aL
ay ty
PAS

THE

P ROTI ISO JIMS

or THE

Pere SOCIETY

OF

Nei Sonar WWoOMEaS

FOR THE YEAR

1954

VOL. LXXIX.

WITH NINE PLATES.
202 Text-figures.

SYDNEY:
PRINTED AND PUBLISHED FOR THE SOCIETY BY

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

and
SOLD BY THE SOCIETY.
1955.

il CONTENTS.

CONTENTS OF PROCEEDINGS, 1954

PARTS J-II (Nos. 371-872).
(Issued 10th June, 1954.)

Presidential Address, delivered at the Seventh-ninth Annual General Meeting,
3ist March, 1954, by J. M. Vincent, B.Sc.Agr., Dip.Bact. The Root-Nodule
Bacteria of Pasture Legumes. (One Text-figure.)

Elections

Inheritance and the Genetic Relationship of Resistance Possessed by Two
Kenya Wheats to Races of Puccinia graminis tritici. By D. S. Athwal
and I. A. Watson

Macadamia ternifolia F. Muell. and a Related New Species. By L. A. S. Johnson

Experimental Crossing of Aédes (Stegomyia) pseudoscutellaris Theobald and
Aédes (Stegomyia) polynesiensis Marks (Diptera, Culicidae). By A. R.
Woodhill Nap oot soUS ts merchspen, A RG ie M RS yas) cee Dem, 2 Moe tee te 8s

Cytological Studies in the Myrtaceae. IV. The Subtribe Huchamaelaucinae.
By S. Smith-White. (Forty-one Text-figures. )

A New Species of Hibbertia Andr. from Western Australia. By A. T. Hotchkiss.
(Plate i and four Text-figures.)

A Comparative Account of the Terrestrial Diatoms of Macquarie Island. By
J. S. Bunt. (Plates ii-iii and twenty-three Text-figures. )

Pages.
i-xxxii

XXxii

1-14
15-18
19-20
21-28
29-33

34-57

CONTENTS.

PARTS III-IV (Nos. 373-374).
(Issued 17th September, 1954.)

The Fossil Diptera of the Tertiary Redbank Plains Series, Queensland. By
E. F. Riek. (Five Text-figures.)

A Second Specimen of the Dragon-fly Aeschnidiopsis flindersiensis (Woodward)
from the Queensland Cretaceous. By EH. F. Riek. (Plate iv and one Text-
figure.)

Notes on the Bacteria belonging to the Rhodobacteriineae Breed, Murray and
Hitchens, and the Chlamydobacteriales Buchanan occurring at Macquarie
Island. By J. S. Bunt. (Plate v.)

The Genus Theobaldia (Diptera, Culicidae) in Victoria. By N. V. Dobrotworsky.
(Nine Text-figures. )

The Inheritance of Inflorescence Characters in Hucalyptus. By L. D. Pryor.
(Hight Text-figures.)

Notes on the Scleractinia or Stony Corals (Coelenterata) of Heron Island,
Queensland. I. A List of the Common Species. By K. E. W. Salter ..

Collenia frequens in Upper Proterozoic Rocks in the Northern Territory of
Australia. By D. M. Traves. (Communicated by Dr. Germaine A. Joplin.)
(Plate vi and one Text-figure.)

Australian Fungi. II. New Records and Revisions. By C. G. Hansford. (Two
Text-figures. )

Notes on Australian Beetles in the Tribe Bolboceratini formerly in the Genus
Bolboceras. By H. F. Howden. (Communicated by Dr. P. B. Carne.)

Antirrhinum Rust, Puccinia antirrhini D. & H. in Australia. By J. Walker.
(Plate vii and two Text-figures.)

lil

Pages.
58— 60
61- 62
63-— 64
65- 78
79-— 89
90-— 94
95- 96
97-141
142-144
145-155

iv CONTENTS.

PARTS V-VI (Nos. 375-376).

(Issued 26th January, 1955.) Pages.

Acarine Parasites of Two Species of Rattus from Brisbane, Australia. By
Robert |DOmMTOW ss a5 9 ce REM RS Mt OtS SCENT teem opiemreie ee me aes 156-158

A Second Species of Holothyrus (Acarina: Holothyroidea) from Australia. By
IROloever ADXoumarntonny, (COG) MNES THR) “oo oo co 06 65 oo 49 40 oc 159-162

The Nature and Significance of Non-reciprocal Fertility in Aédes scutellaris
and Other Mosquitoes. By S. Smith-White and A. R. Woodhill. (Three
Text-figures. ) seo! Felten g buses ic petaty lacet 2 Re eee NOE a EST eo lee crete os eee 163-176

A New Coptotermes and Ahamitermes (Isoptera) from Australia. By F. J. Gay.
(Communicated by Dr. A. J. Nicholson.) (One Text-figure) .. .. .. 177-181

Notes and Descriptions of Australian Chloropidae (Diptera). By Curtis W.
Sabrosky. (Communicated by Mr. D. J. Lee.) (Five Text-figures.) .. .. 182-192

The Culex pipiens Group in South-eastern Australia. III. Autogeny in Culex
pipiens form molestus. By N. V. Dobrotworsky. (One Text-figure.) .. .. 1938-195

An F1 Hybrid between Hucalyptus cinerea F. Muell. and Hucalyptus robusta :
Sint? BysvD a Pry ors \(Onemhext-fi cures) vie lie anew tine Santee nya eum 196-198

A Note on the Occurrence of ‘Anomalous’? Krasnozem in the Richmond-Tweed
Region of New South Wales. By J. W. McGarity and D. N. Munns. (Com-
municated by Dr. W. R. Browne.) (Plate viii and one Text-figure.) .. .. 199-202

Supplementary Notes on the Genus Brachycome Cass. Descriptions of Five
New Australian Species and Some New Locality Records. By Gwenda L.
Davis. (Twenty Text-figures.) Reenter natn hy cra Bee Ra IAMET AE RIN y coaiuted ahs ned 203-210

The Influence of the Physical Properties of a Water Container Surface upon its
Selection by the Gravid Females of Aédes scutellaris scutellaris (Walker)
for Oviposition (Diptera, Culicidae). By A. K. O’Gower. (One Text-figure. ) 211-218

Notes on the Morphology and Biology of Scaptia vicina Tayl. and a New
Species of Scaptia (Diptera, Tabanidae). By Kathleen M. I. English.
(Thirty-four Text-figures.) a Ea Cia uy CGR EIA aa aY arene ay Sto 219-229)

A New Genus and Species of the Tribe Labenini from Australia (Pimplinae,
Ichneumonidae). By Arthur W. Parrott. (One Text-figure.) .. .. .. 230-232

Australasian Cercatopogonidae (Diptera, Nematocera). Part VII. Notes on the
Genera Alluaudomyia, Ceratopogon, Culicoides and Lasiohelea. By

David J. Lee and Eric J. Reye. (Plate ix and thirty-six Text-figures.) .. 233-246
Balance Sheets for the year ending 28th February, 1954 .. .. .. .. XXX11i-xXxXXV
Nestrace of IPTOCCECINES fiisc ute Ge RS eos) ioe eee 0 Oe a Bier ne ear cKORORGV ATL XSI
WEISt Of, Mem berSe fio f ok Resd ck ve) APSO) cee hs 0) cee ire Bical aot eccrs cL ee oxcalavalent
Lists of New Genera, Subgenera, Species, Subspecies and Names GRAD celeste xlix-l
List of Plates CS ae ee en ENE i ir, mn eS Sy eh ea eM Sp ]
Corrigendum ae CME AEP EE he yt cic Pe tS PP one Ane ey <6 1

Index Berke h mae Uyatics prope te,"St- ad fay ale Oe Ra Ot Rc oe li-liv

ANNUAL GENERAL MEETING.
31st Marcu, 1954.

The Seventy-ninth Annual General Meeting was held in the Society’s Rooms, Science
House, Gloucester Street, Sydney, on Wednesday, 31st March, 1954.

Mr. J. M. Vincent, President, occupied the Chair.

The Minutes of the Seventy-eighth Annual General Meeting, 25th March, 1953, were
read and confirmed.

PRESIDENTIAL ADDRESS.

At the outset of this year’s Presidential Address, I feel sure you will want me to
record the very great joy we have all experienced on the occasion of the visit of Her
Majesty, Queen Elizabeth II, to Australia.

In considering the Society’s activities over the past year, it is my pleasant duty to
record once again our great indebtedness to the Honorary Secretaries, Dr. W. R. Browne
and Dr. A. B. Walkom. Each of these gentlemen has continued to give unstintedly of
time and effort, and has brought to bear on the activities of this Society a wealth of
experience and enthusiasm, to its very great benefit. It is most important that we should
not lose sight of the fact that the maintenance of our position at the present level,
especially in connection with the Proceedings, is only possible by virtue of the honorary
services thus rendered. The relatively satisfactory state of the Society’s finances is due
to their careful husbanding by Dr. Walkom, in his capacity as Honorary Treasurer, and
most of all to the fact that, for several years now, we have not had to meet the salary of
paid Secretary and Editor. It is difficult to assess the full measure of our indebtedness
to Dr. Browne and Dr. Walkom; it can only be matched by the satisfaction they must
each derive from seeing the consequent continued prosperity of our Society.

Nor should I omit to make special mention of the excellent services rendered to this
Society by our Assistant Secretary, Miss Allpress. It is Miss Allpress’ detailed knowledge
of Society procedure and organization that smooths the path of its honorary officers.
More than that, however, none of us who has had occasion to seek her help and advice
can fail to have been impressed by her graciousness, and enthusiasm for the Society’s
welfare.

As I proceed then to report on the Society’s activities, might I express your thanks
and my own great indebtedness, to these officers and, indeed, the whole Council for their
untiring efforts and support during 1953.

Parts 1-4 of Volume 78 of the Society’s Proceedings were published in 1953 and
Parts 5-6 in January, 1954. Volume 78 consists of 315 + Ivii pages, 18 plates and 201
Text-figures. Donations towards the cost of publication of papers were made by the
University of Melbourne, Mr. J. W. T. Armstrong, and Mr. K. E. W. Salter.

Library accessions from scientific societies and institutions totalling 1,561 were in
excess of the total for the previous year. Requests for library loans, especially from
interstate, University and C.S.I.R.O. libraries, have been as numerous as in previous
years. Many requests were made for early volumes of sets in the library, which
apparently were not available in these libraries. The Society is giving good service in
being able to accede to these frequent requisitions. The number of subscribers to the
Proceedings has increased and enquiries for complete sets of the Proceedings are
numerous. New exchanges to receive our Proceedings were commenced with Universidade
de Lisboa, Lisboa, Portugal; Institut Scientifique de Madagascar, Tananarive,
Madagascar; Research Council of Israel, Jerusalem, Israel; Agra University, Agra,

A

ii PRESIDENTIAL ADDRESS.

India: to receive Zoological Reprints: Nederlandsche Dierkundige Vereeniging,
Zoologisch Station, Helder, Holland; Zoologisch Museum, Amsterdam, Holland; and to
receive Entomological Reprints: Centro Ecuatoriano de Investigaciones Entomologicas,
Guayaquil, Ecuador. During January, 1954, the Assistant Secretary, with an assistant,
re-arranged the greater portion of the periodicals in the library in order to make
available much needed space for current accessions to the library. Commencement has
been made with the binding of volumes in the library and it is hoped to continue this
during the coming year.

Programmes of special interest were given at the monthly meetings during the year:

April: Lecturette by Professor John A. Moore on “Experimental Studies on the
Evolution of Australian Frogs”;

June: Lecturette by Mr. F. A. McNeill entitled “A Search for Rarities along the
Great Barrier Reef from Gladstone to Cairns”;

July: Lecturette on ‘The Importance of Light in Animal Ecology” by Dr. L. C.
Birch;

August: Lecturette entitled “Oxidative Mechanisms in the Wood-rotting Fungi’ by
Dr. B. J. Ralph; 5

September: Lecturette by Dr. N. C. W, Beadle entitled ‘‘Death of the Mulga, and
Decline in Soil Fertility in the West Darling Country”;

October: Lecture, illustrated by kodachrome slides, by Professor J. Macdonald
Holmes entitled “Journeyings in North Australia”;

November: Lecturette by Dr. L. G. Baas-Becking on “The Role of Henri Derx in
Tropical Biology”’.
We are indebted to and wish to thank all who contributed to these programmes.

Since the last Annual Meeting the names of 11 members have been added to the list,
four have been removed from the list under Rule VII, and four have resigned. The
number of members as at 15th March, 1954, is: Ordinary Members, 207; Life Members,
25; Honorary Member, 1; Corresponding Members, 2; Associate Member, 1; total, 236.
The average attendance at Ordinary Monthly Meetings during the year was 26.

The request of the Linnean Society and various State Royal Societies for financial
help was rejected by the Prime Minister’s Department.

Council decided that, in order to conserve the publication rights of members, no
paper by non-members would be accepted for reading and publication before the July
General Meeting in each year unless costs of publication are fully met from an outside
source.

By decision of the Council a flat rate of £3/10/- per volume will in future be charged
for each volume of the Society’s Proceedings.

At the invitation of the Royal Society of South Australia to nominate a representa-
tive to be present at the Special Meeting on 24th September, 1953, to celebrate the
centenary of that Society, Professor J. B. Cleland was asked by Council to represent this
Society and he consented to do so.

Council has been informed by the Director-General of Education that the Chief
Secretary and Minister for Immigration has approved of the recommendation of the
Fauna Protection Panel, that all applications by overseas institutions or individuals for
permission to obtain specimens of Australian fauna in New South Wales should be
referred to the Panel which will have regard to the requirements of the Australian
Museum when considering conditions to be attached to any licences issued.

The total net return from the Society’s one-third ownership of Science House for the
year was £597.

Under the auspices of the Joint Scientific Advisory Committee a fifth trip was made
in the Kosciusko region from 28th December, 1953, to 10th January, 1954, by a party of

PRESIDENTIAL ADDRESS. lil

four geologists and three botanists. Transport and accommodation were again provided
by the Botany Department of the University of Sydney, Snowy Mountains Hydro-electric
Authority and the Kosciusko State Park Trust. An entomologist of the Department of
Agriculture made a number of trips to the area to study the insect larvae responsible for
the killing of the snowgrass.

In addition to the congratulations extended to members during the year I wish to
extend them to Miss Dorothy Shaw, who left for Canada to take up a Thomas Lawrance
Pawlett Scholarship for two years; and Mr. T. G. Vallance, on the award of a Fulbright
Scholarship to continue his studies in metamorphism at the University of California,
Berkeley, U.S.A.

Linnean Macleay Fellowships.

In November, 1952, the Council reappointed Miss Mary Hindmarsh and Mr. T. G.
Vallance to Fellowships in Botany and Geology respectively for 1953.

Miss Hindmarsh’s resignation was accepted as from 30th April, 1953, to enable her to
visit England for further experience in cytological research. The work done by her
during her four months’ tenure of a Fellowship may be summarized as follows:
Experiments were carried out to determine the effects of metanilamide with p-amino-
benzoic acid. Cell division was inhibited by metanilamide which produced abnormalities
similar to those found previously with sulphanilamide treated roots. The addition of
p-aminobenzoic acid did not reverse metanilamide inhibition. This suggested that
sulphanilamide inhibition of cell division is specific. Further work on the morphological
effects of colchicine on root tips completed that aspect of the investigation.

Mr. Vallance continued his research studies, spent a short period in the field at
Kosciusko and commenced field-work in connection with the new research project on the
pre-Cambrian rocks of the Broken Hill region. His research on the Wantabadgery-
Adelong-Tumbarumba area was published in the Society’s Proceedings during 1953.
Council accepted Mr. Vallance’s resignation from his Fellowship as from 31st July, 1953,
Mr. Vallance having been awarded a Fulbright travel grant through the Institute of
International Education, New. York, enabling him to continue further research at the
University of California, Berkeley, U.S.A.

In November, 1953, the Council appointed Miss Nola Hannon and Miss Ruth Simons
to Fellowships in Botany for 1954.

Miss Hannon proposes to continue her studies on the status of nitrogen in
Hawkesbury Sandstone soils of the Sydney district. She hopes to commence studies on
the drainage waters and to investigate any possibilities of denitrification with the aim in
view of presenting a balance sheet for nitrogen, as well as accounting for its origin and
accumulation in these plant communities; also to elucidate the nutrient status of the
important elements of plant nutrition and to contribute to the general understanding of
nitrogen in soils.

Miss Simons proposes to continue her research on aspects of humus formation from
forest litter, very little work having been done on Australian native vegetation. Various
methods of isolation of fungi have been used and gradually a picture is being built up
of the species inhabiting the litter. Her research will include: further isolations of
fungi; use of natural media prepared from the Casuarina litter itself; completion of
histological studies; examination of material in the late stages of decay; and identifica-
tion of substances set free during breakdown.

Best wishes are extended to both Fellows for a successful year of research.

Macleay Bacteriologist.

During the year 1953-54 Dr. Yao-tseng Tchan was given permission to deliver four
lectures to advanced students in Agricultural Microbiology. His research was concerned
with three main topics: nitrogen fixation, nitrification and soil metabolism, and soil
algae. A large collection of samples by the C.S.I.R.O. Soil Survey Service permitted the
southern geographical limit to the distribution of the genus Beijerinckia to be defined.

iv PRESIDENTIAL ADDRESS.

An investigation of the nitrogen economy of semi-desert soils in the Broken Hill district,
in conjunction with the N.S.W. Department of Conservation and the Botany Department
otf the University, has shown that the non-symbiotic nitrogen-fixing bacteria, though
present, do not play an important role. Nitrogen-fixing blue-green algae may contribute
some nitrogen but rhizobia of native legumes are rare or absent. Twenty soils collected
by the C.S.I.R.O. are under investigation for nitrification and respiration. Although the
oxidation of soil organic matter seemed usual, nitrification was slow even under the
best conditions that could be provided. The direct study of soil algae made possible by
the method of fluorescence microscopy has contributed to a knowledge of their ecology,
the more so as a result of improvements involving a combination with a method of
dark-field illumination and the use of acridine orange as fluochrome. Several papers
have been published in the Proceedings during the year and a preliminary paper, with
Dr. Beadle of the Botany Department, has been submitted to the International Congress
of Botany in Paris. Dr. Tchan was also invited to participate in the Adelaide Soil
Science Conference, organized by C.S.I.R.O.

Your Council is glad to acknowledge the continued and generous support of the
Commonwealth Bank of Australia, the Rural Bank of New South Wales, the Commercial
Banking Company of Sydney, Limited, and the Bank of New South Wales that has
enabled the position to be maintained at a more reasonable financial status than would
have otherwise been the case. It has, however, become apparent to your Council that
more definite and long term measures are essential to obtain remuneration and security
more in keeping with the nature of the post of Macleay Bacteriologist. An approach to
the University of Sydney, suggesting a joint appointment as Macleay Bacteriologist and
University Lecturer, has been favourably received by the Senate of that body and steps
are now being taken to clarify the legal issues with a view to making the necessary
approach to the Equity Court for permission to vary the term of Sir William Macleay’s
will in this regard.

THE Roor-NopULE BACTERIA OF PASTURE LEGUMES.

The root-nodule of the leguminous plant constitutes a symbiosis between plant and
bacterium of the greatest biological and practical significance. It has therefore attracted
the attention of many workers and led to the accumulation of a considerable
bibliography. Fortunately the subject has been well covered by monographs and reviews
and I shall make these my chief landmarks. Fred, Baldwin and McCoy (1932) provided
an excellent basis and later developments have been covered by Wilson (1940), and the
recent review by Allen and Allen (1950).

There is in Australia to-day a mounting interest in pasture improvement, with
leguminous species playing an essential réle in raising nutritive value and improving
soil fertility. The dramatic success of subterranean clover (Trifolium subterraneum) in
the more southern half of the continent and its extension northwards towards a limit
yet to be defined, the proved value of lucerne (Medicago sativa) in diverse situations,
and the possibilities of barrel medic (Medicago tribuloides), have caught the interest of
many farmers. Moreover, economic conditions have been more favourable to a return
of cultivation paddocks to pasture which, properly established and managed, can prevent
and reverse depletion of nitrogen and organic matter,

In this account, I shall be chiefly concerned with the nodulation of Trifolium and
Medicago, genera that include our important pasture species. The subject matter will
also be largely restricted to those aspects of which we at Sydney have had some first-
hand knowledge. It follows then that my examples will almost all be drawn from
Australian experience, particularly in New South Wales.

I shall be using the results of my colleagues so frequently that separate acknowledge-
ment will often be impossible. I should like at the outset then to acknowledge the
valuable contributions of Dr. Hilary F. Purchase, Mrs. Lawrie M. Waters and, more

recently, Miss Kathleen J. Baird and Messrs. Marshall and Bockman to the work that I
shall be describing.

PRESIDENTIAL ADDRESS. v

THE NATURAL OCCURRENCE OF ROOT-NODULE BACTERIA.

As far as is known, the rhizobia that form nodules with true clovers (Trifoliun)
have, like their hosts, been introduced into this country, probably as seed contaminants.
There appears to be no native legume that acts as host for the bacteria that invade
clover. There is at least some degree of cross invasion between members of the medic
group and native species of Trigonella. It seems likely, however, that the strains of
Rhizobium found in the nodules of naturalized medics will more often have had their
origin overseas.

It is possible that rhizobia, recognized as they are chiefly by ability to invade a
legume, occasionally develop from non-invasive soil organisms. This, however, is a
proposition that lacks experimental evidence. Such data as are available show the
absence of clover or medic rhizobia from soils in which those legumes have not
previously been grown or which have not been open to wind-borne contamination.

Experience in areas new to clover in Western Australia (Cass Smith and Pittman,
1938) and South Australia (Strong, 1938) has been suggestive of lack of these rhizobia.
If soil conditions are favourable for the survival and multiplication of rhizobia, dust and
other sources of contamination may be sufficient to account for them being found in
some areas where the appropriate plant has never grown. Similarly conclusions from
nodulated control plants in the vicinity of inoculated plots and results with open pots,
cannot provide critical supporting evidence for the native occurrence of rhizobia. The
evidence for most other agricultural legumes is much the same, but native legumes may
provide strains able to nodulate one or more of the miscellany known as the cowpea
group.

Workers at the School of Agriculture have examined a number of localities in New
South Wales, Queensland and Macquarie Island. Since these have been based on
aseptically collected samples, under conditions of testing that guard against contamina-
tion, a positive result can be accepted as reliable. The negative cases suffer from the
uncertainty associated with small samples, but field experience in many of the same
areas has shown that the rhizobia are, at best, markedly deficient. They are reported as
sparse when some of the sub-samples are negative, others positive: a result that
correlates with patchy nodulation in the field. Cases recorded as fairly abundant or
abundant have rhizobia at least in a hundredfold dilution; such soils have given regular
nodulation of uninoculated seed. It appears that a condition of Rhizobium scarcity is
not uncommon, particularly for lucerne. Our results can be summarized:

Number of Localities with Rhizobia

Absent, or Fairly Abundant
Markedly Deficient Sparse or Abundant
Clover Rhizobia PA, bape 5 6 14
Lucerne Rhizobia .. Ete aes 9 3 3

All the markedly deficient cases for clover rhizobia were from virgin soils, except
in one case where uninoculated seed had been sown recently without any sign of
nodulation. The few rhizobia in red soils at Lismore and in two podsolic soils near
Armidale refiect low clover populations and, in the case of the acid Lismore soil, a
particularly unfavourable environment for rhizobial survival and multiplication (Vincent
and Waters, 1954). A Pilliga soil near Narrabri was also very acid (pH 5-0-5-2) and
deficient in clover rhizobia.

Besides their absence from virgin soils and sparseness in the acid soils at Lismore
and in the Pilliga, the medic rhizobia were deficient, or absent, in the case of several
soils in which the clover rhizobia were reasonably numerous. The same contrast was
found near Parkes (pH 5) where nodulated clover and non-nodulated medic plants were
regularly found in close proximity. Shortage of medic rhizobia at two sites near Dubbo
-was of special interest because these samples came from old wheat paddocks, a few
hundred yards from uncultivated land on which well nodulated burr medic (Medicago
hispida var. denticulata) grew abundantly. Ten to fifteen years of clean cultivated

vi PRESIDENTIAL ADDRESS.

fallow had removed virtually all medics and, in the absence of the host, the medic
rhizobia had become seriously depleted.

Finally, it is of some interest to note that nodulation of members of the “cowpea’’
group—Vigna sinensis, Desmodium, Stylosanthes, and Indigofera—was obtained with
three Queensland soils that lacked rhizobia for clover or lucerne. This undoubtedly
reflected the occurrence of Acacia in those areas.

THE SPECIFIC RELATIONSHIP BETWEEN HOST AND BACTERIUM.

The specitic relationship, or compatibility, that exists between a host plant and a
strain of Rhizobium is undoubtedly a major factor in determining invasion, formation of
a nodule, and maintenance of nitrogen fixation (Fred, Baldwin and McCoy, 1932;
Wilson, 1940; Allen and Allen, 1950; Thornton, 1952; Nutman, 1952). Whilst information
has to be sought keeping one of the two participants constant, it is misleading to
consider either without the other. By themselves such terms as “effective” and
“ineffective”, applied to a strain of Rhizobium, are without meaning. Few, if any, are
fully effective at fixing nitrogen with all the hosts they invade and general ineffectiveness
is also uncommon. It is only in the description of a defined partnership that the terms
have precision. Unfortunately, even nowadays, the literature includes examples of too
sweeping generalizations based on results with a single host species.

It is natural that workers in this field should have been anxious to find other
characters, morphology, biochemical and cultural properties, serology and bacteriophage
sensitivity that might serve as pointers to a strain’s ability to invade and form effective
associations. However, as would be expected from the complicated patterns of host-
bacterium specificity found in almost any collection of rhizobia, these attempts have been
quite unsuccessful.

In this account I shall consider specificity in relation to invasiveness (infective-
ness), relative nodulating success of competing strains, and ability of the nodule to fix
atmospheric nitrogen (effectiveness).

Specific Invasibility.

The fact that certain groups of hosts are nodulated by particular groups of bacteria
has led to the definition of ‘“cross-inoculation” groups (Fred, Baldwin and McCoy, 1932).
Within such groups reciprocal invasibility is common, beyond them unusual. However,
invasion can occur outside accepted groups and invasibility within groups may be
incomplete (Wilson, 1939a, 1944; Kleczkowska, Nutman and Bond, 1944). These excep-
tions are considered by some sufficient to invalidate the inoculation group as a basis for
species definition, or even the description of commercial inocula. Wilson (1939b) noted
a general correlation between cross-pollinating behaviour of the host (hence genetic
diversity), and ability to symbiose with diverse strains of rhizobia. Nutman (1946a,
1952) with red clover, and Aughtrey (1948) with lucerne, have shown how the host’s
genetic constitution can control invasion.

Grafting experiments have given conflicting results. Nutman (1952) has failed to
obtain nodulation of resistant stock by the use of susceptible scion, but recent experi-
ments with interspecific grafts (Bonnier, Hely and Manil, 1952) have been more
successful.

It is possible that the early stages of invasion may occur beyond the usual limits of
cross-inoculation groups without this being followed by nodule development. Allen
and Allen (1950) quote observations made in their laboratory of clover rhizobia invading
as far as the lucerne root cortex without causing meristematic activity.

Specific invasibility in the clover group—Trifolium ambiguum is rarely nodulated
by regular clover strains (Allen and Allen, 1947), but otherwise reactions within the
clover group have been taken as fully reciprocal. However, Baird (1953) found all
degrees of infectivity in a collection of one hundred and ninety isolations from several -
species of clover. One hundred and twenty-three cultures nodulated all four of red (Ca
pratense), white (T. repens), crimson (T. incarnatum) and subterranean Ge sub-

PRESIDENTIAL ADDRESS. Vil

terraneum) clovers on both occasions of testing. Eleven failed to nodulate any host on
either occasion and the remainder were inconsistent either between hosts (13), or
between occasions on the one host (43). Apart from those that were wholly negative,
nodulation was consistent on the species from which the culture was first obtained.
There appear then to be degrees of adaptation between bacterium and host in this group.

Specific invasibility in the medic growp.—Three sub-groups have been revealed
(Table 1).

TABLE 1.
Nodulation Sub-groups in Strains of R. meliloti.

'
|

Number of Strains Nodulating as
Sub-group Sub-group Sub-group
A. B. C. i
Nodulation of :
M. laciniata + — —
M. hispida — | + =
M. sativa + + +
Tsolated from :
M. laciniata 36 ae 31 2 1
M. hispida 75 1
Others 1 31 23

Note.—Here and elsewhere in this account the common burr medic,
M. hispida var. denticulata, has been listed as M. hispida for brevity.
Data from Purchase, Vincent and Ward, 1951b, and Waters, unpublished.

Almost all WM. laciniata isolates were unable to nodulate M. hispida (sub-group A)
and vice-versa (sub-group B). Cultures from nodules on other species also failed to
nodulate M. laciniata, but many of these were also incompatible with M. hispida (sub-
groups B and C). Several of the last group came from M. sativa and one from M.
falcata, growing in diverse localities. Others were stock cultures from Australian and
overseas sources.

There appears then to be a marked degree of incompatibility between M. laciniata
on the one hand, and several species represented by M. hispida on the other. Cross
invasion between them is imperfect and rare. Other hosts, notably M. sativa and
Melilotus alba, are freely invaded by isolates from all species and have been called
“bridging” hosts on that account.

The nodulation of a wider range of species by representatives of sub-groups A and B
is Shown in Table 2. M. rigidula was intermediate between M. hispida and M. sativa,
but most other species were more specific and behaved like M. hispida.

According to Mr. F. W. Hely’s assessment of the pollinating habits of the Medicago
(private communication), the range from bridging hosts to those that are more
specialized in their invasibility (e.g. M. laciniata and M. hispida) correlates in a general
way with decreasing cross-pollination.

These laboratory studies also provide an explanation of the common lack of nodules
on M. laciniata in the field, even when other medics are abundantly nodulated. A survey
undertaken by Mrs. L. M. Waters for the west and north-west of New South Wales
illustrates this:

Number of Plants Percent
Species Examined Nodulated
M. hispida var. denticulata .. .. .. .. 40 88
MER TVUNULITUGM, oe MBE ee oe Wael ei ee 23 87

M. laciniata EER SORY Oak NEMS Soin: AA; 15 33

vill PRESIDENTIAL ADDRESS.

The Host as a Factor in Determining Success in Nodulation from Mixed Inoculum.

It has been found that when several strains of Rhizobium are used as mixed
inoculum, the species of host can exercise a marked selection in determining the strains’
relative success in nodulation. When a mixture of strains 157 and 204 was used, it was
found that most of the nodules on red clover were due to strain 157 whereas strain 204
was chiefly responsible for the nodulation of subterranean clover (Vincent and Waters,

TABLE 2.
Nodulation Behaviour of a Larger Collection of Medicago.

Number of Strains Nodulating in
Tested on. |
Sub-group A. Sub-group Bb.
|
M. laciniata a ays, ft 17 0
M. hispida .. age Hie 0 22
M. murex | 0 22
M. orbicularis ae 1 22
M. ciliaris .. a A a| 1 22,
M. obscura .. aio oo. | 1 22
M. tribuloides | 3 | 22
M. arabica .. | 4 | 22
M. rigidula .. | 13 22
M. sativa ye ate 17 22
Mel. alba .. Se ull 17 22
Total strains tested 17 22

1954). These results were consistent between environments as different as soil and
seedling agar. A more detailed study of the nodulation of four clover species inoculated
with a mixture of five strains of Rhizobium (Vincent and Waters, 1953) showed the
importance of host selection even more clearly:

Host Species Major Nodulating Strains*
Red clover eka See RL SON, MoM rite. aad Reese ti Ee Deel eee LA tate OF
White clover .. .. A EESEE Dentnti ere beta Lale or | etic Beate ora MACS
Crimson clover aoe ae ee a wil) one. RA eee Casement a OM Se Ome con
Subterranean clover aa. ics ies GG lakal © SBML® cia CRC ee Th

* From a mixed inoculum comprising strains 36, 157, 204, 284 and 297.

It appears, moreover, from recent data obtained by Mrs. Waters that this influence
of host extends to variety within species. Yarloop and Mt. Barker varieties of sub-
terranean clover had consistent differences in the relative success of five strains used as
mixed inoculum.

There seems to be no simple relationship between ability to grow in the root’s
surroundings, singly or in competition, and a strain’s competitive ability to form
nodules (Vincent and Waters, 1953). This was also the result in a comparison of growth
in sterilized soil with relative nodulation success in unsterilized soil (Baird, 1953), and
in a simpler tube experiment involving two strains. In the first case three strains that
between them produced only 8% of the identified nodules, grew as well as the other two
that were responsible for 92%. In the second case, although relative growth of the two
strains was about equal, 86% of the nodules were due to one of them.

Our experiments have failed then to support the general validity of the relationship
between growth and nodulation postulated by Nicol and Thornton (1941) on the basis
of the fewer cases they could study with the methods at their disposal. More recent
results by Read (1953), working with Thornton at Rothamsted, have also failed to
demonstrate any regular connection between the two properties. Nor is it possible
always to explain this selective action of the host in terms of the relative invasive power
of the strains when in contact with the host singly, as far as this can be judged by

PRESIDENTIAL ADDRESS. ix

earliness of nodulation or number of nodules produced (Bockman, 1953, unpublished
results).

We do not know, then, what it is that determines the specific ability of the host to
control the proportion of nodules produced by different strains from a mixed inoculum,
or at what stage in the invasion process such factors operate. Our present knowledge
opens up, however, the practical possibility of using mixed inoculum to ensure the
production of effective nodules on each of several host species. This could be done by
providing a mixture such that its members would preferentially nodulate the host with
which they were best able to fix nitrogen. For example, a mixed sowing of red and
subterranean clovers could be treated with inoculum containing strain 157, preferentially
and effectively nodulating red clover, and strain 297, having similar properties for
subterranean clover. Equally well the data illustrate the foolishness of using mixed
inocula without knowledge of the behaviour of the constituents.

Specificity in Nitrogen Fixation.

It is unusual for the one strain of Rhizobium to be markedly effective on all species
of the hosts it is able to nodulate.

In four out of five collections of clover cultures, involving more than 500 isolations
tested against four species: white, red, subterranean and crimson, or ball clover (7.
glomeratum), 10% or generally less were reasonably effective against them all. The one
collection that gave a high proportion (59%) of uniformly effective strains was from a
restricted area. Never in our experience has the one strain been among the best with all
of four hosts. Similarly Parker and Allen (1952) found none of 35 strains markedly
effective with all of a similar grouping of hosts, although 14% combined moderate and
marked effectiveness. None was effective with 7. ambiguum. The more uniform
behaviour of 54 British cultures tested on red, white and alsike (7. hybridum) clovers
(Bond and McGonagle, 1951) is undoubtedly due to the group of host species involved.
The inclusion of species like subterranean, crimson and ball clovers would probably
have led to results similar to those quoted above.

It is even more unusual for the one strain of clover Rhizobium to be uniformly
ineffective. We have encountered two out of five hundred and nine, ineffective against
all of four hosts. Parker and Allen found two in their collection. Again, the larger
proportion of uniformly ineffective strains found by Bond and McGonagle could be
attributed to limitations imposed by their test species. Moreover, a strain that appears
wholly ineffective may yet have to encounter the host to which it is better adapted.

The position in the medic group is similar. 4% of fifty-four strains, able to nodulate
M. sativa, M. hispida and Mel. alba, were reasonably effective with the three. None was
ineffective with all three (Purchase, Vincent and Ward, 1951b). This agrees with data
of Burton and Erdman (1940) who found two out of about thirty strains reasonably
effective with the same three hosts and one ineffective. In our experience, even the
M. laciniata isolates, that were all ineffective with the homologous host, were sometimes
moderately effective with lucerne or Mel. alba.

Effectiveness sub-groups amongst the clovers—The performance of one hundred
strains, approximately twenty from each of five localities in New South Wales, has been
examined for all possible pairings of white, red, subterranean and crimson clovers
(Table 3.)

There is a highly significant positive correlation between the performance of strains
on red and white; crimson and subterranean. Outside these pairs there is either non-
Significant relationship (two pairings involving red clover), or a significant negative
correlation (each of subterranean and crimson clovers with white clover). It has been
our experience that these trends have fluctuated between localities. The Lismore
collection, for example, showed almost complete agreement between the behaviour of a
strain on subterranean and crimson, and the white/red clover relationship was also well
defined (Vincent, 1954). These distinctions were much less clear at Armidale (Baird,
1953). For these reasons it was thought most satisfactory to use a representative
composite collection for the analysis.

x PRESIDENTIAL ADDRESS.

The marked agreement between subterranean and ball clover (Vincent, 1945;
Purchase and Vincent, 1949), and the generally better performance of ball clover isolates
on subterranean and crimson clovers (see p. xiii) justify grouping these three together.
The results of others with alsike clover (Bond and McGonagle, 1951), 7. procumbens
and 7. fragiferum (Parker and Allen, 1952) justify their inclusion with white and red
clover. Our allocation of berseem clover to the subterranean clover sub-group (B) is
quite tentative, being based on three cultures only.

To summarize, the following sub-groupings appear to be justified on the basis of
like response to strains of rhizobia:

Sub-group A Sub-group B Sub-group C
ta)

T. repens T. subterraneum T. ambiguum
T. pratense T. incarnatum

T. hybridum T. glomeratum

T. procumbens T. alexandrinuwm

T. fragiferum

The earlier grouping of red and white and their separation from subterranean
(Strong, 1937) have been justified by these later more comprehensive tests, but was not
in fact sufficiently substantiated by the results with so few strains available at that
time. Moreover, despite these correlations, the extent of the exceptions must not be lost
sight of. As a practical measure, for example, it cannot be assumed that a strain
performing well with white clover will therefore be effective with red or alsike clover.
Apart from all the exceptions that we have encountered, the plant passage experiments
of Allen and Baldwin (1931) included cases in which re-isolations varied independently
in their reaction with white, red and alsike clovers.

TABLE 3.

Relationship Between Pairs of Clover Hosts in Nitrogen Fixation with 100 Strains.
Correlation Significance of
Host Pair. | Coefficient. Relationship.

|
Red-White be ats ml +0:47 Significant (P <0-01)
Subterranean—W hite ot a —0-28 Significant (P<0-01)
Crimson—White uy ae ay, —0-:27 Significant (P <0-01)
Subterranean—Red .. af eral —0:17 Not significant

| (P =0-1-0:05)
Crimson—Red , | =0F15 Not significant

| (P=0-2-0-1)
Crimson—Subterranean : +0-65 Significant (P <0-01)

Effectiveness sub-groups in the medics—Present data do not permit the same
examination of reactions with pairs of hosts in the medic group. It is possible, however,
to compare the reaction of 78 strains on M. sativa and Mel. alba and 65 strains on
M. sativa and M. hispida. There is a highly significant degree of agreement between the
first pair (r=+0-39; P< 0-01), but none between the second (r=-—0:20; P=0-2-0-1).
These results are in good agreement with those of Burton and Erdman (1940) and it
might be considered that the failure of some lucerne isolates to invade M. hispida
(p. vii) is a further manifestation of incompatibility between strains adapted to
lucerne and a more exacting host. Our results with cultures that came from different
hosts, though not as striking, showed the same trend as those of Burton and Erdman.
Lucerne isolates were, on the average, less effective on M. hispida than those that came
from the homologous host or other burr medics (M. arabica, M. minima). It is of
interest, too, to note in the work of Burton and Erdman that, although there was a
general agreement in performance on lucerne and Mei. alba, a culture isolated from
Mel. indica behaved like a M. hispida isolate. This is in agreement with a brief
observation by Strong (1940). Jensen (1942) had some indications, too, that a burr

PRESIDENTIAL ADDRESS. xi

medic sub-group differs from that of lucerne and that M. tribuloides belongs to the
former.

Although cultures obtained from M. laciniata were able occasionally to establish a
moderately effective association with the “bridging” hosts (M. sativa and Mel. alba)
they were generally ineffective with both hosts (Purchase, Vincent and Ward, 19510).

These effectiveness sub-groups might then be postulated, though rather tentatively:

A. M. sativa, Mel. alba, (Mel. officinalis).
B. M. hispida, M. arabica, (M. tribuloides, M. lupulina, Mel. indica).
C. M. laciniata.

THE ABILITY OF “NATIVE” STRAINS TO FIX NITROGEN IN ASSOCIATION WITH PASTURE LEGUMES.

We have had the opportunity of studying the symbiotic behaviour of a large number
of strains, chiefly collected from diverse localities in New South Wales. For a composite
account such as this it is convenient to make use of the nitrogen-free uninoculated
control, and the nitrate control, to facilitate comparisons between testing hosts and
localities.

This has been done by dividing the arithmetic difference between the nitrogen-
containing and the nitrogen-free control into ten equal classes, providing for additional
classes in excess of the nitrate control, and zero and negative classes below the non-
nitrogen control. Class 1 has the value of this control as its lower limit; successive lower
classes are 0, —-1, —2, etc. Hach histogram shows the frequency of strains in each class
expressed as a percentage of the total strains involved in the test.

Trifolium Species.

Cultures have been tested against four clover hosts: white, red and subterranean
clovers common to all, crimson clover used in later surveys, ball clover in the earlier.
Histograms are set out in Fig. 1. The performance of a group of isolates on the four
clovers can be followed horizontally; locality effects for the one testing host, vertically.

Cultures from Lismore were generally very effective with white clover, the common
clover of the area (Vincent, 1954). The most striking contrast was Kirkham which,
although on the central coast near Sydney, has too unreliable a rainfall for good
natural growth of white clover. Instead ball clover and knotted clover (7. striatum)
were fairly abundant. Tichborne, inland and dominated by ball and woolly clover (T.
tomentosum), provided many ineffective as well as some effective strains. Other
Tichborne sub-localities and a collection made between. Parkes and Coonabarabran (not
included in the present set of histograms, because these tests lacked nitrate controls)
had a much larger proportion of strains clearly ineffective on white clover. The Armidale
cultures are of particular interest. Those from the more easterly and elevated ‘‘wet”
localities came from a mixture of naturally occurring clovers that included 7’. repens,
T. dubium, T. procumbens and T. glomeratum. The more western “dry” localities had
mostly 7. glomeratum. Isolates from the “wet” localities were, on the average, superior
with white clover. Taree cultures were rather less effective on white clover than might
have been expected from a higher rainfall coastal area.

The performance of cultures on red clover was similar to white clover. Results with
subterranean clover were, however, quite different. Practically all the Lismore isolates,
and a large part of those from Taree, were ineffective. Those from Kirkham and
Tichborne were largely effective. Cultures from both Armidale sub-localities performed
rather poorly with this host. The best group of isolates did, however, come from a “dry”
area. Crimson clover closely paralleled subterranean clover except that the superiority
of “dry” over “wet” Armidale localities was more clearly shown. Results with ball
clover, which was used for Tichborne isolates, were also very similar to those with
subterranean clover.

The nodulation of uninoculated clovers sown in these areas has agreed quite well
with what would have been predicted from the survey. At Lismore nodules on
uninoculated subterranean and crimson clover have been almost always ineffective; those

xii : PRESIDENTIAL ADDRESS.

on white and red clover generally effective. There were apparently enough effective
strains at Taree to ensure adequate nodulation of subterranean clover at two of the three
sub-loealities, but the third, which appeared worst in the survey, showed a marked
response to seed inoculation (Marshall, 1953). The unsatisfactory prospect for sub-
terranean and crimson clovers at Armidale generally was borne out in field trials and

Wii tal Th gs T= SUBTERRANEAN RED CRIMSON
CLOVER CLOVER CLOVER CLOVER

LISMORE . | | LISMORE
' '
! |

| | | TAREE

1 '
1 i]
ARM!DALE ARMIDALE
‘WET’
' 1
' '

1 ' ' 1 1 i}

TAREE

ARMIDAL ARMIDALE
! ‘DRY’
\
60+ - : ;
t ! !
i= KIRKHAM
aA KIRKHAM IRKHA
w
oO
a
Ww
5 9 207 1 I
1 i) ' 1
1 I
i) 1
i CLUSTER
; i CLOVER
T ICHBORNE | TICHBORNE
: | 1 f 1 t ' t '
! ! t t 1 I 1 '
Cc N c N c N iS N

Figure 1.—The height of each column gives the percentage of the strains tested for each
locality having the relative effectiveness shown. As bases of comparison, C = uninoculated
nitrogen-free control and N = uninoculated nitrate control. Thus a high column near C, means
that many strains are ineffective in nitrogen fixation; a large proportion near N, indicates
effectiveness.

The histograms have been constructed from data due to the following: Lismore, Vincent ;
Taree and Kirkham, Marshall; Armidale, Baird; Tichborne, Purehase.

again there was good agreement with sub-locality observations (Baird, 1953). At
Kirkham the natural nodulation of subterranean clover was satisfactory, as would be
expected from the survey.

The effect of field host—There was a striking effect associated with field host in
two regions where suitable data were available (Table 4). The cultures that were
obtained from white clover at Armidale and Lismore were markedly effective with the

PRESIDENTIAL ADDRESS. xili

same species as testing host; those from other hosts were much less effective. Results
on red clover were very similar to those on white. Subterranean and, more strikingly,
crimson clover, were less effectively nodulated with white clover isolates, more effectively
with isolates from subterranean and ball clovers.

Besides the practical importance of these results, they are also of considerable
interest, showing as they do clear evidence of adaptation between the rhizobia and clover
hosts of an area. As for the means by which this adaptation is achieved, it seems
unnecessary to go beyond the selective advantage achieved by a strain in the presence
of a host with which it can form an effective association. This advantage would be
greatest where a shortage of available nitrogen threw the legume most on its symbiotic
resources and where the soil was unfavourable for the survival of the rhizobium away
from the host. Both of these conditions are known to operate in the Lismore red soils
(Vincent and Waters, 1954), and it is perhaps no accident that it is in the collection
from these soils that adaptation to white clover is closest, to subterranean and crimson,
least. These results also support the grouping of white with red; subterranean with
crimson and ball clovers (p. x) on the basis of effectiveness in nitrogen fixation.

TABLE 4.

Average Performance of Isolates According to Field Host.

Relative Effectiveness* on

Origin of Culture. |
White | Red Subterranean | Crimson
Clover. Clover. | Clover. | Clover.
Lismore :
White clover (a) 53 oh re ys 4-6 | 1-7 0-8 | 0-9
Red clover (6)... 2 ta in | 2-9 1:8 | 0-8 | 0-7
Subterranean and Crimson clovers, ineffective | |
nodules (c) oe - ah a a Beal 1°5 0-7 | 0-9
Subterranean and Crimson clovers, effective | |
nodules (d) .. os a fa: be 2-0 | 1-0 1:5 | 1-8
Significant differences .. ae Be! so ||| 6G, OSC fy 05 CSW \ CSG b, | WG, (0, C
Armidale : |
White clover (a) .. os ee oe areal 6-3 1:6 1:3 1:2
Subterranean clover (db) A A ial 2-6 1:2 1-5 ILO?/
Ball clover (c) .. ae ey: Shy Sof || 3-8 1-3 14: Loz
a

Significant differences .. ae as Pi a>b,c¢ a>b, ec Os C=

|

* Geometric mean of the group compared with nitrogen-free control=1.

Finally, it seems reasonable to generalize to some degree from these findings.
Higher rainfall areas, with white clover as the common natural clover, are likely to have
rhizobia reasonably effective for that host, red clover and other members of sub-group A.
They are likely to harbour a large proportion of strains parasitic or only weakly
effective with subterranean, crimson and ball clovers (sub-group B). Conversely, drier
regions that favour ball clover are likely, if numbers of rhizobia are sufficient, to provide
effective nodulation of sub-group B, though they can be expected to be less satisfactory
with sub-group A.

Species in the Medicago Group.

Whereas most clover cultures in our experience have been able to fix nitrogen in
association with the species from which they were obtained, the position is quite
different with Medicago laciniata. Not only does this species almost entirely resist
invasion by cultures from other medics, but its own nodules are generally ineffective.
None of the 23 strains collected from M. laciniata growing at Curlewis was able to form
a clearly effective association with that host, although some of them did show evidence
of nitrogen fixation with M. sativa and/or Mel. alba (Purchase, Vincent and Ward,
19516). Mrs. Waters has more recently found five strains that appear to be reasonably

xiv PRESIDENTIAL ADDRESS.

effective. These all came from the one region: Trangie and nearby Mungeribar, whereas
a further six ineffective strains came from localities as separated as HEdgeroi and
Temora. It is interesting to note that Fred, Baldwin and McCoy (1932) reproduce in
plate 25 a drawing by Eriksson, 1873, of this species, apparently effectively nodulated.

Isolations from M. hispida growing at Curlewis also contained many that were
ineffective on the homologous host. The collection did as well, or better, with M. sativa
and Mel. alba.

Cultures from other localities and hosts gave a wide range of reaction with WW.
hispida. Isolates from the homologous host included the most effective, while those from
lucerne were rather inferior (Cf. Burton and Erdman, 1940).

There was no apparent difference in average effectiveness of M. hispida and lucerne
cultures on M. sativa, but those from M. laciniata were markedly less effective. A group
of M. minima cultures from localities as far apart as Bathurst and Collarenebri were
outstandingly good with lucerne. Results with Mel. alba were very similar to those from
lucerne; in this case, however, data were not available for the M. minima collection.

SEROLOGY OF THE RHIZOBIA.

Serological methods have long been applied to the study of the root-nodule bacteria
(Fred, Baldwin and McCoy, 1932), but although the early work purported to test the
serological definition of species, in fact neither the method nor number of strains
examined was adequate for the purpose.

In 1940 it seemed worth applying to these bacteria some of the simple refinements
that had proved so useful with pathogenic organisms (Topley and Wilson, 1936).
Bushnell and Sarles (1989) had, indeed, at about that time, come to the same conclusion
and used three types of antigen: whole cell, heated and saline extracted. However, the
results they quoted were almost all restricted to the whole cell and they attempted no
distinction between flagellar and somatic antigens. Two points in agglutination
technique seemed to merit particular attention: (i) distinction between flagellar and
somatic agglutination; (ii) application of antiserum absorption tests. These were
applied in a study of groups of medic and clover rhizobia (Vincent, 1941, 1942).
Experience over a ten-year period was summarized in a further paper in the series
(Purchase, Vincent and Ward, 1951la). Meanwhile Kleczkowski and Thornton (1944) had
used substantially the same methods for clover and pea cultures.

Details of the methods are given in the papers quoted, but the basis of the test
might be briefly stated. There are at least two distinct kinds of antigen associated with
the rhizobial cell: some on the main body of the cell (somatic) and others on the
flagella. If a suspension of motile cells is injected intravenously into a rabbit, each
antigen causes the animal to produce, and excrete into its blood stream, a highly specific
antibody. A week or two after the course of injections is completed, the antibodies in
the blood serum will cause a characteristic clumping (agglutination) of a suspension of
the specific bacterium. Agglutinations due to flagellar antigens can be distinguished
from those due to somatic by the nature of the reaction and the use of specially treated
reagents: cells heated to destroy flagellar antigens, or antiserum heated mildly to remove
somatic antibody.

The application of this distinction reveals the rhizobia as serologically heterogeneous.
Within cross-inoculation groups strains are readily found that fail to cross-agglutinate in
either the flagellar or the somatic reaction. A broader, less specific, grouping of strains
can be made on the basis of flagellar reactions. The somatic reaction is much more
specific and, because different combinations of flagellar and somatic antigens occur, the
combined result of both reactions is an even better basis to strain recognition.

Clover Rhizobia.
The detailed study of 12 strains has revealed at least two flagellar and nine somatic
antigens (Purchase, Vincent and Ward, 1951a).
Larger collections of rhizobia havé mostly contained one or other of these two
flagellar antigens, but at least one other has had to be postulated to account for motile

PRESIDENTIAL ADDRESS. xV

cultures that fail to react with either antibody. Our experience of the frequency of
flagellar antigens can be illustrated by data for one hundred and twenty-six cultures
(Table 5).

TABLE 5.
Frequency of Certain Flagellar Antigens of R. trifolii
(Purchase and Vincent, 1949.)

Flagellar Reaction with Antisera to
Group A. Group B. Frequency.
36 | 61 94 204 46 157
|
+ 3 af ap = = 59
= | = = = ae ae 45
= | — = — - = 19
ar | SF ae as oF a 1
+ | + + a = + 2
| 126
|
|

Purchase (1953) working with a collection of British cultures found the same flagellar
group (A) in the majority of cases, with (B) second. At least three other flagellar
antigens had to be postulated.

Somatic reactions of the one hundred and twenty-six cultures with six antisera,
representing eight somatic antibodies, are shown in Table 6. Somatic reactants to
antisera 61 and 94 were also among the commonest of the British strains studied by
Purchase. As in our experience, reaction with antiserum 157 was rare.

TABLE 6.
Frequency of Certain Somatic Antigens of R. trifolii.
(Purchase and Vincent, 1949.)

Somatic Reactions with Antisera to
|

| | Frequency.
36 | 46 61 94 M5 a|) 204 Antigens.*

|
de ee Ne = = I 11
= | + — — = - TIL 25
= | + + - _ ~ iy AV 7
mea Meg HEEL L Pe 2 — — - V 14
= = + | + — = IL 31
= = = | ae ce at IV | 6
— | — = _ - | + IX 10
~ - —- — = - 22

| We

126

x Based on minimal somatic antigens postulated in Purchase, Vincent and Ward (1951a),
and to explain cross reactions demonstrated by the strains under test.

Amongst the Australian cultures there appeared to be no restriction to combination
between either of the flagellar with any of the somatic antigens. Purchase similarly
found that the British cultures that reacted with antisera to Australian strains 61 and
94 had similar freedom in combinations between the two types of antigen. On the other
hand the British strains that had also been used for the development of antisera showed
significant restrictions in this respect. Particular flagellar antigens tended to occur with
particular somatic antigens.

xvi PRESIDENTIAL ADDRESS.

Amongst the twelve cultures of FR. trifolii studied in detail, the flagellar reaction
alone would have revealed two strains, the somatic alone eight, and the combination of
both, ten. Distinetion of a larger collection was similarly improved by taking account of
both types of antigen (Purchase and Vincent, 1949).

The Medic Rhizobia.

The flagellar groups were not as clearly defined as with FR. trifolii, in that strains
commonly shared a minor antigen. They could, however, be distinguished quantitatively
as shown by these data (abridged from Vincent, 1941):

Flagellar Cross Reactions* of Strains 27 and 47

Antiserum

Antigen 27 47
ie 3,200 50-100
47: 50-100 800-1600

* The values given are the reciprocal of the highest Gilution
of antiserum giving a definite reaction.

In a detailed study of sixteen cultures (Purchase, Vincent and Ward, 1951a) it was
found that thirteen belonged to the “27 flagellar group’, two to the “47 group’, whilst
strain 101 failed to react with either, although positive with its own antiserum. The
“27 flagellar group” has been generally the commonest in our experience (Table 7).

TABLE 7.
Frequency of Certain Flagellar Antigens of R. meliloti.

| Frequency.

Type Flagellar

ia |
Reaction. | Vincent, Hughes and | Purchase, Vincent
| 1941. | Vincent, 1942. | and Ward,
| | 1951b.
|
|
27 | 39 5 68
47 | 7 9 il
Others 1 6 | 20
47 20 | 89

The somatic reactions of the sixteen strains required the formulation of at least
seven somatic groups and fifteen antigens. The relative frequency of reaction with a
range of test sera is shown below for the Curlewis collection (Purchase, Vincent and
Ward, 1951b) (Table 8).

It will be seen that about half the strains could be defined by seven somatic antigens.
Twenty-five, listed as “other various’, were mostly single strains that gave other com-
binations of reaction and six shared no antigen with any of the test strains. The three
strains that gave a somatic reaction with all antisera may possess a group antigen but
are more likely to show the non-reciprocal reaction already found for strains 8, 12 and
126 (Purchase, Vincent and Ward, 195iqa).

In the detailed study of sixteen strains, it was found that all four types of somatic
reaction encountered with the ‘47 flagellar group”, were also found with “27-flagellar
group” strains. Strain 101 which was very distinctive in its flagellar reaction shared a
somatic antigen with strains that occurred in both the other flagellar groups. The
sixteen strains could be put into three groups by flagellar reaction alone, thirteen on
the basis of somatic reactions and fourteen by the use of both.

Stability of Serological Properties.
Other workers have recorded the serological stability of cultures after continued
cultivation and plant passage. We have been able to compare the reactions of some
strains after sub-culturing for ten years (Purchase, Vincent and Ward, 1951a). In

PRESIDENTIAL ADDRESS. Xvii

total only 6% of the cases differed by as much as two fourfold dilutions. Two cases with
R. trifolii involved the flagellar reaction, but the negative reaction on the earlier occasion
may have been due to lack of motility in the antigen suspension. More notice has to be
taken of a later positive somatic reaction between 161 and 61. Discrepancies for R.
meliloti most commonly involved the minor antigen shared between “27” and “47”
groups and need not be regarded as serious. Practically all the inconsistent somatic
reactions were associated with the widespread non-reciprocal reactivity of antigens 8,
12 and 126. The only other case involved the reduction, but not the complete loss, of
titre by antiserum 66 with its homologous antigen.

TABLE 8.
Frequency of Certain Somatic Antigens of R. meliloti.

Somatic Reaction with Antisera to | |

| i | | Fre-
| | | | | Antigens.| quency.
7 47 5a 02) 74 a 84s 166 OM eelOn § | 12 | Sl |) 126
| | |
| | | | | |
+ | | | | TV eer
oy We etal Silly Baba ete Ra te an hapettes Usa
+ | + | | | | | (plop eXcTiTig | eas 8
ee 4 T, U1 8
\ronoan ati | VI 9
| Ista tr | | 1G, Wal 2
| | | te a | XG 8
+ + +] +f] + + | +] + ] 4 + | 4 ? 3
ee eas vee | e
Other various .. 25
89

Whilst variation in the reactivity of stock cultures over a long period is uncommon,
and relatively unimportant, some serious discrepancies have been encountered where
single colony re-isolations have been involved. This step would facilitate detection of
variants. In some cases failure to obtain somatic agglutination with unheated antigen
could be attributed to a Vi-like antigen and its effect could then be overcome by the
simple expedient of heating the cells (Vincent, 1953). However, in other cases heated
cells have failed to react with the homologous antiserum, so it seems there can be
non-reactive variants quite apart from the Vi-like phenomenon. Purchase (1953) has
also recorded the case of two mutants of strain A unable to react with an antiserum to
that strain (see also p. Xxvii).

Relationship to Other Properties.

Specific Invasiveness.—There is no consistent relationship between ability to invade
a host and the serology of the rhizobia. This is illustrated by cases of failure to obtain
cross agglutination between isolates from plants in the same cross-inoculation group
(Refs. Fred, Baldwin and McCoy, 1932, especially Stevens, 1923 and 1925; Vincent,
1941, 1942; Kleczkowski and Thornton, 1944) and even between cultures obtained from
the one plant (Hughes and Vincent, 1942). Cross-agglutination can also occur between
strains unable to invade the same host (Bushnell and Sarles, 1939; Kleezkowski and
Thornton, 1944; Purchase, Vincent and Ward, 1951b). ‘We have also encountered two
cases where loss of invasiveness in clover rhizobia has left their specific agglutinability
unchanged (p. Xxvili).

Although there appeared to be no relationship between serology and association of
strains with species of clover (Vincent, 1942), it appeared to operate in a collection of
medic strains obtained from widely separated localities (Vincent, 1941). Nine out of
twelve cultures obtained from M. hispida reacted like strain 27, whereas only one out of

B

Xvili PRESIDENTIAL ADDRESS.

eleven lucerne cultures showed the same reaction. Differences were also found in the
types of serology of isolates from M. laciniata and M. hispida vay. denticulata, growing
in the same locality (Purchase, Vincent and Ward, 1951b). This distinction could be due
merely to the segregation achieved by the inability of strains infecting one host to invade
the other. The failure of some lucerne isolates to nodulate M. hispida (p. vii) might
also explain serological differences in strains from the two hosts. In neither case does it
seem likely to reflect any causal relationship between serology and invasiveness.

Effectiveness in nitrogen fixation.—There is no evident relationship between effec-
tiveness in nitrogen-fixation and serological constitution. Loss of effectiveness without
serological change (Kleczkowski and Thornton, 1944; Vincent, 1944) precludes any
direct relationship between the two properties. Although Kleczkowski and Thornton
found a partial correlation between flagellar antigens and effectiveness on red clover, we
have not obtained any such effect in more detailed studies of Australian cultures (from
clover, Purchase and Vincent, 1949; and Medicago, Purchase, Vincent and Ward, 19510).

Bacteriophage Susceptibility—Kleczkowski and Thornton (1944) were unable to find
any relationship between the serology and the action of bacteriophage. Marshall (1953),
however, has found that strains of rhizobium ’phage obtained from University soil were
able to lyse only those strains that were related serologically to strains 297 and 298.
However, the larger number of strains that reacted with antisera of 297 and 298 were
phage-resistant. Although such an antigenic constitution appears necessary for
sensitivity to these strains of bacteriophage, it does not guarantee it. This has also been
demonstrated by the phage resistance of certain cultural variants of strains 297 and 298
that have retained their original antigenic properties.

Cultural Properties—Unlike the well known smooth to rough transformation in
Salmonella and Diplococcus pneumoniae, there appears to be little relationship between
the cultural properties of the rhizobia and their antigenic behaviour. Although Israilsky
and Leonowitsch (1933) had difficulty in checking the serology of saline unstable “rough”
variants, it seemed that these retained their ability to react with specific antiserum. I
was able to show, though again with some difficulty because of saline instability, that the
cultural variants encountered by Jensen (1942) retained the serological properties of the
original culture. Since then we have encountered numerous instances of colonial
variation, generally without any evident change in antigenic properties (see p. xxvii).

Application of Serological Methods.

The agglutination reaction almost certainly reflects some fundamental and relatively
constant characteristic of the bacterial cell. In this regard it compares very favourably
with other properties: cultural or biochemical characteristics, invasiveness and effective-
ness. It should therefore provide a ready and reliable method for the recognition and
separation of strains. Examples of its use will be considered.

Serological methods can be used as a means of obtaining information on the dis-
tribution of strains in widely separated areas and as a means of determining the number
of strains that can be recognized within a restricted area, on the one plant and within
the one nodule. Data along these lines are given in a separate section (p. xix).

Besides these uses there are many occasions in the field, glasshouse and laboratory
where it is necessary to be able to identify strains with some assurance. This may be
desirable to check the identity of a collection culture, to compare growth made by two or
several strains when in competition with each other or to identify the strain, or strains,
inhabiting a nodule. For these purposes the method will need to be sufficiently specific
to avoid confusion between strains, dependable, so as not to give false negatives or be
subject to variation in the characters used, and sufficiently simple in requirements of
time, media and equipment to permit adequate numbers to be examined. Finally, it is
important for certain types of experiment that the basis of identification shall not
interfere with the phenomena being investigated. For example, plant tests using strains
of differing effectivity as a means of identification are subject to this limitation.

PRESIDENTIAL ADDRESS. SKINS

On the whole serological methods meet those requirements better than any other.
The specificity that derives from distinction between flagellar and somatic reactions is
very good even in the presence of “native” soil forms. The method has a high order of
dependability, although not entirely free of the risk of false negatives in strains likely
to give non-reactive variants (p. xvii). For typing purposes it can be made quite simple
so that a large number of cultures can be identified at the one time. The lack of relation-
ship between serological and other properties (p. xvii) is an advantage in that labelled
strains can be selected generally with a minimum of experimental limitation.

Dunham and Baldwin (1931) made some use of the method in simultaneous inocula-
tion experiments, but its first extensive use for field studies was due to Thornton and
Kleezkowski (1950) in Britain—extended by Read (1953)—and Manil and Bonnier
(1950) with the collaboration of Tchan and Chaloignac, in Belgium. Our early use was
to study competition between strains in the laboratory (Purchase, unpublished datay
but, more recently, it has been used intensively for field as well as laboratory experiments
(Vincent and Waters, 1953 and 1954; Jenkins, Vincent and Waters, 1954; Baird, 1953;
Marshall, 1953).

THE DISTRIBUTION OF STRAINS OF RHIZOBIA.

As judged by serological identity the one strain of Rhizobium can occur in widely
different parts of the world. Similar, and in some cases identical, strains have been
found in Australia and overseas. R. meliloti strain 7, that came as a stock culture from
Wisconsin, U.S.A., appears to be identical with Australian strains 27 and 62.

An appreciable number of clover cultures were able to give the same flagellar and
somatic reactions as strain 204: a clone from Rothamsted Clover F (Purchase and
Vincent, 1949). British cultures have often shown the same reactions as our strains 61
and 94 (Purchase, 1953). Strains with identical serology have also been found in widely
separated parts of Australia. Clover strains 61 and 111, from Huston and Manildra
respectively, were serologically similar (Vincent, 1942), as were medic strains 27 and 62
from near Sydney and at Roseworthy, South Australia (Vincent, 1941).

Strains in a Restricted Area.—It has been exceptional in studies of the root-nodule
bacteria for the detailed locality of the host plant to be recorded. Besides some data,
chiefly based on effectiveness behaviour (Helz, Baldwin and Fred, 1927; Eckhardt,
Baldwin and Fred, 1931; Leonard and Dodson, 1933), Wright, Sarles and Holst (1930)
have briefiy reported a study of 156 strains of R. japonicum isolated from soils in which.
soybeans had been growing. Without giving details, they stated that they found a
marked tendency for strains from any one soil to fall into the same serological and
cultural group. Although the authors recognized six serological groups it would seem
that this appearance of homogeneity in a soil was likely to result, at least in part, from
failure to distinguish flagellar and somatic agglutinations (see p. xiv).

The heterogeneity of the clover and medic rhizobia that inhabit restricted areas is
apparent from Table 9. A similar result was obtained for two collections, when the
rhizobia were classified according to their ability to fix nitrogen in association with
several hosts (Purchase and Vincent, 1949; Purchase, Vincent and Ward, 1951b).

The mixed nature of the rhizobial population, even in a restricted area, has some
important implications. First it means that a fair number of cultures will have to be:
obtained before the nitrogen-fixing property of the rhizobia of the region can be assessed.
Second, the “native’’ population is likely to provide material for the selective action of a
particular host. In this way the overall population of an area might change its effective-
ness behaviour within a few years.

Multiple Infection of the One Plant.—Despite reports of resistance to invasion by a
second strain, there is little reason to believe that mixed infection of a plant (different
strains in different nodules) is difficult. Dunham and Baldwin (1931) noted that several
earlier workers had found more than one serological type of strain on the same plant.
They themselves were able to obtain ready double infection with two distinct strains
if the cultures were supplied simultaneously. Eight out of sixteen clover plants

xX PRESIDENTIAL ADDRESS.

examined by Hughes and Vincent (1942) harboured more than one serological strain and
the regular occurrence of more than one strain per plant (whether classified serologically
or on effectiveness in nitrogen fixation) was found in larger collections of cultures from
clover (Purchase: and Vincent, 1949) or Medicago (Purchase, Vincent and Ward, 19510).
Burton and Allen (1950), and Baird (1951) also obtained multiple infection by strains
of different nitrogen-fixing capacity.

Multiple Infection in the One Nodule.—Although it is common to find more than one
strain on the one plant, it is unusual to find more than one strain in a single nodule.
Dunham and Baldwin (1931) quote private reports (Sarles and Cray) of double infection
of the one nodule, but were themselves unable to find such a condition. Hughes and
Vincent (1942) made 73 single colony pickings from 12 nodules, mostly from plants
known to harbour different strains in different nodules. No case of multiple strains
within a nodule was encountered. Some factor evidently operates to make the population
within the nodule more homogeneous than between different nodules.

TABLE 9.

Serological Heterogeneity of Isolates from the Same Locality.

| ]
Minimum |
| Number of | Number of
Host. Location. | Tsolates. | Serological | Reference.
Types.
Medicago minima. Gunnedah. | 20 9 Hughes and Vincent, 1942.
T. glomeratum. Tichborne. | 19 8 a ‘, Be
T. repens. | Sydney. 18 6 ss a 5
M. sativa
M. hispida \ | Merrylands. 12 3 FA es
Melilotus alba } | |
T. glomeratum. Tichborne. 94 16 Purchase (1948).
5 | Parkes (26m. N.) 9 7 35
= | Yeoval. 9 5 a
As | Gilgandra (21m. N.). 6 4 ne
. | Coonabarabran (23 m. §.). 8 3 FA
M. hispida. | Curlewis. 63 26 Purchase, Vincent and Ward,
1951b.
M. laciniata. | Curlewis. 26 20 Purchase, Vincent and Ward,
| 1951b. ;

Where, as in the experiment quoted above, the plants were growing in soil, the
uniformity within the nodule might be attributed to the development of separate micro-
colonies in the vicinity of a root hair. If such colonies were discrete and had developed
from a cell of one or other of the strains growing separately in the soil, this might
explain the absence of mixed growth in the single nodule. An agar tube experiment, in
which organisms might be expected to be more intimately mixed, could then be expected
to increase the frequency of nodules with a mixed population. This has in fact been
found with a mixed inoculation (strains 36 and 297) of subterranean clover growing on
seedling agar. Strain 297 was the more successful nodulating strain, being isolated
from 89% of the nodules. Four further single-colony pickings were made from nodules
where the first culture had proved to be strain 36. If there was no restriction on double
occupation of a nodule, the chance for each replicate colony to be strain 297 would be in
the order of 9:1. Although double infection was demonstrated in three of the eleven
nodules examined in this way, the remaining eight nodules (40 colonies) were uniformly
strain 36. It can be concluded that a nodule can contain more than one strain of
Rhizobium but that the frequency of this is much less than the incidence of multiple
infection of the plant itself. It cannot be concluded from these results that the mixed
nodule population arose from double infection of the one root hair. Although the

PRESIDENTIAL ADDRESS. XXi

nodules were chosen to avoid mixing the contents of closely neighbouring nodules, there
was still the chance that a nodule could have occasionally involved coalescence of two
separate infections in its early development (Cf. Dunham and Baldwin, 1931).

THE ESTABLISHMENT OF ADDED INOCULA.

Where “native” rhizobia are deficient, or ineffective in fixing nitrogen with a legume,
it becomes a matter of acute practical importance to secure successful seed inoculation.
Fred, Baldwin and McCoy (1932) quote many instances of benefit due to this practice,
whilst Thornton (1947) has found that many of the rhizobia of British hill pastures
are ineffective on white clover. Documented reports for Australia are not very numerous.
O’Reilly (1937) recorded a marked response to lucerne seed inoculation in the Northern
Tablelands of New South Wales. Strong (1938) showed the importance of inoculation of
subterranean clover, lucerne and field pea in poor soils in South Australia. Our own
experience of localities, either deficient in clover or lucerne rhizobia (p. v) or having
native rhizobia poorly adapted to new clovers (Fig. 1), suggests that the successful
introduction of rhizobia is often likely to be important in pasture improvement pro-
grammes in this country.

TABLE 10.
Comparison of Single Strain Inocula in Lismore Red Soils.
1953—Subterranean
1952—Crimson Clover.* and Crimson Clover.
Inoculum.
Green Weight. | Percentage of Plants | Mean Effective
(Mg./Plant.) | Effectively Nodulated. Establishment.7
Nil 261 0 0
295 240 3 0
284 434 21 6
36 468 33 11
297 469 30 23
298 660 37 15

* Jenkins, Vincent and Waters, 1954.
+ Number of plants effectively nodulated with strain specified in a 6 ft. row (data due to Waters,
private communication).

Determination of success with inoculation has been greatly facilitated and made
more precise by the use of antigenically labelled strains (Thornton and Kleczkowski,
1950; Manil and Bonnier, 1950; Read, 1953; Vincent and Waters, 1954; Jenkins, Vincent
and Waters, 1954).

Strain of Organism.

Read (1953) found that several strains differed in their ability to nodulate red clover
in competition with “native” rhizobia. To some extent the relative performance of
strains was affected by locality. Our group at Sydney has collected data for five strains.
used to inoculate subterranean and crimson clovers in field trials at several centres.
As shown in results from Lismore (Tables 10, 11), strains 297 and 298 were consistently
satisfactory whether used as single inocula, or in a mixture. On the other hand strain
295 was almost always unsuccessful, even when applied singly. Strains 36 and 284
performed fairly well as single inocula, but were generally submerged by 297 and 298
in the mixed. Similar results have been obtained in field and pot tests with Armidale
soils (Baird, 1953; Bockman, 1954, private communication), in field trials with coastal
soils at Taree (Marshall, 1953), and in seedling agar. In a few as yet unexplained
Situations, strains 36 and 284 have shown up to greater advantage. The poor results
with strain 295 might well reflect declining invasive power, in that several sub-strains
have failed to nodulate clovers in pure culture. Strain 46, which performed poorly in
British tests (Read, 1953), has also become non-invasive in pure culture.

xxii PRESIDENTIAL ADDRESS.

