong
to the 1873 period.

After long examination the conclusion has been reached that the gaps in these
skulls are post-mortem effects. There is an extensive literature relating to native
cranial operations, their healing, and subsequent cranial appearances. A comprehensive
review of the subject can be found in Ford, 1937, pp. 471-477, and in Rogers, 1937-38,
pp. 321-340.

The region between Java, Borneo, and the Philippines for over two centuries
provides historically substantiated piracy, head hunting and cannibalism. Malay,
Menankabau and Dyak were outstripped in ferocity by the Sulu marauders.

Trophy skulls were suspended in long houses, by a variety of methods, a knotted
rope through the foramen magnum, through a hole knocked in the vault, or by a basket
framework round the skull (Hose, 1926, pp. 138, 140, 145; Mundy, 1848, p. 196).
Breaking a hole in the temporals bilaterally to suck the blood, and breaking away the
base to secure the brain is substantiated in prehistoric and historic cranial studies.

Crania Nos. 42-45 are almost identical with each other in appearance, and suggest
exposure to weathering and bleaching.

S)

BY N. W. G. MACINTOSH. 181

Geography.—Luzon is the largest of the northern islands of the Philippine Group.
The Mariveles Mountains are in the south-west corner of Luzon. Miklouho-Maclay
obtained a negrito skull in Pilar Village in the Limai Hills in Mariveles.

Cebu is a small island in the middle of the Philippine Group. Sulu Island and
the Sulu Sea are to the south-west, between Timor and the Philippines. Lomblen
is an island north-west of Timor. Solor is an island west of Timor.

AUSTRALIA.
46. Skull and mandible. Derby. N.W. Aust. in ink on left temporal.

Described by Krause under 36 in his series. The skull shows the classical Australian
features, and is a representative of the extreme range of strongly developed bony
prominences, namely, supraorbitals, nuchal lines, maxillary tuberosity, external occipital
protuberance, cone-shaped mastoid, and flat occipital shelf with very pronounced
muscular markings.

47. Skull. Derby. N.W. Australia in pencil on left temporal.
Very similar to cranium No. 46, but a less extreme grade of development of bony
prominences.

48. Skull and mandible. Derby. N.W. Aust. in ink on left temporal.

Described by Krause under 42 in his series. The skull has been treated with red
ochre and fired; some soil adheres to the interior. It is much smaller, lighter and
thinner boned than Nos. 46 and 47. Upper incisors have been subject to ritual removal.
The third molars are unerupted, the basi-sphenoid is not fused and the lower face is
asymmetrical. The Australian features so pronounced in Nos. 46 and 47 are lacking.
Facial and alveolar prognathism is marked. Krause calls it a male skull, but a good
case could be supported for its classification as a female aged about sixteen years.

49. Skull. Derby, N.W. Aus. in ink on left temporal.
Very small; bone disarticulated at the sutures; third molars not erupted. Frontal
suture obliterated. May be a child of nine.

50. Skull and mandibie. Port Darwin. Macleay Museum in ink on frontal and right
mandible. Paper label Sie XIII No. 1648 on right parietal. V in ink on left
mandible.

Described by Krause under 39 in his series. It is a large skull, but lacks the
hypertrophy of bony prominences; the bones are smooth and rounded; the extreme
Australian characters are not obtrusive. A pterion point is present on the right, an
epipteric bone on the left side.

51. Skull and mandible. Cape York Macleay Museum in ink on left temporal and
mandible. Paper label Sie XIII No. 1650 on frontal. IX in ink on left mandible.

Collected by Brazier of Chevert Expedition; Somerset, Cape York, 1875. Described
by Krause under 40 in his series. Epipteric and Wormian bones in the sutures and
orbits. Bony prominences are rounded off. Occipital shelf is undulant.

52. Skull and mandible. Cape York in ink on frontal and right mandible.
Probably obtained by Brazier, Chevert Expedition, Somerset, Cape York, 1875.

53. Skull and mandible. Cape York in ink on frontal. Paper label Sie XIII No. 1651
on frontal, VIII on left temporal, X on left mandible.
Collected by Brazier, Chevert Expedition, Somerset, Cape York, 1875. Described
by Krause under 36 in his series. A very similar cranium to No. 52.

54. Skull and mandible. Cape York Macleay Museum in ink on frontal. Paper label
Sie XIII No. 1652 on frontal. X on left temporal.

Collected by Brazier, Chevert Expedition, Somerset, Cape York, 1875. Described
by Krause under 38 in his series. Very similar cranium to Nos. 51, 52 and 53. These
four skulls are dolichocephalic, large and have reasonable vault height. Like the Port
Darwin skull, they are less rugged than the Derby skulls but show classical Australian

182 CRANIA IN THE MACLEAY MUSEUM,

features. The mandibles were incorrectly linked with the skulls and have been
re-arranged.

55. Skull and mandible. Herbert River in pencil on left temporal (unusual printed
script). Herbert River, Queensland in pencil on frontal.
Probably collected by J. A. Boyd of Ingham, 1876-79. The zygomatics are broken
away. The occipital shelf is flat and the occipital part of the vault is shortened in
the vertical plane. Characteristic Australian features. Skull is weathered.

56. Skull and mandible. Hinchinbrook Island. N.E. Australia in ink on paper label
tied to the skull.
Described by Krause under 43 in his series. A very dolichocephalic skull, the
narrowness is uniform throughout the entire length of the skull. The bones are thin
and light, and there is a lack of excessive bony prominences.

57. Skull. H. Austral. Balona R. 1880 in Miklouho-Maclay’s handwriting in ink on
left temporal. A one ecm. square area indicates the site of a former label.
The calvarium is sawn through and present.

58. Skull. H. Austral. “4 Balona R. 1880 in Miklouho-Maclay’s handwriting in ink on
left temporal. A one cm. square label with the number 35 is pasted on the
glabella.

The calvarium is sawn through and missing. Described by Krause under 34 in his
series; he refers to a colossal canine fossa 10 mm. deep. This skull looks almost a
twin to No. 57. The short vertical face is noticeably in contrast with the wide
bizygomatic. extent.

59. Skull. Cumeroogunja. 1924 in ink on left maxilla.
Individual bones are disarticulated. The bones are thin and delicate. Cumeroogunja
is possibly a station property, location unknown.

60. Skull and mandible. Richmond River Macleay Museum in ink on frontal. Paper
label Sie XIII. No. 1659 on frontal. XI on left temporal and mandible.
Probably collected by Mr. Goodman, 1879. Part of base is missing. The skull is
stained with iron. Small multiple erosions may have been made by ground beetles
or water weathering. A rugged dolichocephalic skull.

61. Skull. Australian p. 1 Kokeruga. M. Tork in ink in Russian script in Miklouho-
Maclay’s handwriting on right temporal.
Serpiginous iron stains on the skull indicate that it had been lying on its left side
in a swamp margin. Multiple erosions are identical in appearance with those of
Skull 60. Collected probably in 1880-81.

62. Skull and mandible. Coolangubrie N.S.W. in ink on frontal. Paper label Sie XIII.
No. 1649 on left parietal.

Exhibited by Masters (1877) and stated to be a known female from Bega.
Coolangubra is about seven miles south-west of Bega. It was described by Krause
under 40 in his series and classified by him as a male. The classical Australian features
are present and bony prominences are moderately rugged. Krause was probably
unaware of the history of the skull. This is a good illustration of the difficulty of
classifying sex with certainty. A gap occurs in the vault between bregma and lambda.
(Vide supra previous discussion.)

63. Skull. Female, N.S.W.? in ink on occipital. 851 in ink on frontal.

The norma verticalis is five-ssided, very wide at the parietal eminences, narrow
at the frontal margins. The skull suggests Melanesian affinities rather than Australian.
64. Skull and mandible. Paper label with the number 1184 on the frontal. A card

with the number. 1184 tucked inside the skull states “portion of a skull found
near Bondi by Mr. F. O’Brien”’.

BY N. W. G. MACINTOSH. 183

The skull is complete, has an excellent set of teeth and shows evidence of weathering
in sand. It is large, characteristically Australian and almost certainly male.
Exceptionally developed alveolar prognathism.

65. Skull and mandible. Paper label with number 1185 on the frontal. A card with
the number 1185 tucked inside the skull states “adult male aboriginal presented
by Sir Charles Nicholson”.

The condition of the skull is identical with No. 64. A smaller, less robust edition

of No. 64. It is much more likely to be a female than a male. In fact Nos. 64 and 65

look like brother and sister.

66. Skull. Old Collection printed in ink on frontal.
An extreme grade of the rugged Australian characteristics, almost approaching
Neanderthaloid type of brow ridge.

67. Skull. No documentation.
Almost certainly Australian. Classical Australian characteristics.

68. Skull. No documentation.
Norma verticalis five-sided, wide parietal bones, narrow frontal. Extreme alveolar
prognathism. Its affinities appear to be those of Melanesia.

69. Skull and mandible. No documentation. A one cm. square area indicates site of
former label, i.e. probably a Miklouho-Maclay specimen.
A well-preserved cranium with some of the integumentary tissues adhering to it
and an excellent set of teeth with no attritional wear. Very dolichocephalic and with
some features suggesting Australian affinities.

70. Skull. No documentation.
Its affinities appear to be very like those of Darnley Island skulls.

Summary.

Documentation which can be accepted is available for crania Nos. 46-65, with the
exception of cranium No. 63, which looks like a Melanesian skull, although carrying the
label “Female, N.S.W.?”’; also cranium No. 59, in which the place name cannot be
located. Of crania Nos. 65-70, which have no label of geographical identity, Nos. 66 and
67 can almost certainly be accepted as Australians, and No. 69 has some Australian
morphological characters. Nos. 68 and 70 remain “Unknown skulls”.

The geographical range or scatter of the 18 crania identified with certainty is very
satisfactory, Victoria, South Australia and Tasmania being the only States not
represented.

For comparison and contrast this small series represents a valuable addition when
studied with the larger numbers of documented Australian crania in the Australian
Museum and the Anatomy Department in Sydney; the last possesses no crania from
some of the sites of the Macleay crania, for example, Derby, Darwin and Hinchinbrook.

The literature concerning the Australian aborigines contains much disagreement.
The writings of Wood Jones would appear to indicate that the Australians form a
homogeneous race which expresses itself over a range from large crania with exaggerated
masculine characters to exceptionally small-headed female crania (Wood Jones, 1934,
Deeoze)) Of the writers who considered the Australians non-homogeneous, it is
sufficient to contrast the following two: Topinard said a smaller dolichocephalic negroid
type belonged to the west, and a larger, less dolichocephalic, non-negroid type occupied
the rest of the continent (Topinard, 1872, p. 211). Birdsell practically reverses this
opinion, claiming a pigmy type in the east, and a Murrayian type and a Carpentarian
type in the rest of the country, so giving the Australian a trihybrid origin (Birdsell,
1946, p. 103; Tindale and Birdsell, 1941, p. 9).

The 18 geographically located Australians in the Macleay Museum present examples
of the extremely rugged large type, the thin-boned lightly ridged type, and intermediate
grades; further, these variations are scattered, not locally concentrated. Crania No. 46
from Derby, No. 50 from Darwin, No. 52 from Cape York, No. 60 from Richmond River,

184 CRANIA IN THE MACLEAY MUSEUM,

No. 64 from Bondi, and No. 66 “Old Collection” are comparable in size, dolichocephaly,
and exaggeration of bony prominences. In addition, however, superimposed on the
basic similarities, there do appear to be local concentrations of modified genotypes.
Howells arrived at this conclusion, although he expressed it in different terms; ‘‘there
are definite regional deviations in a remarkably homogeneous people” (Howells, 1937,
p. 40). As an illustration, the inferior part of the squamous occipital in the large
heavy skulls quoted above is, in the Derby crania, a flat shelf with exaggerated muscle
markings; in the Darwin skull it is uniformly curved and smooth; in the Cape York
crania it is undulant and smooth, and in the Bondi skull the flat shelf reappears.

Apart from this, many miscellaneous subjects for study are available in these
crania, to mention a few examples: epipteric and Wormian bones, a pterion point in
one (Skull 50), metopism, thickness and thinness of cranial bone (a subject on which
the literature is not very adequate), range of cranial capacity—which appears in this
series to be considerable, and so on.

VARIOUS REGIONS.
71. Skull. Marked out in phrenological areas. Probably European.

72. Skull. Sakkara. Egypt printed in ink on the basi-occiput. A one cm. square area
on the glabella indicates the site of a former label.
Apparently a Miklouho-Maclay specimen. No clue to its history is available.
Maclay conducted marine biological observations in the Red Sea region and passed
the Egyptian region several times.

73. Skull and mandible. Hanged, amputated, preserved head. Paper label one cm.
square with the number 49 pasted on the glabella region.
Jimmy Ah Sue, Chinese, hanged at Brisbane (31/5/1880). The brain was removed
by Miklouho-Maclay. The calvarium is missing but scalp and hair are still present.
(See Greenop, 1944, pp. 174, 175, photograph p. 160; also Maclay, 1881, p. 196.)

74. Skull and mandible. Hanged, amputated, preserved head. Paper label one cm.
square with the number 50 pasted on glabella region.
Maximus Gomez, alias Pedro, hanged (21/6/1880). The brain was removed,
dissected and exhibited by Miklouho-Maclay. Bones of the calvarium are missing,
scalp and hair are still present. (Greenop, 1944, pp. 174, 175; Maclay, 1881, p. 196.)

75. Skull. New Zealand in ink on frontal.
76. Skull. New Zealand in ink on frontal.

77. Skull. New Zealand in ink on frontal.
There is no information available for crania Nos. 75, 76 and 77.

78. Skull. Papuan written in pencil on the left parietal. The script was written with
the skull held in the inverted position. A piece of newspaper dated October 14th,
1878, Brisbane, is wrapped round three-teeth and tucked inside the skull.
The bones are bleached by weathering, and the vault stained, suggesting that the
skull was originally lying on its vault in swampy or marshy ground.

79. Skull. Papuan written in pencil on the right parietal. The script was written with
the skull held in the inverted position.
The skull is similarly bleached and the vault similarly stained.

80. Skull. Papuan written in pencil on the right parietal in the inverted position.
The skull is similarly bleached. ;
Crania Nos. 78, 79 and 80 are marked by the same handwriting and it is like

that of Sir William Macleay. These might be specimens from Mokatta Village, Katow

River, described (3/7/1875) by Sir William as lying around the village among refuse

and filth (Macleay Chevert Diary, 1875, p. 38).

81. Skull. No documentation.

Some tissues adhere to the base. Probably female, possibly young, very
brachycephalic.

BY N. W. G. MACINTOSH. 185

FIJI.

82. Skull. Fiji in ink on frontal.

83. Skull. Fiji in ink on frontal.

84. Skull: Fiji Macleay Museum in ink on frontal. Paper label Sie. XIII. No. 1655
on left temporal. Gap in left temporal.

85. Skull. Fiji in ink on frontal.

86. Skull. Fiji in ink on frontal.

87. Skull. Fiji Macleay Museum in ink on frontal. Paper label Sie XIII. No. 1653
on left parietal.

88. Skull. Fiji Macleay Museum in ink on frontal. Paper label Sie XIII. No. 1654
on left parietal.

89. Skull. Fiji in ink on frontal. Facial skeleton missing.

90. Skull and mandible. Fiji Macleay Museum in ink on frontal. Paper label Sie XIII.
No. 1656 on left temporal. VII on left temporal and left mandible, and Fiji.

91. Skull. Fiji in ink on frontal.

92. Skull. Fiji in ink on frontal.

93. Skull and mandible. Fiji in ink on frontal. Mandible fractured post mortem.

94. Skull. Fiji in ink on frontal.

95. Skull and mandible. No documentation.

Summary.

Crania Nos. 82-95 are remarkably similar in size and morphological appearance.
Possibly the majority are female. As a general statement, they are dolichocephalic and
hypsicephalic, with vertical facial outlines and little prognathism, fairly high frontals
and practically no supraorbital prominences.

The Donations Ledger records two Fijian skulls from Archibald Boyd (15/2/1876).

Sir William Macleay’s Diary, 1879, records osteological specimens (human) from
Fiji, Mallicolo and other islands, purchased from Mr. A. Boyd.

Dr. Cox (25/1/1882) refers to purchases from A. Boyd, Fiji, from Espiritu Santo,
Solomons and New Hebrides.

This series of 14 crania accords well with Flower’s description of the cranial
characters of Fijians (Flower, 1880, pp. 153-173).

VITI LEVU.

96. Skull. Vatu Co. on paper label pasted on left temporal.
Face and base missing through weathering.

97. Skull. Vatu Co. E. Coast, Viti Levu Macleay Museum in ink on frontal. Paper
label Sie XIII. No. 1659? on right temporal. Vatu Catho. E. Coast Viti Levu on
paper label on left temporal.

98. Skull. Vatu Co. E. Viti Levu Macleay Museum in ink on frontal. Paper labels on
right temporal Sie XIII No. 1657, and Vatu Co. E. Viti Levu.

99. Skull. Viti Levu Macleay Museum in ink on frontal. Also paper label Sie XIII.
No. 1658.

Summary.

The four crania Nos. 96-99 from Viti Levu are badly weathered and look like
trade specimens. As a contrast group with the previous group they are of some
value; they appear to be more hypsicephalic, particularly No. 99, and more prognathic,
particularly No. 97, than the Fijian Group. No. 98, probably male, has some Australoid
affinities in the facial skeleton and in the inferior part of the squamous occipital.

186 CRANIA IN THE MACLEAY MUSEUM,

VARIOUS MELANESIAN ISLANDS.
100. Skull. Pentecost in ink on paper label pasted on left fronto-parietal.

This skull has been artificially moulded (deformed). The technique appears to
have been circumferential binding in the coronal plane, so that the frontal has been
moderately flattened, the squamous occipital pushed back and up. The binding has
been applied most effectively on the anterior parietals, so that the skull presents in
norma lateralis a moderate hump on the frontal, a more pronounced hump on the
posterior parietals, and a wide saddle between the two humps. This form of deformation
is rather different from, and of less degree than, the Malekulan deformation. This
skull has a metopic suture and a left epipteric bone.

101. Skull. Pentecost Is. New Hebrides in ink on paper label pasted on left fronto-
parietal.

This skull is artificially moulded like No. 100, but to a much less pronounced stage.

102. Skull. Lakoni (Gama) Banks Group in ink on paper label pasted on right fronto-
parietal.
Morphologically this small skull is very like the Fiji crania Nos. 82-95.

103. Skull and mandible. Solomon Is. printed in ink on the frontal.

A well-preserved skull, the mandible is attached by string at the condyles and
mandibular fossae.

Summary.

Crania Nos. 96-103 probably represent purchases from Archibald Boyd between
1876 and 1881. Pentecost and Banks Islands are in the New Hebrides Group, which,
according to most authorities, is populated by at least three racial elements, possibly
four or five. These skulls permit of some comparison with the following series of 26
crania from Mallicolo, also in the New Hebrides.

MALLICOLO.

104. Skull. Mallicolo Macleay Museum in ink on frontal. Paper label Sie XIII. No. 1667
on frontal. III on left temporal.

105. Skull. Mallicolo Macleay Museum in ink on frontal. Paper label Sie XIII. No.
1664 tied to palate. IV on left temporal.

106. Skull. Mallicolo in ink on frontal.

107. Skull. Mallicolo Macleay Museum in ink on frontal. Paper label Sie XIII. No.
1665 on right parietal.

108. Skull. Mallicolo Macleay Museum in ink on frontal. Paper label Sie XIII. No.
1666.

109. Skull. Mallicolo Macleay Museum in ink on frontal. Paper label Sie XIII. No. 1667
on left parietal.

110. Skull and mandible. Mallicolo in ink on frontal. Paper label Sie XIII. No. 1668.
II on mandible and temporal.

111. Skull. Mallicolo Macleay Museum in ink on frontal. Paper label Sie XIII. No.
1669.

112. Skull and mandible. Mallicolo in ink on frontal and on right mandible. IV on
left mandible.

The mandible does not fit cranium No. 105; it undoubtedly belongs to No. 112,
which it fits perfectly. The posterior half of the vault is blackened, and the nares
and orbits are stuffed with fibre.

113. Skull. No documentation. Posterior half of the vault is blackened.
114. Skull. Mallicolo in ink on frontal.
115. Skull and mandible. Mallicolo in ink on frontal.

BY N. W. G. MACINTOSH. 187

116. Skull. Mallicolo in ink on frontal.
117. Skull. No documentation.

118. Skull. Mallicolo in ink on frontal.
119. Skull. Mallicolo in ink on frontal.
120. Skull. Mallicolo in ink on frontal.
121. Skull. Mallicolo in ink on frontal.
122. Skull. No documentation.

Of these 19 crania, three have no name and no label attached. These three, Nos.
113, 117, and 122, have been arranged in the series so that they lie adjacent to the
crania which they most closely resemble. The 19 crania all present artificial moulding
in the same technique but of varying degrees. Nos. 104, 105, 113 and 122 possess a
metopic suture. No. 108 shows advanced obliteration of the cranial sutures. All except
Nos. 108 and 116 present epipteric bones. In eleven crania epipterics occur bilaterally,
in six crania epipteric formation is unilateral, four being left-sided, two right-sided, in
position. The crania all exhibit occipital elongation, sloping up and back from low
flattened frontals; prominent brow ridges, moderately wide zygomatics, high and wide
palates, and facial and alveolar prognathism of moderate degree occur in all these skulls.

123. Skull and mandible. No documentation.
Presents a complete Rambaramp reconstruction, with a wig and feathers on the
vault, and sticks supporting the neck.
124. Skull and mandible. Mallicolo Macleay Museum in ink on frontal region. Sie XIII
No. 1661. in ink on paper label tied to left zygomatic arch.
Rambaramp reconstruction.

N25) Skull and mandible. No documentation.
Rambaramp reconstruction, including wig.

126. Skull and mandible. Mallicolo Macleay Museum in ink on frontal region. Sie XIII.
No. 1662 in ink on paper label tied to left mandible.
Rambaramp reconstruction, including wig.
127. Skull and mandible. No documentation.
Rambaramp reconstruction of the face. A perfect set of teeth.

128. Skull and mandible. Mallicolo in ink on frontal bone.

Rambaramp reconstruction on face and on floor of mouth; remainder of skull bare.
A full set of teeth, upper central incisors have a peg-top appearance unlike those of
No. 127. An epipteric bone is visible on the left side.

129. Skull and mandible. No documentation.
Rambaramp reconstruction.

These seven crania include four without name or label, but they can quite safely
be accepted as belonging to the same series; in morphology and in Rambaramp
reconstruction they are very similar and characteristic of South Malekula.

These Malekulan crania can be fairly well substantiated historically. The Macleay
diary entry of 15/3/1879 records receipt and subsequent purchase from Archibald Boyd,
of human osteological specimens from Mallicolo; on 25/3/1879 Masters exhibited “two
heads of Malicolo Chiefs’. Macleay in his diary of 26/3/1879 records “two skulls of
Chiefs of Island of Mallicolo by Mr. Masters from my Museum’. Harper (1901)
exhibited Malekulan crania; and finally there is the evidence of the serial numbers on
two heads and five bare skulls in sequence from Nos. 1661-1669.

Harrisson refers to the “interesting long-headed folk farther south’ who adorn
the skulls of the dead with bright coloured clay and “good spider’ web hair”. He
refers one to Deacon for a fuller- account (Harrisson, 1937, pp. 417, 423).

Flower, as early as 1881, was fully aware of the functional intent of the recon-
struction when he used the term “Monumental heads”, i.e. memorials in the modelled
likeness of the dead (Flower, 1881, pp. 75-81).

188 CRANIA IN THE MACLEAY MUSEUM,

Deacon gives such a complete description of the manufacture and the ingredients in
the reconstruction that there is no point in repeating it (Deacon, 1934, p. 544). Both
he and Harrisson point out that the face is modelled to resemble the dead as closely
as possible, that degree of care in decoration depends on the former rank of the
owner, and that interest in the reconstructed heads persists for no more than a year,
when they are thrown into the clan ossuary.

The technique of execution in South Malekulan reconstructions is standard and is
rather different from reconstructions in any other region. In colour and amount of
decoration, however, variation does occur. The heads in the Macleay Museum are a
plain black, in the Anatomy Department there are similar elongated heads, with the
Malekulan reconstruction, but the colour is pink. Reconstructions appear to be
confined to male skulls; the women had their own ceremonials, but the Rambaramp is
not part of them.

It may be suggested, therefore, that Masters and Macleay, and possibly Archibald
Boyd also, were not necessarily accurate in referring to crania Nos. 124 and 126 (i.e.
Sie XIII, Nos. 1661-1662) as heads of chiefs. Cranium No. 123 is far more complete
and elaborate in its reconstruction than these two and might possibly be the young
son (perhaps aged 15) of a chief.

Cranial artificial deformation is widely mentioned in the literature, but intensive
work on the subject appears to be less than in many other cranial fields. Among fairly
recent work, Cilento, 1921, described seven deformed crania from New Britain. Hambly
has dealt fully with the craniometry of Ambrym Island, 20 miles east of Malekula,
and proposed to use these undeformed Ambrym crania as a control for comparison with
55 Malekulan deformed crania collected by A. B. Lewis in 1909-13 (Hambly, 1946,
pp. 1, 2). So far as the writer knows, the study has not yet eventuated.

Of approximately 35 artificially deformed crania in the Anatomy Department,
University of Sydney, 13 appear to be definitely from Malekula. There are others in
the Australian Museum, Sydney.

The writer hopes to combine these with the 26 Macleay specimens -and carry out
an intensive study of the Malekulan cranium. A minor start has already been made
at the suggestion of Professor A. N. Burkitt.

Geography.—Malekula was formerly spelt Mallicolo, sometimes Mallicollo; in the
report of Masters’ exhibit it appears as Malicolo. The old navigators give a variety
of spellings—Mannicana, Manicolla, ete. It is the second largest island of the New
Hebrides group, lying south of Espiritu Santo and west of Pentecost, Ambrym and
Epi. The Maskelyne Archipelago is off the east coast. The Banks and Torres
subgroups are at the northern end of the New Hebridean chain.

MUMMIFICATION.

A complete male obtained (16/8/1875) from a village at Treacherous Bay, Darnley
Island, by Mr. Williams, first mate of the Chevert. It is extended on, and lashed to,
a frame of wooden poles and displayed in a glass table case. Photographs of the
specimen have been sent to other museums from time to time. Anatomists including
Sir Grafton Elliot Smith have referred to it.

A complete Peruvian mummy. The technique accords with the classical descriptions
of many writers on Peruvian evisceration and preservation. The body is more or less
rigid in a position of foetal flexion.

MANDIBLES.

There are 13 spare mandibles. They have been tried against all the crania but
do not fit any of them. One has Fiji Macleay Museum in ink on the right ramus, but
does not fit any of the Fijian crania. One has New Zealand in ink on its inferior
border, but does not fit any of the three New Zealand crania. The others have no
labels. The majority are mildewed and non-European in appearance.

BY N. W. G. MACINTOSH. 189

MISCELLANEOUS.

Three complete, skeletons in perished canvas bags; they are in a good state of
preservation and covered with sandy soil, suggesting exhumation. No documentation.
Possibly Darnley Islanders obtained by the Chevert Expedition (1875).

Finally, there are two drawers filled with Aboriginal skeletal fragments, mostly
from coastal New South Wales.

ACKNOWLEDGMENTS.

Professor A. N. Burkitt for translation of Dutch literature, Dr. A. Capell, Lt. Cmdr.
L. Hinchliffe, V.R.D., and Mr. F. Greenop for translations of Russian script, and Miss
S. Orlay for translation of German literature. Personal friends who have lent or given
me Russian and German textbooks. Mr. Rainbow and Miss Dixon of the Australian
Museum, Miss Hunter of the Anatomy Department, University of Sydney, Messrs. Steel
and Burrows of the Fisher Library for suggesting, obtaining and lending relevant
literature. Professor A. N. Burkitt and Professor A. A. Abbie for inspecting -and
commenting on the crania after they had been arranged, and offering further suggestions,
all of which were used. Dr. Murray of the School of Tropical Medicine, and Miss B. Orr
and Mr. Mackenzie of the Geography Department, University of Sydney, for search for
place names. Messrs. Woodhill and Lee, who suggested and made the subject available.
Mr. F. A. Greenop, who gave much time and asistance, both in the Macleay Museum
and in his own office, to the writing of this paper. Mr. J. Henry, Curator of the
Macleay Museum, who cheerfully joined with the writer in moving about one-sixth
of the contents of the Macleay Museum from one place to another. Miss A. Scot
Skirving who typed and re-typed the entire paper. To these my complete
acknowledgment and thanks are due.

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CYCLOSTYLE NOTICE TO STUDENTS.—Undated.

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Mactay, N. von MIKLUCHO, 1875.—Ethnologische Bemerkungen uber die Papuas der Maclay-
Kiiste in Neu-Guinea. Natuurk. Tijdschr., 35: 66-93.

Macray, N. vON MIKLUCHO, 1876.—Meine zweite Excursion nach Neu-Guinea (1874). Natuurk.
Tijdschr., 36:148-179.

Mactay, N. von MIKLUCHO, 1876.—Ethnologische Bemerkungen Uber die Papuas de Maclay-
Kiiste in Neu-Guinea. Natuwurk. Tijdschr., 36: 294-3338.

Maciay, N. voN MIKLUCHO, 1878.—Anthropologische Notizen, gesammelt auf einer Reise in
West-Mikronesien und Nord-Melanesian im Jahre 1876. Z%. Hthn., 10: 99-118.

Mactay, N. DE MIKLOUHO, 1881.-—A Short Resume of the Results of Anthropological and
Anatomical Researches in Melanesia and Australia Mar. 1879-Jan. 1881. Proc. LINN. Soc.
N.S.W., 6: 171-175.

Macuay, N. DE MIKLOUHO, 1881.—Exhibits, Proc. LINN. Soc. N.S.W.; 6:196.

Mactay, N. DE MIKLOUHO, 1881.—Cranial Deformation of New-Born Children of the Island
Mabiak, and other Islands of Torres Straits, and of the Women of the S.E. Peninsula of
New Guinea. Proc. LINN. Soc. N.S.W., 6: 627-630.

Mactay, N. pe MIkKLOUHO, 1884a.—Remarks on a Skull of an Australian Aboriginal from the
Lachlan District. Proc. LINN. Soc., 8: 395-396.

Macuay, N. pp MIKLOUHO, 1884b.—On a Very Dolichocephalic Skull of an Australian Aboriginal.
Proc. LINN. Soc. N.S.W., 8: 401-4038. i

Macuay, N. DE MIKLOUHO, 1886.—Plants in Use by the Natives of the Maclay Coast of New
Guinea. Proc. LINN. Soc. N.S.W., 10: 353-354.

MACLEAY, W., 1875.—Notes on the Zoologicai Collections made in Torres Straits and New
Guinea during the Cruise of the Chevert. Proc. LINN. Soc. N.S.W., 1: 36.

MAcLEAY, W.—Diaries 1873-1881 (Volume 1878 missing). Stored in the Library of Linn. Soc.
N.S. W.

Macueay, W., 1875.—Journal (Diary) of the Chevert Expedition. Stored in the Library of the
Linn. Soc. N.S.W.

MARTIN, R., 1928.—Lehrbuch der Anthropologie. Zweite Auflage. Zweite Band. Jean.

MASTERS, G., 1877.—Exhibition of, and Remarks upon, the Skull of an Aboriginal Female.
Proc. LINN. Soc. N.S.W., 2.

MASTERS, G., 1879.—Exhibits. Proc. LInn. Soc. N.S.W., 4: 244, 25th March.

MIKLUHO-MAKLAY, NIKOLAI NICOLAEVITCH, 1846-1888.—The Australian Encyclopaedia, 2:, 1926,
82.

Munpy, R., 1848.—Narrative of Events in Borneo and Celebes from the Journals of James
Brooke, Esq., Vol. I, London, 96.

MusGRAVE, A., 1932.—Bibliography of Australian Entomology, 1775-1930, 101.

NEW GUINEA PLACE NAMES (claimed as Maclay’s handwriting).—Single sheet of blue foolscap,
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PAPERS AND CORRESPONDENCE.—Unbound and scattered in the Macleay Museum.

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Rocers, S. L., 1937-38.—The Healing of Trephine Wounds in Skulls from Pre-Columbian Peru.
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BY N. W. G. MACINTOSH. 191

THOMAS, O., 1885.—Account of a Collection of Human Skulls from Torres Straits. J. R. anthrop.
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TINDALE, N. B., and BIRDSELL, J. B., 1941.—Tasmanoid Tribes in North Queensland. Results
of the Harvard Adelaide Universities Anthropological Expedition 1938-39. Rec. S. Aust.
WH Roe Up (QD), Be

ToPINARD, P., 1872.—Etude sur les races indigénes de l’Australie. Bull. Soc. Anthrop., Tome 7,
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Visitors’ Book.—Macleay Museum, 1890.

Woop JONES, F., 1934.—Contrasting Types of Australian Skulls. J. Anat., 68: 323-330.

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in the Commonwealth of Australia. Biometrika, 30 (iii-iv), 305-340.

192

THE HAIR TRACTS IN MARSUPIALS.
Part III. DESCRIPTION OF SPECIES, CONCLUDED.

By W. BOARDMAN.
(Department of Zoology, University of Melbourne.)

(Two Text-figures.)

[Read 31st August, 1949.]

INTRODUCTION.

The marsupial pouch young, all from Western Australia, which form the subject
of this communication are part of the collection of the Western Australian Museum,
Perth. I am very appreciative of the kindness of Mr. L. Glauert, Curator of the Museum,
who made available to me both the specimens and his knowledge of marsupials.

Suborder POLYPROTODONTIA.
Family DASYURIDAE.
Subfamily PHASCOGALINAE.
PHASCOGALE, Sp.*

Material.—M.2407-9—three males (length of head and body about 37 mm.), young of
M.2406; Woolleen Station, Murchison District; coll. B. H. Sharpe, 28th March, 1940.

Hair tract pattern as recorded for Dasycercus cristicauda (Boardman, 1946a) is
repeated in this species. The forearm reversal enters into the formation of a convergent
point placed postaxially about a third or more of the length of the forearm from the
elbow. <A triangular area in front of the scrotal stalk similar to that which, in
Dasycercus and Antechinus is the site of a reversal, is sparsely provided with hairs of
indeterminate inclination. However, the general arrangement of the tracts in the
vicinity of the scrotum points to the development of a typical prescrotal reversal
in older specimens.

SMINTHOPSIS CRASSICAUDATA CRASSICAUDATA Gould.

Material—Seven males and five females, probably constituting two litters (crown-rump
length about 17 mm.); Western Australia. ;
As with the subspecies macrura (Boardman,. 19430) and centralis (Boardman,

1946a), the hair tracts are similar in pattern to those of Antechinus maculatus.

Family MyrRMECOBIIDAE.
MYRMECOBIUS FASCIATUS Waterhouse. Text-figs. 1 and 2.

Material.—M.2495-8—a litter consisting of two males and two females (crown-rump length
about 35 mm.); Pardelup Prison Farm; pres. J. A. Thompson, 28th July, 1941. M.2587-90—a
litter consisting of a single male and three females (crown-rump length about 42 mm.) ;
Narrogin; pres. J. B. Higham, 19th July, 1943. M.1161-4—a litter consisting of a _ single
male and three females (length of head and body 71 mm.); The Williams, Williams River;
coll. 18th July, 1929. M.1868—-a male (crown-rump length 53 mm.); South Dale; pres. Miss
Eva Wills, 30th August, 1934. M.2432-5—a litter consisting of two males and two females
(crown-rump length about 42 mm.); Wagin; pres. H. S. Goldsmith, 22nd June, 1930.
M.2226-9—a litter consisting of three males and one female (crown-rump length 29 mm.) ;
The Williams, Williams River; coll. 21st May, 1937.

Wood Jones (1923) has recorded the general arrangement of the hair tracts in
Myrmecobius. Examination of this very fine additional material confirms the original
account and makes possible its extension particularly with reference to the side of the
head and the ventral surface.

* Mr. Glauert informs me that the species is new.

BY W. BOARDMAN. 193

Head and Neck.—Dorsally the flow is caudalwards. On the side of the head the
general direction is also caudalwards, except between the eye and the ear where a
preauricular reversal occurs, resulting in the formation of a divergent interval
(obscured by the stronger growth of the caudally flowing hairs in older specimens).
The preauricular reversal originates in a divergent centre on the tragus. A backwardly
directed feathering reminiscent of that described in the monotreme Tachyglossus
aculeatus (Boardman, 19460) is present in front of the medial canthus; its axis

:
SN
Ah
\
y)

2)

Sc Se oe

NS

lll
HW

Text-figures 1 and 2.
Figs. 1 and 2.—Myrmecobius fasciatus. 1. Lateral view of head, neck and forelimb. The
broken line indicates the caudal margin of the mystacial zone and the dotted line the orbital
ring. 2. Ventral aspect. The circle in front of the root of the tail marks the position

of the cloacal hillock.

makes an obtuse angle with the palpebral fissure. The origin of the feathering is in
what appears to be a divergent centre imperfect in that hairs do not stream forward
from it and situated on the caudal margin of the mystacial zone. Most of the
specimens have a similar but less definite parting in the mid-dorsal line immediately
behind the margin of the rhinarium. Between the mandibles the hair flows cranio-
caudally, but from a level somewhat in front of the angles of the mouth divergence
from the middle line initiates a caudodorsal current on the ventral aspect of the head
and neck. Laterally on the head beneath the eye and the ear and laterally or dorso-
laterally on the neck this diverging current merges insensibly with the general caudal-
wards. stream; further back it flows over the shoulder and upper arm.

Q

194 THE HAIR TRACTS IN MARSUPIALS. PART III,

Trunk.—The tendency for the ventral hair current on the head and neck to diverge
from the middle line is continued on the trunk, so that laterally between the attachment
of the fore- and hind-limbs it follows a curved course dorsally and caudally to merge
with the caudalwards stream on the dorsum. In the female inguinal region the
primitive caudoventral arrangement is not interrupted by the presence of the pouch.
In the male a prescrotal triangle of reversed hairs is found as in the Dasyuridae; the
hair field delimited by the triangle contains hairs having a craniomedial direction.

Limbs.—The forearm has a well-defined reversal postaxially with the convergent
point (occasionally not well developed) at the elbow or placed more proximally on
the arm. The hind-limb shows the undisturbed primitive condition.

Family PERAMELIDAE.
PERAMELES Sp.
Material.—10457-9—three females, probably litter mates (length of head and body
142 mm.); Napier Broome Bay; coll. G. F. Hill, 26th Feb., 1910.
This undetermined species of Perameles is similar to P. nasuta (Boardman, 1943a
and 19430). All three specimens have the gular whorled system single—in two on the
left side and clockwise, in the third on the right side and counter-clockwise.

A forearm reversal of the usual type, that is, postaxial and with the convergent
point distal of the elbow, is present in only two of the three specimens.

Suborder DriproropontTra.
Family PHALANGERIDAE.
Subfamily TARSIPEDINAE.
TARSIPES SPENSERAE Gray.
Material.—M.1888—a female (crown-rump length about 26 mm.); Youngs; pres. M.
Kittner, 26th November, 1934.
A single well-preserved specimen that confirms the previous account (Boardman,
1943b). The submental hairs flow cranially and medially.

Subfamily PHALANGERINAE.
CERCARTETUS CONCINNUS Gould.
Material. M.2638—a male (crown-rump length 24 mm.); Katanning; pres. W. Folland,
7th September, 1944.
No significant differences could be detected from Cercartetus nanus (Boardman, *
1946a).

TRICHOSURUS VULPECULA HYPOLEUCUS Wagner.

Material.—M.224—a female (length of head and body 180 mm.) ; Margaret River District;
coll. L. Glauert, 20th April, 1915. M.1422—a female (length of head and body 197 mm.) ;
Claremont; pres. J. EH. Payne, 11th August, 1931.

A previous account (Boardman, 1946a) of this subspecies based on a single female
at an early stage of hair growth is confirmed. The divergent system on the dorsal
aspect of the head is whorled counter-clockwise. The condition of the tracts in the
female inguinal region is, from an examination of M.224, similar to what has been
figured for the male of the subspecies vulpecula (Boardman, 1946a, Fig. 20) except
that the mid-ventral tuft is not whorled and there is no equivalent of the convergence
on the scrotal stalk; the convergent interval associated with the mid-ventral tuft lies
in the middle line on the margin of the pouch fold. Specimen M.1422, in which the
hair is much longer, at first sight appears not to have a mid-ventral tuft. There is,
however, a convergent interval on the lip of the pouch in the middle line as in M.224
but the cranially flowing current from it is almost immediately taken up in a pair
of centripetal whorls (clockwise on the left, of indeterminate direction on the right)
situated laterally on the lip of the pouch. It would appear probable that the mid-ventral
tuft is in this specimen displaced backwards into the pouch zone, resulting in a
secondary splitting of the whorl as the pouch cavity and lips developed.

BY W. BOARDMAN. 195

Family MAcCROPODIDAE.
Subfamily PorToRoIrNak.
BETTONGIA LESUEUR GRAII Gould.

Material.—10643—a male (length of head and body 175 mm.); Bernier Island, Shark
Bay; pres. O. H. Lipfert.

Head and Neck.—The head and neck show in the arrangement of the tracts a
pattern similar to that described for Potorous tridactylus apicalis (Boardman, 1946a).
A small divergent interval, occasioned by the presence of a divergent centre on the
margin of the upper eyelid just lateral of the medial canthus, occurs immediately in
front of the eye. On the left side a supernumerary centripetal whorl, clockwise, lies
on the oblique ridge formed by the meeting of the backwardly directed facial stream
and the forwardly directed stream from the lateral neck whorl.

Trunk.—Dorsally and laterally the thorax has the hairs primitively disposed; the
arrangement of the tracts ventrally is conditioned by the presence of an axillary
divergent interval as in Bettongia penicillata. Over the flank the primitive caudoventral
trend is maintained. The inguinal region is, with minor variations, as described and
_ figured for Bettongia penicillata (Boardman, 1946a, Fig. 33).

Limbs.—A convergent point associated with a reversed field is present on the
postaxial margin of the forearm somewhat distal of its middle.

Subfamily MaAcROPODINAE.
SETONIX BRACHYURUS Quoy and Gaimard.

Material.—M.310—a male (length of head and body 240 mm.); Western Australia.

Setonix brachyurus differs in only minor details from the account given of
Wallabia agilis (Boardman, 1946a@). There is no small whorl in the vicinity of the
lateral canthus. The convergent centre on the crown shows clockwise whorling. The
nuchal whorled system is probably bilaterally doubled but the hairs are locally
disturbed so that the details of tract formation are obscured. The tracts in the groin
are somewhat simpler than in Wallabia bicolor (Boardman, 19436, Fig. 25) in that the
convergent interval between the scrotum and cloacal eminence is absent.

References.

BOARDMAN, W., 1943a.—On the External Characters of the Pouch Young of Some Australian
Marsupials. Aust. Zool., 10:138-160.

———. 1943b.—The Hair Tracts in Marsupials. actiogle Description of Species. PROC.
LINN. Soc. N.S.W., Ixviii: 95-113.

————,, 1946a.—The Hair Tracts in Marsupials. Part II. Description of Species, continued.
ibid, Ixx: 179-202.

, 1946b.—The Hair Tracts in Monotremes; their Relationship to the Mammalian

Primitive Condition. Univ. Qld. Pap., Dept. Biol., II, 7:7 pp.

JONES, F. Woop, 1923.—The External Characters of Pouch Embryos of Marsupials. INOS i
Myrmecobius fasciatus. Trans. Roy. Soc. S. Aust., 47: 195-200.

196

STUDIES ON AUSTRALIAN MARINE ALGAE. V.

OBSERVATIONS ON AND GEOGRAPHICAL RECORDS OF VARIOUS SPECIES, PARTICULARLY
THOSE OF THE GELIDIUM COMPLEX.

By VALERIE May, M.Sc.
(C.S.I.R.O. Marine Biological Laboratory, Cronulla, N.S.W.*)

[Read 25th May, 1949.]

INTRODUCTION.

This paper is concerned mainly with species of the genus Gelidiwm, various
observations and new records of distribution being reported. In addition, extended
geographical ranges are recorded for certain other algae and the occurrence of tetrasporic
material is noted in a species normally found only in a sterile condition.

Brief mention of certain of these species was made in Notes. and Exhibits on an
Excursion to the Great Barrier Reef (Anon., 1948).

Unless stated otherwise, the specimens discussed are located either in my own
herbarium (quoted as VM), in the National Herbarium of New South Wales, Sydney
(quoted as NSW), in the Herbarium of A. H. S. Lueas, now held at the National
Herbarium of New South Wales (quoted as AHSL) or in the Herbarium of the
University of Sydney, New South Wales (quoted as SYD).

GELIDIUM CAULACANTHEUM J. Ag.
New Record for Australia.

For establishing the specific identity of this species the writer has examined New
Zealand material of G. caulacantheum, e.g., Lindauer’s Alg. N.Z. Exsicc. No. 63 (Herb.
VM) and also a series of specimens kindly sent me by Miss L. Moore of D.S. & LR.,
Botany Department, Wellington, N.Z.

G. caulacantheum approaches G. pusillum (discussed elsewhere in this paper), both
species being relatively small tufted plants, dark in colour, often with terete branches
pinnately branched. However, G. caulacantheum is a larger, more lax plant, with
branching less imbricate, and more regularly pinnate, the branching frequently being
relatively less developed at the base of the frond. G. caulacantheum is nearly terete
and shows less tendency to flattening than does G. pusillum. So far in Australia
G. caulacantheum has been found only in harbour or estuarine waters, whereas
G. pusillum trequents headlands exposed to the ocean and occurs less frequently in
sheltered waters; however, both species have been found growing associated (at
Bradley’s Head, Port Jackson, N.S.W., 20/1/1946; specimens both quoted this paper).

Localities from which G. caulacantheum has been collected in Australia are as
follows:

Locality. Date. Herbarium. Notes.
Port Jackson, N.S.W.— :
Bradley’s Head. 20/ 1/1946 V.M. No. 1196. Both tetrasporic and cystocarpic
plants.
No. 2002-4.
No. 2006.
0 3 23/ 6/1947 V.M. No. 2343-4. Cystocarpic plants.
No. 2345-8.
Mrs. Macquarie’s Chair, Botanic
Gardens. 28/11/1947 V.M. No. 2474-5.
Rose Bay. 28/11/1947 V.M. No. 2476.
Woolloomooloo Bay. —/11/1931 A.H.S.L. Both tetrasporic and cystocarpic
plants.
Clifton Gardens. -/ 1/1914 A.H.S.L.
Farm Cove. —/ 4/1930 A.H.S.L.

* Contribution No. 79 from the laboratory.

BY VALERIE MAY. 197

OBSERVATIONS ON GELIDIUM AUSTRALE J. Ag.

For authentic material of this species I have examined Harvey’s Alg. Aus. Exsicc.
No. 333H Gelidium asperum (specimen in Herb. NSW) from Western Point, Victoria,
which A. and E. S. Gepp (1906) record as being true G. australe J. Ag.

This and other specimens of the species show rhizoidal filaments abundant,
originating first in the cortical region of the thallus. These observations do not accord
with Okamura’s report (1934, p. 55) that an Australian specimen of G. australe was
“void of rhizoidal filaments”.

G. australe has been recorded in Australia as G. asperum by Harvey, among others
(as quoted above). I have not seen authentic material of G. asperum from Australia,
nor apparently had Lucas (1931). Dr. Tore Levring (private correspondence) has
agreed that the G. asperwm recorded by him (1946) from Victoria, Australia, is in
fact G. australe. He says that the limits between these two species “do not seem to
be very sharp”’.

G. australe is known from the southern coasts of Australia, from Tasmania and
from New South Wales; I have also found it as drift in southern Queensland. In
habit the species resembles a fine form of Pterocladia capillacea. Lucas (1914)
surprisingly recorded the species as “characteristic of the ocean surf-flora”, but in
1935 he recorded it from Botany Bay and Port Jackson, i.e., estuarine waters of
New South Wales. The latter record is in accord with the present writer’s experience
that G. australe frequents sheltered waters of harbours and estuaries such as Botany
Bay, Port Jackson and Pittwater in New South Wales.

GELIDIUM CRINALE J. Ag.
Species excluded from Australian Records.

This species was recorded from Queensland, Australia, by A. D. Cotton (1915). In
answer to a request for Queensland material of this species, Miss C. I. Dickenson, of
the Herbarium, Royal Botanic Gardens, Kew, kindly sent me a specimen labelled
G. crinale, and collected by A. J. Banfield, in February, 1910, at Dunk Island, off
Queensland. This is likely to be a portion of the material on which Cotton based his
record. This specimen is not G. crinale; it lacks both the rhizoidal filaments and
apical cell which are characteristic of the genus Gelidium. The specimen quoted is,
in fact, Gelidiopsis intricata (Ag.) Vickers, a species discussed elsewhere in the present
paper. The writer therefore rejects the record of Gelidiwm crinale from Australia.

NoTE ON THE ANATOMY OF GELIDIUM LUuUCASTI May.

In 1944 I described and figured portions of the thallus of this species and
inadvertently used the term “filament” in a different sense from that usually used in
this genus. The structures I figured occur very much nearer to the apices than do
the true rhizoidal filaments (sensu Okamura, 1934). Sections taken further from the
apex of G. Lucasii show that the true rhizoidal filaments originate in the peripheral
layers outside a zone of large cortical cells. Lower down the axis rhizoidal filaments
occur in the medulla also. Thus G. Lucasii follows Okamura’s rule that generally a
cross section of the thallus of a species of Gelidiuwm shows rhizoidal filaments very
evident towards the periphery of the thallus.

GELIDIUM PUSILLUM (Stackh.) Le Jol.
Variation and Distribution.

Bailey (1912) recorded this species from Queensland on the authority of
identification made from Kew, presumably by Mr. A. D. Cotton; since then the species
has been quoted as known from Australia, or North Australia, without more detail
as to location. Lucas (1934) referred to a species “recalling in habit G. pusillum
(Stackh.) Le Jol.” from Low Island, Qsld., but generally there appears to be extremely
little information in the literature about G. pusillum and where and in what forms it
occurs in Australia.

198 STUDIES ON MARINE ALGAE. V,

G. pusillum is a persistent element in our marine flora and is now recorded from a °
number of localities in N.S.W., S.A., W.A., Tas., and also from Lord Howe Island.*

The identification of this species is based primarily on the description and illustra-
tion of Okamura (1934) and also on comparison with Lindauer’s Alg. N.Z. Exsiccatae
No. 238 (Herb. VM).

Of the species of Gelidiwm previously recorded from Australia, G. pusillum is fairly
readily recognized by its small size, felting base and compact habit. The distinctions
between G. pusillum and G. caulacantheum are discussed in this paper under the latter
species.

The Australian specimens of G. pusillum vary greatly, but withal the series shows
continuous gradations with frequent transition stages between forms, so that the whole
series must be regarded as a single biological unit or species. Given only examples of
the extreme forms, one would justifiably consider them as distinct species, but as an
examination of a larger range of specimens shows there is no discontinuity, the
variation must be regarded as of sub-specific significance only.

At one extreme of this range the specimens are fine and terete, while branching
is distant, irregularly alternate or secund. This form is usually found in still waters
associated with mud, but may occur in other locations. This is f. pseudointricata,
discussed below.

With more robust specimens branching is more frequent and more generally
pinnate, and branches are more generally flattened. This is the form usually found
on ocean headlands and is that usually accepted as alone representing G. pusillum
(includes var. conchicola Pice. and Grun.).

At the other end of the range of G. pusillum specimens become very coarse and
much flattened, and the branches may become foliose (includes f. foliacewm Okam.).

The fine terete form of G. pusillum approaches in habit very near to Geliodiopsis
intricata (Ag.) Vickers (= Gelidium intricatum Kuetz.), from which it differs
anatomically (Gelidiopsis having no apical cell or rhizoidal filaments). The description
and illustrations (Levring, 1941) of Gelidium pseudointricata Skottsb. and Levr. suggest
that this species represents merely the fine form of G. pusillum. Examination of
Lindauer’s specimen of G. pseudointricata (Alg. N.Z. Exsice. No. 263, Herb. NSW)
confirms this opinion, and G. pseudointricata Skottsb. and Levr. is consequently now
reduced in status from species to form, becoming G. pusillum f. pseudointricata
(Skottsb. and Levr.), stat. nov.

Gelidium crinale (Turn.) Lamx. is reported as also very near in habit to
Gelidiopsis intricata; it appears to be closely allied to G. pusillum f. pseudointricata
and may actually be identical. I have insufficient material available for determining
this point.

In the following list of Australian collections of G. pusillum the specimens are
arranged according to their form, ranging from finest to coarsest, but it must be
emphasized that there are no distinct breaks between these various forms; rather
the species consists of a series of intergrading forms.

Locality. Date. Herbarium. Notes.
f. pseudointricata—
Sans Souci, Botany Bay, N.S.W. 11/ 3/1945 V.M. No. 555. Tetrasporic.
Tuggerah, N.S.W. 10/ 2/1943 V.M. No. 699. Cystocarpic. On whelks.
Queen’s Lake, Laurieton, N.S.W. 30/ 4/1944 V.M. No. 1146. Cystocarpic. On whelk.
Rose Bay, Port Jackson, N.S.W. 28/11/1947 V.M. No. 2478. Shows range of form.
Lord Howe Island. —/ 5/1982 A.H.S.L. Tetrasporic. Coll. F. A. McNeill.
Pennington Bay, Kangaroo Is., S.A. 16/ 1/1945 V.M. No. 909. Tetrasporic.
Gladesville Bay, Parramatta, N.S.W. —/11/1930 <A.H.S.L. Tetrasporic. Coll. A. H. 8. Lueas.
Narrabeen Head, nr. Sydney, N.S.W. 6/ 5/1945 V.M. No. 744. Tetrasporic.
Fairy Bower, Manly, nr. Sydney, .
N.S.W. 28/ 4/1945 V.M. No. 704.

* Specimens from Kangaroo Island, South Australia, quoted in this paper, were sent to the
writer for identification by H. B. S. Womersley, who has since recorded the occurrence of this
species in his ecological studies (Trans, Roy. Soc. 8S, Aws., 71 (2); 1947, 228-252, Pls. 9-13).

Locality.

Whale Beach, nr. Sydney, N.S.W.

Kangaroo Island, $.A.

Encounter Bay, S.A.

Lane Cove River, Port Jackson,
N.S.W.

Barrenjoey Head, Palm Beach, nr.
Sydney, N.S.W.

Cottesloe, Swan River, W.A.

” ” ” ”

var. conchicola—
Sandy Bay, R. Derwent, Tas.
Snug, Tas.
Long Reef, nr. Sydney, N.S.W.

Balmoral, Port Jackson, N.S.W.

Narrabeen Headland, nr. Sydney,
N.S.W.

Kurnell, Botany Bay, N.S.W.

Newport, nr. Sydney, N.S.W.

Collaroy, nr. Sydney, N.S.W.

Bradley’s Head, Port Jackson, N.S.W.

Forty Baskets, nr. Manly, Port
Jackson, N.S.W.
Careel Head, Whale Beach, nr.

Sydney, N.S.W.

Newport Headland, nr. Sydney, N.S.W.

Sans Souci, Botany Bay, N.S.W.

Fairy Bower, Manly, nr. Sydney,
N.S.W. -

Bondi, N.S.W.

Little Coogee, N.S.W.

Forms approaching f. foliacewm—
Port Hacking, Port Macquarie, N.S.W.
Potato Point, nr. Tuross Lake, Sth.
Coast, N.S.W.
Green Island, nr. Entrance of Conjola
Lake, Sth. Coast, N.S.W.
Stanwell Park, Sth. Coast, N.S.W.
Kiama, Sth. Coast, N.S.W.
Bundeena, Port Hacking, N.S.W.
Narrabeen Head, nr. Sydney, N.S.W.
Fairy Bower, Manly, nr. Sydney,
N.S.W.
f. foliaceum—
North Head, Curl Curl, nr. Sydney,
N.S.W.

BY VALERIE MAY.

Date.
22/ 6/1947

13/ 5/1945

25/ 2/1945

—/10/1926
—/12/1926

—/ 1/1909
—/11/1925
3/ 7/1947

4] 9/1947
6/12/1947

16/11/1944
21/10/1944
11/11/1944
20/ 1/1946
17/11/1945

1/ 1/1946

—/ 4/1945
5/ 5/1946
31/ 8/1947

—/4/1914
-/ 2/1914

31/ 5/1937
26/10/1947

3/ 5/1946

26/ 3/1945
23/ 6/1945
24/ 6/1945
1/ 7/1946
28/ 4/1945

CAULERPA

Herbarium.
V.M. No. 2341.
V.M. No.
A.H.S.L.
V.M. No.

V.M. No.

A.H.S.L.
A.H.LS.L.

A.H.S.L.
A.H.S.L.
V.M. No. 2362.

V.M.
V.M.

No. 2411.
No. 2482.

V.M.
V.M.
V.M.
V.M.
V.M.

195.
Nos. 491, 492.
No. 489.
No. 2005.
No. 1056.
V.M. . 1186.
NOs O
. No. 207
. No. 240

IQ

and N.S.W.

V.M. No.

ANCEPS Harv.

New Record for Australia.

199

Notes.

Tetrasporic.

Shows variation.

Coll. J. B. Cleland.

Mangrove Swamp. On whelk.
Plants taller than usual for this
form.

Collection shows variation. Plants
taller than usual for this form.
Sheet B, tetrasporic.

Tetrasporic. Coll. 1 Banks.

Coll. 1 Banks.

Tetrasporic. Coll. L. Rodway.

Coll. A. H. S. Lucas.

This collection shows a good range
of form.

Tetrasporic.

be)
Tetrasporic.
Tetrasporic. Coll. A. H. $8. Lucas.
Tetrasporie.
Tetrasporic.
Tetrasporic. Plant tall.
Cystocarpic. On mussels.
Tetrasporic. Range of form within

this collection.

Tetrasporic.

Weber Van Bosse (1898) described this species clearly, and my specimens accord
with Harvey’s Friendly Island Algae No. 67 (Herb. NSW).
C. anceps varies greatly in size of plant and also in whether the edges of the

leaves are smooth or dentate.

Of Australian species of Caulerpa, this species approaches

nearest to C. parvifolia Haryv., from which it differs by being larger and having a very

much coarser stolon.

Although collected here some time ago, C. anceps has not previously been recorded

from Australia.

Locality.
Heron Island, Great Barrier Reef, Qsld.

Brown Bay, Cairns, Qsld.
Hayman Is., Great Barrier Reef, Qsld.

Archer Pt., 9 m. North of Cooktown,
Qsld,

Date.
—/ 1/1948
—/8/1935

—/6/1934
—/9/1935

Herbaruim.
V.M. and SYD.
A.H.S.L.

A.H.S.L.
A,H,S,L,

Notes.
Collected by Sydney University
Biological Society. Specimens

Nos. 35 and 35X.

Coll. H. Flecker. Stolon less coarse
than usual.

Coll. F. A. McNeill.

Coll. P. Kelly. Stolon less coarse
than usual,

200 STUDIES ON MARINE ALGAE. V,

SARCONEMA FILIFORME (Sond.) Kylin.
New Record for New South Wales and for Lord Howe Island.
8. filiforme is known from W. and N.EH. Australia and from Tasmania, so that it is
not surprising that it should occur also in New South Wales.

Locality. Date. Herbarium. Notes.
Double Bay, Port Jackson, N.S.W. 30/7/1943 V.M. No. 2604, 2605. Trawled.
Bradley’s Head, Port Jackson, N.S.W. —/4/1918 A.H.S.L. Coll. Hindmarsh.
Lord Howe Island. —/6/1933 A.H.S.L. Coll. F. Perrin and A. H. $8. Lucas

GELIDIOPSIS INTRICATA (Ag.) Vickers.
Definite Records from Australia.

Specimens of the genus Gelidiopsis frequently resemble closely specimens of the
genus Gelidium, but, following Feldmann (1931), the genera may be readily distinguished,
since Gelidiopsis lacks an apical cell and does not develop rhizoidal filaments.

Gelidiopsis intricata is a fine terete tufted plant with irregular branching; in habit
the species recalls strongly both Gelidium crinale (Turn.) Lamour, and Gelidium
pusillum f. pseudointricatum (Skottsb. and Levr.) May, as discussed elsewhere in this
paper.

Gelidiopsis intricata (as Gelidium intricatum Kuetz.) was recorded from “North
Australia”, without further locality, by Grunow (1874) and there appears to be no
more definite locality record of the species from this country, later writers merely
repeating the statement that the species had been recorded from Australia. It appears
to be of value, therefore, to list the Australian specimens of G. intricata which I have
examined, and the locations from which these plants were collected.