Quantity of Inoculum.

It is not likely that there will be any simple relationship between the size of
inoculum and its successful establishment. So much will depend on the conditions the
bacteria encounter in the soil. Optimum conditions for their multiplication could permit
adequate nodulation with few bacteria per seed. Unfavourable conditions, or the
presence of an actively competitive native population, might justify a heavier dose of

TABLE 11.
Relative Success of Strains in Mixed Inocula in Lismore Red Soils.

Percentage Identified as
Host.
36 284 295 297 298
Crimson clover, 1952* 5 0 0 22 73
‘Crimson clover, 1953+ Ke 4 4 0 75 17
Subterranean clover, 19537 .. Ete 6 0 0 80 14

* Jenkins, Vincent and Waters, 1954.
7 Waters, private communication, 1953.

inoculum. The older literature (Fred, Baldwin and McCoy, 1932) includes the conflicting
results that would be expected from such considerations. Some workers found no
increase of nodulation with heavier inoculation; others found considerable improvement.

In this country, Spencer (1950) obtained markedly better nodulation by the use of
massive inoculum in an acid soil. There have been occasions, too, when a heavier
inoculum was advantageous on the acid red soils of the Lismore District (Table 12). On

TABLE 12.

Effect of Inoculum Dose on Nodulation.
(Crimson clover, Lismore red soils, Jenkins, Vincent and Waters, 1954.)

Inoculum Percentage of Plants Green Weight
(Cells/Seed, Effectively at 10 Weeks.
Approx.) Nodulated. (Mg./Plant.)
0 0 260
3,000 26 550
30,000 51 900

the other hand two soils from the Armidale region, known to be more favourable for the
survival of the bacteria, failed to show any consistent improvement over a one-
hundredfold increase of inoculum concentration (Table 13).

Inoculum more concentrated than the usual commercial preparation would seem to
be a worthwhile precaution where the conditions might be less favourable for the
organisms’ survival and multiplication. The maintenance of a large number of viable
cells in the inoculum between the time of its production and use could be important for
the same reason.

Method of Applying Inoculum.

Fred, Baldwin and McCoy (1932) describe progression from the soil transfer method
to pure culture preparations. Nowadays the culture is grown and supplied on agar, or
grown in liquid medium and mixed with finely ground, neutralized peat. Whichever
method is used, the viability of the culture will be affected by its age and the conditions
of storage. Aeration, retention of some water, and cool storage appear most important
{Fred, Baldwin and McCoy, 1932; Hedlin and Newton, 1948).

PRESIDENTIAL ADDRESS. Xxiii

Although the powdered peat method has virtually displaced agar cultures for
commercial purposes, it is by no means sure that it will always be as satisfactory. Two
out of nine commercial peat cultures we recently tested lacked sufficient viable cells, and
in some cases the presence of other organisms appeared to be excessive. A system of
regular checking of commercial cultures is certainly called for.

Heavy surface application of inoculum on established plants has been successful on a
number of occasions in small field trials (cases quoted in Jenkins, Vincent and Waters,
1954). Although a large field trial with lucerne was unsuccessful, probably due to
unfavourable climatic conditions and the use of the inoculum in too fine a spray, the
method has possibilities for plants already established but not nodulated. In fact it
might sometimes be easier to establish a new organism if the plant is already growing
and in an invasible state, than to introduce it with the seed into an unfavourable soil.
Some of our field and pot experiments with the red soils point in this direction.

TABLE 13.

Effect of Inoculum Dose on Nodulation.
(Subterranean clover, Armidale podsolic soils, Baird, 1953.)

Percentage of Nodules Due to Inoculum in
Inoculum. |
(Cells/Seed.
Approx.) Yarrowyck Soil. Abington Soil.
300 63 65
1,500 41 30
3,000 64 | 60
30,000 80 40

Conditions in the Soil.

There are many soils in this State sufficiently acid (pH 5, or less) to be marginal
for the survival and growth of the clover rhizobia, and too acid for most strains in the
medic group (cf. Jensen, 1942). In such cases soil acidity will be a major factor in
determining the survival of rhizobia added with the seed. In the red soil at Lismore
(Vincent and Waters, 1954), it was found that as much as 8 tons to the acre of calcium
hydroxide would be required to provide optimal conditions for rhizobial growth. On the
other hand the addition of as little as 1 ton/acre would permit the bacteria to survive
and grow, though more slowly. Although even this quantity would seem excessive on
economic grounds, drilling seed and lime into the soil together could be expected to
permit an application of as little as 2 to 3 cwt./acre to exercise a favourable effect in the
seed’s immediate environment. }

Harmful effects due to the acidity of superphosphate itself need consideration. The.
importance of avoiding contact between the inoculated seed and acid phosphate before
sowing has long been recognized (Reid, 1930; Pittman, 1935; Cass Smith and Pittman,
1938), but there is also danger of a harmful effect in the drill row, especially in soil that
is already on the acid side of neutrality. The use of “neutralized” superphosphate
(Pittman, 1944) is to be commended. The superiority of the superphosphate plus
dolomite plots in the Lismore field trials (Table 14) would be chiefly due to this factor.
Further results due to Crofts and Jenkins (1953, private communication) show the
same effect (Table 15).

Beside its influence on survival of rhizobia due to change of pH, the Ca*+ ion is
important for the organisms’ growth and for nodulation (refs. quoted in Fred, Baldwin
and McCoy, 1932; Albrecht, 1933, and McCalla, 1937). Spencer (1950) found local
Situations where the same effect appeared to operate and Pulsford (1952) has shown its
importance in the Lismore red soils.

The superiority of superphosphate alone over dolomite alone (Table 14) would be
due to phosphate itself, known to stimulate both partners in the symbiotic association

XXi1V PRESIDENTIAL ADDRESS.

(Fred, Baldwin and McCoy, 19382). A factorial experiment by Pulsford (1952) supports
this conclusion for the Lismore red soils.

The field trials in 1952 (Table 14) also showed a marked depression due to the trace
element mixture. This was almost certainly due to the action of copper which, sown
with the inoculated seed in the drill, could be in toxic concentration at a critical stage

TABLE 14.

Comparison of Fertilizer Treatments.
(Jenkins, Vincent and Waters, 1954.)

Percentage of
Green Weight. Plants

Fertilizer.* (Mg./Plant.) Effectively

Nodulated.
Nil ai bis oe bsg AN Ne 232 i
Superphosphate Ae hs a at 767 48
Dolomite Be oe ae of Ae 574 39
Trace mixture .. ad Su ae Ns 181 2
Superphosphate + dolomite Eis aie ots 1141 57
Superphosphate +trace mixture fe ra 226 &
Dolomite +trace mixture Els ah is 226 16
Superphosphate +dolomite-+trace mixture .. 639 29

Increment due to:

Superphosphate a aS ne ae 388 19
Dolomite .. aa oe ihe ie 294 20
Trace mixture as a ae a —358 —25

* Rates per acre: Superphosphate, 175 lb.; dolomite, 350 lb.; in trace mixture:
copper and zine salts, each 10 lb.; sodium molybdate, 8 oz.

following sowing. Although copper has suggested itself as the most likely toxic com-
ponent of the mixture, it has not yet been possible to test the elements separately. The
concentration in the seed’s immediate environment seems to be critical and it has been
difficult to reproduce well enough the situation encountered in the 1952 field trial. This
experience does, however, suggest the desirability of avoiding direct contact between
inoculated seed and possibly toxic trace elements.

TABLE 15.
Influence of Superphosphate and Calcium Carbonate on Nodulation.

Percentage of Plants Effectively Nodulated.*
Superphosphate (Cwts./Acre).
Plus
CaCO3.
(Cwts./Acre.)

bo
>
for)

0 ike/ 19 12
2 47 38 27
40 53 48 55

* Data due to Crofts and Jenkins, 1953, private communica-
tion; average values for Yarloop subterranean, and crimson
clovers.

The Lismore field trials showed also how the favourable influence of superphosphate
and dolomite and the harmful influence of the trace element mixture could be ascribed
wholly to their effect on nodulation (Table 14). This is supported by the fact that

heavier inocula were better able to withstand the trace element effect (Jenkins, Vincent
and Waters, 1954).

PRESIDENTIAL ADDRESS. XXV

Soil temperature and moisture are obviously important factors in determining the
survival of rhizobia already in the soil or added as inoculum with the seed (Fred,
Baldwin and McCoy, 1932). In this country a report by Swaby and Noonan (1946) cites
cases where too much and too little soil water seemed to be responsible for inoculation
difficulties with canning beans in two successive years. Excess water may limit
aeration and hence the survival of rhizobia and the health of the host. However, white
clover has grown quite successfully in a waterlogged soil at Beerwah (Q.), abundantly
and effectively nodulated just above the water level. High temperatures and desiccation
of surface soil might easily operate against the survival of root-nodule bacteria. This is
a further argument for the practice of sowing a pasture legume directly into the cooler,
moist soil under a grass sward (Breakwell and Jenkins, 1951, 1953), rather than in a
clean seed bed, where the surface soil is likely to become hot and dry.

Competitive Effect of Native Micro-organisms.

Other micro-organisms, especially those of the rhizosphere of the leguminous plant,
will inevitably constitute an important factor in determining the success or failure of
an introduced inoculum. This they might do by inhibiting or stimulating the growth of
introduced rhizobia, by affecting the ease with which the root can be invaded and, if
themselves rhizobia able to invade that host, competing for the production of nodules.

Competition by “native” rhizobia.—Although cases of crop failure due to ineffective
native rhizobia have not been frequently reported, there are undoubted situations where
this will in fact operate (Leonard, 1930; Leonard and Dodson, 1933; Thornton, 1947,
and cases in this account, where the introduction of uninoculated subterranean and
crimson clovers is likely to result in a large proportion of ineffectively nodulated plants).
Where this is the position it is important that the introduced bacteria shall be able to
grow sufficiently in the root environment to give them the chance to nodulate and,
perhaps more important, that they shall have a competitive advantage at the point of
plant invasion. This property is a specific function of host and organism (p. viii).

Invasion by one strain does not itself prevent other nodules being produced by
different strains. There is ample evidence for multiple infection of the one plant,
although organisms within the one nodule are generally the same strain (p. xx). Other
things being equal the effective nitrogen-fixing strain should have a competitive
advantage over the ineffective. The large, long functioning nodules that it produces are
likely to restrict further invasion (Nutman, 1949), whereas early-formed ineffective
nodules will exercise little restriction on further nodulation.

As yet there is little precise information as to the deleterious effect of ineffective
nodules on plants that otherwise carry a fair complement of efficiently functioning
nodules. There is some suggestion (Burton and Allen, 1950) that the simultaneous
presence of effective and ineffective nodules on crimson clover, leads to a plant response
intermediate between those obtained when good and poor strains are applied separately.
There are, however, serious discrepancies in the data quoted by those authors. The
importance of obtaining early effective nodulation was, however, clearly demonstrated.

Effect of the non-rhizobial population.—Besides the records of earlier work on this
subject (Fred, Baldwin and McCoy, 1932), Casas-Campillo (1949) and Allen and Allen
(1950) have provided useful reviews of evidence for antagonism exercised by other
soil micro-organisms towards rhizobia. Most of the evidence is circumstantial and fails
to demonstrate the effect in the soil itself. Some later work, particularly that of
Casas-Campillo with soluble fractions obtained from soils that are inhibitory to rhizobia
and some soil experiments are more convincing. Hedlin and Newton (1948) have shown
that the growth of other organisms in peat and soil cultures can markedly depress the
number of rhizobia.

There are also records of rhizobial stimulation by other organisms (Fred, Baldwin
and McCoy, 1932). Krasilnikovy and Korenyako (1944) have put forward evidence for
“activating” bacteria able to hasten and increase nodulation by R. trifolii under other-
wise aseptic conditions. They also reported stimulation in pot and field trials. Harris

Xxvi PRESIDENTIAL ADDRESS.

(1953) has reported the stimulation of a weakly invasive strain of #&. trifolii in the
presence of other soil bacteria in a tube experiment. Harris uses this to explain an
increase in nodulation by the same strain in non-sterile, compared with sterile, soil. The
general significance of the phenomenon and the practicability of making use of
“activating” bacteria are matters meriting further investigation.

Examples of Success with Seed Inoculation.

The results of field trials in the red soils of the Lismore District (Tables 12, 16, 17)
illustrate the success so far achieved in what can be regarded as a difficult environment
for rhizobia (Vincent and Waters, 1954). Although the degree of success differed
between localities, there was in every case a marked improvement over the uninoculated
controls. In respect of the latter it should be noted that, as indicated by serological tests,
an appreciable proportion of the effective nodulation could represent escapes of inoculum
used for the inoculated plots.

TABLE 16.

Success with Seed Inoculation of Clover Sown in Lismore Red Soils.*

| Number of Effectively Nodulated
Plants per 6ft. Row.
Locality. Treatment.
Subterranean

Clover. Crimson

(Mt. Barker). Clover.
1 Nil. 9 3
Inoculated.+ 28 28
2 Nil. 2 1
TInoculated.+ 12 14
3 Nil. 3 2
TInoculated.+ 12 7

* All with 2 ewt. superphosphate, 2 cwt. dolomite and 7 Ib. ZnSO, and
CuSO, per acre (the last sprayed on before sowing). Data due to Waters, 1953,
private communication.

+ Mixed inoculum of five strains.

Yarloop subterranean clover and vetch were rather more difficult to inoculate (Table
17). In the case of Yarloop this is in agreement with difficulties found elsewhere with
this variety during the same growing season (Carter, 1953) and shown to a less extent
in recent pot experiments (Waters, 1954, private communication). These last experiments
also show that the relative competitive ability of strains may be different between
Yarloop and Mt. Barker subterranean clovers (p. viii). From tests conducted by Crofts
and Jenkins since the last growing season, it seems that some, at least, of the residual
difficulties with seed inoculation in these soils can be ascribed to inadequate neutraliza-
tion of the acidity of superphosphate (p. xxiii).

VARIATION.

It is of considerable theoretical and practical significance that the root-nodule
bacteria show marked variability in all their important characteristics. Demonstration
of this variability is considerably facilitated by the collection of phage resistant mutants
(Kleczkowska, 1950).

There have been several reports of cultural variation in Rhizobium (Israilsky and
Starygin, 1930; Almon and Baldwin, 1933; Israilsky and Leonowitsch, 1933; Jensen,
1942; Kleczkowska, 1950). Although the terms “rough” and “smooth” have been used to
describe such cultural variants, it is doubtful whether this is sound practice in view of
the well known and regular antigenic changes associated with these terms with other
bacteria (Jensen, 1942).

PRESIDENTIAL ADDRESS. XXVii

Kleezkowska (1950) found that phage resistant mutants of four strains all gave
colonial variants, in significantly different proportions between strains (8% to 51%).

Recent examination of seven of our stock cultures of FR. trifolii that had passed
through at least two successive single colony pickings, and had been maintained for
periods ranging from 2 to 13 years, showed at least two distinct colony types in all but
one of them. Serological tests of all colonial types were generally constant, although
one variant gave a weakened flagellar reaction, and variants of two other cultures
revealed the ‘“Vi-like’’ condition recently reported for rhizobia (Vincent, 1953).

Two cultures that were studied more closely yielded four cultural types, listed as A,
B, C and D (strain 298) and A’ and B (strain 297).

TABLE 17.
Success with Seed Inoculation of Legumes Sown in Lismore Red Soils.*
Percentage of Plants Effectively Nodulated.

According to Localities. According to Hosts.
Not Not
Locality. Tnoculated. Inoculated. Host. Tnoculated. Inoculated.
1 32 83 Subterranean Clover:
Mt. Barker. . 12 72
2 1 46 Tallarook 16 84
3 11 66 Yarloop 15 | 43
4 3 70 Crimson Clover 20 82
5 25 66 Barrel Medic 0 62
6 3 61 Vetch 12 48

* Fertilizer treatments as Table 16. Data due to Crofts and Jenkins, 1953, private communication.

Apart from a type A culture (that subsequently produced a type D colony variant
(298/5331)), the sub-strains have retained their characteristics over a period of six
months’ regular sub-culture. This type D colony did not show the associated serological
variation found in the earlier type D variants (see below).

. Serology.

The stability of this property has already been discussed briefly (pp. xvi, xvii). The
occurrence of a “Vi-like” variant, that interferes with the ordinary somatic reaction but
whose effect can be avoided by the use of heat, has also been noted (p. xvii). There
have, however, been other occasions when a proportion of single colony variants have
failed to react with homologous antiserum, even when heated (Vincent and Waters,
1953). These could occur amongst cells growing outside the plant or coming from the
same nodule. Kleczkowska (1950) also found that 8% of her phage resistant colonies
failed to agglutinate with antiserum to the parent culture. It seems that the rhizobia
can vary between a reactive and non-reactive state so far as their somatic agglutination
is concerned.

Colony types A, A’, B and C of strains 297 and 298 retained their original serological
properties, but type D of strain 298 had an antigen not present in the parent strain,
whilst at the same time having its reaction with the homologous antiserum weakened or
lost (Table 18). The more recent variation to type D colony (298/5331) was accompanied
by no such change in serology.

Symbiotic Properties.

Although cultures can generally be maintained on agar for many years without
showing loss of invasiveness or change in nitrogen-fixing ability with a particular host,
some variability is likely to be encountered, with some strains more than others.
Attempts to correlate variation in these properties with the storage medium and

XXViii PRESIDENTIAL ADDRESS.

conditions have yielded inconclusive results. There are, for example, conflicting reports
on the merits of sterilized soil for this purpose (Fred, Baldwin and McCoy, 1932;
Albrecht and Turk, 1930; and Nutman, 19460). Nutman found that nine months’ storage
in sterilized soil gave 30-35% of the cultures ineffective in association with red clover.
Variation in the reverse direction has been more difficult to obtain. However, Nutman
found that two out of 22,000 nodules produced by an ineffective variant yielded an
effective culture and he was also able to obtain a mutant effective on a selected line of
red clover for which the parent culture was ineffective (Nutman, 1952). Kleczkowska
(1950) found a marked degree of variability amongst phage resistant mutants, in these
as well as other properties. About 16% of further single colonies from a sub-strain that
was itself an ineffective variant red clover, failed to nodulate that host. 22% of
phage resistant colonies obtained from the same effective strain as used initially by
Nutman were clearly ineffective in nitrogen fixation with red clover. Kleezkowska also
found variation in the reverse direction less common.

TABLE 18.
Properties of Variants from Strains 297 and 298.

Serology.
Somatic Reactions Nodulation Reactions Phage Reactions.*
with Antisera with Subterranean Lysed by
Colony Type. Sub-strain. ai Clover.
|
94 297 298 1c |) IW |) Ie Wae
A. “ Pearl Drop ”’ en 298/533 == sr ats Early, effective. = = =
A}. 3 “a 297/31 se ae ar > 2 =F = =
Al. eA ae 297/33 sh oP + yy ”» = a ss
B. “Small White’ .. 298/531 = + a Late, ineffective. =P sr =
298/538 AA a + + =
297/32 ak se + 2 > ca aF =
Cae hincr Colonvaaee 298/534 = + =F Early, effective. oF SP =
298/539 99 2 se oF =
D. “Milky Colony” .. 298/536 — se —(+)+ | Early, ineffective. _ — —
298/53-18 = aE (SP) >» » a 7

* Phage tests due to Marshall (1953).
7 Positive agglutination obtained with antigen growth in liquid medium, negative grown on agar slope.

Plant passage has been studied as a means of modifying the effectiveness of rhizobia
(Refs. in Fred, Baldwin and McCoy, 1932, esp. Allen and Baldwin, 1931). It seems likely
that plant passage will select rather than cause variants (Vincent, 1944). The fact that
Nutman (19460) found his cultures to be stable to twelve plant passages, although giving
a large proportion of variants when stored in soil, could reflect differences in two
selection methods.

In our own experience we have had two cultures lose their power of invasion whilst,
in each case, retaining their distinctive serology (cf. Kleczkowska, 1950, and non-
invasive colonial variants studied by Almon and Baldwin, 1933). A reported acquisition
of invasiveness (Krasilnikov, 1941) awaits confirmation.

Colonial variants of strains 297 and 298 showed an interesting correlated variation
in their symbiosis with subterranean clover (Table 18). In the case of the two colonial
forms A (A’) and B, the symbiotic behaviour could be compared for what must have
been independent variations in the two strains. The three A (A’) type variants invaded
early and produced large and effective nodules. The four B type invaded late and gave
multiple bead-like ineffective nodules. Type C colonies formed effective nodules early
and type D, ineffective nodules, but without the delay that characterized type B. All
sub-strains tested against red clover nodulated at about the same time and all showed
the low level of partial effectiveness characteristic of the parent cultures.

PRESIDENTIAL ADDRESS. Xxix

Variation in Bacteriophage Sensitivity.

The rapid outgrowth of a phage resistant culture is characteristic of all kinds of
bacterium-phage behaviour, and occurs as readily in the case of Rhizobium. The variation
from susceptibility to resistance and vice versa appears to occur quite frequently. Laird
(1932) showed that cultures derived from a single cell were likely at any time to consist
of a mixture of resistant and susceptible cells and Kleczkowska (1950) found that up to
30% of single colony re-isolates from a phage resistant suspension were once more
susceptible to that phage. Cultures can, however, be obtained having a consistent
susceptibility or resistance to phage. Undoubtedly these reflect cases where the balance
is tipped towards the preponderance of one or the other type of cell. More equal balance
between the two will result in partial clearing.

Cultural variants A (A’) and B showed similar bacteriophage behaviour within types
and different behaviour between types (Table 18), whether initially from strain 297 or
298. Types C and D similarly showed correlation between colony type and phage
susceptibility.

CONCLUSION.

Although this account has been restricted to aspects coming more directly in the
field of my own experience, I would not like to finish without reminding you that there
are many others concerning the root-nodule bacteria that have been scarcely touched on.
Indeed one has to admit that, despite the advances that have been made in a period of
almost seventy years, we still know remarkably little about the fundamental processes
of the root-nodule symbiosis. What is in fact known about the exact nature of the
specificity between host and organism? What is the mechanism of entry and how does
the host exercise a selective action at this point? What are the detailed stages by which
nitrogen is fixed, and what are the respective roles of host and bacterium in this process?
Answers to questions like these will be forthcoming only as a result of a more determined
attack by investigators with diverse interests and training. So much remains yet to be
discovered that one wonders, not at the progress so far, but at the relative scarcity of
workers in this field. The physiological and genetic approach (Nutman, 1952) has a
great deal to commend it, but at some stage the biochemist too will have to face up more
courageously to the difficulties inherent in a study of such a biological partnership. The
Wisconsin and Helsinki schools have indeed made notable contributions, but, compared
with other fields of biochemical research, biological nitrogen fixation has been neglected.
There are other worthwhile investigations as well. There are indeed so many gaps in
our knowledge that there is work enough for the bacteriologist, geneticist, physiologist,
biochemist and plant nutritionist alike.

References.
ALBRECHT, W. A., 1933.—Inoculation of legumes as related to soil acidity. J. Amer. Soc. Agron.,
25: bi 2-522.
— and Turk, L. M., 1930.—Legume bacteria with reference to light and longevity. Miss.
Agric. Exp. Sta. Res. Bull., 132, 19 pp.
ALLEN, ETHEL, K., and ALLEN, O. N., 1947.—A survey of nodulation among leguminous plants.
Proc. Soil Sci. Soc. Amer., 12: 203-208..
, 1950.—Biochemical and symbiotic properties of the rhizobia. Bact. Rev.,

14 : 273-330.

ALLEN, O. N., and BALDWIN, I. L., 1931.—The effectiveness of rhizobia as influenced by passage
through the host plant. Wis. Agric. Exp. Sta. Res. Bull., 106:56 pp.

ALMON, Lois, and Baupwin, I. L., 1933.—The stability of cultures of Rhizobium. J. Bact.,
26: 229-250.

AUGHTREY, J. D., 1948.—Effect of genetic factors in Medicago on symbiosis with Rhizobium.
Cornell Univ. Agric. Exp. Sta. Mem., 280:18 pp.

BaikD, KATHLEEN J., 1951.—Maultiple infection of clover plants by strains of the nodule organism
in the field. Nature (Lond.), 168:116-117.

, 1953.—Clover root-nodule bacteria in the New England Region. Internal Rep. Univ.
Sydney.

Bonpb, G., and McGonaciz, M. P., 1951.—The effectiveness of strains of the nodule organism
when associated with different species of clover. Ann. Appl. Biol., 38: 246-251.

XXX PRESIDENTIAL ADDRESS.

Bonnier, C., HELY, F. W., and MANIL, P., 1952.—Essai d’adaptation a Soja hispida de souches
de Rhizobium non spécifiques. Influence de greffes sur la spécificité d’hote du genre
Rhizobium. Bull. Inst. Agron. Sta. Rech. Gembloux, 20:137-140.

BREAKWELL, E. J., and JENKINS, H. V., 1951.—Some aspects of pasture improvement in the
Richmond River Area of New South Wales. Proc. Aust. Agrostology Conf., Canberra.

— r , 1953.—A pasture overseeding implement. J. Aust. Inst. Agric. Scez.,
19: 109-110.

Burton, J. C., and ALLEN, O. N., 1950.—Inoculation of crimson clover (Trifolium incarnatum L.)
with mixtures of rhizobia strains. Proc. Soil Sci. Soc. Amer., 14:191-195.

—______ and ErpMAn, L. W., 1940.—A division of the alfalfa cross-inoculation group correlating
efficiency of nitrogen fixation with source of Rhizobium meliloti. J. Amer. Soc. Agron.,
32: 439-450.

BUSHNELL, O. A., and SARLES, W. B., 1939.—Investigations upon the antigenic relationships
among the root-nodule bacteria of the soybean, cowpea and lupine cross-inoculation groups.
J. Bact., 138: 401-410.

Carter, O. G., 1953.—Pasture establishment, plant nutrition and rhizobial investigations in
County of Cumberland, and at Yarraman, North West Slopes of N.S.W. Thesis, University
of Sydney.

CASAS-CAMPILLO, C., 1949.—E]1 antagonismo microbiano en relacion con las bacterias de los
nodulos de las plantos leguminosas. Ciencia, 9:193-199. (C.S.I.R.O. Translation No. 1921.)

Cass SMITH, W. P., and PirrMan, H. A. J., 1938.—The influence of methods of planting on the
effective inoculation and establishment of subterranean clover. J. Agric. W.A., 16: 61-79.

DUNHAM, D. H., and BALDWIN, I. L., 1931.—Double infection of leguminous plants with good and
poor strains of rhizobia. Soil Sci., 32: 235-249.

EckKHARDT, M. M., BALDWIN, I. L., and Frep, E. B., 1931.—Studies of the root-nodule organism
of Lupinus. J. Bact., 21: 273-285.

FrebD, E. B., BALDWIN, I. L., and McCoy, ELIZABETH, 1932.—Root nodule bacteria and leguminous
plants. Univ. Wisconsin, Madison.

Harris, J. R., 1953.—Influence of rhizosphere micro-organisms on the virulence of Rhizobium
trifoli. Nature, 172:507.

HEDLIN, R. A., and Newton, J. D., 1948.—Some factors influencing the growth and survival of
rhizobia in humus and soil cultures. Can. J. Res. C., 26: 174-187.

Hety, G. E., BALDWIN, I. L., and Frep, E. B., 1927.—Strain variations and host specificity of the
root-nodule bacteria of the pea group. J. Agric. Res., 35: 1089-1050.

HuGHES, 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., 67: 142-152.
ISRAILSKY, W., and LEONOWITSCH, KATHARINE, 1933.—Dissoziation bei einigen Bakterienarten II.

Zentr. Bakt. Parasitenk. Infek., II, 88: 216-235 (Allen and Allen, 1950).

and STARYGIN, LyprA, 1930.—Die Dissoziation bei einigen Bakterienarten. Zentr. Bakt.
Parasitenk. Infek., Il, 81:1-11 (Allen and Allen, 1950).

JENKINS, H. V., VINCENT, J. M., and WATERS, LAWRIE, M., 1954.—The root-nodule bacteria as
factors in clover establishment in the red basaltic soils of the Lismore District, New South
Wales. III. Field inoculation trials. Aust. J. Agric. Res., 5: 77-89.

JENSEN, H. L., 1942.—Nitrogen fixation in leguminous plants. I. General characters of root-
nodule bacteria isolated from species of Medicago and Trifolium in Australia. Proc. LINN.
Soc. N.S.W., 67: 98-108.

KLECZKOWSKA, J., 1950.—A study of phage resisting mutants of Rhizobium trifolii. J. Gen.
Microbiol., 4: 298-310.

~-- , NutMAN, P. S., and Bonpb, G., 1944.—Note on the ability of certain strains of
Rhizobium from peas and clover to infect each other’s host plants. J. Bact., 48: 673-675.

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

KRASNILIKOV, N. A., 1941.—Mutability of nodule bacteria. Compt. Rend. Acad. Sci., U.R.S.S.,

ol 3 7G.
and KoRENYAKO, A. I., 1944.—Influence of soil bacteria on the virulence and activity
of Rhizobium. Mikrobiologiya, 13: 39-44 (C.S.1.R.O. Trans. No. 1868).

LairD, D. G., 1932.—Bacteriophage and the root-nodule bacteria. Arch. Mikrob., 3:159-193.

LEONARD, L. T., 1930.—A failure of Austrian winter peas apparently due to nodule bacteria.
J; Amer Soc. Agron., 227217 1-279.

and Dopson, W. R., 1933.—The Effects of nonbeneficial nodule bacteria on Austrian
winter pea. J. Agric. Res., 46: 649-663.

MANIL, P., and BoNNiER, C., 1950.—Fixation symbiotique d’azote, chez le luzerne (Medicago
sativa L.). Bull. Inst. Agron. Sta. Rech. Gemblouxz., 18 : 89-126.

MaArSHALL, K. C., 1953.—Root-nodule bacteria as factors in the establishment of pasture
legumes in milk zone areas of New South Wales. Thesis, Univ. Sydney, Australia.

McCatLia, T. M., 1937.—Behaviour of legume bacteria (Rhizobiwm) in relation to exchangeable

calcium and hydrogen ion concentration of the colloidal fraction of soil. Univ. Missouri
Coll. Agric. Res. Bull., 256: 44 pp.

PRESIDENTIAL ADDRESS. XXXI1

Nicou, H., and THoRNTON, H. G., 1941.—Competition between related strains of nodule bacteria
and its influence on infection of the legume host. Proc. Roy. Soc. (Lond.), B, 130: 32-59.

Nutman, P. S., 1946a.—Genetical factors concerned in the symbiosis of clover and nodule
bacteria. Natwre, 157: 463-465.

, 1946b.—Variation within strains of clover nodule bacteria in the size of nodule
produced and in the “effectivity” of the symbiosis. J. Bact., 51: 411-432.

, 1949.—Physiological studies on nodule formation. II. The influence of delayed
inoculation on the rate of nodulation in red clover. Ann. Bot. N.S., 13: 261-283.

, 1952.—A discussion on symbiosis involving micro-organisms. Host factors influencing
infection and nodule development in leguminous plants. Proc. Roy. Soc. (Lond.), B,
139: 176-185.

— and READ, MARGARET P., 1952.—Symbiotic adaptation in local strains of red clover and
nodule bacteria. Plant and Soil, 4: 57-75.

O'REILLY, M. V., 1937.—Lucerne “inoculation”. An essential to successful establishment on
lighter soil types. Agric. Gaz., N.S.W. Dept. Agric., October : 549-554.

Parker, D. T., and ALLEN, O. N., 1952.—The nodulation status of Trifolium ambiguum. Proc.
Soil Sci. Soe. Amer., 16: 350-353.

PirtMaNn, H. A. J., 1935.—Leguminous crops and their place in the building up of soil fertility.
J. Dept. Agric. W.A., 12:105-116.

, 1944.—Commonwealth of Australia, Patent Spec. 127,817.

PuLsForD, J. S., 1952.—Legume establishment and nutrition on the red basaltic soils of the
Richmond River District of New South Wales. Thesis, Univ. Sydney, Australia.

PURCHASE, HILARY F., 1948.—Studies of clover nodule bacteria with particular reference to their
distribution in New South Wales. Thesis, Univ. Sydney, Australia.

—, 1953.—Studies of the rhizosphere of leguminous plants and its relation to nodule
formation. Thesis, Univ. Lond.

and VINCENT, J. M., 1949.—A detailed study of the field distribution of strains of clover

nodule bacteria. Proc. Linn. Soc. N.S.W., 74: 227-236.

, VINCENT, J. M., and WARD, LAwRIE, M., 1951a.—Serological studies of the root-nodule
bacteria. IV. Further analysis of isolates from Trifolium and Medicago. Proc. LINN. Soc.
N.S.W., 76: 1-6.

—_—, ——_, —, 1951b.—The field distribution of strains of nodule bacteria from species
of Medicago. Aust. J. Agric. Res., 2: 261-272.

READ, MARGARET P., 1953.—The establishment of serologically identifiable strains of Rhizobium
trifolii in field soils in competition with the native microflora. J. Gen. Microbiol., 9:1-14.

Reip, W. D., 1930.—Hstablishment of lucerne root nodules. Further experiments with the
inoculum. N.Z. J. Agric., 41: 310-314.

SPENCER, D., 1950.—The effect of calcium and soil pH on nodulation of T. swbterraneum L. clover
on a yellow podsol. Aust. J. Agric. Res., 1: 374-381.

STRONG, T. H., 1937.—The influence of host plant species in relation to the effectiveness of the
Rhizobium of clovers. J. Counc. Sci. Indus. Res., 10:12-16.

, 1938.—Legume establishment and its function in relation to the development of some
of the poor soils on Kangaroo Island. J. Agric. South Aust., 41: 542-550.

, 1940.—Non-effective associations of nodule bacteria and legumes. J. Aust. Inst.
Agric. Sci., 6: 14-20.

Swasy, R. J., and Noonan, J. B., 1946.—Nodulation of field (canning) beans. New South Wales
Dept. Agric., Misc. Publication No. 3302.

THORNTON, H. G., 1947.—The biological interactions of Rhizobiwm to its host legume. Antony
v. Leeuwenhoek etc., 12: 85-96.

—— , 1952.—A discussion on symbiosis involving microorganisms. Introduction: The
symbiosis between Rhizobium and leguminous plants and the influence on this of bacterial
Stram=, Proc, Roy: Soc, CLond))> By 1393 UV 7aRa76.

and KLECZKOWSKI, J., 1950.—Use of antisera to identify nodules produced by the
inoculation of legumes in the field. Natwre (Lond.), 166:1118-1119.

TopLEY, W. W. C., and WILSON, G. S., 1936.—Principles of bacteriology and immunity. Hdward
Arnold, Lond.

VINCENT, J. M., 1941.—Serological studies of the root-nodule bacteria. I. Strains of Rhizobiwm
meliloti. Proc. LINN. Soc. N.S.W., 66: 145-154.

, 1942.—Serological studies of the root-nodule bacteria. II. Strains of Rhizobiwm
trifolii. Proc. LINN. Soc. N.S.W., 67: 82-86.
, 1944.—Variation in the nitrogen-fixing property of Rhizobium trifolii. Nature (Lond.),
3: 496-497.
, 1945.—Host specificity amongst root-nodule bacteria isolated from several clover
species. J. Aust. Inst. Agric. Sci., 11: 121-127.
, 1953.—Vi-like antigen in clover nodule bacteria. Awst. J. Sci., 15:133-134.
—, 1954.—The root-nodule bacteria as factors in clover establishment in the red basaltic
soils of the Lismore District, New South Wales. I. A Survey of ‘native’ strains. Aust.
J. Agric. Res., 5: 55-60.

XXXii PRESIDENTIAL ADDRESS.

VINCENT, J. M., and WATERS, LAWRIE M., 1953.—The influence of host on competition amongst
clover- root-nodule bacteria. J. Gen. Microbiol., 9: 357-370.

9 _ 1954.—The root-nodule bacteria as factors in clover establishment in the
red basaltic soils of the Lismore District, New South Wales. II. Survival and success of
inocula in laboratory trials. Aust. J. Agric. Res., 5: 61-76.

WILSON, J. K., 1939a.—Leguminous plants and their associated organisms. N.Y. (Cornell)
Agric. Exp. Sta. Mem. No. 221: 48 pp.

—____. 1939b.—A relationship between pollination and nodulation of Leguminosae. J. Amer.
Soc. Agron., 31: 159-170.

—_____, 1944.—Over five hundred reasons for abandoning the cross-inoculation groups of the
legumes. Soil Sci., 58: 61-69.

WILSON, P. W., 1940.—The biochemistry of symbiotic nitrogen fixation. Univ. Wisc. Press,
Madison.

WRIGHT, W. H., SARLes, W. B., and Houst, E. G., 1930.—A study of Rhizobium japonicum
isolated from various soils. J. Bact., 19:39.

The Honorary Treasurer, Dr. A. B. Walkom, presented the Balance Sheets for the
year ended 28th February, 1954. duly signed by the Auditor, Mr. S. J. Rayment, F.C.A.
(Aust.), and his motion that they be received and adopted was carried unanimously.

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

President: F. V. Mercer, B.Sc., Ph.D.

Members of Council: Lilian Fraser, D.Sc.; Professor P. D. F. Murray, M.A., D.Sc.;
G. D. Osborne, D.Sc., Ph.D.; T. C. Roughley, B.Se., F.R.Z.S.; A. B. Walkom, D.Sc.; and
Professor W. L. Waterhouse, M.C., D.Sc.Agr., D.I.C.

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

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

‘LIANSVAL], “UOTT

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XXXVIi

ABSTRACT OF PROCEEDINGS.

ORDINARY MONTHLY MERTING.

dlst Marcu, 1954.
Dr. F. V. Mercer, President, in the chair.

Library accessions amounting to 64 volumes, 359 parts or numbers, 35 bulletins,
11 reports and 27 pamphlets, total 496, had been received since the last meeting.

PAPERS READ (by title only).
1. Macadamia ternifolia F. Muell., and a Related New Species. By L. A. S. Johnson.

2. Inheritance and the Genetic Relationship of Resistance possessed by two Kenya
Wheats to Races of Puccinia graminis tritici. By D. S. Athwal and I. A. Watson.

ORDINARY MONTHLY MEETING.
28th Aprin, 1954.

Dr. F. V. Mercer, President, occupied the Chair.

The President announced that the Council had elected the following office-bearers for
the 1954-55 session: Vice-Presidents: Mr. D. J. Lee, Mr. A. N. Colefax, Mr. S. J. Copland
and Mr. J. M. Vincent; Honorary Treasurer: Dr. A. B. Walkom; Honorary Secretaries:
Dr. W. R. Browne and Dr. A. B. Walkom.

The following were elected Ordinary Members of the Society: Mr. G. N. Baur,
B.Se.For., Dip.For., Gordon, N.S.W.; Mr. R. F. Black, B.Se., Sydney University; and
Mr. J. B. Williams, B.Sc., Sydney University.

Congratulations were offered to Dr. A. B. Walkom on the award of the Royal
Society’s Medal for 1953.

The President referred to the deaths of Mr. K. G. Brown, who had been elected a
member in 1950, on 7th April, 1954, and of Mr. W. A. W. de Beuzeville, J.P., who had
been a member of the Society since 1925, on 28th March, 1954.

Library accessions amounting to 10 volumes, 120 parts or numbers, and 16 pamphlets,
total 146, had been received since the last meeting.

The papers taken as read at the March Ordinary Monthly Meeting were discussed.

PAPERS READ.

1. A Comparative Account of the Terrestrial Diatoms of Macquarie Island. By J. S.
Bunt.

2. A New Species of Hibbertia Andr., from Western Australia. By A. T. Hotchkiss.

8. Cytological Studies in the Myrtaceae. IV. The Subtribe Euchamaelaucinae. By
S. Smith-White.

4. Experimental Crossing of Aédes (Stegomyia) pseudoscutellaris Theobald and Aédes
(Stegomyia) polynesiensis Marks (Diptera, Culicidae). By A. R. Woodhill.

NOTES AND EXHIBITS.

The President, Dr. F. V. Mercer, described some experiments concerning the pollina-
tion mechanism of the Trigger Plant, Stylidiwm species. The column which bears the
anthers and stigma is sensitive when mature. On stimulation the column snaps rapidly
from one side of the flower to the other and at the same time bends through an angle of
about 270°. After about 3-4 minutes the column gradually returns to the initial position,
but requires about another 4-6 minutes before it regains sensitivity. The process can
be repeated many times without the column tiring.

R

XxXxviili ABSTRACT OF PROCEEDINGS.

ORDINARY MONTHLY MEETING.
26th May, 1954.
Dr. F. V. Mercer, President, occupied the chair.

The President offered congratulations to Dr. T. G. Vallance, on obtaining the
degree of Ph.D.; Mr. E. H. M. Haley, Mrs. Beatrice M. Errey, Miss Olga Kooptzoff and
Mr. A. K. O’Gower, on obtaining the degree of M.Sc.; and Miss Elizabeth C. Pope on
her election as a Corresponding Member of the Zoological Society of London.

Library accessions amounting to 27 volumes, 216 parts or numbers, 1 bulletin and 3
reports, total 247, had been received since the last meeting.

PAPERS READ.

1. The Fossil Diptera of the Tertiary Redbank Plains Series, Queensland. By
K. F. Riek.

2. A Second Specimen of the Dragon-fly, Aeschnidiopsis flindersiensis (Woodward)
from the Queensland Cretaceous. By EH. F. Riek.

3. Notes on the Bacteria belonging to the Rhodobacteriineae Breed, Murray and
Hitchens, and the Chlamydobacteriales Buchanan occurring at Macquarie Island. By
J. S. Bunt.

NOTES AND EXHIBITS.

Mr. A. J. Bearup referred to cases of dermatitis which have been reported as due
to immersion in the salt waters of coastal lakes in the Sydney district. One probable
cause has been found in forktail cercariae which develop in Pyrazus australis, a snail
common on sand flats in estuaries and coastal lakes in New South Wales. This forktail
cercaria is the larval stage of a trematode (family Schistosomatidae) which lives in
blood vessels of vertebrates, especially birds. It is capable of penetrating unbroken
skin. The adult form of this parasite probably lives in water birds such as ducks,
swans and gulls. Four of five silver gulls (Larus novae-hollandiae), which are common
on the coastal lakes, have been found with adult schistosomes of the genus Austrobilharzia.

LECTURETTE.
A lecturette entitled “Hair Growth in Animals’ was given by Dr. A. S. Fraser,
Animal Genetics Unit, C.S.1.R.0O.

ORDINARY MONTHLY MEETING.

30th Jung, 1954.

Dr. F. V. Mercer, President, occupied the chair.

The President offered congratulations to Dr. H. G. Raggatt, who had received the
honour of C.B.H.; and to Mr. P. G. Valder, on the award of the Sir Benjamin Fuller
Travelling Scholarship for further study at Cambridge.

Library accessions amounting to 19 volumes, 183 parts or numbers, 56 bulletins, 6
reports and 14 pamphlets, total 278, had been received since the last meeting.

PAPERS READ.
1. The Genus Theobaldia (Diptera, Culicidae) in Victoria. By N. V. Dobrotworsky.
2. The Inheritance of Inflorescence Characters in Hucalyptus. By L. D. Pryor.

LECTURETTE.
A lecturette entitled ‘‘The Cosmic Radiation” was given by Dr. A. J. Herz.

ORDINARY MONTHLY MEETING.
28th Juny, 1954.

Dr. F. V. Mercer, President, occupied the chair.
Mr. Robert Domrow, Brisbane, Queensland, was elected an Ordinary Member of

the Society.

ABSTRACT OF PROCEEDINGS. XXX1xX

The President referred to the death, on 10th July, 1954, of Mr. Ernest G. Jacobs,
who had been a member of the Society since 1917. The President also referred to the
death, in London, of Professor J. P. Hill, who had been an Honorary Member of the
Society since 1923.

The President announced that Dr. N. C. W. Beadle had been elected a member
of Council in place of Professor J. L. Still.

The President offered congratulations to Dr. Mary Hindmarsh on obtaining the
degree of Ph.D. of the University of Sydney.

Library accessions amounting to 7 volumes, 72 parts or numbers, 1 bulletin, 2 reports
and 14 pamphlets, total 96, had been received since the last meeting.

PAPERS READ.
1. Collenia frequens in Upper Proterozoic Rocks in the Northern Territory of
Australia. By D. M. Traves. (Communicated by Dr. Germaine A. Joplin.)

2. Notes on Australian Beetles in the Tribe Bolboceratini formerly in the Genus
Bolboceras. By Henry F. Howden. (Communicated by Dr. P. B. Carne.)

38. Australian Fungi. II. New Records and Revisions. By C. G. Hansford.

LECTURETTE.
A lecturette entitled ‘‘Let’s Talk Turkey” was given by Dr. A. T. Hotchkiss.

ORDINARY MONTHLY MEETING.
25th AueustT, 1954.

Dr. F. V. Mercer, President, occupied the chair.

The President offered congratulations to Dr. N. C. W. Beadle and Dr. A. H. Voisey
on their appointment to the Chairs of Biology and Geology respectively in the University
of New England.

The President announced that Subscribers’ Day and Reunion will be held at
.“Muogamarra” Sanctuary on 11th September, 1954.

Library accessions amounting to 19 volumes, 102 parts or numbers, 37 bulletins, 8
reports and 31 pamphlets, total 197, had been received since the last meeting.

PAPERS READ.

1. Notes on the Scleractinia or Stony Corals (Coelenterata) of Heron Island,
Queensland. -I. A List of the Common Species. By K. HE. W. Salter.

2. Antirrhinum Rust, Puccinia antirrhini D. & H., in Australia. By J. Walker.

NOTES AND EXHIBITS.

Dr. W. R. Browne (Hon. Secretary) exhibited (on behalf of the Society) fourteen
water-colour paintings of native plants painted by, and bequeathed to the Society by,
the late Mr. P. C. W. Shaw, a retired engineer, who died on 8th me cues 1944, at
the age of 84 years, and who had been a member of the Society since 1941.

Dr. G. D. Osborne exhibited specimens and photographs of veins of asbestos (Chryso-
tile) in serpentine from Woodsreef, east of Barraba, showing ribbon structure. From
field study he suggested the probability that the ribbon fibre had developed largely
as a result of spheroidal weathering in the serpentine near the surface of the ground.

Miss K. English exhibited shield coverings of the curious elongated larvae of a
species of scale insect (Coccidae) found on Casuarina at Bantry Bay, Middle Harbour.

Mr. D. K. McAlpine exhibited eggs, larvae, pupae and adults of the fly, Batracho-
myia, which is parasitic under the skin of certain Australian frogs. Very little HS known
of the bionomics and structure of this fly. —

xl ABSTRACT OF PROCEEDINGS.

Dr. Lilian Fraser exhibited a small plant of orange on Seville root stock which
was affected with the aphid-transmitted virus disease, tristeza. This is one of the major
diseases of modern times. It affects oranges on Seville stock, but not trees on rough
lemon, sweet orange or trifoliata rootstocks. Trees on these stocks can carry the virus
symptomlessly. During the last 20 years tristeza has been responsible for the death of
more than 9,000,000 citrus trees in South America and it is at present spreading through
California. It has been present in New South Wales for at least 80 years, but because
the stocks in general use are rough lemon and trifoliata it has no economic importance.

Mr. J. Walker exhibited a specimen of the fungus Podaris pistillaris (lL. ex Pers.)
Morse collected at Bourke, N.S.W. It is a member of the Order Lycoperdales, Family
Tulostomataceae, the main characteristic of which is that the peridium is carried upon
a well developed stem, which in Podawzis traverses the gleba as a columella. The organism
is an interesting one in that it has been described as 32 different species, indicating
its variability. The genus is sometimes known as Podazxon, but the name Podazis has
priority and should be used.

Mr. J. T. Waterhouse exhibited a series of 1,097 original water-colour drawings
of British Fungi by Edwin Wheeler (1833-1909), a homoeopathic druggist of Bristol.
The drawings are of high artistic merit and scientific accuracy and should be of value
for reference in taxonomic work on our own species. They are bound in five albums
and belong to the Fisher Library, University of Sydney.

ORDINARY MONTHLY MEETING.
29th SEPTEMBER, 1954.

Dr. F. V. Mercer, President, occupied the chair.

Mr. G. W. Douglas, Strathmore, Victoria, was elected an Ordinary Member of the
Society.

The President announced that the Council is prepared to receive applications for
Linnean Macleay Fellowships tenable for one year from ist January, 1955, from
qualified candidates. The range of actual salary is £650-£80C, according to qualifications.
Applications should be lodged with the Hon. Secretary not later than Wednesday,
3rd November, 1954.

The President offered congratulations to Dr. D. S. Simonett on obtaining the degree
of Ph.D. of the University of Sydney for a thesis on ‘The Status of Land Utilization
in North Queensland”.

Library accessions amounting to 5 volumes, 78 parts or numbers, 4 bulletins, 2
reports and 2 pamphlets, total 91, had been received since the last meeting.

PAPERS READ.
1. A Second Species of Holothyrus (Acarina: Holothyroidea) from Australia. By
R. Domrow. (Communicated by Dr. I. M. Mackerras.)
2. Acarine Parasites of Two Species of Rattus from Brisbane, Australia. By R.
Domrow. (Communicated by Dr. I. M. Mackerras.)
3. A New Coptotermes and Ahamitermes (Isoptera) from Australia. By F. J. Gay.
(Communicated by Dr. A. J. Nicholson.)

4. The Nature and Significance of Non-reciprocal Fertility in Aédes scutellaris and
Other Mosquitoes. By S. Smith-White and A. R. Woodhill.

NOTES AND EXHIBITS,

Mr. G, H. Hardy exhibited larvae of the family Asilidae (Robber-flies) taken in
garden soil at Katoomba.
LECTURETTE.

A lecturette on “The Biosynthesis of Rubber” was delivered by Dr. Adele Millerd.

ABSTRACT OF PROCEEDINGS. xli

ORDINARY MONTHLY MEETING.
27th Octoper, 1954.
Dr. F. V. Mercer, President, occupied the chair.

Miss Mary B. Macdonald, B.Se., Sydney University, was elected an Ordinary
Member of the Society.

The President announced that the Council is prepared to receive applications for
Linnean Macleay Fellowships tenable for one year from ist January, 1955, from
qualified candidates. The range of actual salary is £650—£800, according to qualifications.
Applications should be lodged with the Hon. Secretary not later than Wednesday,
3rd November, 1954.

Library accessions amounting to 6 volumes, 77 parts or numbers, 1 bulletin and 3
reports, total 87, had been received since the last meeting.

PAPERS READ.

1. The Culex pipiens Group in South-eastern Australia. III. Autogeny in Culex
pipiens form molestus. By N. V. Dobrotworsky.

2. Supplementary Notes on the Genus Brachycome Cass.—Descriptions of Five New
Australian Species and Some New Locality Records. By Gwenda L. Davis.

3. Notes and Descriptions of Australian Chloropidae (Diptera). By C. W. Sabrosky.
(Communicated by D. J. Lee.)

4. Notes on the Morphology and Biology of Scaptia vicina Tayl., and a New Species
of Scaptia (Diptera, Tabanidae). By Kathleen M. I. English.

NOTES AND EXHIBITS.
Dr. N. C. Beadle exhibited a white Waratah, which came from a plant growing on
Hawkesbury Sandstone in the Hunter Valley near the road from Wiseman’s Ferry to
Singleton.
Dr. W. R. Browne exhibited some coloured slides showing the effects of bushfires
on the vegetation of the Kosciusko district.

LECTURETTE.
A lecturette entitled “Plants, Mountains and Laboratories’ was delivered by Dr.
Clark Ashby, Fulbright Fellow in Botany.

ORDINARY MONTHLY MEETING.
24th NovemMsBeEr, 1954.

Mr. J. M. Vincent, Vice-President, occupied the chair.

Miss Beryl M. Cameron, B.Sc., Concord, N.S.W., was elected an Ordinary Member
of the Society.

The Chairman announced that Miss Nola J. Hannon, B.Sc., and Miss Mary B.
Macdonald, B.Sc., have been re-appointed and appointed respectively to Linnean Macleay
Fellowships in Botany for the year 1955. ;

Library accessions amounting to 10 volumes, 225 parts or numbers, 13 bulletins, 1
report and 1 pamphlet, total 250, had been received since the last meeting.

PAPERS READ.

1. A Note on the Occurrence of “Anomalous” Krasnozem in the Richmond—Tweed
Region of N.S.W. By J. W. McGarity and D. N. Munns. (Communicated by Dr. W. R.
Browne.)

2. The Influence of the Physical Properties of a Water Container Surface upon
its Selection by the Gravid Females of Aédes scutellaris scutellaris (Walker) for
Oviposition (Diptera, Culicidae). By A. K. O’Gower.

3. An Fl Hybrid between Hucalyptus cinerea F. Muell, and Hucalyptus robusta Sm,
By L, D. Pryor,

xlii ABSTRACT OF PROCEEDINGS.

4. A New Genus and Species of the Tribe Labenini from Australia (Pimplinae,
Ichneumonidae). By A. W. Parrott.

5. Australasian Ceratopogonidae (Diptera, Nematocera). Part VII. Notes on the
Genera Alluaudomyia, Ceratopogon, Culicoides and Lasiohelea. By David J. Lee and
Eric J. Reye.

NOTES AND EXHIBITS.

Mr. A. J. Bearup exhibited schistosomes of the genus Microbilharzia which had
developed in a budgerigar (Melopsittacus) as a result of experimental infection with
cercariae from Pyrazus australis. The worms were present in the liver and intestinal
veins 22 days after infection. Eggs with miracidia were present in the intestinal wall
and in the faeces. The cercariae were the same as those reported (at the May meeting
of this Society) as the probable cause of “bathers’ itch’ in coastal lakes of New South
Wales.

LECTURETTE.

A lecturette entitled “Field Work in the New Guinea Highlands” was given by
Ellis Troughton, Curator of Mammals, Australian Museum.

1940

1927
1940

1$22
1927

1952
1912

1952

1951

1948

1952

1949
1950

1954
1935
1946
1940

1952
1907

1948
1948

1954
1941
1929
1946
1924
1949
1911
1952
1949
1931

1920

1927

1934

1949

1936

1947

xlili

LIST OF MEMBERS.
(15th December, 1954.)

ORDINARY MBMBERS.
(An asterisk (*) denotes Life Member.)

Abbie, Professor Andrew Arthur, M.D., B.S., B.Sc., Ph.D., c.o. University of Adelaide,
Adelaide, South Australia.

*Albert, Michel Francois, ‘‘Boomerang”’’, 42 Billyard Avenue, Blizabeth Bay, Sydney.

*Allman, Stuart Leo, B.Sc.Agr., M.Sc., Entomological Branch, Department of Agriculture,
Farrer Place, Sydney.

Anderson, Robert Henry, B.Sc.Agr., Botanic Gardens, Sydney.

*Armstrong, Jack Walter Trench, ‘‘Callubri”’, Nyngan, N.S.W. °

Ashton, David Hungerford, B.Sc., 92 Warrigal Road, Surrey Hills, E.10, Victoria.

Aurousseau, Marcel, B.Se., No. 15 Hollycroft Avenue, London, N.W.3, England.

Baas-Becking, L. G. M., Ph.D., D.Sc, C.S.I.R.O., Division of Fisheries, P.O. Box 21,
Cronulla, N.S.W.

Backhouse, Thomas Clive, M.B., B.S., D.P.H., D.T.M. & H., F.R.A.C.P., School of Public
Health and Tropical Medicine, Sydney University.

Baddams, Miss Greta, B.A., B.Sc., Faculty of Science, The University of New England,
Armidale 5N, N.S.W.

Baehni, Professor Charles, Dr.sc., Conservatoire botanique, Université de Genéve, 192,
rue de Lausanne, Genéve, Switzerland.

Baker, Eldred Percy, B.Sc.Agr., Ph.D., Faculty of Agriculture, Sydney University.

*Barber, Professor Horace Newton, M.A., Ph.D., Department of Botany, University of
Tasmania, Hobart, Tasmania.

Baur, George Norton, B.Se.Agr., Dip.For., 24 Bushlands Avenue, Gordon, N.S.W.

*Beadle, Noel Charles William, D.Sc., Botany School, Sydney University.

Bearup, Arthur Joseph, 66 Pacific Avenue, Penshurst, N.S.W.

Beattie, Joan Marion, D.Sc. (née Crockford), c.o. Lake George Mine, Captain’s Flat,
N.S.W.

Bennett, Miss Isobel Ida, Department of Zoology, Sydney University.

Benson, Professor William Noel, B.A., D.Sc., 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, D.Ag.Se., M.Se., Zoology Department, Columbia University, New
York 27, N.Y., U.S.A.

Black, Roger Foster, B.Sc., Department of Botany, Sydney University.

Blake, Stanley Thatcher, M.Sc., Botanic Gardens, Brisbane, Queensland.

Boardman, William, M.Sc., Zoology Department, University of Melbourne, Carlton, N.3,
Victoria.

Brett, Robert Gordon Lindsay, B.Sc., 7 Petty Street, West Hobart, Tasmania.

Browne, Ida Alison, D.Sc. (née Brown), Department of Geology, Sydney University.

Browne, Lindsay Blakeston Barton, Department of Zoology, Sydney University.

Browne, William Rowan, D.Sc., Department of Geology, Sydney University.

Bunt, John Stuart, B.Sec.Agr., Faculty of Agriculture, Sydney University.

Burden, John Henry, 1 Havilah Street, Chatswood, N.S.W.

*Burges, Professor Norman Alan, M.Sc., Ph.D., Professor of Botany, University of Liver-
pool, Liverpool, England.

Burkitt, Professor Arthur Neville St. George Handcock, M.B., B.Sc., Medical School,
Sydney University.

Campbell, Thomas Graham, Division of Economic Entomology, C.S.I.R.O., P.O. Box 109,
City, Canberra, A.C.T.

*Carey, Professor Samuel Warren, D.Sc., Geology Department, University of Tasmania,
Hobart, Tasmania.

Carne, Phillip Broughton, B.Agr.Sci. (Melb.), Ph.D. (London), D.I.C., C.S.I.R.O., Division
of Entomology, P.O. Box 109, City, Canberra, A.C.T.

*Chadwick, Clarence Earl, B.Sc., Entomological Branch, Department of Agriculture, Farrer
Place, Sydney.

Christian, Stanley Hinton, Malaria Control, Department of Public Health, Banz, Western
Highlands, via Lae, New Guinea.

xliv

1932
1946

1947
1901

1931
1946
1942
1947
1908

LIST OF MEMBERS,

*Churchward, John Gordon, B.Se.Agr., Ph.D., 1 Hunter Street, Woolwich, N.S.W.

Clark, Laurance Ross, M.Sc., ¢.o. C.S.I.R.O., Division of Entomology, P.O. Box 109, City,
Canberra, A.C.T.

Clarke, Mrs. Muriel Catherine, M.Sc (née Morris), 122 Swan Street, Morpeth, N.S.W.

Cleland, Professor John Burton, M.D., Ch.M., 1 Dashwood Road, Beaumont, Adelaide.
South Australia.

Colefax, Allen Neville, B.Se., Department of Zoology, Sydney University.

Colless, Donald Henry, 9 Eng Neo Avenue, Singapore 21, Malaya.

Copland, Stephen John, M.Se., Chilton Parade, Warrawee, N.S.W.

Costin, Alec Baillie, 21 Beverley Street, East Doncaster, Victoria.

Cotton, Professor Leo Arthur, M.A., D.Se., 113 Queen’s Parade East, Newport Beach,
N.S. W.

Crawford, Lindsay Dinham, B.Sc., 4 Dalton Avenue, West Hobart, Tasmania.

Davis, Mrs. Gwenda Louise, Ph.D., B.Sc., Faculty of Science, The University of New
England, Armidale 5N, N.S.W.

Day, William Eric, 23 Gelling Avenue, Strathfield, N.S.W.

Deuquet, Camille, B.Com., 126 Hurstville Road, Oatley, N.S.W.

Dobrotworsky, Nikolai V., M.Sce., Department of Zoology, University of Melbourne,
Carlton, N.3, Victoria.

Domrow, Robert, Queensland Institute of Medical Research, Herston Road, Valley,
Brisbane, Queensland.

Douglas, Geoffrey William, B.Agr.Sci., 3 Columban Avenue, Strathmore W6, Victoria.

Drover, Donald P., Institute of Agriculture, University of Western Australia, Nedlands,
W.A.

Durie, Peter Harold, M.Sc., C.S.I.R.O., Veterinary Parasitology Laboratory, Yeerongpilly.
Brisbane, Queensland.

Dyce, Alan Lindsay, B.Sc.Agr., C.S.I.R.O., Division of Entomology, P.O. Box 109, City,
Canberra, A.C.T.

Ealey, Eric H. M., M.Se., Girrawheen, Duffy Avenue, Thornleigh, N.S.W.

Edwards, Dare William, B.Sc.Agr. (Hons.), Forestry Commission of N.S.W., Division
of Wood Technology, 96 Harrington Street, Sydney.

Elliott, John Henry, ‘‘Greengates’’, Pennant Hills Road, Beecroft, N.S.W.

Eindean, Robert, M.Sce., Department of Zoology, University of Queensland, Brisbane,
Queensland.

English, Miss Kathleen Mary Isabel, B.Sec., 2 Shirley Road, Roseville, N.S.W.

Errey, Mrs. Beatrice Mary, M.Sce., 34 Rosedale Avenue, Fairlight, N.S.W.

Frame, William Robert, c/- G. G. Smith & Co. Ltd., P.O. Box 17, Port Moresby, Papua-
New Guinea.

Fraser, Ian McLennan, Ph.D. (Cambridge), School of Medicine, College of Medical
Evangelists, Loma Linda, California, U.S.A.

Fraser, Miss Lilian Ross, D.Sc., ‘““Hopetoun”, 25 Bellamy Street, Pennant Hills, N.S.W.

Garden, Miss Joy Gardiner, B.Sc.Agr., Botanic Gardens, Sydney.

*Garretty, Michael Duhan, D.Sc., “Surrey Lodge’, Mitcham Road, Mitcham, Victoria.

Greenwood, William Frederick Neville, 11 Wentworth Avenue, Waitara, N.S.W.

*Griffiths, Mrs. Mabel, B.Sc. (née Crust), 2 Carden Avenue, Wahroonga, N.S.W.

Griffiths, Mervyn Edward, M.Se., Australian Institute of Anatomy, Canberra, A.C.T.

*Gunther, Carl Ernest Mitchelmore, M.B., B.S., D.T.M., D.T.M. & H. (England), Bulolo
Gold Dredging Limited, Bulolo, Territory of Papua—New Guinea.

Hamilton, Edgar Alexander, 16 Hercules Street, Chatswood, N.S.W.

Hannon, Miss Nola Jean, B.Sc., 22 Leeder Avenue, Penshurst, N.S.W.

*Hansford, Clifford Gerald, M.A., Se.D. (Cantab.), D.Se. (Adel.), F.L.S., Waite Agricultural
Research Institute, Private Bag, G.P.O., Adelaide, South Australia.

Hardy, George Huddleston Hurlstone, ‘““Karambi’’, Letitia Street, Katoomba, N.S.W.

Harker, Miss Janet HElspeth, M.Se., Ph.D., F.R.E.S., Sub-Department of Entomology,
Department of Zoology, University, Cambridge, England.

Hewitt, Bernard Robert, 4 Hillview Street, Lakemba, 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.Sc, Ph.D., Department of Geology, University of Queensland,
Brisbane, Queensland. opine’

Hindmarsh, Miss Mary Maclean, B.Sc., Ph.D., 79 Onslow Street, Rose Bay, 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.

LIST OF MEMBERS. xlv

Horowitz, Benzoin, Eng.Agr.S., Dr.Agr.Sc. (Cracow, Poland), CS3ER-O;, ‘cio; Waite
Institute, Private Mail Bag, Adelaide, South Australia.

Hossfeld, Paul Samuel, M.Sc., 132 Fisher Street, Fullarton, South Australia.

Hotchkiss, Arland Tillotson, M.S., Ph.D. (Cornell), Department of Botany, Sydney
University.

Humphrey, George Frederick, M.Sc., Ph.D., Department of Biochemistry, Sydney
University.

Jacobs, Maxwell Ralph, D.Ing., M.Se., Dip.For., Australian Forestry School, Canberra,
A.C.T.

Jessup, Rupert William, M.Sc., 38 Taylor Street, Armidale, N.S.W.

Johnson, Lawrence Alexander Sidney, B.Sc., c.o. National Herbarium, Botanic Gardens,
Sydney.

Johnston, Arthur Nelson, B.Sc.Agr., 99 Newton Road, Strathfield, N.S.W.

Jones, Mrs. Valerie Margaret Beresford, M.Sc. (née May), Mooloolabel Esplanade,
Narrabeen, N.S.W. E

Joplin, Miss Germaine Anne, B.A., Ph.D., D.Se., Department of Geophysics, Australian
National University, Canberra, A.C.T.

Judge, Leslie Arthur, 87 Eastern Road, Turramurra, N.S.W.

Keast, James Allen, M.Sc., Australian Museum, College Street, Sydney.

Kerr, Harland Benson, B.Sc.Agr., 41 Badminton Road, Croydon, N.S.W.

Kesteven, Geoffrey Leighton, D.Sec., c.o. F.A.O. of United Nations, Viale delle Terme di
Caracalla, Rome, Italy.

Kesteven, Hereward Leighton, D.Sc., M.D., Maroochydore, Queensland.

Kinghorn, James Roy, C.M.Z.S., Australian Museum, College Street, Sydney.

Langford-Smith, Trevor, M.Se., School of Pacific Studies, Australian National University,
Canberra, A.C.T.

Langley, Miss Julia Mary, B.Sc., 17 Clifford Street, Gordon, N.S.W.

Larcombe, Miss Pauline Gladys, B.Sc., 17 Ethel Street, Burwood, N.S.W.

Latter, Barrie Dale Hingston, B.Sc.Agr., 25 Abbott Street, Coogee, N.S.W.

*Lawrence, James Joscelyn, B.Sc., School of Public Health and Tropical Medicine, Sydney
University.

Lawson, Albert Augustus, 9 Wilmot Street, Sydney.

Lee, Mrs. Alma Theodora, M.Se. (née Melvaine), Manor Road, Hornsby, N.S.W.

Lee, David Joseph, B.Sc., School of Public Health and Tropical Medicine, Sydney
University. 4

Lothian, Thomas Robert Noel, Botanic Gardens, Adelaide, South Australia.

Lower, Harold Farnham, Waite Agricultural Research Institute, University of Adelaide,
Private Mail Bag, G.P.O., Adelaide, South Australia.

Lowery, Edwin Sanders, M.A., 13 Francis Street, Northmead, N.S.W.

Macdonald, Colin Lewis, High School, Goulburn 2S, N.S.W.

Macdonald, Miss Mary Beth, B.Sc., Department of Botany, Sydney University. :

Macintosh, Neil William George, M.B., B.S., 32 Benelong Crescent, Bellevue Hill, N.S.W.

Mackerras, Ian Murray, M.B., Ch.M., B.Sc., Queensland Institute of Medical Research,
Herston Road, Valley, Brisbane, Queensland.

*Mair, Herbert Knowles Charles, B.Sc., Botanic Gardens, Sydney.

Marks, Miss Elizabeth Nesta, M.Se., Ph.D., Department of Entomology, University of
Queensland, Brisbane, Queensland.

Martin, Mrs. Hilda Ruth Brownell, B.Sc. (née Simons), 36 Spencer Road, Killara, N.S.W.

Mawson, Sir Douglas, D.Se., B.H., F.R.S., University of Adelaide, Adelaide, South
Australia. :

Maze, Wilson Harold, M.Se., University of Sydney.

McAlpine, David Kendray, 12 St. Thomas Street, Bronte, N.S.W.

McCulloch, Robert Nicholson, D.Se.Agr., B.Se., Roseworthy Agricultural College, Rose-
worthy, South Australia.

McDonald, Miss Patricia M., B.Sc., Dip.Ed., 29 Dee Why Parade, Dee Why, N.S.W.

McKee, Hugh Shaw, B.A., D.Phil. (Oxon.), South Pacific Commission, P.O. Box 9,
Noumea, New Caledonia.

McMichael, Donald Fred, B.Sc., Australian Museum, College Street, Sydney.

McMillan, Bruce, M.B., B.S., 5 Windmill Lane, Bushey Heath, Watford, Hertfordshire,
England. - c

Mercer, Frank Verdun, B.Sc., Ph.D. (Cambridge), Botany School, Sydney University.

Messmer, Mrs. Pearl Ray, 64 Treatts Road, Lindfield, N.S.W.

*Meyer, George Rex, B.Sc., Dip-Ed., Department of Zoology, Sydney University.

*Miller, Allen Horace, B.Sc., Dip.Ed., 46a Wentworth Street, Wallsend 2N, 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.Sc., Ph.D., Department of Biochemistry, Sydney University.

xlvi LIST OF MEMBERS.

1947 Millett, Mervyn Richard Oke, B.A., 19 Avoca Street, South Yarra, Vic.

1946 Millington, Richard James, Waite Agricultural Research Institute, Private Mail Bag,
Adelaide, South Australia.

1949 Minter, Philip Clayton, 49 Cotswold Road, Strathfield, N.S.W.

1952 Monro, John Malcolm, University of New England, Armidale 5N, N.S.W.

1944 Moye, Daniel George, B.Sc., Dip.Ed., 6 Kaling Place, Cooma, N.S.W.

1939 Moye, Mrs. Joan, B.Se. (née Johnston), 6 Kaling Place, Cooma, N.S.W.

1926 Mungomery, Reginald William, c.o. Bureau of Sugar Experiment Stations, Department
of Agriculture and Stock, Brisbane, B.7, Queensland.

1949 Murray, Professor Patrick Desmond Fitzgerald, M.A., D.Se., Department of Zoology,
University of Sydney, N.S.W.

1920 Musgrave, Anthony, F.R.E.S., Australian Museum, College Street, Sydney.

1949 Myers, Kenneth, C.S.I.R.O., Wildlife Survey Section, Field Station, 598 Affleck Street,
Albury, N.S.W.

1947 Nashar, Mrs. Beryl, B.Se., Ph.D., Dip.Ed. (née Scott), c/- 23 Morris Street, Mayfield
West 2N, N.S.W.

25 Newman, Ivor Vickery, M.Sc., Ph.D., F.R.M.S., F.L.S., 1 Stuart Street, Wahroonga, N.S.W.

22 Nicholson, Alexander John, D.Sc., F.R.E.S., C.S.I.R.O., Box 109, Canberra, A.C.T.

5 *Noble, Norman Scott, D.Se.Agr., M.Sec., D.1.C., C.S.I.R.O., 314 Albert Street, East
Melbourne, C.2, Victoria.

1920 Noble, Robert Jackson, B.Sce.Agr., Ph.D., 324 Middle Harbour Road, Lindfield, N.S.W.

1912 North, David Sutherland, 42 Chelmsford Avenue, Lindfield, N.S.W.

1948 O’Farrell, Antony Frederick Louis, A.R.C.Sc., B.Se., F.R.E.S., University of New England,
Armidale 5N, N.S.W.

1950 O’Gower, Alan Kenneth, M.Sc., 59 Brighton Street, Harbord, 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., 6 Magdalen Street, College
Park, South Australia.

1921 Osborne, George Davenport, D.Sc., Ph.D., Department of Geology, Sydney University.

1950 Oxenford, Reginald Augustus, B.Sc., 10 Fry Street, Grafton 3C, N.S.W.

1952 Packham, Gordon Howard, 61 Earlwood Avenue, Harlwood, N.S.W.

1952 Palmer, Rev. Robert George, Box 13, P.O., Glen Davis, 6W, N.S.W.

1954 Parrott, Arthur Wilson, ‘‘Lochiel’, Wakapuaka Road, Hira, R.D., Nelson, New Zealand.

1940 *Pasfield, Gordon, B.Sc.Agr., 20 Cooper Street, Strathfield, N.S.W.

1922 Perkins, Frederick Athol, B.Sc.Agr., Department of Entomology, University of Queensland,
Brisbane, Queensland.

1947 Phillips, Miss Marie Elizabeth, M.Se., Ph.D., 4 Morella Road, Clifton Gardens, N.S.W.

1937 Plomley, Kenneth Francis, ¢c.o. Division of Forest Products, C.S.I.R.O., 69 Yarra Bank
Road, South Melbourne, S.E.1, Victoria.

1985 Pope, Miss Elizabeth Carington, M.Sc., C.M.Z.S., Australian Museum, College Street,
Sydney.

1938 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., 19 Hampden Avenue, Cremorne, N.S.W.

1929 Raggatt, Harold George, D.Sc., 60 Arthur Circle, Forrest, Canberra, A.C.T.

1951 Ralph, Bernhard John Frederick, B.Sc., Ph.D. (Liverpool), A.A.C.I., N.S.W. University
of Technology, Broadway, Sydney.

1952 Ramsay, Mrs. Helen Patricia, M.Se. (née Lancaster), Cnr. Ridge and Sofala Streets,
Portland 6W, N.S.W.

1953 Reye, Eric James, M.B., B.S. (Univ. Qld.), c/- C.S.I.R.O., Veterinary Parasitology
Laboratory, Fairfield Road, Yeerongpilly, Queensland.

1950 Rickwood, Frank Kenneth, B.Sc., Department of Geology, Sydney University.

1946 Riek, Edgar Frederick, B.Sc., C.S.I.R.O., Division of EHconomic Entomology, P.O. Box
109, City, Canberra, A.C.T.

1986 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.O., Private Mail Bag, Homebush, N.S.W.

1245 Ross, Donald Ford, ¢.o. Ross Bros. Pty. Ltd., 545-547 Kent Street, Sydney.

1925 Roughley, Theodore Cleveland, B.Sc., F.R.Z.S., 5 Coolong Road, Vaucluse, N.S.W.

1932 Salter, Keith Eric Wellesley, B.Sc., ‘“Hawthorn’’, 48 Abbotsford Road, Homebush, N.S.W.
1919 *Scammell, George Vance, B.Se., 7 David Street, Clifton Gardens, N.S.W.

1951 Seccombe, Mrs. Lorraine, B.A., 28 Fairweather Street, Bellevue Hill, N.S.W.

1950 *Sharp, Kenneth Raeburn, B.Se., Eng. Geology, S.M.H.E.A., Cooma, 4S, N.S.W.

1948
1930
1947
1953

1952
1953
1916
1943
1945
1937

1926

1898
1932
1949
1935
1952
1911

1940

1950
1949

1944
1943
1946
1921
1950

1952

1949
1917

1930
1940
1934

1952

1909
1946

1947
1930
Pala
1936
1947
1927

L941
1949
1946

1953
1926
1946
1954
1952
1950
1947

LIST OF MEMBERS. xlvii-

Shaw, Miss Dorothy Edith, M.Se.Agr., 119 Bellevue Parade, Hurstville, N.S.W.

Sherrard, Mrs. Kathleen Margaret, M.Sc., 43 Robertson Road, Centennial Park, Sydney.

Shipp, Erik, 59 William Edward Street, Longueville, N.S.W.

Simonett, David Stanley, M.Sc., Department of Geography, The University, Nottingham,
England.

Slade, Milton John, B.Sc., 10 Elizabeth Street, Raymond Terrace, N.S.W.

Smith, Eugene Thomas, 22 Talmage Street, Sunshine, Victoria.

Smith, Miss Vera Irwin, B.Sc., F.L.S., ‘“Loana’’, Mt. Morris Street, Woolwich, N.S.W.

Smith-White, Spencer, B.Se.Agr., 15 Berowra Road, Mt. Colah, N.S.W.

Southeott, Ronald Vernon, M.B., B.S., 13 Jasper Street, Hyde Park, South AustraJia.

Spencer, Mrs. Dora Margaret, M.Sc. (née Cumpston), c/- Public Health Department,
Minj, Western Highlands, New Guinea.

Stanley, George Arthur Vickers, B.Sc., c.o. Messrs. Robison, Maxwell and Allen, 19 Bligh
Street, Sydney.

Stead, David G., ‘“Boongarre’’, 14 Pacific Street, Watson’s Bay, N.S.W.

Stead, Mrs. Thistle Yolette, B.Sc. (née Harris), 14 Pacific Street, Watson’s Bay, N.S.W.

Stevens, Neville Cecil, B.Sc., 12 Salisbury Street, Hurstville, N.S.W.

Still, Professor Jack Leslie, B.Sc., Ph.D., Department of Biochemistry, Sydney University.

Sullivan, George Emmerson, M.Se. (N.Z.), Department of Zoology, Sydney University.

*Sulman, Miss Florence, 11 Ramsay Street, Collaroy, N.S.W. :

Taylor, Keith Lind, B.Sc.Agr., c.o. Division of Entomology, C.S.I.R.O., Box 109, City,
Canberra, A.C.T.

Tchan, Yao-tseng, Dr., és Sciences (Paris), Botany School, Sydney University.

Thorp, Mrs. Dorothy Aubourne, B.Se. (Lond.), ‘“Carinya’”’, Tara Street, Kangaroo Point,
Sylvania, N.S.W.

Thorpe, Ellis William Ray, B.Sc., University of New England, Armidale 5N, N.S.W.

Tindale, Miss Mary Douglas, M.Se., 60 Spruson Street, Neutral Bay, N.S.W.

Tipper, John Duncan, A.M.I.E.Aust., Box 2770, G.P.O., Sydney.

*Troughton, Ellis Le Geyt, C.M.Z.S., F.R.Z.S., Australian Museum, College Street, Sydney.

Tugby, Mrs. Elise Evelyn, B.Se. (née Sellgren), Department of Anthropology and
Sociology, Australian National University, Box 4, G.P.O., Canberra, A.C.T.

Valder, Peter George, B.Sc.Agr., c/- EH.S. & A. Bank, Ltd., West End Branch, 8-12 Brook
Street, London, W.1, England.

Vallance, Thomas George, B.Sc., Ph.D., 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.Se., Botanic Gardens, Sydney.

Vincent, James Matthew, B.Sc.Agr., Dip.Bact., Faculty of Agriculture, Sydney University.

*Voisey, Professor Alan Heywood, D.Sc., University of New England, Armidale 5N,
N.S. W.

Walker, John, B.Sc.Agr., Biological Branch, N.S.W. Department of Agriculture, Box 36,
G.P.O., Sydney.

Walkom, Arthur Bache, D.Se., 45 Nelson Road, Killara, N.S.W.

Wallace, Murray McCadam Hay, B.Sc., Institute of Agriculture, University of Western
Australia, Nedlands, Western Australia.

Ward, Mrs. Judith, B.Sc., Lilac Cottage, Invergarry, Inverness-shire, Scotland.

Ward, Melbourne, Gallery of Natural History and Native Art, Medlow Bath, N.S.W.

Wardlaw, Henry Sloane Halcro, D.Sc., F.R.A.C.1., 71 McIntosh Street, Gordon, N.S.W.

Waterhouse, Douglas Frew, D.Sce., C.S.I.R.O., Box 109, Canberra, A.C.T.

«Waterhouse, John Teast, B.Se., Department of Botany, Sydney University.

Waterhouse, Professor Walter Lawry, D.Sc.Agr., M.C., D.I.C., 30 Chelmsford Avenue,
Lindfield, N.S.W.

Watson, Irvine Armstrong, Ph.D., B.Sc.Agr., Faculty of Agriculture, Sydney University.

Whaite, Mrs. Joy Lilian, c/- Department of Main Roads, Deniliquin 6S, N.S.W.

Wharton, Ronald Harry, B.Sc., Institute for Medical Research, Branch lLaboratory,,.
Kuantan, Pahang, Federation of Malaya.

Whitehouse, Miss Jill Armson, B.Sc., 8 Francis Street, Lindfield, N.S.W.

*Whitley, Gilbert Percy, Australian Museum, College Street, Sydney.

Wilkins, Miss Marjorie Jessie, M.Se., 33 Muston Street, Mosman, N.S.W.

Williams, John Beaumont, B.Sc., Department of Botany, Sydney University.

Williams, Owen Benson, M.Agr.Se. (Melbourne), 47 George Street, Deniliquin, N.S.W.

Willis, Jack Lehane, M.Sc., A.A.C.I., 26 Inverallan Avenue, Pymble, N.S.W.

Winkworth, Robert Ernest, P.O. Box 77, Alice Springs, Northern Territory, Australia

1952

LIST OF MEMBERS.

Womersley, Herbert, F.R.E.S., A.L.S., South Australian Museum, Adelaide, South
Australia.

Wood, Edward James Ferguson, C.S.I.R.O., Marine Biological Laboratory, P.O. Box 21,
Cronulla, N.S.W.

Woodhill, Anthony Reeve, D.Sec.Agr., Department of Zoology, Sydney University.

Zeck, Emil Herman, F.R.Z.S., 694 Victoria Road, Ryde, N.S.W.
Zeck, Mrs. Nance (Anne), 694 Victoria Road, Ryde, N.S.W.

CORRESPONDING MEMBERS.

Jensen, Hans Laurits, D.Se.Agr. (Copenhagen), State Laboratory of Plant Culture,
Department of Bacteriology, Lyngby, Denmark.
Rupp, Rev. Herman Montague Rucker, B.A., 32 Neville Street, Willoughby, N.S.W.

ASSOCIATE MEMBER.
Seccombe, John Edwin, 28 Fairweather Street, Bellevue Hill, N.S.W.

xlix

LISTS OF NEW GENERA, SUBGENERA, SPECIES, SUBSPECIES AND NAMES.

Neonotus
Protasmanina

Austrotheobaldia

acaciae (Physalospora)
acaciae (Stagonospora)
alyxiae (Phyllosticta)

amygdalinum (Microthyrium) ..

apicalis (Aneura?)
arthrocnemi (Puccinia)
australiense (Lembosiopsis)
australiensis (Chlorops)
australiensis (Hendersonia)
australiensis (Malacaria)
australiensis (Nematothecium)
banksiae (Botryosphaeria)
banksiae (Pestalotia)
biparia (Brachycome)
brunneus (Coptotermes)
bundyensis (Culicoides)
bunrooensis (Culicoides)
carpholobi (Septoria)
centaurii (Cystopus)
chadwickii (Neonotus)
clelandi (Lachnella)
clelandi (Gorgoniceps)
clematidis (Asterina)
coronalis (Culicoides)..
constrictus (Holothyrus)
daviesiae (Godronia)
doryphorae (Asterina)
drimidicola (Asterina)
eucalypti (Hlsinoe)
eucalyptorum (Hendersonia)
eyrensis (Brachycome)
fieldiae (Asterina)

Vou. 79.

Genera.
Page
.. 230 Protolimnophila

60

Subgenera.
68, 75 Neotheobaldia

Species.
ao JUlg) fraseri (Balladyna)
bo dlaxs fraseri (Hendersonia)
so algal fraseri (Microthyrielia)
. 102 goodiae (Montagnellina)
. 60 gracilis (Brachycome)
ee 29 grevilleae (Stagonospora)
so db hardenbergiae (Leptosphaeria)
50 ales) heterodendri (Ascochyta)
ao diay! hornseyi (Itajahya)
oo dial imperfecta (Centrocnemis?)
5 alas inclusus (Ahamitermes)
oo daily lomatiae (Stigmatea)
.. 139 lomatie (Monochaetia)
.. 206 longistriata (Hendersonia)
a LNT moreensis (Culicoides)
50) ae) muehlenbeckiae (Monochaetia)
oS muscula (Scaptia)
. 138 myopori (Camarosporium)
Be te nana (Protasmanina)
50 BAD nattaiensis (Culicoides)
so ZAG notelaeae (Lembosia)
oo AlAs) novae-zealandiae (Meliolina)
. 107 occidentalis (Batrachomyia)
oo Cee! oritis (Asterina)
oo dl) ortidis (Stigmatea)
So k24 peculiaris (Pemphigonotus)
.. 109 phaeospora (Microthyriella)
.. 109 rara (Brachycome)
ap iS restionis (Leptosphaeria)
. 135 rubi (Aphanostigme)
.. 207 salicorniae (Camarosporium)
elon samueli (Camarosporium)

Page
5s

68, 69

.. 100
.. 185
so dll
co, abil
.. 206
.. 134
.. 120
.. 132

. 130
56) OY
.. 180
.. L101
.. 140
.. 136
.. 239
.. 140
.. 223

. 137
.. 60.
.. 2387

5 dial
SoS)
.. 196
50 JG)

. 102
.. 186
oo dil
.. 204
did AIL

. 114
oo. UBS
0 UBL

Page Page

samueli (Pseudosphaeria).. .. .. 118 Alene (CAUSUMOC)) 65 65 oo oo Wt
serrata (Hibbertia) Saga i) espa yee uliginosa (Brachycome) .. .. .. 208
Sy OS (COTM NCKANOD)) 60 60 oo oo Llmil victoriensis (Diplotowa) .. .. .. 184
suaedae-australis (Mycosphaerella) .. 122 victoriensis (Theobaldia) .. .. .. 72
superba (Protolimnophila) Pemraae tte) waringi (Culicoides) .. .. .. .. 240
tetraphylla (Macadamia) .. .. .. 15 williamsi (Coniochaeta) .. .. .. 115
tetraspora (Lamprospora).. .. .. 126 williamsii (Linocarpon) .. .. .. 122
tonnoiri (Lasiopleura) Sea ee Miter d betes williwilli (Culicoides) Peet hie Cacho) ZOOM
triodiae (Hendersonia) FS ee mote eal Lai) xanthorrhoeae (Physadalospora) 50 JIAKI)
Subspecies.
frenchi atritarsalis (Theobaldia) .. 73

New Names.
austropalpalis (Culicoides) RG ee) mallochi (Thaumatomyia).. .. .. 185

LIST OF PLATES.
PROCEEDINGS, 1954.

i—Hibbertia serrata, sp. nov.

iv.—Aeschnidiopsis flindersiensis (Woodward).
v.—Bacteria from Macquarie Island.

vi.—Collenia frequens.

vii.—Antirrhinum Rust, Puccinia antirrhini D. & H.
vili.im—“Anomalous” krasnozem at Fairy Hill.

ix.—Wings of species of Culicoides.

CORRIGENDUM (PROCEEDINGS, 1954, Parts 1-2).

Page xxviii, Table 18, Column 3 (headed “94’’), last two lines (D, ‘Milky Colony”),
for — read +.

li

INDEX.

(1954.)

Page.

Abstract of Proceedings ...... XXXVIi-xlii
Acarine Parasites of Two Species of

Rattus from Brisbane, Australia 156
Aédes scutellaris and Other Mos-
quitoes, the Nature and Signifi-
cance of Non-reciprocal Fertility

UTR ere hireurceoasieaycices acieust elicits ieriers 163
Aédes scutellaris scutellaris (Walker),
The Influence of the Physical
Properties of a Water Container
Surface upon its Selection by the
Gravid Females of, for Ovi-

position (Diptera, Culicidae) .. 211
Aédes (Stegomyia) pseudoscutellaris
Theobald and Aédes (Stegomyia)
polynesiensis Marks (Diptera,
Culicidae), Experimental Cross-

ISTE OLE Sete cueher Gee Agee eC REPRE aI 19
Ahamitermes (Isoptera) from Aus-

tralia, A New Coptotermes and .. 177

Annual General Meeting ........... i
Antirrhinum Rust, Puccinia antirrhini
Dae cienine eAuistiraliiae-.- jae sc 145

Ashby, Clark, see Lecturettes.

Athwal, D. S., and Watson, I. A.,
Inheritance and the Genetic
Relationship of Resistance pos-
sessed by Two Kenya Wheats to
Races of Puccinia graminis
BUC LC Teeny aie erect ole eas ke a 1

Australasian Ceratopogonidae (Dip-
tera, Nematocera). VII. Notes on
the Genera Alluaudomyia, Cerato-
pogon, Culicoides and Lasiohelea 233

Australian Beetles in the Tribe
Bolboceratini formerly in the

Genus Bolboceras, Notes on .. 142
Australian Chloropidae (Diptera),

Notes and Descriptions of ..... 182
Australian Fungi. II. New Records

ENaC! IEWAISMOINIS SooboacbocuonooU. Da

Bacteria belonging to the Rhodo-
bacteriineae Breed, Murray and
Hitchens, and the Chlamydobac-
teriales Buchanan occurring at
Macquarie Island, Notes on the.. 63

Balance Sheets for the Year ending
28th February, 1954 .... xxxiii-xxxv

Baur, G. N., elected a member .. xxxvii

Beadle, N. C. W., elected member of
Council, xxxix—see Notes and
Exhibits.

Bearup, A. J., see Notes and Exhibits.

Beetles, Australian, in the Tribe
Bolboceratini formerly in the
Genus Bolboceras, Notes on .. 142

Black, R. F., elected a member .. xxxvii
Brachycome Cass., Supplementary
Notes on the Genus Sd5aC G00 04 203

Page.
Brown, K. G., reference to death .. xxxvii
Browne, W. R., elected Honorary Sec-
retary, xxxvii—see Notes and
Exhibits.
Bunt, J. S., A Comparative Account
of the Terrestrial Diatoms of
Macquarie Island, 34—Notes on
the Bacteria belonging to the
Rhodobacteriineae Breed, Murray
and Hitchens, and the Chlamydo-
bacteriales Buchanan occurring
at Macquarie Island, 63. :

Cameron, Beryl M., elected a member xli
Ceratopogonidae (Diptera, Nemato-
cera), Australasian, VII ....... 233
Chloropidae (Diptera), Australian,
Notes and Descriptions of ..... 182
Colefax, A. N., elected a_ Vice-
IEA RENS HUG CEN DY HAR ares Hag sce Aa see eee Pe ieee XXXVii
Collenia frequens in Upper Protero-
zoic Rocks in the Northern Terri-
Woy Oi ANWR, ooococconouode 95
Comparative Account of the Terres-
trial Diatoms of Macquarie
VISMAING Bee se netsnast pen eee aia eet 34
Congratulations to members iii, xxxvii-xl
Copland, S. J., elected a Vice-Presi-

(OICLON E aetorchescavearaere ts carina Cones icky ha XXXVii
Coptotermes and Ahamitermes (Isop-
tera), a New, from Australia .. 177

Culex pipiens Group in South-eastern
Australia. III. Autogeny in Culex
pipiens form molestus ......... 193

Cytological Studies in the Myrtaceae.

IV. The Sub-tribe Huchamaelau-
CHM AC Selec cio ac sueesy MeN ree me ee Tit rp

Davis, Gwenda L., Supplementary
Notes on the Genus Brachycome

Cals’ Ai ssi OLE RE Seath ir es ae in bk 203
de Beuzeville, W. A. W., reference to
Cea Eh Fae ee eels a ee Hae XXXVIii

Diatoms, Terrestrial, of Macquarie
Island, A Comparative Account
GRAAL Ae MIEN) ics Suey Bae Sesame Rene eae 34

Diptera, Fossil, of the Tertiary Red-
bank Plains Series, Queensland 58

Dobrotworsky, N. V., The Culex
pipiens Group in South-eastern
Australia. III, 193—The Genus
Theobaldia (Diptera, Culicidae)
in Victoria, 65.

Domrow, R., elected a member,
xxxvili— Acarine Parasites of
Two Species of Rattus from Bris-
bane, Australia, 156—A Second
Species of Holothyrus (Acarina:
Holothyroidea) from Australia,
159.

“AM Are cy
a | Ole $2.

lil INDEX.

Page.

Douglas, G. W., elected a member .._ xl
Dragon-fly, Aeschnidiopsis flindersien-
sis (Woodward) from the Queens-

land Cretaceous, A Second
Specimenworsthelacnacecemeciercer 61
IDJARIOINS Gooooocosd00. Xxxii, XxXXvii-xli

English, Kathleen M. I., Notes on the
Morphology and_ Biology of
Scaptia vicina Tayl., and a New
Species of Scaptia (Diptera,
Tabanidae), 219—see Notes and
Exhibits.

Eucalyptus cinerea F. Muell., and
Eucalyptus robusta Sm., An F1
Ely brid) between anes cee ceerrle a 196

Eucalyptus robusta Sm. An Ft
Hybrid between Eucalyptus
cinerea F. Muell., and .......... 196

Eucalyptus, The Inheritance of In-
florescence Characters in ...... 79

Hxchange relations) Sys... coe eee: i

Exhibits—see Notes and Exhibits.

Experimental Crossing of Aédes
(Stegomyia) pseudoscutellaris
Theobald and Aédes (Stegomyia)
polynesiensis Marks (Diptera,
CUlicidae) i VAs eee eee 19

Fl Hybrid between JHucalyptus
cinerea F. Muell., and Hucalyptus
TOOUSEG (SMS,, é:ccacceiea.sastoeneneceieeee 196

Fossil Diptera of the Tertiary Red-
bank Plains Series, Queensland 58

Fraser, A. S., see Lecturettes.

Fraser, Lilian, see Notes and
Exhibits.
MungiseAustralians ll Sansone ers 97

Gay, F. J.. A New Coptotermes and

Ahamitermes (Isoptera) from

AuStraliia Cahn teste tae a ed eeeee 177
Genus Theobaldia (Diptera, Culi-

GiCES)) sha Wau, Sobacoccccu00e 65

Hannon, Nola J., appointed Linnean
Macleay Fellow in Botany for
1954, iii—reappointed for 1955,
xli.

Hansford, C. G., Australian Fungi.

ITT gM ar es at ee ee ae 97

Hardy, G. H., see Notes and Exhibits.

Heron Island, Queensland, Notes on
the Scleractinia or Stony Corals
(Coelenterata) of ............. 90

Herz, A. J., see Lecturettes.

Hibbertia Andr., A New Species of,
from Western Australia ....... 29
Hill, Prof. J. P., reference to death xxxix

Hindmarsh, Mary M., Linnean
Macleay Fellow in Botany, re-
appointed for 1953, iii—Summary
of work, iii—resignation, iii.

Holothyrus (Acarina: Holothyroidea)
from Australia, A Second Species
OL ine cavetaveiace anne tee elie eee neato soo Abe)

Page.
Hotchkiss, A. T., A New Species of
Hibbertia Andr., from Western
Australia, 29—see Lecturettes.

Howden, H. F., Notes on Australian
Beetles in the Tribe Bolboceratini
formerly in the Genus Bolboceras 142

Influence of the Physical Properties
of a Water Container Surface
upon its Selection by the Gravid
Females of Aédes_ scutellaris
scutellaris (Walker) for Ovi-
position (Diptera, Culicidae) .. 211

Inheritance and the Genetic Relation-
ship of Resistance possessed by
Two Kenya Wheats to Races of

Puccinia graminis tritici ...... alt
Inheritance of Inflorescence Charac-
ters in Hucalyptus ............ ao,

Jacobs, E. G., reference to death .. xxxix

Johnson, L. A. S., Macadamia terni-
folia F. Muell., and a Related
NEW SDECLES aren ce ieee 15

Kosciusko Region, Fifth Natural
History Survey made .......... ii

Krasnozem, “Anomalous”, in _ the
Richmond-Tweed Region of
N.S.W., A Note on the Occurrence
Of 2. a ee ene 199

Labenini from Australia (Pimplinae,
Ichneumonidae), A New Genus

and Species of the Tribe ...... 230
Lecturettes:
Ashby, Dr. Clark—Plants, Moun-
tains and Laboratories .......... xli
Fraser, Dr. A. S.—Hair Growth in
SATIIMAISH ce er netee cree es eee XXXVili
‘Herz, Dr. A. J.—The Cosmic
Radiation ss eae. XXXVIli
Hotchkiss, Dr. A. T.—Let’s Talk
AMIBKG<55 2 ic seen ktet se el samme XXxix
Millerd, Dr. Adele—The Biosyn-
thesis of Rubber ............... xi
Troughton, E.—Field Work in the
New Guinea Highlands ........ xiii

Lee, D. J., elected-a Vice-President xxxvii

Lee, D. J., and Reye, E. J., Aus-
tralasian Ceratopogonidae (Dip-

tera, Nematocera). VII ........ 233
Legumes, Pasture, The Root-nodule
Bactenla cota scrseiss seasons, | eee iv
Library accessions ................. i
Linnean Macleay Fellowships:
Reappointments for 1953 ....... iii
Appointments for 1954 .......... iii

Applications invited for 1955 .. xlI-xli
Reappointment and appointment

for 955: cis aacnat «osteo enee xli
Iuist of Members ...-.:..:......- .. Xliii
Tnisteof PlatesSi« cys cach hos Se eee 1

Lists of New Genera, Subgenera,
Species, Subspecies and Names xlix-l

Page
Macadamia ternifolia F. Muell., and
a Related New Species ......... 15
Macdonald, Mary B., elected a

member, xli—appointed Linnean
Macleay Fellow in Botany for
1955, xli.

Macleay Bacteriologist, acknowledge-
ment of financial support from
banks and report re position of,
iv—see also under Tchan, Yao-
tseng.

Macquarie Island, A Comparative
Account of the Terrestrial
Diatoms of, 34—Notes on the
Bacteria belonging to the Rhodo-
bacteriineae Breed, Murray and
Hitchens, and the Chlamydobac-
teriales Buchanan occurring at,
63.

Martin, Mrs. Hilda R., appointed

Linnean Macleay Fellow in
Botany for 1954 ...............

McAlpine, D. K., see Notes and
Exhibits.

McGarity, J. W., and Munns, D. N.,
A Note on the Occurrence of
“Anomalous” Krasnozem in the
Richmond-Tweed Region of
INASSVWist eres etc ses weos te Se eae

Members, List of ...........

Mercer, F. V., elected President, xxxii
—see Notes and Exhibits.

Millerd, Adele, see Lecturettes.

Mosquitoes, The Nature and Signifi-
cance of Non-reciprocal Fertility
in Aédes scutellaris and other ..

Munns, D. N., see McGarity, J. W.,
and Munns, D. N.

Myrtaceae, Cytological Studies in the,
IV

a Cd

Nature and Significance of Non-
reciprocal Fertility in Aédes
scutellaris and other Mosquitoes

New Coptotermes and Ahamitermes
(Isoptera) from Australia

New Genus and Species of the
Tribe Labenini from Australia
(Pimplinae, Ichneumonidae)

New Species of Hibbertia Andr., from
Western Australia ............

Northern Territory of Australia,
Collenia frequens in Upper Pro-
terozoic Rocks in the .........

Note on the Occurrence of “Anom-
alous” Krasnozem in the Rich-
mond-Tweed Region of N.S.W. ..

Notes and Descriptions of Australian
Chloropidae (Diptera)

Notes and Exhibits:

Beadle, N. C. W.—A white Waratah
which came from a plant grow-
ing on Hawkesbury Sandstone in
the Hunter Valley

Bearup, A. J.—Schistosomes of the
genus Microbilharzia which had
developed in a budgerigar
(Melopsittacus) as a result of
experimental infection with cer-
cariae from Pyrazus australis .

ili

199

xliii-xl viii

163

21

163
177

. 230

29

95

199
182

xli

. xiii

INDEX.

liii

Page.

Reference to cases of dermatitis
which have been reported as due
to immersion in the salt water
of coastal lakes in the Sydney
GIStniche teases oa ee

Browne, W. R.—Coloured slides
showing the effects of bushfires
on the vegetation of the
Kosciusko district, xli — (on
behalf of the Society) Fourteen
water-colour paintings of native
plants painted by and bequeathed
to the Society by the late P. C. W.
Shaw, xxxix.

English, Kathleen M. I.—Shield
coverings of the curious elongated
larvae of a species of scale insect
(Coccidae) found on Casuarina at
Bantry Bay, Middle Harbour ..

Fraser, Lilian—A small plant of
orange on Seville root stock
which was affected with the
aphid-transmitted virus disease,
EPISTLE ZA Rey ae eae a iene

Hardy, G. H.—Larvae of the family
Asilidae (Robber-flies) taken in
garden soil at Katoomba .......

Linnean Society (on behalf of)—
see Browne, W. R.

McAlpine, D. K.—Eggs, . larvae,
pupae and adults of the Fly,
Batrachomyia, which is parasitic
under the skin of certain Aus-
tralian frogs

Mercer, F. V.—Description of some
experiments concerning pollina-
tion mechanism of the Trigger
Plant, Stylidium species

Osborne, G. D.—Specimens and
photographs of veins of asbestos
(chrysotile) in serpentine from
Woodsreef, east of Barraba,
showing ribbon structure ....

Shaw paintings, see under Browne,
W. R.

Walker, J— Specimen of the fungus
Podaxis pistillaris (Ll. ex Pers.)
Morse collected at Bourke,
INES EWA Mes cond eS CURE ed avers

Waterhouse, J. T.—Series of 1,097
original water-colour drawings
of British Fungi by Edwin
Wheeler (1835-1909), belonging
to the Fisher Library, University
of Sydney

Notes on Australian Beetles in the
Tribe Bolboceratini formerly in
the Genus Bolboceras

Notes on the Bacteria belonging to
the Rhodobacteriineae Breed,
Murray and Hitchens, and the
Chlamydobacteriales Buchanan
occurring at Macquarie Island ..

Notes on the Morphology and Biology
of Scaptia vicina Tayl., and a
New Species of Scaptia (Diptera,
Tabanidae)

eC

ee

XXXVili

XXX1X

xl

xl

XXXix

XXXVI1i

XXX1X

xl

xl

142

63

liv INDEX.

Page.
Notes on the Scleractinia or Stony
Corals (Coelenterata) of Heron
Island, Queensland. I. A List of
the Common Species ........... 90

O’Gower, A. K., The Influence of the
Physical Properties of a Water
Container Surface upon its Selec-
tion by the Gravid Females of

Aédes scutellaris scutellaris
(Walker) for Oviposition (Dip-
Wares (OwIhiGiGlA®)) sioccocnooudoos 211

Osborne, G. D., see Notes and
Exhibits.

Parrott, A. W., A New Genus and
Species of the Tribe Labenini

from Australia (Pimplinae,

Ichneumonidae) ............... 230
PIL ATeS GETS Ota: pentose cieie: nae cba nus, o's 1
Presidential Address .............. i

Pryor, L. D., An Fl Hybrid between
Eucalyptus cinerea F. Muell., and
EHucalyptus robusta Sm., 196—
The Inheritance of Inflorescence
Characters in Hucalyptus, 79.

Puccinia antirrhini D. & H., Antir-
rhinum Rust, in Australia .... 145

Puccinia graminis tritici, Inheritance
and the Genetic Relationship of
Resistance possessed by Two
Kenya Wheats to Races of ...... i

Raggatt, H. G., congratulations on
the receipt of the Honour of
(Gis) Bil AD ah wee aal ava Arete mana ie Sve Nice pte XXXVIli

Rattus from Brisbane, Australia,
Acarine Parasites of Two Species
Oy aati ae ene Sues RFs AE leet 156

Reye, E. J., see Lee, D. J., and Reye,
wh,

Richmond-Tweed Region of N.S.W.,
A Note on the “Anomalous”
Krasnozem in the .............. 199

Riek, E. F., A Second Specimen of
the Dragon-fly Aeschnidiopsis
flindersiensis (Woodward) from
the Queensland Cretaceous, 61—
The Fossil Diptera of the Ter-
tiary Redbank Plains Series,
Queensland, 58.

Root-nodule Bacteria of Pasture
Te SUIMES Weiets ivazsoot cp home bao cust ae iv

Sabrosky, C. W., Notes and Descrip-
tions of Australian Chloropidae
QDS a) Fete ara eee een ere 182

Salter, K. H. W., Notes on the Sclerac-
tinia or Stony Corals (Coelen-
terata) of Heron Island, Queens-
Mevrn eels pew stvcr sacha ee ae eens Pca ite ge 90

Scaptia vicina Tayl., and a New

Species of Scaptia (Diptera,
Tabanidae), Notes on the
Morphology and Biology of .... 219

Science House, Net return from ... li

_ Second

Page.
Scleractinia or Stony Corals (Coelen-
terata) of Heron Island, Queens-
lanidi’ Notes) one they tie... mee 90
Species of Holothyrus
(Acarina: Holothyroidea) from
Australia). “cu saotee coolers 159
Second Specimen of the Dragon-
fly, Aeschnidiopsis flindersiensis
(Woodward) from the Queens-
land Cretacecousman acco cei 61
Simons, Hilda Ruth, see Martin, Mrs.
Hilda R.
Smith-White, S., Cytological Studies
in the Myrtaceae. IV .......... 21
Smith-White, S., and Woodhill, A. R.,
The Nature and Significance of
Non-reciprocal Fertility in Aédes
scutellaris and other Mosquitoes 163
Supplementary Notes on the Genus
Brachycome Cass., Descriptions
of Five New Australian Species
and some New Locality Records 203

Tchan, Yao-tseng, Macleay Bacteriolo-

gist, Summary of work ........ iii
Theobaldia (Diptera, Culicidae), the
Genusani aVactoniana eee 65

Traves, D. M., Collenia frequens in
Upper Proterozoic Rocks in the
Northern Territory of Australia 95

Troughton, E., see Lecturettes.

Vallance, T. G., Linnean Macleay
Fellow in Geology:

Reappointed for 1953 .......... iii
Summary of work ............. ili
Resienationy 4s. yee een: lili
Vincent, J. M., elected a _ Vice-
President, xxxvii — The Root-
Nodule Bacteria of Pasture

Legumes (Presidential Address),
Iv.

Walker, J., Antirrhinum Rust, Puc-
cinia antirrhini D. & H., in Aus-
tralia, 145—see Notes and
Exhibits.

Walkom, A. B., congratulations on
award of Royal Society’s Medal
for 1953, xxxvii—elected Hon-
orary Secretary and Honorary
Treasurer, Xxxvii.

Waterhouse, J. T., see Notes and

Exhibits.
Watson, I. A., see Athwal, D. S., and
Watson, I. A.

Wheats, Two Kenya, Inheritance and
the Genetic Relationship of Re-
sistance possessed by, to Races
of Puccinia graminis tritici .. 1
Williams, J. B., elected a member xxxvii
Woodhill, A. R., Experimental Cross-
ing of Aédes (Stegomyia) pseudo-
scutellaris Theobald and Aédes
(Stegomyia) polynesiensis Marks
(Diptera, Culicidae), 19—see also
Smith-White, S., and Woodhill,
PN, 18%,

peut
4

INHERITANCE AND THE GENETIC RELATIONSHIP OF RESISTANCE POSSESSED
BY TWO KENYA WHEATS TO RACES OF PUCCINIA GRAMINIS TRITICI.

By D. S. ArHwat and I. A. Watson, The University of Sydney.
[Read 31st March, 1954.]

Synopsis.

The results of genetic studies on the stem rust resistance of Kenya 744 and Kenya 117A
are reported. Their resistance to race 222AB is inherited in a simple manner. The studies on
the relationship of seedling and field reactions are made. In addition to the major gene for
seedling resistance in these varieties modifying factors also appear to influence the field reaction.
The resistance in the two varieties has been found to be allelic.

The use of more races of stem rust in inheritance studies reveals the presence of one more
non-identical gene for stem rust resistance in each of these varieties.

A scheme is also presented for designating genes for stem rust resistance in Kenya wheats.

Introduction.

Wheats of Kenya origin have made important contributions to breeding programmes
designed to evolve varieties resistant to stem rust. Three of these varieties, Kenya 743,1
744 and 745, have been widely used for this purpose in Australia (Watson and
Waterhouse, 1949). Each of them was shown to have a diverse gene for resistance to
stem rust (Watson, 1943; Watson and Waterhouse, 1945). The gene in Kenya 743 was
rendered useless with the appearance of race 126B in 1942 (Watson and Waterhouse,
1949) and one in Kenya 745 became ineffective in 1948 with the appearance of races
222AB and 222BB (Waterhouse, 1952; Watson and Singh, 1952). The third gene present
in Kenya 744 is still effective in giving protection to races of stem rust now prevalent
in Australia and it has been incorporated into the commercial wheat varieties Festival
and Dowerin (Single, 1952). Kenya 117A is another variety resistant to all races of
stem rust in Australia and is being used by wheat breeders (Watson and Waterhouse,
1949; Single, 1952).

Kenya wheats have also been mentioned as important sources of resistance to stem
rust in other countries. A report of the International Wheat Stem Rust Conference held
at Winnipeg, Canada, in January, 1953, indicates the use being made of these wheats in
U.S.A. and Canada. The main source of resistance to race 15B had been Kenya wheats
and their derivatives (Ausemus et al., 1953) and Kenya 117A is being used to incorporate
resistance to this race. In South America, too, Kenya wheats are reported to be among
the most promising sources of resistance to stem rust (Vallega, 1951).

Review of Literature.

A review of literature on the inheritance of resistance in Kenya wheats is presented
below.

Watson (1941) found that Kenya 744 and Kenya 745 each had a single dominant
factor for seedling resistance to races 36 and 56 of stem rust and that the seedling and
field reactions were highly correlated. He also reported that single factors which
governed seedling resistance to race 17 in Kenya 744 and Kenya 745 were non-allelic.
Watson (1943) later carried out more extensive studies on these Kenya wheats, and
found that Kenya 745 had an additional factor for seedling resistance against some
races and that the resistance of Kenya 744 to powdery mildew was governed by a single
dominant factor inherited independently of the gene for stem rust resistance. He found
also that a factor in Kenya 745 inherited independently of the one giving resistance to
races 56 and 15 gave semi-resistance to a very virulent culture of race 15 from Brazil.

1 Varieties carry the Sydney University Accession numbers.
Cc

2 RESISTANCE POSSESSED BY TWO KENYA WHEATS TO RACES OF PUCCINIA GRAMINIS TRITICT,

Abbasi (1949) found that the resistance of Kenya RL 13738 to races 34 and 11 was
governed by single factors which were loosely linked. Hasanain (1949) reported the
resistance of certain Kenya wheats to Puccinia graminis tritici as a trigenic character.

Watson and Waterhouse (1949) studied several crosses involving Kenya varieties
and classified these varieties into three groups represented by Kenya 743, Kenya 744,
Kenya 745. These workers also reported that Kenya 117A which showed resistance to
stem rust in Australia as well as to race 15B in U.S.A. was a variety distinct from Kenya
743, 744 and 745 and of considerable potential value as a source of resistance. Kenya
117A was found (Athwal, 1953) to have a single partially dominant factor for resistance
to some races of Indian stem rust, and its resistance to race 42 was controlled by two
factors.

Materials.

The object of this study was to learn the mode of inheritance of resistance to stem
rust possessed by Kenya 744 and Kenya 117A 1347 and also to investigate the relation-
ship between genes for resistance present in these varieties. The reactions of both the
varieties were influenced by temperature and their range of reaction to different races
is shown in Table 1. The more resistant reaction in each case was obtained at approxi-
mately 60-65°F and the less resistant at approximately 75—-80°F.

TABLE 1.
Range of Reaction of Two Kenya Varieties to Races of Stem Rust.
Reaction to Stem Rust Races.
Variety.
| 126 | 126B |222AB/222BB| 15 15C 21 34 38 40 122 24 21K
=
Kenya 744 .. | R-SR} R-SR | R-SR | R-SR | R-S R-S R-SR | SR-S | R-SR|SR-S |S SS) SS)
Kenya 117A .. | R-SR | R-SR | R-SR | R-SR | R-SR | R-SR | R-SR | R-SR | R-SR | R-SR | R-SR| R-SR|S

R=resistant ; SR=semi-resistant ; S—=susceptible.

Races 15, 21, 34, 38, 40, 122 and 24 of stem rust gave standard reactions on the
differential set recorded by Stakman and Levine (1938). Race 15C differs from race 15
in its ability to attack Yalta, and 21K differs from 21 in its ability to attack Kenya 117A.

In the field Kenya 744 and Kenya 117A are highly resistant to a collection of the
common Australian rusts, viz. races 126, 126B, 222AB and 222BB.

Kenya 744 is an early variety with white chaff, red grain and is half awned. It is
probably identical with a variety known elsewhere as Kenya Standard. Kenya 117A is a
later maturing variety with white chaff, red grain and tip awned. It has a high degree
of resistance under Australian conditions. This variety is of some interest also in that
it shows the usual compatibility with Khapli Emmer (Waterhouse, 1952). However, it
appears to differ in its behaviour to.stem rust from a collection of Kenya 117A imported
from U.S.A. As far as is known the pedigree of Kenya 744 and Kenya 117A 1347 is
A 8 V.12.D(L) x Marquis and BF 4.2D.30.B.2(L) respectively.

Federation 107 and Chinese White 1806 were used as susceptible parents in crosses
with Kenya varieties. Chinese White was susceptible to all the races used in inheritance
studies while Federation showed only partial resistance to race 38 at low temperatures,
60-65°F. The tests on crosses involving Federation were carried out at reasonably high
temperatures against race 38, 70-75°F.

Methods.

Inheritance of resistance was studied in the F,, F., F, and F, generations of crosses
of Kenya 744 and Kenya 117A with a susceptible variety. Race 222AB was first used in
the glasshouse to study the mode of inheritance and later the relationship between the
four races, 126, 126B, 222AB and 222BB, was found from the breeding behaviour of F,

Ww

BY D. S. ATHWAL AND I. A. WATSON.

and F, families. After testing with race 222AB, F, seedlings of the crosses Federation x
Kenya 744 and Federation x Kenya 117A were transplanted in the field. F,; progenies of
these plants were studied as seedlings with race 222AB to check the F, classifications.
F,, lines of these crosses were also grown in the field and classified for stem rust reaction
to mixed inoculum of the above four races. It was thus possible to correlate the seedling
and adult plant reactions.

F,s of the crosses Federation x Kenya 744 and Chinese White x Kenya 744 when
tested with stem rust race 222AB were also classified for reaction to powdery mildew to
confirm the earlier results of Watson (1948).

Fs and F.s of the cross Kenya 744 x Kenya 117A and its reciprocal were studied with
race 222AB in the glasshouse. The seedlings were then transplanted in the field and
notes for stem rust reaction were taken on mature plants. Some of the plants were again
tested with race 222AB in the F, generation to confirm F, results.

In addition to the Australian races of stem rust, race 38 from the United States and’
race 122 from Kenya were used to get more information on gene relationship and the
number of genetic factors for resistance in these varieties.

The following classes were used to record stem rust reaction in the seedling and
mature plant stage.

Seedling reactions: Resistant (R) —; and 1; Semi-resistant (SR)—2, 3°; Intermediate
(Int.)—3°; Segregating (Seg.)—F, or F, line consisting of Resistant
or Semi-resistant and susceptible plants; Susceptible—3 and 4.
Field reaction: Resistant (R)—Almost free from rust; Moderately Resistant (R)—15.
to 25% of the area of stem was infected; Semi-Resistant (SR)—25—40%,
of the stem was infected; Intermediate (Int.)—40-60% of the stem was:
infected; Segregating (Seg.)—-Lines consisting of resistant and!
susceptible plants; Susceptible (S)—The plants were heavily infected
like the susceptible parents.

TABLE 2.

Reaction of F.s of the Cross Federation x Kenya 744 to Powdery Mildew and Stem Rust
Race 222AB.

Reaction to Race 222AB.

Resistant to
Semi-resistant. | Susceptible. Total.
Reaction to mildew :
Resistant 216 80 296
Susceptible 69 19 88
Total 285 99 384

P value for 3:1 ratio for stem rust reaction lies between 0-70 and 0-80.—P value
for 3: 1 ratio for mildew reaction lies between 0-30 and 0-50.—P value for independence
test lies between 0-30 and 0-50.

EXPERIMENTAL RESULTS.
Inheritance of the resistance of Kenya 744.

Glasshouse studies with race 222AB.—8 F, seedlings of the cross Federation x Kenya
744 were tested with race 222AB and the resistance was found to be dominant.

F.s of this cross and of the cross Chinese White x Kenya 744 were inoculated with
race 222AB. After incubation the pots were allowed to stand near wheat plants infected
with powdery mildew. Mildew infection did not interfere unduly with the rust reactions
and it was quite possible to take observations on stem rust and mildew simultaneously.
The data are presented in Tables 2 and 3. Federation and Chinese White were susceptible
to stem rust race 222AB and mildew, whilst Kenya 744 was resistant.

4 RESISTANCE POSSESSED BY TWO KENYA WHEATS TO RACES OF PUCCINIA GRAMINIS TRITICT,

From data in Tables 2 and 3, it can be concluded that a single dominant factor in
Kenya 744 controlled seedling resistance to race 222AB. The resistance to powdery
mildew which was monogenic did not show any association with stem rust resistance.

F, plants of the cross Federation x Kenya 744 representing resistant and susceptible
classes in Table 2, were transplanted in the field. Their F, reactions are compared in
Table 4 with the breeding behaviour of F,, families when tested with the same race of
stem rust.

TABLE 3.

Reaction of F's of the Cross Chinese White x Kenya 744 to Powdery Mildew and Stem Rust
Race 222AB.

Reaction to Race 222AB.

Resistant to
Semi-resistant. | Susceptible. Total.
Reaction to mildew: |
Resistant .. HS ts ae 504 163 667
Susceptible aS a6) 66 170 49 219
Total ate 6 ah 674 212 886

P value for 3:1 ration for stem rust reaction lies between 0-30 and 0-50.—P value
for 3:1 ratio for mildew reaction lies between 0-80 and 0-90.—P value for independence
test lies between 0°50 and 0-70.

The breeding behaviour of F, families in Table 4 shows that five out of 112 F, plants
were wrongly classified. P value for the agreement of F, data with a 1:2:1 ratio is
rather low because a relatively high proportion of resistant F, plants gave progenies
which were homozygous for resistance. If the resistant and segregating F, families are
grouped together the data (83:29) agree well with 3:1 ratio (P 0-80-0-90).

TABLE 4.
The Breeding Behaviour of F; Families from the Cross Federation x Kenya 744 Raised from F, Plants
Tested and Transplanted in the Field.

F; Classification with Race 222AB.

Resistant to
Semi-resistant.| Segregating. Susceptible. Total.
F, reactions with race 222AB:
Resistant to semi-resistant 38 41 1 80
Susceptible at Bra = 4 28 32
Total = ae 38 45 29 112

P value for 1:2:1 ratio=0-05-0-10.

Additional information on the inheritance of resistance to race 222AB was obtained
from tests with a random sample of 158 F, lines of the cross Federation x Kenya 744.
Forty-four lines were homozygous for resistance or semi-resistance, 71 segregated and 43
were homozygous susceptible. The data agree with a 1:2:1 ratio (P 0-30-0-50). The
numbers of resistant and susceptible plants were found in the 71 segregating lines as
well as in the 45 segregating lines of Table 4. A total of 116 lines consisted of 2045
plants of which 1480 were resistant to semi-resistant, 43 intermediate and 522 susceptible.
If the intermediate plants are grouped with the resistant class, the P value for the
goodness of fit to 3:1 ratio lies between 0-50 and 0-70.

BY D. S. ATHWAL AND I. A. WATSON. 5

Field Studies.—F, plants of the cross Federation x Kenya 744 showed slight develop-
ment of rust in the field and they were classed as moderately resistant. Kenya 744
showed complete freedom from stem rust and Federation was susceptible.

Two hundred and sixty-eight F, lines of the Federation x Kenya 744 cross were also
grown in the field. All the four Australian races of stem rust were present and the lines
were classified as resistant to moderately resistant, segregating and susceptible. Table 5
contains the data on the F, lines.

TABLE 5.

Classification of F, Lines of the Cross Federation x Kenya 744 for Their Reaction to Stem Rust in the
Field.

Reaction to Stem Rust in the Field.

| Resistant to |
Resistant. | Moderately | Segregating. Susceptible. | Total.
| Resistant. | | |
|
Parents ¥ Kenya 744 | — | — | Federation | a
F, lines et _: | 78 134 56 | 268
| |

P value for 1:2:1 ratio lies between 0-10 and 0-20.

The field data also indicate the presence of a single major factor for resistance in
Kenya 744. The resistance was partially dominant and appears to be influenced by
modifying genes. Since 744 is a very early variety escapes can probably account for the
larger number of resistant lines as compared to the susceptible ones. Of the 268 lines
151 were tested in the glasshouse with race 222AB and Table 6 shows relationship
between the seedling and field reaction which is fairly satisfactory.

TABLE 6.
Relationship of the Seedling Reaction of Federation x Kenya 744 F, Lines to Their Reaction in the Field.
Seedling Reaction to Race 222AB.
Resistant to |
Semi-resistant.| Segregating. | Susceptible. | Total.
| | |
Reaction to Races 126, 126B, | |
222A B, 222BB in the field : |
Resistant to moderately re- | - |
sistant a i Le 41 | 6 — | 47
Segregating R &S, or R- &S.. 4 56 | 10 70
Susceptible ce be Bo — 4 | 30 34
Total oe ts 45 66 40 | 151

Relationship between races.—F, lines of the cross Federation x Kenya 744, with
known reactions to race 222AB, were tested in the F, generation against races 126, 126B
and 222BB. Twenty-nine lines which were homozygous resistant to race 222AB, bred
true for their resistance against the other three races. Another 22 F, lines homozygous
susceptible to race 222AB also showed the same breeding behaviour against the other
races. From this it can be concluded that the same gene in Kenya 744 was responsible
for resistance to the four races of stem rust.

Studies with race 38.—Preliminary tests made on F, lines of the cross Federation x
Kenya 744 did not show strong correlation between reactions to race 38 and race 222AB.
Fs of the cross Chinese White x Kenya 744 were tested with race 38 to study the mode

6 RESISTANCE POSSESSED BY TWO KENYA WHEATS TO RACES OF PUCCINIA GRAMINIS TRITICI,

ot inheritance of resistance. The data in Table 7 show that Kenya 744 has two factors
for resistance to race 38. One of these factors appears to be only partially dominant.
The plants showing an intermediate type of reaction were grouped with the resistant
and semi-resistant class when interpreting the data. The goodness of fit was not very
satisfactory, probably because some of the plants classified as intermediate for their
reaction to race 38 are genotypically susceptible. The data on F, lines included in the
same table confirm the presence of two factors in Kenya 744.

TABLE 7.

Reactions of F'.s of the Cross Chinese White x Kenya 744, and Fs of the Cross Federation x Kenya 744 to Stem Rust Race 38.

Reactions to Race 38.

| |
Resistant to
Semi-resistant. | Intermediate. | Segregating. Susceptible. Total.
Parents of F.s oe a Kenya 744 — — Chinese White —
Chinese White x Kenya 744 F.s 708 36 — 36 780
Parents of Fys_ .. ae a: Kenya 744 — — Federation —_
Federation x Kenya 744 Fys_ .. 55 — 68 6 129

P value for 15 : 1 ratio for F, data lies between 0:05 and 0-10.—P value for 7: 8: 1 ratio for F, data lies between
0-50 and 0-70.

The reactions of 178 randomly selected F, and F, lines of Federation x Kenya 744
cross to race 38 and race 222AB are compared in Table 8.

These data show that one of the two factors in Kenya 744 which give resistance to
race 38, is the same as the one giving resistance against race 222AB. Figures within
brackets in Table 8 indicate the lines which show reactions not expected on the basis of

TABLE 8.

Distribution of 178 Lines of the Cross Federation x Kenya 744 for Their Reaction to Race 38 and
Race 222AB.

Reaction to Race 38.
Resistant to
Semi-resistant. | Segregating. Susceptible. Total.
| |
Reaction to race 222AB: |
Resistant to semi-resistant 51 (2) | ay 53
Segregating fy NG 24 | 57 (1) 82
Susceptible ae a 8 27 8 43
|
| —— |
Total 0 Ats7e| 83 86 9 178

this hypothesis. This is a minor discrepancy and could happen by chance. If the three
families showing the unexpected behaviour are disregarded, the reactions of the rest of
the lines (51:24:57:8:27:8) can be fitted with expected ratio of 4:2:6:1:2:1 on the basis
of preceding hypothesis. The P value for the goodness of fit lies between 0-30 and 0-50
which is fairly satisfactory.

It is of some interest to find the presence of two factors in Kenya 744 against race
38. Only one of them is effective against Australian rusts. Several other varieties, for
example, Bencubbin, Ford, Gular and Mentana, which are susceptible to stem rust in
Australia show resistance to race 38. Work is in hand at present to relate the resistance
of these varieties to race 38 with that of Kenya 744.

BY D. 8S. ATHWAL ANDI. A. WATSON. 7

Inheritance of the resistance of Kenya 117A.

Glasshouse studies with race 222AB.—6 F, plants of the cross Federation x Kenya
117A were tested with race 222AB and were found to be almost as resistant as Kenya
117A.

Athwal (1953) reported that reactions of F, segregates from crosses involving Kenya
117A were difficult to read and the heterozygous plants showed an intermediate type of

TABLE 9.
Reactions of F.s, Fs and Fs of the Cross Federation x Kenya 117A to Race 222AB.

Reactions to Race 222AB.

Resistant or
Semi-resistant.| Segregating. Susceptible. | Total.
|
| |
Kenya 117A — Federation | a=
Fs 407 = 138 | 545
Fs 24 49 23 96
F,s 15 45 22 | 82
{ |

2

P value for 3: 1 ratio for the F, data lies between 0-80 and 0:90.—P value
for 1 : 2:1 ratio for the F, data lies between 0-95 and 0:98.—P value for 1:2:1
ratio for the F, data lies between 0:30 and 0-50.

reaction. With race 222AB, however, the segregation was clear cut and heterozygous F,
plants were indistinguishable in their reactions from homozygous plants. The data on
F.s, F,3s and F,s of the cross Federation x Kenya 117A are presented in Table 9.

F, data indicate the presence of a single dominant factor for seedling resistance in
Kenya 117A.

The breeding behaviour of F, and F, lines confirms the conclusion drawn from F,
data.

TABLE 10.
The Breeding Behaviour of the Progenies of F, Plants of the Cross Federation x Kenya 117A Tested
and Transplanted in the Field.
|
| Seedling Reactions of F; Families to Race 222AB.

|
Resistant or
Semi-resistant. | Segregating. Susceptible. Total.

F, seedling reactions to race

222AB: |

Resistant to semi- |
resistant Be ae 15 26 — 41
Susceptible ae os — = 11 11
Total fe Pi 15 26 | 11 52

P value for 1: 2:1 ratio for F; reactions lies between 0-70 and 0-80.

Some F, seedlings from the resistant and susceptible classes were transplanted in
the field. Their progenies were again tested with race 222AB to confirm the accuracy of
F, classification. The breeding behaviour of F, families is correlated with the F,
reactions in Table 10.

As the resistance of Kenya 117A is dominant, the progenies of resistant F, plants
were either homozygous resistant or segregating, whilst all the susceptible F, plants
bred true in the F, generation. The F, test showed the correctness of the F, classification.

8 RESISTANCE POSSESSED BY TWO KENYA WHEATS TO RACES OF PUCCINIA GRAMINIS TRITICI,

In Tables 9 and 10, 75 segregating F. lines of the cross Federation x Kenya 117A
consisted of 1375 plants of which 1009 were resistant to semi-resistant, 7 intermediate
and 359 susceptible. By grouping the intermediate plants with the resistant ones, the
data agree with 3:1 ratio with a P value of 0-30—0-50.

Field Studies.—F, plants of the cross Federation x Kenya 117A were only moderately
resistant in the field whereas Kenya 117A maintained its full resistance. One hundred
and sixty F, lines of the same cross were also grown and tested in the field for rust
reaction to a mixture of four races of stem rust. The infection was good and Federation
was severely rusted. The F, lines were more difficult to classify and the segregation was
not as clear cut as expected on the basis of a single dominant factor. Only 11 out of 160
F, lines approximated to the resistance of Kenya 117A. Some lines showed a range of
reaction from resistance to moderate resistance and semi-resistance. Among the lines
which segregated for resistance and semi-resistance were some that were difficult to
separate from lines segregating for resistance and susceptibility. Likewise certain lines
segregating for semi-resistance and susceptibility were hard to separate from homozygous
susceptible lines. In such cases both the alternative classifications were recorded. The
field reaction of F, lines are given in Table 11 and are compared with the seedling
reaction against race 222AB.

TABLE 11.

Reaction of F'; Lines of the Cross Federation x Kenya 117A in the Field and Their Relationship with the Seedling Reactions
Against Race 222AB.

Field Reaction to Mixed Races.
R to R- | Rand SR
R. or or Seg. SR and 8 5. Total.
R to SR. | R and 8. or 8.
Seedling reaction to race 222AB :
Resistant to semi-resistant 11 12 a} 3 — — 39
Segregating ae ae —- ; — 10 63 15 1 89
Susceptible A ane — i — — _- 8 24 32
Total .. be we 11 12 23 66 23 25 160

In cases when two alternative field reactions are given for F, lines in Table 11,
efforts were later made to find out the more probable ones. Of the 23 lines under the
class R & SR or R & S, 17 were recorded as R to SR and 6 as segregating, and of the 23
lines under the class SR & § or S, 13 were recorded as susceptible and 10 as segregating.
Taking the more probable reactions, 160 F, lines in the field can be classified as 11
resistant, 29 resistant to semi-resistant, 82 segregating and 38 susceptible. If the
resistant and semi-resistant lines are treated as one class, the field data will fit 1:2:1
ratio with a P value of 0-90-0-95. The field reaction of F, lines can be best explained
by the presence of one main modifying factor and probably some minor genes which
reduce the effectiveness of the gene for resistance in Kenya 117A in the mature plant
stage. Except for the modifying factors, the seedling and the field reactions appear to
show satisfactory correlation.

Table 11 shows that the resistant F, lines, which all appear to be identical for their
reaction in the seedling stage and are almost as highly resistant as Kenya 117A, show
graded resistance in the field. Approximately 4 of the resistant and semi-resistant
lines in the field or + of all the lines are almost as resistant as Kenya 117A. This is
expected on the basis of the preceding explanation where one main modifying factor
prevents the gene for rust resistance in Kenya 117A showing its full resistance in the
field.

BY D.S. ATHWAL ANDI. A. WATSON. 9

Clark and Smith (1935) reported the presence of modifying factors as it was
difficult to obtain lines with as complete freedom from rust as the Hope and H-44 parents.
Athwal (1953) reported modifying genes in crosses of Federation with Kenya 117A.
Modifying factors probably also affect the field reaction of Kenya 744 as expressed earlier
in this paper.

In backerossing programmes carried out at Sydney University to incorporate the
resistance of Kenya 117A into commercially desirable wheat varieties, it was found
rather difficult to recover the full resistance of Kenya 117A. F, lines from the successive
backcrosses were necessary to insure that the complete resistance was present. Modifying
factors can thus influence the breeding programme. If the full resistance is not obtained
after the first backcross then it probably will not be recovered subsequently.

TABLE 12.
Reactions of F.s and F,s of the Cross Federation x Kenya 117A Against Race 38.

Reactions to Race 388.

Resistant to
Semi-resistant. | Intermediate. Segregating. Susceptible. Total.
Kenya 117A — _ Federation —_—
F,s 236 13 — 12 261
F,s 69 — 90 9 168

Relationship between races.—Selected F, lines of the cross Federation x Kenya 117A

were tested with races 126, 126B, 222AB and 222BB. All the 26 lines which were resistant
and 25 lines which were susceptible to race 222AB behaved similarly with the other three
races. Of the 12 lines which segregated with race 222AB as well as 126 and 222BB, 10
were found to be segregating against race 126B; one of the remaining two lines was
resistant and the other susceptible. From the data it appears that the same gene in
Kenya 117A gives resistance to all the races of stem rust in Australia.

TABLE 13.

Numbers of F, Lines of the Cross Federation x Kenya 117A Observed and Expected in Various Classes
when Inoculated with Two Races of Rust.

Reaction to Race 222AB.
Resistant to
Semi-resistant. | Segregating. Susceptible. Total.
Reaction to race 38:
Resistant to semi-resistant | 39 (40-5) 26 (20-25) 5 (10-125) 70 (70-875)
Segregating — 65 (60-75) 18 (20-25) 83 (81-0)
Susceptible — — 9 (10-125) 9 (10-125)
Total 39 (40-5) 91 (81-0) 32 (40-5) 162

Studies with race 38.—Federation x Kenya 117A F,s were tested with race 38. Table
12 contains the data on F.s and also a random sample of F, lines of this cross.

If 13 F, plants showing intermediate type of reaction are grouped with the resistant
class, the data will give a satisfactory fit to 15:1 ratio with a P value of nearly 0-30.
The F, data agree with 7:8:1 ratio with a P value of 0-50-0-70. Thus it appears that
Kenya 117A also has two factors for resistance to race 38.

Reactions of 162 F, families of the cross Federation x Kenya 117A to races 38 and
222AB are compared in Table 13.

10 RESISTANCE POSSESSED BY TWO KENYA WHEATS TO RACES OF PUCCINIA GRAMINIS TRITICT,

In Table 13 all the lines that were resistant to race 222AB gave the same reaction
against race 38. Of the lines that were segregating and susceptible with race 222AB,
some bred true for resistance against race 38. It shows that one of the two factors in
Kenya 117A for resistance to race 38 is the same as the one giving resistance to race
222AB and the other is independent of it. The figures within brackets in Table 13
indicate the number of lines expected on the basis of this hypothesis. The observed
agrees with the expected with a P value of 0-30-0-50.

Studies on the genetic relationship of the resistance of two Kenya wheats.

Fs of a cross between Kenya 744 and Kenya 117A were tested with race 222AB in
the glasshouse. All the plants were as resistant as the parents in the seedling stage and
nearly as resistant in the field. F,s of a reciprocal cross also behaved in the same
manner.

In the F, generation 1036 plants of the cross Kenya 744 x Kenya 117A and 380 plants
of the reciprocal cross were tested with race 222AB in the glasshouse, but no susceptible
segregate was found. Both the crosses showed segregation for mildew reaction.

TABLE 14.

Reactions of Kenya 744 Kenya 117A Fs and Federation x Kenya 117A Fs to Race 122.

Reactions to Race 122.
Cross and |

Generation. | |
Resistant to |
| Semi-resistant. | Segregating. Susceptible. | Total.
| Kenya 117A — Kenya 744 —
Kenya 744 Kenya 117A F,.. 355 — 123 478
Kenya 117A — Federation =
Federation x Kenya 117A F,.. | 37 93 35 165

P value for 3:1 ratio for the F, data lies between 0-70 and 0-80.—P value for 1: 2:1 ratio
for the F, data lies between 0-20 and 0-30.

Of the 1036 F, plants, 343 were transplanted in the field and their reactions to stem
rust were recorded under conditions of heavy infection. The plants segregated for awn
and other morphological characters. Whereas in the glasshouse the resistant reaction
of all the plants was equal to the parents, in the field some plants showed moderate to
semi-resistant and even intermediate type of reaction. A considerable amount of rust
developed on the plants giving intermediate type of reaction but no fully susceptible
plant was observed. The two parents maintained their resistance. Of the 343 F, plants
in the field, 319 were as resistant or nearly as resistant as parents, 15 showed moderate
to semi-resistance and 9 plants were intermediate for their reaction to stem rust.
One hundred and thirty-one plants representing different field reaction (117 resistant, 10
moderately to semi-resistant and 4 intermediate) were harvested and their progenies
tested with race 222AB in the seedling stage. All the 131 F, families bred true for
resistance and they were all identical in their breeding behaviour in the seedling stage.
The explanation for this is that the genes for resistance to race 222AB and consequently
other Australian races are allelic in Kenya 744 and Kenya 117A. Modifving factors
which are capable of influencing the field reaction of the two Kenya varieties are
probably non-allelic.

As the gene for resistance to race 222AB in Kenya 744 and in Kenya 117A also gives
resistance to race 38 no segregation for resistance to this race is expected in crosses
between these two varieties. Such tests therefore will reveal nothing about the relation-
ship of the second gene each of these wheats appears to possess against race 38. The
reactions to different races of stem rust in Table 1, however, show that the two above
varieties have at least one non-identical gene for rust resistance.

BY D. S. ATHWAL ANDI. A. WATSON. 11

Kenya 744 is fully susceptible to races 122 and 24 at all temperatures tested, whereas
Kenya 117A gives a reaction ranging from resistant to semi-resistant. F.s of the cross
Kenya 744 x Kenya 117A and F, lines of Federation x Kenya 117A were tested with race
122. The data in Table 14 show that a single dominant factor in Kenya 117A governs
resistance to race 122 to which Kenya 744 is susceptible.

TABLE 15.

Numbers of F', Lines of the Cross Federation x Kenya 117A Observed and Expected in Various Classes
when Inoculated with Two Races of Rust.

Reactions to Race 222AB.

|

Resistant to |
Semi-resistant.| Segregating. Susceptible. Total.

Reaction to race 122:
Resistant to semi-resistant 10 (10) 22 (20) 6 (10) 338 (40)
Segregating 21 (20) 51 (40) 19 (20) 91 (80)
Susceptible 8 (10) 15 (20) 8 (10) 31 (40)
Total 39 (40) 88 (80) 33 (40) 160

Reactions of 160 F, lines of the cross Federation x Kenya 117A to race 122 and
222AB are compared in Table 15. The data show that the gene in Kenya 117A giving
resistance to race 122 is quite independent of the gene for resistance against race 222AB.
On the basis of independent segregation of the two genes in Kenya 117A, the breeding
behaviour of F, lines for reactions to races 122 and 222AB can be fitted with a ratio
Of 1:2:1:2:4:2:1:2:1. The figures in parenthesis in Table 15 show the calculated
frequencies.

If the observed and the expected frequencies in Table 15 are compared, the P value
lies between 0-50 and 0-70 indicating a good agreement.

TABLE 16.
Comparison of the Reactions of F, Lines of the Cross Federation x Kenya 117A to Race 122 and 38.

Reaction to Race 122.

|
Resistant to
Semi-resistant. | Segregating. Susceptible. Total.
}
Reaction to race 38:
Resistant to semi-resistant 33 26 8 67
Segregating ne 3 67 19 8¢
Susceptible oe BS 1 — 7 8
Total Le es Bi 93 34 164

Koala is one of the varieties being used at Sydney University to incorporate the
resistance of Kenya 117A into a commercially desirable genotype. Koala is susceptible
to race 122 and 222AB. Sixteen families of the cross Koala? x Kenya 117A which bred
pure for resistance to race 222AB, were tested against race 122. Of the 16 families,
4 were resistant, 3 segregated and 9 were found to be susceptible to race 122. This
indicates that new strains incorporating the resistance of Kenya 117A to Australian
stem rust races would not necessarily show as wide a resistance as Kenya 117A.

In Table 16, F4 lines of the cross Federation x Kenya 117A are compared for their
reactions to race 122 and 38.

12 RESISTANCE POSSESSED BY TWO KENYA WHEATS TO RACES OF PUCCINIA GRAMINIS TRITICT,

It appears from the table that one of the two genes in Kenya 117A against race 38
is the same as the one giving resistance to race 122. One line in Table 16 which was
semi-resistant to race 122 was recorded as susceptible to race 38. The reaction of this
line could not be re-checked, but this discrepancy is believed to be a mistake probably
due to the effect of high temperature at which the tests were carried out against race 38.
Three lines which were resistant to race 122 but segregated with race 38 can be expected
on the basis of chance. The remaining 160 lines fit the expected ratio of 4:2:6:1:2:1
with a P value of 0:30-0:50, which is satisfactory.

Of the two factors in Kenya 117A against race 38, one gives resistance to race
222AB and the other gives resistance to race 122.. On this basis, if the reactions of
F, lines are known against race 122 and 222AB, their reactions to race 38 can be

TABLE 17.

Symbols for Genes Controlling Rust Resistance in Kenya Wheats.

Symbol Previous Symbol Other Varieties Known to
Variety. | Suggested. Particulars. (if Any). Have the Same Gene.
Kenya 743. Sr. Controls resistance to race | F (Watson, 1952). | Eureka, Kenya 1304 (RL1373
Kal 126, 222AB, ete. Minn. 2693), Sweden 12380,
Kenya 1034, Kenya 1037,
(Watson and Waterhouse,
1949).
Kenya 744. Sr. Controls resistance to races | H (Watson, 1952). | Allelic with one gene in Kenya
Kb1 126, 126B, 222A B, 222BB, 117A—Sr. 4d (present paper).
| 38, etc. Kb1
Sr. b Controls resistance to race 38. Allelic! to one gene in Kenya 745.
Ke2 :
Kenya 745. Sr. Controls resistance to races | K,(Watson, 1943).| Kenya 1035, Kenya 1049, Kenya
Kel 126, 126B, etc. 1053, Gabo, Yalta, Kendee,
Charter (Watson and Water-
house, 1949).
Sr. Controls resistance to race 38. | K,.(Watson, 1943). | Allelic! to one gene in Kenya 744.
Ke2
Kenya 117A | Sr. d. Controls resistance to races I (Watson, 1952). | Allelic with one of the genes in
1347. Kbl1 126, 126B, 222AB, 222BB ] Kenya 744—Sr.
and 38. > Ka, Athwal, Kb.1.
Sr. Controls resistance to race | 1953. (Present paper).
Kdl 38 and 122. J

1 Fs of the cross Kenya 744 x Kenya 745 did not show any segregation for susceptible plants against race 38.

predicted. Any line which is resistant to one or both of the races 122 and 222AB
would be resistant to race 38. Lines which segregate against one or both races should
also segregate against race 38. Only those lines would be susceptible to race 38 which
show susceptibility to each of the other two races. The reactions of 156 F, lines of the
cross Federation x Kenya 117A to races 122, 222AB and 38 showed the appropriate
correlation.

It can be concluded that the second gene in Kenya 744 which gave resistance
against race 38 but not race 122 or 222AB must be non-identical to the second gene in
Kenya 117A which was operative against race 38 and 122 but not race 222AB. It is also
probable that genes in Kenya 744 and Kenya 117A governing resistance to four races
of Australian stem rust are identical.

Designation of Genes for Stem Rust Resistance in Kenya Wheats.
Ausemus et al. (1946) proposed the symbols Sr. 1, Sr. 2, etc., for genes responsible
for resistance to stem rust. These workers also recommended that a letter subscript, .
preferably the first letter of the variety, be used as a temporary designation for the
factors until they can be checked against previously established ones.
Kenya wheats have contributed several non-allelic genes for resistance to stem
rust. Taking into consideration the recommendations of the Committee on Nomenclature

BY D. S. ATH WAL AND I. A. WATSON. 13

of Genetic Factors (Ausemus et al., 1946), the writers suggest that these genes be
designated according to the following scheme. K would show that the gene had first
been found in a wheat from Kenya, letters a, b, c, etc., would indicate varieties within
these introductions, and the subscripts 1, 2, 3, etc., would represent different loci when
more than one is known in a variety. Where sufficient evidence is not available at
present about allelic genes being identical, alphabetical superscripts will be used.
According to this scheme the genes so far identified in Kenya wheats are listed in
Table 17.

SUMMARY AND CONCLUSIONS.

Kenya 744 and Kenya 117A were each found to have a single dominant gene for
resistance to 126, 126B, 222AB and 222BB races of stem rust. A satisfactory correlation
was observed between the seedling and mature plant reaction to these races in crosses
of Federation with these varieties, but resistance in the field appeared to be influenced
by modifying factors.

Two dominant factors in Kenya 744 and in Kenya 117A governed resistance to
race 38. One factor for race 38 in each of these two varieties was the same as giving
resistance to races 126, 126B, 222AB and 222BB. The second factor in Kenya 117A
also gave resistance to race 122 to which Kenya 744 was susceptible. It can be concluded,
therefore, that the second factor in Kenya 117A is genetically different from either of
the two factors in Kenya 744.

Kenya 117A is a variety having resistance to wheat stem rust over a wide geo-
graphical area. The genetical studies herein reported give some evidence that independent
genes may operate to give resistance against races in those different areas. The gene
present in this variety which is operative against the Australian rusts is allelic with
the one present in Kenya 744 and is probably identical with it.

Since the second gene present in Kenya 117A and that in Kenya 744 are of no use
against the Australian rusts these genes can be disregarded for wheat breeding purposes
in this country.

The genetical similarity between these two rust resistant parents enables us to
predict that, if a new race of rust occurs in Australia capable of attacking either Festival
or Dowerin, the lines drawing their resistance from Kenya 117A will also be attacked.
It is doubtful whether both varieties should be used concurrently in programmes to
obtain stem rust resistant varieties.

Acknowledgements.
Grateful acknowledgement is made for financial assistance in this work from the
University Research Grant, and from the Commonwealth Rural Credits Development
Fund of the Commonwealth Bank.

References.

ABBASI, FE. H., 1949.—Inheritance of resistance to stem rust in crosses of Premier and Kenya
wheats. Ind. J. Genet. and Pl. Breeding. 9: 7-1T.

ATHWAL, D. S., 1953.—Gene interaction and the inheritance of resistance to stem rust of wheat.
Ind. J. Genet. and Pl. Breeding (in press), Vol. 13, No. 2.

AUSEMUS, E. R., WorRzELLA, W. W., HARRINGTON, J. B., and Reitz, L. P., 1946.—A summary of
genetic studies in Hexaploid and Tetraploid wheats. Jour. Amer. Soc. Agric., 38: 1082-1099.

AUSEMUS, E. R., SUNDERMAN, D. W., Hsu, K. J., and SmirH, D. H., 1953.—Report of the

_ International Wheat Stem Rust Conf., Winnipeg, Canada, 1953 (Mim.).

CLARK, J. A., and SMITH, G. S., 1935.—Inheritance of stem rust reaction in wheat. Jour. Am.
Soc. Agric., 27: 400-407.