Locality. Date. Herbarium. Notes.

Low Is., N. Qsld. —/6/1931 A.H.S.L. Filed as Gelidium sp.

Green Is., off Cairns, N. Qsld. —/6/1931 A.H.S.L. Filed as Gelidium sp.

Dunk Is., Qsld. —/2/1910 V.M. ex KEW. Specimen discussed in this paper
under “‘ Gelidium crinale’’.

Heron Is., Great Barrier Reef, Qsld. —/1/1948 V.M. Coll. by Sydney Univ. Biological
Society. Specimens No. Bl and
B4.

Mooloolaba, Qsld. 24/1/1944 V.M. No. 2602 Drift.

Yorkey’s Knob, Qsld. 5/7/1936 V.M. and N.Q. Nats.*

No. 2002.

SYMPHYOCLADIA MARCHANTIOIDES (Harv.) Falk.
New Record for New South Wales.

This species was described and illustrated by Falkenberg (1901) and by Okamura
(1912). It has been recorded previously as occurring in north-eastern Australia, but
unfortunately none of these specimens is available to me for study.

The present is the first record of S. marchantioides from New South Wales; all
specimens were encrusting and obtainable only at very low tide. My plants are
smaller than, but otherwise agree with, Lindauer’s Alg. N.Z. Exsicc. No. 200 (Herb.
VM).

Locality. Date. Herbarium. Notes.
Balmoral, Port Jackson, N.S.W. 4/9/1947 V.M. No. 2409, 2410. Tetrasporic.
Bungan Island, Bungan Head, nr. 27/1/1948 V.M. No. 2536, 2537. Under rock cave, —6 in. tide.
Sydney, N.S.W. Ecklonia zone.
Little Beach, Narrabeen, nr. Sydney, 26/1/1948 V.M. No. 2545. In shallow, very shaded pool.
N.S.W. Tetrasporic.
Little Beach. Narrabeen, nr. Sydney, 25/1/1948 V.M. No. 2553. Few tetraspores.
N.S.W.
Little Beach, Narrabeen, nr. Sydney, 23/3/1948 V.M. No. 2586. Tetrasporic.
N.S.W.
Little Beach, Narrabeen, nr. Sydney, 23/3/1948 V.M. No. 2587, 2588.
N.S.W.
Fairy Bower, Manly, Sydney, N.S.W. 24/3/1948 V.M. No. 2597. Few tetraspores.

* Herbarium of the North Queensland Naturalists’ Club, Cairns, Qsld.

BY VALERIE MAY. 201

LENOMANDIA CORONATA Lind. and Setch.
New Record for Australia.

Lindauer and Setchell (1946) described and illustrated this species from New
Zealand. It is now recorded as occurring in Australia also. The specimens of two of
the three collections recorded below are slightly smaller than the New Zealand plants
(i.e., Lindauer Alg. N.Z. Exsiccatae No. 74, Herb. NSW and VM).

The Australian specimens were all found growing on rocky ocean headlands and
could be collected only at extra low tide. The apices of the “leaves” vary greatly,
some being stoloniferous, others mucronate, while others have a depressed growing
point surrounded by dichotomous monosiphonous hairs.

Locality. Date. Herbarium. Notes.
Near Sydney, N.S.W.—
S.E. Narrabeen Headland. _ 1/ 7/1946 V.M. No. 2096. Far out rock pool L.T.L.

Little Beach, Narrabeen. 28/12/1947 V.M. No. 2515. Surf-sprayed rock, extra L.T.L.
: Specimens larger than 2096 or

2552.
a5 55 = 25/1/1948 V.M. No. 2552. Rock face, extra L.T.L. Tetra-

sporic.

ACROSORIUM UNCINATUM (J. Ag.) Kylin.
Occurrence of Tetrasporic Material.

A. uncinatum (J. Ag.) Kylin occurs widely distributed, and in Australia is known
from our southern and eastern coasts. Generally, however, this species is found in a
sterile condition. Kylin (1924), who associated the present binomial name, says he
has not seen fertile material of this species. Kuehne (1946) and Nott (1900) regard it
as highly likely that A. uncinatum reproduces itself largely by vegetative means without
spore formation. Lucas (1926, p. 602), speaking of this species in Australia, says:
“Though I have examined hundreds of specimens, I have never found cystocarps and
only once discovered sori, and then only on one segment of a fairly large plant.” He
does not give collecting details of this specimen, but in his herbarium the specimen,
which is quite large, is clearly marked. He collected it at Sandringham, Botany Bay,
N.S.W., in December, 1930.

The only other tetrasporic specimen, other than those quoted below, seen by the
writer, was that collected by Harvey (Alg. Aus. Exsice. No. 298, Herb. NSW), from
Port Fairy, Victoria. We do not know whether Harvey collected his specimen from
the ocean headland or from the estuarine lagoon at Port Fairy.

The present writer has collected A. uncinatum frequently. Most, and largest,
specimens have been trawled from estuarine and harbour waters, but none of these
has been fertile. Tetrasporic specimens have, however, been obtained several times
from ocean headlands, as listed below.

It is suggested, therefore, that ecological conditions affect the production of
tetraspores in A. uncinatum and that the conditions occurring in sheltered waters
favour this process less than do conditions on ocean headlands.

Specimens of A. uNcINATUM Bearing Tetrasporic Sori.

Locality. Date. Herbarium. Notes.
Headland opp. Coila Lake, nr. Tuross, 30/10/1947 V.M. No. 2493. Membranes wide, apices more
Sth. Coast, N.S.W. rounded than usual for the
species.
Headland between Newport and Bungan, 27/12/1947 V.M. No. 2499. Collected at extra L.T.L.
nr. Sydney, N.S.W.
Little Beach, Narrabeen, nr. Sydney, 25/1/1948 V.M. No. 2551. Below “‘browns”’. Extra L.T.L.
N.S.W.
Little Beach, Narrabeen, nr. Sydney, 26/1/1948 V.M. No. 2542. Extra L.T.L.
N.S.W.
Little Beach, Narrabeen, nr. Sydney, 26/1/1948 V.M. No. 2541. On Amphiroa.
N.S.W.

Tathra, N.S.W. 3/1/1943 V.M. No. 2532.

202 STUDIES ON MARINE ALGAE. V.

SUMMARY.

In this paper Caulerpa anceps Harv., Gelidium caulacantheum J. Ag. and Lenomandia
coronata Lind. and Setch. are recorded for the first time as collected from Australia,
the record of the occurrence here of Gelidium crinale is dismissed, while extended
geographic ranges within Australia are recorded for Gelidium pusillum (Stackh.) Le Jol.,
Sarconema filiforme (Sond.) Kylin, Gelidiopsis intricata (Ag.) Vickers, and
Symphyocladia marchantioides (Harv.) Falk. In addition, observations are recorded
on Gelidium australe J. Ag. and Gelidium Lucasii May, and the occurrence of tetra-
sporic material in Acrosorium uncinatum (J. Ag.) Kylin is recorded and discussed.

The new combination is made of Gelidiwm pusillum f. pseudointricata, based on
Gelidium pseudointricata Skottsb. and Levr. -

References.

ANONYMOUS, 1948.—Notes and Exhibits on an Excursion to the Great Barrier Reef. By
members of the Sydney University Biological Society. LINN. Soc. N.S.W., Ixxiii, xliv.
Batiuey, F. M., 1912.—Comprehensive Catalogue of Queensland Plants. Brisbane.
Corron, A. D., 1915.—Cryptogams from the Falkland Islands. J. Linn. Soc. Bot., 43 (290):
137-231, Pls. 4-10.
FALKENBERG, P., 1901.—Fauna und Flora des Golfes von Neapel. Rhodomelaceen. Monographie
Zoologischen Station zu Neapel. Berlin.
FELDMANN, JEAN, 1931.— Remarques sur les genres Gelidiwm Lamour., Gelidiopsis Schmitz. et
Echinocaulon (Kutz.) emend. Rec. Trav. Cryptog. dédiés a Louis Magin, 151-166.
Gepp, A., and GEPP, EH. S., 1906.—Some Marine Algae from New South Wales. J. Bot., 44:
249-261, Pl. 481.
Grunow, A., 1874.—Algen der Fidschi-, Tonga- und Samba-Inseln. Jowr. Mus. Godeffroy, 3
(6) : 23-50.
KUEHNE, P. E., 1946.—Four Marine Algae from Australia and New Zealand. Lloydia, 9 (1):
31-44, Pls. 1-7.
KYLIN, H., 1924.—Studien tiber die Delerseriaceen. Lunds Univ. Arsskr. Nien rey PAW) (C3)) 8 aeallilil.
Taf 1-80.
LEVRING, ToRB, 1941.—Die Meeresalgen der Juan Fernandez-Inseln. The Natural History of
Juan Fernandez and Master Island. Ed. Dr. Carl Skottsberg. Vol. 2.
, 1946.—A List of Marine Algae from Australia and Tasmania. Goteborgs Botaniska
Tradgard, 16: 215-227.
LINDAUER, V. W., and SETCHELL, W. A., 1946.—Note on a New Species of Red Alga, Lenomandia
coronata. Trans. Roy. Soc. N.Z., 76 (1): 66-7, Pl. 1. :
Lucas, A. H. S., 1914.—Marine Algae. Brit. Ass. Adv. Sci. Hdbk. N.S.W., 459-463.
, 1926.—Notes on Australian Marine Algae. III. The Australian Species of the Genus
Nitophyllum. PRoc. LINN. Soc. N.S.W., li (4): 594-607, Pls. 37-45.
————, 1931.—Notes on Australian Marine Algae. VI. Descriptions of Six New Species.
Proc. LINN. Soc. N.S.W., lvi (5): 407-411, Pls. 23-27.
, 1934.—Notes on Australian Marine Algae. VII. The Algae of the Low Islands. Proc.
LINN. Soc. N.S.W., lix (5-6) : 348-50.
, 1935.—The Marine Aigae of Lord Howe Island. Proc. LINN. Soc. N.S.W., Ix (3-4):
194-232, Pls. 5-9.

May, VALERIB, 1944.—Studies. on Australian Marine Algae. 1. The Corrected Name for
Pterocladia pectinata (A. and HE. S. Gepp) Lucas. Proc. Linn. Soc. N.S.W., Ixix (5-6):
226-228.

Nott, C. P., 1900.—Nitophylla of California. Proc. Calif. Acad. Sci., 3, Bot. 2:1-62, Pls. 1-9.

OxkAmouRA, K., 1912.—Icones of Japanese Algae. 2. Tokyo.

WEBER VAN BOssk, A., 1898.—Monographie des Caulerpes. Ann. Jard. Bot. de Bwitenzorg, 15
(2) : 243-401, Pls. 20-34.

PLATE III.

Proc. Linn. Soc. N.S.W., 1949.

Australian Rust Studies.

PLATE Iv.

Proc. Linn. Soc. N.S.W., 1949.

Australian Rust Studies.

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203

THE STRATIGRAPHY OF THE LOWER MARINE SERIES OF THE PERMIAN
SYSTEM IN THE HUNTER RIVER VALLEY, NEW SOUTH WALES.

By G. D. Ossporne, D.Se., Ph.D., Reader in Geology, University of Sydney.
(Two Text-figures.)

[Read 28th September, 1949.]

Contents.

Page

Introduction ss CR Ee Rn ea) es Ae hat Se ete hak tes cm Tey Uae acne Wy ja serait anemere OB
mae otrer Marine Series) in, ghee AEN a lok Wis ha ge PT ee tN EN We nA ee Del BABAR eG = en MP ere seesi An’) 3)
Geographic Distribution .. . IAT SOUS en EN ee PC ae MANES ae, nayreh tee cued av Agmyew on! fajicete vow ie (1)
Areas of Regional Dylon: Fai de Ee AG eee an cssa eee OO
Relations between the Lower Marine Series end ine renee series Le Eyer ag pees Watt tora OG
Palaeogeographical Setting .. oe Me tr At Rae hei 2 Mee i ies sete ee aay et 206
Stratigraphy of the Lower Marine Geries Be PERE aden aE My fa IN nonin d” RTA Li Ac hayes Sunray eee OL (4)
The Lochinvar Stage MO Si eee ted Lao Pepe ae hcrmul (tat tse OA et ed aehe AE Gomme ete Ort
nnerAl lan dales Stage mney cus fo ciewe fess) ec ne Sh oll, Be oo Daubert uccty Gal oF oo Zail)
The Rutherford Stage ee PID A Ne ame SP Seg Na em Ae a aN ee tm eto Ce ABD ca GA has Ses ee kay De
The Farley Stage Ha RIPE Se RTS aro oe Ros oe teste Heo LG

The Lower Marine Rocks of ihe Gramisp Conner Baein SNe) eh epee Siriaas ama eon? Ad LY
The Lower Marine Rocks of the Muswellbrook District ..° .. .. .. .. .. ©. .. 218
CornelationsMotmoCCllOnsimtdts ites ts eee tard tek Pei cu cieme iy ito Sopa LAE UN a es eee NS
Summary of the Stratigraphy eaiiyits ORDEAL dA arin ey ot | PL err eansrevt a O)
Possible Underground Extensions of the Tornoe MEwatas ‘Goss SHEEN Teach Mee eRe eal’ luteus ete gO.
TLFISREGS,. ONE: <TSOVSISIU IST ct | argon) alee 4 more meter )t atom Wt Opes Ua aut hs RAO Ta APT to oT RR ee Vee ah?
FVCLCRENICCS mae RE Tata, Pee es eis 4 Ie alps hota ceccmed lA Laat en ae wabssh mate” Sheen eterae TOMS SV as cane lee oer

INTRODUCTION.

During several field seasons between 1939 and 1946 the stratigraphy of the Lower
Marine Series in the Middle and Lower Hunter Valley has been examined and carefully
measured. This series has not received much detailed field investigation in the past,
although it has been the subject of much discussion and correlation by stratigraphers
and palaeontologists.

The chief aim of the writer has been to establish, as completely as possible in the
time available, the detailed succession in many areas, and to study the facies variation
from place to place.

Numerous sections have been measured, and some correlation of these has been
attempted. Much information concerning the structure of the Lower Marine Series has
accrued from the field-study, particularly concerning the area between Bishops Bridge
and Ravensfield, and also in the Pokolbin district, but the writer does not intend to
discuss tectonics beyond the mention of certain structural relations between the various
stages of the Lower Marine, where the consideration of these relationships assists in an
understanding of the stratigraphy. The tectonic evolution of the Lochinvar Dome, and
particularly the special character of the Pokolbin section of the Dome, are matters of no
small importance in any structural survey of the region, and these subjects will be
reserved for separate communication.

The field-work occupied about twelve weeks and during this aggregate period
approximately 500 carefully selected samples were obtained. Apart from a considerable
amount of work on foot, about 1500 miles of traverse were made along transport routes.

Some generai idea of the geographic distribution of the various stages is obtainable
from inspection of the accompanying map (Text-fig. 1).

Ss

204 STRATIGRAPHY OF THE LOWER MARINE SERIES,

DISTRIBUTION
of the
CRANKY LOWER MARINE
CORNER
SHIRES

ce ©se ewe

ROTHBURY -.

Farley Stage
Rutherford Stage
MMM «Allandale Stage
(4244) Lochinvar Stage

SiGAINE Ons caleroe nd Ho Vaat ROSES

Text-figure 1.

BY G. D. OSBORNE. 205

It must be stressed that not only was there insufficient time for the mapping of
the various horizons throughout the basin, but it was found that with the exception of
the Ravensfield sandstone, and the basalts, many of the strata varied in lithology along
the strike so as to make difficult any correlation for survey purposes.

In running the sections specimens were taken at very short stratigraphical intervals,
often from points five feet (stratigraphically) apart, and occasionally with even smaller
intervals. Of course in many places where the lithology was fairly constant for some
considerable thickness, the frequency of the specimens was less.

In the past the Lower Hunter region has received a great deal of attention from
geologists chiefly because of the extreme scientific interest of the rocks in the area,
and also because of the occurrence within the Permian System of the Greta and
Newcastle Coal Measures. The writer does not propose to review in detail the course
of geological investigation in the area as this has been done in other works. It will be
sufficient to remind the reader that the most important work in the past was the
establishment of the sequence and the elucidation of the structure of the Permian rocks
of the great Lochinvar structural province and associated areas, by the late Professor
Sir Edgeworth David in his survey of the Hunter River Coalfield, begun about 1884 and
continued at intervals until the publication of the Hunter River Memoir (1907) and
even in later years. Professor David made it clear in his writings that a detailed study
of the Lower Marine Series was not undertaken by him and his associates since the
chief task was the mapping and examination of the coal measures.

In 1913 A. B. Walkom made some detailed investigations of certain Lower Marine
sections in connection with a study of the stratigraphy of the Maitland—Branxton district.

A considerable amount of palaeontological research extending over a long period of
time has been done upon both megascopic and microscopic fossils from the area (see
reference list). d

In dealing with the Carboniferous—Permian problem, H. G. Raggatt and H. O.
Fletcher (1937) reviewed the Permian stratigraphy in the Hunter River district and
suggested the term Allandale Stage for about 800 feet of strata formerly grouped: in the
upper part of the Lochinvar Stage. The stratigraphical thicknesses given in their Lower
Marine Section (1937, p. 153) must now be superseded by the data available from the
writer’s detailed investigation.

The writer does not think it necessary in the stratigraphical section of this paper to
enumerate all the fossils recorded from the respective areas under discussion. It will
be apparent from the following pages that the correlations of various horizons in the
Hunter Basin can be effected by means of a few (often one or two) prominent species
amongst the faunas, and thus complete lists of fossils from any two equivalent horizons
are not always necessary for comparative purposes. It will thus be the practice here
to refer occasionally to the most important or most abundant fossils found on leading
horizons.

However, at the end of the paper lists of megascopic fossils from the various stages
are given. These are taken mainly from Walkom’s paper and are not intended to be
absolutely complete. They are listed for the purpose of saving the time of any future
worker dealing with the Lower Marine.

With regard to the micro-fauna of the series, the samples of the various horizons
were collected in partially weathered or friable condition, from which some studies have
already been made by Miss Irene Crespin (see list of references). Particular attention
was given to collection of material from the foraminiferal limestones and other marly
rocks near Pokolobin and elsewhere.

THE LOWER MARINE SERIES.
Geographic Distribution.

The Lower Marine Series outcrops over a region of about 200 square miles in the
Lower Hunter district. A small area of volcanic rocks of the Lower Marine occurs near
Muswellbrook. The Lower Hunter outcrops include a large area within the girdle of the
outcrop of the Greta Coal Measures, extending from a little to the north of Maitland

206 STRATIGRAPHY OF THE LOWER MARINE SERIES,

right round the flanks of the Lochinvar Dome marking out an irregular semi-elliptical
boundary which ends on the north-west in the neighbourhood of Branxton and Hlderslie.

In addition to the main Lower Marine province there are the Paterson—Mindaribba
Basin composed partly of Lower Marine rocks, and the country east of Paterson where
the series has a much reduced development. Again, in the Cranky Corner Basin, north
of the Hunter River, Lower Marine rocks outcrop in a continuous subcircular belt
forming the lowest of the three Permian divisions of that locality.

Areas of Regional Development.

It is possible to divide the area of the Lower Marine upon the basis of the develop-
ment of certain distinctive lithological facies of sedimentation. Thus we may recognize
the following geographic subdivisions:

(i) Gosforth—Lochinvar District.
(ii) Paterson—Seaham District.
(iii) Aberglasslyn Estate and its environs.
(iv) EHelah—Comerfords District.
(v) Farley—Ravensfield—Bishops Bridge—Rutherford areas.
(vi) Allandale and Harpurs Hill.
(vii) Old Rothbury—Rothbury-—Greta District.
(viii) Pokolbin—Milfield District.
(ix) Cranky Corner Basin.

The Muswellbrook District Lower Marine forms a separate entity, geographically

and lithologically.

Relations between the Lower Marine Series and the Kuttung Series:

The general relationship of these two series is that of conformity, as there is no
marked divergence in the strikes of the two series at any place, nor, with the exception
perhaps of the section at Pokolbin, any noteworthy difference in the amount of dip.
Small differences of dip, such as do occur, are those to be expected where overlap has
been in progress. Near Lochinvar there appears to be no break whatever between the
uppermost terrestrial Carboniferous rocks and the lowermost Permian.

The section at Pokolbin was interpreted by Professor David as showing a strong
unconformity between Lower Marine and Carboniferous. The present writer, however,
considers that the structural relations are the result of overlap followed by the epi-
Permian folding of Lower Marine sediments which had been deposited upon an eroded
Carboniferous basement during an uplift of Kuttung islands in the Lower Marine sea.
‘This will be referred to again in a later section.

The general overlap in the Hunter Region is clearly displayed by the features
shown on Professor David’s map. Starting in the Luskintyre-Gosforth district and
proceeding east or west along the margin of the Carboniferous province, one can trace
the gradual encroachment by successively younger beds of the Lower Marine over the
Kuttung floor. Near Raymond Terrace the effect is so strong that Upper Marine rocks
rest directly on Kuttung tillites.

Palaeogeographical Setting.

Fortunately the Lower Marine Series in the Lower Hunter forms a fairly well-
defined province, and in any investigation of the stratigraphy and lithological variation
of the rocks, one can obtain an understanding of the palaeogeographical conditions
within the confines of the present development.

It appears that the land in late Kuttung time did not stand very high above the sea.
The invasion of the Kuttung land by the sea began in the neighbourhood of Gosforth.
Gradually the Lower Marine Series began to increase in development, and, as W. R.
Browne (1926) has pointed out, the region of maximum sedimentation was along the line
which later formed the axis of greatest uplift.

We may note that there was probably no high ground adjacent to the region until
the beginning of the Allandale Stage (see below), when a distinct epeirogenic episode

BY G. D. OSBORNE. 207

occurred, elevating some of the Carboniferous floor of the Lower Marine geosyncline,
and producing islands that now constitute the inliers of Blair Duguid and Pokolbin.

Following this uplift erosion gradually wore down the Kuttung hills, and sinking
allowed the accumulation of much sediment. Eventually the Lower Marine sea disap-
peared from the region as a result of the operation of two factors, (i) the silting of
the deposition-area, and (ii) actual retreat.

STRATIGRAPHY OF THE LOWER MARINE SERIES.

The general succession given by Professor David in the Hunter River Memoir
(1907) was as follows:
Farley Stage, with Ravensfield Sandstone at base.
Lochinvar Stage, with Lochinvar Shales at base.
Approximate total thickness, 4,800 feet.
Arising out of the writer’s detailed work, the following classification is used in this
paper:
Farley Stage (with Ravensfield Sandstone at base).
Rutherford Stage (with Bishops Hill tuff at base).
Allandale Stage (with Allandale Conglomerate at base).
Lochinvar Stage (with Gosforth Shales at base).

THE LOCHINVAR STAGE.
1. The Gosforth Section.

W.R. Browne and W. S. Dun (1924) discussed the stratigraphy of the basal portions
of the Permian System as exposed at Gosforth, and at that time (following Professor
David) regarded the Lochinvar Shales (with glacial erratics) as the basal unit in the
System. In recent years, in connection with the completion and revision of the David
Book on the Geology of Australia, Dr. Browne (verbal communication) has finally placed
the Lochinvar Shales as the topmost horizon in the Carboniferous Kuttung Series. Thus
the lowest horizon in the Permian is that of the shales and mudstones with Hurydesma
hobartense, which here will be called the Gosforth Shales. Immediately succeeding the
shales is a well-developed sandstone and sandy tuff carrying plant stems and showing
pebbly phases. This rock persists along the strike for a distance of about eight miles
and is about 80 feet thick. In places it carries marine fossils, Dielasma being the most
abundant, and Spirifer also common. Next succeeding are some olive-green calcareous
mudstones which are followed by 600 feet of basalt. This is overlain by 200 feet of shale
which passes into calcareous material and eventually merges into a local bed of limestone
about 10 feet thick which is exposed in a creek a little to the south-east of Gosforth
Church. This bed carries quite a varied fauna, the chief type being Fenestella internata,
F. fossula, Martiniopsis subradiata, Spirifera aff. tasmaniensis, Deltopecten linaeformis,
Aviculopecten englehardti, Aviculopecten mitchell, Chaenomya sp., Moeonia sp. (from
list given by Browne and Dun).

The limestone is overlain by buff-coloured sandstone and tuffaceous sandstone which
are about 150 feet in minimum thickness and are overlain by decomposed vesticular basalt
and basic tuffs. This basic rock is exposed in a quarry just north of the main road
about 1:8 miles from the junction with the Gosforth Road. Here the section is:

Heet
AaAromrine-Sr ai Med stuits gas Meee eco, est Sate Tees bree y Cane e saee hoa 20
Sub-spheroidal basic tuff Ae ei WM Rete eA UeaeR ty 1 Kad a 2 Ext WOAS TS 15
Olive-green mudstone Pea AMA UE etal pi crricah awe tey Oyen iter bare sana d 10
DECOMPOSED ASAE wy wayece nee acne: eee sk ou fennel seth Vay tial tea larectd 10
Buckledmrm wd stone: (pha wciscnmeer Otis tunes tie hase eat ria ese M SS dt Ae 15
Spheroidaltibasallti i 5s scons oeae es treet! reall hie by eo ig ee Ge Med apd OO

The dip here is EH. 15-25 S. at 50°.

These basaltic rocks form the second group of flows, etc., in the lower part of the
Lochinvar Stage, and they can be traced from the Gosforth Road across the northern
road to a hill to the south-east of Lochinvar.

208 STRATIGRAPHY OF THE LOWER MARINE SERIES,

The record section may be taken as typical of the lower part of the Lochinvar Stage
for the region lying just east of the axis of the Lochinvar Dome. A summarized state-
ment follows:

Feet
Basalt and basic tuffs with interbedded mudstone .. .. .. .. 170
Sandstone andmcuttacecouss: Sand Stone merriment niet mmr nes 1()
TAIMEStONE: Ge. Rea 82 bet Gn Soa SEAR eT RR OP A Oe Wed 10
ShalesiwithwcaleareoulsieshalemaycacuO 0 aye mee eminent ep et e271)
Basalt ack ietstan oye h obaee’ SRS RG Re ees ois ay 6 SUPA A se Nila ree, GOO
Plant-bearing sandstone and tuff, with marine fossils .... .. 80
Gosforth shales with Hurydesma hobartense .. .. .. .. .. 9

Total ..1,219 Feet

Further details of the second group of basalts and associated strata are available
in the property near the Gosforth Road, west of “Anambah’. In a low-lying patch of
country along a small stream three distinct basalt flows are interbedded with mudstone.

2. Lochinvar Section.

This section begins on pebbly tuff just south of a creek to the north of the Roman
Catholic Convent, and is continued south to Lochinvar Station where the nose of the
Lochinvar anticline occurs, and then west along the railway line, linking up with over-
lying Allandale Stage. This is a fairly complete section through the Lochinvar Stage,
but in its compilation a certain amount of generalization has been employed. Thus some
blue sporangia-bearing cherts exposed beside the main northern road to the west of
Lochinvar are incorporated in the statement of the section.

The lowest unit is a pebbly tuff, 50 feet thick, followed by conglomerate 300 feet, and
then sandstone 200 feet. This is succeeded by 150 feet of basalt which is on the
horizon of the Village of Gosforth basalt. Then follow basic tuff 50 feet thick, 10 feet
of hard siliceous tuff and about 10 feet of sporangia tuff or chert, leading into a consider-
able amount of sandy shale. Overlying the shale is the second basalt horizon and then
more shale succeeded by 150 feet of fine shaly sandstone, the top of the bed being crowded
with Ptycomphalina in narrow bands. Drift specimens of Gangamopteris also occur
here. This is an important bed in being the lowest horizon in New South Wales from
which the Gangamopteris—Glossopteris Flora has been recorded. It is also the first —
horizon since the plant-bearing tuff of Gosforth to yield marine fossils. Fine shaly
sandstone separates the two Ptycomphalina bands from one another. The next unit is a
shaly mudstone, free from fossils, and 200 feet thick. The section now is obscured for
about 200 feet, and then comes the third basalt horizon of the Lochinvar Stage. This is
200 feet thick approximately, and is seen on the east side of the road in thoroughly
weathered condition. This is followed by 200 feet of sandy shale with odd erratics,
seen in the railway cutting.

On the main northern road between Lochinvar and the Allandale turn-off these last-
mentioned beds are strongly fossiliferous in one or two zones, details of which are given
in the fossil lists below.

Pursuing the railway section westward one finds next a shale unit about 80 feet
thick, with distinctive bluish fine-grained bands carrying plant remains. In this unit
are scattered erratics of quartzite, porphyry and slate up to two feet in diameter. This
is considered to be the second glacial horizon in the Lower Marine succession. Next
comes the fourth basalt horizon in the Lower Marine.

Referable to this horizon are the patches of basic soil seen on the road from Sawyers
Gully to Lochinvar Station. Clay shale and ferruginous grit now follow for 300 feet
with a few erratics at the top. Then come spheroidally weathering basic tuff 100 feet
thick, and pebbly shales 50 feet thick. This completes the Lochinvar Stage as the
succeeding rocks are at the base of the Allandale Stage. About 200 feet from the top
of the section just described is the only Lower Marine, a glendonite horizon in the
Hunter region.