HASANAIN, S. Z., 1949.—Resistance of certain Kenya wheats to Puccinia graminis tritici race
15B as a trigenic character. Absts. Pap. Agri. for Sect. 1st Pakistan Sci. Conf., 1949, pp. 43
(Abst.). (P.B.A. 1950, 20: 240.)

SINGLE, W. V., 1952.—Conf. Cereal Breed. and Geneticists, Wagga, N.S.W., 1951 (Mim.).

STAKMAN, EH. C., and LEvINE, M. N., 1938.—Analytical key for the identification of physiologic
races of Puccinia graminis tritici U.S. Depart. Agri. Div. Cereal Crops and Diseases and
Minn. Agri. Exp. Sta. (Mim).

VALLEGA, J., 1951.—Consideration of race 15B of Puccinia graminis tritici. Inform. Invest.
Agric. (IDIA) B. Aires, 1951, 45:14-17 (Abst.). (P.B.A. 1952, 22: 1098.)

14 RESISTANCE POSSESSED BY TWO KENYA WHEATS TO RACES OF PUCCINIA GRAMINIS TRITICT.

WATPRHOUSE, W. L., 1952.—Australian Rust Studies. IX. Physiologic race determinations and
survey of cereal rusts. Proc. LINN. Soc. N.S.W., 77: 209-258.

-, 1952.—Australian rust studies. X. Further work with Khapli Emmer wheat, an

outstanding source of stem rust resistance. Proc. LINN. Soc. N.S.W., 77: 331-336.

Watson, I. A., 1941.—Inheritance of resistance to stem rust in crosses with Kenya varieties of
Triticum vulgare Vill. Phytopath, 1941, 31, 6: 558-560.

, 1943.—Inheritance studies with Kenya varieties of Triticwm vulgare Vill. PRoc. LINN.
Soc. N.S.W., 68: 72-90.
——, 1952.—Conf. Cereal Breed. and Geneticists, Wagga, N.S.W., 1951 (Mim.).

WATSON, I. A., and SINGH, D., 1952.—The future for rust resistant wheat in Australia. Jour.
Aust. Inst. Agri. Sci., 18: 190-197.

WATSON, I. A., and WATERHOUSE, W. L., 1945.—A third factor for resistance to Puccinia
graminis tritici. Nature, Lond., 1945, 155: pp. 205.

; , 1949.—Australian Rust Studies. VII. Some recent observations on wheat
stem rust in Australia. Proc. LINN. Soc. N.S.W., 74:113-131.

————

MACADAMIA TERNIFOLIA F. MUELL. AND A RELATED NEW SPECIES.
L. A. S. JoHNSoN, National Herbarium of New South Wales, Botanic Gardens, Sydney.
[Read 31st March, 1954.]

Synopsis.

The two chief economic species of the Proteaceous genus Macadamia, ‘“‘Queensland Nut’,
have been much confused in the past. The distinctions between these are worked out and it is
shown that the correct name for one is M. ternifolia whereas the other, hitherto undescribed, is
now named WM. tetraphylla.

There are in cultivation two common species of Macadamia (Proteaceae), both
known under the names of “Queensland Nut’, “Popple Nut’, and variants of the latter.
These have been greatly confused, and misleading statements are common concerning
the difference, or lack of it, between them. This has been due to several causes; firstly,
semi-juvenile stages of one species resemble the mature stage of the other in the
possession of toothed leaves; secondly, two states of the first species have been described
under two different names; and thirdly, the second species has not been described or
named at all. :

The first species under consideration is the type species of the genus, M. ternifolia
F. Muell., of which M. integrifolia Maiden & Betche is a synonym. This has toothed
leaves in its earlier stages and often flowers in those stages, but in the fully mature
stage the leaves are entire (whence the name M. integrifolia). This species is found in
Queensland from Maryborough south to Beechmont and possibly in New South Wales,
and various strains are cultivated. The second species, which has toothed leaves at all
stages, has been wrongly known as M. ternifolia in New South Wales and overseas. It
appears to grow naturally only on the far North Coast of New South Wales and the
adjoining part of Queensland, but is also commonly cultivated. Those who know both
species as living plants, calling the first WM. integrifolia and the second (wrongly) M.
ternifolia, have naturally been puzzled by statements (e.g. that of Francis, 1951, p. 91)
to the effect that it is doubtful whether M. integrifolia is even varietally distinct from
M. ternifolia. These statements are, however, technically quite accurate as made by
those who have (correctly) applied the name WM. ternifolia to dentate-leaved states of the
first species.

The species may be distinguished as follows:

1. Leaves mostly in whorls of 4, always regularly spinose-dentate with about 35-40 teeth
on each side, subsessile or on petioles not longer than 0-2 em., lamina 7-25 em. long,
+ truncate at the base, with 13-20 pairs of main lateral nerves. Inflorescence markedly
loose-pubescent, often arising from older wood as well as new shoots, flowers Somewhat
purplish, seed surface somewhat wrinkled or with shallow depressions .... M. tetraphylla.
1*. Leaves in whorls of 3 or sub-alternate, never in whorls of 4, in earlier stages irregularly
spinose-dentate with up to 10 teeth on each side, in later stages entire, petiolate with
petioles 0-4-1-5 cm. long, lamina 5-15 em. long, tapered at the base, with 7-12 pairs of

main lateral nerves. Inflorescence sparingly puberulous, usually near the ends of
slender shoots, flowers cream, seed surface smooth ...................... M. ternifolia.

MACADAMIA TETRAPHYLLA L. Johnson, sp. nov.
Typification: Lismore, N.S.W., T. G. Hewitt, IX.1909 (N.S.W. 25513), fl. HoLoryre.
Arbor mediocris staturae vel frutex altus, saepe multicaulis, cortice dense lenti-
cellato. Ramuli verticillati, teretes dense lenticellati. Folia ramuli uniuscujusque
Superiora quaternata, infima opposita, media ternata, sessilia vel subsessilia (petiolis
0-2 cm. longis, brevioribusve) oblongo-oblanceolata, 7-25 cm. longa, 2-:0-4:5 cm. lata,
patentia, rigidiuscule coriacea, novella plus minusve colorata, glabra vel infra in costa

16 MACADAMIA TERNIFOLIA F. MUELL. AND A RELATED NEW SPECIES,

media praesertim sparsissime pilosa pilis fuscis vel pallidis perpaucis, costa media
prominens praesertim infra, pares nervorum primariorum lateralium 13-20, venuli
coarctate reticulati et praesertim supra prominuli; apex laminae subacutus vel acutus,
mucronatus, basis plus minusve truncata vel arcte contracta non sensim attenuata;
margines semper regulariter antrorse spinoso-dentati dentibus utrinque 35-40, spinis
0-15-0-2 em. longis.

Inflorescentiae axillares pseudoracemosae patentes vel subnutantes omnibus partibus
pubescentibus, 12-30 ecm. longae, rhachidis parte infima pedunculiforma 2-5 cm. longa.
Bracteae minutae caducae, irregulariter dispositae vel quasi-verticillatae, quaeque flores
duos in pedicellis patentibus circiter 0-2—0-3 cm. longis subtendens. Perianthium pallide
lilacinum 0:6 em. longum extus pubescens, intus glabrum, limbo post anthesin revoluto;
filamentorum pars libera circiter 0-025 cm. longa, antherae 0-1 cm. longae, pallidae,
longitudinaliter dehiscentes; glandulae in discum cupuliformem coalitae; ovarium
sessile villosulum, stylus cum ovario 0-8—-1-0 cm. longus viride pallens infra pubescens
supra glaber ad apicem anguste claviformis, stigma verum terminale minutum. Folliculus
in pedicello brevissime crassiusculo, apiculato-globosus circiter 2-0-—>3-0 em. diametro,
glaber, paulo rugulosus, tarde dehiscens. Semen unicum apiculato-globosum, 1-5-—2-0 cm.
diametro, subfuscum vel laete castaneum, aliquanto nitidum, plus minusve rugosum vel
laeviusculum, edule.

Specimens examined.—Queensland: Moreton District: Numinbah Valley, 2 miles
south of Beechmont road turn-off, J. H. Beaumont, 1.1954 (NSW. No. 26573), “furthest
north of M. tetraphylla in this valley’, fr.; Advancetown, Numinbah Valley, J. H.
Beaumont, i.1954 (NSW. No. 26574), veg.; Natural Arch, Numinbah Valley, J. H.
Beaumont, 1.1954 (NSW. No. 26572), fr. New South Wales: North Coast: Near
Mullumbimby, J. Farrell, 17.ix.1919 (NSW. No. 25504), fl.; Burringbar, Pope, ii.1897
(NSW. No. 25503), old infl.; Ballina to Bangalow, J. H. Maiden and J. L. Boorman,
1.1903 (NSW. No. 25505), young fr.; Nimbin, HE. Cheel, 13.ix.1926 (NSW. No. 25502), fl.
buds; Lismore, Rothwell, xi.1906 (NSW. No. 25500), veg; Lismore, T. G. Hewitt, ix.1909
(NSW. No. 25513), fl. Holotype; Richmond River, (NSW. No. 25506), fl.; Clarence
River, (NSW. No. 25501), fil.; without locality, —— (NSW. No. 25507), ff.
Cultivated: Botanic Gardens, Sydney, E. Betche, ix.1901 (NSW. No. 25508), “pink-
flowering”, fl., ix.1901 (NSW. No. 25509), “‘white-flowering’’, fl., L. A. S. Johnson, ii.1951
(NSW. No. 25512), “bushy shrub 12 feet, branching from base, fl. were mauve or pale
lilac”, fr.; Ashfield, E. Cheel, 28.x.1929 (NSW. No. 25511), veg.

The distribution of this species and the occurrence of hybrids are discussed under
M. ternifolia. The maximum size of the species in its natural habitat, lowland rainforest
or rainforest margins, is not known, though quite massive trunks are said to be formed.
In cultivation it is often quite shrubby, several-stemmed from the base, and of densely
bushy habit. The characters which distinguish it are given in the key above, and also
under M. ternifolia. Variation in thickness of testa is notable, especially in cultivated
plants. The flowers are usually pale lilac but occasionally pink or whitish. Occasional
specimens are found, especially in cultivation, in which the leaves on most branches
remain ternately rather than quaternately arranged, but these plants are absolutely
typical in other respects and the ternate arrangement does not indicate hybridity unless
accompanied by other intermediate features. On all such plants I have been able to find
some shoots exhibiting the characteristic tetraphyllous condition.

This is the more common of the cultivated species in New South Wales, and in this
State as well as in Hawaii it has been regarded as M. ternifolia. Mueller’s type, however,
and the M. ternifolia of Queensland botanists, is the following species.

MACADAMIA TERNIFOLIA F. Muell.
In Trans. Phil. Inst. Vict., ii, 1858: 72.
Typification: “In forests on the Pine River of Moreton Bay. Hill and Mueller.”
Part of this material, without precise indication of locality or collector is NSW. No.
25491 and agrees entirely with Mueller’s plates (1.e.).

BY L. A. S. JOHNSON. 17

Synonymy: Helicia ternifolia F. Muell., Fragm. Phytogr. Austral., II, 1860: 91; VI,
1868: 191. (As published in 1860 it was without any reference to Macadamia ternifolia,
and though apparently based on the same material, was described as a new species.)

Macadamia integrifolia Maiden et Betche in Proc. Linn. Soc. N.S. Wales, XXI,
1897: 64. (Holotype: NSW. No. 25495, see below.)

M. ternifolia F. Muell. var. integrifolia (Maiden at Betche) Maiden et Betche, l.c.,
XXIV, 1899: 150.

Note.—The description given by Bentham, Fl. Austral., V, 1870: 406, under WM.
ternifolia is based on material of both this species and M. tetraphylla. Many subsequent
interpretations followed this.

The chief characters by which this species differs from M. tetraphylla are set out in
the key above. Additional differences are the more slender, less prominently lenticellate
branchlets, the usually greater breadth of the leaves proportional to their length, the
paler coloration of young leaves, the occasional production of racemes on leafless shoots
resulting in a compound inflorescence and the usually somewhat longer pedicels.

Specimens examined.—Queensland: Moreton District: See under Typification, above;
Goonabah Creek, foot of Tamborine Mountain, W. Binsted per J. H. Beaumont, i.1954
(NSW. No. 26568), veg.; North Beech Mountain (Beechmont), J. H. Beaumont, i.1954
(NSW. No. 26570), ‘wild tree, in pasture (cleared rainforest)’, veg.; The Pocket,
Numinbah Valley, J. H. Beaumont, i.1954 (NSW. No. 26571), veg.; Beech Mountain,
above The Pocket, J. H. Beaumont, 1.1954 (NSW. No. 26569), “Furthest south that I have
seen M. ternifolia in south-east Queensland”, fr. Cultivated: Botanic Gardens, Sydney,
EH. Cheel, 20.viii.1925 (NSW. No. 25496), fl. buds, —— x.1896 (NSW. No. 25497), fl. buds,
, Xi.1896 (NSW. No. 25498), juvenile, L. A. S. Johnson, 5.ii.1954 (NSW. No. 26582),
“bushy tree (single stem but branching low and stooling) 15-20 ft., infructescences
pendulous, hidden amongst foliage. Young shoots not coloured.” fr.; Oahu, Hawaiian
Islands, H. M. Curran, No. 117, iv.1911 (NSW. No. 26687), fl.

M. ternifolia, like the preceding species, is found in lowland rainforest but its
distribution is more northerly. At the present time plants are definitely known only as
far south as the Numinbah Valley, north of the Macpherson Range. Dr. J. H. Beaumont
states that he has not been able to find any plants of this species south of this area.
However, the following specimens bear New South Wales localities: North Coast:
Murwillumbah, Greer, iv.1931 (NSW. No. 25492), old infl.; Wilson’s Creek, Richmond
River, C. Moore No. 32, —— (NSW. No. 25493), fl. buds; Lismore, R. White, x.1926
(NSW. No. 25494), fl.; Camden Haven, C. Moore? 1850-60 (NSW. No. 25495), fl.

Nos. 25492 and 25494 could well be from cultivated trees, but No. 25493 bears the
note “plentiful” which would seem to indicate a natural occurrence, though Dr. Beaumont
states that only M. tetraphylla is known in the Wilson’s Creek district. This specimen
has a few of its leaves in whorls of four and there is a suggestion that M. tetraphylla
may have played a part in its ancestry, though it is much closer to WM. ternifolia and
certainly not a first generation hybrid. No. 25495 presents a problem since it is rather
unlikely that the species would have been cultivated at Camden Haven at this early date.
Possibly there was a confusion of localities by the collector. Camden Haven is much
further south than any other known locality for either species.

M. ternifolia apparently attains a height of 70 feet or so under optimum conditions,
but is frequently a much smaller plant, and often several-stemmed. The variability in
leaf-dentition for which it is notorious seems to be largely a matter of physiological age.
The toothed leaves are always quite different from those of M. tetraphylla. Occasional
whorls of four leaves may be found in very young plants, but their occurrence in mature
individuals is an indication of hybrid origin.

It seems very likely that M. minor F. M. Bail. (in Queensl. Agric. Journ., XXV,
1910: 11) with which M. lowii F. M. Bail. (l.c., KXVI, 1911: 127) appears to be
synonymous, is merely a local form or an ecotype of M. ternifolia, but field study is
desirable to settle this point. I have seen the following specimen: Kin Kin, Queensland,

D

18 MACADAMIA TERNIFOLIA F. MUELL. AND A RELATED NEW SPECIES.

Cc. T. White, i.1916 (NSW. No. 25725), fr. This does not appear to differ significantly
from many specimens of WM. ternifolia.

Hyprips.—The following specimens are intermediate in all respects between WM.
ternifolia and M. tetraphylla. One group is found somewhat to the north of the known
limit of the latter species and it is possible that M. tetraphylla has been “swamped” in
that area by hybridization with M. ternifolia. The other group is from the known zone
of contact of the two species. The specimens exhibit considerable variation which
suggests the existence of a hybrid swarm, involving more than one generation of
hybridity. The hybrids show intermediate conditions in petiole length, leaf shape and
dimensions, number of marginal teeth and of lateral nerves, roughness of seeds and in
general aspect. Flowers have not been seen. The number of leaves in the whorl is
usually three, occasionally four. Variation on one plant may be considerable.

Specimens examined.—Queensland: Moreton District: Goonabah Creek, foot of
Tamborine Mountain, W. Binsted per J. H. Beaumont, i.1954 (NSW. No. 26576), fr.,
(NSW. No. 26577), veg., (NSW. No. 26575), veg. [nearer M. ternifolia]; Walla Walla,
Tamborine Creek, H. Welch per J. H. Beaumont, i.1954 (NSW. No. 26578), “old trees
over 100 years old. 7 ft. in circumference at base of clump”. fr.; Numinbah Valley,
J. H. Beaumont, i.1954 (NSW. No. 26580), veg.; Beechmont (cult.), J. H. Beaumont,
1.1954 (NSW. No. 26579), fr.

It is noteworthy that these hybrids are of restricted occurrence and that there is no
general intergradation of the species which differ in a number of characters and are
quite constant throughout their area, though apparently interfertile.

My thanks are due to Dr. J. H. Beaumont of the Hawaii Agricultural Experiment
Station for renewing my interest in these two species of Macadamia, which are the
basis of the Macadamia nut industry, and amongst the natural populations of which
Dr. Beaumont is seeking promising plants for use in breeding and selection programmes.

Reference.
FRANCIS, W. D., 1951.—Rain Forest Trees of Australia.

19

EXPERIMENTAL CROSSING OF AEDES (STHGOMYIA) PSEUDOSCUTELLARIS
THEOBALD AND AEDES (STEGOMYIA) POLYNESIENSIS MARKS
(DIPTERA, CULICIDAE).

By A. R. WoopHIL1,

Dept. of Zoology, University of Sydney.

[Read 28th April, 1954.]

Synopsis.

The results of experiments in crossing Aédes (Stegomyia) pseudoscutellaris Theobald with
Aédes (Stegomyia) polynesiensis Marks are described, and it is shown that the two species
are not genetically isolated, since under laboratory conditions small numbers of fertile hybrids
are produced using either species as the female parent. These hybrids have been bred through
to the Fe generation and appear to be normal in every way.

INTRODUCTION.

In previous papers (Woodhill, 1949, 1950) accounts have been given of crossing
experiments with various species and sub-species of the “‘scutellaris group” of mosquitoes
from Melanesia and Polynesia. Recently E. N. Marks (1951) described a new species,
A. polynesiensis, with a wide range from Fiji through the Ellice Islands, Samoa, the
Society Islands and the Tuamotu Islands. This had previously been confused with
A. pseudoscutellaris, which is recorded only from Fiji; the two species are separated
by Marks on small differences in the scaling of the scutal angle and in the setae of
the basal lobe of the male coxite.

During 1953 eggs of A. polynesiensis were received from Papeete, Tahiti, and a
laboratory colony was successfully established, thus enabling crosses between the two
species to be carried out.

CrossING EXPERIMENTS.

The general technique was similar to that described in previous papers, approxi-
mately 130 females and 150 males being used in each experiment, and records being
kept until egg production ceased, this usually covering a period of from two to three
weeks. The following table gives the results.

Experiment Number of Eggs Percentage of
Number. Species and Sexes. Produced. Eggs Hatched.
il © Pol. xo Pseud. 1223 4-8
2 2 Pseud. x g Pol. 229 5-6
3 © Pol. xo Pseud. 1723 36-6
4 2 Pseud. xg Pol. 530 ; 19-4
5 © Pol. xd Pol. 6582 95-2
6 © Pseud. x 3d Pseud. , 5948 91-8
DISCUSSION.

It will be seen that crossing occurred between the two species to a limited extent,
using either species as the female parent, but that more eggs were produced from
? Pol. x g Pseud. than in the reciprocal cross. Total egg production and the percentage
ef viable eggs was much lower than in a normal colony as exemplified in Experiments
5 and 6.

DD

20 EXPERIMENTAL CROSSING OF AIDES.

The hybrids produced were bred through to the Ff, generation and behaved normally
in every way. Dr. E. N. Marks kindly examined a series of F, hybrids from 9 Pol. x ¢
Pseud. and these showed considerable variation, but on the whole were intermediate
between the parent forms. It would appear therefore that the two species are not
genetically isolated, and observations indicate that the low egg production and low
viability of the eggs is due to infrequent copulation as compared with a normal colony.
As shown previously (Woodhill, 1949), virgin females of this group will deposit sterile
eges but in greatly reduced numbers.

However, where the two species occur together, as in Fiji, it is possible that some
crossing would take place under natural conditions and that intermediate forms would
occur in the field. It is highly desirable that a close study of the two species in Fiji
should be made, and it is hoped that a colony of Fiji polynesiensis will be established
in order to compare their behaviour (in crossing with pseudoscutellaris) with those
from Tahiti. On the above evidence it would perhaps be more accurate to consider the
two forms as sub-species, rather than to give them full specific rank.

Acknowledgements.
The author’s thanks are due to Dr. Backhouse, Dr. Bonnet and Mr. Nelson for
obtaining eggs from Tahiti and Fiji, and to Dr. Marks for examining the hybrids.

References.

Marks, E. N., 1951.—The Vector of Filariasis in Polynesia: A Change in Nomenclature.
Ann. Trop Med. & Parasit., 45, No. 2: 137-140.

WoopHILuL, A. R., 1949.—A Note on Experimental Crossing of Aédes (Stegomyia) scutellaris
scutellaris Walker and Aédes (Stegomyia) scutellaris katherinensis Woodhill. Proc. LINN.
Soc. N.S.W., 74, Pts. 5-6: 224-226.

, 1950.—Further Notes on Experimental Crossing within the Scutellaris Group of
Species. Proc. LINN. Soc. N.S.W., 75, Pts. 5-6: 251-258.

21

CYTOLOGICAL STUDIES IN THE MYRTACEAE. IV.
THE SUB-TRIBE EUCHAMAELAUCINAE.
By S. SmirH-WHiITtTE,
Department of Botany, University of Sydney.
(Forty-one Text-figures. )
[Read 28th April, 1954.]

Synopsis.
Chromosome numbers are reported for 40 species, representing all genera in the subtribe

Euchamaelaucinae.
_ One or several reducing series in chromosome number occur within the tribe, the haploid

complement of 11, found in Chamaelaucium and some species of Verticordia, probably being
primitive. Both Verticordia and Darwinia include several different basic numbers, and both
may prove to be unnatural groups. In Verticordia, polyploid series occur on several of the

base numbers.

It is necessary to accept the Huchamaelaucinae as a cytologically specialized and derived
group within the family. As a consequence, the author’s earlier view of the phylogeny of the
family is destroyed, and Andrews’ earlier conception is strengthened.

INTRODUCTION.

In previous papers (Smith-White, 1942, 1948, 1850) it has been claimed that the
basic chromosome number in the Myrtaceae is x = 6. Although the haploid number in
the family is almost uniformly eleven, this view was initially founded upon observations
of the occurrence of secondary association between bivalent chromosomes during meiosis
in pollen mother cells and the presence of several nucleoli in meiotic interphase and
microspore nuclei. The inference gained some support from records of haploid 12 in
the related family Melastomaceae, and appeared to be greatly strengthened by the
discovery of an actual haploid 6 in Darwinia and Actinodium.

As a consequence of the hypothesis it was necessary to infer the cytological
primitiveness of the Chamaelaucoideae, or at least of the Huchamaelaucinae, and perhaps
polyphyletic origins of the 11-genom in each of the three tribes of the family. Andrews’s
view (1913) of the evolutionary development of the family was considered to be untenable.

Atchison (1947), unaware of the cytological constitution of Darwinia, believed
that the remarkable uniformity of chromosome number in the family indicated the
ancient and monophyletic origin of the family itself, and of its characteristic chromo-
some constitution.

Studies by Thomas and Revell (1946) have greatly reduced the significance of
secondary association for the inference of secondary polyploidy, and the value of
nucleolar number for such inference has also been reduced with an increasing knowledge
of the nucleolar cycle in nuclear division. Consequently the author’s earlier view now
rests mainly upon the occurrence of the haploid number of 6 in Darwinia.

OBSERVATIONS.

A summary of the genera and species of the Euchamaelaucinae which have been
examined, and of the chromosome number determinations made, is given in Table 1.
For convenience, earlier records are also included. Identifications have been made
by reference to the Blackall Flora of Western Australia (unpublished), which was
made available by Dr. Grieve, and the species names used are those given by Gardner
(1931). Taxonomically, the genus Verticordia is difficult, and doubt of some identi-
fications is unavoidable. Herbarium material has been lodged in the Departmental
Herbarium.

22 CYTOLOGICAL STUDIES IN THE MYRTACEAE. IV,
TABLE 1.
Chromosome Numbers in the Euchamaelaucinae.
Chromosome
Numbers.
Genus and Species. Ace. State." Localities. Reference.
Number.?
n. 2n.
Actinodium cunninghamir — W.A Albany. 6 12 S.-W. 1950.
Schau.
Darwinia, Sect. Genetyllis.
D. hypericifolia (Turez) | 52/73. W.A Toolgenup, Stirling Ra. 6 — Text-fig. 1.
Domin.
D. speciosa (Meissn.) Benth. | $.A.52/105. W.A Blackwood, S8.A.,° ex. 6 12 Text-figs. 2, 3.
Stirling Ra.
D. collina Gardn. .. | S.A.52/102. W.A. 59 5 a 6 = Text-fig. 4.
D. leiostyla (Turez) Domin. | $.A.52/9. W.A. Blackwood, S.A.,° ex. 6 —
Bluff Knoll, W.A.
D. citriodora (Endl.) Benth. | 50/47. W.A. Lesmurdie. 6 12 S.-W. 1950.
D. taxifolia A. Cunn. 50/63. N.S.W. Blackheath. 6 = S.-W. 1950.
var. intermedia (Cunn.) | 50/15. N.S.W. Kuring-gai, Loftus. 6 — S.-W. 1950.
Cheel.
var. grandiflora Benth. 50/18. N.S.W. Helensburgh, Bulli. 6 = S.-W. 1950.
var. biflora Cheel 50/23. N.S.W. Epping, Mt. Colah. 6 — S.-W. 1950.
D. grandiflora Baker 50/28. N.S.W. Berowra. 6 = S.-W. 1950.
D. “ mesembryanthemoides *** | 53/8. N.S. W. Kariong. 6 — Text-fig. 5.
D. vestita (Endl.) Benth. 50/105. W.A. Albany, Frenchman’s 9 — Text-figs. 6, 7.
Bay, Esperance Road.
D. pauciflora Benth. 52/40. W.A Mullewa. 9 = Text-figs. 8, 9.
D. diosmoides (D.C.) Benth. | 50/112. W.A Albany, Frenchman’s _— 12
Bay.
D. fascicularis Rudge 49/72. N.S. W. Gordon, Loftus, Kuring- 6 12 S.-W. 1950.
gai.
D. sp. §.A.52/7. S.A. Blackwood, S8.A.,°> ex. = 12 Text-fig. 10.
south-east districts.
Darwinia, Sect. Schau-
mannia.
D. micropetala Benth. 8.A.52/6. S.A. Pt. Lincoln. 7 14 Text-fig. 11.
Homoranthus virgatus | 53/13. N.S. W. Evans Head. 9 = Text-figs. 12-14.
A. Cunn.
HL. flavescens A. Cunn. — N.S. W Wellington.* 9 —
H. darwinioides (Maid. & | 51/12. N.S. W Lee’s Pinch. 9 = Text-figs. 15, 16.
Betche) Cheel (=Ryl-
stonea cernua Baker).
Verticordia, Sect. EKuverti-
cordia.
V. densiflora Lindl. 50/. W.A. Cannington. 11 =
V. plumosa (Desf.) Domin. 50/82. W.«.A. Gleneagle, Albany. 11 — Text-fig. 17.
50/106.
$.A.52/103.
V. sp. (near plumosa)® 50/87. W.A. Nedlands. 11 — Text-fig. 18.
V. brownii (Desf.) D.C. 50/64. W.A. Coorow, Southern Cross. 9 = Text-figs. 19, 20.
50/123.
V. nitens (Lindl.) Schau. .. | 50/34. W.A. Gnangara. 8 — Text-figs. 21, 22.
50/88.
V. grandiflora Endl. 50/58. W.A. Coorow. 6 12 Text-fig. 23.
V. chrysantha Endl. $.A.52/110. W.A. Blackwood, $.A.° 16 —_— Text-fig. 24.
V. preissii Schau. .. 50/60. W.A. Coorow. 8 a Text-fig. 25.
50/63.
50/124.

1 Reference to Herbarium sheets, Botany Dept. Herbarium.
2 State to which species are native.
* Transplants from nature, grown in Miss E. Ashby’s garden, Blackwood, $.A.
The name used is from an unpublished manuscript by the late Mr. Blakely.

4 An undescribed species.

5 Transplants from nature, grown by Mr. G. Altofer, of Wellington, N.S.W.
6 7Plants growing in the gardens of the University of Western Australia.

The material supplied by Miss
Baird in 1949, and mentioned in Smith-White (1950), was probably taken from these plants.

bo
(ee)

BY S. SMITH-WHITE.

TABLE 1.—Continued.

Chromosome Numbers in the Euchamaelaucinae.

Chromosome
Numbers.
Genus and Species. Acc. State.” Localities. Reference.
Number.?
| | n. 2n.
Verticordia acerosa Lindl. 50/46. W.A. Lesmurdie. 8 — Text-figs. 26, 27.
V. oxylepis Turez.’ .. | 50/86. W.A. Nedlands. 8 —_ Text-figs. 28, 29.
V. huegellii Endl. .. ao tt COVE W.A. Lesmurdie. 8 — Text-figs. 30, 31.
V. insignis Endl. 5 on || OEY W.A. Coorow. 9 — Text-figs. 32, 33.
V. habrantha Schau. bo || Beal. W.A. Chester Pass. 18 — Text-figs. 34, 35.
V. roew Endl. te .. | 52/56. W.A. Busselton. 9 —= |
V. monadelpha Turez. .. | S.A.52/1010. W.A. Blackwood, S.A.* c. 18 —
Verticordia, Sect.
Catocalyptra. |
V. picta Endl. tee oc |) BOB}, W.A. | Coorow. 11 — Text-fig. 36.
| 50/59.
V. drummondii Schau. .. | 52/. W.A. Morowa. 11 =
V. pennigera Endl. ee sAE52ie W.A. Blackwood, $.A.? 11 —
V. grandis Drummond .._| 50/122. W.A. Watheroo. 22 — Text-figs. 37, 38.
Pileanthus peduncularis — W.A. Coorow. 11 = Text-figs. 39, 40.
Endl.
Chamaelaucium uncinatum — W.A. Sydney, cult. 11 = 8.-W. 1950.
Schau.
Ch. drummondii Meissn ... —- W.A. Coorow. 11 —
Ch. axillare F. Muell. .. | 9.4.52/112. W.A. Blackwood, S.A.* } dak Text-fig. 41.

* Reference to Herbarium sheets, Botany Dept. Herbarium.

* State to which species are native.

* Transplants from nature, grown in Miss E. Ashby’s garden, Blackwood, S.A.

* An undescribed species. The name used is from an unpublished manuscript by the late Mr. Blakely.

° Transplants from nature, grown by Mr. G. Altofer, of Wellington, N.S.W.

° 7 Plants growing in the gardens of the University of Western Australia. The material supplied by Miss
Baird in 1949, and mentioned in Smith-White (1950), was probably taken from these plants.

The genus Darwinia was divided into two sections by Bentham (1866). In the
section Genetyllis, haploid 6 is the usual chromosome number, but two species, D. vestita
and D. pauciflora, which are morphologically allied, are each haploid 9, a number
characteristic of the small eastern genus Homoranthus. The section Schaumannia is
represented only by the South Australian D. micropetala, which is unique in having
a haploid number 7.

In the genus Homoranthus, H. flavescens was considered by Bentham to be identical
with H. virgatus. H. darwinioides was given generic status under the name Rylstonea
by Baker (1898), and both Baker and Maiden and Betche (1898) believed that it was
closely related to Verticordia. All species are haploid 9.

Verticordia was also divided into two sections by Bentham—Huverticordia and
Catocalyptra. In the former, gametic numbers of 6, 8 and 11, and ploidy on base 8
and base 9 are reported, but the occurrence of these numbers does not conform to the
taxonomic arrangements given either in Bentham’s Flora Australiensis or in the
unpublished Blackall flora of Western Australia. The confusion in the previous report
of the chromosome number of V. plumosa (Smith-White, 1950) has been resolved. The
material forwarded by Miss Baird in 1949, which gave a count of n = 8, was probably
taken from the plant 50/86, growing in the gardens of the University of Western
Australia. This plant has been identified by Mr. C. A. Gardner, Government Botanist
in Western Australia, as V. oxylepis. The cultivated material forwarded by Miss Eardley,
of the University of Adelaide, was grown by Miss Ashby at Blackwood, and is similar to
material of V. plumosa collected at Albany.

CYTOLOGICAL STUDIES IN THE MYRTACEAE, LV;

In
tetraploid, n =

99
- 24.

and Chamtielaucium.

eo — © @ ®
o 6.6 See oe 4S) @°e
Se % @° 7. SBC o °
; 2 ~ = A
3
°¢ 6, : @
66° “o6's 28°
@ to es
5 6 5

Text-figures 1-23.

1, Darwinia hypericifolia,

1-A. 2, D. speciosa, 1-M.
4, D. colina, 1-M. 5, D

5, D. “mesembryanthemoides’’, 1-M.

3, D. speciosa, somatic metaphase.
8, D. pauciflora, 1-M. 9, D. pauciflora, 2-M.

6, D. vestita, 1-M. 17, D. vestita, 2-M.

10, Darwinia sp. 52/7. 11, D. micropetala, 1-M.
12, Homoranthus virgatus, 1-M. 13, H. virgatus, 1-A, side view. 14, H. virgatus, 2-M. 15,

H, darwinioides, diakinesis. 16, H. darwinioides, 1-M. 17, Verticordia plumosa, 2-M. 18, Verti-
cordia sp. 50/87, 1-M. 19, V. brownii, 1-M. 20, V. brownii, 1-M, side view. 21, V. nitens, 1-M.

The large bivalent is particularly conspicuous. 22, V. nitens, 2-M. 23, V. grandiflora. (All
figures x 2000.)

Verticordia Section Catocalyptra, the gametic number is 11, and V. grandis is
Eleven is also the base number for the smaller genera Pileanthus

BY S. SMITH-WHITE. 25

DISCUSSION.

Chromosome numbers in the Huchamaelaucinae have significance for the systematics
of the subtribe, and have an important bearing on the interpretation of the phylogeny
of the Myrtaceae, and its historical development in Australia.

The taxonomy of the Huchamaelaucinade.—In Darwinia (n = 6, 7, 9) and Verticordia
(n = 6, 8, 9, 11) there is a similar pattern of variation in chromosome number, and

Text-figures 24-41.

24, V. chrysantha, 2-M. 25, V. preissii, 1-M.
Side view. 28, V. oxylepis, 1-M. 29, V.-oxvylepis, 2-M. 30, V.
huegelli, 2-M. 32, V. insignis, 1-M. 33, V.
habrantha, 2-M. 36, V. picta, 2-M.

peduncularis, 1-M. 40, P. peduncularis, 2-M. 41, Chamaelauciwm awillari, 2-M. (All figures
x 2000.)

26, V. acerosa, 1-M. 27, V. acerosa, early 1-A,
huegellii, diakinesis. 31, V.
insignis, 2-M. 34, V. habrantha, 1-M. 35, V.

37, V. grandis, 1-M. 38, V. grandis, 2-M. 39, Pileanthus

26 CYTOLOGICAL STUDIES IN THE MYRTACEAE, Iv,

Homoranthus (n = 9) connects the two larger genera at the 9-chromosome level.
Morphologically, Darwinia and Verticordia are distinguished by calycine characters.
In the former, the five sepals are entire, but in the latter the sepals are deeply divided
each into five plumose lobes, or into numerous hair-like parts. In Homoranthus, each
sepal is produced into a single plumose bristle, or, in H. darwinioides, into five bristles.
Both morphologically and cytologically, Homoranthus links the two larger genera.
Bentham (1866) indicated that the genera of the Huchamaelaucinae could be arranged
in a linear morphological series as follows: Actinodium, -Darwinia § Genetyllis, -Darwinia
§ Schaumannia, -Homoranthus, -Verticordia § Euverticordia, -Verticordia § Catocalyptra,
-Chamaelaucium.

On the basis of cytological and morphological evidence, two alternative views of the
phyletie status of Darwinia and Verticordia may be proposed.

(a) The two genera, as at present established, are sound, and the chromosome
number series present in each represent cases of parallel chromosomal evolution.

(0) The numerical patterns belong to the same series, and the phyletic status of
the two genera is unsound. Parallel evolutionary development of calycine form in
several lineages is required. A multiple origin of particular characters in a group
involves no serious genetic difficulty, since various species must carry many homologous
genes with similar mutational tendencies. The occurrence of similar mutations in
different species of a genus is well known in Drosophila, Triticum, Hordeum, Gossypium,
and in fact in almost every genus which has been investigated genetically. Such
parallelisms provide the basis for the “Law of Homologous Variations” enunciated by
Vavilov (1935). On the other hand, parallel series in chromosomal evolution have been
demonstrated in Crepis (Babcock, 1949).

Choice between the two alternatives must await further cytological and systematic
study, but the following suggestions are offered towards the revision of the group.

1. No taxonomie revision of Darwinia or Verticordia should be attempted without
the incorporation of cytological data.

2. The 9-chromosome species in Darwinia should be separated from that genus,
and perhaps combined with Homoranthus.

3. The genus Verticordid, as at present accepted, is a composite one and should
be divided. The subdivision of the genus given by Bentham, however, is not likely
to provide a satisfactory basis.

Verticordia grandiflora may be more akin to species of Darwinia, and the 11-chromo-
some species in Verticordia § Catocalyptra are allied to Chamaelaucium, and should be
separated from other species of the present genus.

The phylogeny of the Myrtaceae.——Any consideration of the origin and evolution
of the Myrtaceae must take into account the origin of the 11-chromosome genom which
is generally characteristic of the family. The basic chromosome number in the Rosales,
from which most systematists derive the Myrtales, is seven. Darlington and Mather
(1949) consider that the primitive haploid number in the Angiosperms was seven, and
Stebbins (1950) has suggested that many groups of woody Angiosperms with haploid
numbers of 11, 13 and 19 have been derived from basic numbers of 6 or 7 by amphi-
diploidy and secondary polyploidy.

The basic number of 11 in the Myrtaceae is almost certainly a derived one, but
any hypothesis on the evolutionary development of the family must depend on the
status conceded to the 6-genom found in Darwinia. Three possibilities may be considered.

1. The 6-chromosome constitution in the Huchamaelaucinae is primitive, and the
11-genom has arisen only once. This view would mean that the 11-chromosome species
of the Huchamaelaucinae, the Leptospermoideae and the Myrtoideae are all related at
the 11-chromosome level. The view cannot reasonably be maintained.

2. The 6-chromosome genom is primitive, and the 11-chromosome genom has arisen
on several or many occasions in different lineages. Independent origins in the Eucha-
maelaucinae, in the other subtribes of the Chamaelauceae, in the Leptospermoideae,
and in the Myrtoideae would be a minimum requirement. The method of origin would
involve polyploidy to give a haploid number 12, followed by the loss of a chromosome

BY S. SMITH-WHITE. 27

or of a centromere. This view would lead to an expectation of haploid numbers of 12
within the family, but Atchison’s report (1947) of this number in some species of
Eucalyptus, even if correct (Smith-White, 1948), is of no significance in this respect.
It would indicate only a secondary derivation of 12 chromosomes from the 11-genom
characteristic of Hucalyptus. The hypothesis does not take into account the intermediate
haploid numbers of 7, 8 and 9 found in Darwinia and Verticordia, and must now be
rejected.

3. The 11-chromosome genom is basic to the family, although having an earlier
derivation from a lower number, and the chromosome complements of 6, 7, 8 and 9
found in the Euchamaelaucinae have arisen by reduction. Reducing series in chromosome
numbers are not infrequent in many groups of Angiosperms (Darlington and Mather,
1949: Darlington and Janaki, 1944; Stebbins, 1950). Darlington (1937) has shown how
such series arise from structural change and centromere loss. Tobgy (19438) has actually
demonstrated this process in Crepis.

Hypothesis 3 is, consequently, much the most probable. On this basis, the chromosome
number series in Darwinia and Verticordia represent either a single, or several parallel
reducing series, 11—~(10)—9—8—>7—6, and the morphological series presented earlier
should be read from right to left, i.e. from Chamaelaucium to Actinodium, and not in
the opposite direction.

It follows, if this hypothesis is accepted, that the Huchamaelaucinae represent a
group of genera which are both morphologically and cytologically specialized and
derived. Andrews’s view on the evolution of the Australian Myrtaceae receives con-
firmation rather than denial from the cytological evidence, and Atchison’s conclusion
that the cytological data support an ancient and monophyletic origin of the family
is strengthened.

A new problem, however, arises. Why is it that, whereas chromosome constancy,
with a low frequency of even such easy changes as polyploidy, has been a feature
of the Myrtaceae, one small subtribe, with a relatively restricted distribution, has been
able to undergo repeated steps in chromosome diminution? Perhaps the answer to this
problem may be found in Darlington’s suggestion (Darlington and Janaki, 1944) that
liability to chromosomal alterations is dependent on the characteristics of chromosome
architecture. The presence of heterochromatin close to the centromeres would perhaps
allow a better chance of survival for individuals suffering centromere loss.

The historical development of the Huchamaelaucinae in Australia.—Crocker and
Wood (1947) have suggested that the floras of eastern and western Australia have been
isolated from each other practically continuously since the Miocene period. On the
basis of cytological and geographical evidence and inference, it is possible to relate
the development of the Huchamaelaucinae to this time scale.

The absence of vicarious species within genera, and the existence of vicarious genera
such as the eastern Homoranthus and the western Verticordia, in the eastern and western
regions suggest that considerable evolution at the species level has occurred since the
development of the main intracontinental migration barrier. If slow rates of evolution
are characteristic of woody Angiosperms (Stebbins, 1950), the migration barrier must
be relatively ancient in origin, i.e. Miocene rather than Pleistocene, and the thesis of
Crocker and Wood is supported. However, the 6-chromosome genom in Darwinia and
the 9-chromosome genom in the Darwinia-Homoranthus-Verticordia complex are pan-
Australian, and consequently their origins must have antedated this barrier, i.e. they
must be placed in the early Miocene or in the Hocene. Harly in the development of the
Huchamaelaucinae, after the floristic isolation of Australia, and after their differentiation
from the rest of the Myrtaceae, but before the floristic bisection of the continent, the
subtribe must have suffered important changes in chromosome structure permitting
the occurrence or survival of interchange and numerical diminution.

SUMMARY.
Additional chromosome number determinations are reported for the subtribe EHucha-
maelaucinae in the family Myrtaceae. Data are now available for 40 species, representing
all genera and subgenera in the tribe.

28 CYTOLOGICAL STUDIES IN THE MYRTACEAE, IV,

The data demonstrate the existence of one or several reducing series in chromosome
number within the subtribe, and especially in the larger genera Darwinia and Verticordia.
These genera may prove to be unnatural groups, and there is need for their revision on
a basis of both morphological and cytological evidence.

It is necessary to accept the Euchamaelaucinae as a cytologically specialized and
derived group within the family. As a consequence, the author’s earlier view of the
phylogeny of the family is destroyed, and Andrews’s conception is supported.

Acknowledgements.

This work was made possible by grants from the University Research Fund in
1950 and 1952, for study in Western Australia. Grateful acknowledgement is made to
Dr. Brian Grieve and his staff, of the Botany Department, University of Western
Australia, for the use of the facilities of that department. I am in debt to Mr. Stoate,
Conservator of Forests, W.A., and to Mr. J. Harding, of the Forestry Department, for a
ereat deal of assistance in field work. My thanks are also due to Miss EH. Ashby, of
Blackwood, South Australia, for access to her collection of Western Australian plants.

References.

ANDREWS, E. C., 1913.——The Development of the Natural Order Myrtaceae. Proc. LINN. Soc.
N.S.W., 38:529-568.

ATCHISON, E., 1947.—Chromosome Numbers in the Myrtaceae. Amer. J. Bot., 34: 159-164.

Baspcock, BE. B., 1949.—The Genus Crepis.. Pt. 1. Univ. Calif. Publ. Bot., Vol: 21.

Baker, R. T., 1898.—On a Supposed New Genus of the Natural Order Myrtaceae. Proc. LINN.
SoCs INGSHW 233 7@BWSn qi.

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

Crocker, R. L., and Woop, J. G. (1947).—Some Historical Influences on the Development of
the South Australian Vegetation Communities and their Bearing on Concepts and Classi-
fication in Ecology. Trans. Roy. Soc. S. Aust., 71: 91-136.

DARLINGTON, C. D., 1937.—Recent Advances in Cytology. Second Ed. Churchill, London.

and JANAKI AMMAL., 1944..-A Chromosome Atlas of Cultivated Plants. Allen and
Unwin, London.
and MatuHemr, K., 1949.—Hlements of Genetics. Allen and Unwin, London.

GARDNER, C. A., 1931.—Enumeratio Plantarum Australiae Occidentalis. Govt. Printer, Perth.

MAIDEN, J. H., and BretcHst, E., 1898.—Four New Species of N.S.W. Plants. Proc. Linn. Soc.
INESAVWEM EZ onlin ato):

SMITH-WHITE, S., 1942.—Cytological Studies in the Myrtaceae. i. Microsporogenesis in Several
Genera of the Leptospermoideae. Proc. LINN. Soc. N.S.W., 67: 335-342.

, 1948.—Cytological Studies in the Myrtaceae. ii. Chromosome Numbers in the Lepto-
spermoideae and Myrtoideae. Proc. LINN. Soc. N.S.W., 73: 16-36.

, 1950.—Cytological Studies in the Myrtaceae. iii. Cytology and Phylogeny in the
Chamaelaucoideae. Proc. LINN. Soc. N.S.W., 75: 99-121.

STEBBINS, G. L., 1950.—Variation and Evolution in Plants. Colwmbia University Press, New
York.

THOMAS, P. T., and REVELL, S. H., 1946.—Secondary Association and Heterochromatin Attrac-
tion. i. Cicer arietinum. Ann. Bot., N.S., 9:159-164.

Topey, H. A., 1943.—A Cytological Study of Crepis fuliginosa and C. neglecta and their Fi
Hybrid, and its Bearing on the Mechanism of Phylogenetic Reduction in Chromosome
Number. Jour. Genet., 45: 67-111.

Vavitov, N. I., 1935.—The Law of Homologous Series in the Inheritance of Variability.
Translation by Kk. S. Chester. Chron. Bot., 13:56-94 (1950).

29

A NEW SPECIES OF HIBBERTIA ANDR. FROM WESTERN AUSTRALIA.
By A. T. HorcnHxtss,
Department of Botany, University of Sydney.
(Plate i; four Text-figures. )
[Read 28th April, 1954.]

Synopsis.
A new species of the genus Hibbertia Andr. of the family Dilleniaceae from Western
Australia is described.

The genus Hibbertia Andr., with about 140 species, is the largest of some 11 genera
in the family Dilleniaceae. Hoogland (Blumead, 7, 1952, 1-145) places Hibbertia in the
subfamily Dillenioideae. It is generally regarded as possessing some of the most
primitive floral characters of that subfamily. Some species with completely actino-
morphic flowers, with stamens o and all fertile, and carpels ~, show this particularly
well. The genus Hibbertia is known to occur in Madagascar, western and southern
New Guinea, within the peripheral areas of Australia and in New Caledonia.

HIBBERTIA SERRATA, Sp. NOV.

Type specimen: Pemberton, W.A., A. T. Hotchkiss, September 2, 1953; holotype in
N.S.W. National Herbarium, isotypes in Sydney University Herbarium, Department of
Agriculture Herbarium, Perth, Cornell University Herbarium.

Frutex erectus, pubescens, odoratus 1-1-5 m. altus; caules teretes: rami majores
crassi et erecti; secundarii rami graciliores et ascendentes: cortex ramorum seniorum
rubido-fuscus, pubescens, longitudine diffissus, denique particulis longis laxis disjunctis.
Folia simplicia, exstipulata, alterna, sessilia, mollia, rhomboido-cuneata, leviter pubes-
centia infra, nervo medio et marginibus prope base villis simplicibus praecipue pubes-
centibus; margines plani; nervus medius subtus conspicuus et protusus; nervi secundarii
angulum 45° nervo medio formantes; basis folii tumidula et leviter amplectans. Folia
ramorum majorum 5-9 cm. longa x 12-27 mm. lata, marginibus serrato-dentatis, apicibus
mucronulatis; folia ramorum secundariorum 15-25 mm. longa x 3-4 mm. lata, marginibus
serratis vel integris. Inflorescentia brevis, arcta, axillaris; florae parvae, sessiliae,
2 (rare 3) bracteolis latis, fuscis, seariis, cuspidatis ca. 3-5 mm. longis x 2:5 mm. latis,
et 3 bracteis linearibus, foliaceis ca. 5-7 mm. longis; sepala 5, petala includentia; sepala
ovata, acuminata 6-9 mm. longa x 2:5-3 mm. lata, 3-nervata, pubescentia, imbricata,
persistentia et fructos juvenes includentia; petala 5, late obovata, incisa, flava, caduca,
8 mm. longa x 6 mm. lata; stamina ca. 20 (5-27), libera, ca. 4 mm. longa, circum
gynoecium in fasciculis cum carpellis alternantibus disposita; poris 2 obliquis sub-
apicalibus; staminodia vel nulla vel ca. 3-6 (2-17), filiformes, clavata vel in forma
semi-staminis, staminibus indefinite disposita. Carpella 2 vel 3, libera, ca. 5 mm. longa
x 2 mm. lata, glabrata, ovata, sessilia, folliculos maturitate formantia; stylus ca. 2-5 mm.
longus, ab axe directus; stigma simplicissima; placentatio parietalis; ovuli 2; arillus
albus, prope semen maturum tegens; semen globosum, laeve, fulgidum, rubido-fuscum,
ca. 1:5 mm. diametro.

An erect pubescent, odorous shrub, 1-1-5 m. high; stems terete, the major branches
coarse and erect, the secondary branches more slender and ascending; the bark on
older branches reddish-brown, pubescent, splitting longitudinally and finally becoming
loosened in long strips. Leaves simple, exstipulate, alternate, sessile, soft, slightly
pubescent beneath with simple hairs especially on the midrib and on the margins near
the base, rhomboid-cuneate to elliptical; margins flat; midrib prominent, protruding
on the lower surface; secondary veins forming an angle of 45° with the midrib; base

30 A NEW SPECIES OF HIBBERTTA,

of the leaf swollen and slightly stem-clasping. Leaves on major branches 5-9 ecm. long
x 12-27 mm. broad, margins serrate-dentate, apices mucronulate; leaves on secondary
branches 15-25 mm. long x 3-4 mm. broad, margins serrate or entire.

Flowers rather small, closely sessile, crowded in short axillary inflorescences; below
each flower are 2 (3) small, brown, scarious, broad and cuspidate bracteoles about
3-5 mm. long x 2-5 mm. broad; below which are 3 linear, foliaceous bracts about 5-7 mm.
long; calyx of 5 sepals enclosing the petals: sepals ovate, acuminate, 6-9 mm. long x
2-5-3 mm. broad, 3-nerved, pubescent, imbricate, persistent and closed again in the
young fruit except for the scarious spreading tips; petals 5, broadly obovate, deeply
notched, 8 mm. long x 6 mm. broad, yellow, ephemeral; stamens about 20 (5-27), free,

B

Text-fig. 1.—Hibbertia serrata, sp. n. <A, Large leaf from a major stem, natural size;
B, Undersurface of same leaf showing venation; C, D, small leaves from a secondary branch,
natural size.

about 4 mm. long, placed all round the gynoecium, usually in clusters alternating with
the carpels; dehiscence of stamens subapical from 2 oblique pores opening from 4
chambers; staminodia either none or about 3-6 (2-17), filiform, clavate or half-stamen
in size, placed indefinitely with the stamens. Carpels 2 or 3, free, about 5 mm. long x
2mm. broad, glabrous, ovate, sessile, ripening into a follicle; style about 2-5 mm. long,
directed abaxially, stigma very simple; ovules two, attached to opposite sides of the
adaxial placenta, usually only one maturing; arillus white, nearly covering the mature
seed; seed spherical, smooth, shiny, reddish-brown, about 1:5 mm. in diameter.

The crushed leaves and stems have a peculiar odour suggestive of certain shrubs in
the Compositae. The arrangement of the stamens at once places this species in the
section Cyclandra. Large, flat, thin, pinnately-veined leaves with serrate margins are
seldom found in this section (or the genus Hibbertia) outside the subsections Hemi-
hibbertiae and Bracteatae. The possession of bracts and also staminodia are features
common to both these groups, but the closely sessile condition of the flowers indicates
a position in the Bracteatae for this species. Within this group it is allied to H. montana

BY A. T. HOTCHKISS.

Text-figs. 2-4. Hibbertia serrata, sp. n.

2—A, Broad, brown bracts beneath the flower (flower removed); A’, The same bracts

beneath a flower bud; B, Linear foliaceous bracts; C, Bud arising above a bract; D, Sepals
of flower bud.

3.—A, Diagram showing arrangement of floral parts, staminodia in solid black; B, Petal;

C, Foliaceous bract; D, Broad, brown bract; H, A stamen with 3 staminodia; F, Sepal showing
part of indumentum; G, Carpel.

4.—A-A, Transverse section through anther after anthesis; B, Stamen showing oblique,
subapical pores.

31

32 A NEW SPECIES OF HIBBERTIA,

Steud. and the closely related species H. argentea Steud., H. pilosa Steud., and H.
quadricolor Domin, all of which differ from H. montana principally in the amount, type
and position of their indumentum. H. serrata is quite different from the typical form of
H. montana whieh has a low straggling habit with several stems; a conspicuous
indumentum, especially on the sepals; small, thick, mostly entire leaves; and large
flowers with large, conspicuous, shining, brown bracts. H. montana var. major Benth.
(including H. ovata Steud. placed here by Bentham in FU. Aust.) is described in Fl. Aust.
as ‘‘Larger and more branched and often more or less hirsute, with long spreading hairs.
Leaves usually larger, on luxuriant shoots often above 14 or 2 in. long, broad and
coarsely toothed, almost all less contracted at the base than in the normal form, and
closely sessile. Some specimens of this variety look so different from H. montana, with
their coarse habit, long spreading hairs, and broad-toothed leaves, that I at first retained
them as a distinct species; but they pass into the smaller forms through so many
intermediates, that I have been quite unable to draw any definite limits between them.”
Later authors have commented on this var. major and have perhaps enlarged the original
concept of Bentham to include an increased amount of variation. Domin (1921-1922.
Vest. Kral. Ces. Spol.) says “var. major Benth. . .. W.A.:—Yallingup and Cape
Naturaliste, A. A. Dorrien-Smith. A very peculiar form entirely distinct from the
typical H. montana. Its leaves are of very thin texture, on the upper surface glabrous
and slightly hairy with scattered appressed hairs beneath, oblong-elliptical, the middle
leaves about 8 cm long and almost 2-5 em broad, very coarsely dentate. The scarious
scales are broad but very short and fall off very early. The sepals are slightly hairy,
the petals are large. Very near to our form are the specimens collected by Cecil Andrews
(among granite rocks by Helena River, 17 m. east of Perth, 8 XI. 1902, W.A. FI. First
Coll. No. 14), which, however, are still more hairy and have somewhat smaller flowers.
Hardly another Hibbertia-species could be found in which the indumentum, the size,
form, dentation, texture of leaves, the size of the bracts and flowers would vary in such
a degree; nevertheless Bentham is right in uniting all these forms, because there are
transitions between them.” It is of course impossible to say what Bentham would have
done with these forms collected by Dorrien-Smith and Andrews. Bentham made no
reference to the texture of the leaves, the size or falling of the bracts, or the size of the
flowers in distinguishing var. major from the typical H. montana. Ostenfeld (Dansk.
Vidensk. Selskab. 1921) also refers to specimens of var. major collected at Darlington
(No. 302; 28 Aug. 1914, Cecil Andrews) and says: ‘“‘This variety is very different from
the true H. montana, and ought perhaps be taken as a separate species.” He gives no
description of the specimens.

- About half the flowers examined contained no staminodia but in other flowers from
the same plant staminodia were numerous and occasionally exceeded the stamens in
numbers. The staminodia have no definite position being placed usually outside the
stamens but occasionally among or inside the stamens. The significance of staminodia
in this group is uncertain. Bentham in Fl. Aust. records staminodia in no species of
the Bracteatae except H. mylnei. Domin records staminodia in H. quadricolor Domin.
The presence of staminodia in some species of the Bracteatae may illustrate a transitional
tendency between the Bracteatae and the Hemihibbertiae where staminodia are always
present. The presence of staminodia in H. serrata is correlated with the number of
carpels as seen in the accompanying table.

I have concluded that H. serrata should be erected as a new species very distinct
from H. montana and its allies. H. montana var. major is intermediate in some
characters but does not approach H. serrata as closely as it does to H. montana with
which it appears to intergrade. H. serrata would include the tall robust forms which
have thin, coarsely serrate leaves, short inconspicuous bracts, smaller flowers and more
or less indumentum. It is probable that the specimens of Dorrien-Smith and Andrews,
cited by Domin whose description indicates that they have thin leaves and differ in only
minor ways from the type of H. serrata, also belong to this new species.

BY A. T. HOTCHKISS. ah

The first collections of blooming material were made on 2nd September, 1953; for -
later collections of flowering material made on 8th January, 1954, I am indebted to Mr.
John Meachem, Department of Forestry, Pemberton. This species was found in the karri
forest (Hucalyptus diversicolor F.v.M.) on the sandy coastal area in south-west Western
Australia, the type locality being near the roadside in a reserve of karri trees near
Pemberton, Western Australia.

No. of Carpels No.of Stamens No. of Staminodia

per Flower. per Flower per Flower (by
(by clusters). stamen clusters).
iL. 2 8-13 0—0
2: 2 7-10 0-0
3. 2 6-7 0-0
4. 2 6-9 0-0
5. 2 6-11 0—0
6. 2 7-12 0-0
Uo 2 6-11 0-3
8. 2 6-9 0-3
8). 3 6-7-8 0—0—0
10. 3 6—6-—8 0—0—-0
ial, 3 8—2-—7-1-9 0—0—0—0-0
12. 3 7-6-7 3-1-1
Se 3 5-9-8 0-0-2
14. 3 3-4-2 2—4-6
15). 3 5-7-8 1-1-0
16. 3} 0-2-3 6—6—5
Wife 3 3-2-2 5—6—-6
18. 3 4-8-7 2-0-0
il)e 3 6—2-7 0-5-1
20. 3 4—4-6 0—4—-2

I wish to thank Mr. C. A. Gardner, Government Botanist, and Mr. R. D. Royce,
Senior Botanist, for making available to me the facilities of the State Herbarium, Perth,
and also Mr. J. Harding and the Dept. of Forestry, Perth, Western Australia, for their
kindness in providing transportation from Perth to Pemberton.

EXPLANATION OF PLATE IT.

Hibbertia serrata, sp. nov. Photograph of type specimen.
(Photograph by Woodward-Smith. )

A COMPARATIVE ACCOUNT OF THE TERRESTRIAL DIATOMS OF
MACQUARIE ISLAND.

By J. S. Bunt,! Microbiology Laboratories, School of Agriculture, The University of
Sydney.

[Read 28th April, 1954.]

Synopsis.

A brief account is given of the work carried out in this field by past expeditions and
a systematic account has been prepared comprising the species found in samples of soil, water
and fluvio-glacial sediments at Macquarie Island. The terrestrial diatoms already recorded in
the antarctic and sub-antarctic have been listed for comparative purposes following the
precedent of Peragallo (1921). Finally, the importance of these organisms in the soils has
been discussed and consideration has been given to the influence of the past and present
environment on the species occurring in the various habitats examined. Certain differences
existing between the past and present diatom floras of Macquarie Island have been indicated
and the reasons underlying the affinities, distribution and origin of these organisms in antarctic
regions have been investigated. Diagrams and photomicrographs of every species listed from
Macquarie Island have been included.

INTRODUCTION.

Far less attention has been paid to the terrestrial diatoms of the antarctic and
sub-antaretic regions than to those which occur in the oceans of southern latitudes,
although collections have been made since the earliest stages of antarctic exploration by
scientific bodies.

Probably the first examination of terrestrial species was made by Ehrenberg (1854)
from a sample of soil collected by Hooker in 1843 at Cockburn Island (64° 12’ lat. S.;
59° 41’ long. O.G.) during the voyage of the “Erebus” and “Terror” under the command
of Sir J. C: Ross. Later, in 1854, Ehrenberg published the results of his examinations
of samples from Kerguelen Island, Falkland Island and Cape Horn. As the result of
an expedition to Kerguelen Island in 1874-5 to observe the transit of Venus, some algal
material was collected by the Rev. A. EH. Haton and described by Reinsch (1876, 1879).
During the voyage of H.M.S. “Challenger” (1873-6), H. N. Moseley collected further
samples at Kerguelen Island and some at Marion Island, a description of the diatoms
being published by O’Meara (1876). From Cape Horn, Petit (1888) described 37 species
and Reinsch (1890) described 19 species from South Georgia. In 1900, Cleve published a
list comprising 62 species from Magellan.

The National Antarctic Expedition :(1901—4), under Scott, collected the first diatom
samples from the antarctic mainland in the vicinity of Cape Adare and McMurdo Bay.
The results were published by Fritsch (1912). Several forms were also gathered by
Borchgrevinck about the same time and described by Holmboe (1902). Rudmose Brown
returned material from the South Orkneys collected during 1902-4 and the types were
described by Fritsch (1911). James Murray with Shackleton’s expedition (1907-9)
collected a good deal of material at Ross Island and South Victoria Land, the systematic
account being prepared by W. and G. 8S. West (1911). Peragallo (1921) published a
valuable account of the diatoms of Graham Land and included lists of species described
from other antarctic localities by several authors. More than 240 species or varieties
from Patagonia were described by O. Miiller (1909). Fritsch (1917) described a small
number of diatoms collected by the “Terra Nova” Expedition (1910) under Scott.

1 Biologist, Australian National Antarctic Research Expedition.

BY J. S. BUNT, 35

The samples that have been examined in the preparation of this account were
collected by the author at Macquarie Island during 1951-2 whilst he was engaged as
biologist with the Australian National Antarctic Research Expedition under the leader-
ship of Mr. P. G. Law.

It will be seen from a map of the antarctic that the present collection is important
since it is representative of the only land in sub-antarctic latitudes between Kerguelen
and the southern tip of South America and the closest link with Cape Adare and South
Victoria Land from which other collections have been gathered.

Although it is suspected, for reasons to be given later, that the lists of species from
some of the other antarctic islands are not complete, it should be possible, with the
information at present available, to make certain postulations regarding the differences
and affinities existing between the diatom floras of these regions. Further, with a rather
detailed knowledge of the ecological environments of Macquarie Island, some account
may be given of the possible factors influencing the local distribution of the various
species.

- PREPARATION OF THE SPECIMENS FOR HXAMINATION.!

Subsamples of the soil, peat or mixed algal material were placed in Kjeldahl flasks
and boiled in concentrated nitric acid until all the organic matter was destroyed. The
flasks were then filled with triple-filtered tap water and the sediment allowed to settle.
The supernatant liquid was removed by decantation and the process was repeated until
the washings were no longer acid. A small quantity of the sediment was spread on a
no. 0 coverslip and dried carefully before mounting in canada balsam on a microscope
slide.

It is considered that, since mixed slides facilitate comparison between species and
thus aid in identification, the preparation of single species mounts is of doubtful value,
especially in the light of the arbitrary separation of many variable types into specific
groupings.

Each slide was scanned carefully at least four times with both high power dry and
oil immersion lenses. It is believed that very few, if any, species have been missed from
the account which follows. The table showing the degree of abundance of the various
species isolated from the soil has resulted only from estimates and is not based on
strictly quantitative methods.

NOTES ON THE SAMPLES.

a. Soils.

1. A mineral soil from the feldmark; primitive; derived from till; pH 6-8; %
organic carbon 3-6; % nitrogen (total) 0-3; C/N 12-0; soil adhering to underside of
pads of the moss, Dicranoweisia antarctica (C.M.) Par.; 1000 feet above sea level;
continually wet; soil temp. rarely higher than 4°C.

2. An organic soil from the feldmark; localized to cushions of Azorella selago Hook.
forming incomplete cover of the surrounding till; pH 6-2; % org. C 12-5; %N (total)
0-78; C/N 16-0; 700-800 feet above sea level; continually wet; soil temp. rarely higher
than 5°C. ;

3. An organic soil from the sub-glacial herbfield, Gadget Gully; water table
constantly at or, in places, above the surface; pH 7-1; % org. C 18-5; % N (total) 1-56;
C/N 12:0; 500 feet above sea level; soil temp. in Feb. 6:0°C.

4, Wet’ tussock grassland bordering beach, Hasselborough Bay; bare boggy ground
between tussocks; frequented by seal elephants; pH 4-3; soil temp. Feb. 8-4°C. Following

1 From continued direct observation of soil and water samples and in vitro studies, it is
believed that most of the species described in this paper are living constituents of the flora
in the environments from which they were collected. This opinion is further supported
by the fact that practically no frustules of the species which abound in the plankton of the
surrounding seas and which must be blown on to the island, were found in the terrestrial
samples. Fragilariopsis antarctica Castr. is a notable exception, but this species was observed
only rarely.

36 TERRESTRIAL DIATOMS OF MACQUARIE ISLAND,

figures from similar soil no. 2276 from Piper (1938): % org. C 18-6; % total N 1-58;
C/N 11-6.

5. Sub-glacial herbfield; peaty organic soil continually wet; pH 5-8; % org. C 24-1;
% total N 2-19; C/N 11-0; 750 feet above sea level; soil temp. Feb. 5-4°C.

6. Wet tussock grassland, Buckles Bay; black beach sand in partly exposed root
masses of Poa foliosa Hook. f.; well-drained area bordering beach; pH 4-3; % org. C 1-3;
% total N 0-11; C/N 11-8; soil temp. Feb. 8-0°C.

7. Small pockets soil around roots of Puccinellia macquariensis Cheesm. growing on
rocks jutting into sea, Buckles Bay; % org. C 4-4; % total N 0-69; C/N 6-4; pH 7-2;
soil temp. Feb. 8-0°C.

8. Wet tussock grassland; deep hill peat, moist to wet; pH 4-4; % org. C 44-6;
% total N 3-33; C/N 13-4; soil temp. Feb. 4-8°C; 600 feet above Half Moon Bay; Poa
foliosa Hook. f. dominant.

9. Wet tussock grassland; wet, deep, hill peat; pH 4-0; % org. C 43-6; % total N
3:33; C/N 13-1; soil temp. Feb. 4-8°C; 100 feet above Half Moon Bay; Poa foliosa
Hook. f.: Stilbocarpa polaris A. Gray co-dominants.

10. Sub-glacial herbfield; very deep waterlogged peat; pH 4-7; % org. C 39-4; %
total N 1-79; C/N 22-0; soil temp. Feb. 9-0°C; 30-40 feet above sea level, Handspike
Point.

b. Lakes and Ponds.

1. Small pools in sub-glacial herbfield, Half Moon Bay (coll. B. W. Taylor 31.1.51).

2. Source of small creek in feldmark; one mile west Mt. Elder, 500 feet above sea
level (coll. B. W. Taylor 1951).

3. Fresh-water pool in sub-glacial herbfield, Hasselborough Bay (coll. B. W. Taylor,
Sept. 1950).

4. Stagnant pool in sub-glacial herbfield, Handspike Point (April 1952).

5. Small lake on plateau, elevation 650 feet, one mile south Scoble’s Lake (coll.
B. W. Taylor, 6.2.51).

6. Stagnant pool, sub-glacial herbfield, Hasselborough Bay (coll. B. W. Taylor,
Bell).

7. Repeat of no. 6, Handspike Point (coll. B. W. Taylor, 19.3.51).

8. Stagnant pool, wet tussock grassland, north end Isthmus, (17.8.51).

9. Repeat of no. 8, (17.8.51).

10. Bottom sediment, Prion Lake, (19.1.52).

11. Shallow water, mixed algal material, Prion Lake, (21.1.52).

c. Fluvio-glacial Deposits.

1. Half-way Hill. A deposit of banded sediments regarded by Mawson (1943) as
fluvio-glacial and dating from the end of the last Ice Age. Interbedded, are several
layers of lignite and diatomaceous earth. Samples 99, i, j, k, m, n represent these strata
in order of increasing depth.

2. Lusitania Bay. Deposits probably representing the same period as those from
Half-way Hill and about 25 feet thick (mostly gravel). Samples 12B3 and 12B4 are
from the upper and lower layers of the first peaty stratum from 5 to 6 feet below the
surface. Sample 12D was taken from the second peat layer several feet lower down the
profile.

3. North Mountain. Fossil peat or lignite of unknown age in a deposit up to 2 feet
thick-and underlying the surface 2 feet of till; 800 feet above sea level. Samples 18d,
e, f, j were taken at intervals of 6 inches down the profile from 24 inches to 42 inches.

4. Flat Creek alluvial or fluvio-glacial deposits. Samples 26i, j, k, 1 and 27 a-d
represent peaty strata from the surface of the exposed creek bank to a depth of about
8 feet.

5. Raised coast terrace, Hasselborough Bay. Samples F1, 2, 3, 5, 6, 9, 10 were taken
at 6 inch intervals through the deep peat of this recently uplifted coastal area from the
surface to the point where the peat meets the old beach sediments. No diatoms were
found in samples F4, 7 and 8.

BY J. S. BUNT. 37

TAXONOMY.

As long ago as 1855, Gregory warned against the widespread practice of multiplying
specific names amongst the diatoms without adequate justification. Mangin (1914) has
discussed the considerable variation in form exhibited by certain types of diatoms
collected in antarctic waters and has shown that the frustular structure is not necessarily
constant. Similar work has been published by Wilson (1927) as a result of investiga-
tions with Cocconeis scutellum Eh. Mann (1937), in pointing to the arbitrary boundaries
which often exist between species and even genera and the “misty condition” of the
taxonomy generally, has not classified his list of antarctic diatoms into higher groups
but has arranged the genera and species into alphabetical order, “irrespective of possible
relationships”. This method has been followed in the present paper.

An examination of the list of species will show that no attempt has been made to
change established names even though they represent types with indistinct boundaries.
Explanatory notes have been added in these cases and, wherever possible, probable
affinities have been indicated. The object has been to encompass the aims of nomen-
clature review without actual name-changing which could bring about some confusion.
For comparative purposes, however, many species from the antarctic appearing as
Navicula in the literature have been given the more recent generic name Pinnularia Eh.

Diagrams or photomicrographs of every species listed from Macquarie Island have
been included with details of size, etc., in the accompanying index to enable any errors
which may have been made to be more readily detectable by future workers. The lists
of synonyms which have been appended to many of the species described include the
authority’s name in brackets following each list. The systematic account which follows
comprises all the species observed by the author in the samples examined.

List oF MACQUARIE ISLAND DIATOMS.
ACHNANTHES Bory.

Achnanthes affinis Grun.—Hustedt in Rab. Kryptogamenflora, Bd. 7, Ht. 2, p. 381.
Synonym: Microneis affinis Cleve. (Hust.)

Achnanthes Biasolettiana Grun. (Text-fig. 7).—Hust. in Rab. Krypt., Bd. 7, Ht. 2,
p. 379. Synonyms: Synedra Biasolettiana K.,.Achnanthes Biasolettiana f. minuta Grun.,
Microneis Biasolettiana Cleve ?, Achnanthes Biasolettiana var. perminuta V.H. (Hust.)

Distribution: Falkland Island, Cape Horn, Magellans. (Antarctic records.)

Achnanthes brevipes Ag. var. intermedia K. (Pl. ii, Figs. 1 and 2).—Hust. in Rab.
Krypt., Bd. 7, Ht. 2, p. 425; W. and G. S. West, Brit. Ant. Exped. Reps., vol. 1, Diyas
Pl. xxvi, figs. 126-127 (1907-9). Synonyms: Achnanthes intermedia K., Achnanthes
subsessilis K., Achnanthes multiarticulata Ag., Achnanthes turgens Eh., Achnanthes
carpensis K., Achnanthes subsessilis var. ovalis Dippel, Achnanthes loczyi Pantocsek.
(Hust.)

Distribution: Cape Adare, Ross Island, South Victoria Land, Macquarie Island
(Antarctic records).