BY G. D. OSBORNE. 209

A tabular statement of the section follows:

Feet

‘PebblyaSHaAlesmie rd slewmnvori wckeah cscs Muses. cate bustph ih tavcres citreus slog Ul at. 50
Spheroidal tuff .. .. enee0,0)
Clay shale and Rormuninote Brit sti some “Gumnsies ee ine fon e300
Basalt and basaltic tuff, @ourth horizon) 5. .. .. -.. -. -- 00
Shale with scattered erratics et fi ee eee Nene ee 80
Sandy, tuffaceous shale withierratics) <5 5.992: 9.4) s.—=.. -. 200
Basalt (third horizon) .. eetised malate, wy oa O
(Hiatus in section, ee onably aie. 6 SPEIS)) te eee Late aise 0.0
Shaly mudstone ts BP Sel) copa Min eRe ere vue suet 20,0
Ptycomphalina Bed No. 2 pA TN cee eas 0 LS bee a rea eh LR ee 1
Sandy shale Bie ems Paws ch Were Deka. 9 ea eit oes ees 18
Ptycomphalina Bed NO. rf POEs cain eee L Mocige aNSe. eoichts DSL ae 1
Fossiliferous shaly sandstone 18 Soe Ss PO Reo en noe elo
Basalt (second horizon) Me cS acess” Fons, Uhh Cdeet lars sy eee tte 50
Sandy shale EE WEY ope Mee cs TA, Siszs. y SRE UAIESY ccsal piadaa asta OO
SPOKANE Amb UL terre em aye yeaies Sree iymetarcPel Uiiusenry siti ple eae) @ (eyes ats 10
TSieWRebe, WOLDES Glo clots eta aee RLS ioe Gascon Meee Ce Ci neh ils | ER Nra tees PUR Chey ara 10
IBFIGMO Wblie Go. o oe 2 bere adhe Gee Sane eee 50
Basalt Gene cdaieraal) al 1 inozizan Be aR eich ed aac Ue esa
Sandstone eeawiere-w bent eyes pictnr yea eta Aetna Pec ey ome, 200,
Conglomerates sume sce a ise) cusp Gond-e Uiket ay eae ser cee 300.
Pebbly tuff SRP MEd arate seth a ence eat any nae ett Lon ne eee 50

Total .. 2,740 Feet

3. Helah—Comerfords Section.

This section takes in the interval from the base of the Series up to basalt which here
immediately underlies the Allandale Stage. On top of the Carboniferous Lochinvar
Shales, as seen in Helah Creek, comes mudstone containing plant remains for 25 feet,
followed by buff sandstone for 100 feet. Then comes the first horizon of sporangia tuff,
which is 60 feet thick. Pebbly mudstone and sandstone now succeed, and are followed
by bluish sporangia tuff (second horizon), also 60 feet thick. It is to be expected that
the seasonal development of the sporangia would be reflected in the zonal and restricted
character of their occurrence in these rocks.

The next group of rocks, 600 feet thick, comprises tuffaceous sandstone and tuff,
some almost of quartzitic texture, and a very thin intercalated flow of basalt. These
clastic rocks cross the Helah—Rosebrook Road and make an imposing scarp on the left
bank of the Hunter River, where they dip E.S.E. at 18°.

Basalt flows and associated basic tuffs aggregating 400 feet now succeed, and lead
up to the base of the Allandale Stage. A summary of the section is as follows:

Feet.
Basalt and tuff Soon ae EE ee ea ee eh: Poe ely Sets gta ed ().()
Tuffs and tuffaceous ictOne Aree Gee eae ne toe eon cay acme cess, ko UlO),
Mudstonewand= tuft minterbed ded as aie) laren oe ysienreennn 2.3.0
Basalt SENSE awes Fe Tike aeRO Re oe ARN EARS SPN CARI Be Nroe hye mode — Vana ok 10
Coarse tuff ses Se Ee a ER ye os 4 Kar Oma are ee os 50
Sporangia tuff or hen (No. 2) MA COCE Nc leche - ee epee Rar. Sie 60
Mudstone .. . cb hl Oe eacbn ts Shin were cote et. ics Gt eae a eee ane 50
Pebbly Sandstone ae Pee ee ey aS eee OU eh la a ie 60
Sporangia chert (No. 1) SPM he BRS rtd eR LOA Water? (Satan hee s 60
Buff sandstone .. .. Ae en wl ey epee) Werks Weenie re eeyy dice) dey ty pe OO
Mudstone with plant remains BEET 2d MRO TPR RENT SIS ae ee SEALE: eee Tefen” 25
Total ..1,395 Feet

4. Section from Paterson—Tocal Region.

West of Paterson township overlapping of Lower Marine units brings certain
sporangia-bearing cherts into contact with underlying Carboniferous tillite. These
cherts are 25 feet thick and are followed by basalt for 20 feet and then more bluish
sporangia cherts 150 feet thick, and some dense blue cherty rocks with plant stems, which

210 STRATIGRAPHY OF THE LOWER MARINE SERIES,

pass into a band of sporangia-tuff (third horizon) overlain by more tuff and a peculiar
felsite breccia recalling the Kuttung volcanic facies. The rocks in this region are disposed
in a flat syncline trending more or less east and west, the south side of the structure
being exposed immediately north of Webbers Creek Railway Bridge.

Although the present section is taken at no great distance from the Helah region
and in the same structural unit (the Mindaribba Basin) one finds several differences in
the sequence. The most important of these is that the LOSSES) at Comerfords are evidently
higher stratigraphically than those at Paterson.

The Paterson section may be summarized as follows:

Feet
Melsite: Dreeciay i520 x in Lee yee pe eee yore. beet ee halen een 90
Coarse tuff .. Scale Se ook OEE be Che oe bee ennd ah a Ris 50
Fine sporangia- ane ne. 3) See ee eee Bes cretion Fst Sey eceea ah Brsirs 30
Plant stems in mudstone Ea aarti ieema ssl censienmin s cabin" 31's) 8 50
Basalt : APNE SAN otrtnaet ch saeT tee Ul corey Meee raat lle cy PAI\N)
Sporangia- tuft ane 2) i, Svbua dagen tab uae Rt nts koe ea RECT eer aN so. LB)
Basalt 5 Sa Re Bae lena ee dens i. hace ee ne ort ae ee 20
Sporangia- tuft (No. 1) Sete, (ba SERS RE aes Weert Oe a hel 25

Total 2. 61'5 Meet

From the sections detailed above a generalized statement has been prepared as
shown in Text-figure 2.

The consideration of the Lochinvar Stage in the Cranky Corner Basin will be made
at a later stage in this paper.

THE ALLANDALE STAGE.

The strata grouped into this Stage are distinguished by the following features:

1. They appear to have been developed about the time of some physical change in
the Hunter region. This was localized in several places and amounted to uplift of the
floor of the deposition-area, giving islands from which coarse conglomerates were
derived.

2. They comprise two palaeontological horizons, the more important Hurydesma
cordatum bed, and the zone rich in Aviculopecten and Deltopecten. These organisms
were able to subsist in the coarse gravelly, littoral environment. (In four out of five
places where the Hurydesma bed occurs in the Lower Hunter the rock is a conglomerate.)

3. Distinctive tuffaceous rocks broadly grouped in the past under the title of the
“Harpurs Hill Sandstone” indicate an outburst of explosive vulcanicity more pronounced
than any that had taken place hitherto in Lower Marine time.

Thus the separation of these beds from the rest of the Lower Marine and the
recognition of the Allandale Stage is justified, particularly also because of the importance
of the Eurydesma horizon in the correlation of the Permian beds of Eastern Australia.

1. The Allandale Section.
(a) Preliminary.
This section has been compounded from exposures available as follows:
(i) In the railway cuttings to the east of Allandale Station, where the rocks
form part of the western side of the Lochinvar anticline;
(ii) from the old road between the station eastward to the overbridge
(iii) from the Allandale cross-road.

The beds found in the railway cuttings can be traced north-westerly to the cross-
road leading from the main Northern Road to Allandale Station, and beyond this. North-
west from the cross-road the conglomerate thins out and the tuffaceous sandstone is
strongly developed as may be seen beside the road at the summit of the rise near
Harpurs Hill. About 12 chains further west along the main road some coarse tuffs and
sandstones belonging to the Stage and overlapping the Harpurs Hill horizon are
artificially exposed in a small quarry, just north of the road. Here fossils are abun-
dantly developed.

BY G. D. OSBORNE. maalal

As regards the palaeontological horizons, there are, in addition to the well-known
Eurydesma bed, and the important Pecten horizon, several richly fossiliferous zones,
the prominent genera being Martiniopsis, Keenia, Spirifer, and Fenestella (see footnote,
Deal).

LOWER MARINE SECTIONS

RUTHERFORD
STAGE
ALLANDALZ Total Thickmess

STAGE 1170 ft.

Total Thickness BARLEY

: : STAGE
4 Pecten Bed a ae Total Thickness
Black Tuff . 985 Feet

LOCHINVAR :
-|Conglomerate

STAGE

mop aly rbickness Harpurs Hill
Shale ie

Spheroidal
Tuff

Coarse Congte.
with Burydesma | : Shale and “7 7" "| Mudstone &
=i Muds tone an Sandstone
[xxuxnax| Dark Tuff . —|Martiniopsis
Allandale : Seca yeou-
Shale and © 0 oo| Conglomerate : F phalina
Grit with (Allandale : 5 || BEE ins <==] Martiniopsis Bed
erratics District) -- Grits
Muds tones
Basalt No.4 z F Sandy Shale

Shale with Basie Tuff ‘ Spacer

: *: stone
erratics with Pectens clap pane pera matd Gases is
Sandy Shale

with Erratics

Basalt No.3 Sands tone

ae eal d Tuffaceous

Grits
Sandstone

Fenestella
Limestone

Ptycomphalina |; -L.| Tuff and Sandstone
Tate I: .-' | Sandstone

Limestone
Fossiliferous se eae a “| Sandstone

Sandy Shale
Basalt No.2 Basalt

Conglomerate =: 50 Ft comphalino
with Eurydesma pc Bed
Shaly S.S. (Pokolbin

District) Bluish

Sands tone
Sporangia Tuff —- - ; ;
Tuff : > : | Fine
—. : =] Sandstone
Basalt Noel ae LS aN Beret
- --- e-.-] Sandstone

ee --2-| Ravensfield
andstone Sandstone
(Plants) : 2 ;

L.S.
Bryozoal Tuff
Tuffs and Bishops Hill
Conglomerate Bed.

Gosforth
Shales

Text-figure 2.

212 STRATIGRAPHY OF THE LOWER MARINE SERIES,

(0) Details of the Section.

We begin with 80 feet of conglomerate which eeetiee the topmost shaly beds of
the Lochinvar Stage. This is succeeded by dark greenish-black tuff 20 feet thick, and
then by 150 feet of very coarse conglomerate. This conglomerate is split in only one
place by a band of tuffaceous sandstone of the “Harpurs Hill” type, 15 feet thick. This
conglomerate contains the famous Hurydesma Bed, great numbers of the lamellibranch,
together with Keenia, Notomya, Edmondia and other types, making a regular shell-
bank deposit eighteen inches thick.

Succeeding the conglomerate whose pebbles are almost entirely of albitized
amygdaloidal andesite derived from the Kuttung inlier to the south, come 50 feet of
the true Harpurs Hill tuff overlain by 20 feet of soft brown shale with erratics. Next
follow 80 feet of conglomerate, similar in structure and lithology to that exposed in
the more easterly of the two railway cuttings. Succeeding the conglomerate is a dark
greenish-black tuff, obviously of the same general facies as the “Harpurs Hill” type.
Towards the top of this unit one finds the Pecten bed. This is about three feet in
maximum, but 14 feet in average, thickness, and contains numerous examples of
Deltopecten together with fragmental material partly representing broken-up shells of
Spirifera and Martiniopsis. Fenestella is also prominent on this horizon, which is
followed by the topmost unit of fine conglomerate, 10 feet thick.

The section on the Allandale cross-road indicates clearly that glacial conditions
were present not far from the Hunter region in Allandale time, this being the third
glacial epoch of the Lower Marine Series.

The Allandale Section is tabulated thus:

Feet

Pebbly mudstone SURE ath Steen toy come oem eee) See en MS ear 10
Tuff with Pectens .. aay Meee ak oe 2 eS hn = dose Per SD 3
Blackish-green tuff aig te. Cal lenctith! Polis) crams eeeey yt eeren aes ales 50
Conglomerate (andesite Boulders) AIS aN cares ae iets sO ie bea 80
Soft brown shales with numerous erratics ce eh BA eta 3 20
Tuffaceous sandstone (Harpurs Hill type) ae 50
Heavy Conglomerates with 15 ft. tuff band ain Thue desi

MOTIZ OT AEE Saeed ae Ca ae Sayin Ron ates Oe cacti Re eee ananL tye au prema Ey ()
Greenish-black tuff ete Ane Vie erica ie ee, nets Un) Case ens 20
Conelomenater cea ea iis ye eee Re aise Lev Aree ps See 80

Total .. 463 Feet

2. The Allandale Stage in the Pokolbin District.

The Pokolbin area is very complicated geologically, and the elucidation of the
detailed stratigraphy and structural relations of the Kuttung and Permian rocks will be
possible only after much more field work has been carried out.

The somewhat special nature of the structural features of the Pokolbin region as
a whole is due to the following:

(i) Permian sedimentation probably did not take place until the beginning of
the Allandale Stage.
(ii) The conglomerates accumulated upon an uneven floor during conditions
that gave rise to strong overlap.
(iii) The folding of the Lochinvar anticline was attended by the development
of local structural irregularities due to the Permian sediments being
squeezed around and against the rigid inliers of Kuttung rocks.

It is somewhat difficult in this district to know just where to draw the line between
the Allandale and Rutherford Stages, but by tracing the latter from areas of favourable
exposure it is clear that the only units definitely to be assigned to the Allandale Stage
at Pokolbin are the conglomerates and sandstones which in places mantle the lower
slopes of the Kuttung inliers, and the overlying basalts and basic tuffs.

The conglomerate, 200 feet thick, is very distinctive, possessing a fair variety of
volcanic pebbles, derived from the Kuttung floor. The coarse andesitic matrix contains

BY G. D. OSBORNE. 213

Eurydesma cordatum as well as other typical Lower Marine forms. Excellent exposures
are seen in many parts of the area. The sandstone is fairly coarse and definitely
tuffaceous in places, being 300 feet thick.

The succeeding basalt is obviously of submarine development, being sometimes
vesicular, sometimes amygdaloidal with zeolites, and in places showing pillow structure.
The maximum thickness is 400 feet. It is overlain by basic tuffs approximately 100 feet
thick containing many Pectens. Overlying these come the foraminiferal and bryozoal
marly limestones of the Rutherford Stage.

3. The Allandale Stage at Comerfords.

In the neighbourhood of the cutting where the Helah Road crosses the North Coast
Railway the Allandale Stage is exposed in the form of strong outcrops of a coarse, ill-
assorted shingle and tuff deposit in which Hurydesma, Spirifer, Fenestella and Delto-
pecten are abundantly developed. This bed is about 20 feet thick, and rests on a
basement of basalt. It is clear that here the Hurydesma and Deltopecten horizons have
coalesced. Immediately following the tuff is a fine-grained felspathic rock which
suggests volcanic activity coeval with that which produced the tuffs of Harpurs Hill
and Pokolbin.

THE RUTHERFORD STAGE.

These strata, in so far as they occurred on the eastern side of the area, were referred
to by Professor David as the Rutherford Shales, being regarded as the upper portion of
the Lochinvar Stage. The present writer has instituted a separate Stage because of the
following reasons:

(i) Detailed study of the areas of this Stage has provided much information
not available when Professor David wrote his Memoir. This indicates that
the rocks are fairly definitely marked off stratigraphically and lithologically
from the underlying Allandale Stage on the one hand, and from the over-
lying Farley Stage on the other, except perhaps in the case of the topmost
part of the Rutherford Stage on the eastern side of the Lochinvar anticline.

(ii) Within the Stage there is a greater variation of facies-development than
in any other part of the Lower Marine in the Hunter region. Thus in the
east the rocks are mostly shales, definitely non-calcareous, except for very
local concretionary structures. In the Pokolbin—Rothbury—Illalong districts,
however, there is extensive development of marly shales, foraminiferal and
bryozoal limestones.

(iii) In the Stage are two well-marked fossiliferous zones which are of distinct
stratigraphical value. These are:
(a) Foraminiferal and Ostracodal marl and limestone containing also
Bryozoa and other fossil remains, this being the horizon of the well-
known Pokolbin unit described by Chapman and Howchin (1905).
This occurs about 200 feet above the base of the Stage.

(b) Sandy shale with abundant remains of Spirifer, Martiniopsis and
Fenestella. This occurs about 700 feet above the base of the Stage.

A. B. Walkom noted two horizons of limestones in the rocks which I have designated
the Rutherford Stage, and as far as I can make out from his paper, the two horizons
given above are identical with his zones (see Walkom, 1913). However, he did not deal
with the rocks of the marly type, rich in foraminifera, which are abundant in the Old
Rothbury-—Black Creek—Chick Hill District.

Through the country stretching to the south and south-west of a line from Allandale
to Red Hill Trigonometrical Station, and particularly to the west of Black Creek these
horizons can be traced, and break the monotony of the rest of the Stage which is formed
chiefly of buff-coloured sandy shale and sandy tuff. The foraminiferal limestones do
not appear anywhere on the eastern side of the Lochinvar structure, but the other zone
marked by many Brachiopods and Fenestella has been located at two places on the

214 STRATIGRAPHY OF THE LOWER MARINE SERIES,

eastern side. These are near Bishops Bridge, and in the bank of Black Creek in portion
25, Parish of Allandale on the edge of the Parish. The latter outcrop is identical in
appearance and fossil assemblage with that in Portion 63, Parish of Rothbury.

The basal unit is a pebbly tuff with Bryozoa, which shows a marked lithological
contrast with the underlying Allandale Stage. It is here called the Bishop’s Hill
Horizon, because of its strong development in that area.

SECTIONS IN THE RUTHERFORD STAGE.

It will now be appropriate to take the details of development along various section-
lines. In the case of the intricate area around Pokolbin and Mt. View a generalized
section has been determined.

1. The Rutherford Section in the Pokolbin District.
The following has been prepared from areas adjacent to one another:

Feet.

Sandstone (probable equivalent of Ravensfield) Se oe cee: —
Sandy mudstone with abundance of Martiniopsis and

Ptycomphalina ra Oe th ote oF gee Elita egy Me ay ASE ee AY ieee al Win Fda A oem ante) (0)
Sandy shale A Gateee FAA eee chante sae tps weer in eran MLE ()
Ferruginous cemaetane with iMantinionsts Sete Ge ey om RE eco 10
Sandstone ie Sa mua een AAU Ce aN ested oh UR ey 10
Sandy limestone Aaah emestena ie haps hy a keee yt coeetice La sae te) se eae ee 10
Shaly sandstone BITSY be SNe ENE Ss Se eee rae Search Ae ok My ial am onal oh cpt LNG)
Sandy limestone SEU EERE eb eh | Rema eG ae be eal eo OSes [in et RSIS Renee NN EAS 10
Basalt ah SOIL SAMs ae ene ace SE MIAPON ” PANS ye, Banh otek Sr acen EDC)
Sandy shale itt arnatios Be gst ta PE) EO ey ane ates Son een we 200)
Fine sandstones and tuffs Hee SS ne te eal se ae WE TO a 35
Foraminiferal limestones aah hee Aint eye 2 eaten See Sea Noes 20
INGA OS ISH IBiAyOVORNl WenEE 55 aol Boll ee.’ oe) oo on os oc 25

Total ..1,170 Feet

In the neighbourhood of Jackson’s Hill and to the north the broad succession is:

Feet.
Sandy shale with erratics .. . Pe ieee Si eevmnmece | Pretive nee,
Sandstone with Martiniopsis and Spir ifer pot PeUNARE SAM RR! pulse bere tedbilly 30
ShalyasandstonemwitheMenvestellan wean bled nln ean tee eet 50
Sandstones, often ferruginous Si iy Rel OA eater taeicun bole ookod Be eae Eee OU.
Marls and Ostracod limestone eM Sy cerca eee htleler adel ce} ies caiseann mtn (0)
Notaly = 38 0lneet

Near Pokolbin itself, where Foraminifera were first collected, a little to the west of
“Ben Han” vineyard, and in the neighbourhood of ‘‘Maluna” in Portion 112, Parish of
Pokolbin, we have the following section:

Feet.
Foraminiferal Limestone with Bryozoa and Pectens oth, picke, Meee 4
Fossiliferous shale .. .. soatgray Meee sun tetas tevree apa 5
Limestone with Fenestella and SEenOnOnG” SO, Sach aero anie a cee oh ts BA Ry keg 7A)
Blue limestone .. .. ne baeaet ga eas 5
Mudstone with Ine VeCnieniGn apne aren peodal IRONING. G5 oa os 10
Total .. 44 Feet

These strata succeed the Allandale Stage.

The detail of the Foraminifera which occur in the topmost bed can be obtained from
a perusal of the Memoir by Howchin and Chapman (1905).

There are several places in the broad belt of Rutherford Stage rocks north of
Pokolbin where interesting sections are available.

Thus a cutting on the main road from Pokolbin to Branxton at a point about five
miles south of Branxton exposes the following section of the Rutherford Stage:

BY G. D. OSBORNE. 215

Feet.
Fossiliferous sandstone with marked band rich in Martiniopsis
and Spirifer se REN AP hou caEmeke. | isis 25
Finely bedded shales sittin enell erential Ee Matyi st een vk meee 25
Sandstone .. . Cie Oe cre Ue revaMbres a lnietesemeiale: sic foie i cele 20
Pebbly ironstone aan foaaile WEE fie: CSCS MRUetc gent ET, oes feats thew 2
Hines shalestwithiescatvteredmpebblesems. | srcusmcism asi anise eye ler 20
Totaly... 92 Feet

The highest unit in the above section is the uppermost of the two fossil zones
already referred to at some length above.

Between Mt. View and Pokolbin on the road running west of Jackson’s Hill there
are many exposures of the foraminiferal limestones and associated rocks. In the
cutting near portion 106, Parish of Pokolbin there occurs a beautiful polyzoal limestone,
which is approximately the equivalent of the ‘‘Ben Ean” horizon. The section on the
roadside is as follows:

Feet.
Polyzoal limestone .. .. Miata UPA pa otic Hay ese Wie ait S US wrt bey tice Melia 152
Marly shale, with small foseils Sher yu ib tS a oc SRA pe eee Meera il)
Limestone with Stenopora .. . S Sees Gee GES Cervantes meee erat 12
Calcareous shales with Wenesicua te FAA; si gHahY donepseriaie sy ais 12
Polyzoal limestone crowded with small fossiley seeks Bite ate 4G 10
Totals: 61 Feet

The only place on the west side of the Lochinvar anticline where the sandy phases
of the Rutherford are developed to the exclusion of the calcareous types is along the
right bank of the Hunter River to the N.E. of Greta and east of the Leconfield Road.
Here in a cliff and scarp-section one can note the presence of sandy rocks, mostly
unfossiliferous, but possessing one band with fragments of Fenestella.

Passing over to the eastern side of the area we find a section of the Upper Ruther-
ford exposed on the main Northern Road about three miles from Maitland. Beginning
opposite the gate of a small cemetery we have—

Feet.
Brownesandstones GRavensfield) horizon), 5. a8 6s oc) es —
Pebbly reddish sandstone Oe, Sia RE Ge Sch ae: ERG” eee aie 15
Tele! TSO! WRURAOCSCOWE GENTOO Go aco oo 6n coo od oOo oD 10
Current bedded sandstone a Saat LepeO RARE SED EARS Tone eh y e 25
Brown sandstone with shell fneients abt ESTE RO EP ENN, Del Leask 34
Pebbly sandstone .. nie ACEO SEES Mice Pr MIRE CDR eK oar 12
Whitish shale with Piycomphatina Pia Mere eI erg Le ere ne ome ee Nile ees 10
Buff sandstone .. .. Memo meen ealohte, hore etme taints catch ale 12
Fine grained sandstone atin Spir ifer UMAR Sirst hed acute ReGen sateen ctr is wtelss 15)
IEVAO\ AN WUNIEKOCOUIS SENNOKMONKS Gh do ° op oe con 2 65 G0. 65 oo 40
Total .. 173 Feet

A fuller section of the Rutherford Stage in this eastern part of the province is given
in the cuttings along the railway line between Lochinvar and Farley.

Starting at a point east of Lochinvar and near the 125 mile peg on the railway
line, the section begins on very soft and poorly exposed shales, but to the east stronger
outcrops are seen and we have the following:

Feet.

Shales ra RE aie otek enone Ga ee 80
Cherty shale iki ‘eainy plant. atenis b Sab make A nly 4 etal Geyer ial BeciaaeD teste 60
Shinolsicome Tien Ganev il vols 54° (gok abl Go “om ool. “ae dine os 30
CHICAIeSOUIE ComocnOneinr SONI soso Soot “Gol “oo Mba le6 eae 60
Hard sandstone SER esto 7 nabloy LIS et Mil Ponies Rae mers DNS rd satin ee one 35
Bluish chert oa) us SUBLE ze Maree Pathe Caen ene es na eeks 40
Decomposed bluish mudetone ae Teoh AC OST es eat eee: 20
Shaly mudstone with shell eernentan Pe a ey Sole SPs baer Ly nT 30
Sandy mudstone ne 40
Alternating shales and sane n ee Sain occasional, “aiken remains

andeoddmlayerserich lnweManCintOpSisaas sees else) ee SO

Totaly escwomomiecc

216 STRATIGRAPHY OF THE LOWER MARINE SERIES,
Reviewing the data for the Stage it is seen that the maximum thickness is 1,170 feet.

THE FARLEY STAGE.

This Stage, the topmost of the Lower Marine Series, extends from the base of the
Ravensfield Sandstone to the base of the Greta Coal Measures. The maximum thickness
measured by the writer is 985 feet, but in some places the Stage dwindles to about
125 feet thick.

The persistence of the Ravensfield Sandstone has made it relatively easy to refer
to their correct stratigraphical positions certain sandy beds which occur associated,
both above and below, with that sandstone. Although there are many arenaceous
horizons in the Rutherford and Farley Stages which approximate in general lithology
to the Ravensfield Sandstone, certain features about the latter give it a character by
which a worker thoroughly accustomed to it is able, in almost every case, to distinguish
it from similar horizons. This sandstone was mapped by Professor David as an
indicator horizon.

As regards the extent of the Ravensfield Sandstone, it is interesting to note that
it ends abruptly in two places at least. One locality is near Mt. View and the other is
north of Rutherford.

On “Aberglasslyn’”’, Nicholson’s Quarry has exposed a greenish-black to buff sand-
stone which weathers brown in a manner suggesting the Ravensfield, and so regarded
by some workers. A close scrutiny of the available exposures, however, has led the
writer to regard it not as the Ravensfield horizon but as equivalent to some unit in the
section seen on the left bank of the Hunter River opposite “‘Aberglasslyn”’.

Distribution and Sections of the Farley Stage.

This Stage is the most consistent, lithologically, of all the groups of sediments in the
Lower Marine. The beds are dominantly sandy with occasional shaly layers and are
often decalcified, interbedding of rocks of differing resistance giving characteristic
topographic expression. The Farley Beds have an areal distribution greater than that of
any other group.

The stage is exposed in and around the village of Farley, and stretches to the
south in a broad belt through the Bishops Bridge and Sawyers Gully district. From
Sawyers Gully to the south and south-west the Stage is well represented, and occupies
high country in the Mt. View—Milfield districts. On the west of the Mt. Bright inlier it
is cut out by the Matthews Gap Fault, but reappears on the eastern side of the fault
in the country north of Pokolbin. These rocks are well exposed in Black Creek west of
Rothbury. Further north from here they are prominently developed on the northern
and western slopes of Molly Morgan Ridge and the high country running north from
the Trigonometrical Station. Further north the Farley Stage is found immediately
east of Greta, and again well exposed east of the Leconfield Road, and near Dalwood
crossing of the Hunter River. The next place northward is in the Cranky Corner Basin
where the Stage is much diminished.

In the north-east of the province near Comerfords and west of Bolwarra this Stage
is represented by about 300 feet of strata underlying the Greta Measures. In this case
the Farley beds thin out quickly and eventually become overlapped by higher beds.
Marine fossils are abundant on some horizons. Apart from the fauna of the Ravensfield
Sandstone, there is a fairly wide variety of type, as shown by the list given below, some
thin ironstone bands being crowded with Martiniopsis and Spirifer. Conularia is also
quite common in certain gritty ferruginous rocks, and Ptycomphalina is abundant on
two zones which occur at widely separated places.

Sections in Farley Stage.

Three sections are taken to illustrate the stratigraphical features.
(1) Farley Station to Farley Road Corner.

This gives a completle sequence, and may be stated as follows:

BY G. D. OSBORNE.

Mudstones and brown sandstones
Shale crowded with Martiniopsis
Tuffaceous grit

Bluish mudstone

Bluish sandy shale ..

Sandstone with Martiniopsis ae Ser ycompnalne
Plant-bearing tuff

Bluish-grey tuffaceous grit
Sandstone :

Shale rock with Pivcenronalind
Bluish sandstone

Fine buff sandstone

Pebbly sandstone

Ravensfield sandstone

(2) On Bushland track south of Mt. View.

The section begins about eight chains from the Mt. View school on a sandy unit
which is just above the foraminiferal limestone at the top of the Rutherford Stage, the

Ravensfield sandstone having temporarily failed.
Section:

Buff and brown sandstones .. =
Shaly sandstone with Ptycomphalina
Mudstones with plant stems ..
Sandstone with Martiniopsis ..
Concretionary Limestone

Sandstones with Martiniopsis pana
Calcareous shales with fossils

Sandy and slightly tuffaceous beds
Gritty calcareous tuff with Fenestella

(3) Farley Stage on the extreme west of the Province.

North of the Pokolbin inliers (where the Farley Stage is excluded by the Matthews
Gap Fault), the strata of this division emerge as the fault swings a little to the west of
north and transgresses successively younger beds at the surface outcrop. The lithology
of the Farley representatives here is similar to that of the beds on the Sawyer’s Gully
road. A section was measured from the outcrop of the Ravensfield Sandstone beside
the bridge over Black Creek where crossed by the Pokolbin—Branxton road. The details
are as follows:

Sandstone and grits

Grey sandstone : :

Blue sandy shale with Tete remains ..

Shale ,
Fine blue shale cartin Fa eats ei Brachiopods
Sandstones with plant remains

Blue shales with Spirifer

Bluish-grey mudstone

Sandy tuffaceous rocks arith worm encke.

Buff sandstones with pebble beds, occasional Bal, vers Ant

some fragments of shells
Ravensfield sandstone

Total

Total

Total

Feet.
200
12
20

2
v

20
100
15
220
155
10
100
30
70
20

985 Feet

The following is a summary

Feet.

600
10
100
5

10
50
10
100
10

895 Feet

100
10
12

430
20

737 Feet

THE LOWER MARINE ROCKS OF THE CRANKY CORNER BASIN.

This basin is entirely divorced from the rest of the Permian province.
north side the Permian are brought against the Carboniferous by the Webbers Creek

217

of the

On its

218 STRATIGRAPHY OF THE LOWER MARINE SERIES,

Fault, a fracture connected with the Hunter Overthrust System. Elsewhere the
relations of the Lower Marine to the underlying Kuttung is one of essential conformity.

The chief interest of the Basin is that, although lying to the west of the probable
axis of early sedimentation in the Lower Marine trough, the Lower Marine displays a
condensed section of representatives of the Lochinvar, Allandale, Rutherford and possibly
Farley Stages, the total thickness being about 810 feet.

A. B. Walkom described the strata of the Basin in 1912, and the present investigation
extends the data given in his paper.

The lowest beds are coarse plant-bearing sandstones exposed on the N.E. corner of
the Basin against the Webbers Creek Fault. In the extreme N.W. of the area blue chert
occurs which is to be correlated with other occurrences as discussed below. Over a good
deal of the area of the Lower Marine outcrop there are flows of amygdaloidal basalt
which often rest directly upon the Kuttung Series. Beginning with the basalt in the
S.W. there follows bluish chert about 180 feet thick. The next 200 feet comprises several
fossiliferous bands of shale, the most significant being two Fenestella horizons, separated
by grey shale. Small fragments of lammellibranchs occur in the beds overlying the
upper Fenestella bed and then follow sandstones with well preserved Spirifer and
Martiniopsis. On top of these comes the Hurydesma Bed. This unit is crowded with
thousands of shells, which are more abundantly developed here than anywhere in the
Hunter region. The matrix of the mass is a greenish tuffaceous grit or sandstone.
Overlying this zone is a buff-coloured series of tuffaceous sandstones and grits with
almost a quartzitic appearance. These are partly Rutherford and partly Farley. On top
of these rocks come the Greta conglomerates. The section may be tabulated thus:

Feet.
Tuff and sandstone, etc. .. et aes 14 A ETE AR CU a Seto. | crore ey eer eG)
Hurydesma bed Dea) Ae eetgens = clave iil atade ) A (cue) iy SNe mA Bete ten AROrS Ue cay SRDS 15
Yellowish shale Bar ENG oe Wace ko Steet toe a oe etek tae eae ce ee aed tte 50
Blue shale .. .. Pe ee S 3 os Sn PMR Ta ah ERS Nene fea thee ALS
Fossiliferous GRO SR Rs 5 a Nt cs hrcm yest tad isla: See ae ee tae 10
Blue chert with small Teameltoranicher Bed vac EN Saher oy Ee 20
Massive: SANGStONED: sim aeekte hee escie sian Diciiseiaiarcie thea” aspen) Vue omen ne 30
Fenestella bed No. 2 ite cos co eR te ae ees aoe a eae 5
Grey shales as TERRA LA NE ah NVI Cosine, Rede Pe hi ee ne 50
Fenestella bed No. i is Sx tee eh sikeas Sig so Pau ekg WA ret Mote oo eee 5
Mudstones in eee itr Susie, tone Oe ARE ero ee MEO 40
Olive green eretOne Se MR nee se aT Um rer met | aerate ie ag TA 10
Blue cherts Be ricaaiciee ee eee sh hahaa Seem ethan Bais ae oe JLS@
Basalt Be DAMS TIMP hohe oe Mae ort Ee ly ton Oiaa ENE ibs eum aLOA |)
Plant-bearing Sandctonen aydjen Percy aiaeese feaael ay feyea Ma aa ae ee mate eel 00)

Total .. 810 Feet

THE LOWER MARINE IN THE MUSWELLBROOK DISTRICT.

From a brief examination of the Lower Marine at Muswellbrook, the writer can add
practically nothing to what H. G. Raggatt (1929) has reported. Thus there are two
main sets of rock comprising the Gyarran Series and forming a relatively small area
of outcrop. These are amygdaloidal basalts (identical with those of the Lower Hunter),
overlain by rhyolitic lavas, breccias and tuffs. The former are about 600 feet thick and
the latter 115 feet. There are no sediments worthy of examination, as the tuffs are
thoroughly igneous.

CORRELATIONS OF SECTIONS.

A certain amount of correlation can be effected between sections in the various
Stages from fairly widely separated districts, but considerable difficulty has been
encountered because of the scarcity of outcrops in low-lying country and the unreliable
character of the dips shown in surface expression of some strata.

The Lochinvar Stage.
The plant-bearing sandstone so well developed in the Gosforth district is probably
to be correlated with the coarse tuffaceous rock with plant stems at the N.E. of the

BY G. D. OSBORNE. 219

Cranky Corner Basin and with some plant-tuffs at Helah. (The question of the
correlation of the basalts is left until later.) The blue sporangia cherts which occur
in the roadside exposures west of Lochinvar are regarded as the equivalent of one of
the Paterson Horizons, probably the lowest which outcrops on the road west of the
township. The similar cherts in the North Coast Railway cutting near Quarry Creek,
Tocal, are the equivalent of the highest of the chert horizons on the Helah road section.
The blue cherts on the north-west and in the southern part of the Cranky Corner Basin
are the equivalent of the rocks on the road west of Lochinvar, corresponding also to
the similar rocks from Butterwick and Dunns Creek district, east of the Paterson River.

The Fenestella beds in the Cranky Corner Basin, seen best in the creek below Mr.
Thomas’ property, are probably on the horizon of the marine shales outcropping on the
Northern Road one mile west of Lochinvar, and also of the similar strata overlying the
Ptycomphaiina bed near Lochinvar station. Fenestella is particularly abundant in this
horizon near Lochinvar.

The Allandale Stage

The Hurydesma cordatum horizon is developed at five places. In two of these
Pectens are commonly associated with the Hurydesma, mostly in the beds immediately
overlying the conglomerates. This is the case at Comerfords and at Pokolbin and is
due to convergence in the sequence whereby 280 feet of strata have been cut out.

The lithology of the matrix to the Hurydesma fossils is variable. At Pokolbin and
Allandale the rock is a coarse conglomerate with tuffaceous cement; on the south side
of Blair Duguid a sandy tuff with large glacial erratics; at Cranky Corner a greenish
tuffaceous sandstone, and at Comerfords a very gritty impure calcareous tuff with
irregular boulders.

The Harpurs Hill tuffaceous sandstone of the Allandale district can be correlated
with the tuff overlying the Pecten Beds at Comerfords and with the basic tuff on the
road near Portion 193, Parish of Pokolbin. Greenish tuff associated with basalt on
“Aberglasslyn” is also to be placed at this horizon.

The Rutherford Stage.

Mention has already been made of the essential contrast in the general lithology of
this Stage as shown by its outcrops on eastern and western sides of the province. In
spite of this change, correlation can be carried out by using the horizon of concretionary
limestone carrying abundant Spirifer and Martiniopsis. Along the sector from Red Hill
and the Molly Morgan Ridge through to Pokolbin and Mt. View correlations can be made
between detailed sections of the marls and bryozoal limestones. Thus one unit which
can be detected in many sections is a grey limestone with much Fenestella but also an
abundance of tubular Stenopora.

The Farley Stage.

Apart from the Ravensfield sandstone which is so uniform and persistent there are
two zones rich in Martiniopsis and one with Ptycomphalina which help to link the
widely separated Pokolbin—Milfield and Farley districts. These three horizons are well
displayed on the Farley road, the main Northern Road, the Sawyer’s Gully Road, and
on the bushland track south-west of Mt. View. Also two of the horizons can be traced
in the bed of Black Creek west of Old Rothbury.

Horizons and Correlations of the Lower Marine Basatits.

An exhaustive petrological examination and much more field-work are required
before many of the problems peculiar to these rocks are solved. Concerning their
correlation it is clear that they do not occupy constant stratigraphical positions.

The section from Gosforth to Lochinvar Station and westward on to Allandale shows
the presence of four horizons of flows and associated tuff, and examination of the small
quarry to the east of Lochinvar reveals that the second group of basic rocks is composite,
embracing at least two and perhaps three flows.

av

220 STRATIGRAPHY OF THE LOWER MARINE SERIES,

Almost all the flows are or have been vesicular, and many are now amygdaloidal.
The textural features, however, are of no use in linking flows stratigraphically.

The basalts of Lochinvar Village, Gosforth, Anambah and near Lochinvar Station
can be correlated satisfactorily amongst themselves. The basalts of the Cranky Corner
Basin are regarded as being on the same horizon as that in the village of Lochinvar
outcropping on the roadside. This view is based on the relations between the basic
rock and the sporangia cherts in the two places.

Difficulty arises in attempting to correlate the great quantity of basalt about
Paterson and Northern Tocal with the flows near Comerfords and a little to the north
thereof.

At Comerfords the main group of vesicular and massive basalt underlies the
Eurydesma bed, and this is well above the highest sporangia cherts. At Paterson the
bulk of the basalt is between the two main sporangia chert horizons, and this suggests
a partial correlation of the Paterson units with the second group of flows at Lochinvar.

In the neighbourhood of “Aberglasslyn” a thick mass of amygdaloidal basalt and
basic tuff appears to be stratigraphically above the Comerfords flows. This immediately
suggests some linking with the somewhat similar rocks which overlie the Hurydesma
beds at Mt. View—probably the highest flows (stratigraphically) in the Lower Hunter
River areas.

Tt is of interest to note that as one goes northward up the Hunter Valley one finds
the basalts becoming younger in general, although their petrological characters remain
fairly constant.

The Lower Marine basalts of Muswellbrook, to which reference has already been
made, are very similar petrologically to the Lochinvar Village type, but any direct
correlation between the two districts is impossible.

Speaking generally about these basic rocks, it would appear that for a considerable
portion of Permian time the Hunter region was underlain by an alkaline basic magma
from which lava was drawn off at successive stages, and that basic tuff was frequently
developed, but chiefly in association with the later flows.

SUMMARY OF THE STRATIGRAPHY.

From the foregoing details of the succession we arrive at the following summarized
statement of the sequence, giving maximum thicknesses for the various Stages:

Feet.
MATIC Y Ti OtAR eb aie) ete ites so mie ee ce ee tks bine Aye ctr aeons sale fege, eee 985
RUCHESrfOrd Stas aes while oh Meher Me chee dees DT eG vidverod oraehtrl) Tee greamsb ie orate bel an] ()
Allandale Stage iy BUA IA are elas Mah es San meNly eran PAA pty aarp aml Go bac ap seen) C)
Lochinvar Stage be aie BY ean lays ae as See, Tp loo ata ate eed ar AO)
Total .. 5,895

POSSIBLE UNDERGROUND EXTENSIONS OF THE LOWER MARINE SERIES WEST AND SOUTH
OF THE HUNTER VALLEY.

Considering the Lochinvar Stage and its position in the sequence of groups in the
geosyncline, I would emphasize the probability that a great development of basalt exists
underground between the Lower Hunter and the Muswellbrook area. This is based on
the following criteria:

(a) The increase of Lower Marine basalt in regions west of the meridian of the
Lochinvar—Gosforth zone.

(bv) The abundance of volcanic rock and the absence of true sediment in the
Lower Marine Sections at Muswellbrook.

(c) The evidence of basalt ascending stratigraphically as one goes westerly from
the Lochinvar—Cessnock line.

In view of the heavy sedimentation implied in the Rutherford Stage of the country
between Old Rothbury, Pokolbin and Mt. View to Milfield, the writer is inclined to the
view that in a prolongation of the axis of the Lochinvar Dome and particularly along
the trend of the Pokolbin section of the Dome, we may expect to find fairly extensive

BY G. D. OSBORNE. 22

development of that Stage to the $.S.W. The Allandale Stage is marked by much basic
rock, but one does not expect any considerable development of the sediments of the
Allandale horizons, in the country south and west of the Lochinvar Anticline.

In the general problem of development of the Lower Marine beyond the Hunter,
the significance of the Cranky Corner Basin must be appreciated. In that sequence
we find a condensed section, showing a continuity of stratigraphical development
(qualitatively, although not quantitatively) over a large section of the Permian trough
of sedimentation, because the Cranky Corner Basin lies to the west of the strike of
the main axis of sagging in the trough. Thus, traced westward from the region lying
north of Pokolbin, the Lower Marine Series may be expected to thin out so as to give
very much reduced thicknesses for at least the Lochinvar, Allandale and Rutherford
Stages.

The most important underground extension of the Rutherford Stage would be to
the south.

Coming now to the Farley Stage, we note considerable development of strata on the
extreme west of the Hunter Province. It would appear that this upper arenaceous part
of the Lower Marine succession would be the most likely of all units to show a
sustained extension of lithological type underground to the west.

In conclusion, it appears that the typical Lower Marine section beneath the country
to the west and south-west of the Lochinvar Dome would be basements of basalt with
little Lochinvar Stage material, followed by Rutherford Stage in fair development in
the south, and less well developed in the north, overlain by a considerable thickness
of the arenaceous Farley Beds.

List oF FOSSILS OF THE LOWER MARINE SERIES.

In the foregoing text reference is made in many places to certain fossil-types which
are characteristic of well-known horizons, and some remarks upon the general nature
of the faunal content of the various groups of strata are given. In this section lists
are provided for purposes of general reference.

These lists are for the most part based upon the palaeontological data in Walkom’s
papers (1912, (a) and (0)), to which are added names of fossils more recently described
or recorded. The Foraminifera, Bryozoa and Ostracoda are those described in recent
years by Crespin and Crockford (see references).

It is hoped that the grouping of fossils into lists for the respective Stages may
assist future workers. The data are not absolutely exhaustive as the numerous records
of Permian fossils in N.S.W. Mines Department Reports have not been revised in the
light of modern palaeontological knowledge and classification.

THE LOCHINVAR STAGE.

(a) The Lower Portion in the Gosforth District.

Crinoid stems and ossicles, Polypora internata, Fenestrellina* fossula, F. sp., Seminula sp.
nov,, Martiniopsis radiata, Spirifera aff. tasmaniensis, Deltopecten linaeformis, Aviculopecten
englehardti, A. tenwicollis, A. mitchelli, Chaenomya sp., Conocardium sp. nov., Moeonia sp. nov.,
Orthoceras sp.

In the plant-bearing sandstone, Spirifera of the striati group, Dielasma, Conularia,
Platyschisma.

(b) The Main Part of the Lochinvar Stage.

Tribrachiocrinus sp., indeterminate crinoid, crinoid stems, Fenestrellina (?) internata,
F. (?) fossula, Stenopora tasmaniensis, Spirifer duwodecimcostata, 8S. stokesi, S. avicula, 8S.
vespertilio, S. tasmaniensis, Martiniopsis subradiata var. cf. morrisii, Productus cora var.
farleyensis, Strophalosia jukesi. Chonetes (?) sp., Hdmondia (?) noblissima, Chaenomya sp.,
Merismopteria sp. nov., Aviculopecten sprenti, A. tenwicollis, A. englehardti, Deltopecten subquin-
quelineatus, D. farleyensis, Moenia sp., Pleurophorus, Notomya (?), Pachydomus, Mourlonia
rotundatum, Ptycomphalina triflata, P. nuda, Platyschisma, Keenia, Conularia levigata.

* Dr. Ida Brown informed me that she had recently heard from Mrs. Beattie (Joan
Crockford) that Dr. M. K. Elias intends to use Fenestella as a valid name (as used for 112
years). An application for suspension of Rules of Zoological Nomenclature for the generic name
Fenestella Lons. 1839 was submitted to the International Commission on Zoological Nomen-
elature by G. E. Condra and M. K. Elias (Jowr. Palaeont. 15, 1941, 565-566).

222 STRATIGRAPHY OF THE LOWER MARINE SERIES.

The Allandale Stage.

Hyperamminoides sp. cf. proteus, Hyperamminoides acicula sp. nov., Ammodiscus multi-
cinctus, Ammobaculites woolnoughi sp. nov., Calcitornella stephensi, Frondicularia woodwardi,
Geinitzina triangularis, Crinoid stems, Fenestrellina fossula, F. dispersa, Polypora pertinaz,
Dyscritella restus, D. porosa, Stenopora. spiculata, S. etheridgei, S. johnstoni, Dielasma hastata,
D. sacculus, Martiniopsis subradiata var. cf. morristi, Spirifer vespertilio, S. stokesi, S. tasmani-
ensis, S. clarkei, Solenopsis sp., Chaenomya etheridgei, C. sp., Allorisma curvatum, Aviculopecten
tenuicollis, A. squamuliferus, A. mitchelli, A. sprenti, Deltopecten illawarrensis, D. fittoni,
D. linaeformis, Hurydesma cordata, Aphanaia sp., Modiola crassissima, Pleurophorus, Orthonota
sp., Notomya sp., N. cuneata, Pachydomus antiquatus, P. laevis, P. ovalis, Orthonycia altum,
Platyceras n. sp., Hdmondia (?) noblissima, Palaearca subarguta, Merismopteria macroptera,
M. n. sp., Avicula intumescens, Ptycomphalina triflata, P. morrissiana (?) Keenia platyschis-
moides, Platyschisma oculus, P. depressa, Conularia inornata, C. laevigata, Huomphalus, Bairdia
nyei, Cavellina kulnurensis.

The Rutherford Stage.

Foraminifera, Stenopora tasmaniensis, Fenestella, Protoretepora, Polypora, Spirifer
duodecimcostata, S. tasmaniensis, Martiniopsis subradiata, Productus sp., Aviculopecten
mitchelli, A. tenuicollis, A. sprenti, A. squamuliferus, Deltopecten farleyensis, Moeonia carinata,
Pachydomus, Ptycomphalina (?), Platyschisma, HEuomphalus (?).

The Farley Stage.

(a) The Fauna of the Ravensfield Sandstone.

Lasiocladia, Palaeaster clarkei, P. stutchburri, P. giganteus, Fenestella fossula, Dielasma
cymboeformis, D. biundata, D.* sp., Spirifer tasmaniensis, S. duodecimcostata, Cyrtina,
Martiniopsis subradiata, Solenopsis, Carimorphia (?), Chaenomya mitchelli, C. etheridgei,
Hdmondia (?) noblissima, Aviculopecten squamulferous, A. profundus, A. tenwicollis, A. sprenti,
A. mitchelli, Deltopcten linaeformis, D. subquinquelineatus, D. farleyensis, D. fittoni, Hurydesma
cordata, Moeonia carinata, Pleurophorus, Pachydomus, Astartila corpulenta, Platycerus altum,
Platyschisma, Ptycomphalina triflata, Conularia inornata, Hyolithes lanceolatus, Agathiceras
micromphalus, Orthoceras.

(b) The Fauna of the Upper Part of the Farley Stage.

Textularia eximia, Hyperamminoides sp. cf. proteus, Ammobaculites woolnoughi, Dielasma
sacculus, D. cymbaeformis, D. biundata, D. amygdala, D. inversa, Spirifer duodecimcostata,
S. stokesi, S. tasmaniensis, Martiniopsis subradiata, var. morrisii, var. konincki, Productus cora,
var. farleyensis, P. fragilis, Rhynchonella, Chonetes, Cardiomorpha gryphoides, Edmondia (?)
noblissima, Aviculopecten squamuliferus, A. tenwicollis, A. sprenti, A. englehardti, Aphanaia sp.,
Mytilus bigsbyi, Modiolopsis, Moeonia, Pleurophorus sp., P. gregarius, Stutchburia farleyensis,
Pachydomus, Platyschisma oculus, P. rotundatum, Conularia inornata, Goniatites micromphalus,
Nuculana waterhousei.

Acknowledgements.

For permission to publish some stratigraphical information in this paper, which resulted
from a geological assignment with Oil Search Ltd., Sydney, the writer expresses his thanks.
He also gratefully acknowledges financial help from the Commonwealth Research Grant
administered by the University of Sydney.

References.

BROWNE, W. R., and WALKom, A. B., 1911.—The Geology of the Eruptive and Associated Rocks
of Pokolbin, N.S.W. Jour. Roy. Soc. N.S.W., 45: 379.
BROWNE, W. R., and Dun, W. S., 1924.—The Stratigraphy of the Basal Portion of the Permo-
Carboniferous System in the Hunter River District. Jowr. Roy. Soc. N.S.W., 58: 198.
Browne, W. R., 1926.—The Geology of the Gosforth District, N.S.W., Part I. Jour. Roy. Soc.
N.S.W., 60: 213-277.
Browne, W. R., and WuHitr, H. P—The Hypersthene Andesite of Blair Duguid, near Allandale,
N.S.W. Jour. Roy. Soc. N.S.W., 60: 372.
CHAPMAN, F., and HowcuHin, W., 1905.—A Monograph on the Foraminifera of the Permo-
Carboniferous Limestones of N.S.W. Mem. Geol. Surv. N.S.W., Pal. 14.
CRESPIN, I., and Parr, W. J., 1941.—Arenaceous Foraminifera from the Permian Rocks of
N.S.W. Jour. Roy. Soc. N.S.W., 74: 300.
CrEsPIN, I., 1945.—Some Permian Foraminifera from Eastern Australia. Proc. Roy. Soe. Q’ld,
56: 23-30.
—, 1945.—Permian Ostracoda from Eastern Australia. Proc. Roy. Soc. Q’ld, 56: 31-36.
CrocKForD, J., 1940.—Permian Bryozoa of Eastern Australia, Part 1. Jour. Roy. Soc. N.S.W.,
TES BET
, 1940.—Permian Bryozoa of Hastern Australia, Part 2. Jour. Roy. Soc. N.S.W., 74: 502.
, 1942.—Permian Bryozoa of Eastern Australia, Part 3. Jour. Roy. Soc. N.S.W., 76: 258.

————., 1945.—Stenoporoids from the Permian of N.S.W. and Tasmania. Proc. Linn. Soc.
N.S.W., 70 (1-2): 924.

223

Davip, T. W. E., 1907.—The Geology of the Hunter River Coal Measures, N.S.W. Mem. Geol.
Surv. N.S.W., Geol. 4.

De Koninck, L. G., 1898.—Description of the Palaeozoic Fossils of N.S.W. Mem. Geol. Surv.
N.S.W., Pal. 6.

RaGeatt, H. G., 1929.—Report on the Singleton-Muswellbrook Coalfield. Ann. Rept. Dept. Mines,
N.S.W., pp. 100-104.

and FuprcHer, H. O., 1937.—A Contribution to the Permian-Upper Carboniferous

Problems. Rec. Austr. Mus., 20: 150.

Watkom, A. B., 1912.—Geology of the Permo-Carboniferous System in the Glendonbrook
District. Proc. Linn. Soc. N.S.W., 38: 115-145.

—, 1912.—Stratigraphical Geology of the Permo-Carboniferous System in the Maitland-

Branxton District. Proc. LINN. Soc. N.S.W., 38: 146-159.

THE COTYPES OF FORDONIA PAPUENSIS MACLHBAY.

By ArtHur LoveripGe, Musum of Comparative Zoology, Cambridge, Mass.
(Communicated by S. J. Copland.)

[Read 26th October, 1949.]

Recently, like Boulenger (1896, Cat. Snakes Brit. Mus., 3, p. 23), I suggested (1948,
Bull. Mus. Comp. Zool., 101, p. 388) that the brown water snakes described by Macleay
(1877, Proc. Linn. Soc. N.S.W., II, p. 35) might be synonymous with Fordonia leucobalia
(Schlegel) and should be re-examined. Through the courtesy of J. Henry, Esq., Curator
of the Macleay Museum, and the kindness of Mr. S. J. Copland of Sydney, I have been
enabled to do this.

Macleay designated no type, merely referring to “several specimens’. However, the
character he stresses—exclusion of the upper labials from the orbit—occurs only in the
solitary male (M.M., 1466), while the scale counts and measurements furnished by
Macleay (for one snake only) coincide most nearly with the male which should, there-
fore, be regarded as the type or lectotype. The only other character cited by Macleay
as distinguishing his papuensis was the number of midbody scale rows which he gave
as 22, actually there are 25 in all four snakes! Apparently no grounds remain for
regarding papuensis as even subspecifically distinct from leucobalia when we take into
account all available data of this widely distributed species.

The Macleay series (M.M., 1463-6) consists of a male and three females having
preocular 1; postoculars 2; upper labials 5, the third entering the orbit (M.M., 1463-4),
or the second and third (M.M., 1465), or labials entirely excluded from the orbit by an
extension of the postocular (M.M., 1466); lower labials 7, first three in contact with
the anterior sublinguals; midbody scale rows 25; ventrals 146-152; anals 2; subcaudals
27-33 (the highest number being present in both sexes) pairs, or a few (one to six)
single.

A NOTE ON EXPERIMENTAL CROSSING OF AEDES (STEGOMYIA) SCUTELLARIS
SCUTELLARIS WALKER AND AEDES (STEGOMYIA) SCUTELLARIS
KATHERINENSIS WOODHILL (DIPTHRA, CULICIDAE).

By A. R. Woopnitt, Department of Zoology, University of Sydney.
(One Text-figure.)

[Read 28th September, 1949.]

INTRODUCTION.

The “scutellaris group” of mosquitoes has been recorded from a wide area including
parts of Polynesia, Melanesia, the Philippines, North Australia, the Netherlands East
Indies and the Andaman Islands. Numerous varieties and subspecies have been
described by earlier workers and considerable confusion existed, but in recent years our
knowledge of the group has been greatly enhanced by the work of Farner and Bohart
(1945), Stone and Farner (1945) and Stone (1947). These authors have straightened
out the previous confusion in nomenclature and have described twelve forms, giving
each of these full specific status. In addition, a form from North Australia has been
described by the author (Woodhill, 1949) as a subspecies of A. scutellaris Walker. Some
forms of this group have also been recorded (as A. variegata) from Celebes (Bonne—
Wepster and Brug, 1932), Sumatra (Brug, 1931) and the Andaman Islands (Barraud,
1927)), but much more knowledge is required of this western portion of its range before
any separation into species or subspecies can be made.

It will be seen from Text-figure 1 (based on the work of the authors mentioned
above) that the ‘scutellaris group” presents a very interesting problem in speciation.
The differences used to separate the species consist of small variations in the markings
of the thorax, legs and proboscis and in slight variations in the structure of the basal

ae palate | ean ae

poullis/ |
Saipan
Guam hensillt

Marshall _| Is |

Mor shaslens1s

pLseuooscutclaris

{ * \ A ts
° Christmas | rs : c eee ;
a Society Is.
Sy} (S) }! ela calce

andrews! F sculéloris kohverinensis scutelaris
; scufelaris

iS . AUSTRALIA

Hii ian

DISTRIBUTION of Aedes scutellaris Walk. and closely related species.

a

Forrescens ongae

OW DAVIES

Text-figure 1.

BY A. R. WOODHILL, 225

lobes of the male genitalia. It has not been found possible to separate the larvae of any
* of the species, with the exception of A. horressens. The breeding habits and general
ecology of all the species are similar, as far as is known, throughout the whole range.
Farner and Bohart conclude from the differences in the genitalia and from the fact that
two distinct forms may occur in the same area that they should be considered distinct
species. However, further study may possibly reveal intermediate forms, and consider-
able light should be shed on the problem by attempts at crossing the different forms.
It was with this in view that laboratory crossings were made by the author between
A. scutellaris scutellaris Walker (= A. scutellaris Walker of Stone) and the newly
discovered A. scutellaris katherinensis Woodhill.

EXPERIMENTAL PROCEDURE AND. RESULTS.

Laboratory cultures of scutellaris from Lae, New Guinea, and of katherinensis from
Katherine, Northern Territory of Australia, were set up in small cages 12 inches by
10 inches by 10 inches, supplied with raisins and given a blood feed twice weekly. The
' temperature was maintained at 80°F. and the humidity at 70% to 80%. Eggs were
collected on filter papers immersed in water and the larvae were fed on pulverized dog
biscuit. Under these conditions, abundant eggs were obtained and continuous cultures
maintained without difficulty. Crossing was carried out by setting up pupae in individual
tubes and sexing the adults before liberation in cages. There was thus no possibility
of females being fertilized before liberation in the appropriate cages. Crossing was
carried out by means of bulk lots of 40 to 100 males and females. Crossing of individual
males and females was found to be very difficult and was abandoned. The results are
shown in Table I.

TABLE 1.
Results of Crossing A. scutellaris scutellaris and A. scutellaris katherinensis.

|
| Numbers of Sexes |
Number | and Subspecies. Approximate | Number | Percentage
of Number | Period of Egg of Eggs | Eggs
Experiment. | of Eggs | Production. | Hatched. | Hatched.
; scutellaris. katherinensis. | Produced | |
1 | 6063 4999 | 1970 | 28/8/48 to 19/4/48 | 0 0
2 3799 50gd 1490 26/3/48 to 19/4/48 | 1490 | 100
3 555d 999° | 2160 20/8/48 to 4/9/48 | 0 | 0
4 6229 56g dg 1900 21/8/48 to 15/9/48 | 1900 100
5 = | 64 virgin 99. 340 7/5/48 to 27/5/48 0 0
| |

It will be seen from Table 1 that @ scutellaris x ¢ katherinensis gave normal
numbers of fertile eggs. These eggs were subsequently bred through to the adult stage,
and perfectly normal Fl, F2 and F3 generations of adults were obtained. The recip-
rocal cross, however, proved completely sterile, no development of the embryo taking
place within the egg. In these sterile crosses copulation was frequently observed, and
ten females selected from Experiment No. 3 on 4th September, 1948, when dissected,
showed living spermatozoa in the spermathecae. It will also be noted that small
numbers of sterile eggs were deposited by virgin females of katherinensis, and that
mating provided an obvious stimulus to egg production even when the eggs were
sterile. While there are numerous examples in the literature of sterility or partial
sterility in Fl adult hybrids, this complete sterility of Fl eggs in one cross while the
reciprocal cross is completely fertile down to the F3 generation, appears to be unusual.
The only similar record is that of a cross between A. aegypti and A. albopictus, made
by Downs and Baker (1949). These authors state that albopictus males crossed with
aégypti females gave fertile progeny to the F2 generation, but that the reciprocal cross
was unsuccessful. In this unsuccessful cross twenty-four females were dissected, but

226 EXPERIMENTAL CROSSING OF AEDES (STEGOMYIA) SCUTELLARIS SCUTELLARIS WALKER.

only one showed spermatozoa, although copulation had been observed, and only a few
eges were deposited.

Examination of the chromosomes in the testes of scutellaris and katherinensis has
not so far shed any light on the problem. The only morphological difference between
the two subspecies is a distinct line of white scales on the anterior surface of the mid-
femur, this white line being present in katherinensis and absent in scutellaris. The F1
adults of the fertile cross all show an intermediate condition, i.e., a few scattered white
scales, while in the F2 adults the character referred to above is either present, inter-
mediate or absent. In one series of F2 adults the line of white scales was present in
85 individuals, showed an intermediate condition in 181, and was absent in 99, suggesting
a 1:2:1 ratio.

CONCLUSIONS.