The structure of the frustules was found to be somewhat variable, the central area
of the valves being either open or bearing an isolated line of punctae on only one side.
Sometimes forms intermediate between these forms were observed. The valves tend to
be more rhomboid than the types illustrated by Hustedt and W. and G. §S. West; the
indentations in the valve walls at the edge of the central area are not obvious.

Achnanthes exigua Grun.—Hust. in Rab. Krypt., Bd. 7, Ht. 2, p. 386. Synonyms:
Stauroneis exilis K., Stauroneis quadrata Heribaud, Stauroneis tylophora Reichelt,
Microneis exigua Cleve, Cocconeis exigua Torka. . (Hust.)

Distribution: Magellans. (Only other antarctic record.)

Achnanthes exilis K.—Hust. in Rab. Krypt., Bd. 7, Ht. 2, p. 378. Synonyms:
Achnanthes Leibleinii Ag., Achnanthidium subhungaricum Gutwinsky, Achnanthes
subhungaricum DeToni. (Hust.)

Distribution: Kerguelen. (Only other antarctic record.)

38 TERRESTRIAL DIATOMS OF MACQUARIE ISLAND,

Achnanthes lanceolata (Breb.) Grun. (PI. ii, fig. 3).—Hust. in Rab. Krypt., Bd. 7,
Ht. 2, p. 408. Synonyms: Achnanthidium lanceolatum Breb., Achnanthes haynaldi vay.
oblonga-elliptica Schwarschmidt, Stauroneis truncata Schum., Achnanthidium lanceéo-
latum var. genuina A. Mayer, Achnanthidium lanceolatum var. minima A. Mayer,
Achnanthidium lanceolatum var. conspicua A. Mayer, Achnanthes lanceolata f. typica,
f. pura and f. semipura A. Mayer, Achnanthidium lanceolatum var. rhomboidalis A.
Mayer, Achnanthes lanceolata var. maxima Per., Achnanthes pseudoantigua Per.,
Achnanthes pagesi Per. (Hust.)

Distribution: World-wide.

Hustedt lists several varieties and forms of this species which exhibits considerable
range in shape and size. Although a certain amount of slight variation has been
observed in the Macquarie Island material, it is doubtful whether identification beyond
the level of species is necessary.

Achnanthes linearis Grun.—Hust. in Rab. Krypt., Bd. 7, Ht. 2, p. 378. Synonyms:
Achnanthidium lineare W. Smith, Achnanthidium jackii Rab., Achnanthes linearis var.
jackii Grun. (Hust.)

Distribution: Magellans. (Only other antarctic record.)

Achnanthes minutissima var. cryptocephala Grun.—Sch. At., t. 410, figs. 51-53;
Hust. in Rab. Krypt., Bd. 7, Ht. 2, p. 377.

Distribution: Europe, Asia, Arctic. (From DeToni.)

Should be compared with Achnanthes exilis K., Achnanthes linearis (W. Smith)
Grun. and Achnanthes affinis Grun. The Macquarie Island material has been found to
agree best with Achnanthes minutissima var. cryptocephala Grun. but the boundary line
between this and the other three species does not appear to be very well defined.

Note.—The determination of A. affinis, A. exigua, A. exilis and A. minutissima var.
eryptocephala should not be regarded as definite. The material to which these names
were assigned was extremely fine-structured and difficult to examine even with an oil
immersion lens system. Therefore the names allocated should be taken as indicating
only the most probable taxonomic position. The existing synonymy reflects the difficulties
which surround the identification of such material. Diatoms belonging to this group
appeared to be present in all the samples examined, but only as minor constituents of
the total populations. Mention of their presence has been restricted to this section.

Achnanthes subsalsa Peterson (PI. ii, fig. 7).—Hust. in Rab. Krypt., Bd. 7, Ht. 2,
p. 401. Should be compared with Cocconeis apiculata A.S. in Sch. At., t. 198, fig. 41.

AMPHORA Ehrenberg.

Amphora delicatissima Krasske (Text-fig. 19).—Hust. in Die Siisswasser-flora
Mitteleuropas, Ht. 10, p. 346, fig. 635, (1930).

CocconEIS Ehrenberg.
Cocconeis placentula Eh. (Text-fig. 23).—Hust. in Die Stissw. Mitt., Ht. 10, p. 189,
fig. 260, (1930).
Distribution: Falkland Island. (Only other antarctic record.)

CosciINopiIscus Ehrenberg.

? Coscinodiscus spp.—Small fragments of discs were occasionally observed, but never
a complete specimen. The material has been associated for convenience with this genus,
five species of which have been recorded previously in the antarctic.

CYMATOPLEURA W. Smith.

Cymatopleura solea Breb. var. regula (Eh.) Grun. (Text-fig. 10)—Hust. in Die
Stssw. Mitt., Ht. 10, p. 426, fig. 8230, (1930); Sch. At., t. 276, figs. 10, 11 (C. regula (Eh.)
Ralfs). Synonyms: Cymatopleuram regulam (Eh.) Pritch., Cymatopleura regula (Eh.) |
Ralfs, Surirella regula Eh., Cymatopleura parallela W. Smith. (DeToni.) ‘

Distribution: Mexico, Europe. C. solea Breb. recorded from Graham Land.

BY J. S. BUNT.

(tm,

pe

Text-figs. 1-23.

1, Pinnularia nivorum Per. var? 26 x 5 w. 13 costae/ 10 uw. 2, Navicula mutica Kutz.
13 x 5 w. 10 striae/ 5 w. 3, Stauroneis pygmaea Kreiger. 24 x 5 uw. 30 costae/ 10 uw. 4, Navicula
mutica Kutz. var? 16 x 6 uw. 24 striae/ 10 uw. 5, Navicula seminulum Grun.. 21 x 6 uw. 16-18 costae/
10 uw. 6, Pinnularia microstauron (BHh.) Cleve var. Brebissonii (Kutz.) Hust. 19 x 7 uw. 20 costae/
10 w. 7, Achnanthes Biasolettiana Grun. 10 x 3:5 w. 23-25 costae/ 10 u.
Bh. 7 x 4 uw. 10 costae/ 10 pw. 9, Pinnularia (interrupta?) W. Smith.

10 w. 10, Cymatopleura solea Breb. var regula (EKh.) Grun. 40 x 7 uw. 11, Pinnularia fasciata
Lagerst. 34 x 7 mw. 20 costae/ 10 uw. 12, Frustulia rhomboides (EKh.) DeToni. 100 x 22 uw 13,
? Surirella engleri O.M. forma angustior O.M. 80 x 7 w. 14, Fragilaria bicapitata A. Mayer.
50 x 4 w. 17 costae/ 10 w. 15, Navicula mutica Kutz. var? 19 x 8 w. 15 striae/ 10 uw. 16, Pinnularia
Atwoodii Per. 30 x 9 w. 15 costae/ 10 uw. 17, Neidium affine (Eh.) Cleve. 48 x 10 yw. 28 striae/
10 uw. 18, Rhopalodia gibberula (Eh.) O.M. var baltica O.M. Length 48 uw. 19, Amphora deli-
catissima Krasske. 26 x 3 uw. 30 costae/ 10 mw. 20, Pinnularia borealis Eh. 50 x 10 uw. 6 costae/
10 w. 21, Fragilaria capucina Desm. 63 x 4:5 uw. 15 costae/ 10 uw. 22, Gomphonema angustatum

(Kutz.) Rab. var. producta Grun. 31 x 8 uw. 12 costae/ 10 w. 23, Cocconeis placentula Eh. 25 x 16 uw.
25 striae/ 10 wp.

8, Fragilaria pinnata
30 x 9 mw. 16 costae/

In all figures, 10 w = approximately 10 mm.
EE

40 TERRESTRIAL DIATOMS OF MACQUARIE ISLAND,

CyMBELLA Agardh.
Cymbella pusilla Grun. var. ? (Pl. ii, fig. 5)—Hust. in Die Stissw. Mitt., Ht. 10,
p. 354, fig. 646, (1930).
The type which occurs on Macquarie Island has consistently parallel striae and
differs in this respect from the type figured and described by Hustedt in which the
striae are somewhat radiate.

DIATOMELLA Greville.
Diatomella Balfouriana Grey. (Pl. ii, fig. 8).—Hust. in Die Stissw. Mitt., Ht. 10,
p. 214, fig. 3112; (1930); Brit: Inf, p. 810, Bl IV; figs! bij 52) Synonyms Gnammdto-
phora ? Balfouriana W. Smith, Disiphonia australis Eh. (DeToni.)
Distribution: World-wide.

DIPLONEIS Ehrenberg.
Diploneis Smithii (Breb.) Cleve. (Pl. ii, fig. 6).—Hust. in Die Stissw. Mitt., Ht. 10,
p. 253, fig. 402, (1930); Sch. At., t. 7, figs. 14-22. Synonyms: Navicula Smithu Breb.,
Navicula elliptica W. Smith. ~
Distribution: World-wide.

Eunorra Ehrenberg.

Hunotia alpina (Naeg.) Hust. (PI. ii, fig. 12).—Hust. in Die Stissw. Mitt., Ht. 10,
Dewls sa tiger2 52.) (930) aS Cle Atte mite 2 Ole TIS Se aiamos

Distribution: Magellans. (Only other antarctic record.)

The apical swellings were slightly more developed in the Macquarie Island specimens
than in the figure shown by Hustedt.

Hunotia lunaris Grun. (PI. ii, fig. 11).—Hust. in Die Siissw. Mitt., Ht. 10, p. 183,
(1930). Synonyms: Pseudoeunotia lunaris (Eh.) Grun., Synedra lunaris Eh., Exilaria
lunaris Hass., Hxilaria curvata K., Hunotia curvata Lagerst., Synedra campyla Hilse.
(DeToni.)

Distribution: World-wide.

Having 22-25 striae/ 10u, and being only 3u wide, this diatom agrees more closely with
the variety subarcuata (Naeg.) Grun., but the apices are bulbous rather than tapered.
Varietal assignation has therefore been omitted.

Eunotia trinacria Krasske. (PI. ii, fig. 10).—Hust. in Die Stissw. Mitt., Ht. 10, p. 176,
figs. 221a-d, (1930).

FRAGILARIA Lyngbye.

Fragilaria bicapitata A. Mayer. (Text.-fig. 14).—Hust. in Die Stissw. Mitt., Ht. 10,
p. 148, fig. 148, (1930).

Fragilaria capucina Desm. (PI. ii, figs. 13, 14; Text-fig. 21)—Hust. in Die. Stissw.
Wott, Jett, IO, yo, Nexes tie, IAG, (Clg e))) 3 Ire, Ihe, , 700, JBI ID, wigs, tage, 7étes Jel, SONY,
figs. 1, 2; Sch. At., t. 298; W. Smith, Synops. Brit. Diat., vol. 2, p. 22, Pl. XXXV, fig. 296.
Synonyms: Staurosira capucina Borzsc., F. ventriculus Eh., F. pectinalis Lyngb., F.
tenuis Ag., fF. rhabdosoma Eh., F. diophthalma Eh., F. multipunctata Eh., F. bipunctata
Eh., F. angusta Eh., F. scalaris Eh., F. fissa Eh., F. sepes Eh., F. pusilla Breb., F. corrugata
K., Nematoplata argentea Bory, Nematoplata capucina Bory, Bacillaria pectinalis
Nitzsch., Diatomosira pectinalis Trev., Bacillaria multipunctata Eh.

Distribution: World-wide, including Falkland Island and Marion Island in the
antarctic.

The great variability of this species is indicated by the synonymy. (From DeToni.)

Fragilaria Harrison (W. Sm.) Grun. (PI. ii, fig. 9)—Hust. in Die Siissw. Mitt.,
Ht. 10, p. 139, fig. 132, (1980); Sch. At., t. 296, figs. 6-18; W. Smith, Synops. Brit. Diat.,
vol. 2, p. 18, Pl. LX, figs. 373, 374. Synonyms: Odontidium Harrisonii W. Sm., Biblarium
leptostauron Eh., Diatoma Harrisonii Cleve, Staurosira Harrisonii Cleve. (DeToni.)

Distribution: Common in northern hemisphere.

Fragilaria pinnata Eh. (Text-fig. 8).—Hust. in Die Siissw. Mitt., Ht. 10, p. 142, figs.
141la-d, (1930); Sch. At., t. 298, figs. 47-74.

BY. ws UBUD: 4}

Fragilaria virescens Ralfs. (Pl. ii, fig. 15)—Hust. in Die Stissw. Mitt., Ht. 10, p.
1A tic 144, i930) =) Sch. “At. t. 297%, many figs, Synonyms: Ff. undata W. Sm., F.
constricta Eh., F. construentum (binodum) Eh., F. pectinalis Eh., F. equalis Heib.,
Diatoma virescens Hass., F. confervoides Grev., Diatoma sulphurescens Ag., Staurosira
equalis Cleve and Miiller. (DeToni.)

Distribution: Ross Island and South Victoria Land. (Only other antarctic records.)

F. virescens, according to Hustedt, is 12—120u long and 5-10u across the valve. The
forms examined differ from these measurements, being only 3—4u across the valve. The
length, 40—45u, is fairly constant. Occurrence is restricted to fossil peats.

Fragilaria (virescens ?) Ralfs. (Pl. ii, fig. 4)—This material is 15-204 in length,
3—4u across the valves and has 21-23 striae/10u. It represents a type quite distinct from
that listed above and occurs soth as a fossil and as a living constituent of the present
algal flora. It has been placed with some doubt as F’.. virescens.

i FRAGILARIOPSIS Hustedt.

Fragilariopsis antarctica Castr. (Pl. ii, fig. 18).—N. Ingram Hendey, “The Plankton
Diatoms of the Southern Seas’ in Discovery Reports, vol. XVI, 1937, p. 332; Sch. At.,
t. 299, figs. 9-14. Synonyms: Fragilaria antarctica Castr., Denticula tenuis K. var.
antarctica Fritsch. (Hendey.)

Distribution: Very common antarctic oceanic diatom. Also Cape Adare, Bouvet
Island and Heard Island.

FrRustutia Agardh.

LE USHICLUCCM IAITILE sabe (ele ti tes) 9) Preity, lmnite. ps 924. Synonym: Synedra
frustulum K. (Pritchard.)

Only outline and raphe visible. Identification doubtful.

Frustulia rhomboides (Eh.) DeToni. (Text-fig. 12)—Hust. in Die Stissw. Mitt.,
Ht. 10, p. 220, fig. 324, (1930). Synonym: Vanheurckia rhomboides Breb. (DeToni.)

Distribution: World-wide.

GOMPHONEMA Agardh.

Gomphonema angustatum (K.) Rab. var. producta Grun. (Text-fig. 22).—Hust. in
Die Stissw. Mitt., Ht. 10, p. 373, fig. 693, (1930).

The figure shown in Sch. At., t. 234, fig. 26 does not agree with that of Hustedt.
Schmidt, in the same table, presents a series of figures of G. angustatum K. and it is
clear that there is wide and continuous variation exhibited by the species so that the
naming is probably somewhat artificial.

Gomphonema intricatum K. (Pl. ii, fig. 17) Sch. At., t. 235, fig. 15. Synonyms:
G. dichotomum K., G. vibrio Eh., G. cygnus Eh. (DeToni.)

Distribution: World-wide.

In this case the figure shown by Hustedt does not agree with Schmidt’s Atlas. Again,
the most appropriate authority has been chosen.

Gomphonema parvulum (K.) Grun. (PI. ii., fig. 16)—Hust. in Die Siissw. Mitt.,
Ht. 10, p. 372, fig. 7138a, (1930); Sch. At., t. 234, fig. 2. Synonyms: Sphenella parvula K..,
Gomphonema minutissimum Breb., Gomphonella parvula Rab., Sphenella rostellata K.,
Gomphonema rostellatum Rab., Gomphonema Lagenula (K.) Rab., Gomphonema
rostratum W. Sm., Gomphonema tenellum W. Sm. (DeToni.)

Distribution: World-wide.

HANTZSCHIA Grunow.

Hantzschia anphiozys (Eh.) Grun. (Pl. ii, fig. 28).—Hust. in Die Siissw. Mitt.,
Ht. 10, p. 394, fig. 747, (1930). Synonyms: Hunotia amphioxys Eh., Navicula amphioxys
Westend, Nitzschia amphioxys W. Sm. (DeToni.)

Distribution: World-wide.

Specimens typical of Hustedt’s fig. 747 are of infrequent occurrence. Most of the
material has closer affinities with the variety maior Grun. (Hustedt’s fig. 749) although
the size range is well below the range quoted for this variety. For this reason, varietal
subdivisions have been avoided.

42 TERRESTRIAL DIATOMS OF MACQUARIE ISLAND,

Menosrra Agardh.

Melosira granulata (Eh.) Ralfs. (Pl. ii, figs. 20a and b).—Hust in Die Stissw. Mitt.,
Ht. 10, p. 87, fig. 44, (1930). Synonyms: Gallionella granulata Eh., Gallionella marchica
Eh., Gallionella procera Eh., Gallionella tenerima Eh., Melosira ordinata K., Orthosira
punctata W. Sm. (DeToni.)

Distribution: World-wide.

? Melosira spp.—Small dises, usually with undefinable structure, have been placed
for convenience with this genus. The possibility that they may represent one or more of
the following genera (Thalassiosira Cleve, Cyclotella K., Stephanodiscus Eh.) should not
be overlooked.

NaviIcuLa Bory.

Navicula mutica K. (vars ?) (Text-figs. 2, 4, 15).—Hust. in Die Siissw. Mitt., Ht. 10,
p. 274, fig. 453a-e, (1930). Synonyms: Stauroneis Dumontii Breb., Stawroneis polymorpha
Lagerst., Navicula Semen Eh., Stauroneis Semen Eh., Stauwroneis Cohnii Rab. (DeToni.)

Distribution: World-wide.

With the variation in form of this species shown by Hustedt it is difficult to under-
stand how there can be any justification or necessity for erecting the separate species
Navicula muticopsis V.H. (W. and G. S. West, Fresh Water Algae, Brit. Ant. Exp.,
1907-9, vol. 1, pt. 7, p. 283, Pl. XXVI, figs. 121-124; Peragallo, Diatomées d’Eau Douce,
Expedition Antaretique Francais, 1908-10, p. 17, Pl. 1, fig. 40), Navicula muticopsiforme
W. and G. S. West (these authors in Brit. Ant. Exp. as above, p. 284, Pl. XXVI, fig. 131),
Navicula dicephala Eh. (F. E. Fritsch, Fresh Water Algae, Brit. Ant. Exp., 1910, Bot.,
pt. 1, p. 14, Pl. 1, fig. 32) and Navicula globiceps Greg. (W. and G. S. West in Brit. Ant.
Exp. as above, p. 284, Pl. X XVI, fig. 135; F. E. Fritsch, Fresh Water Algae, Nat. Ant.
Exp., 1901-4, vol. VI, p. 52, Pl. III, figs. 154, 155).

Navicula dicephala Bh., as described by Fritsch, has punctate striae and so does not
agree with the Navicula dicephala Eh. described by Hustedt (Die Stissw. Mitt., Ht. 10,
p. 302, fig. 526) in which the striae are unbroken. Fritsch, in fact, expresses his doubts
regarding the naming, and states that the diatom named by him Navicula muticopsis
VES Gea binna Soc Bot Xe: p. 336, Text-fig. B) is identical with Navicula dicephala
Eh. There is some disagreement between Navicula globiceps Greg. (Fritsch, Nat. Ant.
Exp., 1901-4, vol. VI, p. 52) with punctate striae and Navicula globiceps Greg. (W. and
G. S. West, Brit. Ant. Exp. as above) with broken striae. Fritsch’s reference is Gregory’s
original description so that the name by W. and G. S. West may be misapplied.

The following list of probable affinities should be noted:

Navicula mutica var. ventricosa (K.) Cleve very similar to Navicula globiceps
Greg.

var. Cohnii (Hilse) Grun. very similar to N. muticopsiforme
W. & G.S.W.

var. truncata Per. identical with N. muticopsis V.H.
var. binodis Hust. symmetric form of WN. mutica var.
cymbelloides Per.

Navicula mutica, var. truncata Per., var. Cohnii (Hilse) Grun, and var. nivalis (Eh.)
Hust. have been observed in the samples from Macquarie Island.

Navicula (rostellata ? K.) (Pl. ii, fig. 21).—Sch. At., t. 47, figs. 28, 29. Navicula
rostellata K., as listed by Hustedt (Die Stissw. Mitt., Ht. 10, p. 297, fig. 502) has unbroken
striae. Synonym: Navicula rhyncocephala K. var. rostellata Grun. (Cleve and Grunow;
DeToni.) The species is queried because of the disagreement between Schmidt and
Hustedt.

Navicula seminulum Grun. (Text-fig. 5).—Hust. in Die Stissw. Mitt., Ht. 10, p. 272,
fig. 443. .

Distribution: Cape Adare. (Only other antarctic record.)

BY J. S. BUNT. 43

NeIpium Pfitzer.
Neidium affine (Eh.) Cleve. (Text-fig. 17).—Hust. in Die Siissw. Mitt., Ht. 10,
p. 242, fig. 376, (1930); W. Sm., Synops. Brit. Diat., I, p. 50, Pl. XVI, fig. 143 (Navicula
affinis Eh.); Sch. At., t. 49, figs. 20-23. Synonyms: Navicula iridis var. affinis Eh.,
Navicula affinis Eh., Navicula ampliata Eh. (DeToni.)
Distribution: World-wide.
This species was identified directly from a slide mount prepared by West.

OPEPHORA Petit.
Opephora Martyi Heribaud. (PI. ii, fig. 22).—Hust. in Die Stissw. Mitt., Ht. 10, p. 132,
fig. 120, (1930).

PINNULARIA Ehrenberg.

Pinnularia appendiculata (Ag.) Cleve. (PI. ii, fig. 23).—Hust. in Die Stssw. Mitt.,
Ht. 10, p. 317, fig. 570a, (1930). Synonyms: Frustulia appendiculata Ag., Cymbella
appendiculata Ag., Navicula obtusa Eh.

Distribution: Graham Land, Falkland Island, Cape Horn. (Antarctic records.)

The similarity between this species and Pinnularia molaris Grun. should be noted.

Pinnularia Atwoodii Per. (Text-fig. 16).—Peragallo, Diatomées d’Eau Douce,
Expedition Antarctique Francais, (Bot.), p. 20, Pl. I, fig. 17 (1908-10).

Distribution: Quinipiac River, U.S.A. (Only other record.)

Peragallo points out affinities between this species and Pinnularia nivorum Per.
(Exp. Ant. Franc., as above, Pl. I, fig. 18).

Pinnularia borealis Eh. (Text-fig. 20).—Hust. in Die Siissw. Mitt., Ht. 10, p. 326,
fig. 597, (1930); Sch. At., t. 45, figs. 15-21. Synonyms: Pinnularia latestriata Greg.,
Pinnularia chilensis Bleisch, Pinnularia hebridensis Greg., Navicula borealis Eh.
(DeToni.)

Distribution: Graham Land, South Orkneys, Cockburn Island, Cape Adare, Falkland
Island, Cape Horn, Magellans, Kerguelen Island. (Antarctic records.) World-wide.

Pinnularia brevicostata Cleve. (Pl. ii, fig. 29).—Hust. in Die Stissw. Mitt., Ht. 10,
p. 329, fig. 609, (1930); Sch. At., t. 48, figs. 26, 27.

The species is somewhat variable since fig. 609 (Hust.) does not agree very well with
those in Schmidt’s Atlas, which correspond better with the Macquarie Island type. The
presence of a central gap in the costae and the straightness of the sides of the valves
are the sources of the variation. This species should be compared with Pinnularia
hemiptera K. (Sch. At., t. 43, fig. 28) and also with Pinnularia montana Hust. (Sch.
At., t. 389, fig. 6).

Pinnularia cardinalis (Eh.) W. Sm. (PI. ii, fig. 30).—Hust. in Die Stissw. Mitt.,
Ht. 10, p. 337, fig. 621, (1980); Sch. At., t. 44, figs. 1-2; W. Sm., Synops. Brit. Diat.,
I, p. 55, Pl. XIX, fig. 166. Synonyms: Stauroneis cardinalis Eh., Stauroptera cardinalis
Eh. (DeToni.)

Distribution: World-wide.

The specimens examined were 75u long and 124 wide compared with 150-320u long
and 35-454 wide. (Hustedt.)

Pinnularia divergens W. Sm. (PI. iii, fig. 1) —Hust. in Die Stissw. Mitt., Ht. 10, p.
323, fig. 589 (1930); W. Sm., Synops. Brit. Diat., I. p. 57, Pl. XVIII, fig. 177. Synonym:
Stauroptera divergens Kirch. (DeToni.)

Distribution: Cape Horn. (Only other antarctic record.)

Pinnularia divergentissima (Grun.) Cleve. (Pl. ii, fig. 26).—Hust. in Die Siissw.
Mitt., Ht. 10, p. 320, fig. 581, (1930); Sch. At., t. 388, figs. 23, 24.

The material examined was not constricted subapically to the extent shown by
Hustedt.

Pinnularia fasciata Lagerst. (Text-fig. 11).—Hust. in Die Siissw. Mitt., Ht. 10, p. 316,
fig. 569, (1930).

Pinnularia interrupta ? W. Sm. (Text-fig. 9)—Hust. in Die Siissw. Mitt., Ht. 10,
p. 317, fig. 573a, (1930); Sch. At., t. 45, figs. 67, 69, 70-72, 75, 76, 79; W. Sm., Synops.

44 TERRESTRIAL DIATOMS OF MACQUARIE ISLAND,

Brith DLA ley Ds DO lee eri comemlicseh Synonyms: Navicula mesolepta Eh. var.
stauroneiformis Grun., Pinnularia mesolepta var. interrupta W. Sm. (DeToni.)

There is considerable variety within this species principally in the disposition of
the striae, Hustedt’s figure having the closest affinity with the material examined in
which the striae bend like ribs to the edge of the valve. In this respect the Macquarie
Island specimens are quite distinct, but the variation within the species makes the
erection of a new species or variety of doubtful value, especially as there appears to be
no distinct boundary between this species and Pinnularia mesolepta (Kh.) W. Sm. (Sch.
At., t. 45, fig. 62) and Pinnularia Braunii (Grun.) Cleve (Sch. At., t. 45, figs. 77, 78).

Pinnularia lata (Breb.) W. Sm. (PI. iii, fig. 2).—Hust. in Die Stssw. Mitt., Ht. 10,
Dp: 324, fie. 595) (1930) We om. Synops) Brit) Dia wl spea 555 elk exeVvsllin shoo Glin SCs
At., t. 45, figs. 5-8. Synonyms: Frustulia lata Breb., Navicula pachyptera K., Pinnularia
pachyptera Eh., Pinnularia megaloptera Eh., Pinnularia suecica Eh., Pinnularia’
pleurophora Eh. (DeToni.)

Distribution: Graham Land, Magellans. (Antarctic records.) World-wide.

Hustedt and W. Smith show slight bulge in valve walls opposite central area. The
specimens examined were identified with a slide mount prepared by West.

Pinnularia microstauron Eh. (Pl. ii, fig. 27; Pl. iii, fig. 4)—Hust. in Die Stissw.
Mitt., Ht. 10, p. 320, fig. 582, (1930); Sch. At., t. 44, figs. 14, 16, 34, 35 and t. 45, figs. 31-34.
Synonyms: Stauroneis microstauron Eh., Stauroptera microstauron Rab. (DeToni.)

Distribution: Falkland Island, Magellans, Kerguelen Island. (Antarctic records.)
World-wide.

This species is highly variable. The Macquarie Island specimens were identified
from a slide mount prepared by West, which contained a wide range of forms.

Pinnularia microstauron (BHh.) Cleve var. Brebissonii (K.) Hust. (Text-fig. 6).—
Hust. in Die Stssw. Mitt., Ht. 10, p. 321, fig. 584, (1930). Synonyms: Navicula Brebissonii
K., Frustulia bipunctata Breb., Navicula bipunctata Bory., Pinnularia stauroneiformis
W. Sm., Pinnularia Brebissonii Rab., Stauroptera Brebissonii Kirch. (DeToni.)

Distribution: Europe, Norway.

Pinnularia molaris Grun. (PI. ii, figs. 24, 25).—Sch. At., t. 44, fig. 54; Hust. in Die
Stssw. Mitt, Ht. 10, p. 316, fig. 568, (1930). Synonym: Pinnularia macra Grun.
(Schmidt. )

Distribution: Graham Land. (Only other antarctic record.)

Should be compared with Pinnularia divergens W. Sm. var. sublinearis Cleve (Sch.
At., t. 44, fig. 54) and Pinnularia leptosoma Grun. (Sch. At., t. 388, figs. 13-15).

Pinnularia nivorum Per. var.? (Text-fig. 1)—The closest related type that could be
found for these specimens was Pinnularia nivorum Per. (Peragallo, Diatomées d’Kau
Douce, Exp. Ant. Franc., 1908-10, Bot., p. 20, Pl. I, fig. 18.) The material has therefore
been placed as a possible variety of this species.

RHOPALODIA O. Miiller.

Rhopalodia gibberula (Eh.) O.M. var. baltica O.M. (Text-fig. 18).—Sch. At., t. 2538,
1, BB

STAURONEIS Hhrenberg.

Stauroneis acuta.W. Sm. (PI. iii, figs. 3a, b, ¢)—Hust. in Die Siissw. Mitt., Ht. 10,
Dp: 259, fig. 415, (1930); Sch. At., t. 241, fic. 4; W. Sm., Synops. Brit. Diat., 1, p: 59)
Pl. XIX, fig. 187; Prit. Inf., p. 914, t. vii, fig. 76. Synonym: Plewrostauron acutum
(W. Sm.) Rab. (DeToni.)

Distribution: World-wide.

Stauroneis anceps Eh. var. hyalina Brun and Per. (PI. iii, fig. 8).—Sch. At., t. 242,
i, lil.

In Hustedt (Die Siissw. Mitt., p. 256, fig. 408) no punctation is shown on the
striae and the apices are more acute than in Schmidt’s Atlas and the Macquarie Island
specimens. Stauroneis anceps Eh. has been recorded in the antarctic from Cape Adare,
Ross Island, South Victoria Land, Cape Horn and Kerguelen Island.

BY Je Se BUND 45

Stauroneis parvula (Grun.) Jan.—Hust. in Die Stissw. Mitt., Ht. 10, p. 260, fig. 417a,
(1930). Synonym: Pleurostauron parvulum (Jan.) Grun.

Stauroneis pygmaea Kreiger. (Text-fig. 3).—Hust. in Die Stissw. Mitt., Ht. 10, p. 257,
fig. 409, (1930).

SURIRELLA Turpin.

Surirella angustata K. (Pl. iii, fig. 5).—Hust. in Die Stissw. Mitt., Ht. 10, p. 435,
figs. 844, 845, (1930); Sch. At., t. 23, figs. 39-41 (Surirella angusta K.). Synonyms:
Suriraya ovalis Breb., var. angusta K., Surirella apiculata W. Sm. ;

Distribution: Cape Adare, Ross Island, South Victoria Land, Cape Horn. (Antarctic
records. )

Surirella bifrons K. (Pl. iii, fig. 7).—Sch. At., t. 22, figs. 5, 8, 10, 11, 12 and t. 23,
ines, Il, Be

The type from Macquarie Island fits figure 12, t. 22 best. There is some variation
in form with this species and it should be compared with Surirella linearis W. Sm.
(Sch. At., t. 23, fig. 27), Surirella saxonica Auersw. (Sch. At., t. 22, fig. 1) and Surirella
tenera Greg. (Sch. At., t. 23, figs. 7-9).

? Surirella Engleri O.M. forma angustior O.M. (Text-fig. 13).—Sch. At., t. 245, fig. 14.

This is a large planktonic species more than 200u long. Its affinities with Surirella
linearis W. Sm. are indicated in Schmidt’s Atlas. Although the specimens examined are
only 75-80u long, they fit the above species very well.

SyYNEDRA Ehrenberg.
? Synedra vaucheriae K. (Pl. iii, Fig. 6).—Hust. in Die Stissw. Mitt., Ht. 10, p. 161,
figs. 192a-c, (1930); Sch. At., t. 305, figs. 18-31.
Distribution: Kerguelen Island. (Only other antarctic record.)
The determination is doubtful as only isolated frustules were found. Typical star-
like aggregates were, however, observed in fresh samples. Comparison should be made
with Fragilaria intermedia Grun.

ENVIRONMENT AND ITS EFFECT ON THE PAST AND PRESENT DIATOM FLORA OF
MACQUARIE ISLAND.

The diatoms of Macquarie Is. may be considered as occurring in three major
ecological environments; the soil, the water of the ponds, lakes and watercourses, and a
habitat now non-existent but, as will be shown, of considerable interest and represented
by the organic and diatomaceous strata of the fluvio-glacial sediments of the closing
ice age.

In table I have been listed the species of diatoms and their estimated frequencies
of abundance in the 10 soils already described. That there are differences between soils
and varying frequencies of any one species from soil to soil is immediately clear. A
closer examination reveals that five of the soils have the same dominant or co-dominant
species (Pinnularia molaris Grun.) but that no such comparison may be applied to the
remaining five. This information is presented in table V and further emphasized by
table VI showing the number of species common to 1, 2, 3, ...410 soil habitats. Only
4 types are common to five or more soils, and of these, 3 are dominants, viz. Pinnularia
molaris Grun., Pinnularia Atwoodii Per. and Hanteschia amphioxys Grun. Of the
remaining dominant species, 4 occur in two soils, 4 occur in three soils and 3 occur in
four soils, dominance in each case, however, being restricted to one habitat. The forms
grouped arbitrarily as Melosira spp. were found only occasionally in soil no. 8 but have
been used to characterize this situation. 43% of the species were found as minor
constituents in single habitats.

From these observations, it is apparent that the soil, as an ecological environment,
has marked effects on the size and complexity of the diatom population. In certain
soils (e.g. no. 3) they form an important part of the microflora because of their photo-
synthetic activities and the effect of these activities on other groups of microorganisms,
whilst in other soils, notably no. 6, they are practically non-existent. A full assessment

46

Achnanthes Biasolettiana Grun.
—— brevipes Ag. var. inter-
media K.
— lanceolata Breb.
subsalsa Peterson
Coscinodiscus sp.
“Cyumbella pusilla ? Grun.
Diatomella Balfouriana Grev.
Diploneis Smithii (Breb.)
Cleve j fe
Eunotia lunaris Grun.
trinacria Krasske
Fragilaria bicapitata A. Mayer
pinnata Eh. ne oe
Fragilariopsis antarctica Castr.

2 Frustulia minuta Rab.
rhomboides (Eh.) De Toni
Gomphonema angustata (K.)
Rab. var. producta Grun.
Hantzschia amphioxys (Eh.)
Grun.

Melosira granulata (Eh.) Ralfs
spp.

Navicula mutica K.

(rostellata ? K.)

seminulum Grun. :

Neidium affine (Eh.) Cleve ..

Pinnularia appendiculata (Ag.)
Cleve.

TERRESTRIAL DIATOMS OF MACQUARIB ISLAND,

|

TABLE J.
The Diatom Flora of Macquarie Island Soils.

3 (0).
4 (vr).

2 (vr), 3 (0), 4 (vr).

4 (0), 5 (vr).

4 (vr), 7 (vr).

3 (rt).

1 (f), 2 (0), 3 (vr).

1 (vr), 2 (vr), 3 (a),
5 (vr).

3 (r), 10 (a).

3 (7), 9 (vr), 10 (a).

7 (c).

1 (vr), 2 (vr).

2 (vr), 4 (vr), 5 (vr),
7 (vr), 9 (0), 10 (vr).

3 (0).

1 (vr), 2 (vr), 3 (f).

3 (vr).

1 (0), 2 (r), 3 (0),
4 (vr), 5 (0), 7 (a),
10 (f).

5 (r).

3 (r), 8 (0),
10 (c).

1 (x), 4 (vr), 5 (vr).

3 (vr).

5 (vr).

1 (vr), 5 (vr).

83 (69);

9 (f),

Pinnularia Atwoodii Per.

borealis Eh.
brevicostata Cleve pa
cardinalis (Eh.) W. Sm.

divergens W. Sm.
divergentissima
Cleve me
fasciata Lagerst.
interrupta ? W.Sm.
—— lata (Breb.) W. Sm.

(Grun.)

—— microstauron Eh.

Hust.
molaris Grun.

var. Brebissonii (K.)

nivorum ? Per. var.? ..

Rhopalodia gibberula OM.
var. baltica O.M.

Stauroneis acuta W. Sm.

anceps. var. hyalina Brun.

and Per.

parvula ? (Grun.) Jan...

pygmaea Kreiger

Surirella angustata K.

bifrons K.

Englert
-angustior O.M.

2 Synedra vaucheriae K.

2 (O.M.) B

2 (zr), 3 (0), 4 (vr),
5 (0), 7 (£).

1 (vr), 2 (2), 3 (c).

5 (vr).

1 (@), 3 (vr), 5 (x),
9 (vr).

1 (vr), 3 (0).

3 (c), 5 (vr), 9 (vr),

2 10),

1 (f), 3 (vr), 7 (c).

| 9 (f), 10 (0).

1 (vr), 2 (0), 3 (ce),
5 (¢).

IE X(0)5 210); S(O) >
4 (vr).

9 (zr).

lL @s 4 @, 8 @)-
4 (vr), 5 (f), 8 (vr),

| 9 (a), 10 (a).

i 1 (f), 2 (x).

3(vr).

| a (vr).

3 (vr).

1 (vr).

9 (zr).

1 (ec).

2 (vr), 3 (r).
3 (r).

| 4 (0).

Key to symbols: a, abundant; f, frequent ; c, common; 0, occasional; r, rare ; vr, very rare.
The numbers 1-10 refer to the soil samples (described above, p. 35) in which the species were found.

TABLE II.
The Diatom Flora of Macquarie Island Lakes and Ponds.

Achnanthes Biasolettiana Grun.
brevipes Ag. var. inter-
media K. oes
subsalsa Peterson
Amphora delicatissima Krasske
Coscinodiscus sp. a ©:
Cymatopleura solea Breb. var.
regula (Kh.) Grun.
Cymbella pusilla ? Grun.
Diatomella Balfouriana Grev.
Diploneis Smithii (Breb.) Cleve
Eunotia alpina K.
lunaris Grun.
trinacria Krasske
Fragilaria bicapitata A. Mayer
capucina Desm.
pinnata Eh.
virescens ? Ralfs
Fragilariopsis antarctica Castr.
2 Frustulia minuta Rab.
Frustulia rhomboides (Eh.)
De Toni ay te
Gomphonema intricatum K.
parvulum (K.) Grun.
Hantzschia amphioxys (Eh.)
Grun. a

OT ee Se SO

1,5, 11.

6.

2,5, 10, 11.
2, 10, 11.
2,8, 10.
10, 11.

5, 10, 11.
Ty Wil.

gS
EP
=

1, 2, 4, 5, 10.

Melosira spp. ..

Navicula (rostellata ? K.) A

Neidium affine (Eh.) Cleve ..

Pinnularia appendiculata (Ag.)
Cleve

—— Atwoodii Per.

borealis Eh.

brevicostata Cleve

—— cardinalis (Eh.) W. Sm.

divergens W. Sm.

—— divergentissima
Cleve Soe

lata (Breb.) W. Sm.

fasciata Lagerst.

microstauron Bh.

molaris Grun.

nivorum ? Per. var. ?

Stauroneis anceps var. hyalina
Brun. and Per.

2 Surirella Engleri f. angustior
O.M.

bifrons K. 5

2 Synedra vaucheriae K.

»

(Grun.)

ye) 2, By, 7 Sy
itil,

5, 10, 11.

5, 10, 11.

10.

3, 6, 8, 9, 10, 11.
10, 11.

1.

The numbers 1-10 refer to the lakes and ponds (listed on p. 36) in which the species occur.

BY J. S. BUNT. 47

of the implications of these findings cannot at present be made. Our knowledge of the
interbiotic relations of the various groups of soil-inhabiting organisms is most inadequate
and is frequently restricted to isolated observations. Rotifers, for example, have been
observed by the author apparently subsisting on soil algae growing on an artificial agar
medium. This illustrates a small link in a complex food-chain which has been discussed
in a most constructive manner by Birch and Clark (1953) using forest soils for their
particular studies. Certain free-living nematodes are also known to be algal feeders
and the author has found (unpub. results) that this group frequently occurs in large

TABLE III.
The Fossil Diatoms of the Deep Peats and Fluvio-glacial Sediments of Macquarie Island.

|

Species. Peat Sample Number.
Achnanthes Biasolettiana Grun. = Ae so. || 2th Os Gs
brevipes Ag. var. intermedia K. ms .. | 99, km, n, 12B3, 12B4, 12D) 187, 26k, 27c, F1, F9, F10-
lanceolata Breb. .. Se. ah Bo 2 eo 2 Ba. 2D 187. 27s .¢e) Hy ho:
Coscinodiscus sp. a as ae sa |) IDB
Cocconeis placentula Eh. bn so We 5 Ty Lbs Wop
Cymatopleura solea Breb. var. regula (Eh. ) Ganuni. .. | 12B4,
Cymbella pusilla ? Grun. Be ae an oa CYR, UID, YAtldy G5 dal,
Diatomella Balfouriana Grev. ts ae .. | 91,9, k, n, 12B3, 12B4, 12D, 18), 267, k, 27d.
Diploneis Smithii (Breb.) Cleve m3 es so | thd P05 1) Ub PWD; @ GE 1,
Eunotia trinacria Krasske Ke ae be so |) JAD
Fragilaria capucina Desm. ane oes Sas .. | 9m, 12D.
Harrisonti (W. Sm.) Grun. Ba at .. | 99,9, k, m,n, 12B3, 12B4, 12D, 18), 261, 270, c.
—— pinnata Eh. oes aie ie ei .. | 99, 7,9, k, m,n, 12B4, 12D, 18), 261, 27c.
—— virescens Ralfs a: oe ee ais ao || OY Gp dy Hy Gly Wy ASB
virescens ? Ralfs .. = oe a .. | 9m, 12B4, 12D, 18), 26k, 1, 27a, b,c, d, F1, F2, F3.
Fragilariopsis antarctica Castr. a: 3h .. | 91,9, k,n, 18D, 263, k, 27a, b, F3.
Frustulia rhomboides (Eh.) De Toni .. . | 26k, 27c.
Gomphonema angustatum (K.) Rab. var. PrOnea Gannta 99,9, Kk, 12B4, 12D. ~
—— intricatum K. oe is as Se .. | 99, k, m, n, 12B4, 12D, 187, 27b, ec, d, F1, F3, F10.
parvulum (K.) Grun. is et aa .. | 99-n, 12B3, 12B4, 12D, 27¢c, F1, F3, F10.
Hantzschia amphioxys (Eh.) Grun. .. 35 ~-. | 12B4, 12D, 261, 27b, c, F1.
Melosira spp. a ass ye ae a .. | 9i-n, 12B3, 12B4, 12D, 189, 269, k, 27a, c, F1—9.
Navicula mutica K. a ee a .. | 99g, 12B4.
Neidium affine (Eh.) Gitar = 3% ae .. | 12B4, 12D.
Opephora Martyi Heribaud ai a oY so |) Che Ui Wy py UAIDE Arar
Pinnularia brevicostata Cleve .. a an .. | 12B4, 12D, 27a.
—— cardinalis (Eh.) W.Sm. .. se Ss .. | 12B4, 12D, 18), 26k, 1, 275, c.
—— divergens W.Sm. .. ar us ae .. | 12B4, 12D, 261, F1.
—— lata (Breb.) W. Sm. Bee ue re .. | 94,9, n, 12B4, 12D, 18d, 26k, 27a.
—— microstauron Eh. an oe ee, sa || Gy, IPB WARY IPM), wes, Ga, PA, Bld, @; Gi Ieil, IS}, A)
molaris Grun. a ae co || JBI, 1NSy, IDO),
Rhopalodia gibberula var. otiiien O. Ny pe 55 |) PAVE Uh
Stauroneis acuta W.Sm. bis ee .. | 9%, k, 12B4, 26k, l, 270, c, d.
anceps var. hyalina Brun. and Re. 33 Pe 2 Bas TADS 845 26 2a0sC
Sune bifrons K. oe es oo || PAG Rs
Engleri (O.M.) f. minignctiien O. M. a ae | 26K:
2 Synedra vaucheriae K. ae ah sh oo || W2ID,

The numbers refer to the samples from the fluvio-glacial deposits listed on p. 36.

numbers in the soils of Macquarie Is. The ultimate effect of such relationships on the
soil and thence on the growth of higher plants rests with bacterial decomposition. Should
a particular group, such as the diatoms, be prominent, this provides, in itself, a ready
source of energy for bacteria but this food source may not undergo direct decomposition,
being diverted to other channels such as nematodes so that the eventual composition of
the soil organic matter is affected, depending on the nature and complexity of the food
chain or path of breakdown.

The influence of soil conditions is complex and there is sufficient variation in the
soils examined, both physically and chemically, to explain in some part, the differences
in the diatom floras. Variations in local climate are also important partly from the

48 TERRESTRIAL DIATOMS OF MACQUARIE ISLAND,

TABLE IV.
List of Antarctic Diatoms.

Achnanthepyla Bongraini Per. il; Cymbella antarctica Cleve LOE
—— Gainii Per. as Be | ile aspera Bh. .. we 56 | IO)
Achnanthes antarctica Per. | ile cistula Kirch. oe be ule
—— affinis Grun. ve 45 4) BE cuspidata Kk. aes Yd 9.
australis Eh. a seal willis cymbiformis Ag... S53 ils
—— Biasolettiana Grun, ; 7, 8, 10, 18. —— Khrenbergii K. i Ses 8, 9.
—— brevipes Ag. var. interme Hi gastroides Eh. is sg |) 1,

K. a a ae 55 4, 6, 18. lanceolata Kirch. .. ae 1
—— Charcotii Per. ie te ily —— microcephala Grun. aie \\ 110);
—— coarctata Grun. var. eli- naviculaeformis Auers. .. | 10.

minata Lag. f. antarctica Per. | 1. pusilla Grun. oe Phe 6, 13 (2).
—— (coarctata var.?) distorta spuria Cleve ae no || JO)

Per. Ei : ee ae ile Diatoma anceps Kirch. As il,

exigua Guin she 56 |) aos asi —— Ehrenbergii K. var. grande
CHS Keer ate soy) des Tey Vo 1Bies : ae tis Oo ty lil,
hungarica Grun. .. ae 9. elongatum Ave, Se ae 4,
—— inflata Grun. af Ze By Co grande W. Sm. re vo |) Uae
—— lanceolata Breb. .. 54 ale} rhombicum OPM... edd | ele
linearis Grun. les 10, 13. Diatomella Balfouriana Grev. .. | 138.
minutissima K. var. eniitie- Diploneis elliptica Cleve Mee 9, 10.
cephala Grun. .. ws sa |). bBo —— ovalis var. oblongella Cleve ils
muscorum Per. Ih, Smithii (Breb.) Cleve bo.) IS}
== var. minor Per. ile Denticula tenwis K. var. antarctica
—— Semen Eh. .. Us Fritsch 4,
subsalsa Peterson 13. thermalis K. var. minor 2;
Amphora gracilis Eh. 12. Encyenema gracilis Bab. 10.
hyalina Eh. Lo —-—- lunula Grun. 9.
libyca Eh. Ue, So Epithemia sorex Kk. 1.
—— ovalis K. ae 2. —— turgida K. ae ile
— var. gracilis V.H. Gi, all, —— var. zebrina Rab. Ife
a pediculus V.H. 10. —— zebra K. bee iby Go
delicatissima ? Krasske 113° —— —— f. minor V.H. il,
Amphiprora minor Greg. 2" Eunotia alpina kK. 10, 13.
parva Reinsch 15 —— arcus Eh. ne 8, 10, 11
Amphiprora Pockornyana Genin, 12: —— var. hybrida Grun. .. 9.
Biddulphia arcticum Brightw. 6. attenuata Cleve 10.
sp. : 2° Camelus Eh. 8.
Campylodiscus magnus Remecn 12. —— Cygnus Eh. iil,
Cocconeis borealis Eh. was exigua Rab. 10.
costata Greg. PA, —— Falklandiae Eh. To
——— var. imperatria (Eh. 2) Bs gracilis W. Sm. WA. tS}5
—— var. pacifica Eh. 3: impressa Eh. ne Shy sled
—— lhtigiosa V.H. 6. incisa Greg. pee so It JO),
—— placentula Eh. Oey 13 lunaris Grun. Ba as; i, ile,
== var. lineata V.H. 8. —— major Rab. ae xo {| al),
striata Eh. (Diploneis —— monodon Eh. “A promi azlale

elliptica ?) Bats Sib He teky II obesa Cleve “eS a 9.

Coscinodiscus decipiens Grun. . . 4, obtusiuscula Cleve Be 9.
griseus Grey. 4, —— parallela Eh. es 8, 9.
lacustris Grun. 4, pectinalis Dilwyn 12.
lentiginosus Jan. 6. —— Rab. var. stricta Rab. 9
radiatus Bh. A 2 quaternaria Eh. 8
2 Coscinodiscus sp. at aa. te det robusta Ralfs od 9.
Cyclotella comta K. ae le — var. papilio Rab. .. 9.
Meneghiniana K. var. ternaria Cleve Se oo |] iL
plana Fritsch hye se al, tridentata Eh. as so || IOs
operculata K. é ee 4. —— tridentula Eh. ae =e 1, 8, 10.
Cymatopleura solea W. ont a il, var. ventricosa Per. .. 1.
Breb. var. regula trinacria Krasske . . Ave ila

(Eh.) Grun. .. xo |) 18% triodon Eh. te 3 il.
Cymbella americana A. Sch. aa 9. Fragilaria bicapitata A. Newer 13}.

—— capucina Desm. .. als 7,13, 14

The numbers refer to the following records: 1. Peragallo, 1908, Graham Land; 2. Fritsch, South Orkneys ;
3. Hooker, Ehrenberg, Cockburn Island; 4. Fritsch, 1901-4, Cape Adare; 5. Fritsch, 1910, Cape Adare;
6. W. and G. S. West, 1907-9, Ross Island, South Victoria Land; 7. Ehrenberg, Falkland Island; 8. Ehrenberg,
Cape Horn; 9. Petit, Cape Horn; 10. Cleve, Magellans; 11. Ehrenberg, Kerguelen; 12. Reinsch, 1874-9, Ker-
guelen ; 13. Bunt, Macquarie Island; 14. O’Meara, Marion Island.

B Yaa Je See WINEDS 49

TABLE LV.—Continued.
List of Antarctic Diatoms.—Coutinued.

Fragilaria construens Gru. 8 Navicula antarctica (Kh.) De Toni 11.
curta V.H. ee i lean. —— Atomus Grun. ve va ives
Eunotia Eh. ? ne alas australis Eh. ae Be 8.

— Harrisonii (W. Sm.) Guin Heelies —— bacillum Eh. $45 Se Bes 10),

2 linearis Castr. .. oval ages —— hisulcata Lag. Ft PAN Pe ic

—— mutabilis Grun. .. fae, —— Boudeti Per. he Bent eats

— obliquecostata V.H. sa lay @: brachysira Breb. .. Bee ell Os
pinnata Eh. oe pr Cciey, carassius Eh. es ak | 14.
tamuicolisstleibs iy.) 1) yeh a4: > Clemens es ee oe ae

— var. antarctica W. & —— Charlatii Per. 55 So tl) te

Gao West 6 as alee? —— chiliensis K. ae Buea asl
virescens Ralfs. es ae 6, 13 —— completa Cleve 5h | 10.
virescens ? Ralfs .. oo || 1a —— crassinervia Breb. co ||

Fragilariopsis antarctica Castr. 13. —— curta Per. : Ae beny ag

2 Frustulia minuta Rab. Bye aloe —— curta Per. var. elon, ain Bore lWeeail

Frustulia rhomboides (Eh.) —— cuspidata Kk. ans erally rch

De Toni ae. aro de abet —— cymatopleura W. & G. 3.

Gomphonema CERI RTERIED (K.) West .. Be ae ite 4, 6.

Rab. var. producta Grun. .. | 13. —— Dactylus K. os ae 9.
bicapitatum OM. .. ao) see —— var. ? bt AE 9,
augur Eh. .. we Sat) dal, decurrens Kh. Ups
auritum Braun... 53 |} Os —— dicephala En. irs So) By 05 il

— Brebissonii K. Si poll dep —— elliptica K. var. Portonoiiiee
catena Eh. a ars Hae IPOS ie iG A
constrictum Eh. es il, ergadensis Ralfs st Me 9.
dichotomum K. ie, ee tlie Ga — Falklandiae Eh. .. Be Uo
gracile Eh. .. . Up iu, Gastrum Bh. Ay pial
intricatum K. ie sa |} 18% glaberrima W. & G.S. West | 6.
Kamtschaticum Grun. var. —— Godfroyi Per. poo Weal bs

antarctica Per... : uy ils hungarica Grun. var. |
longiceps Eh. Le pa || dats capitata Eh... eo a6 > By, Se

—— montanum Schum.. . pis De —— icostauron Grun. ae | il.

—— Mustela Eh. ! aa 9. ial,
parvulum (K.) Grun. oo |) UB} Tene Giewe ‘2 op Hf dle

Hantzaschia amphioxys (Eh.) Grun.| 3, 4, 6, 7, 8, 13. leptogongyla Eh. .. oe Uo

var. brasiliana Grun. limosa K. .. rls ae Uo kOe
f. minor : ue By ile ¢—— lucidula Grun. .. ae 2.

—- var. capensis Grun. 1. —— Magellanica Cleve fot IO.

— var. minor Per. il, microsphenia Eh. .. We

—— —— var. uticensis Grun. ils minima Grun. | il

— var. xerophila Grun. k. —— minutissima Grun. Ss elle

—— ( var. ?) antarctica Per. iL, : —— muralis Grun. aS iL,
elongata Grun. 4, 6, 10, 11. —— murrayi W. & G.S. West 4, 5, 6.

Hemiaulus ambiguus Jan. 6. —— var. elegans W. &

Larirella diaphana Bleisch IP G. S. West 4, 5, 6.

Mastogloia imperfecta Cleve 10. —— mutica K. ae oA 1, WBS =

Melosira crenulata K. iL, —_ var. sporangialis Per. hs

= var. lavis Grun. 8. —— —— var. capitulata Per. 1.
Dickiei K. 1. —— —— yar. cymbelloides Per. 1
distans (Eh.) K. Bf 6. —— —— var. truncata Per. 1
granulata (Eh.) Ralfs se) |) ale3, — var. ventricosa Cleve 1
Roseana Rab. U, muticopsiforme W. & G.S.
setosa Grey. il, West F 4,5, 6

—— sol K. De muticopsis Vv .H. il, By Gy 5 ©

—— varians Ag. PB, == var. capitata Per. .. Ile

—— spp. 455, a3: neglecta K. 7

Navicula aequalis Eh. ewlelie —— nobilis K. 7
affinis Eh. A a Up Se peraustralis W. & G. S.
amphirhynchus Eh. fs. West oe 6.

minor and major ite pa || ies peregrina K. By iil,
amphisbaena Bory. . 8, 11. perlepida Grun. S¥s 6
amphisphenia Eh. 8. —— pisiculus K. ue att 9.

The numbers refer to the following records: 1. Peragallo, 1908, Graham Land; 2. Fritsch, South Orkneys ;
3. Hooker, Ehrenberg, Cockburn Island; 4. Fritsch, 1901-4, Cape Adare; 5. Fritsch, 1910, Cape Adare ;
6. W. and G. S. West, 1907-9, Ross Island, South Victoria Land; 7. Ehrenberg, Falkland Island; 8. Ehrenberg,
Cape Horn; 9. Petit, Cape Horn; 10. Cleve, Magellans; 11. Ehrenberg, Kerguelen; 12. Reinsch, 1874-9, Ker-
guelen; 13. Bunt, Macquarie Island; 14. O’Meara, Marion Island.

50 TERRESTRIAL

DIATOMS OF MACQUARIE

ISLAND,

TABLE I1V.—Continued.

List of Antarctic Diatoms.—Continued.

Navicula platalea Eh. . .

8. Pinnularia Legumen Eh. le Ye Alta

—— platystoma Bh. Ss nond|. tb macilenta Eh. Ur tsdytt), Abi!

—— pleuronoctes Eh. Ney major K. 9, 10.

—— producta W. Sm. 9. mesolepta Eh. 9.
pterophaena (Eh.) De Toni 11. —— microstawron Eh. (i LOM S:

—— quinquenodis Grun. 1. — var. Brebjssonii (K. ) |
radiosa K. 6, 10. Hust. a Nes 13.

— var. acuta Grun. tts molaris Grun. (P. macra

—— rhomboides (Eh.) (Van- Grun.) 1, 13.

heurckia) : 9, 10 —— nivorum Per. 1.

— var. niimleurardes 10. — var. ? 13.
rhyncocephala K. f. 6. —— —— yar. elongata eer iv
(rostellata ? K.) 13. —— nodosa Eh. 9.

Ms —— stauroneiformis Per. 1.

semicruciata A. Sch. 9. stauroptera Grun. .. 10.

seminulum Grun. 4, 13. —— stauropteroides Fritsch 4,

serians K. 10. —— Thiebaudi Per. 1.

Shackletoni w. & Gs S. West 4,6 viridis K. a 7, 8, 9, 10, 11, 12.
—- var. pellucida W. & a var. commutator Gaui 9, 10.

S. West 4, 6. viridula W. Sm. 12.

sphaerophora K. 10. spp. (Eaton, Reinsch) 12.

streptoraphe Cleve var. Rhoikoneis interrupta Per. i

styliformis Cleve Wo Rhoicosphenia curvata Grun. 10.

—— subcapitata Greg. var. Rhopalodia gibba O. Muell. 1, 10.

stauroneiformis Pet. .. iL. gibberula (Eh.) O.M. var.

—— subtilissima Cleve . . 10. baltica O.M. BE
tabellaria K. (A); all, ventricosa O.M. 1.

— transversa A. Sch. 9. Stauroneis acuta W. Sm. 13.
vespa Eh. 11. amphilepta Eh. 10.

SHO. 00 Be on WP, anceps Eh. 4, 6, 9, 12
Neidium affine (Eh.) Cleve 118%, — var. birostris Eh. Uo Ails
Nitzschia Frauenfeldii var. ant- var. hyalina Brun.

arctica Per. 1. and Per. 13.
frustulum Grun. 10. goeppertiana pieicen IPA,

palea W. Sm. : ah ile 2 parvula (Grun.) Jan. 13.

subtilis (K.) Grun. var. —— phoenicenteron Eh. Ue ft, Oy tal, ales

(glacialis or paleacea) Grun. 6. var. gracilis J.B. &

Odontidium hyemale Lyng. 12 (O’Meara).’ Per. A an pa omlalte

Opephora Martyi Heribaud 13. Semen Eh. .. ele

Pinnularia appendiculata (Ag.) pygmaea Kreiger 1183,

Cleve 1, fo te 1B} Stenopterobia anceps Breb. 10.

—— Atwoodii Bow 13. Surirella angustata K. 4, 6, 9, 13

borealis Eh. 115 7, 8h 4S 5 th OS bifrons K. 13.
10, 11, 13 constricta Eh. ‘ mn 9.

= var. dilatata Eh. Up Bp il. Engleri (O.M.) f. an-

— Braunii Grun. 2. gustior O.M. 13.
brevicostata Cleve 13. euglypta Eh. Ho
—— Brebissonii K. Se ioe esa! 25 —— Falklandiae Eh. 7.
— var. diminuta V.H... 2. —— Guatamalensis Eh.. . 10.
cardinalis (Eh.) W. Sm. 11835 insularum Eh. To
—— Depauxii Per. il linearis W. Sm. 10.
divergens W.Sm. .. ae 9, 13. —— Maluwinensis Eh. tho
—— —— yar. elliptica Grun. .. | 10. —— microcora Eh. Zo
divergentissima (Grun.) splendida Eh. Te
Cleve 13. — var. tenera Greg. 10.
gibba K. 7, 8, 10, 11. striatula Turp. To
Greg. 4, 6. Synedra vaucheriae K. 12.

fasciata Lagerst. 13. 2 Synedra vaucheriae K. 13.

intermedia Lagerst. il, Synedra sp. : 2.

var. antarctica Per. .. 1 Tabellaria filecsnllogm (Roth. ) K. 6.

2 interrupta W. Sm. 13. Trachyneis aspera (Eh.) Cleve 6.
lata (Breb.) W. Sm. 1, 10, 13. Tropidoneis laevissima W. &

—— —— var. integra Per. 1. G.S. West 6.

—— —— var. minor Grun. 1.

The numbers refer to the following records: 1. Peragallo, 1908, Graham Land; 2. Fritsch, South Orkneys .
3. Hooker, Ehrenberg, Cockburn Island; 4. Fritsch, 1901-4, Cape Adare; 5. Fritsch, 1910, Cape Adare;
6. W. and G. S. West, 1907-9, Ross Island, South Victoria Land; 7. Ehrenberg, Falkland Island; 8. Ehrenberg,
Cape Horn; 9. Petit, Cape Horn; 10. Cleve, Magellans; 11. Ehrenberg, Kerguelen; 12. Reinsch, 1874-9, Ker
guelen; 13. Bunt, Macquarie Island; 14. O’Meara, Marion Island.

BY J. S. BUNT. 51

direct effect exerted, such as percentage of time the ground is frozen and duration of
each freeze, and partly from the type of vegetation occurring in any given area and, at
Macquarie Is., this depends principally on exposure to wind. The type of plant
community, under these conditions, largely influences the type of soil which develops.

It has already been noted that Pinnularia molaris Grun. is a dominant species in
soils 2, 3, 5, 9 and 10 which are in many respects dissimilar. This diatom is clearly an
adaptable type, since it also appears as a minor constituent in three other soils. Its

TABLE V.
Diatoms Dominant in Ten Macquarie Island Soils.

Soil
Number. Dominant Species.
1 Diatomella Balfouriana Grev., Pinnularia fasciata Lagerst., Pinnularia
nivorum Per. var. ?
2 Pinnularia molaris Grun.
33 Diploneis Smithii (Breb.) Cleve, Frustulia rhomboides De Toni, Pin-
nularia molaris Grun.
4 Achnanthes subsalsa Peterson.
5 Pinnularia molaris Grun., Pinnularia lata (Breb.) W. Sm.
6 None.
7 Hantzschia amphioxys (Eh.) Grun., Pinnularia Atwoodii Per.
8 Melosira spp.
9 Pinnularia molaris Grun., Pinnularia interrupta W.Sm., Melosira spp.
10 Eunotia lunaris Grun., Eunotia trinacria Krasske, Pinnularia molaris
Grun.

absence from soil no. 6 may be explained mainly by the relative lack of moisture and
from soil no. 7 by the undoubted salinity of this situation on rocks constantly receiving
sea spray. For many of the other species an explanation of the distribution patterns
may not be so simple for, unlike the vascular flora which is controlled by fairly readily
definable factors, chiefly exposure to wind and position of water table, the number of
controlling variables is large and frequently cannot be measured. In this connection

TABLE VI.
Distribution of Diatoms in Macquarie Island Soils.

Number of Soil Number of
Environments. Species.
1 20
2, 9
3 li
4 6
5 1
6 | 1
7 1
8 1
9 and 10 —

one should mention several factors discussed by Pringsheim (1950), viz. the possible
influence of simple competition by other organisms in the same environment for
available nutrients and the not uncommon production of antibiotic substances by many
of these organisms to which some algae are susceptible. Certain instances of antibiotic
activity in soil, once subject to doubt by some workers, have been quoted by Stevenson
and Lochhead (1953) and many of the fungi from Macquarie Is. produce these sub-
stances. Furthermore, some algae are more or less dependent on growth substances and

OZ TERRESTRIAL DIATOMS OF MACQUARIE ISLAND,

vitamins which may be produced and excreted by other microorganisms so that complete
autotrophism is not an invariable characteristic of this group. Such factors should be
included with the more commonly considered effects of pH, salinity, etc.

Table II lists the species of diatoms found in samples from the ponds, lakes and
watercourses of the island. The general distribution pattern shown in Table VII agrees
rather well with that for the soils. Only three species were found common to 5 or more
habitats and 36% were recorded from single situations. The greatest variability occurs
in the ponds of the raised coast terrace sub-glacial herbfields, but certain affinities were
apparent in the remaining samples examined. Of the 9 species found in a plateau creek
(sample no. 2), 5 occur in Prion Lake; the 12 species in the small lake (sample no. 5)
all oceur in Prion Lake, the bottom mud of which contained only 4 more species than a
sample taken in shallow water near the shore. The two samples (nos. 8 and 9) from
ponds in the wet tussock grassland contained few species, four of which were common
to both habitats.

TABLE VII.
Distribution of Freshwater Diatoms.
Number of Freshwater Number of
Environments. Species.

1 | 15

2 | 9

3 11

4 | 4

5 | 1

6 | 1
to. 8 —

9 1
10 to 11 —

The factors influencing the growth of diatoms in water are probably somewhat.
different from those which apply in soil. The characteristics of the waters sampled
showed considerable variation. Within the sub-glacial herbfields of the coast one finds
a range of water environments including muddy, foul-smelling stagnant ponds supporting
a dense and complex population of microorganisms (often dominated by purple and
filamentous sulphur bacteria), clear almost stagnant pools of varying size rich in algae
and water-tolerant or hydrophilic vascular plants such as Callitriche antarctica and
Ranunculus biternatus and small, slow-flowing, ill-defined freshwater runners passing as
surface drainage from the inland edge of the herbfields to the sea. Thus, at one extreme,
one might expect the complexity of a soil environment and at the other situations in
which factors such as oxygen tension, pH and simple nutrient supply are of greatest
importance.

The simplicity of the diatom flora found in the small, stagnant pools of the wet
tussock grassland is probably influenced to a large extent by the saline conditions
existing so close to the sea. A sample of rain water collected in this vicinity was found
to contain 0-88 gm. of NaCl/litre. As yet, little is known of the limnology of the
plateau lakes. Geologically there are several types; water-filled deep rock-basins of
glacial origin such as Prion Lake, small tarns and rather more extensive ponded
depressions such as Island Lake. These bodies of water do not always have outlets (e.g.
Prion Lake) and are frozen over during the winter. The ecological environment, then,
exhibits a certain diversity. Prion Lake, which was investigated briefly by Lindholm,
Jerums and the author in February 1952, is more than 100 feet deep near the centre,
with steep walls and a narrow bench platform to which most of the scant plant life is
restricted. The bacterial content is low, the summer temperature of the surface is
6°—8°C. and the pH close to 7-0. 0-068 gm. of chlorides was recorded from Scobles Lake

Or
ww

BYGMdis ace UIE.

further to the north. Copepods were the only prominent animal organisms taken in
plankton hauls. The climatic peculiarities of the island are considered to be the chief
factors limiting a wider diversity amongst the diatoms (and other forms of life) of the
lakes since the biosphere is limited largely to the shallow water where the physical
environment is more or less uniform in every body of water, small or large, on the
plateau. This view may require revision if a full-scale limnological programme is under-
taken at a later date.

In the fluvio-glacial sediments, differences such as occur in the soil and water are
not so apparent. The species recorded are listed in table III and it will be seen from
table VIII that 10 of the 37 species occur in 10 or more of the strata examined. This
points to a greater uniformity of conditions at the time the sediments were being formed
than exists today. Of greater significance, however, are several differences between
what may be designated the old and the new floras, viz. the absence particularly of
Fragilaria Harrisonii (W. Sm.) Grun. and Fragilaria virescens Ralfs, but also of
Cocconeis placentula Eh.t from the present-day flora, whereas the former two species
were abundant and widespread at the close of the last ice age. They were not found in

TABLE VIII
Distribution of Diatoms in the Fluvio-glacial Sediments.

Number of Number of Number of Number of

Strata. Species. Strata. Species.
1 6 9 1
2 5 10 1
3 3 11 2
4 2 12 1
5 2 13 4
6 2 14 | 1
7 3 19 i 1
8 3

the deep peat of the raised coast terrace which has been uplifted and colonized in
comparatively recent times. It is difficult to postulate a satisfactory explanation for
the disappearance of these once prominent forms since 33 of the 37 fossil species still
constitute part of the flora. It might be suggested that the conditions which once
favoured their growth no longer prevail and yet this seems rather unusual when one
considers the severity of the past climate, and that Fragilaria virescens Ralfs has been
found at Ross Is. (antarctica). Further, both species have been described from Europe
which would indicate tolerance to a wide range of conditions.

Another possibility is that some intervening process or event has caused their
extinction. The strata examined all underlie gravelly sediments of varying thickness
which may indicate a further brief period of glaciation or an extremely wet period
accompanied by very active erosion. Both might have been sufficient to bring about a
marked alteration or even elimination of the diatom flora. Why then should two of the
most prominent species be lost? In the event of complete extinction, which seems
doubtful, it does not seem likely that practically the same flora should be reintroduced
by such a haphazard agency as wind or even flying birds with fixed migratory habits, —
which should not be ignored as carriers.

In any event the dominant flora has changed in a striking fashion and the possi-
bility of similar changes in the vascular flora should not be overlooked by workers in
that field, already investigated hypothetically by Cheeseman (1919), Rudmose Brown
(1912) and others.

Finally, some comparison must be made between the diatom floras of the three
major environments. A total of 46 species was found in the soil, 43 in the water and 37

+The author has since found this species in a single sample from a lake on the plateau.

54 TERRESTRIAL DIATOMS OF MACQUARIE ISLAND,

in the old sediments. Twenty-five species were common to all three environments. The
marked similarity between the soil and freshwater floras is a clear reflection, partly of
the high water status of the soils and partly of the relative lack of diversity of species
in the freshwater environments, and shows that an ecological grouping of the species
such as has been suggested by Petersen (1935) would be of little value under the
conditions existing at Macquarie Island.

THE AFFINITIES, DISTRIBUTION AND ORIGIN OF TERRESTRIAL DIATOMS IN ANTARCTIC AND
SUBANTARCTIC REGIONS.

Table IV, which lists the species recorded from subantarctic islands and the
antarctic mainland, is summarized in table IX. About 340 species and varieties com-
prising 43 genera are included. Fifteen genera are represented by only one species each,
15 by 2-5 species, 3 by 6-10 species and 10 by 11 or more species. From the information
in table X it will be seen that there are no major affinities between the diatom floras of

TABLE IX.

Statistics of the Species and Genera of Diatoms Recorded in the Antarctic.

| Number of
| Number of Number of Number of Species and
Geographic Location. | Species and Species. Genera. | Varieties not
| Varieties. | | Recorded Elsewhere.
|

Graham Land .. a ce | 74 54 17 | 61
South Orkneys Ae a5 als) 15 10 11
Cockburn Island Re ae 8 7 5 2
Cape Adare a oa oh 29 27 12 13
Ross Island and South Victoria

Land ae At ae 32 30 17 18
Falkland Islands ae oes 51 49 13 20
Cape Horn ae 53 ys 68 63 16 36
Magelans as an ale 54 52 nA 34
Kerguelen Island aa ee 56 53 17 30
Macquarie Island ae oe 60 59 24 41
Marion Island .. af ae 5 5 4 4
Bouvet Island aS oe 1 1 1 =
South Georgia a a 19 19 10 ==

|

the various zones explored. Only five species are common to Macquarie Is. and Cape
Adare and South Victoria Land immediately to the south. Hight species are common
to the Magellans and Falkland Is. and 4 species are common to the Magellans and
Graham Land. These slight affinities could well be explained by chance in view of the
overriding differences which exist in the records.

The reason for this situation must be sought in the methods by which diatoms have
been introduced to these relatively isolated environments. The principal agent is
believed to be wind, for, although migratory birds must also be regarded of some
importance, the lack of affinity existing between the species at the various stations
implies the more accidental process of aerial dissemination. Hooker (1843) recorded
diatoms in dust collected aboard ship off the Cape Verde Islands and Chapman and
Grayson described 23 species in “red rain” at St. Kilda, Victoria, in 1903. More recently
van Overeem (1937) and Polunin and Kelly (1952) have collected by direct methods
the living spores of various organisms carried by air currents in the upper atmosphere.
Many of the species recorded on the antarctic mainland appear not to be part of the
living population but air-borne frustules which have been deposited and preserved in
the ice.

It is clear, then, that with wind as the principal agent of introduction, few affinities
could be expected since a great number of species appear to be capable of successful

BYAULS- BUNT: 55

growth under the severest conditions. The resistance of these organisms to partial
desiccation during transport is probably an important factor influencing survival. It is
also reasonably certain that the present diatom floras are more or less stable and have
been long established in their various environments. This is indicated partly by the
presence of local varieties which are not found in other situations. Achnanthes brevipes
Ag. var. intermedia K., for example, a species common on Macquarie Is., has been shown
to possess certain minor characteristics which distinguish it from the same organism
occurring elsewhere. Similar cases are quite common in the literature. It is shown
further by the comparative lack of diversity of the recorded lists of species and by the
fact that there has not been a very great increase in the number of species on Macquarie
Is. since the close of the last ice age. Also the improbability of a newly-introduced
species competing successfully in an already well-developed community is well recog-
nized. The nature of the climate, of course, is a further limiting factor.