It would appear from the above experiments that if scutellaris and katherinensis
were present in the same area they would interbreed and produce a population showing
all degrees of intergradation between the two forms, since, as far as is known, they
occupy similar ecological niches, unless some unknown biological factor prevented them
from mating under field conditions. Nevertheless, some differentiation must have taken
place, as evidenced by the sterile reciprocal cross, and it is therefore concluded that
one is justified in regarding these two forms as subspecies. It is hoped to carry out
further studies to determine whether there are any inherent differences in the two
subspecies in relation to temperature and humidity.

References.

Barraup, P. J., 1927.—Revision of the Culicine Mosquitoes of India, Part 23. Ind. Journ. Med.
RES.) lib 6b3e:

Bouwart, R. M., and INGRAM, R. L., 1946.—‘‘Mosquitoes of Okinawa and Islands in the Central
Pacific.” Dept. of the Navy, Bureau of Medicine and Surgery, Washington, D.C.

BONNE WEPSTER, J., and Bruce, S. L., 1932.—The Subgenus Stegomyia in Netherland India.
Geneesk. Tijds. Ned. Indie. Bd. 2: 39-119.

Brue, 8. L., 1931.—Arch. f. Hydrobiol. Suppl. Bd. ix: 1.

Downs, W. G., and Baxer, R. H., 1949.—Experiments in Crossing Aédes (Stegomyia) aegypti
Linnaeus and Aédes (Stegomyia) albopictus Skuse. Science, 109, No. 2826, 200-201.

FARNER, D. S., and Bouart, R. M., 1945.—A Preliminary Revision of the Scutellaris Group.
U.S. Naval Medical Bull., 44 (1): 21-37.

STONE, A., 1947.—A Topotypic Male of Aédes scutellaris Walker. Proc. Ent. Soc. Washington,
ae) (33) 3 Se

STONE, A., and FARNeER, D. S., 1945.—Further Notes on the Aédes scutellaris Group. Proc. Biol.
Soc. Washington, 58: 155-162.

WooDHILL, A. R., 1949.—A New Subspecies of Aédes (Stegomyia) scutellaris Walker (Diptera-
Culicidae) from Northern Australia. Proc. LINN. Soc. N.S.W., Ixxiv, 140-144.

227

A DETAILED STUDY OF THE FIELD DISTRIBUTION OF STRAINS OF
CLOVER NODULE BACTERIA.

By Hinary F. PurcuaAser and J. M. Vincent, School of Agriculture,
University of Sydney.

(One Text-figure.)

[Read 26th October, 1949.]

INTRODUCTION.

Although the difference in nitrogen fixing ability of different rhizobial strains is
well known, and this property shows considerable specificity in relation to the nature
of the testing host, little information appears to be available on the uniformity or
otherwise of strains from within a smaller or larger area in the field.

An earlier detailed study (Hughes and Vincent, 1942), using antigenic properties as
a guide to strain identity, has shown that isolations from the same species on the one
farm can yield many distinct strains and that different nodules on the same plant are
often inhabited by distinct serological types.

The present paper reports the results of similar and more extended studies, chiefly
using the pattern of effectiveness with four testing clovers as criterion of strain identity.
At the same time, serological data have been obtained for a large part of the isolated
strains. Attention has also been given to possible relationships between the nature of
the field host and behaviour with testing host, and the extent to which grouping within
the four testing hosts appears to be valid. ;

For the present purpose, involving the simultaneous testing of a large number of
cultures, an agar tube method is convenient and permits adequate differentiation between
effective and ineffective strains.

EXPERIMENTAL.
Source of cultures.
Isolates were obtained in the usual way from clover plants collected according to
the following schedule:

Series. Locality. Field Host. Year
4) Tichborne,’?) 4 sub-localities and paired large and T.G. 1947
small nodules.
QQ) Parkes to Coonabarabran, 5 sub-localities and paired T.G. 1947
large and small nodules.
3 Tichborne®) sub-locality 1. T.G. 1948
4 Tichborne,'?) single additional sub-locality. T.S.(a) T.R.(a)
T.G.(b) T.T.(b) 1948

() Serological data available.

(2) At ** Woodbine’, the property of Mr. C. Watson.

(a) From plants sown as sterilised seed.

(b) From plants sown as sterilised seed and from naturally occurring plants.

T.G.=Trifolium glomeratum L.; T.S.=Trifolium subterraneum L.; T.T.=Trifolium tomentosum L.
Trifolium repens L.

GB) ME pe

It will be observed that in series 1-3 the field host has been restricted to the one
species. The soil at Tichborne is classified as red brown earth, and is relatively acid
(pH approx. 5). Along the Parkes—Coonabarabran route soils are mainly red brown
earths, with occasional patches of darker soils of higher alkalinity (e.g., soil from
sublocality 5 was of this type). No detailed soil surveys have been made of this area.

Testing method.
Commercial seed of four clover species (Trifolium glomeratum, T. subterraneum,
T. pratense L., and T. repens), sterilized with 1/500 mercuric chloride, germinated

U

228 FIELD DISTRIBUTION OF STRAINS OF CLOVER NODULE BACTERIA,

aseptically after the fifth washing and inoculated with the appropriate rhizobial strain,
was planted at the top of an agar medium having the following composition:
CaHPO, 1 gm.; K,HPO, 0:2 gm.; MgSO,.7H.O 0:2 gm.; NaCl 0-2 gm.; FeCl, 0-1
gm.; agar 8 gms.; water 1 litre. pH was adjusted to 6:5 with sodium hydroxide. In
some experiments a nitrate control tube was included, containing 0:05% KNO,.

Complete bacteriological control was aimed at throughout and only one out of eighty
tubes used as uninoculated controls in many different experiments was found to be
contaminated by rhizobium: mould and actinomycete development was consistently
observed in a small number of cases and some tubes had to be discarded because of bad
infection. In later experiments previously tested effective and ineffective strains were
included for comparison.

A series of isolates was tested at the one time, provision being made for the analysis
of block effect on replication and the position effect within the tube in relation to light
source. Interaction terms were examined by comparison with error variance associated
with the individual plant, using the F test. Estimates of effectiveness were based on
the mean of single plant wet weights, known to be highly correlated with nitrogen
content and much simpler to determine, transformed to logarithms to allow greater
homogeneity of errors.

Criterion of effectiveness.

An improvement of 1-5 times the mean growth of control plants was selected as the
mean level of effective strains. To equalize the chances of wrongly classing a mean
either way, averages showing an improvement at least 0-5 x log 1:5 + log control mean
were classed as effective (E), the others being regarded as not different from the control
and therefore, ineffective (I). The probability of the Type I (and Type II) errors is
determined from the t value (one tail) using the interaction term (blocks x organisms)
variance to determine the standard error of the mean deviation.

‘

Classification of strains.

In the effectiveness tests these are classed according to the combinations of gradings
assigned to each testing host. Serological groupings are arrived at in the usual way
(Hughes and Vincent, 1942) according to the detailed flagellar and somatic reactions. In
the present case more antisera were available than in the earlier paper.

RESULTS.
GENERAL REVIEW OF EFFECTIVENESS TESTS.

Log weight mean values for each isolate have been grouped in classes and set out
in Text-figure 1 to provide a histogram pattern for each host tested with each series of
strains.

Certain features are common to all hosts:

(1) Uninoculated controls for each host are fairly consistent in different experi-
ments, except for subterranean clover in series 1, where larger sized seeds were used.

(2) Of the two check strains tested in series 1, 3 and 4, the ineffective strain 4/12S
has been consistent in its behaviour; strain 3/2L has been always effective, but has
varied to some extent in its more detailed performance between experiments.

(3) Plants provided with nitrate have also varied in the growth attained. With
the small seeded varieties, it is likely that insufficient nitrate was provided to supply
an excess of nitrogen during the whole of the experiment.

Probability of the Type I (and Type II) errors (approximate values tabulated
below) varies considerably for the different hosts, but is fairly consistent between
experiments:

Host. Series.
1 2 3 4
Trifolium glomeratum ve thd >25% 20% WAS 30%
T. subterraneum.. Bie ne 5% 1:5% 12% 7%
T. pratense .. ae ate a 12% 15% YM WIM

T. repens +. ae +2 ais 22% AYE 27% 2%

BY HILARY F. PURCHASE AND J. M. VINCENT. 229

In general, the variances are least for subterranean clover and highest for ball and
white clovers, and this is reflected in the magnitude of the errors which are directly
related to the variances.

GROUPING OF STRAINS ACCORDING TO EFFECTIVENESS PATTERNS.
A summary of effectiveness patterns observed in the localities investigated is provided
in Table 1.

Parallel reactions on test plants.

Statistical tests have shown that there is a better than chance parallelism of
behaviour between responses on the paired hosts, ball and subterranean clover; red and
white clover, despite the 75% disagreement observed between reactions of the second
pair in the 1948, Tichborne sublocality 1 isolates. Concurrence between the two groups
is no better than chance. It is interesting to observe the high proportion of cultures
effective on the four hosts. (Strain A, Table 1.)

TABLE 1.
Frequencies of Strains from Various Localities Grouped According to Effectiveness Behaviour on Four Hosts.
Tichborne.
Parkes-Coonabarabran.
a Testing Host. Sub-locality. Percentage
= General
5 Sub-locality. P-C. | Total.
1947. 1948. | 1947 | Tich- Total. Tich-
Te Gra Ee Se Dae sky. Total. | borne borne.| P.C.
To tals |e 2 ee ee
HN BY ol Beit a ah th &
PAG OB ea Hee stearate de (Ol Wude (ne 160 27 94 GN Bay 2 11 105 49 26
B/E 1h) |] TB) y] Ge fata eb ee | pe 11 21 32 1 i it 8 7 39 17 17
C TH 4) Jd) aa E i} Bi} 8 21 | 8 7 36 iL] B 83). al 8 44 19 19
1D) Jd) fp aby | a I 4/3 il 8 9 |) ey) Bog 15 24 5 36
IB} |] 1D) | dL E E Wy) 1 1 4 by 5 3 0
a 1d ae I E 1 1 1 J <1 0
(By i) ADs eal 1p} |} IL 1 1 2 2 2 1 0
jel} 13 |} I I I il | 2 2; 5 5 5 3 0
i I E E | E 1 1 1 i <i 0
J I E | I E 1 1 1 1 <il 0
Kea Aa: Bee | Ti 0 0 0 0 0
1b) fal 13) |) JE I 1 1 ft 1 <i 0
MI I E E 1 i 1 1 1 2 <1 c<il
ING |e I Daa 0 0 0 (0) 0
@) 1 I I E 0 0 0) 0 0)
Py I I if I iL if a 1 2 3} | 3 2 (0)
if
Totals ee 191 42 233
i}

E=effective reaction. [=ineffective reaction.

Results for a single block are quoted for Tichborne 1947 series tested on 7’. glomeratum. A marked difference
in reaction was shown in the second block (where the testing conditions were somewhat different from all other tests)
which would give a high proportion of ineffective on ball clover, a result not found elsewhere.

Testing hosts: Trifoilum glomeratum=T.G.; T. subterraneum=T.8.; T. pratense=T.P.; 7. repens=T.R.

Comparison of major localities.

On the 1947 results, Tichborne shows a significantly greater dispersion of strains
over the effectiveness patterns; the Parkes—Coonabarabran series, on the other hand,
is practically restricted to patterns A—D, D being represented more than at Tichborne.
However, the results for Tichborne in sublocality 1/1948, and the same sublocality in
1947, are markedly different and throw doubt on the validity of attributing the observed

behaviour merely to localities.

UU

230 FIELD DISTRIBUTION OF STRAINS OF CLOVER NODULE BACTERIA,

Comparison of sublocalities in 1947.

It is not possible, on the basis of the full effectiveness patterns, to detect a sublocality
effect, but if the comparison is restricted to reaction on white clover it will be seen that
a significantly high proportion of ineffective strains was isolated from Tichborne sub-
locality 4 and Parkes—Coonabarabran sublocalities 3, 4 and 5.

Paired plant isolates.

Isolates with the same effectiveness patterns were treated as identical for the
comparison of pairs from each plant examined. It has been found that the incidence
of identical pairs (11/38 for Tichborne, and 7/20 for Parkes—Coonabarabran) was no
greater than would be expected by chance, having regard to the distribution of effective-
ness patterns in each series.

There is thus no evidence that invasion of a plant by one strain piaces any restriction

on its invasion by a second strain.

Isolates from large and small nodules.

No more than random variation was observed in the incidence of effectiveness
groups for isolates from large and small nodules. However it was noted from the wet
weight yields in the 1947 Tichborne series that isolates from smaller nodules gave a
significantly lower return on the homologous host than those from large nodules.
Amongst the Parkes—Coonabarabran isolates, the reverse situation was encountered
against white clover. However, those differences have not a very high order of
significance and it would be difficult to generalize from them.

Strains collected by different field hosts.

On the basis of effectiveness patterns, isolates from T. subterraneum and T. repens
(all from sterilized seed), 7. tomentosum and T. glomeratum (including plants from
sterilized seed and naturally growing specimens) showed ho significant differences
attributable to field host (Table 2).

TABLE 2. .
Frequency of Different Effectiveness Patterns among Isolates from Four Fieid Hosts.

Effectiveness Pattern with
Testing Host. Field Host.
s< Strain:*: Total.
T.G. T.S. T.P. | TR. a aR |) eae, | ae,
A E E En leek 14 5 21 20 60
B E E E I 4 2 2 3 rl
C E E I E 4 4 8
D E E I I | 1 1
E E I E E | 1 1
P I I I I 1 1
|
Total ee ek ARS CAMB orale oy 20a lesa arated Sua i MCA ED at Wane

T.T.=Trifolium tomentosum. (Other abbreviations as given in Table 1.)

SEROLOGICAL TESTS.

Isolates from Tichborne 1947 and Parkes—Coonabarabran were tested for flagellar

and somatic agglutination, against six antisera of known constitution. Nineteen different
serological types were distinguished (Table 8).

BY HILARY F. PURCHASE AND J. M. VINCENT. 231

TABLE 3.
Type Reactions with Six Antisera, Based on Flagellar and Somatic Agglutination, and their Frequency among 126 Isolates.

Reaction with Serum.
Serological
Index. Frequency.
36 46 61 94 157 204
Flagellar type a H = H H — H 59
Somatic type a, ar = = = = = 11
” ” Qe re a a ee egy ae 13
on as ve = _ 4e = = = 6
” ” Us = ey at + core re 11
” ” as Say an sai ats far rr 2
oF) ” as ar = cad = = ar 10
” ” a, =n Ge = = = Fa 6
Flagellar type b he -_ H — — H _ 45
Somatic type b. eS - + = — — — i
9 ” bs -- = = + _— — ih: 5
2 ” ba as a + ar = = 20
” ” bs = a a al = = 4
. m9 (Og Bt _ — _ — — _ 3
a Os ae - + ao - — — 6
Flagellar type c .. sé = = = — - — 19
Somatic type c, i = + = = — — 2
9 ” C3 = rs <F a ea oy 3
” ” C7 oe iy <a a ag mat 13
9 mp Oh Me = =F ar a or = 1
Flagellar type d a0 h h h h h h 1
Somatic type d, te _ + = — = — 1
Flagellar type e .. Bic H = H h H h 2
Somatic type e .. a0 = +r = = = — 2
H, h, — indicate strong, weak and no flagellar reactions, respectively.
+, — indicate presence or absence of somatic reactions.

Comparison of major localities. ;
The distribution of some major serological types differed between the Tichborne and
Parkes—Coonabarabran series (Table 4, cols. 6 and 7).

Comparison between sublocalities.
No heterogeneity could. be demonstrated amongst the sublocalities of Parkes—
Coonabarabran, but was denfiite in the Tichborne sublocalities (Table 4, cols. 2-5).

Paired plant isolates.

In only one out of the four sublocalities of the Tichborne series was a greater than
chance number of similar serological pairs encountered (5 observed as against 1-7
expected). No significant pairing was encountered among the Parkes—Coonabarabran
isolates.

Isolates from large and small nodules.

There was a highly significant correlation between the size of the nodule and
incidence of serological strains (Table 5), but before any general significance can be
attached to this finding, it seems reasonable to correct for the effect of having taken sets
of isolates from restricted areas where all similar strains might easily have arisen fairly
recently from the one parent. For this reason figures of Table 5 are adjusted so that a
particular strain is represented once only for a particular sublocality. There is now no
better than a chance distribution of antigenic types in large and small nodules although
it needs to be noted that the small number of cases available for the analysis makes
significance difficult to demonstrate.

232 FIELD DISTRIBUTION OF STRAINS OF CLOVER NODULE BACTERIA,

TABLE 4.
Distribution of the Most Frequent Serological Types in Tichborne Sublocalities and Parkes-Coonabarabran Major Locality.
Tichborne Sublocality. |
Serological | Tichborne Parkes- General
Type. Total. Coonabarabran. Total.
il 2 3 4 |
ay 6 2 1 — 9 2 11
As — — — 8 8 5 13
a4 — — 2 1 3 8 11
OG ae ct — 1 4 2 7 3 10
a residua ae — 1 2 6 9 5 14
Motal aww tae 6 4 9 17 36 23 59
Dba sce <a — 9 1 — 10 10 20
b residual 9 5 6 = 20 5 25
MOWED Ge 9 14 7 — 30 15 45
ait |
i}
Cr, aes ae — — a 6 13 — 13
e residua il 1 1 1 4 | 2 6
Total ¢ ay 1 1 8 rd 17 2 19
d — 1 _ — 1 2 3
General total 16 20 24 24 84 42 126
TABLE 5.
Proportion of the Most Frequent Serological Types Found in Large and Small Nodules.
Gross Figures. With Sub-locality Correction.
Serological
Index.
Large Small Large Small
Nodules. Nodules. : Nodules. Nodules.
ay on a ays 9 (5:9) 7) (2 1))} 5 (8-7) i, (283)
Qy ae a se 12 (6-9) L_ (6:1) 5 (32%) (23)
Qs ae By Mie 8 (5:9) 3 (=) By (41983) v3 {(P4017/)
As a Sa rae (55983) 3 (47) 5 (4:9) & (eal)
b, As Ud ag 7 (10-6) 13 (9-4) 4 (5-5) & (Bob)
Cy ae ve ae 1 (6:9) 25 (G)-ah) 1 (1:8) 2 (22)
Others af ne 23 (25-5) 25 (22-5) 8 (9:1) 7 (5:9)
Totals ae 67 (67-0) 59 (59-0) 33 (33-0) 21 (21-0)

Figures in brackets represent expected values, calculated from cross totals.

Relationships between serological and effectiveness reactions.

Table 6 shows that serological characters might appear to be partially correlated
with effectiveness patterns, but when account is taken of a possible locality effect even
this imperfect correlation is reduced to an insignificant degree,

BY HILARY F. PURCHASE AND J. M. VINCENT. 233

No correlation could be established between flagellar or somatic reactions separately
and effectiveness patterns, when a correction for locality was applied in a similar
manner.

TABLE 6.
Distribution of the Strains from Major Serological Groups in the Major Effectiveness Patterns.
Gross Figures. With Sub-locality Corrections.
Effectiveness Patterns. Effectiveness Patterns.
Serological
Index. |
A B Cc D Others. A B C D Others.
|
|
ay. Tf (Bed © (Mes WW Glosral Coil 83 alevpy cb ame a) (akeran@), (@louly ak (Glo) sy (Glo)
Qs. 1 (4:0)| 8 (3-0)| 0 (1:°6)| 2 (2-4)) 2 (2-0)) 1 (2-4)) 4 (1-8)) O (1-2)| 2 (1-8)] 2 (1-8)
Qs. (0) (Goel “ob aL Glee) 7 (cabal (Glory (cab) 4s Glory) al (alcal)yiac® (alo) al (ater)
Oe OSD) A 123) 20 ae 2) 45 (9) 3" (5) 0 (216) 1 (221) 2) (23)) 4 (2-0)) 3° (2:0)
Over 14 (@oP))) BY (o@)] PA (Heed) 2 (Soro) 1k (Sioab) (45) (P4583) PAs (ako) mak (Aaa) ab (oR)
O56. iN Sen} 0) (87) 3) (27) "8" 5) 2) (222) 45 (-9) |) OF (26) 2), (22 1)|| 2) (1-3), 2) (20) "4 (220)
Others .. .. {18 (14-2)]11 (10-4)| 7 (5-7)| 5 (8-6)| 5 (7-1)]14 (9:8)| 8 (7:3)| 5 (4:6)} 4 (6-9)| 4 (6-9)
Totals .. |88 (38-0)/28 (28-0)/15 (15-0)/23 (23-0)/19 (19-0)|24 (24-0)/19 (19-0)/12 (12-0))18 (18-0))18 (18-0)

Figures in brackets represent expected values.

DISCUSSION.

There is considerable evidence that most of the rhizobia isolated from clover in
these wheat soils are capable of fixing nitrogen with 7. glomeratum, which is the most
common clover found growing naturally. The prevalence of rhizobia effective on the
common natural host may have some relation to selective influences associated with it,
but this is by no means certain.

To some extent the observations of Strong (1937), on a degree of similarity of
effectiveness behaviour on red and white clover, and dis-similarity on subterranean
clover, has been confirmed (with ball clover being placed in the latter group, in accord-
ance with the findings of Vincent, 1945), but the groups are by no means mutually
exclusive, as has already been pointed out by Jensen and Vincent, 1941, and Vincent,
1945.

Relationships between effectiveness patterns and serological grouping are at best only
partial, and are no more than would be expected in studying members of a localized field
population probably having a fairly recent common origin. With similar serological
types encountered in widespread localities, it can be considered that they are sufficiently
removed in time from a common origin to have afforded greater opportunity for change
in their behaviour on the host plant.

A change in the field population with time has been observed with regard to
classification of strains based on (a) effectiveness patterns (Tichborne, sublocality 1,
1947, compared with 1948) and (b) serological types (1947 Tichborne data compared
with those found earlier by Hughes and Vincent, 1942).

The occurrence of different serological types on the same plant was observed by
Hughes and Vincent (1942); it is now assured that in general strains with different
serological characters or different effectiveness patterns can be obtained from the one
plant. Where identity between serological characters from two isolates from the one
plant was observed in one locality, it probably indicates no more than that localization
may have occurred within the small area (possibly in the rhizosphere). There is
definitely no suggestion that the plant has any specific discriminating effect.

Difficulty was experienced in isolating strains with any particular effectiveness
characteristics from larger or smaller nodules. This is probably due to the complications
introduced by having nodules of different ages (cf. Nutman, 1946).

234

— tor

O71 Nv 3W

2)

LIM

LHSISM

FIELD DISTRIBUTION

NUMBER

OF STRAINS OF CLOVER NODULE BACTERIA,

OF STRAINS

Text-figure 1.

BY HILARY F. PURCHASE AND J. M. VINCENT. 235

The final picture that emerges from these investigations is that even within the
immediate environs of the single plant there is a diverse rhizobial population charac-
terized in the present instance by a general ability to fix nitrogen in association with
the common clover of that area. Isolates from one or two nodules cannot give any real
idea of the nature of the existing rhizobial population at any one time, much less its
constitution on a second occasion. It would seem reasonable to expect that subterranean
clover would be effectively nodulated if sown in an established ball clover area, but in
the introduction of another clover species—such as red or white—into such an area,
seed inoculation with a strain of known performance on these hosts (preferably also
effective on ball and subterranean clovers) would be indicated. In such a case the
ability of the introduced strain to establish and maintain itself in competition with local
types would be of some importance even though initially the mass effect of inoculation
would probably ensure nodulation predominantly by the introduced strain. Further
studies in competition between strains are planned.

SUMMARY.

Effectiveness patterns on four clover hosts and serological classification have been
used to compare a large number of isolates from clovers growing in representative New
South Wales wheat soils. Isolates were collected in such a way as to permit comparisons
on the basis of locality, sublocality, the individual plant and on the basis of nodule size
on the original field host.

Differences in effectiveness behaviour were observed between isolates from Tichborne
and those from Parkes—Coonabarabran in 1947, and between cultures from one of the
Tichborne sublocalities in two successive years. Serological types were heterogeneously
distributed in different localities. No distinction could be made in effectiveness behaviour
between ball clover isolates and isolates from other clover species sown in the Tichborne
area.

A significant concurrence of effectiveness was noted between tests on ball and
subterranean clovers, and white and red clovers using all the clover strains of Rhizobium,
but agreement within each of these host groups was by no means complete.

No evidence was obtained of any restriction on the invasion of a plant by strains
having different effectiveness patterns, but in one sublocality more identical serological
types were obtained from paired nodules than could be expected by chance.

Isolates from small nodules showed somewhat lower ability to fix nitrogen on the
homologous host than strains from large nodules, but results were inconsistent and little
significance could be attributed to them. Certain serological types were found confined
mainly to a particular nodule size but this observation could be mainly attributed to the
localization of such strains to areas where they could have arisen from a single parent
rhizobium.

No correlation between effectiveness patterns and serological behaviour could be
observed above that attributable to the collection of isolates from small areas where
certain rhizobial strains, probably of recent common parentage, were predominating.

The rhizobial population even within a restricted area is likely to be diverse and a
few samples can give only a poor idea of its nature. The practice of seed inoculation
may often be advisable to ensure nodulation with a suitable strain, but data on the
strain’s competitive ability would also be desirable.

Text-figure 1.
Frequencies of strains producing responses of host plants falling within specified mean
logarithmic weights limits, for each series of four host species.

Abbreviations: T.G., Trifolium glomeratum; T.S., T. subterraneum; T.P., T. pratense ;
T.R., T. repens.

1, 2, 3, 4, series numbers for sets of strains.

I, ineffective strain, 4/12S; E, effective strain, 3/2L; C, uninoculated control (two tubes
per series) ; N, 0:05% KNO,.

Broken line represents the division between effective and ineffective responses.

236 FIELD DISTRIBUTION OF STRAINS OF CLOVER NODULE BACTERIA,

ACKNOWLEDGEMENTS.

Thanks are due to Dr. D. B. Duncan for advice in the handling of statistical data. The
authors also wish to thank Miss J. Logan and Mr. W. Hurry for technical assistance throughout
the course of the work, Dr. I. A. Watson and Mr. C. Watson for their help in the collection of
plants, and Mr. B. L. Jones for carrying out a large part of the serological tests. This work
has been assisted by a grant from the Rural Bank of New South Wales.

References.

HueGuHeEs, D. Q., and VINCENT, J. M., 1942.—Serological Studies of the Root Nodule Bacteria. iii.
Tests of Neighbouring Strains of the Same Species. Proc. LINN. Soc. N.S.W., Ixvii, 142-152.

JENSEN, H. L., and VINCENT, J. M., 1941.—Specificity between Host Plant and Strain of
Rhizobium trifolti. Austr. J. Sci., 3: 169.

NutMAN, 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.

Strone, T. H., 1937.—The Influence of Host Plant Species in Relation to the Effectiveness of
the Rhizobium of Clovers. J. Coun. Sci. Indus. Res., 10: 12-16.

VINCENT, J. M., 1945.—Host Specificity amongst Root Nodule Bacteria Isolated from Several
Clover Species. J. Austr. Inst. Agric. Sci., 11: 121-127.

xxiii

ABSTRACT OF PROCEEDINGS.

ORDINARY MONTHLY MEETING.
30th Marcu, 1949.

Dr. R. N. Robertson, President, in the Chair.
Library accessions amounting to 46 Volumes, 344 Parts or Numbers, 14 Bulletins,
3 Reports and 11 Pamphlets, total 418, had been received since the last meeting.

PAPERS READ.

1. The Distribution of Formic and Alcohol Dehydrogenases in the Higher Plants,
with particular Reference to their Variation in the Pea Plant during its Life-cycle. By
Daphne C. Davison.

2. The Importance of Formic Dehydrogenase in the Oxidation Mechanisms of
Pisum sativum. By Daphne C. Davison.

3. Revision of the Genus Brachycome Cass. Part II. New Zealand Species. By
Gwenda L. Davis.

4. Australian Formicidae. New Genera and Species. By J. J. McAreavey, S.J.

5. On Australian Species of Creophilus (Coleoptera: Staphylinidae). By W. O.
Steel. (Communicated by J. W. T. Armstrong.)

6. On Australian Dermestidae. Part V. Notes and the Description of Four New
Species. By J. W. T. Armstrong.

ORDINARY MONTHLY MEETING.
27th Aprit, 1949.

Dr. R. N. Robertson, President, occupied the Chair.

The President announced that the Council had elected Dr. A. B. Walkom to be
Honorary Treasurer for the Session 1949-50.

The President also announced that the Council had elected Dr. Ida A. Brown, Mr.
A. R. Woodhill, Dr. G. D. Osborne and Dr. Lilian Fraser to be Vice-Presidents for the
Session 1949-50.

Mr. L. B. Burton Browne, Rose Bay; Mrs. EH. M. Campbell, Hunter’s Hill; Mr. J. H.
Elliott, Neutral Bay; Professor P. D. F. Murray, Sydney; Mr. J. K. Pollak, Glenfield;
Mr. M. D. Wilson, Lindfield; and Mrs. Anne Zeck, Ryde, were elected Ordinary Members
of the Society.

Library accessions amounting to 14 Volumes, 86 Parts or Numbers, 18 Bulletins,
4 Reports, and 6 Pamphlets, total 128, had been received since the last, meeting.

PAPERS READ.

1. The genus Dawsonia. By Alan Burges.
2. Graptolites from Tallong and the Shoalhaven Gorge, New South Wales. By
Kathleen Sherrard, M.Sc.
EXHIBITS.

Mrs. P. Messmer exhibited a number of cushion plants, collected during a recent
visit to Cradle Mountain, Tasmania. These included: Abrotanella forsterioides Hook.,
Dracophyllum minimum F.v.M. (this species is indistinguishable from Abrotanella when
not in flower and very closely resembles Donatia), Pterygopappus Lawrencii Hook.,
Phyllachne Colensoi Berggr. These are distinguishable from the mat-forming plants
such as Raoulia Meredithae F.v.M. (= Hwartia Meredithae F.v.M. (Beau.) ), Pentachronda
pumila, by the hard interior of the cushions, exclusion of light and air, and content of

XXiv ABSTRACT OF PROCEEDINGS.

packing materials, and are characterized by their hard exterior as adaptation to climate
and capacity for holding water as a temperature control for the protection of the under-
lying root system.

The Secretary exhibited lava from the new (February, 1949) flow at Mt. Ngauruhoe,
New Zealand.

ORDINARY MONTHLY MEETING.
25th May, 1949.

Dr. R. N. Robertson, President, occupied the Chair.

Mr. Kenneth Myers, West Kogarah, was elected an Ordinary Member of the Society.

The President announced that Miss Judith Balmain and Miss Adele Millerd,
Linnean Macleay Fellows in Biochemistry, had recently obtained the degree of M.Sc.
at the University of Sydney.

The President also announced that the Rev. H. M. R. Rupp had been awarded the
Royal Society of New South Wales’ Clarke Memorial Medal for 1949 for his distinguished
contributions to knowledge of Australian Orchidaceae, and that Professor W. L.
Waterhouse had been awarded the Medal of the Royal Society of New South Wales and
the Medal of the Federal Council of the Australian Institute of Agriculture for his
research on cereals and other aspects of agricultural science.

The President stated that authors of papers could obtain their original drawings
one month after publication of the paper but that the manuscripts would be retained
by the Society for three years.

Library accessions amounting to 3 Volumes, 84 Parts or Numbers, 2 Bulletins, 2
Reports, 29 Pamphlets, total 120, had been received since the last meeting.

PAPERS READ.

1. Studies in Australian Marine Algae. V. Studies on and Geographical Records
of Various Species, particularly those of the Gelidium Complex. By Valerie May.

2. Notes on the Morphology and Biology of a New Species of Tabanus (Diptera,
Tabanidae). By Kathleen M. I. English.

3. Revision of the Genus Brachycome Cass. Part III. Description of Three New
Australian Species and some New Locality Records. By Gwenda L. Davis.

4. Notes on Microspore Types in Tasmanian Permian Coals. By J. A. and Roma
Dulhunty.

NOTES AND EXHIBITS.

Mr. A. Musgrave exhibited a collection of insects recently secured by him in the
vicinity of Hobart, Tasmania, and gave a brief account of the history of entomology in
Tasmania.

Miss M. Hindmarsh demonstrated the difference between normal cell division and
division in a cell treated with colchicine, which prevents spindle formation. Upsetting
cell division in this way increases chromosome numbers and produces polyploid cells.

The June and July Monthly Meetings were not held because of lighting restrictions
caused by the Coal Miners’ Strike.

ORDINARY MONTHLY MEBRTING.
31st Aueust, 1949.

Dr. R. N. Robertson, President, occupied the Chair.

The following were elected Members of the Society: Messrs. Eldred P. Baker,
University of Sydney; John H. Burden, Chatswood; Miss Olga Kooptzoff, Paddington;
Professor Charles EH. Marshall, B.Sec., Ph.D., D.Se., F.G.S., M.I.Min.E., University of
Sydney; Miss Isabel J. McPhail, B.Sc., Armidale, N.S.W.; Messrs. Allen H. Miller,
Punchbowl; Philip C. Minter, Strathfield; Miss Hilary F. Purchase, B.Sc.Agr., University
of Sydney; Mrs. Dorothy A. Thorp, B.Sc., Kangaroo Point; Miss Betty M. B. Turnock,
B.Sc., Armidale, N.S.W.; Mr. Thomas G. Vallance, Sutherland.

ABSTRACT OF PROCEEDINGS. XXV

The President congratulated Dr. J. L. Still on his appointment to the Chair of
Biochemistry in the University of Sydney.

A subcommittee for Phenological Observations was formed, its members being
Messrs. A. Musgrave, HE. H. Zeck, Miss Muriel Morris, and Dr. H. S. McKee.

Library accessions, amounting to 18 Volumes, 293 Parts or Numbers, 2 Bulletins,
11 Reports and 27 Pamphlets, total 351, had been received since the last meeting.

PAPERS READ.

1. The Hair-tracts in Marsupials. Part III. Description of Species concluded. By
W. Boardman.

2. Crania in the Macleay Museum. By N. W. G. Macintosh.

3. Australian Rust Studies. VII. Some Recent Observations on Wheat Stem
Rust in Australia. By I. A. Watson and W. L. Waterhouse.

4. A New Subspecies of Aédes (Stegomyia) scutellaris Walker (Diptera, Culicidae)
from Northern Australia. By A. R. Woodhill.

NOTES AND EXHIBITS.

Miss Muriel Morris described the flotation mechanisms of certain planktonic

organisms with illustrations from samples of plankton collected from the Hawkesbury
Estuary.

ORDINARY MONTHLY MEETING.
28th SEPTEMBER, 1949.

Dr. R. N. Robertson, President, occupied the Chair.

The following were elected Members of the Society: Messrs. Phillip B. Carne,
Canberra, A.C.T.; Justus E. Humpoletz, B.Sc., Roseville; Harold F. Lower, Adelaide,
S. Australia; Neville C. Stevens, Hurstville; Mrs. Lucy M. Willings, B.A., Cronulla;
Mr. Edward J. F. Wood, Cronulla.

The President announced that the Council is prepared to receive applications for
Linnean Macleay Fellowships tenable for one year from 1st January, 1950, from qualified
eandidates. Applications should be lodged with the Secretary not later than Wednesday,
2nd November, 1949.

Library accessions amounting to 13 Volumes, 59 Parts or Numbers, and 5 Bulletins,
total 77, had been received since the last meeting.

PAPERS READ.

1. The Stratigraphy of the Lower Marine Series of the Permian System in the
Hunter River Valley, N.S.W. By G. D. Osborne.

2. A Note on Experimental Crossing of Aédes (Stegomyia) scutellaris scutellaris
Walker and Aédes (Stegomyia) scutellaris katherinensis Woodhill (Diptera, Culicidae).
By A. R. Woodhill.

Talk by Dr. N. W. G. Macintosh on “Crania in the Macleay Museum”’.

ORDINARY MONTHLY MEETING.
26th OctToBEr, 1949.

Dr. R. N. Robertson, President, occupied the Chair.

Miss J. W. Newman was elected a member of the Society.

The President referred to the death of Mr. W. J. Enright, who had been a member
of the Society since 1914.

The President announced that the Council is prepared to receive applications for
Linnean Macleay Fellowships tenable for one year from 1st January, 1950, from qualified
candidates. Applications should be lodged with the Secretary not later than Wednesday,
2nd November, 1949.

Library accessions amounting to 19 Volumes, 106 Parts or Numbers, 1 Bulletin, 6
Reports, and 2 Pamphlets, total 134, had been received since the last meeting.

Vv

XXVi ABSTRACT OF PROCEEDINGS.

PAPERS READ.

1. The Cotypes of Fordonia papuensis Macleay. By A. Loveridge. (Communicated
by S. J. Copland.)

2. A Detailed Study of the Field Distribution of Strains of Clover Nodule Bacteria.
By Hilary F. Purchase and J. M. Vincent.

NOTES AND EXHIBITS.

Methods used in the Study of Plant Enzymes. By Miss A. Miliera.

The principles and use of the Thunberg and manometric techniques were described
and their application to the investigation of the succinoxidase system of the potate
tuber (Solanum tuberosum L.) discusseé.

Hifect of Vitamins on Fungi. By Mr. I. M. Fraser.

Mr. I. M. Fraser briefly discussed the existence of vitamin deficiencies in the fungi,
exhibiting as an example of this a graph showing the effects of small amounts of biotin
on the growth of Neurospora sitophila Shear and Dodge. In work on the cultivated
mushroom, Psalliota hortensis (Cooke) Lange forma albida Lange, he has shown that
in addition to possessing a deficiency for thiamin, its growth is very markedly increased
under certain conditions by the native growth hormone of higher plants, indole —3—-
acetic acid. A graph showing the effect of a series of concentrations of indole —3— acetic
acid was exhibited.

ORDINARY MONTHLY MEETING.
30th NovemMBeErR, 1949.

Dr. R. N. Robertson, President, in the Chair.

Mrs. Joy L. Whaite, Panania, N.S.W., was elected an Ordinary Member of the
Society.

The President announced that Miss Muriel Morris, Miss Mary Hindmarsh, and Miss
Adele Millerd had been reappointed to Linnean Macleay Fellowships for 1950.

The President also announced that the Society had been asked to advise the Fauna
Protection Panel on choosing suitable areas for dedication as faunal reserves.
Suggestions from members would be welcome.

Library accessions amounting to 17 Volumes, 201 Parts or Numbers, 21 Bulletins,
10 Reports and 1 Pamphlet, total 250, had been received since the last meeting.

The meeting then took the form of a joint Symposium with the Society for Experi-
mental Biology of New South Wales on: “The Surface of the Bacterial Cell.”

Dr. R. J. Swaby discussed the nature of superficial structures on bacterial cells, as
determined by their appearance under the light microscope after differential staining,
and under the electron microscope before and after metal shadowing. This presented
the morphological evidence for believing that flagella, capsules, cell walls and the Gram
positive layer exist.

Miss P. M. Rountree took the cell surface of the Pneumococcus as a model of a
typical bacterial cell. Information obtained by the bacteriologist, chemist, physical
chemist and geneticist was correlated and discussed. It was concluded that, apart from
its capsule, the chief antigenic component of the cell surface is a polysaccharide-lipid-
protein complex. This is an expression of the biological specificity of the cell and is
genetically determined.

Before opening the subject for general discussion, the Chairman, Dr. R. N. Robertson,
referred to the physical properties of the surface, including its ionic properties and its
passive permeability to substances in solution. The ability of the cell to bring about
the active transport of dissolved substances was mentioned and the general biological
significance of these properties was stressed.

A general discussion followed.

XXVil

LIST OF MEMBERS.
(15th December, 1949.)

ORDINARY MEMBERS.

Abbie, Professor Andrew Arthur, M.D., B.S., B.Se., Ph.D., c.o. University of Adelaide,
Adelaide, South Australia.

*Albert, Michel Francois, ‘““Boomerang’’, 42 Billyard Avenue, Elizabeth Bay, Sydney.

*Allman, Stuart Leo, B.Se.Agr., M.Sc., Entomological Branch, Department of Agriculture,
Farrer Place, Sydney.

Anderson, Miss Beverley I., 19 Kareela Road, Chatswood, N.S.W.

Anderson, Robert Henry, B.Se.Agr., Botanic Gardens, Sydney.

Armstrong, Jack Walter Trench, “Callubri’’, Nyngan, N.S.W.

Ashby, Professor Eric, D.Sc., D.I.C., F.L.S., Botany Department, The University,
Manchester, England.

Aurousseau, Marcel, B.Sc., c.o. Mr. G. H. Aurousseau, 16 Woodland Street, Balgowlah,
N.S.W.

Baddams, Miss Greta, B.A., B.Sc., New England University College, Armidale, N.S.W.

Balmain, Miss Judith Hope, M.Sc., School of Agriculture, Sydney University.

*Beadle, Noel Charles William, D.Sc., Botany School, Sydney University.

Bearup, Arthur Joseph, 66 Pacific Avenue, Penshurst, N.S.W.

Beattie, Mrs. Joan Marion, M.Se. (née Crockford), Bradley Street, Cobar, N.S.W.

Benson, Professor William Noel, B.A., D.Sce., 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.Sc., M.Se., Department. of Zoology, Sydney University.

Blake, Stanley Thatcher, M.Sc., Botanic Gardens, Brisbane, Queensland.

Boardman, William, M.Sc., University of Melbourne, Mildura Branch, Mildura, Victoria.

Bradhurst, Miss Peggy Joan, B.Sc., 25 Belgium Avenue, Roseville, N.S.W.

Brett, Robert Gordon Lindsay, B.Sec., 7 Petty Street, West Hobart, Tasmania.

Brough, Patrick, M.A., D.Sc., B.Sc.Agr., Botany School, Sydney University.

Brown, Miss Ida Alison, D.Se., Department of Geology, Sydney University.

Browne, Lindsay Blakeston Barton, 34 Kent Road, Rose Bay, N.S.W.

Browne, William Rowan, D.Sc., Department of Geology, Sydney University.

Browning, T. O., Waite Agricultural Research Institute, Adelaide, South Australia.

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.

Burgh, Henry Bertram, 42 Warialda Street, West Kogarah, N.S.W.

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, Council for Scientific and Industrial Research, Box 109,
Canberra, A.C.T.

Carey, Miss Gladys, M.Sc., 32 Rawson Street, Epping, N.S.W.

*Carey, Professor Samuel Warren, D.Sc., Geology Department, University of Tasmania,
Hobart, Tasmania.

Carne, Phillip Broughton, B.Agr.Se. (Melb.), Division of Entomology, C.S.I.R.O., Box 109,
P.O. City, Canberra, A.C.T.

Carne, Walter Mervyn, c.o. Department of Commerce and Agriculture, Reliance House,
Flinders Lane, Melbourne, Victoria.

Carroll, Miss Dorothy, B.A., B.Sc., Ph.D., D.I.C., Science House, 157 Gloucester Street,
Sydney.

*Chadwick, Clarence Earl, B.Sc., Entomological Branch, Department of Agriculture, Farrer
Place, Sydney.

Cheel, Edwin, 40 Queen Street, Ashfield, N.S.W.

Christian, Stanley Hinton, c.o. Department of Public Health, Port Moresby, Papua-
New Guinea.

Churchward, John Gordon, B.Sec.Agr., Ph.D., 1 Hunter Street, Woolwich, N.S.W.

Clark, Laurance Ross, M.Sc., c.o. Council for Scientific and Industrial Research, Box 109,
Canberra, A.C.T.

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.

Cooper, Roy Percy, F.F.C.A., 14 Third Avenue, Willoughby, N.S.W.

* Life Member.

XXViili LIST OF MEMBERS.

1942
1947
1908
1928
1946

1929
1945
1948
1936

1934
1925
1937
1927
1948

1937
1926
1946

1948
1941

1949
1943
1947
1930

1947
1948
1948
1930
1939

1935
1944

1936
1939

1928
abs)aby/
1947
1932
1947,
1930

1938
1943
1946
1930
1943

1932
1942

1917
1938

1947
1945
1907

1948

Copland, Stephen John, B.Se., 7 Creewood Street, North Strathfield, 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.
Craft, Frank Alfred, B.Se., 10 Bank Street, Wellington, N.S.W.

Crust, Miss Mabel, B.Sc., 21 Silex Road, Mosman, N.S.W.

Dakin, Professor William John, D.Sc., 9 Kuring-gai Avenue, Turramurra, N.S.W.

Davis, Mrs. Gwenda Louise, B.Sc., New England University College, Armidale, N.S.W.

Davison, Miss Daphne Claire, M.Sc., ‘‘Carinya’”’, Waratah Street, Palm Beach, N.S.W.

Day, Maxwell Frank, Ph.D., B.Sc., Council for Scientific and Industrial Research,
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.

Dulhunty, John Allan, D.Sc., Department of Geology, Sydney University.

Dumigan, Edward Jarrett, 10 High Street, Toowoomba, Queensland.

Durie, Peter Harold, B.Sc., Regional Pastoral Laboratory, C.S.I.R. Mail Bag, Armidale,
N.S.W.

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.

Elliott, John Henry, 8 Shellcove Road, Neutral Bay, N.S.W.

Ellison, Miss Dorothy Jean, M.Sc., Abbotsleigh College, Wahroonga, N.S.W.

Endean, Robert, 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., 14 Milray Avenue, Wollstonecraft, N.S.W.

Fraser, Miss Lilian Ross, D.Se., ‘“‘Hopetoun”’, 25 Bellamy Street, Pennant Hills, N.S.W.
Frohlich, Mrs. Frances Marie Veda, D.Sc., 40 Smith Street, Summer Hill, N.S.W.

*Garretty, Michael Duhan, M.Sc., 477 St. Kilda Road, Melbourne, S.C. 2, Victoria.

Greenwood, William Frederick Neville, c.o. Colonial Sugar Refining Co. Ltd., Lauto} :,
Fiji.

Griffiths, Mervyn Edward, M.Sc., Australian Institute of Anatomy, Canberra, A.C.T.

Gunther, Carl Ernest Mitchelmore, M.B., B.S., D.T.M., Bulolo, New Guinea.

Hamilton, Edgar Alexander, 16 Hercules Street, Chatswood, N.S.W.

Hardy, George Huddleston Hurlstone, “‘The Gardens’, Letitia Street, Katoomba, N.S.W.

Harker, Miss Janet Elspeth, B.Sc., New England University College, Armidale, N.S.W.

Harris, Miss Thistle Yolette,’°B.Sc., 14 Pacific Street, Watson’s Bay, N.S.W. i

Henderson, David Leonard Wylie, L.S.M.I.S., “Craigstone”, Woodford, 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.Se., Botany School, University of Sydney, N.S.W.

Holland, Victor Wallace, 6 Warrenball Avenue, Newtown, 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, Engr.Agr.S., Dr.Agr.Se. (Cracow, Poland), Waite Institute, Private
Mail Bag, Adelaide, S.A. ¢

Hossfeld, Paul Samuel, M.Sc., 132 Fisher Street, Fullarton, South Australia.

Humphrey, George Frederick, M.Sc., Department of Biochemistry, Sydney University.

Jacobs, Ernest Godfried, ‘‘Cambria’’, 106 Bland Street, Ashfield, N.S.W.

Jacobs, Maxwell Ralph, D.Ing., M.Sc., Dip.For., Commonwealth Forestry Bureau, Canberra,
INK OMAN,

Johnson, Lawrence Alexander Sidney, 178 Beecroft Road, Cheltenham, N.S.W.

Johnston, Arthur Nelson, B.Sce.Agr., Hawkesbury Agricultural College, Richmond, N.S.W.

Johnston, Professor Thomas Harvey, M.A., D.Sc., F.L.S., University of Adelaide, Adelaide,
South Australia.

Joklik, Wolfgang, 74 New South Head Road, Vaucluse, N.S.W.

* Life Member.

LIST OF MEMBERS. KOKO

Jones, Mrs. Valerie Margaret Beresford, M.Sc. (née May), Mooloolabel Esplanade,
Narrabeen, N.S.W.

Joplin, Miss Germaine Anne, B.Sc., Ph.D., ‘‘“Huyton’”’, 18 Wentworth Street, Eastwood,
N.S.W.

Jopling, Alan Victor, B.Sc., B.E., 28 Cliff Street, Manly, N.S.W.

Judge, Leslie Arthur, No. 1 Bridge Road, Hornsby, N.S.W.

Keast, James Allen, 313 West Botany Street, Rockdale, N.S.W.

Kesteven, Geoffrey Leighton, B.Sc., Special Commissioner’s Office, South East Asia, Cathay
Building, Singapore, Malaya.

Kesteven, Hereward Leighton, D.Sc., M.D., The Hospital, Cooktown, Queensland.

Kiely, Temple Baylis, 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.

Langford-Smith, Trevor, M.Sc., Ministry of Post-War Reconstruction, Canberra, A.C.T.

Larcombe, Miss Pauline Gladys, B.Se., 17 Ethel Street, Burwood, N.S.W.

Lascelles, Miss June, M.Sc., Department of Biochemistry, University of Oxford, Oxford,
England. ;

Lawrence, James Joscelyn, B.Sc., 91 Boundary Street, Clovelly, N.S.W.

Lawson, Albert Augustus, 9 Wilmot Street, Sydney.

Lee, Mrs. Alma Theodora, M.Sc. (née Melvaine), 16a Raglan Street, Mosman, N.S.W.

Lee, David Joseph, B.Sc., School of Public Health and Tropical Medicine, Sydney
University.

Liddell, Miss Jean Joyce, University of Melbourne, Mildura Branch, Mildura, Victoria.

Lothian, Thomas Robert Noel, Botanic Gardens, Adelaide, South Australia.

Lower, Harold Farnham, 7 Avenue Road, Highgate, Adelaide, South Australia.

Macintosh, Neil William George, M.B., B.S., Department of Anatomy, University of
Sydney, N.S.W.

Mackerras, David, 5 Edwards Bay Road, Mosman, N.S.W.

Mackerras, Ian Murray, M.B., Ch.M., B.Se., Queensland Institute of Medical Research,
Herston Road, Valley, Brisbane, Queensland.

*Mair, Herbert Knowles Charles, B.Se., 5 Collaroy Street, Collaroy Beach, N.S.W.

Manefield, Tom, Jr., 14 Maida Road, Epping, N.S.W.

Marks, Miss Elizabeth Nesta, M.Sc., Biology Department, University of Queensland,
Brisbane, Queensland.

Marshall, Professor Charles Edward, B.Se., Ph.D., D.Se., F.G.S., M.I.Min.E., Department
of Geology, University of Sydney, N.S.W.

Martin, Donald, B.Se., Stawell Avenue, Hobart, Tasmania.

Mawson, Sir Douglas, D.Sc., B.E., F.R.S., University of Adelaide, Adelaide, South
Australia.

Maze, Wilson Harold, M.Se., Sydney University.

McAreavey, John Joseph, S.J., St. Patrick’s College, Cathedral Place, E. Melbourne, C.2,
Victoria.

McCulloch, Robert Nicholson, B.Sc.Agr., B.Se., Roseworthy Agricultural College, Rose-
worthy, South Australia.

McKee, Hugh Shaw, B.A., D.Phil. (Oxon.), Council for Scientific and Industrial Research,
Private Bag, P.O., Homebush, N.S.W.

MeMillan, Bruce, 171 Lawson Street, Hamilton, Newcastle, N.S.W.

McPhail, Miss Isabel Jean, B.Se., New England University College, Armidale, N.S.W.

McRoberts, Miss Helen May, B.Sc., New England University College, Armidale, N.S.W.

Mercer, Frank Verdun, B.Se., St. John’s College, Cambridge University, Cambridge,
England.

Messmer, Mrs. Pearl Ray, 64 Treatts Road, Lindfield, N.S.W.

Middleton, Bertram Lindsay, B.A., M.D., Bridge House, Murrurundi, N.S.W.

Miller, Allen Horace, 1281 Canterbury Road, Punchbowl, N.S.W.

Miller, David, Ph.D., M.Sc., F.R.S.N.Z., F.R.E.S., Cawthron Institute, Nelson, New Zealand.

Millerd, Miss Alison Adele, M.Se., Department of Biochemistry, Sydney University.

Millett, Mervyn Richard Oke, B.A., ‘‘Urlingford’’, Cammeray Road, Cammeray, N.S.W.

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, B.Sc., Department of Zoology, Sydney University.

Moye, Daniel George, B.Sc., Dip.Ed., Warragamba Dam, via Wallacia, N.S.W.

Moye, Mrs. Joan, B.Sc. (née Johnston), Warragamba Dam, via Wallacia, 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.Sc., Department of Zoology,
University of Sydney, N.S.W.

* Life Member.

XXX LIST OF MEMBERS.

1920 Musgrave, Anthony, F.R.E.S., Australian Museum, College Street, Sydney.
1949 Myers, Kenneth, 1 Station Street, West Kogarah, N.S.W.

1925 Newman, Professor Ivor Vickery, M.Sc., Ph.D., F.R.M.S., F.L.8., Department of Botany,
University of Ceylon, Colombo 3, Ceylon.

1922 Nicholson, Alexander John, D.Sc., F.R.E.S., Council for Scientific and Industrial Research,
Box 109, Canberra, A.C.T.

1935 *Noble, Norman Scott, D.Se.Agr., M.Se., D.I.C., Council for Scientific and Industrial
Research, 314 Albert Street, Hast Melbourne, C.1, Victoria.

1920 Nobile, Robert Jackson, B.Sc.Agr., Ph.D., 324 Middle Harbour Road, Lindfield, N.S.W.

1912 North, David Sutherland, 42 Chelmsford Avenue, Lindfield, N.S.W.

1942 O’Brien, Brian Robert Alexander, Department of Anatomy, Sydney University.

1948 O’Farrell, Antony Frederick Louis, A.R.C.Se., B.Sc., F.R.E.S., New England University
College, Armidale, N.S.W.

1927 Oke, Charles George, 34 Bourke Street, Melbourne, C.1, Victoria.

1927 Osborn, Professor Theodore George Bentley, D.Sc., F.L.S., Professor of Botany, University
of Oxford, Oxford, England.

1921 Osborne, George Davenport, D.Sc., Ph.D., Department of Geology, Sydney University.

1947 Paramonov, Sergey Jacques, D.Sc., Division of Economic Hntomology, C.S.I.R., Box 109,
Canberra, A.C.T.

1940 Pasfield, Gordon, B.Sc.Agr., 20 Cooper Street, Strathfield, N.S.W.

1922 Perkins, Frederick Athol, B.Se.Agr., Biology Department, University of Queensland,
Brisbane, Queensland.

1947 Phillips, Miss Marie Elizabeth, M.Sc., 4 Morella Road, Clifton Gardens, N.S.W.

1937 Plomley, Kenneth Francis, 50 Domain Street, South Yarra, Melbourne, Victoria.

1948 Pollak, Mrs. Audrey, 33 Arundel Street, Glebe, N.S.W.

1949 Pollak, John Kurt, 33 Arundel Street, Glebe, N.S.W.

1935 Pope, Miss Hlizabeth Carington, M.Sc., Australian Museum, College Street, Sydney.

1918 Priestley, Professor Henry, M.D., Ch.M., B.Sc., 54 Fuller’s Bridge Road, Chatswood, N.S.W.

19388 Pryor, Lindsay Dixon, M.Sc., Dip.For., Parks and Gardens Section, Department of the
Interior, Canberra, A.C.T.

1949 Purchase, Miss Hilary Frances, B.Sc.Agr., School of Agriculture, University of Sydney,
N.S. W.

1929 Raggatt, Harold George, D.Sc., 485 Bourke Street, Melbourne, Victoria.

1947 Richards, Miss Barbara Vivian, B.Sc., 10 Small Street, Bondi Junction, N.S.W.

1946 Riek, Edgar Frederick, B.Sc., Council for Scientific and Industrial Research, Box 109,
Canberra, A.C.T.

1940 Robbins, Mrs. Elizabeth Marie, M.Sc. (née Basnett), 344 Railway Terrace, Guildford,
N.S.W.

1936 Roberts, Noel Lee, 43 Hannah Street, Beecroft, N.S.W.

1932 Robertson, Rutherford Ness, B.Sc., Ph.D., Food Preservation Research Laboratory,
C.S.1.R., Private Mail Bag, Homebush, N.S.W.

1245 Ross, Donald Ford, c.o. Ross Bros. Pty. Ltd., 545-547 Kent Street, Sydney.

1925 Roughley, Theodore Cleveland, B.Sc., F.R.Z.S., Chief Secretary’s Department, Box 304,
G.P.O., Sydney.

1932 Salter, Keith Eric Wellesley, B.Sec., ‘““Hawthorn’’, 48 Abbotsford Road, Homebush, N.S.W.

1919 *Scammell, George Vance, B.Sc., 7 David Street, Clifton Gardens, N.S.W.

1947 Scott, Miss Beryl, B.Se., Department of Geology, University of Tasmania, Hobart, Tas.

1928 Selby, Miss Doris Adeline, M.Sc., M.B., 11 Locksley Street, Killara, N.S.W.

1948 Shaw, Miss Dorothy Edith, B.Sc.Agr., 119 Bellevue Parade, Hurstville, N.S.W.

1930 Sherrard, Mrs. Kathleen Margaret, M.Sc., 43 Robertson Road, Centennial Park, Sydney.

1947 Shipp, Hrik, 59 William Edward Street, Longueville, N.S.W.

1948 Smith, Robert, B.Se. (Agric.), Institute of Agriculture, University of Western Australia,
Nedlands, Western Australia.

1916 Smith, Miss Vera Irwin, B.Sc., F.L.S., ‘Suoana’’, Mt. Morris Street, Woolwich, N.S.W.

1943 Smith-White, Spencer, B.Sc.Agr., 7 Merriwa Street, Gordon, N.S.W.

1945 Southcott, Ronald Vernon, M.B., B.S., 12 Avenue Road, Unley Park, Adelaide, South
Australia.

1937 Spencer, Mrs. Dora Margaret, M.Sc. (née Cumpston), 16 Attunga Street, Woollahra,
N.S.W.

1942 Spencer, Terence Edward, 16 Attunga Street, Woollahra, N.S.W.

1926 Stanley, George Arthur Vickers, B.Sc., c.o. Messrs. Robison, Maxwell and Allen, 19 Bligh
Street, Sydney.

1898 Stead, David G., “Boongarre’’, 14 Pacific Street, Watson’s Bay, N.S.W.

* Life Member.

1949
1935

1948
is )akal

1940

1949

1944
1943
1946
1921
19/49

LIST OF MEMBERS. XXxi

Stevens, Neville Cecil, 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., New England University College, Armidale, N.S.W.
*Sulman, Miss Florence, ‘““Burrangong”, McMahon’s Point, N.S.W.

Taylor, Keith Lind, B.Sc.Agr., Forestry Commission, Division of Wood Technology, 96
Harrington Street, Sydney.

Thorp, Mrs. Dorothy Aubourne, B.Sc. (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.1I.H.Aust., Box 2770, G.P.O., Sydney.

*Troughton, Ellis Le Geyt, C.M.Z.S., F.R.Z.S., Australian Museum, College Street, Sydney.

Turnock, Miss Betty May Blanche, B.Sc., New England University College, Armidale,
N.S.W.

1949
IG)aLY

1930
1940
1934

1909
1946
1930
1911

1936
1947
1928
1927
1941
1941
1946
1926
1946
1949
1949

1947
1947

1949
1934

1949
1932

1949
1936

1923

1902
1949

1942
1943

Vallance, Thomas George, 57 Auburn Street, Sutherland, N.S.W.

Veitch, Robert, B.Sc., F.R.E.S., Department of Agriculture and Stock, William Street,
Brisbane, Queensland.

Vickery, Miss Joyce Winifred, M.Sc., Botanic Gardens, Sydney.

Vincent, James Matthew, B.Sc.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., P.O. Box 127, Katanning, Western Australia.

Ward, Melbourne, Gallery of Natural History and Native Art, Medlow Bath, N.S.W.

Wardlaw, Henry Sloane Halcro, D.Se, F.A.C.I., Department of Physiology, Sydney
University.

Waterhouse, Douglas Frew, M.Sc., Council for Scientific and Industrial Research, Box
109, Canberra, A.C.T.

Waterhouse, John Teast, B.Se., Department of Botany, Sydney University.

Waterhouse, Lionel Lawry, B.E., ‘“Rarotonga’’, 42 Archer Street, Chatswood, N.S.W.

Waterhouse, Professor Walter Lawry, D.Sec.Agr., M.C., D.I.C., Faculty of Agriculture,
Sydney University.

Watson, Irvine Armstrong, Ph.D., B.Sc.Agr., Faculty of Agriculture, Sydney University.

Wearne, Walter Loutit, c.o. James Gilberg & Co., 143 Elizabeth Street, Sydney.

Wharton, Ronald Harry, B.Sc., Institute of Medical Research, Kuala Lumpur, Malaya.

*Whitley, Gilbert Percy, Australian Museum, College Street, Sydney.