TABLE X.

Distribution of Diatoms in Antarctic Regions.

Number of Number of
Areas. | Species.
|
1 224
2 43
3 16
4 9
5 or more 3

It should be mentioned here that some of the collections from subantarctic islands
are believed not to be complete. Those from Marion and Bouvet Islands are obviously
only isolated records, but. Kerguelen is a larger island than Macquarie, with a slightly
more favourable climate and offers a much wider range of environments and yet only
56 species and varieties have been identified. A similar situation applies to the
Falklands and Magellans which have a further advantage in their close connection with
South America.

Finally, some mention should be made of the differences which exist between the
diatoms of the arctic and antarctic. Mann (1937) has explained the reasons for the
greater complexity of marine forms in the antarctic than in the arctic by the openness
of the Southern Ocean and its connections with the other large oceans of the world. A
somewhat similar argument may be applied in reverse for the terrestrial diatoms. Most
land masses in the antarctic and subantarctic are extremely isolated. This is not so in
the arctic. Further, the climate in southern latitudes is far more severe (Cheeseman,
1919) than in the north and it is the combination of these two factors, severe climate
and geographic isolation, that may be held responsible for the paucity of the diatom
floras in these southerly regions.

Acknowledgements.

The author wishes to express his appreciation for the continued financial support of
A.N.A.R.E. since his return to Australia in 1952 and for the permission which was given
to return to Macquarie Island during the 1953 change-over operations. His thanks are
also due to the University of Sydney for provision of laboratory facilities in the Faculty
of Agriculture, to Mr. J. M. Vincent for criticism of the manuscript and to Mr. B. W.
Taylor, A.N.A.R.E. biologist, for making available samples of freshwater algae. Finally,
he wishes to record his thanks for the helpful advice and assistance given by Professor
L. M. Baas Becking, Mr. E. J. Ferguson-Wood and Mr. L. Crosby of the Fisheries Division
of C.S.1.R.O. at Cronulla, New South Wales.

56 TERRESTRIAL DIATOMS OF MACQUARIE ISLAND,

Bibliography.

Biren, L. C., and CLARK, D. P., 1953.—Forest Soil as an Ecological Community, with special
reference to the fauna. Quart. Rev. Biol., 28, 1: 13-36.

CHAPMAN, F., and GRAYSON, H. J., 1903-4.—On “red rain’, with special reference to its occur-
rence in Victoria. With a note on Melbourne dust. Vict. Naturalist, 20:17.

CHEPSEMAN, T. F., 1919.—The Vascular Flora of Macquarie Island. Australasian Antarctic
Hapedition—1911-14. Scientific Reports, Series C.—Zool. and Bot., VII, 3.

CLEVE, T. P., 1900.—Expédition Suédoise au détroit de Magellan, sous les ordres de Nordenskiold.
Incomplete reference from Peragallo, 1921.

DETON!I, B., 1891-4.—Sylloge Algarum omnium hucusque cognitarum, II, Bacillarieae.

TSHRENBERG, E. C., 1854.—Mikrogeologie (Leipzig) p. 2, t. XXXV, A.I., p. 262, t. XXXV, A.II,
pp. 287-9.

FritscH, F. E., 1911.—Fresh water algae collected in the South Orkneys by Mr. R. N. Rudmose
Brown of the Scottish National Antarctic Expedition, 1902-4. J. Linn. Soc. Bot., XU: 293-338.

——— , 1912.—Fresh Water Algae. National Antarctic Eapedition, 1901-4.. Natural
History, VI.

, 1917.—Fresh Water Algae. British Antarctic Haxpedition, 1910. Botany, Pt. I,
pp. 1-16.

GREGORY, W., 1855.—On a remarkable group of diatomaceous forms, with remarks on shape
or outline as a specific character. Trans. Mic. Soc. London, 3: 10-15.

HENDEY, N. INGRAM, 1937.—The plankton diatoms of the southern seas. Discovery Reports,
W, 2SWIlS 105 BZ4,

HoLMBOER, J., 1902.—Navicula mutica K., aus dem antarktischen Festlande. Nyt Mag. f. Natur-
videnskab, XL: pp. 209-222, pls. 3-4.

Hooker, J. D., 1839-43.—The Botany of the Antarctic Voyage of H.M. Discovery Ships
“Hrebus” and “Terror”. (Flora Antarctica.) Algae (by W. H. Harvey and J. D. Hooker).
Vol. I, pp. 175-193 (1845); Vol. II, pp. 454-519 (1846-7).

Hustept, F., 1930?.—in Rabenhorst’s Kryptogamenflora, Bd. 7, Ht. 2.

, 1930.—Die Stisswasser-flora Mitteleuropas, Ht. 10, Diatomeae (Jena).

MANGIN, L., 1914.—Sur le polymorphisme de certaines diatomées de 1l’Antarctique. Compt. Rend.
Acad, Sci., Paris, 159: 476-484.

MANN, A., 1937.—Diatoms. Australasian Antarctic EHapedition, 1911-14, Scientific Reports,
Series C, Zool. and Bot., Vol. I, pt. I.

Mawson, D., 1943.—Macquarie Island, its Geography and Geology (from the records of L. R.
Blake). Australasian Antarctic Expedition, 1911-14, Scientific Reports, Series A, Vol. V.

MUuuer, O., 1909.—Incomplete reference from Peragallo, 1921.

C’MeEaArRA, E., 1876.—On the Diatomaceous Gatherings made at Kerguelen’s Land by H. N.
Moseley, H.M.S. “Challenger”, J. Linn. Soc., Bot., XV: 55-59.

PERAGALLO, M., 1921.—Diatomées d’Eau Douce. Eapédition Antarctique Frangais, 1908-10.

PETERSON, J. Boyer, 1935.—Studies on the Biology and Taxonomy of Soil Algae. Dansk. Botanisk
Arkiv, 8: 1-172.

Perit, P., 1888.—Diatomées récoltées dans le voisinage du Cap Horn. Mission scientifique dw
Cap Horn. T.V. Botanique.

Piper, C. S., 1938.—Soils from Subantarctic Islands. Section 1. An examination of soils from
Possession, Heard, Kerguelen and Macquarie Islands. B.A.N.Z.A.R.E., 1929-31. Scientific
Reports, Series A, Vol. II, pt. 7.

POLUNIN, N., and Keuuy, C. D., 1952.—Arctic Aerobiology. Fungi, bacteria, etc., caught in
the air during flights over the geographical North Pole. Natwre, 170, No. 4321.

PRINGSHEIM, E. G., 1950.—The Cultivation of Algae. Hndeavour, 9, No. 35: 138-143.

PRITCHARD’S Infusoria, London, 1861.

REINSCH, P. F., 1876.—Species ac Genera nova Algarum aquae dulcis, quae sunt inventa in
speciminibus in expeditione Vener. transit. hieme 1874-75 in insula Kerguelensi a clar
Eaton collectis. J. Linn. Soc., Bot... XV: 205-221.

———, 1879.—Fresh water algae collected by the Rev. A. E. Haton. Algae aquae dulcis
Insulae Kerguelensis. Phil. Trans. Roy. Soc. London, 168: 65-92.

, 1890.— Die Siisswasseralgenflora von Stid-Georgien, in G. Neumayer, Die dewtschen
Expeditionen und ihre Ergebnisse, Bd. I1: 329-365, tabb. 1-4.

RuDMOSE Brown, R. N., 1912.—The Problems of Antarctic Plant Life. Botanical results of the
Scottish National Antarctic Expedition.

Scumipt, A., 1874-1938.—Atlas der Diatomaceenkunde. Fortgesetzt von M. Schmidt, F. Fricke,
O. Miiller, H. Heiden und F. Hustedt. Aschersleben-Leipzig.

SmitH, W., 1853-56.—Synopsis of the British. Diatomaceae. London. (Vols. 1 and 2.)

STEVENSON, I. L., and LocHHEAD, A. G., 1953.—The use of a percolation technique in studying
antibiotic production in soil. Canad. J. Bot., 31: 23-27.

VAN OVEREEM, Marig A., 1937.—On Green Organisms Occurring in the Lower Troposphere.
Recueil des Travaux Botaniques Neerlandais, 34: 388. q

West, W. and G. S., 1911.—Freshwater Algae. British Antarctic Expedition, 1907-9, Scientific
Reports, Vol. I, pt. VII.

Wiuson, O. T., 1927 Asymmetric variation in Cocconeis scutellum Eh. Am. J. Bot., 14: 267.

a,

Proc. Linn. Soc. N.S.W., 1954. PLATE I.

Hibbertia serrata, sp. nov.

PLATE It.

Proc. Linn. Soc. N.S.W., 1954.

AROS

2 REGS.

Diatoms from Macquarie Island.

PLATE It,

Proc. Linn. Soc. N.S.W., 1954.

Diatoms from Macquarie Island.

BY J. S. BUNT. 57

EXPLANATION OF PLATES II-III.

Plate ii.

1, Achnanthes brevipes Ag. var. intermedia Kutz. 73 x 20 uw. 10-11 striae/ 10 mw x 1000.
2, A. brevipes var. intermedia. 58 x 20 mw. 9-10 striae/ 10 w. x 1000. 3, Achnanthes lanceolata
(Breb.) Grun. 28 x 9 w. 12 costae/ 10 uw. x 2000. 4, Fragilaria (virescens?) Ralfs. 13-20 x 3 mu.
23 costae/ 10 uw. x 700. 5, Cymbella pusilla Grun. var.? 33 x 5:7 uw. 16 costae/ 10 mw. x 700.
6, Diploneis smithii (Breb.) Cleve. 32 x 15 w. 12 striae/ 10 uw. x 2000. 7, Achnanthes subsalsa
Peterson. 18 x 8 uw. 14 costae/ 10 uw. x 1000. 8, Diatomella Balfouriana Grey. Length 15 uw.
21 costae/ 10 uw. x 1000. 9, Fragilaria Harrisonii (W. Smith) Grun. 21 x 13 uw. 10 costae/ 10 wm x
700. 10, Hunotia trinacria Krasske. 16 x 2 uw. approx. 20 costae/ 10 w. x 700. 11, Hunotia lunaris
Grun. 36 x 3 w. 22-25 costae/ 10 uw. x 700. 12, Hunotia alpina (Naeg.) Hust. 36 x 3 uw. 15 costae/
10 w. x 700. 138, 14, Fragilaria capucina Desm. 34-53 x 3 w. 17 costae/ 10 w. x 1000. 15, Fragilaria
virescens Ralfs. Length 40 uw. 16-18 costae/ 10 uw. x 700. 16, Gomphonema parvulum (Kutz.) Grun.
382 x 9 w. 13-14 costae/ 10 uw x 1000. 17, Gomphonema intricatum Kutz. 54 x 10 uw. 12 costae/
10 w x 1000. 18, Fragilariopsis antarctica Castr. 30 x 8 uw. 5-6 striae/ 10 uw. x 1000. 19, ? Frus-
tulia minuta Rab. 25 x 4 mu. x 1000. 20a and b, Melosira granulata (Eh.) Ralfs. 20a, 11 x 7-5 wy.
x 2000. 20b, 27 x 4 w x 700. 21, Navicula (rostellata?) Kutz. 40 x 5 w. 18 striae/ 10 um.
x 1000. 22, Opephora Martyi Heribaud. Length 14 uw. 10 costae/ 10 uw. x 2000. 23, Pinnularia
appendiculata (Ag.) Cleve. 40 x 6 uw. 14 costae/ 10 w. x 700. 24, Pinnularia molaris Grun.
37 x 5 uw. 16 costae/ 10 wu. x 1000. 25, P. molaris. 44 x 6 w. 15 costae/ 10 uw. x 1000. 26, Pinnularia
divergentissima (Grun.) Cleve. 36 x 6 w. 11-12 costae/ 10 uw. x 700. 27, Pinnularia microstauron,
Eh. 48 x 8 uw. 16 costae/ 10 w. x 1000. 28, Hantzschia amphioxys (Eh.) Grun. 75 x 10 uw. 23 costae/
10 w. x 2000. 29, Pinnularia brevicostata Cleve. 78 x 15 wu. 9 costae/ 10 mw. x 1000. 30, Pinnularia
cardinalis (Eh.) W. Smith. 74 x 12 yw. 10 costae/ 10 uw. x 1000.

Plate iii.

1, Pinnularia divergens W. Smith. 120 x 22 uw. 8-9 costae/ 10 uw. x 2000. 2, Pinnularia lata
(Breb.) W. Smith. 85 x 20 w. 4 costae/ 10 w. x 2000. 3a, b, c, Stauroneis acuta W. Smith.
143 x 28 mw. 15 striae/ 10 w. 3a and 3c, x 1000. 3b, x 2000. 4, Pinnularia microstauron Eh.
60 x 12 w 15 costae/ 10 uw. x 1000. 5, Surirella angustata Kutz. 74 x14 uw. x 1000. 6, ? Synedra
vaucheriae Kutz. 24 x 4 uw. 16 costae/ 10 uw. x 1000. 7, Surirella bifrons Kutz. 100 x 28 uw. x 1000.
8, Stawroneis anceps Eh. var. hyalina Brun and Per. 68 x 13 uw. 20 striae/ 10 uw. x 1000.

58

THE FOSSIL DIPTERA OF THE TERTIARY REDBANK PLAINS SERIES,
QUEENSLAND. i

By E. F. Riek.
(Five Text-figures. )
[Read 26th May, 1954.]

Synopsis.
A new genus and species of Tipulidae and a new monotypic genus and another two new
species of Mycetophilidae are described.

An earlier paper (Riek, 1952) described the Mecoptera and Neuroptera of this bed.
This paper deals with the small collection of about twenty-five recognizable specimens:
of Diptera. Most of them belong to the Mycetophilidae and have a wing length usually
less than 4:0 mm. One Tipulid species has a wing of 10:0 mm. and there is a fragment
of a moderately-sized Brachycera probably referable to the Muscoidea.

Family TIPULIDAHE.
Subfamily LIMNONIINAE.
Tribe HEXxATOMINI.
Subtribe PSEUDOLIMNOPHILARIA.

The Tipulids are slender-bodied flies commonly called crane-flies. They are
distinguished on wing venation from most other Nematocera by retaining two long,
simple anal veins. In many species there is a closed discal cell.

The family is a very large one, subdivided into three subfamilies, with many tribes
and lesser categories. The subfamily Limnoniinae, with the largest number of genera,
contains small, fragile species, most of which are referred to the tribe Hexatomini.

Genus PROTOLIMNOPHILA, NOv.

Genotype, Protolimnophila superba, sp. nov.

Wing: Very similar to Archilimnophila Alexander, 1934. Compared with Prolimno-
phila, its discal cell basally is more truncate, the branches M, and M, are more widely
separated, and all of M is more sinuous. The m-cu cross-vein slopes in the opposite
direction and CuP almost, if not completely, reaches the wing margin.

PROTOLIMNOPHILA SUPERBA, SD. NOV.
(Text-fig. 1.)

Wing: Length 10 mm.; wing fairly broad, not narrowed markedly at base, apex
rounded; characters as outlined in the generic diagnosis; m-cu cross-vein well before
the forking of M;,,, stem of M,,. strongly angled at the r-m cross-vein, M;,, and CuA
widely separated; M,,. arched towards Rs after the cross-vein to M,; M, and M, well
separated, arched strongly at origin; CuP distinct, close to CuA and almost, if not
completely, reaching the wing margin; 1A and 2A straight, except at wing margin.

Type.—Holotype wing, F.14344, in the University of Queensland, Department of
Geology Collection.

Type Locality and Horizon.—Redbank Plains Fish Bed, Lower Tertiary, at type
outcrop. :

Only the perfect holotype wing of this species is known.

Family MYCETOPHILIDAE.
Moderately small, rather delicate, slender flies with elongated coxae.
Wings relatively large: Sc present, though sometimes very short; Rs normally
simple but occasionally with an upper branch; M,,. normally forked and M,,, often

BY E. F. RIEK. 59

appearing as a branch of CuA and free of M,,,, though sometimes connected to CuA
by a cross-vein; CuP very reduced; 1A distinct or reduced, 2A normally absent.

There are a number of subfamilies based mainly on the wing venation or at least
separable on wing venation. Two of the subfamilies are known to occur in this deposit.

Subfamily DIroMyYIINAE.
This rather primitive subfamily is characterized by the forked Rs in which R, is
quite long, and Sc is short and ends freely. In other groups where Rs is forked R,
is very short and Sc is long and ends in the costa.

Genus CENTROCNEMIS Philippi, 1865.
A species based on a fragmentary new specimen, lacking Sc, is referred to this
genus tentatively.

CF .
~~

Text-figs. 1-5.

1, F.14344, holotype wing of Protolimnophila superba, sp. nov. x 4 ca. 2, F.14345A, holotype
wing fragment of Centrocnemis ? imperfecta, sp. nov. x 9 ca. 3, F.14347A, holotype wing of
Protasmanina nana, sp. nov. x 9 ca. 4, F.14348, wing of Protasmanina nana. x 9 ca.. 5,
F.14361A, holotype wing fragment of Aneura ? apicalis, sp. nov. x 9 ca. y

CENTROCNEMIS ? IMPERFECTA, Sp. NOV.
(Text-fig. 2.)

Wing: Length of fragment 5:0 mm., indicating a total length of 6:5 to 7-0 mm.;
R, curved slightly round towards the apex; Rs forked about the middle of its length,
upper branch only gradually diverging away from lower branch, approaching close
to R, at the middle of its length; M,.. forked slightly before Rs with the lower branch
continuing the line of the basal stem; origin of M,,, not preserved; CuA slightly
sinuated towards the wing margin.

Type—Holotype wing F.14345 A-B in the University of Queensland, Department of
Geology Collection.

Type Locality and Horizon.—Redbank Plains Fish Bed, Lower Tertiary, at type
outcrop.

There is a second imperfect specimen of 4:5 mm. which may belong to this species.

Subfamily ScIoPHILINAE.

One of the larger subfamilies, differing from the typical Mycetophilinae in usually
possessing a long Sc ending in the costa. Mg,,, is fused basally to CuA and Rs is simple.

The species described below probably both belong in the tribe Sciophilini, in which
Sc is nearly always long and R,, beyond the origin of Rs, is several times as long as.
the r-m cross-vein, which is usually oblique. In this tribe macrotrichia are present on
the wing membrane, but the preservation of the material does not permit this being
ascertained.

‘60 FOSSIL DIPTERA OF THE TERTIARY REDBANK PLAINS SERIES, QUEENSLAND.

Genus PROTASMANINA, NOvy.

(Genotype, Protasmanina nana, sp. nov.

Wing: Very similar to Tasmanina Tonnoir, 1929, but having a distinct 2A reaching
nearer the wing border than 1A. Sec ending in C after the origin of Rs, with a cross-
vein to R,; Rs transverse from its origin to the r-m cross-vein, which is more or less
longitudinal, about equal to the stem of M,,.; M,,, arising from CuA more basad than
the forking of M,,.; CuP long but weak; 1A ending well before the wing margin; 2A
strong, longer than 1A.

PROTASMANINA NANA, Sp. nov.
(Text-figs. 3-4.)

Wing: Length 3:5 to 4:0 mm., with holotype 4:0 mm.; wing fairly broad, apex
rounded; forking of M,,. close to origin of M,,.; M;,, arising from CuA just before the
first forking of M; R, and CuA strongly convex; CuP very concave, not quite reaching
the wing margin; 2A very strong; humeral cross-vein rather oblique.

Type.—Holotype wing F.14347 A-B in the University of Queensland, Department of
Geology Collection.

Type Locality and Horizon.—Redbank Plains Fish Bed, Lower Tertiary, at type
outcrop.

There are many specimens of this small fly. In some of the better preserved
specimens where R, stands out strongly convex it is hard to distinguish Sc, which lies
deep in a groove relatively close to C. For this reason F.14347, which shows Sc clearly,
has been chosen as holotype even though the stems of Rs and 2A are not so clearly
preserved.

Genus ANEURA Marshall, 1896.

Genotype, Aneura boletinoides Marshall, 1896.

Wing: Sc without an apical cross-vein to R,; M,,. forking towards the wing margin
so that the stem is longer than the branches and longer than the r-m cross-vein, which
itself is rather long; only 1A distinct.

The genus is recorded from New Zealand and Tasmania. An imperfectly known
species from this bed is considered tentatively in this genus.

ANEURA ? APICALIS, Sp. NOV.
(Text-fig. 5.)

Wing: Length of fragment 2-5 mm., indicating a total length of from 3-5 to 4:0 mm.;
only the apical portion preserved; M,,, forking rather close to the margin so that the
stem is considerably longer than the branches; r-m cross-vein not half the stem of
M,,.; Se extending beyond the origin of Rs.

Type.—Holotype wing fragment F.14361 A-B in the University of Queensland;
Department of Geology Collection.

Type Locality and Horizon.—Redbank Plains Fish Bed, Lower Tertiary, at type
outerop.

There are two other fragmentary specimens.

Suborder BRACHYCEHRA.
Superfamily MUSCOIDEA.
There is a fragmentary specimen of a wing, F.14364 A-B, belonging very probably
in this superfamily. It is mentioned merely because it is the only fragment of the
Brachycera from these beds.

; References.

CURRAN, C. H., 1934.—The Families and Genera of North American Diptera. 512 pp.

Epwarps, FE. W., 1921.—A Note on the Dipterous Subfamily Ditomyinae with Descriptions of
New Recent and Fossil Forms, Ann. Mag. Nat. Hist. (9), 7: 431-36.

RiEK, EH. F., 1952.—The Fossil Insects of the Redbank Plains Series. Part 1. Univ. Qland.
Papers (Geol.), n.s. 4: 1-14.

Tonnorr, A. L., 1929.—Australian Mycetophilidae. Synopsis of the Genera, Proc. Linn. Soc.
N.S.W., 54: 584-614.

61

A SECOND SPECIMEN OF THE DRAGON-FLY AEHSCHNIDIOPSIS
FLINDERSIENSIS (WOODWARD) FROM THE QUEENSLAND
CRETACEOUS.

By EH. F. RIK.
(Plate iv; one text-figure.)
[Read 26th May, 1954.]

Synopsis.
A second, complete, hindwing of the dragon-fly Aeschnidiopsis flindersiensis (Woodward)
is described. The triangle and associated areas of the wing are of unusual shape and differ
from the interpretation of those structures given by Tillyard from the imperfect holotype.

This second specimen of the species consists of the reverse impression of a complete:
hindwing, somewhat larger than the holotype hindwing and having a length of 55 mm.
and a greatest breadth of 20 mm. The veins are clearly defined except for the apical
four or five millimetres and irregularly so along the posterior border. There is slight
crumpling of the anal area.

The specimen preserves clearly the whole of the triangle and associated areas of
the wing, and these differ somewhat from the interpretation given by Tillyard from
the imperfect holotype.

Family AKSCHNIDIIDAE Handlirsch.
Subfamily AESCHNIDIINAE Tillyard, 1917, Carpenter, 1932.
Genus ArscHNIDIOPSIS Tiliyard, 1917.

Genotype, Aeschna flindersiensis Woodward, 1884.

Hindwing only: The genus differs from Aeschnidium Westwood in the incomplete
arculus, in having an excessively narrowed triangle and in the triangle being
quadrangular. It differs from Urogomphus in the structure of the lower end of the
triangle and in the first of the anal veins.

AESCHNIDIOPSIS FLINDERSIENSIS (Woodward), 1884.
(Plate iv; Text-fig. 1.)

Aeschna flindersiensis Woodward, 1884, Geol. Mag., n.s., 1: 337-339.

Aeschnidium fliindersiense Handlirsch, 1908, Die fossilen Insekten, etc.: 667.

Aeschnidiopsis flindersiensis Tillyard, 1917, Proc. Linn. Soc. N.S.W., 42: 691.

Type.—Hindwing in the Queensland Geological Survey, Brisbane.

Type Locality—Flinders River Beds, N. Queensland, Cretaceous.

A second hindwing of this species is in the Collection of the Geology Department,
University of Queensland (F.3162).

Description of Specimen F.3162: Length 55 mm. Costa somewhat thickened over
most of its length (to the pterostigma). There is a supplementary vein developed
between C and Sc from a double row of cellules. It begins well before the level of the
arculus, possibly from the base of the wing, and continues right to the nodus, but
there it is slightly zig-zagged. This vein is continued for a short distance beyond the
nodus. Se is well separated from the costa and continues strongly to the nodus..
Unfortunately there is slight fracturing at this point. The vein is continued through
the nodus, at first quite strong and straight, but approaching the pterostigma it
becomes zig-zagged and weaker and disappears a little before the pterostigma, where:
R approaches most closely to the costa. R a strong vein gradually approaching the
costa, particularly beyond the nodus, but deflected down rather strongly around the
pterostigma; pterostigma not sharply defined and traversed by cross-veins; Rs and M
divides at the arculus into the concave anterior branch Rs and a convex posterior

62 AERSCHNIDIOPSIS FLINDERSIENSIS FROM THE QUEENSLAND CRETACEOUS.

branch MA; arculus incomplete below (MA and CuP not connected except by normal
cross-veins). |The basic subdivisions of this area are as figured by Tillyard, but his
naming of the veins differs from that adopted here. The vein designated Ms is an
intercalated convex vein of the radial field, IR,. Distally a convex vein is intercalated
between R. and R, and shorter ones close to the wing margin. CuP is a fairly strong,
concave vein, running below the arculus to join the triangle at its proximal angle;
there it turns sharply downwards, at right angles to its previous course, forming the
proximal side of the triangle itself. At the lower end of the triangle it turns again
abruptly almost through a right angle and passes in a strong arch to the wing margin,
giving off one or two short branches on its lower side. The anal vein is a strongly-
marked, convex vein, running in a gentle curve from the base towards the proximal
angles of the triangle, where it is connected to CuP by a cross-vein. Branches of A
arising as a pectinate series on its lower side. All branches strongly curved, the
distal two appearing more as a forked vein with the stem of the most distal incorporated
in the subtriangle. The complete structure of the triangle is outstanding. First, the
triangle is really quadrangular, being closed at the lower end by a distinct but short,
almost transverse cross-vein which is in line with a similar one closing the subtriangle

Text-figure 1.—F.3162, Aeschnidiopsis flindersiensis (Woodward). Triangle and associated
structures. x 2 ca.

below. The triangle is very long and narrow, lying transverse to the length of the
wing. Its proximal margin is formed by portion of the concave vein CuP. Its upper
margin is a strong cross-vein connecting CuP to the strongly convex MA. At this
cross-vein MA forks into two convex veins, the lower branch MA, forms the distal
margin of the triangle and runs almost parallel to the cencave CuP to the lower end
of the triangle, which is clesed by another quite short cross-vein. Beyond the lower
end of the triangle MA, turns abruptly through more than a right angle and merges
into the wing membrane, becoming lost in the intercalated concave and convex branches
lying between MA, and CuP, though it is possible that it may still persist in one of
these branches. (These branches are designated Mspl in Tillyard’s figure.)

The subtriangle is bounded proximally by A, above by a strong cross-vein to CuP
and below by another, very short, cross-vein to CuP. Its proximal border is not
clearly defined, as there are one or two short branches running from the stem of A
into the area but not reaching its lower border.

The species is of interest mainly for its abnormal triangle and in being one of the
few known Cretaceous insects.

References.
CARPENTER, F. M., 1932.—Jurassic Insects from Solenhofen. Annals Carnegie Mus., 21 (3) : 97-129.
HANDLIRSCH, A., 1908.—Die fossilen Insekten, etc.: 667.
TrtuyarpD, R. J., 1917.—Mesozoic Insects of Queensland No. 2, Proc. Linn. Soc. N.S.W.,

42: 676-692.

WoopwarD, H., 1884.—On the Wing of a Neuropterous Insect from the Cretaceous Limestone
of Flinders River, North Queensland, Australia, Geol. Mag. (ms), 1: 337-339.

PLATE IV.
Aeschnidiopsis flindersiensis (Woodward), hindwing, F.3162. 1, x 2 ca.; 2, basal portion, x 4 ca.

63

NOTES ON THE BACTERIA BELONGING TO THE RHODOBACTERIINEAE BREED,
MURRAY AND HITCHENS, AND THE CHLAMYDOBACTERIALES BUCHANAN
OCCURRING AT MACQUARIE ISLAND.

By J. S. Buntr,* Australian National Antarctic Research Hxpedition.
(Plate v.)
[Read 26th May, 1954.]

Synopsis.

The presence is recorded of purple photosynthetic bacteria, filamentous sulphur bacteria
and “iron” bacteria under subantarctic conditions. This note is based on observations made
during a broad microbiological survey conducted by the author as a member of the 1951-52
field party of the Australian National Antarctic Research Expedition to Macquarie Island.

A. The CHLAMYDOBACTERIALES Buchanan.
1. The “Iron” Bacteria.

The climate of Macquarie Island is cold and wet (Mawson, 1943). In addition to
numerous creeks, there are a great many small ponds, especially in areas where the
water table is close to the surface of the ground, and the soils, even on the steeply
Sloping parts of the island, are sometimes constantly saturated with drainage water
from higher areas. In such situations, brown gelatinous masses or flocculent reddish-
brown sediments of ferric hydroxide are very common. An examination of a small
quantity of this material proved to be rich in a species of Crenothrix Cohn.

According to Halvorsen (1931), these bacteria occur in situations which would allow
spontaneous precipitation of iron hydroxides. They do not carry the reaction beyond
the point which would be reached in their absence, so that, although they are abundant,
their importance is a matter of some doubt. Further, Halvorsen (1931) has shown
that heterotrophic soil bacteria exert a significant influence on the precipitation of iron
by affecting the environmental conditions in the soil, and considers that their importance
in this respect has been under-estimated. The author has found that several types ot
heterotrophic bacteria common throughout the soils of Macquarie Island are capable of
precipitating iron hydroxide readily from ferric ammonium citrate. It is suggested
that these specific types, by their direct action, and the general heterotrophic population,
by its indirect influence, are the most important microbial agents causing iron deposition
under these conditions.

2. Filamentous Sulphur Bacteria.

Representatives of this group were found under both marine and fresh-water
conditions. A bottle of sea-water, collected at 11 fathoms in Haselborough Bay and
stored at about 10°C. for several months, developed a very fine veil-like growth attached
to the walls of the jar. A stained smear showed the organism to have the morphology
of Thiothriz marina Molisch. This species was subsequently found in salt-water rock
pools, often associated with decomposing seaweed.

A similar growth was occasionally observed in slowly moving fresh water, especially
in the Finch Creek sub-glacial herbfields. The thick, felted, pale yellow mass appeared
to consist of several species of Thiothrix Winogradsky, the dominant type occurring
as long, unbranched filaments (Plate v, fig. 2) which break up into segments 1—2y long
and about 0-54 wide. The second type (Plate v, fig. 3), which has filaments up to 3u
wide and of varying length, may be Thiothrix nivea (Rab.) Winogradsky. Both
organisms were found to be Gram-negative.

* School of Agriculture, University of Sydney.

64 BACTERIA OCCURRING AT MACQUARIE ISLAND.

B. The RHODOBACTERIINEAE Breed, Murray and Hitchens.

Purple photosynthetic. bacteria are sometimes to be seen in muddy pools in the
sub-glacial herbfields. They are most noticeable in the boggy areas of the north-west
coast. An attempt was made to cultivate members of this group, using five different
samples of soil. A sodium sulphide medium (Allen, 1950) was used to stimulate growth
of the Thiorhodaceae and a medium containing sodium butyrate and yeast extract
(Allen, 1950) for the aerobic and anaerobic Athiorhodaceae. A purple-pigmented
organism (Plate v, fig. 1) developed from only one of the samples, a bog soil from the
sub-glacial herbfields 500 feet above sea level, inoculated into the sodium butyrate
medium and incubated in the light under anaerobic conditions. The bacteria were
Gram-negative, 3-bu long by 2u wide, and may be a species of Rhodopseudomonas
Kluyver and van Niel.

The presence of purple bacteria in visible quantities was not realized until the
author made a further brief visit to Macquarie Island in March, 1953, when a special
search was made to supply material for Professor Baas Becking, of the C.S.1I.R.O.
Fisheries Division, Cronulla. It seems highly probable that a more detailed study by
Professor Baas Becking may uncover a greater diversity of types and provide some
information regarding their physiological activities.

References.

ALLEN, O. N., 1950.—Experiments in Soil Bacteriology. Burgess Pub. Co., Minn.

BerGHy’s Manual of Determinative Bacteriology. Sixth Hdition.

HALVORSEN, H. O., 1931.—Studies on the Transformation of Iron in Nature. III. The Effect of
CO, on the Equilibrium in Tron Solutions, Soil Sci., 32:141-65.

MAwson, D., 1943.—Macquarie Island, Its Geography and Geology, Australasian Antarctic
Expedition, 1911-41, Scientific Reports, Series A, Vol. V.

EXPLANATION OF PLATE V.
1, ? Rhodopseudomonas sp. in enrichment culture. Gram stain. (3-5 x 2u.)
2, Thiothrix sp. Filaments and individual segments. Smear from material collected in the
field. Gram stain.
3, Thiothrix nivea ?. Filaments in smear of sample collected in the field. Gram stain.

THE GENUS THEOBALDIA (DIPTERA, CULICIDAE) IN VICTORIA.

By N. V. Dosrotworsky, Georgina Sweet Fellow in Economic Entomology,
Zoology Department, University of Melbourne.

(Nine Text-figures. )

[Read 30th June, 1954.]

Synopsis.

Three species of the genus Theobaldia, T. frenchi Edw., T. hilli Edw., and T. inconspicua
. Lee have been previously recorded from Victoria. To this list are now added T. littleri Taylor,
T. victoriensis, n. sp., and T. frenchi atritarsalis, n. subsp.

The adults of 7. hilli, T. frenchi and T. littleri are redescribed and descriptions are given
ef the previously unknown immature stages of these species. é

On the basis of the structure of the larvae, 7. victoriensis, n. sp., is placed in the subgenus
Culicella. For T. hilli Edw. and JT. frenchi Edw., a new subgenus, Neotheobaldia, and for
T. littleri Taylor a new subgenus, Auwstrotheobaldia, are proposed.

An account is given of the biology of the Victorian species of Theobaldia.

INTRODUCTION.

Mosquitoes of the genus Theobaldia are widely distributed in the northern hemi-
sphere mainly in temperate regions; a few species extend into the tropics in Africa
and North America. In the southern hemisphere the genus is represented only by two
species in Africa, one in New Zealand and seven in Australia.

The six species hitherto known from Australia are: J. atra Lee, from Western
Australia, T. inconspicua Lee, from New South Wales and Victoria, 7. hilli Edw. and
T. frenchi Edw., from Victoria, 7. weindorferi Edw., from Tasmania, and 7. littleri
Taylor, from Tasmania and New South Wales.

The original descriptions of these species, except the first two, were inadequate in
many respects, so that accurate identification was difficult. Lee’s (1937) revision went
some way towards clarifying the situation, but he was hampered by having only a few
specimens for examination. With an abundance of material of all the Victorian species,
some collected in the field and some obtained by laboratory breeding, it has been
possible to make a full analysis of their distinctive features and to prepare descriptions
of the immature stages, which were previously unknown except in the case of
T. inconspicua. This paper also includes a description of a new species, T. victoriensis,
n. sp., and information on the biology and distribution of the genus in Victoria.

BIOLOGY.

The Australian species of Theobaldia are bush mosquitoes. They are present in
scrub along creeks, in the bush, and are especially numerous in upland forests, where
the humidity is higher and the temperature lower than in the open country. Their
inability to withstand high temperature and low humidity severely restricts their
distribution. Thus, while they are widely distributed in south-eastern Victoria, in
the north they are practically confined to hilly and mountainous country. T. inconspicua
is more adaptable than the other species and is found at lower altitudes and in more
exposed situations.

The three species, 7. hilli, T. frenchi and T. victoriensis, are man-biting mosquitoes
and become very troublesome in late summer and autumn. During the day they fiy
close to the ground and confine their attacks mainly to the legs. : After sunset they
rise higher, biting the arms but almost never the face. TJ. inconspicua and T. littleri
do not attack man.

This difference in feeding habits tends to give a false impression of the relative
abundance of the various species. Adults of 7. inconspicua are not common in the
field, but judging from larval numbers, it is by far the most abundant form. The
larvae are very numerous in rock and ground pools shaded by trees or grass.

66 THE GENUS THEOBALDIA IN VICTORIA,

In spite of a sustained search the larvae of the other species have rarely been seen
in the field. 7. littleri was found breeding in only one place. This was in Sherbrooke
Forest, in a pool under, and largely covered by, the roots of a fallen tree. The
temperature of the water, even in summer, did not rise above 13-14°C. VT. littleri and
T. inconspicua were breeding here during two successive years; on one occasion this
pool also yielded two larvae of 7. victoriensis. This species was also found at Ringwood,
where, in company with 7. inconspicua and T. hilli, it was breeding in a small, deep,
semi-permanent pool shaded by long grass and trees. TJ. victoriensis and T. hilli larvae
were found here only during the winter of 1952 and probably it is not a typical breeding
place for them. It seems likely that they breed, for the most part, in the underground
tunnels of the land crayfish (Hngaeus spp.); this may also apply to TJ. frenchi, the
larvae of which have not been seen in nature.

On the basis of colour and behaviour the larvae of these five species fall into two
groups. Those of 7. inconspicua and T. littleri are brownish and show the behaviour
typical of mosquito larvae, in that, when disturbed either mechanically or by the
passage of a shadow over the surface of the water, they move to the bottom of the
container. In the other species the larvae are milky-white. They show no response to
moving shadows and when disturbed either move just below the surface or, if originally
on the bottom, move towards the surface. Their lack of pigment and their behaviour
suggest that they normally live in underground water.

In 7. hilli mating occurs during the day. Normally the males do not swarm but
are seen, in small numbers, flying about in the grass close to the ground. Coupling
sometimes occurs with resting females but, more usually, is initiated while both sexes
are in flight and is completed on the grass. After sunset the number of “searching”
males increases but they do not become numerous. However, 7. hilli does occasionally
swarm. One such swarm was observed by Mr. A. Neboiss, at 4 p.m. on May 4th, 1953.
It consisted of 150-200 males flying 2-3 feet above the ground. Mating of the other
species has not been observed; presumably it occurs at night.

T. inconspicua is the only one of the five Victorian species which deposits egg
rafts. These have been collected in the field and can also be readily obtained in the
laboratory from engorged females. I have never found the eggs of the other species
in nature and for a long time failed to get oviposition in cages. Later work showed
that this was a result of maintaining the mosquitoes at too high a temperature, 20—22°C.
When engorged females were kept in an ice chest at 8-10°C., they oviposited regularly,
16-29 days after feeding. In the laboratory 7. hill, T. frenchi and T. victoriensis
deposited eggs singly on moist filter paper above water level. The eggs cannot with-
stand desiccation; when laid on the sides of the glass vessel, instead of on filter paper,
they invariably perished. After feeding on human blood the number of eggs varied
from 28 to 60; an engorged female of 7. frenchi caught in the field laid 90 eggs.

When eggs are kept in the laboratory at temperatures of 17—20°C. some eggs
remain viable for a long period without hatching. Thus in one experiment only
8 out of 60 eggs had hatched after 39 days. The unhatched eggs were then kept for
24 hours at 25°C. without any more hatching; but when they were transferred to a
refrigerator all the viable eggs hatched within two hours. In many cases hatching of
larvae from these diapausing eggs occurred when the temperature was lowered from
17°C. to 10-11°C., but with a further fall to 2°C., hatching ceased; it was completed
when the temperature was raised to 10-11°C. ;

In the laboratory, growth of the larvae is slow. They require clean water and at
temperature of 17-19°C. take about two months to reach the fourth stage. At higher
temperatures few survive.

T. inconspicua and T. frenchi maintain reproductive activity throughout the year;
the second species was recorded as biting in July (temperature 13:5°C.) and August
(temperature 8:5°C.).

T. littleri, T. hilli and T. victoriensis hibernate as larvae, mainly in the fourth
stage, and the spring generation of 7. hilli and T. victoriensis appears in October.

BY N. V. DOBROTWORSKY. 67

Genus THEOBALDIA Neveu-Lemaire.

Theobaldia Neveu-Lemaire, 1902, C.R. Soc. Biol., 54: 1331. Type, annulata Schr.
Culicella Felt, 1904, N.Y. State Mus. Bull., 79: 391c. Type, dyari Coq. Culiseta Felt,
1904, N.Y. State Mus. Bull., 79: 391c. Type, absorbrina Felt. Theobaldinella Blanchard,
1905, Les Mostiques, p. 390. Type, annulata Schr. Pseudotheobaldia Theobald, 1907,
Monog. Cul., 4: 271. Type, niveitaeniata Theo. Climacura Howard, Dyar and Knab,
1915, Mosq. N.C. Am., 3: 452. Type, melanura Coq. Allotheobaldia Brolemann, 1919,
Ann. Soc. Ent. France, 88: 90. Type, longiareolata Meg. Theomyia Edwards, 1930,
Bull. Ent. Res., 21: 303. Type, fraseri Edw.

Characters of the Genus.
(Adapted to Victorian species.)

Adult. The head has narrow, curved, upright, forked scales on the vertex and flat
scales laterally. The eyes are almost touching. The proboscis is moderately long.
The palpi of the male are about as long as the proboscis and more or less hairy. The
palpi of the female are always short, with a vestige of a fourth segment.

The thorax has strong acrostichal and dorsocentral bristles; the scales are always
narrow and curved. The scutellum has narrow scales and long border bristles. The
anterior pronotal lobes have a few strong bristles and several shorter ones, and in
some species a few narrow curved scales. The posterior pronotal lobes have bristles
and, in most species, narrow curved scales.

The spiracular bristles are fine and few in number; post-spiracular scales are
usually absent (a few minute ones are present in 7. littleri).

The sternopleura has on the lower part a few strong bristles, several shorter ones
and scales. The pre-alar has a patch of bristles. The upper posterior part of the
mesepimeron has a patch of erect hairs and, below this, a patch of flat scales or fine
hairs; there are two or three strong lower mesepimeral bristles, a few short ones, and
scales.

The legs are dark, with pale reflections apically in some species. The claws on the
anterior legs of the males are toothed; in the females all are simple. Pulvilli are absent.

The wings are unspotted. The venation is very uniform; the subcosta extends
beyond the end of the radius; the fork cell of the second longitudinal vein is longer
than its stem; the cross-veins are well separated. The outstanding scales of the wings
are narrow; on the base of the radius there are, on the upper surface, a few long
hairs, and on the lower surface a group of hairs and scales in front. The squama has
a long dense fringe, and the alula long narrow scales.

The abdomen is always unbanded, blunt tipped, and with a non-retractile eighth
segment. Hypopygium of male: the coxite is rather long; its basal lobe varies in
size and in the extent to which it is separated from the coxite; the tip of the lobe
bears a number of hairs and spines. The style is simple, long and slender, with a
terminal spine. The paraprocts have a few terminal teeth.

Pupa. This is similar to that of Aédes; the trumpet is always short, with a large
oblique opening. The distal margin of the paddle is smooth or very finely spiculate.

Larva. The siphon is rather long. All species, except 7. littleri, have the tuft,
which may consist of a single hair, at the base of the siphon. The comb consists of a
large patch of small teeth.

Subgeneric Division.

The various subgenera of Theobaldia, although clearly defined in the larval stage,
are often difficult or impossible to recognize in adult specimens. Thus Edwards (1923)
Suggested that the two Australian species then known belonged to the subgenus
Culicella, but pointed out that this placing was tentative pending an examination of
larval stages. Subsequently (1932) and still without access to larval material, he
placed all the species in the subgenus Climacura.

With the discovery of the larval stages the Victorian species can be definitely
assigned to their subgenera. As Lee (1937) showed, J. inconspicua belongs to the

68 THE GENUS THEOBALDIA IN VICTORIA,

subgenus Culicella; this is also the case with 7. victoriensis. T. hilli and T. frenchi,
on the one hand, and 7. littleri on the other, require new subgenera.*

Edwards (1932) defined the subgenus Culicella, on larval characters, as follows:
“Head large and broad. Antennae long and stout, with a large tuft well beyond middle;
two of the apical bristles very long and somewhat removed from the tip. Mouth
brushes very large. No air sacs in thorax. Siphon long and tapering, with one pair
of basal tufts; pecten consisting of spine-like teeth only, no fine hairs distally. Comb-
teeth in a large patch. Anal segment ringed by the plate (in fourth stage); several
tufts piercing the plate ventrally before the brush. Outer dorsal hair of anal segment
branched. Gills rather long, pointed.”

The larva of 7. victoriensis agrees with this description, but two of the apical
antennal bristles are not removed from the tip; the outer dorsal hair of the anal
segment is unbranched. Not more than one tuft pierces the anal plate before the brush.

NEOTHEOBALDIA, n. Subg.—Larva (Fig. 6): The head is not very large. The antennae
are of moderate length; the tuft is well beyond the middle; none of the apical bristles
are removed from the tip. The siphon is of moderate length, with one pair of basal
tufts; the pecten teeth are in the form of hairs. The anal segment is ringed by the
plate; no tufts pierce the plate before the brush. The outer dorsal hair of the anal
segment is unbranched. The anal papillae are not large.

This subgenus is erected for T. hilli and T. frenchi. It is close to Theobaldia, s. str..,
but is distinguished by the absence of a row of hairs following the pecten and by the
form of the pecten teeth.

AUSTROTHEOBALDIA, n. Subg.—Larva (Fig. 9): All the hairs are simple. The head is
very large and broad. The antennae are long; two of the apical bristles are removed
from the tip. The siphon is long and tapering; the siphonal tuft consists of a two-
branched hair, or a single one, about half-way along the siphon. The pecten consists
of triangular teeth. The comb has a long row of small scales and two or three
irregular short rows of long scales. The outer dorsal hair of the anal segment is
branched. The anal papillae, which are not large, are pointed.

This subgenus is erected for 7. littlerit. It differs from all other subgenera of
Theobaldia in that all the hairs are simple and the siphonal tuft is half-way along
the siphon.

The Victorian members of the genus Theobaldia can be distinguished from other
genera by the unbanded, blunt-ended abdomen, the absence of pulvilli and the presence
of spiracular hairs. Within the genus the males of the various species are easily
distinguished, particularly by their hypopygial characters, but the females of T. hilli
and T. frenchi are hardly distinguishable from one another and in fact cannot always
be separated with certainty. The larvae of these two species are also very similar.

Key to Victorian Species of Theobaldia.
Adult Males.

1. Proboscis pale beneath; last two segments of palpi, in living specimens, bent back-
2/21 60 kia are ae ee ee meme on iee ts tales, As) uN. |. 6G /Ginldcn alas oes Gumbel! 6 inconspicud.
2. Last segments of tarsi with pale reflections
Tarsi entirely dark

we lL

3. Upright scales on vertex pale, basal lobe of coxite long and well separated from
CHE DASE) eee hs nels hand ek eee Soe ees Cert 12. Ne Ae re ne ernen eis littleri.
Upright scales on vertex dark; basal lobe separated only towards the tip ...........

frenchi atritarsalis.

4. Base of tarsal segments with pale scales. Coxite with black scales and golden bristles
dorsally, andedensexeoldishwhainrsmaveniticallllyaammn eis aie nena nai neraie iene aaa victoriensis.
Proximal segments of tarsi dark scaled, distal segments pale

Oo

* A new Victorian species of Theobaldia has recently been found by Mr. G. Douglas. On
the structure of the larva, it belongs to the subgenus Climacura, which hitherto has been known
only from North America. i

BY N. V. DOBROTWORSKY. 69

Palpi slightly longer than proboscis, densely hairy apically. Coxite more than twice
as long as broad; basal lobe separated only towards the tip ................. frenchi.
Palpi slightly shorter than proboscis, with scanty and relatively short hairs. Coxite
only about twice as long as broad; basal lobe is large, well separated and reaching
me arlyaetOmth eaitipmotethenGCOmiter . i468 Sevag isle eo ksisasaele ao Sir shetetels ood seMeiele = 4: 4aud oi hilli.

O1

Per OOOSCISmD Alem DCM Delian. pts rte aerate cocoa sire sie) sal eure Deriel sviekis) sen ellecencns, vs scovelattorelc inconspicuda.

EOD OSCISMDIACK IDET Cathie ercia etl ara Peete rete doteeets eer cet cbarcte ee otale oc Abberley ake ts ee 2

PDE SIMeMtIne lyAcGanks falecvetsesth sess oF oi Gus A cin aL Mee Tee lehahn eleea eydensd ro ks eks JGR eekly el So eed 3
IMAIStESeLIN echt Ontarsio within palesnrenections ace aes - see's. - cee ce esses came 4

3. Vertex with pale scales ........ PN Ee heh eae den tsi EUR ee CTS. ci MMO es Fol led. littleri.
Wiprichtyscalesimonusviertexs=da rk garrwrw aes cocks cohesehe tect ae Beas elie « frenchi atritarsalis.

Ae SAsSerOt tarsal Sseeiments) with) palewscailes) Sai... .2.2. eae oo. a ee ce victoriensis.
IBASE Ol UARSEYL GESINeIOS jyAloaowhe TRV SERIES Gobdavonouebouuunod oad canpeecubudeouune 5

5. Upright scales on vertex mostly pale. Venter usually with more or less conspicuous
TNE CTAN MD AUCH ESO fm VAC Kets CALC Sie ores isis Ci cueho cis tale cistron a ase mes cheat cc os MBO cba ckie Dec ch oupeeene hilli.
Upright scales on vertex mostly dark. Venter pale scaled .................... frenchi.

Larvae (fourth stage).

il, JElibe iUbEe mene Tae, ToaMlolkolkes Gye hola Gag oocoouuvocodadecbuUdobuocomuahoe bo e.s littleri.
TSP HOBIE ENE LORIE) CONE SIO) NON at eh Bueno otorctalny co cseH ON neo GacNBuntcsa Aoi ie torah le ca EN) Eero i HERS RCE ORR eames ete 2

Pameectenmconsistines of spine=lkemteeths . eine a5 cts melee siemss hokey cee + clots oc cleitlela sche sere 3
WECECNECONSISES AO hel aTs OWe © ls NAMING S) geet “iA, Sirsviays = syish Saal eg Se i cite Tove ces Site ohne) aos) eee ues oh Mae us sb anece s 4

3. Larva milky-white in colour. Siphon tapering, anal papillae very long .... victoriensis.
Larva brown. Siphon slightly swollen in middle. Anal papillae short .. inconspicua.

4, SOOM ebhae Alnol ABRido) lovee waKel aS ho os oc A colo Go Gomme Od God ulo co oleclold ¢c.0 6 mao del hilli.
PipPWOnalen cunt Ofte Sime: wai aes howe os, od eareteueucyeneea osc euiseeleuel etemonsucmeelied sr clicice iene eases <1 frenchi.

DESCRIPTION OF SPECIES.
Subgenus NEOTHEOBALDIA, nh. subgen.
THEOBALDIA HILLI Edwards.

Edwards, F. W., 1926, Bull. Hnt. Res., 17: 111.

Type—The type series from Beaconsfield, Victoria (6.XI and §8.XII.1923), is in
the British Museum.

Distinctive Characters.—The Male. The upright scales on the vertex are pale’
becoming dark towards the neck. The proboscis and palpi are clothed with blackish
scales with violet reflections. The palpi are slightly shorter than the proboscis; the
apex of the shaft and the last two segments bear scanty and relatively short hairs;
the terminal segment is about half as long as the penultimate. The thorax is brown
with pale golden scales, whick are rather larger and lighter than those of T. frenchi.
The bristles are dark brown. The anterior pronotal lobe has three-four strong black
bristles, several shorter pale golden ones, and a few narrow curved scales. The
posterior pronotum bears narrow curved scales and four or five pale golden strong
proepimeral bristles. The scutellum has five-seven strong black border bristles on each
lobe. There are two pale spiracular bristles. The lower part of the sternopleura
bears a long strong bristle and, below this, one or two shorter ones, several short fine
ones and pale scales. The mesepimeron has a patch of pale scales and short bristles
towards the middle; there are two strong bristles (the upper is shorter) on the lower
part. The legs are black-scaled with conspicuous knee spots; the femora are pale
beneath; the ends of the tarsi have light brown or pale reflection. The wing length is
35-36 mm. The first to seventh tergites are black-scaled with violet reflection; the
eighth has black and pale scales. The colour of the venter is variable; usually the
first to fourth sternites are black-scaled; the fifth to seventh have pale scales apically
and these extend forward progressively so that the seventh sternite may have only
a median patch of black scales; the eighth is black-scaled. But in some specimens the
venter. is pale with more or less conspicuous median patches of black scales on all
sternites. The hypopygium (Fig. 1): The coxites are relatively short, being only
about twice as long as broad. The basal lobe is large, well separated, and reaches
nearly to the tip of the coxite; its tip has dense, long curved hairs. The aedeagus is
rather large and strongly chitinized with four strong teeth and four shorter ones; on

70 THE GENUS THEOBALDIA IN VICTORIA,

the distal end there are about thirteen fine hairs. The lobes of the ninth tergite are
prominent, with 9-10 curved hairs.

The females are similar to the males but differ as follows: The upright scales on
the vertex are usually darker than in the male; in some specimens they are almost
black. The last two segments of the tarsi are pale. Wing length is 3-5-3-8 mm. The
upper fork cell is four times as long as its stem. The tergites are black-scaled with

reflections; the fifth-seventh tergites have some pale scales on their apical
The venter is pale with, or without, median patches of black scales.

violet
corners.

SSESESS
SS %
~—

Figs. 1—4.—Male terminalia.
1, 7. hilli; 2, T. frenchi; 3, T. victoriensis; 4, T. littleri.

The Pupa. This is milky-white with black hairs. The trumpet is short, with a
large and very oblique opening, the margin of which is fringed. Seta A of the VIII
segment of abdomen has four to six branches. The paddle (Fig. 5, B) is broadly oval
with a smooth margin. At the tip of the midrib there are two setae, one very small
and single, the other about three times as long and consisting of three or four branches.

BY WN. V. DOBROTWORSKY. val

The Larva. The fourth stage larva (Fig. 6) is milky-white in colour with black
setae. The head is not very large; it is rounded, pale and almost transparent; the
antennae have the subapical hairs slightly removed from the tip; the tuft consists of

Figs. 5-7.

5, Paddle of pupa: A, 7. littleri; B, T. hilli;-C, T. victoriensis; D, T. inconspicua. 6, T. hilli
Edw. Head, terminal segments and mentum of larva. 7, Eggs of Theobaldia. A, B, T. incon-
spicua (two views); C, T. hilli; D, T. frenchi; E, T. victoriensis. a and b show the structure
of the exochorion.

non-plumose hairs. The anterior frontal setae nave three-five branches, the inner frontal
four-six, middle frontal three-four, and the outer frontal five-eight. The sutural seta
(e) is single, and the trans-sutural (f), which is one-third to one-half the length of

co |
bo

THE GENUS THEOBALDIA IN VICTORIA,

(ec), has two-three branches. Prothoracic chaetotaxy: The first, second, fourth, fifth
and sixth are single, the third has two-three branches, the seventh has two branches,
and the eighth is a single minute tuft with about six branches. Abdomen: The hairs
on segments I-VIII are short except for three lateral hairs on the first segment and a
single lateral hair on the following five. - The pentad hairs on the eighth segment:
a usually has three branches, rarely four, 8 is single, y has three plumose branches,
5 is single and e has two to three simple branches. There are about 90 comb teeth
forming a large triangular patch. The inner dorsal brush consists of nine to eleven
branches and a single long outer hair. The ventral brush has twelve to fourteen tufts.
The anal papillae are pointed and of about the same length as the saddle. The saddle
tuft is short with three or four branches. The siphonal index is 5:0—5-4. The siphon
has a tuft of two hairs at the base. The pecten consists of a row of seven to
eleven hairs.

The eggs (Fig. 7, c) are elongate-oval, silvery in colour, with a black base and a
short transparent stem. The egg is about 0-88 mm. long with an index of about 4:3.

Distribution.—A total of 50d and 8099 have been examined from the following
localities: Ringwood, Christmas Hills, Beaconsfield, Healesville, Head of Cement Creek,
Acheron River, Grace Burn Creek, Nyora, Tarra Valley Park, Stony Creek (Gippsland),
Franklin River (N. Dobrotworsky), Trafalgar South, Upper Pakenham, Thompson,
Tarago River, Dollar (near Meeniyan), Boolarra, Mirboo, Gembrook (A. Neboiss).
T. hil is confined to Victoria.

THEOBALDIA FRENCHI Theobald.
Culex frenchi Theobald, Mon. Cul., 2: 66 (1901).
Type.—The type series of four females from Victoria (locality not stated) are in
the British Museum.
The male was described by F. W. Edwards (1926) from Sassafras, Victoria.

Distinctive Characters—The Male. Most of the upright scales on the vertex are
dark. The palpi are slightly longer than the proboscis. Apically the shaft and the
last two segments have dense, long hairs. The last segment is about half as long as
the penultimate. The palpi and the proboscis are clothed with dark brown scales.
The thorax is reddish and is clothed with pale golden scales; the bristles are black.
The anterior pronotal lobes have two or three black bristles, some pale golden ones
and a few narrow pale scales. The posterior pronotal lobes have a few narrow pale
scales and four-five dark proepimeral bristles. There are one or two spiracular bristles.
There are six-seven border bristles on each lobe of the scutellum. The sternopleura has,
on the lower part, a patch of scales, a few pale hairs, one dark strong bristle and a
second shorter one below. There are two lower mesepimeral bristles; the middle part
of the mesepimeron has a patch of scales. The legs are clothed with dark brown scales;
the tarsi are pale apically.

The abdomen is clothed dorsally with black scales, except the last segment, which
has pale golden scales; the venter is clothed with pale, or black and pale scales.
Hypopygium (Fig. 2*): Each lobe of the ninth tergite has a patch of nine-fifteen
curved hairs. The coxite is more than twice as long as broad; the basal lobe is
about two-thirds the length of the coxite; it is separated only towards the tip, which
bears a tuft of rather short hairs. The aedeagus is rather large and strongly chitinized,
with tour large and three small teeth.

The female is similar to the male, but differs as follows: Yhe anterior pronotal
lobes have four-five strong black bristles. Hach lobe of the scutellum has about eight

*TIt will be noted that this figure is very similar to that given by Lee (1937, p. 296,
Text-fig. 3) for T. weindorferi. Through the courtesy of the Division of Entomology, C.S.I.R.0O.,
{ have been able to examine the type of JT. weindorferi; the mounted terminalia correspond
exactly with Lee’s Text-figure 4, labelled 7. frenchi. There can be no doubt that in Lee’s paper,
Text-figures 3 and 4 were transposed. Text-fig. 3 should refer to T. frenchi and Text-fig. 4 to
4d’. weindorfeni.

BY N. V. DOBROTWORSKY. 73

border bristles. The patch cf scales on the pleurae is more distinct. The legs are
darker, almost black above; the femora and tibia are pale beneath. There are
conspicuous knee spots of pale scales; the last three segments of the tarsi are pale.
The venter is pale. Wing length is 3-8-5-0 mm. The upper fork cell is 3-0—4-5 times
the length of the stem.

The Pupa. This is very similar to the pupa of 7. hilli.

The Larva. The fourth stage larva is similar to that of 7. hilli but can be dis-
tinguished from it as follows: The frontal setae are more branched, the pentad hair y
on the eighth abdominal segment usually has 4—5 branches, the basal siphonal tuft is
represented by a single hair. (In 7. hilli the basal tuft always has two branches.)

The eggs (Fig. 7, D) are similar to those of 7. hilli; they are about 0-7 mm. long,
with an index of 3:4—3-5.

Distribution—-A total of 15¢¢ and 12099 were examined from the following
localities: Sherbrooke, Kalista, Grace Burn Creek, Healesville, Head of Cement Creek,
Tarra Valley Park (N. Dobrotworsky), Menzies Creek, Tanjil South (A. Neboiss). As
far as is known, TJ. frenchi is confined to Victoria.

THEOBALDIA FRENCHI ATRITARSALIS, n. subsp.

This subspecies is clearly distinguished from the type by its general darker colour;
the thorax is brown, the proboscis, the palps and the legs are clothed with almost
black seales; the legs also are dark apically. The male palpi are even more hairy
than those of the type, and the shaft apically has about thirty long hairs. The male
genitalia are identical with that of the type.

The pupa, larva and eggs are identical with those of the type.

Types.—The holotype male, allotype female and a paratype series from Stony Creek
(Gippsland) are in the collection of the National Museum, Melbourne.

Distribution.—A total of 5¢¢ and 4499 have been examined from the following
localities: Stony Creek (Gippsland), Franklin River, Kalimna, Tarra Valley Park
(N. Dobrotworsky), Boolarra, Mirboo North, Thompson, Hiawatha, Tyres River, Gould
(A. Neboiss).

Subgenus CULICELLA Felt.
THEOBALDIA VICTORIENSIS, D. Sp.

Types.—The holotype male and allotype female were bred from larvae collected at
Ringwood, Victoria, 3.8.52. These, together with a paratype series of 10¢¢ (three from
Ringwood 3.8.52 with associated larval and pupal skins, five from Tarra Valley Park
5.5.53, two from Boolarra 8.5.53) and 1099 (six from Ringwood 1.1.52, 7.1.53, 1.3.53 and
6.4.53, two with associated larval and pupal skins, two from Tarra Valley Park 5.3.53,
two from Healesville 30.1.53), are in the collection of the National Museum, Melbourne.

Description of Adult—Holotype ¢. The head scales are pale; a few upright ones
become dark toward the neck. The palpi are as long as the proboscis. The shaft bears
about 10 long hairs; the last two segments have long but relatively sparse hairs; the
terminal segment is about half as long as the penultimate. The proboscis and the
palpi are clothed with black scales with violet reflections. The thorax is brown, with
pale golden scales; the small bristles are pale, the strong ones are pale at the base
and dark at the end. The anterior pronotal lobe has pale golden bristles and a few
narrow curved scales; the posterior pronotum has a number of curved scales and five
pale golden proepimeral bristles. There are two fine pale spiracular bristles. The
scutellum has narrow curved pale scales, with four-five border bristles on the lateral
lobe and eight on the central. On the lower part of the sternopleura there are fine
hairs, a few narrow scales, one strong bristle and several shorter ones. The mesepimeron
has a number of fine hairs towards the middle and two strong lower mesepimeral
bristles (upper is shorter). The legs are dark brown with inconspicuous knee spots.
The tarsi: the second to fourth segments of the forelegs and the first to third of the
mid- and hindlegs are pale at the base; the last two segments of the tarsi are covered
with pale scales. The wing length is 4 mm. The abdomen is unbanded. The tergites
are clothed with black scales with violet reflections and pale hairs. The apical border

G

74 THE GENUS THEOBALDIA IN VICTORIA,

of the eighth tergite has pale scales. The venter is also black-scaled but the fourth-
seventh sternites have pale scales apically. The hypopygium (Fig. 3): The coxites have
long pale goldish hairs, black scales dorsally, and dense pale goldish hairs ventrally
on the inner face.

Paratypes %. The series of 10 paratype males differs from the type in colouring.
The venter in some specimens is pale with large median patches of black scales. Each
lobe of the ninth tergite has a patch of ten-twelve curved hairs. The coxites are more:
than twice as long as broad. The basal lobes of the coxites are hairy; they are about
two-thirds of the length of the coxite and are separated from them only at the tip,
which bears a tuft of long hairs. Wing 4:0—4:-1 mm.

Allotype 9. This differs from the holotype as follows: The forked upright scales
on the head are pale only in front and gradually become black towards the neck. The
strong bristles on the thorax are black. The anterior pronotal lobes have three strong
black bristles, several golden ones and a few narrow scales. There are three pale
spiracular bristles. The sternopleura has one strong bristle and several shorter ones.
The mesepimeron has a patch of hairs towards the middle and one strong and three
short lower mesepimeral bristles. The legs are almost black, with violet reflections.
There are conspicuous knee spots. The first three segments of the tarsi have pale
basal rings; the fourth and fifth are pale; the third segment of the hind-tarsi is also
pale apically. Wing: The upper fork-cell is 3-8 times as long as its stem; wing length
is 4.7 mm. The venter is pale-scaled with inconspicuous median patches of black scales.

Paratype 9°. The series of 10 paratype females do not show much variation; the
upright scales on the head are predominantly pale in some specimens and become dark
only towards the neck. The mesepimeron has a patch of hair, a few narrow scales
and two-three strong lower mesepimeral bristles. The length of wing is 3-8—5:1 mm.;
the upper fork cell is three-four times as long as its stem. The median patches of
black scales on the venter may be large.

The Pupa. The trumpet is similar to that of JT. hilli. Seta A of VIII abdominal
segment has three or four plumose branches. The paddle (Fig. 5, C) is broadly oval
with a smooth margin and a few minute denticles slightly removed from the margin.
At the tip of the midrib there are two setae; one is small and single, the other is twice
as long and usually two-branched.

The Larva. The fourth-stage larva (Fig. 8) is milky-white with black setae. The
head is large, oval, pale, almost transparent and with very small eyes. The antennae are
long, thin and curved, with a tuft of fourteen-sixteen slightly plumose hairs. The
anterior frontal setae are single or double, the inner frontal consist of four to five
hairs, the middle frontal of two, and the outer frontal of six-seven. All these except
the anterior frontal are plumose. The sutural seta (e) has two branches and the
trans-sutural (f), which is half as long, has two-three branches. Prothoracic chaetotaxy:
The third seta is single or two-branched, the fourth is short and has two or three
branches, the fifth and sixth are single, the seventh has three branches, and the eighth
is minute with about four branches. Abdomen: The hairs on segments I-VIII are
very short, except three lateral ones on the first segment and a single lateral hair on
segments II-VI. The pentad hairs on segment VIII: a has four-five slightly plumose
branches, 8 is single, y has two-three plumose branches, 6 is single, and e has two
branches. The comb teeth form a large patch. The inner dorsal brush consists of
seven-eight tufted hairs and a single long outer hair. The ventral brush has about
fifteen tufts. The anal papillae are slender, pointed and twice the length of the saddle.
The saddle tuft, which has two-five branches, is small. The siphon is long, slender,
pale at base but gradually becoming brown apically. The siphonal tuft is at the base
and has two branches. The siphon index is 6-3-6-8. The pecten consists of ten to
thirteen spine-like teeth.

The eggs (Fig. 7, HE) are similar to those of 7. hilli, but have more pointed ends:
the index is 2-:8—3:2, the length 0-8 mm.

BY N. V. DOBROTWORSKY. 75

Distribution.—Besides the type series, a total of 10¢¢ and 7099 have been examined
from the following localities: Ringwood, Christmas Hills, Healesville, Grace Burn
Creek, Head of Cement Creek, Meeniyan, Tarra Valley Park, Franklin River, Nyora,
Stony Creek (Gippsland), (N. Dobrotworsky), Boolarra, Thompson, Upper Pakenham,
Menzies Creek, Kinglake, Dollar (near Meeniyan) (A. Neboiss).

THEOBALDIA INCONSPICUA Lee.

Lee, D. J., 1937. Proc. Linn. Soc. N.S.W., lxii: 294-298.

Type.—The type series trom Tinderry and Mittagong. Holotype and allotype are
in the Museum of the Commonwealth Scientific and Industrial Research Organization,
Canberra.

As the adults and larva were carefully described by Lee (1937) only a brief
description is given here.

Adult. The upright scales on the vertex are dark brown, some in front are pale.
The proboscis is dark brown above and pale beneath. The male palpi are as long as
the proboscis. The penultimate segment has scanty and relatively short hairs; the last
has only a few long hairs on the inner side at the basal part. The last two segments
of the palpi, in living specimens, are bent backwards. The anterior and posterior
pronotal lobes are devoid of scales. The tarsi are entirely dark. Ventrally the abdomen
is clothed with dark scales basally becoming pale goldish apically. The male hypopygium:
The coxites taper and are more than twice as long as broad; the basal lobes are very
small and imperfectly separated from the coxites. The aedeagus has three large
teeth.

The pupa is brown; the trumpet is short with a very large and oblique opening.
Seta A of the eighth abdominal segment has two or three branches. The paddle
(Fig. 5, D) is oval, with minute denticles on its surface and larger ones at the margin.
At the tip of the midrib there are two setae; one is minute and single, the other is:
single or two-branched.

The Larva. The fourth-stage larva is brown. The head is large; the antennae are
long. The anterior and middle frontal setae are single, the inner frontal seta consists
of two branches. Prothoracic chaetotaxy: The first, second, fifth, sixth and eighth setae
are single; the third, fourth and seventh setae each consists of two branches. The
hairs on the abdomen are well developed. The eighth abdominal segment: The comb
teeth form a large patch; the siphon is long, slightly swollen in the middle, with an
index 5:0-5-5. There is a single hair at the base of the siphon. The pecten conists of
9-10 serrate teeth.

The eggs (Fig. 7, A, B) are deposited in rafts, which are rounded or oval in
shape with raised margins, and look like a basket. The egg rafts, which contain up
to 217 eggs, exceed 4 mm. x 2 mm. The eggs are black, thick at one end and tapering
to the other; the length of egg is 0:9-1:0 mm., with index of about 4-0.

Distribution.—A total of 30¢¢ and 4092 have been examined from the following
localities: Ringwood, Melbourne, Kalista, Sherbrooke, Healesville, Tarra Valley Park,
Kalimna, Violet Town (N. Dobrotworsky), Shoreham (F. Drummond), Menzies Creek
(A. Neboiss). The species also occurs in New South Wales (Lee, 1937) and Tasmania
(Sulfur Creek, K. W. Dillon).

Subgenus AUSTROTHEOBALDIA, nh. subg.
THEOBALDIA LITTLERI Taylor.

Chrysoconops littleri Taylor, 1913-1914, Trans. Ent. Soc. Lond., p. 702. Theobaldia
littleri Taylor, Lee, 1937, Proc. Linn. Soc. N.S.W., lxii: 295.

Type.—The type female, from Mt. Arthur near Launceston, is in the School of
Tropical Medicine, Sydney. The presumed males, described by Lee, one from Barrington
Tops, N.S.W., another from National Park, N.S.W., are in the Macleay Museum.

Distinctive Characters.—Adult. The male. The upright scales on the head are
pale. The palpi are slightly shorter than the proboscis; the shaft has six to eight long:
apical hairs; the last segment, which is about twice as long as the penultimate, bears:

6 THE GENUS THEOBALDIA IN VICTORIA,

dense long hairs. The palps and the proboscis are clothed with brown scales. The
thorax is brown and bears pale golden scales; the strong bristles are black, the short
ones, pale golden. The anterior pronotal lobe has three to four strong black bristles,
several pale shorter ones and narrow curved scales. The posterior pronotum has
narrow curved scales, and five-six pale golden proepimeral bristles. The scutellum has
four-six border bristles on the lateral lobe and six on the central. There are three
very fine spiracular bristles. The postspiracular area has four-five minute scales. The
sternopleura has one strong bristle, several shorter ones and flat pale ‘scales. The
mesepimeron has a patch of flat scales towards the middle, and two strong and a few
short lower mesepimeral bristles. The legs are black without knee spots; the femora
are pale beneath. The tarsi are entirely dark brown. Wing length is 3-8—4-:0 mm.

The wing veins are covered with brown scales. The underside of the subcostal
vein has hairs and some narrow scales proximal to the humeral cross-vein. The upper
corner of the alula has four-five scales. The abdomen: i1st-7th tergites are black
with violet reflections and pale bristles; the VIIIth tergite is clothed with a mixture of
black and light brown scales. The venter is light brown, with or without lateral
triangular areas of brown and black scales; the VIIIth sternite is entirely black-scaled.
The hypopygium (Fig. 4): The coxites are almost three times as long as broad. The
basal lobes are about two-thirds of the length of the coxite and are separated for
nearly their whole length; they bear rather strong spines. The lobes of the ninth
tergite are not prominent; each bears four-seven bristles. The aedeagus is rather large
and strongly chitinized, with three strong teeth and, under them, a few fine bristles.

The female differs from the male as follows: The anterior pronotal lobe has about
six strong black bristles. There are three strong black proepimeral bristles and a
couple of shorter ones. The scutellum has six black bristles on the lateral lobe and
seven on the central one. The postspiracular area has seven to eight minute scales.
The sternopleura has one strong bristle and a shorter one. Wing length is 3-8-4:5 mm.
The upper fork cell is 3-2—4:0 times as long as its ‘stem. The upper corner of the alula
has about eight scales. The fifth-seventh tergites have some pale brownish scales
laterally. The venter is pale with light brown reflections.

The Pupa. ‘The trumpet is short, with a large and very oblique opening, the margin
of which is fringed. Seta A of the eighth abdominal sternite is single. The paddle
(Fig. 5, A) is oval with minute denticles on its margin; at the tip of the midrib there
are two single setae.

The Larva. The fourth-stage larva (Fig. 9) is reddish-brown; all the hairs are
non-plumose. The head is broad and pale; the antennae are long, thin and curved, with
a tuft of about seven hairs. The anterior frontal setae are single, the inner frontal
have two or three long branches, the mid-frontal are single and longer than the inner
frontal, and the outer frontal has two branches. The sutural and the trans-sutural
setae (e and f) have two-three branches. The mental plate has a central tooth and
about ten lateral teeth. The thorax: Prothoracic chaetotaxy: the third, fourth and
seventh have two branches; the remaining setae are single. The abdomen: The hairs
on segments I-VII are well developed; the lateral hairs consist of two or three branches.
The pentad hairs on the eighth segment are long: a has eight or nine branches, 8 is
single, y has three to five branches, 6 is single, and e has three branches. The lateral
comb consists of sixteen-thirty small scales in a basal row, and eight-fifteen long scales
forming a patch. The inner dorsal brush has five to six branches and one long and one
or two short outer hairs. The ventral brush has ten or eleven tufts. The anal papillae
are narrow and usually equal in length to the saddle. The saddle hair is single and
as long as the saddle. The siphon is long and slender, with an index of about seven.
The hair tuft consists of one or two hairs about half the way along the siphon; there
is no basal hair tuft. The pecten, which does not reach the hair tuft, consists of
13-14 small triangular teeth.

The First-stage Larva. The position of the siphonal hair tuft (a single hair) is
variable; in some specimens it is near the middle of the siphon, as in the fourth-stage
larva, but sometimes it arises closer to the base.

BY N. V. DOBROTWORSKY.

KN

aN
SSS
\

2 aS

Figs. 8-9.

8, T. victoriensis, n. sp. Head, terminal segments and mentum of larva.
Head, terminal segments and mentum of larva.

wi
<I
\ A Wh ffi {} hd
‘ i ai mii | Me J
ZE
ZE- Se

77

9, T. littleri Tayl.

78 THE GENUS THEOBALDIA IN VICTORIA,

In several species of Theobaldia, e.g. T. inconspicua, T. victoriensis and European
forms, the siphonal tuft of the first instar larva lies some distance from the base,
alongside the pecten. In the second and later instars it shifts to the typical basal
position. In 7. littleri the shift in position is in the opposite direction.

Distribution—In Victoria 7. littleri is known only from Sherbrooke Forest. It
has also been recorded from Tasmania (Taylor, 1913) and New South Wales (Lee, 1937).

Acknowledgements.

For asssistance in the preparation of this paper the writer is grateful to Dr. F. H.
Drummond, Zoology Department of the Melbourne University; to Dr. S. J. Paramonov,
C.S.I.R.O., Canberra, for lending the type series of J. weindorferi; to Mr. P. F.
Mattingly, British Museum (Nat. Hist.), for comparison of 7. frenchi with the type
series; to G. Douglas, Lands Department, Melbourne, for his assistance in collecting
mosquitoes in Gippsland; to Mr. A. Neboiss, Lands Department, Melbourne, for the
presentation of mosquitoes of this genus collected in many parts of Victoria.

References.

Dyar, H. G., 1928.—The Mosquitoes of the Americas, pp. 241-250.

EHpWARDS, F. W., 1921.—A Revision of the Mosquitos of the Palaearctic Region. Bull. Hit.
Research, 12: 2638-351. :

————, 1924.—A Synopsis of the Adult Mosquitos of the Australasian Region. Bull. Ent.
Res., xiv: 351-401.
——, 1925.—Mosquito Notes. Bull. Ent. Res., xvii: 101-131.

———, 1932.—Genera Insectorum, fase 194.

—, 1941.—Mosquitoes of the Ethiopian Region, iii.

IKENT, N. E., 1953.—Mosquito Survey in the Melbourne Area. Vict. Nat., 70:117-21.

Lre, D. J., 1937.—Notes on Australian Mosquitoes (Dip. Cul.). IV. The Genus Theobaldia, with
description of a New Species. Proc. Linn. Soc. N.S.W., Ixii: 294-98.

, 1944.—Notes on Australian Mosquitoes (Dip. Cul.). V. The Genus Armigeres and

New Species of Armigeres, Theobaldia and Culex. Proc. LINN. Soc. N.S.W., lxix: 215-25.

TAYLOR, F. H., 1914.—The Culicidae of Australia. I. Trans. Ent. Soc. London, pp. 683-708.

THEOBALD, F. V., 1901.—Monograph of the Culicidae. II.

ag

THH INHERITANCE OF INFLORESCENCE CHARACTERS IN HUCALYPTUS.
By L. D. Pryor.
(Kight Text-figures. )
[Read 30th June, 1954.]

Synopsis.

The inflorescence of Hucalyptus is considered to be a dichasium contracted to give the
outward appearance of an umbel. Inflorescences of either 3 or 7 flowers are very common in
the genus. The 7-flowered unit is found to be partially dominant to the 3-flowered cluster.
Irregular combinations between 3 and 7 flowers sometimes occur in the Fl and often in
segregates in later generations.

To understand the inheritance in hybrid combination of inflorescence characters in
Hucalyptus it is necessary first to know the normal features of this structure in the
genus. A review ot the literature shows there is a good deal of confusion. This has
persisted probably for three main reasons: firstly, because the method of ordinary
herbarium preservation generally results in the blurring or loss of the characteristic
arrangement by which the basic inflorescence pattern might be determined; secondly,
special field collection and observation designed to reveal these characters is necessary
to secure material which can be interpreted easily; and, thirdly, from the point of
view of descriptive botanical work aimed mainly at systematic ends, it is of minor
value in diagnosis to separate, in the description, an umbel from an “umbel-like’”’ cyme,
or a three-flowered umbel from a three-flowered cyme.

Review of Literature Relating tc Eucalyptus.

Bentham (1863) simply regarded the flowers of Hucalyptus as “in umbels or heads
. rarely reduced to a single sessile flower’, and Mueller (1882) in “EKucalyptographia”
briefly described the flowers as occurring “in single or paniculated umbels .. .”.

Maiden (1924, I, 9) quotes Tate as saying, “the usual form of inflorescence is an
umbel which by lengthening of the axis passes to a panicle or corymb’. At the same
time Maiden (ibid.) quotes Naudin, in particular with regard to a key for a group of
some 50-60 species raised in France, in which he makes two contrasting groups, one
having an inflorescence of “cymes or umbels’, the other having panicles or corymbs;
and later (VI, 446), Naudin’s work is again quoted, in which in various groups of
species he refers to the inflorescence as “‘cymes or three-flowered umbels” as opposed
to “axillary umbels with three to eleven flowers”.

The determination of the inflorescence as a cyme or an umbel is hampered in
Hucalyptus by the fact that the bracts and bracteoles, if they do occur, have been found
in few species and are very early deciduous. They are, however, present in at least
some Corymbosae, and Maiden (ibid., VI, 467) found that with #H. ficifolia* ‘“‘The bracts
and bracteoles alternate, and they show six segments ... each umbel shows seven
buds, and it may be that one bracteole has aborted or it was so early deciduous that it
was not seen’. Maiden makes it clear, however, that he considers the matter incon-
clusive and that later study may elucidate matters with regard to the inflorescence. He
says: “Doubtless botanists will give special attention to bracts and bracteoles in the
future and may be able to offer useful generalization.”

Blakely (1934) simply says that the flowers are ‘in umbels or heads, usually
pedunculate, rarely reduced to a single sessile flower .. .”.

Naudin’s view has been renewed and developed, however, by Blake (1953) recently,
in which he first comments on the inflorescence by saying that “the primitive unit of

* Nomenclature as in Blakely, ‘““A Key to the Hucalypts’”’.

80 INHERITANCE OF INFLORESCENCE CHARACTERS IN EUCALYPTUS,

inflorescence appears to be a three-flowered cyme pedunculate in the axils of the leaves
of an ordinary vegetative twig. The cyme may be reduced to one or two flowers, but in
most species it has been altered to an umbel of few or many flowers. In many species
paniculate inflorescences have resulted from the shortening of the internodes of the
twigs accompanied by reduction and finally the loss of the leaves at least in the upper
part.”

It will be noted in passing that this suggestion as to the origin of the paniculate
inflorescence is the reverse of that suggested by Tate, recorded above. Blake goes on
to say: “The primitive three-flowered cyme persists (or sometimes occurs as a result
of secondary reduction?) as a regular feature of many quite dissimilar species .. .
and is occasionally met with in many others; reduced cymes are usual in such unrelated
species as #H. globulus and EH. macrocarpa ....’ Blake’s remark in parentheses is
apparently the first suggestion that the three-flowered inflorescence of Hucalyptus is
perhaps a reduced cyme.

Blake makes one other relevant note of interest in commenting on the inflorescence
in Myrtaceae from a phylogenetic point of view. He points out that the characteristic
inflorescence for two groups within the Myrtaceae, namely, the Myrteae and Metro-
siderinae, is a more or less terminal, dichotomous (or trichotomous) cyme, particularly
in the Australian-Malaysian region.

From the above it will be seen that there is considerable confusion of ideas as to
how the inflorescence of Hucalyptus should be regarded.

The Inflorescence of Dicots.

It seems that study of the inflorescence has been impeded in various ways since
the time of Linnaeus to the extent that the understanding of its structure has, in many
cases, come much later than one might expect from the general development of botanical
knowledge.

A recent paper by Rickett (1944) on the classification of inflorescences gives a
thorough review of the position. He points out that views with regard to the nature
of the inflorescence have varied considerably since the time of Linnaeus, and have
been obscured at different times, particularly by philosophical treatment, such as
resulted from Goethe’s views. Amongst other things, this led to the use of the
character of centrifugal and centripetal order of blooming as a basis in classification,
whereas it has become clear that this is a matter of physiology rather than structure,
and must be discarded as a basis of classification and understanding of inflorescence
structure.

When progressive study of the inflorescence began to take place, this was finally
summarized by Hichler, since when there has been further steady progress.

Rickett draws attention to the fact that, “In dealing with complex clusters it is
noticeable how often one encounters a dichasial group of flowers as a sort of unit
cluster, the entire inflorescence being composed of many such units variously related”’.

Rickett suggests that a dichasium be supposed as the simplest and most primitive
form of inflorescence. “By dichasium”’, he says, “I use the word in its strict sense to
mean a cluster formed by an apparent dichotomy beneath a terminal flower—in its
simplest form consists of three flowers.” Rickett further emphasizes that he does not
imply that the simple dichasium is necessarily earlier in evolutionary history than
the more complicated dichasium. The first inflorescence may already have been a much-
branched system. The transition is equally easy in either direction. “From a primitive
dichasium may be derived also without extensive changes, bostryches, cincinnae and
the rest, and by shortening of the internodes, many of the ‘umbellate’ and ‘capitate’
clusters characteristic of Liliaceous and Cornaceous genera.”

He says, further, that the dichasium might well be the basic inflorescence pattern
of a very large number of dicots. As will be shown later, this suggestion has particular
application in considering Hucalyptus. From an original dichasium he postulates the

BY L. D. PRYOR. rel

steps in evolution of an apparently simple inflorescence from the primitive dichasial
type to be perhaps as follows: (a) the limitation of an original dichasium to a few
flowers; (0) grouping of branches bearing leaves and terminated by dichasia on a
common axis—a grouping which involves the shortening of the branches and of the
internodes between them; (c) reduction of leaves to bracts.

After discussing the way in which many “umbels” may arise from the dichasium,
he says that “Since our terminology is to be used by systematists who cannot always
postpone their conclusions for microscopic investigation, it would be wise to use the
term ‘umbel’ for all clusters whose branches arise from a common point’. But he
adds in addition that where the structure of an ‘‘umbel” is understood, it might further
be designated as sympodial or monopodial.

Of the many studies that have been made of inflorescences in different groups,
it is interesting to note that of Woodson (1935), who postulated the dichasium as the
starting point for infiorescences of the Apocynaceae, from which he was able to
demonstrate how all the types of inflorescences found in that family might be derived.

The Basic Characters of the Eucalyptus Inflorescence.

From field observation of suitable material, particularly flower clusters soon after
they have emerged from the two covering bracts, it is clear that the inflorescence of
Hucalyptus is considerably more regular than has been thought. There are good reasons
for supposing that while it has in many cases the outward appearance of an umbel,
it is in fact a dichasium, in which the axes below the position of the bracteoles are
extremely contracted or eliminated so that the true form is not at once apparent. It
may well be postulated that there is a basic branching pattern in Hucalyptus perpetuated
by buds which are decussate and opposite. This system, which may never have existed
in the neutral state, is expressed in one form to give a leaf-bearing system, and in
another to give a flower-bearing system.

Jacobs (n.d.) has shown how this is the basic pattern in the leaf-bearing system,
where the leaves of Hucalyptus, while appearing alternate and two-ranked, are part of
a modified decussate and opposite system of branching. In a parallel but alternative
way it seems that the basic branching pattern has been modified to suit the flower-
bearing structure so that a flower cluster with the form of an umbel has been derived
from the basic branching which is decussate and opposite.

From the phylogenetic point of view it is interesting to note that a dichasial
inflorescence at once fits the pattern of the Australian members of the groups Myrteae
and Metrosiderinae, and makes Hucalyptus homologous with many members of these
groups in this respect.

Angophora affords an interesting comparison. Bentham describes Angophora as
having “flowers in umbel-like cymes arranged in terminal corymbs’. . The illustrations
of Angophora intermedia (Fig. 1, a, 6) make this quite plain, and the briefly persistent
bracts and bracteoles clearly indicate that the inflorescence is a dichasium.

If it is supposed that the Hucalyptus flower clusters are derived phylogenetically by
reduction from a preceding inflorescence which was an indefinitely prolonged dichasium,
the number of flowers in each inflorescence at different degrees of reduction, assuming
it is completely regularly branched, could be 1, 3, 15, 31, 63, 127, and so on, as shown
in Figure 2, c.

If it is assumed that in some circumstances, however, reduced branching occurs at
some stage (presumably due to crowding following contraction) by the elimination of
one side of the dichotomous branching due to suppression of both buds facing each
other from separate parts of the ramification across the now confined space, the
numbers of flowers in the flower cluster could be 1, 3, 7, 11, 15, 23, 35, ete., arranged in
the pattern as shown in Figure 2, a (the Xs show the suppressed buds).

As a further possibility, if, where, due to crowding in the same way, there is
space for which two adjoining buds are competing, instead of both being suppressed
as is supposed in the previous case, one has developed to fill the available space and

$2 INILERITANCE OF INFLORESCENCE CHARACTERS IN EUCALYPTUS,
the other remained rudimentary, a pattern of numbers could be 1, 3, 7, 13, 21, 31, and
so on, and the arrangement as shown in Figure 2, Db.

It is of interest, then, to examine the inflorescences of various species to see how
they conform with the numbers possible under various kinds of dichasial branching
if unmodified or, alternatively, if modified in the two simple ways set out above. It
will be noticed that the numbers 15 and 31 can be derived in two ways. The likely
numbers which could therefore result from a regular branching are 1, 3, 7, 11, 15 and
31, with a somewhat lesser frequency of 21 and 23.

Fig. 1.—a, b, Showing two variations of the dichasium of Angophora intermedia. It will
be noted that unlike Hucalyptus the axis below the bracts persists in some degree. c, Showing
the bracteole scars; A, B, C, corresponding with successive branches of the dichasium.
d, e, H. ficifolia: d, The typical seven-flowered dichasium showing the branching order A, B, C;
e, Showing the bracteole scars which correspond with dichasial branching, A, B, C. ff, EH.
Macarthuri, a typical seven-flowered dichasium. The branching order is shown by the letters
IN t@ Cr

Fig. 3.—a, Eucalyptus macrorrhyncha, a species which often has eleven-flowered inflores-
cences ; branching order is indicated from A to D. b,c, A segregate from H. elaeophora (seven-
flowered), H. viminalis (three-flowered). Note in b the three-flowered cyme produced below
the irregular four-flowered cyme in the same axil. d, E. pauciflora showing a typical fifteen-
flowered dichasium with the order of branching indicated by the letters A to E.

It must be remembered that, after emergence, the buds are very easily lost from
the inflorescence from failure to grow or by accident, and therefore the material must
be carefully examined. If branching is regular, it is likely that a regular geometric
pattern can be seen in the flower cluster if examined at a suitable stage. There seems
no reason to assume that regular branching will occur indefinitely. One branch of a

BY L. D. PRYOR. 83

dichasial cyme could, at least‘ theoretically, develop irregularly at any stage, and
therefore any number of buds might be produced without supervening accident.

In addition to the actual number of buds and their geometric pattern, further
evidence of the dichasial nature of the flower cluster can be derived by studying the
orientation of the inflorescence. Ii any secondary twisting is eliminated, the plane
of the central bud and the first pair of lateral buds in the dichasium must be at right
angles to that of the axis and the subtending leat or bract.

A further point which would aid in the assessment of the inflorescence as a
dichasium is displayed if bracts or bracteoles are present, because the disposition of
these then must be definite.

If Hucalyptus inflorescences are examined with the above in mind, it is noted that
a few species regularly have solitary flowers on peduncles, particularly #. tetraptera,
HE. globulus, E. macrocarpa, and that this occurs occasionally in a good many other
species, aS an aberrant development. The single flower might be regarded as the
ultimate stage of reduction from an indefinite dichasial cyme, which would be preceded

DMA@) G80
2
a 2 @-O-@ O-O-@
| : : @2'@ @1e@
3) @ = Tease Re aH
COO G00 eecleee
2 8 6 @

Fig. 2.—Diagrammatice patterns of dichasial branching. a, Branching limited where
-opposing primordia have space between them for only one bud, and it is assumed that both
are suppressed. 6, Where there is space for only one bud between opposing primordia but
where it is assumed this is filled by one bud while the other remains suppressed. c, Where all
buds in a completely branched dichasium develop.

by a three-flowered cyme in which the central flower marks the terminal shoot, with
the two lateral flowers arising from the axils of opposite bracts, so that the three buds
lie in one plane. As explained above, because of the phyllotaxy of Hucalyptus, and if
the inflorescence is a dichasium, the plane of these three buds should be at right angles
to the plane of the leaf and stem axis which subtend the inflorescence. It is a fact
that the three-flowered inflorescence which occurs as an exclusive character in some
Species in all the major groups of the genus, is characterized by having the three
flowers lying in one plane, and in this plane being at right angles to the plane
containing the leaf petiole and main axis.

The next stage in complexity above this is a seven-fiowered dichasium in which
two pairs of lateral buds are produced from the axils of the pair of bracts or bracteoles
carried by both members of the pair of the original lateral flowers of the three-fiowered
inflorescence. The planes in which each of these three occur (although not the same)
must lie at right angles to the plane of the original three-flowered inflorescence.

It is found that the seven-flowered inflorescence is common in Hucalyptus—indeed
perhaps the most common unit inflorescence in all groups except the Renantherae,
where larger numbers are common, but that the descriptions have been inadequate to
make this point clear.

84 INITERITANCE OF INFLORESCENCE CHARACTERS IN EUCALYPTUS,

The configuration of the flower cluster in many species, e.g. HE. Macarthuri (Fig-
1, f), indicates that the seven-flowered inflorescence conforms with what is expected in
a dichasial cyme, and that the longer pedicel which sometimes distinguishes the
central, terminal flower on the main axis of the progressive parts of the cyme is
clearly indicated in the position correct for a dichasium.

The seven-flowered cyme is characteristic also of many of the Corymbosae, and in
the case of EH. ficifolia, which has been examined, it is found that, as mentioned by
Maiden, bracteoles are present. The disposition of these can be examined and is found
to correspond precisely with what is required of a dichasium. If the outer pair of
bracts which envelop the inflorescence and related to the central flower of the cyme
are disregarded, there are, and should be if it is dichasial, 12 bracteoles (or bracts)
present as Maiden found, and it was only by assuming the inflorescence was a monopodial
umbel that he felt a seventh bracteole was missing (Fig. 1, d, e).

With regard to the earlier suggestion that as the number of flowers in the
inflorescence increases, lack of space in the condensed dichasium may lead one to
expect that instead of the dichotomous branching of the dichasium continuing, this:
may, after a certain stage, extend as monochasial branching by one member of the
pair of buds being suppressed, the following is of interest. In many specimens of
E. macrorrhyncha (Fig. 3, a) the inflorescence is regular, with the buds geometrically
arranged, and containing 11 flowers. The disposition of the flowers is congruous with
what is required of a dichasial cyme in which, after the seven-flowered stage, only one
axis is developed on each of the four points from which expansion would take place
by two opposite buds, so that four flowers only are added at the next stage instead of
eight, bringing the number to eleven instead of fifteen (as in Fig. 2, a).

In the case of the fifteen-flowered cyme, it is observed in many trees of H. pauciflora
(Fig. 3, d), if careful examination is made, that the buds are also arranged in a
regular geometrical pattern. This pattern is found to conform readily with what would
be expected if the eleven-flowered inflorescence were expanded by similar monochasial
branching (as in Fig. 1, a) to the next unit, a fifteen-flowered inflorescence. It will be
noted in descriptions that a good many species are described as having up to fifteen
buds per inflorescence, and it may well be that the maximum number represents those
eases of dichasia in which there has been no loss of buds during development or
subsequent preservation.

In both #. macrorrhyncha and E. pauciflora it is observed that while in many of
the trees examined the inflorescences are predominantly eleven- and fifteen-flowered
respectively, there are some inflorescences on the same tree which are apparently
quite regular and are fifteen- or eleven-flowered respectively, although the common
maximum number is the reverse and, moreover, that on some individuals there is a
tendency, as in #. macrorrhyncha, for them to be predominantly eleven-flowered, whereas
in other individuals there are mixed eleven- and fifteen-flowered clusters. Both species
also may have a few regular seven-flowered clusters. It seems, therefore, that as the
total number increases, the number of buds is less definite both for the individual and
the species.

An examination was also made of H. Moorei, in which a fairly regular arrangement
can be observed (Fig. 4), and as far as may be deduced from the placement of the buds
in relation one to the other, several patterns of branching are present which lead to
flower clusters with numbers around 20 to 24. The possible organization of these is
demonstrated from reconstruction of some average flower clusters in Figure 4.

It is clear, therefore, that there is scope for more precise description of the
Eucalyptus inflorescence and little doubt that the unit flower cluster is a contracted
dichasium, generally with a definite number of flowers, of which 1, 3, 7, 11 and 15 are
the most common.

Blake (1953) has described the way in which such unit flower clusters can be
brought together on one leafy axis and by the elimination of the leaves a compound
inflorescence, which is usually described as a panicle or corymbose panicle, derived. If
this is studied as in Figure 5 for H. polyanthemos, it is seen that the unit dichasia are

BY L. D. PRYOR. 85

brought together in a compound inflorescence, the secondary arrangement of which
corresponds with the branching pattern developed in the leaf-bearing system. It is
derived, therefore, clearly in the manner in which Blake suggests.

At this point it is interesting to reconsider Tate’s description that “the usual
form of inflorescence is an umbel, which by lengthening of the axis passes to a panicle
or corymb”’. This must be altered and reversed and the usual form of inflorescence

Fig. 4.—EHucalyptus Moorei showing typical inflorescences and the probable pattern of
ssome of them diagrammatically.

Fig. 5.—EHucalyptus polyanthemos. The individual inflorescences are three- or seven-flowered.
‘The arrangement of these is shown in 6b to be opposite and decussate with the pairs separated
in the pattern characteristic of branches. The pairs are indicated by the letters A, B, C, D.

described as “a contracted dichasium which as a unit is often brought together in a
‘secondary paniculate or corymbose arrangement of the clusters by the shortening of the
leafy axis and the elimination of leaves”.

It is worth noting that Rickett says there is no real need to require that the
term ‘‘umbel” be dropped from use, provided in that use it does not obscure the true
‘structure of the flower cluster. The term might be adequately applied when necessary
in Hucalyptus, if it is further qualified, as a “sympodial”’ umbel. On the other hand,

86 INHERITANCE OF INFLORESCENCE CHARACTERS IN EUCALYPTUS,

there is a need to review all species to describe the consistency within the species of
both the number of buds in the flower cluster and their definite structural arrangement.
At present, those occurring singly or in threes are generally known; species with
seven-flowered clusters can often be deduced, but those with higher numbers are
usually obscure.

Before proceeding to examine the inheritance of inflorescence characters, it is
necessary to understand clearly the rea! structure of the inflorescence in these terms.

Inflorescence Characters in Inheritance.
Deductions about the inheritance of inflorescence characters in synthetically
produced F1 hybrids are so far confined to an examination of the Fl, H. rubida x
E. Maideni. E. rubida has exclusively three-flowered dichasia in its inflorescence, which

‘ A

/

: J

deni x Rubida

Fig. 6.—EH. Maideni x E. rubida (E. Maideni seven-flowered; E. rubida three-flowered )
a synthetic Fl hybrid. Note the basal pair of cymes on the H. Maideni x EH. rubida specimen
are three-flowered. The lower of the two has lost two of the buds accidentally.

E Rubida

of course may often be reduced accidentally to two or even one, but in these cases the
scars at the point of bud attachment are always to be seen. H. Maideni has a seven-
flowered dichasium which is usually well displayed because it is robust.

In the F1 hybrid it is noted that the great majority of the inflorescences are seven-
flowered dichasia which suggest this character is approaching complete dominance in
inheritance. However, on about 10% of the unit shoots carrying inflorescences (usually
about four or five pairs) the first pair at the base of the shoot is of three-flowered
dichasia (Fig. 6). In most cases the next pair of inflorescences is of seven-flowered
dichasia, but in a few cases there are exceptions, and it is clear, from the arrangement
of the buds, that various combinations may develop as by the addition of one lateral

BY L. D. PRYOR. 87

bud to the three-flowered group, on one side only giving a four-flowered irregular
cyme. Alternatively, one may develop monochasially on each side only giving a five-
flowered cyme with the start of monochasial branching on either side; or a five-flowered
cyme by the addition of two buds on one side at the base of one of the lateral buds
in the original three, forming one normal half of a seven-flowered cyme, but remaining:
as in the three-flowered condition on the other half.

Fig. 7.—A segregate from #H. cinerea x E. maculosa showing three-flowered cymes in the
basal pair of flowering axils, and an irregular four-flowered cyme showing the addition of the
third order of branching on one side of the cyme and the missing buds (dotted) on the other.

Fig. 8—A segregate from EF. cinerea x H. Blakelyi. Note the three-flowered cyme in the
basal flowering axil, and also showing irregular branching by the elimination of one (C) branch
cn either side of the dichasium.

It seems, therefore, that the character of the seven-flowered dichasium is not
wholly dominant. It is noted that when the three-flowered clusters occur, shoots
carrying them often tend to be grouped, two or three near together on the one branch
of the previous order. This suggests that the determinant of the three-flowered cyme
is able to overcome the otherwise dominant expression of the seven-flowered inflorescence
in some parts of the plants, and particularly in the basal or sub-basal axils of the
flowering shoot. ;

SS INHERITANCE OF INFLORESCENCE CITARACTERS IN EUCALYPTUS,

Some further evidence is adduced from a study of segregating populations derived
from naturally occurring hybrids. Only the female parent is certainly known in
these cases, so the information lacks the precision of the study of a synthetic Fl
hybrid. In particular, evidence is available from segregating progenies of H. cinerea x
HK. Blakelyi, EH. cinerea x EH. maculosa and H. viminalis x EH. elaeophora. E. maculosa
has consistently as its inflorescence a seven-flowered cluster, and H. Blakelyi on many
trees has a seven-flowered group, but sometimes is eleven-flowered. In segregating
progenies from these combinations individuals are found in which, because they
approach the #. cinerea parent more closely, the three-flowered dichasia are more
common than sevens or irregular groups. They nevertheless follow closely, with few
exceptions, the pattern in which the basal axils carry three-flowered inflorescences and
the upper ones on the one flowering shoot carry cymes with more than three flowers
and in the various combinations mentioned.

In the F1 hybrid, the state so far reached is that there is a first flowering comprised
of about 100 inflorescences and a second of about 1,000, and the transition from the
three-flowered cyme to the seven-flowered is usually abrupt. The number of cases in
which there are intermediates is few. In segregating populations the intermediates
become much more frequent, and the various combinations mentioned giving four, five
and even six-flowered irregular cymes (Figs. 7, 8). These facts support the conclusion
which has been reached in the field from the examination of supposed hybrid trees,
that very commonly where the inflorescence of one of the parents is a three-flowered
dichasium and that of the other a seven or more flowered dichasium, some flowering
shoots can be found in which in the basal pair of inflorescences each is three-flowered.
It seems it may be taken as a rule that the presence of three-flowered clusters in the
basal position of a single shoot indicates hybridism.

A further point of apparent regularity is noticed in segregates when, as happens
regularly in a few species, but rarely in most, the concealed bud in a leaf axil produces
a flower cluster as well as the naked bud, so that there are two inflorescences in the
one leaf axil.

If the dichasium from the naked bud is seven-flowered or an irregular combination
of four to six flowers, the complementary dichasium from the concealed bud is usually
three-flowered (Fig. 3, 0), even though this now may be towards the distal end of the
flowering shoot. In one case also in a segregate of H. viminalis x EH. elaeophora the
repeated appearance of a single-flowered cluster from the concealed bud in similar
relation to a three-flowered dichasium from the naked bud has been observed (Fig. 5, D).

Further studies of known F2 populations and back-crosses, when they can be
obtained, give promise of interesting results. It should be noted in those species
which have the three-flowerd dichasium as a characteristic inflorescence that this is
seldom deviated from, and therefore presumably the population comprising the species
is homozygous for this character, since the seven-flowered dichasium (if a generaliza-
tion is risked from the present limited material) is able to express itself in the Fl
with substantial partial dominance.

Summary.

It is deduced that the inflorescence of Hucalyptus is a dichasium, contracted to give
the outward appearance of an umbel. The inflorescence is usually geometrically and
structurally regular and in a large number of species is limited either to three or
seven flowers. In an Fl hybrid between a three- and seven-flowered species, the latter
character is partially dominant. The three-fiowered condition nevertheless is often
found in the basal pair of axils of the flowering shoot of the F1, though the large
majority of the inflorescences are seven-flowered.

In segregating hybrid populations the three-flowered cluster appears in a similar
position, and also often from the concealed bud if this produces flowers. The develop-
ment of irregular combinations between three and seven flowers occurs sometimes in
the Fl, and often in segregates, especially those which exhibit a preponderance of
characters from the three-flowered parent.

BY L. D. PRYOR. 89

References.

BENTHAM, G., 1863-1878.—Flora Australiensis, London.

BuAKE, S. T., 1953.—Australian Journal of Botany, 1, No. 2, 1953.

BLAKELY, W. F., 1934.—A Key to the Eucalypts.

Jacoss, M. R., n.d.—The Primary and Secondary Leaf Bearing Systems of the Hucalypts and
Their Sylvicultural Significance. Commonwealth Forestry Bureau Bulletin, No. 18,
Canberra, n.d.

MAIpEN, J. H., 1924.—A Critical Revision of the Genus Eucalyptus. Government Printer, N.S.W.

MUuELLER, F. v., 1882-1884.—Eucalyptographia. Government Printer, Victoria.

RIcKETT, H. W., 1944.—The Classification of Inflorescences. Botanical Review, 10: 187-231.

Woopson, R. E., Jr., 1935.—Observations on the Inflorescence of Apocynaceae. Ann. Missouri
Botanical Garden, 22, No. 1: 1-48.

90

NOTES ON THE SCLERACTINIA OR STONY CORALS (COELENTHRATA)
OF HHRON ISLAND, QUEENSLAND.

I. A LIST OF THE COMMON SPECIES.
By K. E. W. Satter, Department of Zoology, University Sydney.
[Read 25th August, 1954.]

Synopsis.

About fifty species of Scleractinia are listed in the present paper from a specified locality
not far distant from the southern limits of the Great Barrier Reef. References to the original
descriptions are given and literature used for identification is cited. A new species of
Turbinaria has been discovered but has not been described in the present paper, and for
the first time Coscinaraea has been recorded from the Great Barrier Reef.

INTRODUCTION.

Heron Island is situated about 50 miles north-east of Gladstone, on the coast of
Queensland, close to the Tropic of Capricorn. It is a cay some 800 yards in length
and 300 yards across, and is composed of coral sand, cast up by wind and sea and
partly lithified. It is one of a dozen similar coral cays which together comprise the
Capricorn Group. Many cays at various stages of development are to be found along
the Great Barrier Reef, and maps of forty of these, including Heron, were made by
F. EK. Kemp in 1936 (Steers, 1938).

By contrast with the size of the island, the sub-littoral reef from which it has arisen
is five or six miles long and elliptical in shape. It is edged by a broad reef-crest thickly
cemented by Lithothamnion. This reef-crest is only exposed at low water and it is
then found to encircle a shallow lagoon; however, it is still too deep to wade over the
greater part of this, even when the tide recedes. Prevailing winds and the sea built
up this island on the north-west of Heron Reef at a distance of some two thousand
feet from the broad reef-crest. A shallow, sandy anchorage was also formed between
the island foreshores and the surf-swept crest. Except where indicated, all corals listed
in the present paper were collected in these shallows north of the island between the
sandy beach and the immediate boundary of the reef.

In November, 1925, the Capricorns were visited by members of the Royal Zoological
Society of N.S.W. A most interesting account of the expedition was presented
to that Society by Anthony Musgrave (1926), and it is recorded there that the first
naturalist to visit Heron Island? was J. B. Jukes (1847) during the voyage of H.M.S. Fly
in 1843. Other parties of zoologists visited the islands of the Capricorns in 1904 and
1910. Charles Hedley listed the following twelve Scleractinia from North-west Islet®
(Musgrave, 1926, pp. 249-250): Seriatopora hystrix, Pocillopora bulbosa, Stylophora
pistillata, Goniastrea pectinata, Fungia scutaria, Psammocora gonagra, Acropora
decipiens, Acropora abrotanoides, Acropora surculosa, Acropora brueggemanni, Acropora
baeodactyla, Goniopora tenuidens.

Coral faunas of the Pacific Islands have been studied in considerable detail. This
applies in particular to Murray Island, the Low Isles, American Samoa, Fiji, the
Hawaiian Islands, Laysan and Fanning Islands. Heron Island is not far distant from
the southern extremity of the Great Barrier Reef and is very exposed. Though the
locality studied has been well combed by tourist collectors and was swept by a severe

123° 27’ south.

2Tt received the name of Heron Island before the publication of his narrative in 1847,
but when Jukes landed on 12th January, 1843, the island had no name.

2A eay of the Capricorn Group visible from Heron Island.

BY K. E. W. SALTER. 91

cyclone in February, 1948, the list presented here is of interest since it records some
of the more common species growing so far southward.

In all, a total of about fifty species of coral are recognized in the present paper,
and fer the first time Coscinaraea is recorded from the Great Barrier Reef. A growth
form of Favia speciosa occurs at Heron Island identical with the growth form of that
species found in such abundance at Moreton Bay, Queensland. An undescribed species
of Turbinaria was discovered trom Heron Reef and it is intended to describe this in a
later paper.

This is the first of two papers resulting from field work done in January of 1948,
1950 and 1952. The material collected in 1948 and 1950 was classified into genera and
exhibited at the June meeting of the Linnean Society (Salter and Besly, 1950), while
additional species were discovered in 1952 and identifications completed. In 1952
population counts were made from twelve extended traverses in the same locality. A
later paper will show the distribution and relative abundance of the more common
species of coral occurring at Heron Island.

Acknowledgements.

I wish to acknowledge the assistance given to me by those members of the Sydney
University Biological Society who helped organize expeditions to the Great Barrier
Reef. I should like to express my fullest appreciation of the help given me in the field
in 1948 and again in 1950 by Miss Mary Besly, of the University of Sydney, and for
her subsequent collaboration in the determination of coral genera. Finally I wish to
express my indebtedness to Professor John W. Wells, of Cornell University, Ithaca, N.Y.,
for his valued assistance in checking identifications, for recognition of certain doubtful
species, and for sending me literature unprocurable in Sydney.

List oF STONY CORALS.

Class ANTHOZOA Ehrenberg, 1834.
Subclass HH XACORALLIA Haeckel, 1896.
Order SCLERACTINIA Bourne, 1900.
Suborder ASTROCOENIIDA Vaughan & Wells, 1943.
Family ASTROCOENIIDAE Koby, 1890.
Stylocoeniella armata (Hhrenberg), 343: 1834.

Family SHRIATOPORIDAE Milne Edwards & Haime, 1849.
Stylophora pistillata (HWsper), 1797: 75.
Seriatopora hystrix Dana, 1846: 521.
Pocillopora damicornis (Linnaeus), 1758: 791.

Family ACROPORIDAE Verrill, 1902.
Acropora pulchra (Brook), 1891: 468.
Acropora nasuta (Dana), 1846: 453.
Acropora digitifera (Dana), i846: 454.
Acropora hyacinthus (Dana), 1846: 444.
Acropora squamosa (Brook), 1892: 4638.
Acropora hebes (Dana), 1846: 468.
Acropora humilis (Dana), 1846: 483.
Acropora scherzeriana (Brueggemann), 1878: 397. (Cf. with humilis.)
Acropora brueggemanni (Brook), 1893: 145.
Acropora palifera (Lamarck), 1816: 260. (Cf. with brueggemanni.)
Astraeopora myriophthalma (Lamarck), 1816: 260.
Montipora erythraea Marenzeller, 1906: 58.
Montipora foliosa Pallas, 1766: 333 (?).
Montipora venosa (Ehrenberg), 1834: 118.

Suborder FUNGIIDA Dunean, 1884.

Family AGARICIIDAE Gray, 1847.
Pavona sp.

92 STONY CORALS OF HERON ISLAND, Q.,

Family SIDERASTREIDAE Vaughan & Wells, 1943.
Coscinaraea monile (Forskal),* 1775: 133.

Family THAMNASTERIIDAE Vaughan & Wells, 1948.
Psammocora contigua (Esper), 1797a: 81.
Psammocora profundacella Gardiner, 1898: 537.
Psammocora exesa Dana, 1846: 57.

Family FUNGIIDAE Dana, 1848.
Fungia scutaria Lamarck, 1801: 370.
Fungia fungites (Linnaeus), 1758: 793.
Fungia sp.°

Family PORITIDAE Gray, 1842.
Goniopora sp.
Porites, several spp.

Suborder FAVIIDA Vaughan & Wells, 1943.
Family FAVIIDAE Gregory, 1900.
Subfamily FAVIINAE Vaughan & Wells, 1943.
Favia valenciennesi (Milne Edwards & Haime), 1850: 124.
Favia pallida (Dana), 1846: 224 (= doreyensis M.E. & H., 1850: 168).
Favia speciosa (Dana),° 1846: 220.
Favia stelligera (Dana), 1846: 216.
Plesiastrea versipora (Lamarck), 1816: 264.
Favites abdita (Hillis & Solander), 1786: 162.
Favites virens (Dana), 1846: 228.
Goniastrea benhami Vaughan, 1917: 277.
Platygyra daedalea (Ellis & Solander), 1786: 163.
Leptoria gracilis (Dana), 1846: 261.
Hydnophora exesa (Pallas), 1766: 290.

Subfamily MONTASTREINAE Vaughan & Wells, 1943.
Leptastrea purpurea (Dana), 1846: 239.
Cyphastrea serailia (Forskal), 1775: 135.
Echinopora lamellosa (Esper), 1797b: 65.

Family OCULINIDAE Gray, 1847.
Galaxea fascicularis (Linnaeus), 1767: 1278.
Galaxea musicalis (Linnaeus) ,‘ 1767: 1278.
Acrhelia horrescens (Dana), 1846: 392.

Family MERULINIDAE Verrill, 1866.
Merulina ampliata (Ellis & Solander), 1786: 157.

Family MUSSIDAE Ortmann, 1890.
Lobophyllia corymbosa (Forskal), 1775: 137.
Symphyllia agaricia Milne Edwards & Haime, 1849: 255.

Suborder DENDROPHYLLIIDA Vaughan & Wells, 1943.
Family DENDROPHYLLIIDAE Gray, 1847.
Balanophyllia sp.
Tubastraea manni (Verrill), 1866: 30.
Turbinaria peltata (Wsper), 1797: 27.
Turbinaria, undescribed species.

4Wirst record of Coscinaraea from Great Barrier Reef.

5 Locality: One Tree Island.

6One example was found of a growth form of F. speciosa which was identical with the
growth form of this species found abundantly in Moreton Bay, Queensland.

7From a specimen in the Eleanor Chase collection, North-West Islet.

BY K. E. W. SALTER. 93

Bibliography.

BERNARD, HENRY M., 1896.—The Genus Turbinaria, the Genus Astraeopora. Catalogue
Madreporian Corals in British Museum (Natural History), IL: 1-106.

———, 1897.—The Genus Montipora, the Genus Anacropora. Catalogue Madreporian Corals in
British Museum (Natural History), II1: 1-192.

BourRNE, G. C., 1900.—Anthozoa, in Treatise on Zoology, II, by Ray Lankester.

Brook, GEORGE, 1891.—Description of new species of Madrepora in the collection of the British
Museum. Ann. Mag. Nat. Hist., (6), 8: 468.

, 1892.—Preliminary description of forty new species of Madrepora in the collection of
the British Museum. Ann. Mag. Nat. Hist., (6), 10: 463.

, 1893.—The genus Madrepora. Catalogue Madreporian Corals in British Museum
(Natural History), I: 1-212.

BRUEGGEMANN, F., 1878.—Neue Korallen-Arten aus dem Rothen Meere und von Mauritius.
Abhandl. Naturw. vereins, Bremen, Bd. V: 397, Pl. viii.

CROSSLAND, CYRIL, 1952.—Madreporaria, Hydrocorallinae, Heliopora and Tubipora. Great
Barrier Reef Haped., 1928-29, Scientific Reports, VI, 3: 85-257.

Dana, J. D., 1846.—Zoophytes. U.S. Exploring Expedition, 1836-42, Commanded by Charles
Wilkes, U.S.N., VII: 1-740, with folio Atlas (1849) of 61 plates.

DUNCAN, P. MaArTIN, 1884.—A revision of the families and genera of the sclerodermic
Zoantharia, Ed. and H., or Madreporaria (M. rugosa excepted). J. Linn. Soc. Lond.,
XVIII: 104-5.

Epwarps, H. MILNE, and HAIME, J., 1848-1851.—Recherches sur les polypiers. Mém. IV.
Monographie des Astraéides. Ann. Sci. Nat. Zool. (3). F. valenciennesi, X, Pl. 9, fig. 3,
XII: 124; S. agaricia, XI: 255; F. doreyensis, XII: 168.

, mm, 1849.—Mémoire sur les polypiers appartenant aux groupes naturels des
zoanthaires perforés et ses zoanthaires tabules. Acad. Sci. Paris, Compte Rendu, 29: 262.

EX}HRENBERG, C. G., 1834.—Die Corallenthiere des rothen Meeres. Beitrige zur physiologischen
Kenntniss der Corallenthiere im Allgemeinen. K. Ak. Wiss. Berlin, Abh., 1832: 225.

ELuIs, J., and SoOLANDER, D., 1786.—The natural history of many curious and uncommon
Zoophytes collected from various parts of the globe by the late John Ellis, systematically
arranged and described by the late Daniel Solander. 157, Pl. 41, figs. 1 and 2; 162, Pl. 50,
fig. 2; 163, Pl. 46, fig. 1.

Esper, BE. J. C., 1797a.—Die Pflanzenthiere in Abbildungen nach der Natur mit Farben
erleuchtet nebst Beschreibungen. Th. I, 75, Pl. 60; 81, Pl. 66. Nurnberg.

——., 1797b.—Fortsetzungen der Pflanzenthiere, etc. Th. I, 65, Pl. 58, figs. 1 and 2; 27, Pl.
42. Nurnberg.

ForRSKAL, PETRUS, 1775.—Descriptiones animalium .. . quae in itinere orientale observavit.
Hauiiae, 133, 1385 and 137.

GARDINER, J. STANLEY, 1898.—On the fungid corals collected by the author in the South
Pacific. Proc. Zool. Soc. Lond., 537, Pl. 45, fig. 3.

GRAY, J. E., 1842.—Synopsis, British Museum (44th ed.), 135.

— , 1847.—An outline of an arrangement of stony corals. Ann. Mag. Nat. Hist... (1),
oes WAS,

GreGory, J. W., 1900.—The Corals. Jurassic fauna of Cutch. Pal. Ind. (9), 2, 2: 66.

HAECKEL, ERNST, 1896.—Systematische Phylogenie der Wirbellosen Tiere, Berlin, xviii: 720.

HOFFMEISTER, J. EDWARD, 1925.—Some corals from American Samoa and the Fiji Islands.
Carnegie Inst. Wash., XXII, No. 343: 1-90, 23 Pls.

JUKES, JOSEPH B., 1847.—Narrative of the Surveying Voyage of H.M.S. Fly commanded by
Capt. F. P. Blackwood, R.N., during years 1842-1846. JLond., 1: 6.

Kosy, F., 1890.—Monographie des polypiers jurassiques de la Suisse. Schweiz. pal. Gesellsch.
Abhand., ix: 570.

LAMARCK, J. B. P. D&E, 1801.—Polypes A rayons. Systeme des animaux sans vertébres.
Paris. ii: 370.

, 1816.—Polypiers lamelliferes. Histoire natwrelle des animaux sans vertébres. Paris.
ii: 260-4.

LINNAEUS, CAROLUS VON, 1758.—Systema naturae. I. Regnum animale, 10th ed.: 791.

—, 1767.—Systema naturae, 2, 12th ed.: 1278.

MARENZELLER, EMIL VON, 1906.—Riffkorallen. Expeditionen S.M. Schiff Pola in das Rote Meer.
Denkschr. k. k. Akad. Wiss. Wien., Math. Naturwiss. Kl., 80: 58, Pls. 22, 23.

MATTHAI, GEORGE, 1914.—A revision of the recent colonial Astraeidae possessing distinct
corallites. Trans. Linn. Soc. Lond., 2nd Ser., Zool., 17, Pt. 1: 1-140, 38 Pls.

———, 1928.—A monograph of the recent Meandroid Astraeidae. Catalogue Madreporian Coral
in British Museum, 7: 1-288, 72 Pls.

MUSGRAVE, ANTHONY, 1926.—The Biology of North-West Islet. Aust. Zool., 4, Pt. 4: 199-255.

ORTMANN, A., 1890.—Die Morphologie des Skeletts der Steinkorallen in Beziehung zur
Koloniebildung. Zeitschr. wiss. Zool., 50: 315.

PaAuLas, P. S., 1766.—Hlenchus zoophytorum. The Hague: 2906, 333.

SALTER, K. E. W., and Brsty, M. A., 1950.—Proc. LINN. Soc. N.S.W., L-XXV, Pts. 5-6, Abstract
of Proc., xxv.

94 STONY CORALS OF HERON ISLAND, Q.

Strerrs, J. A., 19388—Detailed notes on the Islands surveyed and examined by the Geographical
Expedition to the Great Barrier Reef in 1936. Reports of the Great Barrier Reef
Committee, IV, Pt. III: 61.

VAUGHAN, T. WAYLAND, 1907.—Recent Madreporaria of the Hawaiian Islands and Laysan.
Smithsonian Inst. U.S. Nat. Mus., 59: 1-222.

—— —, 1917.—Some Corals from the Kermadec Islands. Trains. € Proc. N.Z. Inst., XLIUX: 277.

——-——, 1918.—Some Shoal-water Corals from Murray Island, Cocos-Keeling Islands, and
Fanning Island. Carnegie Inst. Wash., IX, 213: 51-234, Pls. 20-93.

VAUGHAN, T. WAYLAND, and WELLS, J. W., 1943.—Revision of the Suborders, Families, and
Genera of the Scleractinia. Geol. Soc. Amer., Special Papers, No. 44.

VERRILL, A. E., 1866.—Synopsis of the Polyps and Corals of the North Pacific Exploring
Expedition, with descriptions of some additional species from the West Coast of North
America. Proc. Hssex Inst., II1: 30; 1V: 9.

———, 1902.—Variations and nomenclature of Bermudian, West Indian, and Brazilian Reef
Corals, with notes on various Indo-Pacific Corals. Conn. Acad. Art. Sci., Tr., 11; 163.
YABE, H., and SuGiyAmMA, T., 1933.—Notes on three new Corals from Japan. Jap. J. Geol.

Geogr., IX, 1-2, Tokyo: 11, Pl. 2, figs. 1-4.

YABE, H., SuGiyaAmMa, T., and HEa@ucHi, M., 1936.—Recent Reef-building Corals from Japan
and the South Sea Islands under the Japanese Mandate. I. Sci. Rep. Tohoku Imp. Univ.,
2nd Ser. (Geol.), Special Vol. 1: 1-66, 59 Pls.

95

COLLENIA FREQUENS IN UPPER PROTEROZOIC ROCKS IN THE NORTHERN
TERRITORY OF AUSTRALIA.
By D. M. TRAVES.
(Communicated by Dr. Germaine A. Joplin.)
(Plate vi; one Text-figure.)
[Read 28th July, 1954.]

Synopsis.
Specimens of a Pre-Cambrian alga were collected from two localities in the north-western
part of the Northern Territory in 1952. These specimens closely resemble the form Collenia
frequens described by Walcott, and Fenton and Fenton, from the Belt Series of North America.

Cryptozoan frequens was first described in 1906 by C. D. Walcott. In 1914 he trans-
ferred the species frequens into the genus Collenia. Fenton and Fenton in 1931, 1933
and 1937 described specimens of Collenia frequens Walcott from the Belt Series of Upper
Proterozoic sediments of North America.

, >
. YrerGowns 5
¥ SMave /i// |
| GQ” ; ia Q
iH Be “ossi/ localities +
ae NORTH E/|RN |Z
+
at Tennant C) POA |
re a4 op
Baee| Do ReRIT THOR y: le
De, lz
An [2
e +
ica
3 a Ata, WIGS i.
<4 |
\ ie

rr Fa ee eee NT vag rene le

Text-fig. 1.—Locality map.

The Australian specimens attributed here to C. frequens Walcott were collected from
two localities in the north-western part of the Northern Territory in 1952. At both
localities small outcrops of silicified Upper Proterozoic rocks form inliers in the Antrim
Plateau Volecanics of Lower Cambrian age. The first locality is at Top Spring at the
junction of the Montejinni and Murrenji Tracks and the second locality is seven miles
west of Catfish Yard, south of Wave Hill Station.

96 COLLENIA FREQUENS IN UPPER PROTEROZOIC ROCKS.

Description.

The colonies are elongate-conical with each colony consisting of conical laminae
from 1 to 2 mm. apart, which are grouped into layers of 3-15 mm. The apices of the
laminae are rounded and most specimens have a hole or tube through the axis of the
cone. This is well illustrated in Plate vi, figure 1.

Most specimens have a similar taper with an apical angle of approximately 50° and
a ratio of the diameter to the length of the cone of approximately 1-1. However two
specimens have a different taper with apical angles of approximately 30° and a ratio
of -5.

Diameters range from less than 2-5 cm. (1 inch) to greater than 61 cm. (2 feet). In
most cases the outside laminae spread over other colonies.

At both localities the colonies have formed a bioherm which has been surrounded by
the Antrim Plateau Volcanics. The inliers are completely silicified and, in places, rock
erystal has grown within the structures.

Remarks.

The Australian specimens of 0. frequens compare well with the description given by
Fenton and Fenton (1937) and the specimens shown in their plate 15 bear a striking
resemblance to the one in Plate vi, figure 3. However, they do not mention the presence
of an axial tube, which is a prominent feature of the specimens from the two Northern
Territory localities. This tube may have been filled or hidden by different modes of
preservation, but it would be interesting to re-examine the specimens from the Belt
Series of North America to see if a central tube did exist. Also no apical angles of the
cones were given and, as specimens of two different angles were found in the Northern
Territory, the complete range of taper is not known.

Plate vi, figure 4 shows the rounded apical end of C. frequens which easily dis-
tinguishes it from the cone-in-cone structure found in some sediments.

Although C. frequens is the nearest described form it is well to remember that in
the case of Proterozoic algae the general opinion is that each form may be built up of
several species of algae. However, forms may still be used for stratigraphical correlation
and sub-division.

References.

FENTON and FENTON, 1931.-—Journ. Geol., 39: 685.

; , 1933.—Geol. Soc. Am. Bull., 44: 1135-1142.
: —, 1937.—Geol. Soc. Am. Bull., 48 : 1813-1970.
WatLcoTt, C. D., 1914.—Smithsonian Misc. Coll., 64:112.

EXPLANATION OF PLATE VI.
1. C. frequens on top of bioherm at Top Spring, N.T.—2. ©. frequens on side of bioherm at
Top Spring, N.T.—3. Section of C. frequens. x 4.—4. ©. frequens, showing rounded apical
end. x 3.

oF

AUSTRALIAN FUNGI.
II. NEW RECORDS AND REVISIONS.
By C. G. HANSForD, Sc.D., Waite Agricultural Research Institute, University of Adelaide.
(Two Text-figures. )
[Read 28th July, 1954.]

Synopsis.
One hundred fungi are dealt with in this paper; some represent new species, others new
records for Australia or for parts thereof, and notes are included on a number of species
previously recorded here.

This paper deals with a number of Australian Fungi sent to me from various
Herbaria here, and by a number of collectors in different parts of the continent. The
origin of each collection is detailed below, and I have to thank the authorities and
collectors concerned for their most valuable assistance in endeavours to revise the
Fungus Flora of Australia. I can only hope that the present paper will stimulate other
collectors to more activity in this field. The numbering of the species is continued from
the first paper of this series, published in these PRocEEDINGS, 78:51—82, September 1953.

PHYCOMYCETES.
(62) CySTOPUS CENTAURII Hansf., n. sp.

Sori conidiiferi foliicoli et caulicoli, albi, ovoidei vel rotundati, subinde confluentes
et elongati, usque ad 1-5 mm. longi, primo epidermide inflato tecti, demum illo disrupto
subnudescenti, pulverulenti. Conidiophora erecta, hyalina, 20-40 x 12-18u, stipata,
simplicia. Conidia acrogena, catenulata, globosa vel subcuboidea, hyalina, levia,
continua, 18-264 diam., interdum membrana intus annulato-incrassata. Oogonia in
mesophyllo et in cortice caulis dispersa, globosa vel ovoidea, usque ad 70 x 0p; oosporae
singulae, ex fulvo brunnescentes postremo subopacae, globosae, 50-60u diam., plerumque
grosse uni-guttulatae; episporio 5—6u cr., dense reticulato-sulcato.

Hab. in foliis Centaurii spicati, The Chalet, Kosciusko, New South Wales, A. Costin
59.

The conidial sori occur on leaves, less commonly on the stems, and are white and
pulverulent when mature, surrounded by the broken epidermis, up to 1-5 mm. long on
the stems when confluent, covered at first by the inflated epidermis. The conidiophores
form a close palisade at the base of the sorus, and are erect, simple, hyaline, continuous,
20-40 x 12-18u, forming simple chains of spores at the apex, without interstitial cells.
The young spores are globose, but soon become almost cuboid from mutual pressure, and
are hyaline, smooth, continuous, 18-264 diam., sometimes showing an indistinct internal
transverse annular thickening of the wall. The oogonia occur in the leaf mesophyll
and also in the cortical tissue of the stem, being loosely scattered, globose to ovoid,
thin-walled, up to 70 x 50u; each contains a single oospore, which is at first pale
yellowish, but soon darkens first to brown and finally to almost opaque black-brown.
Mature oospores are globose, 50-60” diam., usually containing a single large oil globule;
the epispore is 5-6u thick and closely reticulate with furrows descending 3—4u into the
wall.

This is close to C. swertiae (Berl. & Rom.) Sacce., as given in Syll. Fung. 21:858, 1912,
but the Australian species differs in its larger conidia and oospores.

ASCOMYCETES.
(63) ELSINOE TRISTANIAE Hansf., n. sp.
Maculae epiphyllae, usque ad 70 mm. diam., rufobrunneae, concentrice indistincte
zonatae, zona lutea 5 mm. er. circumdatae. Mycelium in mesophyllo ex hyphis hyalinis,

98 AUSTRALIAN FUNGI, II,

ramosis, septatis, 1-5-3u cr. compositum, sub cuticula folii pseudoparenchymatice
aggregatum et laminam continuam usque ad 30u cr. efformans. Conidiophora dense
fasciculata, erumpentia, erecta, simplicia, hyalina, continua, 10-25 x 1-5—2u4. Conidia
singula, acrogenea, hyalina, continua, ovata, levia, circa 3 x 1-54. Asci in stroma
subcuticulare irregulariter dispersi, singuli vel laxe gregarii, globosi vel ovati, sessiles,
8-spori, circa 35u diam., aparaphysati. Sporae hyalinae, conglobatae, muriformes,
transverse 7-septatae, in medio leniter constrictae, leves, 20-25 x 10-11p.

Hab. in foliis Tristaniae confertae, Chatswood, New South Wales, July 1935, L.
Fraser; Murwillumbah, Fraser, Feb. 1951.

Leafspots epiphyllous, up to 70 mm. diam., becoming red to brown in the centre,
with a yellow-buff border about 5 mm. wide, somewhat concentrically zonate; sometimes
producing an indefinite brown discoloration on the lower side of the leaf. The mycelium
penetrates the mesophyll as fine, branched, hyaline, septate hyphae 1-5—3y thick, inter-
cellular and without haustoria in the host cells. A complete sheet of closely interwoven
similar hyphae, up to 30u in depth, is formed between the upper epidermis and cuticle,
the latter being raised and broken in places by the tufts of conidiophores formed above
this subcuticular stroma; viewed from the surface the raised pattern of the cuticle shows
as irregularly reticulate lines 50-1004 wide. The conidiophores are densely fasciculate,
forming a palisade covering the somewhat thickened mycelial stroma, erect, unbranched,
10-25 x 1-5-2u, hyaline, continuous, forming single conidia at the apex. Conidia not
adherent in chains, hyaline, ovate, continuous, smooth, thin-wailed, about 3 x 1:5u.

The asci are scattered irregularly throughout the subcuticular stroma beneath the
layer of conidiophores, usually single, but sometimes in loose groups separated by mycelial
hyphae, globose to ovate, thin-walled, sessile, 8-spored, about 35u diam.; at maturity
each elongates through the stroma towards the surface, when the free apex discharges
the spores. Paraphyses none. Spores conglobate in the ascus, hyaline, slipper-shaped,
with usually 7 transverse and 1—2 longitudinal septa, constricted at the middle septum,
the upper half slightly wider than the lower, smooth, 20-25 x 10-llu, rather strongly
refractive.

(64) ELSINOE EUCALYPTI Hansf., n. sp.

Maculae amphigenae, irregulares, leniter ruguloso-elevatae, atrobrunneae, in centro
griseo-brunneae, acute limitatae, usque ad 10 mm. diam., vel confluentes. Mycelium ex
hyphis hyalinis, ramosis, septatis, intercellularibus compositum, mesophyllum totum
folii penetrantibus et sub epidermide in stromam plectenchymaticam aggregatis usque
ad 90u cr., demum erumpente et conidiifera. Conidiophora dense stipata, erecta,
simplicia, continua, hyalina, usque ad 20u longa; conidia singula, acrogena, hyalina, ovata,
continua, levia, 4-6 x 3-4u. Asci primo in greges minutos laxos immersos evoluti,
globosi, demum late ellipsoidei, sessiles, usque ad 45 x 28, apice incrassati (5-6y),
8-spori. Sporae conglobatae vel multiseriatae, hyalinae, clavulato-ellipsoideae utrinque
rotundatae, 20-26 x 7-8u, transverse 3- et longitudinaliter 1-septatae, in medio leniter
constrictae, leves, refringentes.

Hab. in foliis Hucalypti sp. indet., Rocky River, Kangaroo Is., South Australia, Feb.
1920, WARI 2010,* leg. T. Osborn.

Leafspots amphigenous, irregularly rounded and elevated, rugulose, sharply defined,
dark brown with greyish-brown centre, up to 10 mm. diam. or confluent, showing through
the leaf on the opposite side as a smooth, dark brown, somewhat raised spot. The
internal mycelium penetrates the whole mesophyll as intercellular hyaline septate, much
branched hyphae, without haustoria, and aggregates beneath the epidermis to form a
plectenchymic stroma up to 90 thick in places; the epidermis eventually ruptures
irregularly and is shed to expose the surface of the stroma, which is then covered with
a dense palisade of conidiophores. Conidiophores erect, simple, continuous, hyaline,
forming conidia singly at the apex, up to 20u long by 2-3u thick. Conidia hyaline, ovate,
smooth, continuous, 4-6 x 3-4u. The asci are first formed in small loose groups within

* WARI = Waite Agricultural Research Institute.

BY C. G. HANSFORD. 99

the stroma, by extension these groups become irregularly and rather widely confluent;
the asci are single and separated by the stroma hyphae, but are eventually formed in
large numbers and then appear in two or three indefinite layers, more or less contiguous,
or separated by ‘“‘septate paraphyses” which represent the remains of the stroma hyphae.
Single asci are first globose, becoming wide ellipsoid, sessile, aparaphysate, 8-spored, up
to 45 x 28u, the wall thickened around the apex to 5-6u; at maturity each elongates to
reach the surface of the stroma, discharges its spores and collapses to make room for
others. Spores conglobate or multi-seriate in the ascus, hyaline, slipper-shaped with
rounded ends, 20-26 x T7-8u, transversely 3- and longitudinally 1-septate, slightly
constricted at the middle septum and the upper half often slightly wider than the lower,
smooth, strongly refractive.

The above two species of Hlsinoe are the first of this genus to be recorded on
indigenous host-plants in Australia; the genus is becoming better known and its distribu-
tion on native flora now includes most parts of the tropics. Both of these Australian
species have typical Sphaceloma conidial stages.

(65) MATIREELLA TASMANICUM (Mass.) Hansf., n. comb.

= Dimerosporium tasmanicum Mass., Kew Bull. 1898: 129; Sacc. Syll. Fung., 16:410,
1902.

This fungus forms small black to greyish raised spots on one or both sides of the
phyllodes, up to 1 mm. diam., at first covered by the epidermis and consisting of a
subcuticular plate of subhyaline cells in 2-3 layers, up-to 304 thick in places,, thinner
towards the edge. The cells are elongated perpendicularly to the leaf surface, and
measure 5-15 x 3-6u, at first hyaline, then turning dark brown in spots, finally becoming
almost entirely brown to black. There is no further penetration of the host, and no
haustoria are formed in the epidermal cells. At several points this subcuticular plate
bursts through the cuticle and forms an ectostroma covering the whole area above the
plate, black, up to 350u high, somewhat rough and containing ascigerous loculi; later
other perithecioid loculi are formed around the edge of the ectostroma, free from each
other and from the remains of the original locules. These secondary loculi have smooth
walls, with an indistinct apical opening, and are irregularly depressed-globose, up to
300u diam.; the wall consists of several outer layers of dark brown, rounded-polygonal
cells, the outermost 10-154 diam. and about 8 deep, the inner cells smaller and with
thinner walls, gradually passing inwards into hyaline tissue, which in early stages may
more or less completely fill the interior. The hymenium of asci and paraphyses covers
the basal two-thirds of the mature loculus, and consists of numerous asci directed
towards the apical opening, mixed with very numerous, flexuous, filiform, continuous,
hyaline paraphyses about ly thick, rather gelatinous and intertwined to form a soft
tissue, apparently unbranched. The paraphyses appear to arise from the whole inner
surface of the loculus. Asci obclavate to cylindric, rounded at the apex, shortly nodose-
stipitate, 8-spored, up to 80 x 20u, the wall around the apex is in immature asci
thickened up to 3u, but no sign of an interior canal or pore was observed, nor any trace
of a double wall. Spores irregularly 2-seriate in the ascus, clavate, light brown, smooth,
1-septate and slightly constricted, 18-21 x 8—9u, the upper cell somewhat larger than the
lower, both ends rounded.

The cavity of each loculus is lysigenous in origin, and the stromatic wall in mature
loculi is about 25 thick in the upper part. The original or primary loculi are irregularly
dehiscent, and rapidly break up and fall away, so that the fully mature ascostroma
consists of a ring of perithecioid loculi surrounding their remains.

On phyllodes of Phyllocladus rhomboidalis, St. Crispin’s Well, Tasmania, Rodway
367 in Herb. Univ. of Tasmania; sent to me through the courtesy of Prof. Barber.

(66) MELIOLINA NOVAE-ZEALANDIAE Hansf., n. sp.

Plagulae hypophyllae, usque ad 10 mm. diam., vel late confluentes, atrae, dense
velutinae. Mycelium ex hyphis atrobrunneis, irregulariter ramosis, exhyphopodiatis,
repentibus compositum, supra stomata folii stomopodia efformans et mesophyllum

100 AUSTRALIAN FUNGI, II,

penetrans. Setae myceliales numerosae, erectae, simplices vel irregulariter furcatae,
ramulis usque ad 120u longis, apice obtusae vel leniter attenuatae, usque ad 440y alt. et
8-llu er. Perithecia dispersa, atra, depresso-globosa, usque ad 5004 diam. et 250 alt.,
sursum setulosa; setae numerosae, simplices, atrae, obtusae, usque ad 250y longae;
paries perithecii extus verrucosus, parenchymaticus, cellulis rotundato-polygonalibus,
leniter prominentibus, atrobrunneis, intus in massam hyalinam mollem transeuntibus.
Asci sat numerosi, basales, erecti, ovati vel late clavati, apice rotundati, nodoso-stipitati,
circa 150 x 50u, 8-spori. Paraphyses hyalinae, septatae, 3-54 cr., evanescentes. Sporae
2-3-seriatae, cylindraceo-ellipsoideae, utrinque obtusae, 3-septatae, leves, 45-55 x 18-21p.

Hab. in foliis Metrosiderodis excelsae, Rangitoto Is., Auckland, New Zealand,
Dingley, July 1950, in Herb. Division of Plant Diseases, Auckland, N.Z.

The colonies are hypophyllous, up to 10 mm. or more in diam. or widely confluent,
black, densely velvety. External mycelium of dark brown hyphae creeping over the leaf
and forming stomopodia over the stomata, from which the internal mycelium enters the
leaf and penetrates most of the mesophyll as intercellular, hyaline, septate hyphae
which do not form haustoria in the host cells. Mycelial setae numerous, erect, mostly
simple and obtuse to somewhat attenuate at the apex, up to 440 x 8-10y, or some forked
irregularly with branches up to 120u long. Perithecia scattered amongst the mycelial
setae, black, depressed-globose, the upper half bearing numerous erect-spreading simple
black obtuse setae, up to 250u long; perithecia up to 500u diam. and 250u high, opening
by a rounded pore at the vertex; the wall consists of an outer layer of rounded-polygonal
cells which are slightly prominent, hence the surface appears slightly verrucose, passing
internally into a hyaline mass of soft parenchyma, at first filling the whole interior.
Within this soft tissue the fairly numerous asci develop from the base, replacing the
original ground tissue, the remains of which form the more or less evanescent septate
“paraphyses”. Asci widely clavate to ovate, rounded at the apex and when young
thickened there to 10u, nodose-stipitate below, 8-spored, about 150 x 50u. Spores cylindric
to slightly ellipsoid, obtuse at the ends, dark brown, 3-septate, slightly constricted,
45-55 x 18-21u, the cells approximately equal in length and the middle ones not noticeably
swollen; end cells with the subterminal and sub-basal narrow hyaline bands usual in
this genus.

(67) MELIOLINA CLADOTRICHA (Lev.) Sydow, Ann. Mycol., Berlin, 12:5538, 1914.

= Meliola cladotricha Lev. = Meliola octospora Cooke. = Meliolinopsis octospora
(Cooke) Beeli. = Meliolina octospora (Cooke) v. Hoehnel. = Meliolina sydowiana Stev.
(ex descr.). = Meliola arborescens (Syd.). Meliolina arborescens (Syd.) Syd. = Meliolina
yatesti Syd. = Meliolinopsis yatesii (Syd.) Beeli. = Meliolina radians Syd.

In my previous account of the genus Meliolina (Proc. Linn. Soc. London, 157:145,
1944-5), I retained the three species M. cladotricha, M. octospora and M. arborescens as
distinct. I have recently been able to examine the type specimen of M. cladotricha on
Syzygium sp., Borneo, in Herb. Paris, and am unable to distinguish it from M. octospora.
Also in recent years I have examined a series of specimens which have indicated that
there is no real difference between M. cladotricha and M. arborescens, aS some specimens.
have proved intermediate between these. I have seen two collections of M. cladotricha
from Australia: Bailey 506 on Tristania sp., Brisbane 1886, in Herb. Kew; and Fraser
191, on Syzygium paniculatum, Dorrigo, New South Wales.

(68) MELIOLINA MOLLIS (B. & Br.) v. Hoehnel, Sitzb. K. Akad. Wiss. Wien, 128:557, 1919..
= Meliola mollis B. & Br. = Dimerosporium mollis (B. & Br.) Sace.
On Acmena smithii, National Park, New South Wales, June 1935, L. Fraser 222.

(69) BALLADYNA FRASERI Hansf., n. sp.

Plagulae amphigenae, usque ad 4 mm. diam., densae, atrae, breviter velutinae.
Mycelium radianto-reticulatum, ex hyphis atrobrunneis, 6—7u cr., cellulis plerumque
15-20u longis, irregulariter ramosis compositum. Hyphopodia continua, alternata vel
unilateralia, irregulariter digitata vel pulvinata, integra vel saepius sinuosa vel

BY C. G. HANSFORD. 101

sublobata, 8-12 x 5-8u. Setae myceliales numerosae, erectae, simplices, obtusae,
subrectae, opace atrae, 2-—3-septatae, usque ad 140 x 7—-9u deorsum non vel lenissime
tumidae, extus leniter granulosae. Perithecia lateralia, breviter stipitata, nigra, globosa,
usque ad 100u diam., apice poro rotundato, circa 154 diam., pertusa; paries verisimiliter
unistratosus, parenchymaticus, cellulis angulosis, circa 154 diam.; deorsum hyphis
radiantibus ornata. Asci aparaphysati, 8-10, in ordine maturescentes, collabascentes,
late ovati, 8-spori, sessiles, usque ad 50 x 25u, in maturitate usque ad 80u longi. Sporae
inordinatae vel subparallelae, oblongae, obtusae, 1-septatae, leniter constrictae, leves,
18-22 x 7-8u, brunnescentes, cellula superiore subglobosa, inferiore longiore et deorsum
leniter attenuata.

Hab. in foliis Canthii coprosmoidei, Hastings R., New South Wales, L. Fraser, April
1952.

Colonies amphigenous, to 4 mm. diam., dense, black, shortly velvety. Mycelium
closely radiating-reticulate, of dark brown hyphae 6—7u thick, the cells mostly 15-20u
long, branching irregular, rarely opposite, at acute angles. Hyphopodia unicellular,
alternate or unilateral, digitate to irregularly pulvinate, concolorous with hyphae, entire
or more often with sinuous outline or sublobate, 8-12 x 5-8u. Mycelial setae numerous,
erect, opaque black, simple, obtuse, more or less straight, 2—3-septate, up to 140 x 7-9un,
the base not or scarcely swollen, the surface slightly granulose, the wall not thickened.
Perithecia arising as short erect lateral branches from the mycelial hyphae, of which
the terminal cell develops into the perithecium, the lower cell remaining as a short
stalk; mature perithecia black, globose, up to 100u diam., with indistinct round apical
pore about 154 diam.; the perithecial wall appears to consist of only a single layer of
dark brown to almost greenish-black angular parenchyma, the cells about 154 diam., but
smaller towards the vertex, smooth on the surface, with some cells around the sides and
base growing out into radiating ‘hyphal struts” to support the developing perithecium.
No setae on the perithecia. Asci basal, formed from branching ascogenous hyphae,
aparaphysate, 8-10 in each perithecium, maturing in succession and collapsing after
spore discharge, broadly ovate, becoming more elongate at maturity, 8-spored, sessile,
about 50 x 25u, elongating to about 80u. The apex of the immature ascus is thickened
to 8u, with a slight central internal canal; no definite evidence of a double-layered wall
was found. Spores multiseriate to sub-parallel in the ascus, oblong with obtusely
rounded ends, becoming brown, 1-septate, slightly constricted, smooth, 18-22 x 7-8u; the
upper cell distinctly shorter than the lower and subglobose, the lower cell slightly
attenuate to the rounded base. —

This species differs from all others in the genus in the greater number of asci in
each perithecium, and also in the unequally-septate spores.