Wilkins, Miss Marjorie Jessie, B.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.

Wilson, Ralph Dudingston, M.Se., M.Se.Agr., Biological Branch, Department of Agricul-
ture, Box 36A, G.P.O., Sydney.

Windridge, Mrs. Judith, B.Sc., Department of Geology, Sydney University.

Winkworth, Robert Ernest, Botany Department, University of Melbourne, Carlton, N.3,
Victoria. !

Wolf, Miss Ruth, M.Se., 19 Birriga Road, Bellevue Hill, N.S.W.

Womersley, Herbert, F.R.E.S., A.L.S., South Australian Museum, Adelaide, South
Australia. 5

Wood, Edward James Ferguson, C.S8.I.R.O., Marine Biological Laboratory, P.O. Box 21,
Cronulla, N.S.W.

Woodhill, Anthony Reeve, B.Sc.Agr., Department of Zoology, Sydney University.

Zeck, Mrs. Anne, 694 Victoria Road, Ryde, N.S.W.
Zeck, Emil Herman, 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.

Broom, Robert, M.D., D.Sc., F.R.S., Transvaal Museum, Pretoria, Transvaal, South Africa.

Jensen, Hans Laurits, D.Sc.Agr. (Copenhagen), State Laboratory of Plant Culture,
Department of Bacteriology, Lyngby, Denmark.

Rupp, Rev. Herman Montague Rucker, B.A., 24 Kameruka Road, Northbridge, N.S.W.

Waterhouse, Gustavus Athol, D.Sc., B.E., F.R.E.S., c.o. Mrs. Millett, Illoura Avenue,
Wahroonga, N.S.W.

* Life Member.

XXXii

LIST OF GENERA AND SPECIES DESCRIBED AS NEW IN THIS VOLUME.

(1949.)
Genera.
Page.
Schizopelta (Myrmicinae) .. .. .. .. 14
Stenothorax (Myrmicinae) .. .. .. .. 3
Species.
Page.
armstrongi (Anochetus) .. .. .. 1 marginatus (Dacxyon)

armstrongi (Dolichoderus) CEN a Ae niger (Monomorium)
armstrongi (Camptonotus) seein suet) nynganensis (Plagiolepis) ..
armstrongi (Monomorium) LETS LO, obovata (Brachycome)
arrowi (Anthrenocerus ) er He elip Me OG orarius (Tabanus)

DEuNea KEMElOPWOTAUS)i se on ee ee) petrophila (Brachycome)
capitalis (Xiphomyrmex) .. .. .. 6 rostratus (Diplograptus)
elesanisia@StigmnGcnos) sna ee eee: sanguinea (Camptonotus)
faleatai(SGniZoperta) uae ae ee all silvicolum (Trogoderma)

katerinae (Stenothorax) 3 stigmacrophilus (Anthrenocerus )
katherinensis (Aédes scutellaris) 141 stolonifera (Brachycome)
macqueeni (Neoanthrenus) 111 tricolor (Metapone)

List of Plates.
Proceedings, 1949.

i-ii—Graptolites from Tallong and Shoalhaven Gorge.
iii-iv.— Australian Rust Studies.

XXxili

GENERAL INDEX.

(1949.)
Page Page
Abstract of Proceedings .............. Saal. DOO OMEG, Wns (CMI 2b oocobboocnusccs 83
Aédes (Stegomyia) scutellaris Walker Dermestidae, Australian, Pt. V ........ 107

(Diptera, Culicidae), A New Sub-

species of, from Northern Australia 140
Armstrong, J. W. T., Australian Dermes-

LCA CH Mm ATs by Vet wetter h esiiee hensycey /oleteita ion
Australian Dermestidae. Part V ......
Australian Formicidae, New Genera and

SDEGCTCS iat tr t A se cte Ne iat aa 1
Australian Marine Algae, Studies on,

Pt. V. Observations on and Geo-

graphical Records of Various
Species, particularly those of the
Gelidiwm Complexe 8 oa. ee ose 196
Australian Rust Studies. VII. Some
Recent Observations on Wheat Stem
IR WISE iba) AQIS ao eats odo ubod 113
Australian Species of Creophilus (Cole-
optera: Staphylinidae) ............ 57
Bacterial Cell, Surface of; Symposium
CVA R SEs ees O ocead anal Lerers cedeuabe lee XXvVi
Baker, HE. P., elected a Member ....... XXiV
Balance Sheets for the Year ending 28th
ING eENA TNL) eee odooocee dos XX-Xxii
Balmain, Miss J., Linnean Macleay
Fellow in Biochemistry, M.Sc.
ESTE Ce iat nis suate eines terencuc ver naee econ XxiV
Boardman, W., The Hair-tracts in Mar-
supials, Pt. III. Description of
Species, concluded ................ 192
Brachycome Cass., Revision of the
GleMISR RET a tees sehr omy ea 97
Brachycome Cass., Revision of the
Genus seb Chol ae aye aie ven cies ean) a5 145

Brown, Ida A., elected a Vice-President, xxiii
Browne, L. B. Burton, elected a Member, xxiii

Burden, J. H., elected a Member ...... XXiV
Burges, A., The Genus Dawsonia ...... 83
Campbell, Mrs. E. M., elected a Member, xxiii
Carne, P. B., elected a Member ........ O07
Clover Nodule Bacteria, a Detailed
Study of the Field Distribution of
RSALELEPEIN TY CUS VISCO) GS es ca ae span AT Uae NP Ua ek 227
Council Members for 1949-50 .......... xix
Crania in the Macleay Museum ........ 161
Creophilus (Coleoptera: Staphylinidae),
Australian, Speciesvor 323445... .). 57

Davis, Gwenda L., Revision of the
Genus Brachycome Cass. Part II.
New Zealand Species .............. we

Davis, Gwenda L., Revision of the
Genus Brachycome Cass. Part III.
Description of Three New Aus-
tralian Species and some New
WocalityaRVecond Sayan hes sees ee

Davison, Daphne C., Distribution of
Formic and Alcohol Dehydrogenases
in the Higher Plants, ete. ......... 26

Davison, Daphne C., Formic Dehydro-
genase in the Oxidation Mechanisms
of Pisum sativum, Importance of .. 37

145

Dulhunty, J. A., and Dulhunty, Roma,
Notes on Microspore-types in Tas-
manianweermian: Coals, 3. 2.5% yc:

Effect of Vitamins on Fungi .......... Xxvi
HTC CET OMS rhe ee ties esr eRe aty iene larsed melee tps
Elliott, J. H., elected a Member ....... Xxili
English, Kathleen M. I., Notes on the
Morphology and Biology of a New
Species of Tabanus (Diptera,
TPA DANITUC LEAs te etek hae ste sine Mine

Enright, W. J., reference to death of ..
Experimental Crossing of Aédes (Steg-
omyia) scutellaris scutellaris Walker
and Aédes (Stegomyia) scutellaris

katherinensis Woodhill 224

Formic and Alcohol Dehydrogenases in
the Higher Plants, with particular
Reference to their Variation in the
Pea Plant during its Life-cycle .... 26

Formic Dehydrogenase in the Oxidation

‘Mechanisms of Pisum _ sativum,

Lm pontan COvOPiy Aiea aeehe Sen canes Be
Formicidae, Australian, New Genera and

NSH) OY eXOUC S11 ik es en gregory ite eR ae a ec 1
Fordonia papuensis Macleay, The Co-

LY DESO been ae esretertts) aenereteust shelielcroyehers 223
Fraser, I. M., Effect of Vitamins on

SUT tee tae ee is ait cds patel salssese alte Paes cae XXV1
Fraser, Lilian, A Gummosis Disease of

(QUEL ieke- cap sea oO gidid Game oincroen eto Vv

Fraser, Lilian, elected a Vice-President, xxiii

Graptolites from Tallong and the Shoal-
haven (GOLLE yes ies Soke eens 62

Gummosis Disease of Citrus, A ........ Vv

Hair-tracts in Marsupials. Pt. III.

Description of Species, concluded .. 192
Hindmarsh, Miss M., Demonstration of
normal and abnormal plant cell
GEVAISTONS ey es ce anal ast atene ee. XXiV
Humpoletz, J. E., elected a Member .... xxv
Kooptzoff, Miss O., elected a Member .. xxiv
IG ONE WAY EXCOEREIIOMIE S665 555k05000¢ xxiii-xxvi
Linnean Macleay Fellowships—
Alteration in conditions of award .... ii
Appointments! for 1949) ~.. 3s. 5 iii
Appointments for 1950 ............. XXvi
Loveridge, A., The Cotypes of Fordonia
DAUDWENSTS MAC Caves ita eae 223

Lower, H. F., elected a Member ........ XXV
Lower Marine Series, The Stratigraphy
of, of the Permian System in the
Hunter River Valley, N.S.W........ 203

XXXIV

Page.
McAreavey, J. J., Australian Formicidae 1
Macintosh, N. W. G., Crania in the
Maicleaya Museum aes crea sees eee
McPhail, Miss I., elected a Member ... xxiv
Marshall, Prof. C. E., elected a Member xxiv
May, Valerie, Studies on Marine Algae.
V. Observations on and Geographical
Records of Various Species, particu-
larly those of the Gelidium Complex 196

IMEMBETESS HEISET Ofer a Amine i teste t Mites en XXVili
Members, notes of interest of ........ iii, iv
Messmer, Mrs. P., Cushion Plants from
AD AUS TALIA a oa aS VON ES Se I fee Xxill
Meyer, K., elected a Member .......... XXiV
Microspore Types in Tasmanian Permian
COATS MG Lk EOP, CE CRU ETL Cee. as 132
Miller, A. H., elected a Member ........ XXiv
Millerd, Miss A., Linnean Macleay
Fellow in Biochemistry, M.Sc.
GeSTOG H GAG eh eae pan aeeone XXiV
Minter, P. C., elected a Member ...... XXiV
Morris, Miss M.—
Exhibit, flotation mechanisms in
olan tomas obo eis piyhe Geer ese XXV
Linnean Macleay Fellow ............ ii
Murraye 2 rot eaDy a pheanelectede a
FO OUCS aU aye ARI A are i ORIN WR A EE oC RAN Xxiii
Musgrave, A., exhibit of insects from
Aran ep aoaly a ieab Zeta a ie Nm cited eet XXiV
New Genera and Species, List of ..... XXXili

Newman, Miss J. W., elected a Member, xxv
Notes on the Morphology and Biology of
a New Species of Tabanus (Diptera,
MAWAMTCAES) rea tieeN acne Mrneatyeos cere
Obituaries—
E. C. Andrews
MES sD al Geb! risky onsen km mies Pisce Vv
Jee DaiMie Com sir.) is ee ea a ea Vv
Brae Nig IVC KiGite ci) ae Siaer ipa gluten ce POOR ea. Vv
R. E. Turner Vv

Original Drawings and Manuscripts,
TA HVUEN, WO) JAH MONE. 5 cS co gduccbunbe. XXiV
Osborne, G. D., The Stratigraphy of the
Lower Marine Series of the Permian
System in the Hunter River Valley,

2) e} sell eige eile te (el eliehielie) elie! (ehelielelieieliels

INS SS Wits comes SAU AAS AED oe dada. Ranh Ee Ba 203
Hlected a Vice-President ............ XXili
Phenological Observations, Subcom-
mMIttee: LOM... eet es aR XXV
Plates whist votes, uch. luego nen lek rut XXXiV
Pollak, J. K., elected a Member ...... XxXili
iPresidentialivAddressiss donee eee i
Purchase, Miss Hilary F., elected a

Memb CR tei. s ik as eee XXili
Purchase, Hilary F., and Vincent, J. M.,
A Detailed Study of the Field Dis-
tribution of Strains of Clover
NodwmleSsBacteniayae hn ee ee

Robertson, R. N., elected President for
1949-50

PARA OS Capt nic tenets Senne tan aca Xxix

Rupp, Rev. H. M. R., award of Clarke
MemonialiMedalige pea. oe. . XXiV

Rountree, Miss P. M., Contribution to
SVMDOSTUMA ee At Ne cen eee Tite XXVi1

Page.
Sherrard, Kathleen, Graptolites from
Tallong and the Shoalhaven Gorge,

INES Wie es aes Sess eo eg New tere ee a 62
Society for Experimental Biology, N.S.W.
VOoye WIGS nhS Wall 5o5¢5c0ce5c000c XXVi
Steel, W. O., Australian Species of
Creophilus (Coleoptera: Staphy-
Vind da @))). ee et eee cate Rei Paes ae 57
Stevens, N. C., elected a Member ...... XXV
Still, J. L., appointment to Chair of
Biochemistry os aa aioe oleae: XXV
Stratigraphy of the Lower Marine Series
of the Permian System in the
Hunter River Valley, N.S.W. ...... 203

Studies on Australian Marine Algae. YV.
Observations on and Geographical
Records of Various Species, particu-
larly those of the Gelidium Complex 196

Swaby, R. J., Contribution to Sym-
DOSIWIN: SERS sek eG ae Se eT be XXvVi

Symposium on “The Surface of the
Bacterial: (Celso. cers tere ey seas XXV1

Tabanus (Diptera, Tabanidae), Notes on
the Morphology and Biology of a

INEWriSDECIES *Of ee eile OS ry ha tae 153
Tasmanian Permian Coals, Microspore
ATV. DGS = TI SSE ete SEI a LDA ats a 132
Thorp, Mrs. D. A., elected a Member ... xxiv
Tindale, Miss M., Linnean Macleay
IMG LW O Ws oie ne Ro Dope tae. eed aes iii
Turnock, Miss B. M. B., elected a
MEGTIOD ET cae eee cece ee ae XXiV
Vallance, T. G., elected a Member ...... XXiv
Vincent, J. M., and Purchase, H. F., A
Detailed Study of the Field Distri-
bution of Strains of Clover Nodule
Bacteria. ie terial) Cee a dues 227
Vitamins, Effect of, on Fungi ......... Xxvi
Walkom, A. B., elected Honorary
TRTCASUT ETI COLES io oEE OS Seis ann XXili
Waterhouse, W. L., award of Medal of
Royal Society of N.S.W. .......... XXiv
Waterhouse, W. L., and Watson, I. A.,
Australian Rust Studies. VII .... 113
Watson, I. A., and Waterhouse, W. L.,
Australian Rust Studies. VII 5 dlalg
Whaite, Mrs. J. L., elected a Member .. xxvi
Wheat Stem Rust, Some Recent
Observations on, in Australia ..... 113

Willings, Mrs. L. M., elected a Member .. xxv
Wilson, R. D., elected a Member ...... Xxili
Wood, HE. J. F., elected a Member ......

Woodhill, A. R.—

A New Subspecies of Aédes (Steg-
omyia) scutellaris Walker (Diptera,
Culicidae) from Northern Australia 140

Elected a Vice-President ........... Xxiil

A Note on Experimental Crossing of
Aédes (Stegomyia) scutellaris scu-
tellaris Walker and Aédes (Steg-
omyia) scutellaris katherinensis
Woodhill

A
c
ie

5 ated,

mf ae

\
‘
‘ay
rere
Sr

(Issued 15th June, 1949.)

Vol. LXXIV. Nos. 341-342.
Parts 1-2.

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Vice-Presidents:
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~

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PROCEEDINGS, LXXIV, PARTS 1-2, 1949,

CONTENTS.

Presidential Address. Seventy-fourth Annual General Meeting: A Gummosis
Disease of Citrus in Relation to its Environment. By Lilian Fraser, D.Sc.

Elections

Balance Sheets for the Year ending 28th February, 1949
\
Australian Formicidae. New Genera and Species. By J. J. McAreavey, S.J.
(Seventy Text-figures.) :

The Distribution of Formic and Alcohol Dehydrogenases in the Higher Plants,
with particular Reference to their Variation in the Pea Plant during its
Life-cycle. By Daphne C. Davison, M.Sc. (Nine Text-figures.)

‘The Importance of Formic Dehydrogenase in the Oxidation Mechanisms of
Pisum sativum. By Daphne C. Davison, M.Sc. (Two Text-figures.) ..

On Australian Species of Creophilus (Coleoptera: Staphylinidae). By W. O.
Steel. (Communicated by J. W. T. Armstrong.) (Nine Text-figures)

Graptolites from Tallong and the Shoalhaven Gorge, New South Wales. By
Kathleen Sherrard, M.Sc. (Plates i, ii; thirty-three Text-figures.)

The Genus Dawsonia. By Alan Burges, M.Sc., Ph.D. (Twenty-six Text-figures.)

Revision of the Genus Brachycome Cass. Part II. New Zealand Species. By
Gwenda L. Davis, M.Se. (Twenty-four Text-figures. )

On Australian Dermestidae. Part V. Notes and the Description of Four New
Species. By J. W. T. Armstrong. (One Text-figure.)

Pages.
i-xVili
xix

XX-XxXii

1-25

26-36

97-106

107-111

Way

Vo re Lxxv. pie Nos. 343-344.

‘Parts 34,

Pe BINGS

_ LINNEAN SOCIETY

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Marine Bisigotcal L
MAE) Faia be et A

MAY 15 (85t

WOODS HOLE, Mass
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CONTAINING PAPERS READ IN MAY=AUGUST.

With two plates.
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The Linnean Society of New South Wales

LIST OF OFFICERS AND AND COUNCIL, 1949-50.

: President: E ;
R. N. Robertson, B.Sc., Ph.D.
Vice-Presidents: ;
A. R. Woodhill, B.Se.Agr.
Lilian Fraser, D.Sc.
Hon. Ree errs ASB: Walkom, D.Sc.

Secretary: Dorothy Carroll, B.A., B.Sc., PhD, 2 EC.)

Council:

G. D. Osborne, D.Sc., Ph.D.

R. N. Robertson, B.Sc., Ph.D.- _

T. C. Roughley, B.Sce., F.R.Z.S.

EK. Le G. Troughton, C.M.Z.S., F.R.Z.S.

J. M. Vincent, B.Sc.Agr., Dip.Bact.

A. B. Walkom, D.Sc.

H. S. H. Wardlaw, D.Sc., F.A.C.I..

Professor W. L. Waterhouse, M.C.,
D.Se.Agr., D.I.C.

A. R. Woodhill, B.Se.Agr.

Ida A. Brown, D.Sc.
- G. D: Osborne, D.Sc., Ph.D.

R. H. Anderson, B.Sc.Agr.

Ida A. Brown, D.Sc.

W. R. Browne, D.Sc.

*Professor N. A. Burges, M.Sc., Ph.D.

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

A. N. Colefax, B.Sc.

Ss. J. Copland, B.Sc.

Lilian Fraser, D.Sc.

Professor J. Macdonald Holmes,
Ph.D., F.R.G.S., F.R.S.G.S.

D. J. Lee, B.Sc.

B.Sc.,

Auditor: 8. 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 Shaftesbury Avenue, London,
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. 1897 XXII |} 10 O 8 6 9 0 12 6 1920 XLV /}10 9 Ths 9° 0 11 0O
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DESCRIPTIVE CATALOGUE OF AUSTRALIAN FISHES.
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Notics To AUTHORS. 3

Authors are requested to assist in facilitating the Society’s publishing work by observing
the following recommendations when preparing papers for submission to the Council. Perusal
of the PROCEEDINGS will show the general style to be adopted.

Manuscript.—Manuscripts should be double-space typed with a margin of at least one inch on

_ the left-hand side, top and bottom of each page. The title should be spaced clearly above
the text on the first page. The original should be submitted and a copy retained by the
author for checking proofs. A Table of Contents, on a separate sheet, should accompany
each manuscript; this is not necessarily for publication, but will serve to show the proper
relation of the headings. A manuscript when submitted to the Council should be complete
in every detail, both with regard to text, references and illustrations, for any extensive
alterations or corrections made on the proofs, if allowed, will be at the author’s expense.
No words except generic and specific names, and those to be printed in italics, should be
underlined. An Abstract of the paper should accompany the manuscript.

References.—References should be carefully checked by the author, who is alone responsible
5 for their accuracy. They should be listed alphabetically at the end of the manuscript, and
should be cited in the text by the author’s name, e.g., Bullough (1939) or (Bullough, 1939) ;
and in the list thus:

BuLLouGH, 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
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Tabulations.—Tables should be numbered consecutively and referred to specifically in the text
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' reduced to one-third, one-half or one-quarter the original size, giving a block 8 x 5 in. or
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PROCEEDINGS, LXXIV, PARTS 3-4, 1949.

CONTENTS.

Australian Rust Studies. VII. Some Recent Observations on Wheat Stem Rust
in Australia. By I. A. Watson and W. L. Waterhouse. (Plates iii, iv)

Notes on Microspore-types in Tasmanian Permian Coals. By J. A. and Roma
Dulhunty. (One Text-figure.) .. :

A New Subspecies of Aédes (Stegomyia) scutellaris Walker (Diptera, Culicidae).
from Northern Australia. By A. R. Woodhill. (Four Text-figures.) ..

Revision of the Genus Brachycome Cass. Part III. Description of Three New
Australian Species and some New Locality Records. By Gwenda L. Davis.
(Seven Text-figures. )

Notes on the Morphology and Biology of a New Species of Tabanus (Diptera,
‘Tabanidae). By Kathleen M. I. English. (Fifteen Text-figures.) a6

Crania in the Macleay Museum. By N. W. G. Macintosh ~

The Hair Tracts in Marsupials. Part III. Description of Species, concluded.
By W. Boardman. (Two Text-figures.) .. .

Studies on Australian Marine Algae. V. Observations on and Geographical
Records of Various Species, particularly those of the Gelidium Complex.
By Valerie May

Pages.

113-131
132-139

140-144

145-152

153-160

161-191

192-195

196-202

(Issued 15th December, 1949.)

Vol. LXXIV. Nos. 345-346.
Parts 5-6.

THE

PROCEEDINGS

OF THE

LINNEAN SOCIETY

OF

New SOUTH WALES

FOR THE yeanViaring Biolagizal ieee ea LUh y

vA En Ph CLS 4

| MAR 26 1951
WOODS HOLE, MASS.

Parts 5-6 (Pages 203-236; (= BH HD) SS
CONTAINING PAPERS READ IN SEPTEMBER-NOVEMBER,

ABSTRACT OF PROCEEDINGS, LIST OF MEMBERS AND
GENERAL INDEX.

SYDNEY:

PRINTED AND PUBLISHED FOR THE SOCIETY BY
AUSTRALASIAN MEDICAL PUBLISHING CO. LTD.,
Seamer Street, Glebe, Sydney,
and
SOLD BY THE SOCIETY,

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Registered at the General Post Office, Sydney, for transmission
by post as a periodical.

Agent in Hurope:
David Nutt, 212 Shaftesbury Avenue, London, W.C.2.

The Linnean Society of New South Wales

LIST OF OFFICERS AND COUNCIL, 1949-50:

President:
R. N. Robertson, B.Sc., Ph.D. |
Vice-Presidents:
Ida A. Brown, D.Sc. A. R. Woodhill, B.Se.Agr.
G. D. Osborne, D.Se., Ph.D. Lilian Fraser, D.Sc.
Hon. Treasurer: A. B. Walkom, D.Sc.

Secretary: Dorothy Carroll, B.A., B.Se., Ph.D., D.I.C.

~

Council:

R. H. Anderson, B.Sc.Agr. G. D. Osborne, D.Se., Ph.D.
Ida A. Brown, D.Sc. R. N. Robertson, B.Se., Ph.D.
W. R. Browne, D.Sc. T. C. Roughley, B.Sc., F.R.Z.S.
Professor N. A. Burges, M.Sc., Ph.D. EH. Le G. Troughton, C.M.Z.S., F.R.Z.S.
Dorothy Carroll, B.A., B.Sc., Ph.D., D.L.C. J. M. Vincent, B.Sc.Agr., Dip.Bact.
A. N. Colefax, B.Sc. A. B. Walkom, D.Sc.
S. J. Copland, B.Sc. H. S. H. Wardlaw, D.Sc., F.A.C.I.
Lilian Fraser, D.Sc. ‘ Professor W. L. Waterhouse, M.C.,
Professor J. Macdonald Holmes, B.Sc., ‘D.SeAgr., D.1.C.

Ph.D., F.R.G.S., F.R.S.G.S. A. R. Woodhill, B.Se.Agr. \

D. J. Lee, B.Se.
Auditor: S. J. Rayment, F.C.A. (Aust.).

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.

te s. d. s. d. s. d. s. d. s. d. s. d. s. d. s. d.
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1877 II 4 0 4 0 4 0 — 1900 XXV 8 0 10 6 10 6 17 6
1878 Til OD Meise 0) 6 0 a6 1901 XXVI | 10 O 9 0 bie7O) LNGO:
1879 IV 6 0 6 0 8 0 6 6 1902 XXVII 0) Thon 7631 150
1880 Vv. 6 6 — 16 Wane 1903 XXVIII 9 0 1206 14 0 TBA)
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1882 Vit eG 10 0 5 0 10 0 1905 xXxXX 625024 atO Nn 12-6 8 05
1883 Vill | 10 0 BO ai 8 0 1906 XXXT | 12 6 12 6 12 6 15 0
18384 IX 8 0 127570 25 0 25 0 1907 XXXIT 8 0 8 6 15 0 18 0
1885 aXe) AD OF ENG 15 0 17 6 1908 XXXIIT 7 0 9 0 14 0 12 6°
1886 Ij| 10 6 12 0 13 0 12726 1909 XXXIV | 12 O 1750 14 0 16 6
1887 IT 7 0 8 0 12 0 PAO 1910 XXXV/|i11 0 tO Gan0) 12.36
1888 Til | 15 0 24 0 20 O 18 0 1911 “XXXVI 9 6 9 6 9 6 10 0
1889 Iv | i1 0 16 0 19 0 tO 1912 XXXVII 8 6 25 0 12 6 15 0
1890 ae oe) 9 0 9 0 9 0 1913 | XXXVIII | 14 0 7 6 6 0 13 0
1891 VI | 10 O 9 6 ATO) Wee 1914 XXXIX | 13 O 17 0 25180 19 0
1892 VII 6 6 4 6 8 0 8 0 1915 XL /|17 O 12°56 10 0 dO
1893 Vill 5 0 INE YG) 6 0 9 0 1916 XLI |} 10 0 12 0 TO 19 0
1894 TX |} 120 12 0 13 0 8 0 1917 XLIT | 14 0 9 0 L2G 16 6
1895 DG NETIE YEA) 8 6 10 0 | 12 0 1918 XLITI | 20 0 14 0 21 0 19 0
1896 XXI 9 0? 6 6 nO 27 6 1919 XLIV | 12 6 eG LS S10
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DESCRIPTIVE CATALOGUE OF AUSTRALIAN FISHES. By William Macleay, F.L.S. [1881]. A few
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Notice To AUTHORS.

Authors are requested to assist in facilitating the Society’s publishing work by observing
the following recommendations when preparing papers for submission to the Council. Perusal
of the PROCEEDINGS will show the general style to be adopted.

Manuscript.—Manuscripts should be double-space typed with a margin of at least one inch on
the left-hand side, top and bottom of each page. The title should be spaced clearly above
the text on the first page. The original should be submitted and a copy retained by the
author for checking proofs. A Table of Contents, on a separate sheet, should accompany
each manuscript; this is not necessarily for publication, but will serve to show the proper -
relation of the headings. A manuscript when submitted to the Council should be complete
in every detail, both with regard to text, references and illustrations, for any extensive
alterations or corrections made on the proofs, if allowed, will be at the author’s expense.
No words except generic and specific names, and those to be printed in italics, should be
underlined. An Abstract of the paper should accompany the manuscript.

References.—References should be carefully checked by the author, who is alone responsible
for their accuracy. They should be listed alphabetically at the end of the manuscript, and
should be cited in the text by the author’s name, e.g., Bullough (1939) or (Bullough, 1939);
and in the list thus:

BuLLouGH, 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
of Scientific Periodicals.

Tabulations.—Tables should be numbered consecutively and referred to specifically in the text
by number. Hach table, provided with a heading descriptive of the contents, should be
\ submitted on a separate sheet. |

Illustrations.

Plates.—The size should not exceed 7% x 5 in. except if the subject will bear reduction. A
number of small photographs should be arranged to make one plate. Photographs should
show good contrast and be printed on glossy paper.

Line drawings will, as far as possible, be printed in the text. Drawings should be made on
White board or stiff white paper with Indian ink. It is advisable to arrange a number
of text-figures, all to bear the same reduction, on a single large sheet, which can be
reduced to one-third, one-half or one-quarter the original size, giving a block 8 x 5 in. or
4 x 5 in. without loss of essential detail. Hach separate figure should be clearly numbered
and all lettering should be plain and large enough to be clearly readable when reduced.
If co-ordinate paper is used for graphs it should be blue-lined. The explanation of Text-
figures should be supplied on a separate sheet; but the explanation of Plates should be
given at the end of the manuscript. Text-figures and Plates should be numbered consecu-
tively and referred to in the text by number. Indicate in the manuscript where each
Text-figure or group of figures is to be inserted.

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, LXXIV, PARTS 5-6, 1949.

CONTENTS.
Pages.
The Stratigraphy of the Lower Marine Series of the Permian System in the
Hunter River Valley, New South Wales. By G. D. Osborne, D.Sc., Ph.D.
(Two Text-figures. ) TIGER ee SUR RS aol Nave Bae SMa aaah Cita Ni ie Svat ae Se aM CR ames (tore
~-The Cotypes of Fordonia papuensis Macleay. By Arthur Loveridge. (Communi-
cated by S. J. Copland.) PSC Ue Seen ae EU MIA Cau A PULL Ly PURI dR aan aA AS UR hans 223

A Note on the Experimental Crossing of Aédes (Stegomyia) scutellaris .—
scutellaris Walker and Aédes (Stegomyia) scutellaris katherinensis
Woodhill (Diptera, Culicidae). By A. R. Woodhill. (One Text-figure.) .. 224-226

A Detailed Study of the Field Distribution of Strains of Clover Nodule Bacteria.

By Hilary F. Purchase and J. M. Vincent. (One Text-figure.) Pes NaN Meek=ooO
Abstract of Proceedings AeA a Ny A OMbral Wake Cera Na Sek ST Na onan aA AS : SS ARRAN Ru XX11i-xxvi
Mist of? Memberss ey se ely ie oe ae Re ee nl aE aie a XXVii-Xxxi
MistwafvGeneraiand GSpecies, hoe. co sie ee fee AS ek ee ak tn eee naa el er XXxli
List of Plates AAR re Heal k ce aR CE BUS ONT LU IAA hay CaN LT ANY sy AMIN Guest Sope tg AIA XXxii
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