(70) MELIOLA ARUNDINIS Pat., Journ. de Bot., 1897: 348.
On Phragmites vulgaris, Johnstone R., Queensland, 1889, S. Johnson (in Herb.
National, Victoria).

(71) STIGMATEA LOMATIAE Hansf., n. sp.

Plagulae epiphyllae, usque ad 5 mm. diam., subinde confluentes, leves, in sicco
luteae, leniter elevatae. Mycelium subcuticulare, brunneum, ex hyphis 5-7y er., cellulis
usque ad 12u longis, ramosissimis compositum, pelliculam unistratosam efformans.
Haustoria nulla. Thyriothecia dispersa, rotundata, atra, levia, nitentia, subcuticularia,
usque ad 150u diam.; paries superior unistratosus, atrobrunneus, radiatus, poro centrali
aperta. Paries inferior subhyalinus, tenuis, ex hyphis 2—3u ecr., intertextis compositus.
Asci suberecti, centripetaliter directi, ellipsoidei, sessiles, usque ad 75 x 18u, 8-spori.
Sporae 2-3-seriatae, oblongo-clavulatae utrinque rotundatae, 1-septatae, constrictae, leves,
brunnescentes, 20-22 x 8—9u, cellula inferiore leniter angustiore.

Hab. in foliis Lomatiae arborescentis, Williams R., New South Wales, L. Fraser
204, p.p.

Colonies epiphyllous only, up to 5 mm. diam., or sometimes confluent, smooth,
yellowish-brown when dry, possibly yellow-green when fresh, slightly raised. Mycelium

102 AUSTRALIAN FUNGI, II,

subcuticular, forming a pellicle of one layer of brown hyphae 5-7u thick, the cells up to
12u long, much branched, the septa not thickened and not much darker than the
remainder of the cell walls. No haustoria were found, nor further penetration of the
host. Thyriothecia scattered, rounded, black, convex, smooth, shining, subcuticular in
development, up to 150” diam.; upper wall convex, indistinctly radiate, opening by
central irregular pore, composed of cells similar to those of the mycelium; lower wall
hyaline to subhyaline, thin, of interwoven hyphae 2-8 thick, passing into the internal
ground tissue. At first the loculus is filled with a loose fibrous tissue of hyaline
“paraphyses” 2u thick, amongst which the asci develop; possibly these paraphyses are
also connected to the upper wall, though this could not be definitely proved from my
sections. The asci are suberect, directed towards the apical pore of the thyriothecium,
ripening in succession, ellipsoid, sessile, at first rather thick-walled, rounded at the
apex, 8-spored, up to 75 x 18u. Spores 2-3-seriate, oblong-clavulate, the ends obtusely
rounded, 1-septate, constricted, smooth, becoming brown, 20-22 x 8—-9u, the lower cell
slightly. narrower than the upper.

(72) STIGMATEA ORITIDIS Hansf., n. sp.

Plagulae epiphyllae, atrae, leves, rotundatae, usque ad 1 mm. diam. vel confluentes.
Mycelium subcuticulare, pelliculosum, irregulariter radiatum, ex hyphis dilute olivaceis,
ramosissimis, 8-12u cr., cellulis 5-10u longis, subinde etiam longitudinaliter vel oblique
1-septatis, septis incrassatis, atrobrunneis, compositum. Thyriothecia laxe dispersa,
rotundata, subcuticularia, levia, atra, usque ad 3504 diam., circa 1004 alt.; paries
superior unistratosus, opacus, indistincte radiatus, poro irregulariter stellato centrali
apertus; paries inferior hyalinus, haud radiatus, ex hyphis intertextis 2-3u crassis
compositus, tenuis. Asci versus poro thyriothecii directi, cylindracei, apice late
rotundati et usque ad 6 incrassati, 8-spori, deorsum breviter nodoso-stipitati,
60-90 x 18-21u. Paraphyses paucae, filiformes, hyalinae, circa 70 x 2-34. Sporae oblique
2-3-seriatae, oblongo-ovoideae, atrobrunneae, 1-septatae, leniter constrictae, saepe

4-cguttulatae, 23-27 x 8-9u, cellula superiore leniter crassiore, episporio tenuiter punctato,
haud verrucoso.

Hab. in foliis Oritidis lancifoliae, Bogong High Plains, Middle Creek, Victoria, Jan.
1950, leg. Mrs. Ducker.

Colonies epiphyllous, black, smooth, orbicular, up to 1 mm. diam., becoming larger
by confluence. Mycelium in or under the cuticle, eventually forming a solid plate of one
layer of pale olivaceous, much branched hyphae 8—12u wide, the cells 5-10u long, often
with one longitudinal or oblique septum in the wider cells, the septa thick and black-
brown. Thyriothecia subcuticular, loosely scattered, circular, Smooth, black, shining,
up to 3504 diam. and about 100u high in the centre, not fimbriate around the edge;
upper wall of one layer of opaque black hyphae indistinctly and irregularly radiating
from the centre, with an indistinct lining of much compressed hyaline delicate cells,
dehiscent by irregularly stellate fissures extending outwards from the centre, not
reaching the margin, and so appearing as a rather wide irregular central pore; basal
wall of hyaline, delicate, interwoven hyphae, not radiate in structure, thin. Asci fairly
numerous, ripening in succession, cylindric with broadiy rounded apex, which in
immature asci is thickened up to 6u, without evident interior canal, not definitely
bitunicate, suddenly narrowed below into a short nodose foot, 8-spored, 60-90 x 18—21yu.
A few paraphyses are present, hyaline, filiform, ? continuous, about 70 x 2-34. Spores
irregularly and obliquely 2—3-seriate, oblong-ovoid with obtuse rounded ends, dark brown
when fully mature, 1-septate, slightly constricted, often 4-guttulate, 23-27 x 8—9u, the

upper cell slightly wider than the lower; epispore finely and indistinctly punctate but not
roughened.

(73) MicrRoTHYRIUM AMYGDALINUM CkKe. & Mass., Grevillea, 19:90, 1890.

On leaves of Hucalyptus spp., South Australia and Tasmania; the original collection
was made around Spencers Gulf, S. Australia. The only recent collection I have seen

BY C. G. HANSFORD. 103

was made at Meningie, S. Austr., by Mr. L. D. Williams, and is preserved at the Waite
Institute as WARI 3519.

Free mycelium none. Thyriothecia superficial, gregarious or scattered, black-brown,
up to 180u diam., amphigenous, circular, flattened, membranaceous, easily secedent.
Upper wall of subopaque dark brown radiating hyphae 5—6u thick, the cells 5-14y long,
not fimbriate at the margin, irregularly dehiscent or with an irregular central aperture.
Lower wall pale olivaceous, of interwoven hyphae 3—4u wide, with short cells, thin. The
interior is at first filled with a loose fibrous tissue of branched, filiform, septate
paraphysoids, 1—2u thick, irregularly interwoven, arising from the lower wall and
doubtfully also connected to the upper wall. Through this loose tissue a few wider
hyphae (4-5u, with cells 4-74 long) ramify at the base, and the asci arise as erect
branches from them, apparently without crozier formation. Asci up to 50 x 18z,
ellipsoid-saccate, thickened up to 7u at the apex and with a central interior canal when
immature, later the wall becoming thinner and the canal disappearing; 8-spored at
first, but often only 2—4 spores mature; the base sessile or very shortly nodose-stipitate.
Spores irregularly arranged, oblong-fusoid with obtuse rounded ends, hyaline, 1-septate,
not or indistinctly constricted, straight, smooth, up to 18 x 6u.

No penetration of the host leaf by this fungus has yet been found by me, but as it
appears to be restricted to the host-genus Hucalyptus, it is probable that in some manner
it is a definite parasite of the leaf.

(74) SEYNESIA MICROTHYRIOIDES (Wint.) Theiss., Osterr. Bot. Zeitschr., 63:125, 1913.

= Asterina microthyrioides Wint., Hedgwigia, 25:94, 1885.

On Hucalyptus niphophila, Kosciusko, New South Wales, A. Costin 118, April 1947.

Thyriothecia closely scattered on indefinite brownish areas of the leaf, up to 20 mm.
diam., hypophyllous. There is a subcuticular layer of hyaline mycelium covering the
whole infected area, 1—2 cells thick, and connected at many points with the basal wall
of the thyriothecium, directly through the cuticle by fine hyaline filaments. The outer
cavities of the stomata beneath the thyriothecia are filled with dark plugs of mycelium,
but these do not penetrate between the guard cells, and apparently are not connected
with the subcuticular mycelium. Individual thyriothecia are rounded, up to 3004 diam.
and: 80-100u high in the centre, black, smooth, flattened-conoid with a central irregular
pore in the upper wall, about 20u diam.; superficial and often confluent into small
groups. There is no external free mycelium. Upper wall of thyriothecium consisting of
a single layer of radiating dark brown opaque hyphae 5-6u wide, not fimbriate at the
margin, lined with an indefinite layer of hyaline compressed parenchyma, to which the
“naraphyses” are connected. Lower wall of 1-3 layers of interwoven delicate hyphae, in
section appearing parenchymatous, closely adpressed to the cuticle of the host leaf,
passing internally in young stages into a tissue of erect, branched, septate, hyaline
hyphae, of which a few remain as “paraphyses” between the maturing asci. Asci
numerous, ripening in succession, at first erect, then directed towards the apical pore of
the thyriothecium, wide cylindric to saccate, widely rounded and when young thickened
at the apex to 5u, subsessile, 8-spored, up to 70 x 20u. Spores irregularly 2—3-seriate,
oblong-clavulate with obtuse rounded ends, straight or slightly bent, becoming brown,
1-septate and slightly constricted, smooth, 20-23 x 7—8u, the upper cell slightly shorter
and wider than the lower.

This species was also collected at Meningie, S. Australia, by L. D. Williams, WARI
3529, on Hucalyptus sp.

(75) SEYNESIA BANKSIAE P. Henn., Hedwigia 1903: 78.

The specimen on Banksia ornata, Kuitpo, S. Australia, collected by Samuel and |
mentioned in Trans. Roy. Soc. S. Australia, 46:175, 1922, has been re-examined.

Beneath the colonies there is a subcuticular mycelium of hyaline, septate, much
branched hyphae, connected to the basal wall of the superficial thyriothecia at many
points directly through the cuticle by fine filaments, and at other points penetrating

104 AUSTRALIAN FUNGI, II,

through the cuticle by similar filaments, enlarging above the cuticle into single brown
stellate cells or into groups of these. Around some colonies in this collection there are
a few superficial brown hyphae extending outwards over the leaf, up to 100 long,
septate, little and irregularly branched, and these hyphae bear alternate or unilateral
hyphopodia, which are globose to hemispheric, entire, 5-10 x 5u. An extreme example of
the development of this mycelial development is shown by a collection made by C. T.
White, in Feb. 1909 on Banksia latifolia, Moreton Island, Queensland, now in the
Queensland Herb.; until Samuel’s collection had been examined in detail, I had pro-
visionally determined the Queensland specimen as a new species “Asterina banksiae’’.

Here the colonies are epiphyllous, thin, effuse and confluent, smooth. Mycelium of
substraight, brown, main hyphae with cells 20-45 x 3-5-4-5u, bearing alternate or
unilateral hyphopodia, and with numerous wavy to very crooked, irregularly branched,
often fasciculate, secondary hyphae without hyphopodia. These secondary hyphae are
very irregular, concolorous or slightly paler than the main hyphae, indistinctly septate,
2-5-4-5u wide, forming a rather close, patchy reticulum. Hyphopodia continuous,
subglobose to ovate, sessile, entire, 5-8 x 5u. Thyriothecia few, loosely scattered, circular,
convex, black, up to 170% diam., not fimbriate at the margin; upper wall subopaque
black-brown, composed of radiating hyphae 3-5—-4-54 wide, the cells up to 12u long,
splitting radiately to the margin at maturity into a few widely triangular segments, the
fissures originating from a central irregular pore about 30u diam. Lower wall indistinct,
subhyaline, not radiate, very thin. Asci rather few, apparently aparaphysate, subglobose
to widely ellipsoid, sessile or very shortly nodose-stipitate, 8-spored, 30-40 x 20-28yu.
Spores conglobate, oblong with rounded ends, 1-septate, slightly constricted, smooth,
13-16 x 6-7u, the cells subequal. Germination is by formation of a hyphopodium near
one end, followed by mycelial growth from the same or from the opposite cell.

I am unable to distinguish generically between these two specimens, which in my
opinion rightly belong to the genus Asterina, but still have to re-examine authentic
collections of Seynesia banksiae P. Henn., before it can be definitely stated that this is
identical.

(76) THALLOCHAETE BAILEYI (B. & Br.) Hansf., comb. n.
= Asterina baileyi B. & Br. = Asterella hakeae McAlpine.

Colonies on leaves of Hakea spp., rounded to elongate, black, at first thin, but
becoming dense and very shortly velvety. External mycelium of flexuous dark brown
hyphae 5—7u thick, the cells 10-30u long, very irregularly branched and forming minute
knots and tufts, which later coalesce; hyphopodia none. Mycelial setae more or less
erect, straight or bent, simple, obtuse, dark brown, septate, up to 70 x 5-7u, the basal
cell slightly inflated, densely scattered on the external hyphae in most collections,
though in others scant or even absent, especially in young colonies. The surface
mycelium is connected at short intervals to other hyaline or pale brown hyphae within
the cuticle, often in two or three indefinite layers, the outermost about 15-20u deep
within the cuticle, and the innermost in close contact with the cells of the epidermis,
mostly running along the grooves between these cells, not penetrating deeper within the
leaf and not forming haustoria. The outermost inflated stomatal cavity is often filled
with dark plugs of mycelium, which may connect directly with the intracuticular hyphae,
but do not penetrate down to, or between, the guard cells.

Thyriothecia closely scattered, and later often becoming confluent at the edges, black,
circular in outline, glabrous or sometimes partially covered with mycelial hyphae
bearing setae, up to 280u diam., in maturity about 90u high in the centre; the upper
wall opaque dark brown, the outer layer of radiating hyphae 4-64 wide, with cells up to
15u long, not fimbriate at the margin, with 2—4 inner layers of thin-walled, paler cells,
much compressed and connected internally to the loose fibrous tissue of ‘paraphyses’’,
which at full maturity break from this upper connection by gelatinization, and thus
become free filaments. The upper wall is dehiscent by an irregular, somewhat stellate

BY C. G. HANSFORD. 105

central aperture. Lower wall of thyriothecium hyaline or subhyaline, of loosely inter-
woven hyphae 13-2u thick, somewhat gelatinous, closely adpressed to the host cuticle.
Asci basal, developing singly in succession and forcing their way through the loose
tissue of paraphyses towards the central aperture of the upper wall, broad cylindric,
becoming ovate, subsessile, up to 55 x 30u, thin-walled, 8-spored. Spores conglobate,
oblong-clavulate with rounded ends, dark brown with transverse lighter band on each
cell, 1-septate, constricted, smooth, 20-24 x 10-12u, the upper cell broader than the
lower and subglobose. Paraphyses filiform, septate, hyaline, sometimes furcate, apparently
formed from the “ground tissue’ of the thyriothecium.

On Hakea dactyloides, Sydney, 1869, leg. Maiden (type of Asterella hakeae McAlp.),
in Herb. Dept. Agric., Victoria; on Hakea leucoptera, Ooldea, Aug. 1922, T. G. Osborn;
on H. vittata, Port Lincoln, July 1952, N. T. Flentje; on HA. elliptica, King George’s
Sound, W. Australia, Morrison 34, Dec. 1898, in Herb. Kew; on Hakea spp., Bellair
National Park, S. Australia, L. D. Williams, J. Brown, etec., in Herb. Waite Institute.

This fungus appears to be widely distributed in Australia and fairly common on
species of Hakea. Under a lens there is considerable diversity in colony appearance
between the collections included above, but I am unable to separate them into species
or even varieties.

(77) LEMBOSIOPSIS AUSTRALIENSE Hansf., n. sp.

Maculae amphigenae, usque ad 10 mm. diam. vel confluentes, margine zono atro- vel
rufo-brunneo, 1-13 mm. er. circumdatae, centro griseo-brunneae, subzonatae, epidermide
folii per fissa irregulares disrumpente. Mycelium internum ex hyphis dilute brunneis,
ramosis, septatis, intercellularibus, 3u cr. compositum, mesophyllum totum penetrans,
haustoria nulla. Mycelium externum nullum. Thyriothecia superficialia, saepius
irregulariter circinatim disposita, vel dispersa, atra, linearia vel X-Y-formia, usque ad
1000 x 100-150, convexa, levia, longitudinaliter dehiscentia, demum late aperta; paries
superior opace niger, ex hyphis radianto-parallelibus, 3-44 crassis compositus, margine
leniter fimbriatus; paries inferior subopace atrobrunneus, circa 30y cr., ex hyphis inter-
textis compositus, intus in subhymenio pallidiore transeuns. Asci numerosi, erecti,
clavati, circa 40 x 10-124, sursum late rotundati, subsessiles, 8-spori. Sporae 2—3-seriatae,
parallelae, oblongae utrinque rotundatae, 1-septatae, lenissime constrictae, hyalinae, leves,
10-13 x 3-5-4-5u, cellulis subaequalibus. Paraphyses numerosae, filiformes, simplices,
ip cr., ascos leniter superantes et epithecium tenuum viridum formantes.

Hab. in foliis Hucalypti cosmophyllae, Cape Jervis Peninsula, South Australia, Jan.
1924, WARI 2070, leg. G. Samuel (typus); in foliis H. marginatae, Kuitpo, S. Australia,
G. Samuel; in foliis H. spec. indet., Cape Jervis Peninsula, WARI 2067, leg. G. Samuel;
Aldgate, S. Australia, Oct. 1912, WARI 2097, leg. T. G. Osborn.

Leafspots amphigenous, on either side of the leaf or penetrating through to both
surfaces, up to 10 mm. diam. or confluent and irregular, surrounded by a dark red-brown
marginal zone 1-1-5 mm. wide, the centre dark greyish-brown and somewhat zonate,
with one or two broken rings of black thyriothecia, and the epidermis soon breaking up
by irregular fissures. The mycelium penetrates the whole mesophyll of the older spots,
but is more limited when these are restricted to one side of the leaf; hyphae brownish,
septate, intercellular, about 3u thick, much branched, very difficult to observe by reason
of the discoloured dead host cells filled with red-brown amorphous matter. There is no
superficial mycelium, apart from the very slightly fimbriate margins of the thyriothecia,
thus differing from L. eucalyptina Petr. & Syd., which occurs on Hucalyptus spp. in
South Africa. The internal mycelium passes out between the epidermal cells, and may
in places form a slight subcuticular aggregation, or this may be absent, the hyphae then
passing direct through the cuticle to contact with the superficial thyriothecia at several
points in their lower surfaces. Thyriothecia usually irregularly circinate, with one or
two broken rings on each leafspot, or sometimes irregularly scattered towards the centre
of the spot, linear, straight or bent, or X- or Y-shaped, up to 1000 x 100-1504 convex,
black, smooth, longitudinally dehiscent by a wide cleft. Upper wall opaque black, of

I

106 AUSTRALIAN FUNGI, II,

radiating-parallel hyphae 3-44 thick, the margin slightly fimbriate; lower wall opaque
dark brown, about 30u thick below the loculus, of interwoven dark hyphae, passing
above into a lighter subhymenial layer. Asci numerous, erect, clavate, about 40 x 10-13y,
broadly rounded above, tapering to a subsessile base, 8-spored. Spores 2—3-seriate and
parallel, oblong, rounded at both ends, 1-septate in middle and sometimes very slightly
constricted, hyaline, smooth, 10-13 x 34—434u, the cells approximately equal. Paraphyses
numerous, filiform, simple, about lu thick, slightly exceeding the asci and forming a thin
greenish epithecium.

The main point of difference from L. euwcalyptina Petr. & Syd. is the absence of any
external mycelium; no conidial stage was found.

(78) ASTEROLIBERTIA CRYPTOCARYAE (Cooke) Hansf., n. comb.

= Asterina cryptocaryae Cooke in Herb. Kew. = Asterinella cryptocaryae (Cooke)
Theiss., Broteria, 10:107, 1912. = Prilleuxina cryptocaryae (Cooke) Ryan, Illinois Biol.
Monogr., 17:79, 1939.

On Cryptocarya meissneri, Comboyne, New South Wales, L. Fraser, Jan. 1935.

Colonies amphigenous, very thin and scarcely visible, up to 5 mm. diam., smooth.
Mycelium of substraight or finely sinuous, dark brown hyphae 4—5y thick, the cells
15-25u long, sparingly and irregularly branched, very loosely reticulate, quite superficial
and devoid of hyphopodia or of specialized ‘‘node cells’, but some cells forming
epidermal haustoria and marked with minute pores in their lower surface next the
leaf, where the filament penetrating into the cuticle is attached. Thyriothecia loosely
scattered, circular, flattened convex, up to 3004 diam., black, smooth; margin crenate,
not fimbriate; the upper wall of radiating dark brown hyphae with cells 8-12 x 4un,
splitting radiately at maturity and the centre falling away. Lower wall indistinct,
hyaline, passing into the fibrous internal tissue of branched, hyaline, much interwoven
hyphae 3u thick, simulating paraphyses. Asci fairly numerous, ripening in succession,
borne as lateral branches of ascogenous hyphae ramifying through the internal fibrous
tissue, erect, wide ellipsoid, broadly rounded at the apex, suddenly contracted below into
a short, wide stipe, which is often bent or almost lateral, 8-spored, aparaphysate, up to
100 x 35u. Spores multiseriate, more or less parallel, clavulate-oblong with obtusely
rounded ends, 1-septate, slightly constricted, smooth, becoming uniformly dark brown,
25-32 x 11-13y, the lower cell slightly smaller than the upper. Germination occurs at
the base of the lower cell, direct into mycelial hyphae, the first cell formed producing a
haustorium in the epidermis. No conidial stage found.

(79) ASTEROLIBERTIA CRYPTOCARYAE Var. NODULIFERA Hansf., n. var.

Plagulae plerumque epiphyllae, tenues, usque ad 3 mm. diam. vel late confluentes,
leves. Mycelium ex hyphis atrobrunneis, 4—5u ecr., cellulis 15-20u longis, opposite
acuteque ramosis, laxe reticulatis compositum. Hyphopodia intercalaria, saepe cellulis
mycelii regulariter alternantia, tumulos globosos minutos sub hyphis mycelii efformantia,
8-104 diam. Thyriothecia laxe dispersa, atra, rotundata, usque ad 400u diam., margine
crenata, haud fimbriata; parietes et structura interior illis typi consimiles. Sporae
atrobrunneae, oblongo-clavulatae utrinque rotundatae, leves, 1-septatae, leniter constrictae,
30-36 x 11-14y.

Hab. in foliis Cryptocaryae rigidae, Bulga, New South Wales, L. Fraser 61 (typus) ;
in foliis C. patentinervis, Williams, R., N.S.W., L. Fraser, April 1952.

Colonies mainly epiphyllous, thin, to 3 mm. diam. or confluent over the leaf, smooth.
Mycelium of substraight dark brown hyphae 4—5u thick, the cells mostly 15-20 long,
branching usually opposite at rather acute angles, loosely reticulate. Hyphopodia
intercalary, often regularly alternating with normal mycelial cells, appearing as a
globose swelling on the lower side of the parent cell and not cut off by a septum, 8—-10u
diam. Thyriothecia loosely scattered, black, circular, up to 400u diam., the margin
crenate, not fimbriate; upper wall convex, of dark brown radiating hyphae, the cells up
to 15 x 4-5u, dehiscent by a central irregular pore about 20u diam., from which a few

BY C. G. HANSFORD. 107

stellate fissures later radiate, the centre finally secedent. Lower wall indistinct, hyaline,
_ passing into a loose tissue of branched hyaline hyphae filling the interior, in which the
asci are formed, and which degenerates at maturity. Asci widely ovate, sessile, 8-spored,
aparaphysate, up to 100 x 45u, ripening in succession. Spores multiseriate and somewhat
parallel in ascus, becoming dark brown and sometimes with a transverse lighter band
around one or both cells, smooth, oblong-clavulate, the ends obtusely rounded, 1-septate,
slightly constricted, 30-36 x 11-144. Germination is by the formation of a hyphopodium
at the basal end of the spore, then by lateral growth of mycelium from this cell.

The most obvious difference from the type is in the appearance of the hyphopodia.

(80) ASTERINA FIELDIAE Hansf., n. sp.

Plagulae epiphyllae, tenuissimae, usque ad 2 mm. diam., leves. Mycelium ex hyphis
undulatis vel flexuosis, brunneis, 3—4u crassis, indistincte septatis, laxe reticulatis
radiantibus compositum. Hyphopodia alternata vel unilateralia, continua, coralloideo-
pulvinata, 6-10u diam. Thyriothecia laxe dispersa, atrobrunnea, convexa, usque ad 100u
diam., subinde 2-connata, margine haud fimbriata; paries superior radiatus, atrobrunneus,,
cellulis 4-8 x 4u, in segmentis angustis stellatim dehiscens. Asci pauci, aparaphysati,
globosi vel ellipsoidei, 8-spori, sessiles, 22-28 x 20-25u. Sporae brunnescentes, oblongae
utrinque rotundatae, 1-septatae, fortiter constrictae, cellulis subglobosis, subaequalibus,.
episporio levi vel subtiliter granuloso, 12-15 x 7-8u. Conidia non visa.

Hab. in foliis Fieldiae australis, Mt. Wilson, New South Wales, June 1932, L. Fraser 5
(typus); Blackheath, N.S.W., June 1935, L. Fraser 210.

Colonies epiphyllous, mixed with those of Irenopsis fieldiae Hansf., very thin, up to
1 mm. diam. Mycelium of wavy to crooked brown hyphae 3-44 thick, indistinctly
septate, irregularly branched, loosely radiating-reticulate. Hyphopodia alternate or
unilateral, continuous, coralloid to pulvinate with wavy outline, 6-104 diam. Thyrio--
thecia loosely scattered, dark brown, convex, up to 100u diam., usually separate but
sometimes 2-connate, not fimbriate at the margin; upper wall of radiating dark brown
hyphae with cells 4-8 x 4u, splitting radiately at maturity into narrowly triangular-
segments to the margin. Asci few, aparaphysate, globose to ellipsoid, 8-spored, sessile,
22-28 x 20-25u. Spores conglobate, becoming brown, oblong with broadly rounded ends,
1-septate and rather deeply constricted, the cells subglobose and equal, the surface from
nearly smooth to finely granulose, 12-15 x 7-84. The lower wall of the ascoma is.
hyaline, of indistinct structure. Pycnidia and pycnospores not seen.

(81) ASTERINA CLEMATIDIS Hansf., n. sp.

Plagulae amphigenae, usque ad 0-5 mm. diam., saepius numerosae et confluentes;.
subdensae, leves. Mycelium ex hyphis flexuosis, brunneis, 3-5—5u er., cellulis 15-20u
longis, opposite vel irregulariter ramosis, subdense reticulatis compositum. Hyphopodia
alternata vel opposita, 1-septata, 9-124 longa, recta vel curvata, cellula basali cylindracea,
2-4u longa, cellula apicali subglobosa vel anguloso-sublobata, apice saepe truncata,
8-9 x 4-8u. Thyriothecia subaggregata, rotundata, nigra, convexa, levia, usque ad 100u
diam.; paries superior radiatus, atrobrunneus, cellulis 5-10 x 3-4u, margine crenatus
vel leniter fimbriatus; paries inferior hyalinus, indistinctus; in maturitate prope
marginem in segmentis angustis stellatim dehiscentia. Asci circa 8, aparaphysati, globosi
vel ovati, sessiles, 8-spori, circa 30 x 20-25u. Sporae conglobatae, atrobrunneae, oblongae
utrinque obtusae, 1-septatae, leves, 15-17 x 7—8u, cellulis aequalibus.

Pyenidia thyriotheciis consimilia; pycnosporae subglobosae vel _ piriformes,
atrobrunneae, transverse hyalino-zonatae, leves, 13-15 x 7—9u.

Hab. in foliis Clematidis glycinoidis, National Park, New South Wales, L. Fraser 27
(typus), 176.

Colonies amphigenous, minute, to 0-5 mm. diam., usually numerous and widely
confluent, rather dense, smooth. Mycelium of flexuous brown hyphae, 3-5-5 thick, the
cells mostly 15-204 long, branching opposite or alternate, rather closely reticulate—
radiating. Hyphopodia alternate or opposite, 2-celled, 9-124 long, straight or bent:

108 AUSTRALIAN FUNGI, II,

stalk cell cylindric, 2-44 long; head cell from subglobose to angulose or sublobate,
often truncate at the apex, 8-9 x 4-84. Thyriothecia closely scattered but usually
separate, circular, black, convex, smooth, up to 1004 diam., the margin crenate or
slightly fimbriate; upper wall of dark brown radiating hyphae, the cells 5-10 x 3—-4y,
dehiscent by numerous radiate fissures nearly to the margin into narrow triangular
segments. Asci about 6-8, aparaphysate, globose to widely ovate, sessile, 8-spored, about
30 x 20-25u. Spores conglobate, dark brown, oblong with obtusely rounded ends, 1-septate
and constricted in the middle, the cells equal and subglobose, smooth, 15-17 x 7-8y.

Pyenidia similar to the thyriothecia; pycnospores subglobose to piriform, dark
brown with transverse median subhyaline band, from which germination occurs, smooth,
13-15 x 7-9u.

{82) ASTERINA DENSA Syd. var. AUSTRALIENSIS Hansf., n. var.

Plagulae hypophyllae, tenues vel densae, usque ad 5 mm. diam., saepe late con-
fluentes. Mycelium ex hyphis atrobrunneis, subrectis vel flexuosis, 4-5 cr., cellulis
plerumque 20-304 longis, irregulariter ramosis, reticulatis compositum. Hyphopodia
alternata vel laxe dispersa, haud opposita, breviter digitata integraque vel irregulariter
sinuoso-curvata, rarius lobata, versiformia, 6-13 x 4-9u, continua, rarius in ramulis
mycelii terminalia. Thyriothecia subaggregata, plerumque discreta, rotundata, usque ad
200u diam., nigra, convexa, levia, margine haud vel leniter fimbriata, hyphis fimbriarum
usque ad 60u longis, tortuoso-radiantibus; paries superior radiatus, atrobrunneus, cellulis
4-12 x 4u, in maturitate stellatim ad marginem dehiscens et parte centrali secedente;
paries inferior dilute brunneus vel subhyalinus, radiatus, tenuis. Asci circa 6,
aparaphysati, globosi vel late ovati, sessiles, 4—8-spori, 40-50u diam. Sporae conglobatae,
oblongae utringue rotundatae, 1-septatae, fortiter constrictae, 23-28 x 12-14u, cellula
superiore crassiore, subglobosa, episporio sublevi vel dense subtiliter verruculoso.
Pyenidia non visa.

Hab. in foliis Pittospori undulati, Mt. Warning, New South Wales, L. Fraser 231
{typus); Williams R., N.S.W., L. Fraser 78; Cambewarra Mt., N.S.W., Rodway, Dec. 1934.

Colonies hypophyllous, thin, becoming dense, up to 5 mm. diam., often widely
confluent. Mycelium of dark brown, substraight to crooked hyphae 4—5u thick, the cells
mostly 20-304 long, irregularly branched, becoming closely reticulate. Hyphopodia
alternate or more scattered, not opposite, continuous, from short digitate and entire
to irregularly sinuous-bent or lobed, versiform, 6-13 x 5—9u, rarely formed at the ends
of short mycelial branches. Thyriothecia closely scattered but usually separate, roughly
circular, up to 200 diam., black, convex, smooth, the margin not, or slightly fimbriate
and then the fringing hyphae tortuous-radiating, up to 60u long. Upper wall of radiating
dark brown hyphae with cells 4-12 x 4u, splitting radiately to the margin into
triangular segments and the central parts soon secedent. Lower wall of pale brownish
to subhyaline radiating hyphae, irregular, thin. Asci about 6, aparaphysate, ripening in
‘succession, globose to oblong, 4—-8-spored, about 40-50u diam. Spores conglobate, oblong
with obtusely rounded ends, 1-septate and rather deeply constricted, the upper cell
usually subglobose and slightly wider than the lower, surface from nearly smooth to
closely and finely verruculose, 23-28 x 12-14u, the lower cell 10-124 wide. Pyecnidia not
seen.

(83) ASTERINA DICTYOLOMATIS P. Henn., Hedwigia 43:372, 1904.

On Hvodia micrococca, Williams R., New South Wales, L. Fraser 211.

Colonies epiphyllous, rarely hypophyllous and then small, up to 3 mm. diam. or
numerous and widely confluent, thin, smooth. Mycelium of substraight to flexuous
brown hyphae 4—5y thick, the cells mostly 15-20u long, branching opposite or irregular,
loosely reticulate. Hyphopodia alternate or unilateral, short digitate or slightly lobate,
continuous, straight or bent, rounded at apex, 6-10 x 5-84. Thyriothecia numerous,
closely scattered, rounded, black, convex, up to 140u diam., the margin loosely fimbriate
with the fringing hyphae tortuous-radiating, up to 100u long; upper wall almost

BY C. G@. HANSFORD. 109

hemispheric, dark brown, of radiating hyphae with cells 4-12 x 4u, at maturity radiate-
dehiscent almost to the margin into narrow triangular segments; lower wall of sub-
hyaline, indistinctly radiating, rather loose hyphae. Asci aparaphysate, globose to
widely ovate, sessile, 8-spored, up to 40” diam., maturing in succession and filling the
cavity of the thyriothecium. Spores conglobate, becoming dark brown, oblong with
obtuse ends, 1-septate, rather deeply constricted, the cells equal and subglobose, or the
upper very slightly wider than the lower, epispore smooth or very finely and closely
granulose to verruculose, 18-22 x 9-10u. Pyenidia and pycnospores not seen.

(84) ASTERINA DORYPHORAE Hansf., n. sp.

Plagulae amphigenae, usque ad 5 mm. diam., vel effusae et late confluentes, tenues,
leves. Mycelium ex hyphis subrectis vel flexuosis, atrobrunneis, 6—7u ecr., cellulis.
plerumque circa 16 longis, irregulariter ramosis, reticulatis compositum. Hyphopodia.
alternata, unilateralia vel rarius opposita, continua, hemisphaerica, integra, 8-10” diam.
Thyriothecia dispersa, atra, convexa, levia, rotundata, haud fimbriata, usque ad 260.
diam.; paries superior ex hyphis sinuosi-radiantibus, pellucide atrobrunneis compositus,
cellulis 5-12 x 4-6u, mox stellatim prope marginem dehiscens; paries inferior hyalinus,
ex hyphis tortuoso-radiantibus, 3-5u cr. compositus, compressus et cuticulam folii
adhaerens. Asci numerosi, in ordine maturescentes, subglobosi, sessiles, 4—8-spori,
usque ad 60 xX 40u; paraphyses numerosae, hyalinae, filiformes, obsolete septatae et
ramosae, mucoso-difluentes. Sporae conglobatae, oblongae utrinque obtuse rotundatae,
1-septatae, constrictae, dilute brunneae, leves, 30-33 x 14-16u, cellulis subaequalibus.

Hab. in foliis Doryphorae sassafratis, National Park, New South Wales, L. Fraser 26
(typus); Clyde Mtn., N.S.W., L. Fraser 180.

Colonies amphigenous, up to 5 mm. diam. or effuse and widely confluent over the
leaf, thin, smooth, becoming denser. Mycelium of substraight to flexuous dark brown
hyphae 6-7u thick, the cells mostly about 15u long, branching irregular, becoming
closely reticulate. Hyphopodia alternate, unilateral or rarely opposite, hemispheric,
entire, 1-celled, 8-10u diam. Thyriothecia scattered, black, convex, smooth, more or less
circular in outline, not fimbriate at margin, up to 260u diam.; upper wall of translucent.
dark brown radiating-wavy hyphae with cells 5-12 x 4-6u, early splitting by radiate
fissures into many narrow triangular segments nearly to the margin, and exposing the
internal tissue of asci and paraphyses; lower wall hyaline, of tortuous hyphae 3-5«
thick, flattened and adherent to the host cuticle. Asci numerous, ripening in succession,
subglobose, sessile, 4—8-spored, up to 60 x 40u; paraphyses numerous, hyaline, filiform,
indistinctly septate and branched, becoming mucose. Ascus wall at first thickened,
without evident apical canal. Spores conglobate, oblong with rounded ends, 1-septate,.
constricted, pale brown, smooth, 30-33 x 14-16u, the cells subequal or the upper slightly
larger. In germination the spore forms a hyphopodium at one end, and the opposite:
cell collapses, while the germinating cell becomes dark brown. The mycelium is closely
similar to that of Clypeolella and Schiffnerula, but no trace of mycelial conidia was.
found.

(85) ASTERINA DRIMIDICOLA Hansf., n. sp.

Plagulae plerumque hypophyllae, tenues, usque ad 5 mm. diam. vel effusae et
confluentes, leves. Mycelium ex hyphis brunneis, subrectis vel flexuosis, 4-5-5-5u er.,
cellulis plerumque 20-30 longis, irregulariter ramosis, laxe reticulatis compositum.
Hyphopodia alternata, opposita vel dispersa, continua, breviter digitata vel sublobata,.
7-10 x 6-84. Thyriothecia dispersa, atra, suborbiculata, usque ad 200u diam., subinde
2-connata; paries superior atrobrunneus, radiatus, cellulis 4-10 x 4u, stellatim prope
marginem dehiscens et parte centrali secedente; paries inferior indistinctus, hyalinus..
Asci circa 6, aparaphysati, ovati vel subglobosi, sessiles, 8-spori, usque ad 60 x 35-40.
Sporae conglobatae, oblongae utrinque. rotundatae, i1-septatae, constrictae, opace
atrobrunneae, leves, 26-29 x 14-15u, cellulis aequalibus. Conidia nulla.

Hab. in foliis Drimidis insipidae, Hastings R., New South Wales, L. Fraser, April.
1952.

110 AUSTRALIAN FUNGI, IT,

Colonies mostly hypophyllous, thin, to 5 mm. diam. or effuse and widely confluent,
smooth. Mycelium of substraight to flexuous dark brown hyphae 4-5—-5-5u thick, the
cells mostly 20-30u long, branching irregular, loosely reticulate. Hyphopodia alternate,
opposite or scattered, continuous, short digitate or sublobate, 7-10 = 6-8. Thyriothecia
seattered, black, convex, circular, the margin crenate, not fimbriate, up to 200 diam.,
sometimes 2-connate at the edges; upper wall dark brown, of radiating hyphae with cells
4-10 x 4u, stellate-dehiscent to the margin into broad triangular segments, the centre
soon secedent. Lower wall indistinct, hyaline. Asci about 6, apparently aparaphysate,
ovate to subglobose, sessile, 8-spored, up to 60 x 35-40u. Spores conglobate, oblong with
rounded ends, 1-septate, constricted, opaque dark brown, smooth, 26—29 x 14—-15y, the cells
equal. No conidia seen.

(86) ASTERINA KNYSNAP Doidge, var. AUSTRALIENSIS Hansf., n. var.

Plagulae epiphyllae, usque ad 4 mm. diam., densae, leves, facile secedentes et
maculas luteas relinquentes. Mycelium ex hyphis subrectis atrobrunneis, 6—7u ecr.,
cellulis plerumque 15-20u longis, irregulariter ramosis, primo laxe reticulatis, com-
positum, demum subsolidum. Hyphopodia alternata, unilateralia vel circa 5% opposita,
plerumque leniter antrorsa, continua, ellipsoidea vel subglobosa, integra, 7-12 x 6-8y.
Thyriothecia subaggregata, rotundata, atra, convexa, levia, usque ad 2004 diam., subinde
2-connata et elliptica; paries superior atrobrunneus, radiatus, cellulis 5-12 x 4-6y,
margine in laminis hypharum tortuoso-radiantibus extensus, haud fimbriatus; paries
inferior subhyalinus vel dilute brunneus, ex hyphis tortuosis 2-34 cr., septatis, com-
positus, haud radiatus. Thyriothecia stellatim dehiscentia. Asci subglobosi vel late
ovati, aparaphysati, pauci, 8-spori, usque ad 60 x 40u, sessiles. Sporae multiseriatae vel
conglobatae, opace atrobrunneae, oblongae utrinque obtusae, i1-septatae, constrictae,
28-35 x 14-15u, cellulis aequalibus, subglobosis, episporio subtiliter denseque granuloso.
Conidia nulla.

Hab. in foliis Canthii coprosmoidis, Hastings R., New South Wales, L. Fraser, April
1952.

Colonies epiphyllous, to 4 mm. diam., dense, smooth, easily secedent and leaving a
pale brown leafspot. Mycelium of substraight dark brown hyphae 6—7y thick, the cells
mostly .15-20u long, branching irregular, forming at first a loose network of hyphae,
which becomes almost solid in mature colonies by further branching and by development
of thyriothecia. Hyphopodia alternate, unilateral or about 5% opposite, often slightly
antrorse, continuous, short ellipsoid to subglobose, entire, 7-12 x 6-84. Thyriothecia at
first closely scattered, rounded, black, convex, smooth, up to 200u diam., soon extending
to form a continuous sheet beneath the mycelium, sometimes 2-connate and then elliptic;
upper wall of substraight dark brown radiating hyphae 4—6u thick, the cells 5—12u long,
passing at the margin into sheets of tortuous-radiating hyphae, not fimbriate. Lower
wall of tortuous pale brown to subhyaline hyphae 2-3 thick, septate, not radiate.
‘Young thyriothecia are filled with a loose tissue of much branched, septate, hyaline
hyphae, amongst which the few asci develop. Asci subglobose to wide ovate, aparaphysate,
8-spored, up to 60 x 40n, at first thickened to 6u around the widely rounded apex, sessile.
Spores multiseriate or conglobate, opaque dark brown, oblong, obtuse, 1-septate, con-
stricted, 28-35 x 14-15u, the cells equal and subglobose, epispore finely and closely
granulose. Conidia not seen. The thyriothecia are dehiscent by stellate fractures, the
central parts falling away.

(87) ASTERINA ORITIS Hansf., n. sp.

Plagulae hypophyllae, usque ad 4 mm. diam., tenues, aegre perspicues, leves.
Mycelium ex hyphis brunneis, flexuosis, 4u cr., cellulis circa 254 longis, irregulariter
ramosis, laxe reticulatis compositum. Hyphopodia alternata vel dispersa, digitata, apice
rotundata, plus minusve sinuoso-curvata, integra, 8-13 x 5-6y. Thyriothecia laxe
dispersa, atra, convexa, orbiculata, usque ad 200u diam., margine leniter fimbriata,
hyphis fimbriarum tortuoso-radiantibus usque ad 50u longis. Paries superior
atrobrunneus, radiatus, cellulis 5-12 x 4u, stellatim dehiscens et parte centrali secedente.

BY ©. G. HANSFORD. — 111

Paries inferior viridulus, ex hyphis sinuoso-radiantibus 2-5-3-5u4 cr. compositus. Asci
usque ad 15, in ordine maturescentes, aparaphysati, in massa fibrosa hyalina immersi,
erecti, late ellipsoidei, sessiles, usque ad 60 x 30y, 8-spori. Sporae conglobatae,
atrobrunneae, ellipsoideae utrinque obtuse rotundatae, 1-septatae, leves, 20-24 x 10-12n,
cellula inferiore distinete minore. Conidia non visa.

Hab. in foliis Oritis excelsae, Williams R., New South Wales, May 1953, L. Fraser 109.

Colonies hypophyllous, to 4 mm. diam., very thin and scarcely visible, smooth.
Mycelium of flexuous to crooked brown hyphae 4 thick, the cells about 25u long,
branching irregular, loosely reticulate. Hyphopodia alternate, unilateral or scattered,
digitate with rounded apex, more or less sinuous-bent, continuous, 8-13 x 5-6u. Thyrio-
thecia loosely scattered, black, convex, circular, up to 200% diam., the margin slightly
fimbriate, the fringing hyphae tortuous-radiating, up to 50u long; upper wall of dark
brown radiating hyphae, the cells up to 12 x 4u, at maturity stellate-dehiscent into broad
triangular segments and finally the centre falling away; lower wall greenish hyaline,
of wavy radiating hyphae 2:5-3:5u wide. Asci up to 15, ripening in succession, embedded
in an indistinctly fibrous tissue of hyaline hyphae, without distinct paraphyses, erect,
broadly ellipsoid, sessile, up to 60 x 30u, 8-spored. Spores conglobate, becoming dark
brown, ellipsoid with obtuse ends, 1-septate below the middle, smooth, 20-24 x 10-12u,
the lower cell distinctly smaller than the upper. Germination not observed. No
pycnidial stage seen.

(88) ASTERINA CINNAMOMICOLA Hansf., Proc. Linn. Soc. London, 157:202, 1945.

On Cinnamomum sp., Russell R., NSW., 1892, Von Mueller, in Herb. National,
Victoria.

This specimen corresponds closely to the type, described on Cinnamomum from
Ceylon.

(89) ASTERINA SP.

On Veronica derwentia, Kinglake, Victoria, 1930, leg. D. Dixon.

Colonies epiphyllous, black, thin, smooth, confluent over the whole leaf. Mycelium
of brown hyphae 4—6u thick, undulate to irregularly flexuous, septate at intervals of
20-30u, branching irregular, forming a loose network of wavy-polygonal meshes. Hypho-
podia alternate or unilateral, from pulvinate to elongate digitate, more or less lobed and
irregularly bent forwards or backwards, rarely with a stalk cell, 8-12 x 6-9u. Setae none.
Thyriothecia closely scattered, numerous, more or less filling the meshes of the mycelium
towards the centre of the colony, commencing as a radiate plate of brown hyphae about
5u thick, the cells towards the centre nearly cubical, those of the margin up to 15y long,
margin crenate, not fimbriate; mature thyriothecia up to 85u diam., irregularly circular
in outline, subopaque; basal membrane indistinct, hyaline; upper wall opaque dark
brown, irregularly dehiscent and the remains becoming more or less erect around the
asci, which are embedded in mucus derived from the hyaline “paraphysoids”. Asci 3-8,
subglobose, 8-spored, 25-30u diam. Spores conglobate, oblong with rounded ends, brown,
smooth, 1-septate, constricted, the cells nearly equal or sometimes the lower slightly
longer and narrower, 12-15 x 5-7u. Pycnidia similar to the thyriothecia, stellate-
dehiscent, rather few. Conidia oblong with rounded apex, very slightly apiculate at the
base, continuous, dark brown, smooth, 12-16 x 7—8u, with a transverse median hyaline
band.

Several species of Asterina have been described on Veronica spp. in the past, from
various parts of the world, and until this Australian specimen can be compared with
authentic material of these, it seems best not to give it a specific name.

(90) LEMBOSIA NOTELAEAE Hansf., n. sp.

Plagulae plerumque epiphyllae, tenues, atrae, usque ad 4 mm. diam. Mycelium ex
hyphis brunneis, subrectis vel flexuosis, 2—4u cr., obsolete septatis, irregulariter ramosis,
subdense reticulatis compositum. Hyphopodia in hyphis primariis efformantia, tenuiter
dispersa, alternata vel unilateralia, hemisphaerica vel pulvinata, continua, 4-7 x 6—-8y.

112 AUSTRALIAN FUNGI, II,

Thyriothecia in centro plagularum aggregata, linearia, usque ad 700 x 150u, saepe
confluentes et subinde Y-formia, atra, convexa, levia, margine irregulariter crenata, haud
fimbriata; paries superior opace atrobrunneus, ex hyphis radiantibus 3-44 cr. com-
positus, in maturitate longitudinaliter rimosus; paries inferior hyalinus, fibrosus,
tenuissimus et in texto centrali hypharum hyalinarum transeuns. Asci numerosi, erecti,
sessiles, late ellipsoidei, usque ad 40 x 20u, 8-Spori, aparaphysati. Sporae conglobatae,
brunnescentes, oblongae utrinque obtuse rotundatae, l1-septatae, leniter constrictae,
leves, 18-14 x 5—-7u, cellulis aequalibus. Conidia nulla.

Hab. in foliis Notelaeae venosae, National Park, New South Wales, L. Fraser, 13D
(typus); in foliis N. reticulatae, Church Point, N.S.W., L. Fraser, Aug. 1934.

Colonies mostly epiphyllous, scattered, thin, black, up to 4 mm. diam., with central
group of thyriothecia. Mycelium of brown hyphae 2—4u thick, the secondary hyphae
paler, indistinctly septate, substraight to very crooked, closely reticulate. Hyphopodia
mostly on the main, darker hyphae, alternate or thinly scattered, continuous,
hemispheric to irregularly pulvinate, 4-7 x 6—-8u. Thyriothecia closely grouped in centre
of colony, linear, up to 700 x 150u, often confluent and sometimes Y-shaped, black,
convex, smooth; margin irregularly crenate, not distinctly fimbriate; upper wall of
opaque dark brown, irregularly radiating hyphae 3—4u thick, splitting longitudinally at
maturity; lower wall hyaline, fibrous, very thin and indistinctly separated from the
internal tissue of hyaline hyphae. This tissue fills the whole interior in young thyrio-
thecia, consisting of tortuous-erect, interwoven, branched and indistinctly septate hyphae
2-2-5u thick, the erect ends simulating paraphyses between the young asci, and
becoming brownish at the tips in the opened ascomata. Asci numerous, erect, sessile,
broadly ellipsoid, up to 40 x 20u, 8-spored. Spores conglobate, long remaining hyaline,
finally brown, oblong with rounded ends, 1-septate and slightly constricted at the middle,
smooth, 13-14 x 5-7u. No conidial stage seen.

(91) MicroTHYRIELLA PHAEOSPORA Hansf., n. sp.

Thyriothecia superficialia, epiphylla, levia, applanata, brunneo-atra, orbiculata, usque
ad 280% diam., margine pallidiora pelliculosaque. Mycelium liberum nullum. Asci in
textu laxo hypharum hyalinarum singulariter subdenseque immersi sub pellicula
myceliale, globosi, tenuiter tunicatae, 15-184 diam., 8-spori, sessiles. Sporae conglobatae
vel subparallelae, clavulatae, 1-septatae, leves, brunnescentes, 10-12 x 4-5—5u.

Hab. in foliis Knightiae excelsae, Kapiti Is., Wellington, New Zealand, leg. H. H.
Allan, November 1936.

The thyriothecia are completely superficial, epiphyllous, thin, flat, dark brown,
circular, up to 2804 diam., the wide margin much paler and consisting of very pale
yellowish-brown wavy-reticulate hyphae about 2-34 wide, indistinctly septate, in a
single layer, agglutinate to form a pellicle which extends over the whole ascoma,
becoming darker towards the centre. Here it covers a rather loose weft of tangled
hyaline hyphae 2u thick, much branched, in which the asci are embedded singly,
though rather closely, apparently each in its own “locule”. The asci are globose, sessile,
thin-walled, 15-184 diam., 8-spored. Spores conglobate to nearly parallel, clavulate,
1-septate, slightly constricted, becoming dark brown when fully mature, 10-12 x 4-5—-5yn,
the upper cell wider than the lower and subglobose, the lower slightly attenuate to the
rounded base.

Formerly this would have been placed in Hremotheca, but this genus is now con-
sidered not to be sufficiently distinct from Microthyriella.

(92) MIcROTHYRIELLA FRASERI Hansf., n. sp.

Thyriothecia hypophylla, subdense dispersa, discreta, dilute brunnea, usque ad 350u
diam., in sicco rugulosa, applanata. Mycelium liberum nullum. Paries superior
membranaceus, plectenchymaticus, pellucide brunneus, levis, ex hyphis 2-5—4u er.,
indistincte septatis compositus. Paries inferior et nucleus ex hyphis hyalinis 2-3» er.,
irregulariter ramosis, laxe intertextis, tenuiter tunicatis compositi. Asci singulariter

BY C. G. HANSFORD. 113

dispersi, globosi, sessiles, usque ad 30u diam., aparaphysati, 8-spori, tunica circa 2y cr.
Sporae conglobatae, oblongae utrinque obtusae, hyalinae, 1-septatae, lenissime con-
strictate, 12-17 x 6-7u, cellulis subaequalibus.

Hab. in foliis Canthii coprosmoidis, Hastings R., New South Wales, L. Fraser, April
1952.

There is no free mycelium in this species and each thyriothecium apparently
represents the complete development of a single unit; they are closely scattered,
hypophyllous, discrete, closely adpressed to the host cuticle, thin and flattened, astomous,
pale brown, becoming rugulose when dry, up to 350u diam. The upper wall is translucent
brown, membranaceous, passing at the edge into the mycelial margin of a single layer
of agglutinate hyphae forming a pellicle, in the centre wavy-plectenchymatous, smooth,
the hyphae 2-5-4u thick, indistinctly septate. The lower wall and the internal tissue
grade into each other, consisting of hyaline, irregularly branched, loosely interwoven
hyphae 2-34 wide, indistinctly septate, forming a loose tissue in which the asci are
scattered singly. Asci globose, sessile, up to 30u diam., 8-spored, aparaphysate, the wall
rather thick (—2u) and brittle at maturity. Spores conglobate, oblong with obtuse ends,
1-septate, very slightly constricted, smooth, hyaline, 12-17 x 6—7yu, the cells more or less
equal.

Neither in this, nor in the preceding species could any evidence of penetration of
the host be observed in sections; both appear to be saprophytic in habit.

(93) TRICHOPELTIS SP.

On Cryptocarya meissneri, Comboyne, New South Wales, L. Fraser, January 1935.

Colonies epiphyllous, black, dendritic, thin and loose, up to 5 mm. diam. or confluent,
smooth. Mycelium of parallel hyphae along the centre of each strand, regularly
divergent to both sides, brown, septate, 3-54 thick, the cells 10-124 long; each strand
consists of a single layer of hyphae laterally agglutinate and up to 100” wide, the
surface finely granulose and the ends of the hyphae almost perpendicular to the sides
of the strand. The thyriothecia develop as a widened and thickened area of the strand,
with a central orifice, the upper wall apparently of mycelial origin and not distinguish-
able from the rest of the strand, the margin entire or finely crenate, circular, up to 300u
diam. or elongated along the strand; central pore more or less circular, 10-154 diam.
Lower wall hyaline, indistinct. The body of the thyriothecium is filled with a loose
tissue of hyaline hyphae 2-5 thick, septate into short cells which easily separate under
pressure, and amongst which the asci develop around the sides, pointing obliquely to the
central pore. Asci irregularly ovate-ellipsoid, sessile, rounded and when immature
somewhat thickened around the apex, 8-spored, aparaphysate, up to 40 x 10u. Spores
multiseriate and obliquely parallel in ascus, hyaline, clavulate with rounded ends,
2-septate, the upper cell largest and widest, smooth, not constricted at the septa,
10-12 x 3y.

It is possible that this may be 7. pulchella Speg., of which I have not seen authentic
material. It appears to be completely saprophytic in habit, and certainly is not hyper-
parasitic upon other fungi.

(94) DIMERIELLA LUDWIGIANUM (Sacc.) Hansf., n. comb.

= Dimerosporium ludwigianum Sacc., Hedwigia, 1889, and in Syll. Fung., 9:405, 1891.

The original collection was made by Tepper on Lagenophora stipitata (L. Billardiert),
Mt. Lofty, South Australia; it was re-collected by N. T. Flentje on the same host at Port
Lincoln, S. Aust., in July 1952.

Colonies amphigenous, dark brown to black, effuse and often confluent over most
of the leaf, when single up to 3 mm. diam., thin. Mycelium of exhyphopodiate brown
hyphae 2-5-3-5u thick, the cells up to 25u long, copiously and irregularly branched,
forming a loose network over the leaf, the hyphae flexuous to irregularly bent and
connected at intervals direct through the cuticle to a subcuticular mycelium. This
latter forms almost a continuous plate beneath the external colony, consisting of hyaline

114 AUSTRALIAN FUNGI, II,

hyphae 2-34 wide, the cells 5-8u long, compressed vertically and often in 2-3 layers; no
haustoria are produced and no other form of penetration of the host leaf was found on
either surface. The perithecia are closely scattered on the external mycelium, and are
sessile, black, globose, membranous, up to 100” diam. and about 90” high, smooth; from
the upper half and the sides arise 10-20 spreading setae, simple, straight or somewhat
irregularly bent, septate, dark brown, 30-250 x 3-5-5u, the shorter ones having obtuse
to very slightly clavulate apices, the longer ones often descending to the leaf surface
and passing outwards into mycelial hyphae. The perithecial wall consists of an outer
dark layer of angular parenchyma, the cells polygonal and up to 154 diam., lined with
one or two layers of much compressed hyaline cells; at first closed, later opening by a
rounded apical pore about 10u diam., surrounded by slightly protruding subhyaline
periphyses. Asci basal, numerous, erect, elongate-ellipsoid, broadly rounded at the apex,
not noticeably thickened there, nodose-stipitate below, 8-spored, up to 40 x 9-12u.
Paraphyses numerous, filiform, hyaline, continuous, 1-1-5 thick, not connected above
to the perithecial wall. Spores 2-seriate, fusoid with rounded ends, 1-septate, very
slightly constricted, the cells sub-equal, 10-12 x 3u (immature and hyaline). No
discharged spores were found in this material, nor any germinating on the leaf surface,
so that it remains unknown whether they become olivaceous when fully mature; judging
from analogy with other similar leaf-parasites, they probably remain hyaline.

As far as the writer is aware, this fungus has remained a mycological puzzle since
its original collection, and it is therefore particularly fortunate that it has now been
rediscovered and provisionally assigned to a position in modern classification; the old
genus Dimerosporium contained a very heterogeneous collection of species now dis-
tributed amongst many different genera of Microthyriaceae and Sphaeriales, and as a
genus has now been totally discarded.

(95) APHANOSTIGME RUBI Hansf., n. Sp.

Mycelium hypophyllum, in tomento folii parasiticum, ex hyphis dilute olivaceis,
flexuosis, septatis, ramosis, 2-34 cr., exhyphopodiatis compositum; haustoria nulla.
Perithecia laxe dispersa, superficialia, atra, globosa vel leniter ampullacea, usque ad
120u diam. et alt.; paries indistincte parenchymaticus, extus granulosus et deorsum
hyphis mycelii vestitus, superne setis erecto-patentibus, numerosis, atrobrunneis, rectis
vel leniter flexuosis, simplicibus, obtusis, usque ad 60 x 2-34 ornatus. Ostiolum conicum,
poro 10-154 diam. pertusum, primo cellulis hyalinis mollibus implectum. Asci
aparaphysati, basales, numerosi, cylindracei, apice rotundati, subsessiles, 8-spori, usque
ad 70 x 10u. Sporae 2—3-seriatae, hyalinae, fusoideae, rectae vel curvulae, leves, 3-septatae,
haud constrictae, utrinque attenuato-rotundatae, 18-22 x 3-—4u.

Hab. in foliis Rubi moluccani, Williams R., New South Wales, L. Fraser 112.

This fungus is parasitic upon the leafhairs on the lower surface of the host leaf;
its mycelium consists of pale olivaceous, crooked, septate, much branched hyphae 2-34
thick, ramifying over the leaf surface between the hairs, with branches penetrating these
and forming small mycelial knots in the lumen. There are no haustoria or hyphopodia,
and no other penetration of the leaf tissue was detected. The perithecia are scattered
loosely on the external mycelium amongst the leafhairs and are black, globose to
somewhat flask-shaped, up to 120u diam. and high, with a blunt conical ostiole, which
is at first filled with a plug of delicate hyaline parenchyma; the external surface is
closely and finely dark-granulose, thus obscuring the parenchymatous structure beneath;
the lower part is clothed with mycelial hyphae extending outwards, and the upper half
bears numerous erect-spreading setae. These setae are straight to slightly flexuous,
simple, obtuse and paler at the apex, apparently continuous, up to 60 x 2-3. Asci
numerous, basal, aparaphysate, cylindric, subsessile, rounded but not noticeably
thickened at the apex, 8-spored, up to 70 x 10u. Spores 2-3-seriate, hyaline, fusoid with
rounded ends, straight or slightly bent, smooth, 3-septate, not constricted, 18-22 x 3—4u.

This parasite is very inconspicuous at first, but in later stages the tomentum of the
leaf is turned reddish in colour; no leafspots are produced.

BY C. G. HANSFORD. 115)
(96) MELIOLINOPSIS MELIOLAE (Stev.) Petrak, Sydowia, 5:334, 1951.
= Perisporium meliolae Stev. = Meliolina meliolae (Stev.) Stev. = Phaeophrag-
meriella meliolae (Stev.) Hansf.
Parasitic on Jrene kiraiensis on Doryphora sassafras, National Park, New South
Wales, Fraser 26.

(97) CONIOCHAETA WILLIAMSI Hansf., n. sp.

Perithecia dispersa vel subgregaria, nigra, globosa, superficialia, usque ad 300y
diam., superne subdense setosa; setae nigrae, continuae, rectae vel curvulae, simplices,
usque ad 30u longae, basi 4-6 cr., Sursum ad apicem acutem attenuatae; paries
perithecii fragilis, carbonaceus, pluristratosus, opace atrobrunneus, cellulis polygonalibus,
intus pallidioribus. Asci numerosi, cylindracei, apice rotundati, deorsum leniter
attenuati, usque ad 80 x 8u, 8-spori, tenuiter tunicatae. Sporae oblique 1-seriatae, ovatae,
brunneae, leves, continuae, 7-9 x 4-5:5u. Paraphyses numerosae, filiformes, hyalinae,
simplices, continuae, ascos aequantes vel leniter superantes, lu er.

Hab. in ligno, Meningie, South Australia, WARI 3533, leg. L. D. Williams.

Perithecia closely scattered or gregarious, black, superficial, up to 300u diam.,
globose, rather densely setose above; setae black, 30 x 4—6u, straight or bent, continuous,
gradually attenuate from the base to the acute simple apex. Perithecial wall brittle
carbonaceous, of several layers of black polygonal thick-walled cells, paler and thinner
inside. Asci numerous, cylindric with rounded apex, not thickened, attenuate below
into a short stipe, 8-spored, up to 80 x 8u. Spores obliquely 1-seriate, ovate with obtuse
ends, brown, smooth, continuous, 7-9 x 4-5-5u. Paraphyses numerous, filiform, hyaline,
simple, continuous, equalling or slightly exceeding the asci, lu thick.

(98) PHYLLACHORA ERAGROSTIDIS Doidge, Bothalia 4:430, 1942.
On Hragrostis infecunda, Two Wells, South Australia, Robertson, April 1953.

(99) NEMATOTHECIUM AUSTRALIENSIS Hansf., n. sp.

Mycelium in plagulis Ireninae acmenae parasiticum, ex hyphis hyalinis, 2-3 cr.,
indistincte septatis, ramosissimis, dense reticulatis compositum. Perithecia superficialia,
laxe dispersa, globosa vel leniter depressa, usque ad 250u diam., leves, apice irregulariter
lateque aperta, extus hyphis mycelii laxe vestita; paries atrobrunneus, dense granulosus,
indistincte parenchymaticus, unistratosus. Asci numerosi, erecti, clavati, apice late
rotundati, usque ad 45 x 10u, deorsum breviter lateque stipitati, 4-8-spori. Paraphyses
numerosae, filiformes, simplices, hyalinae, usque ad 70u longae. Sporae parallelae positae,
eylindraceae utrinque rotundatae, dilute olivaceae, leves, usque ad 35 x 3yu, indistincte
5-septatae, haud constrictae.

Hab. in foliis Acmenae smithii, Myrtle Gully, New South Wales, August 1934, L.
Fraser 159.

This fungus is a hyper-parasite of Irenina dcmenae, and its mycelium forms a thin
pellicle over and between the hyphae of its host; the hyphae are hyaline, 2-3 thick,
much branched, indistinctly septate, agglutinate and closely reticulate. Perithecia
loosely scattered on the mycelium, globose to somewhat flattened, covered outside with
loose hyaline mycelial hyphae, up to 250u diam., not setose, becoming irregularly and
widely open at the apex when mature. Perithecial wall dark brown, closely granulose,
the structure very indistinct, but apparently composed of meandering hyphae forming
a single pseudoparenchymatous layer. Asci very numerous, basal, erect, clavate with
broadly rounded apex, up to 45 x 10u diam. at the apex, attenuate below into a short
wide stipe, containing 4-8 spores; the wall thin and not conspicuously thickened around
the apex. Paraphyses very numerous, filiform, simple, hyaline, continuous, up to 70u
long by about lu thick. Spores parallel in ascus, cylindric with rounded ends, or very
slightly attenuate downwards, becoming pale olivaceous and indistinctly 3-septate when
fully mature, not constricted, smooth, up to 35 x 3u.

The same fungus has been observed on other species of Meliolaceae collected by
Dr. Fraser in New South Wales.

116 AUSTRALIAN FUNGI, II,

(100) MALACARIA AUSTRALIENSIS Hansf., n. sp.

Mycelium ex hyphis hyalinis 1-5-2u er., ramosis, reticulatis, subagglutinatis com-
positum, illud Jreninae fraserianae vestiens. Perithecia superficialia, laxe dispersa,
globosa, in siceco subcollabascentia, ex fulvo fuscescentia vel nigrescentia, usque ad
140u diam., extus levia vel hyphis mycelii laxe vestita. Paries perithecii subtranslucens,
lentus, parenchymaticus, circa 15-20u er., extus ex cellulis rotundato-angulosis 10 x 5y,
et intus ex cellulis hyalinis compressibus compositus. Asci numerosi; paraphysati, late
clavati, apice rotundati, deorsum attenuati, basi truncati (3-4 ecr.), usque ad 40 x 13y,
8-spori. Paraphyses numerosae, filiformes, irregulariter ramosae, hyalinae, ascos
superantes. Sporae parallelae, tenuiter clavulatae, apice rotundatae, deorsum lenissime
attenuatae, continuae, usque ad 28 x 2-5-3u, utrinque appendiculatae; appendices.
setiformes, solidae, hyalinae, rectae vel curvulae, 4-5 x ly; sporae maturae dilute
olivaceo-brunneae, in massa atrobrunneae.

Hab. in plagulis Zreninae fraserianae in foliis Cryptocaryae glaucescentis, Williams
R., New South Wales, L. Fraser 208.

Mycelium of hyaline hyphae 1-5—-2u thick, much branched and closely investing
those of the host fungus, forming a thin network between these, somewhat agglutinate.
Perithecia scattered, superficial, globose, becoming flattened when dry and sometimes.
collapsing, from dirty yellowish becoming fuscous to almost black as the spores mature,
up to 140u diam., the surface smooth or with scattered mycelial hyphae originating
half-way up the wall. Wall of subtranslucent brown rounded-polygonal cells about
10 x 5u, smooth on the exterior surface, enclosing one or more layers of hyaline
flattened cells, the whole wall tough, not brittle, in texture and 15-20 thick. Asci and
paraphyses forming an almost solid nucleus extruded intact when the perithecium is.
compressed; the hymenium covering the basal two-thirds of the loculus, with the asci
directed towards the apical pore. Asci broadly clavate, widely rounded at the apex (to
134 wide), attenuate to the truncate base (3-44 wide), up to 40u long, 8-spored; wall
rather thin, not thickened at the apex. Paraphyses numerous, filiform, irregularly
branched, hyaline, exceeding the asci, 1-1-5u thick, not septate. Spores parallel in
ascus, narrowly clavulate, rounded at the apex, very slightly attenuate downwards to
the rounded base, continuous, up to 28 x 2-5-3, at first hyaline, becoming greenish-
brown, in mass almost black, and furnished at each end with a solid, hyaline, straight
or slightly bent bristle, 4-5 x 1p.

(101) GIBBERELLA FUJIKUROI var. SUBGLUTINANS Edwards, Agr. Gaz., N.S.W., 44:85, 1933.
On old rust sori on Suaeda australis, Meningie, South Australia, WARI 3490,
leg. L. D. Williams.

(102) MASSARINULA PHYLLODIORUM McAlpine, Proc. Linn. Soc. N.S.W., 28:555, 1903.

On Acacia myrtifolia, Mt. Burr, S. Australia, Aug. 1922, WARI 2002, leg. J. G. Wood.

Leafspots penetrating the whole leaf thickness, round, grey with dark brown
margin, up to 2 mm. diam., not secedent. Mycelium of septate, hyaline hyphae,
much branched, intercellular, without haustoria, penetrating and killing the whole
mesophyll, but not extending beyond the margin of the leafspot. Ascomata amphigenous,,.
1—4 on each spot, subepidermal, globose, up to 250u diam., becoming slightly erumpent
with age, the epidermis sloughing off above them, black, hard and stromatic; in early
stages consisting of an outer part of thick-walled, dark brown, angular cells, passing
gradually into a hyaline, pseudoparenchymatous centrum of delicate thin-walled cells.
At the base of this centrum the asci develop singly, in a broken single layer, and enlarge
upwards at the expense of the centrum tissue, till at maturity they appear to be
separated by thin partitions of septate, more or less compressed “paraphysoids”. The
ascoma eventually opens by irregular fracture of the apex of the outer, dark, stromatic
wall. Asci about 12, erect, clavate, up to 180u long, the apex widely rounded and 40-50u.
diam., gradually narrowed to the basal stipe, which is usually bent and about 10 thick,
25-304 long; ascus wall 1-1-5 thick in the lower part, thickened around the apex to 5y,

BY C. G. HANSFORD. 117

with a central “internal pore’; asci 8-spored, aparaphysate. Spores parallel, overlapping,
hyaline, fusoid, slightly bent, rounded at both ends, 1-septate and slightly constricted in
the middle, smooth, 55-65 x 12—14u, the cells equal.

(103) MONTAGNELLINA GoopIAE Hansf., n. sp.

Perithecia epiphylla, dispersa, immersa, glabra, atra, 150-250u diam. et 300 alt.,
globosa vel ampullacea, haud prominula, membranacea; paries circa 20—-25u er., deorsum
subhyalinus, sursum atrobrunneus, apice ad 40u incrassatus et ad stroma subepidermale
atrobrunnea confluens, pluristratosus, parenchymaticus. Centrum loculi ex pseudo-
paraphysibus hyalinis, 3—-4u cr., septatis, simplicibus vel furcatis, laxe intertextis
compositum. Asci basales, numerosi, erecti, aparaphysati, cylindracei vel clavati,
apice rotundati, deorsum in stipitem brevem attenuati, 8-spori, 80-100 x 9-20u, tunica
sursum incrassata usque ad 3—-4u, poro interno centrali praedita. Sporae 1-seriatae vel
plus minusve oblique 2-seriatae, hyalinae, ellipsoideae, obtusae, leves, continuae,
12-15 X 7—8u.

Hab. in foliis Goodiae medicagineae, Rocky River, Kangaroo Is., South Australia,
Oct. 1922, leg. T. G. Osborn, WARI 1997.

Perithecia scattered over the older leaves, which are somewhat yellowed, but not
on definite leafspots, epiphyllous, immersed, glabrous, about 3004 deep and 150-250u
diam., globose to flask-shaped, with short neck not protruding above the epidermis,
black, membranous. Perithecial wall about 20-25. thick, in the deeper parts subhyaline,
towards the apex becoming dark brown on the exterior, and thickened up to 40u, at
the apex confluent with a stroma developed between the flat epidermal cells and the
palisade tissue, composed of dark brown pseudoparenchyma, not filling the epidermal
cells to form a true clypeus; wall of perithecium of concentric flattened pseudo-
parenchyma in several layers. The centrum consists of pseudoparaphyses joined both
at the apex and the base to the innermost wall layer, usually simple though sometimes
branched, 3-4 thick, septate, hyaline, with delicate walls. Into this rather loose tissue
the asci develop from the base of the loculus; numerous, erect or convergent towards
the apex of the loculus, gradually crushing and replacing the original centrum, finally
appearing aparaphysate, cylindric to clavate, rounded at the apex, with attenuated basal
stipe portion, 8-spored, 80-100 x 9-20u; the wall thin, except around the apex where it is
thickened to 3-44 and has a central internal pore. Spores 1-seriate or more or less
obliquely 2-seriate, hyaline, ellipsoid, smooth, continuous, 12-15 x T-8u. Internal
mycelium copious, of septate, hyaline, branched hyphae 2—4u wide, intercellular, without
haustoria.

The fungus resembles a Phyllachora, but the perithecia are always single and have
no true clypeus; the ostiole is lined with very short hyaline periphyses, with a pore
about 204 diam.

(104) BorrRYOSPHAERIA BANKSIAE, Hansf., n. sp.

Ascomata hypophylla, in greges 2-10 dense aggregata, nigra, subglobosa, levia, usque
ad 470u diam. et alt., apice poro minuto pertusa, interdum lateraliter connata .vel
2-loculata. Loculi subampullacei, circa 300u alt. et 270u diam., collo conoideo, circa
100% longo et basi 100u diam., sursum in poro apicali 30u diam. contracto. Asci numerosi,
aparaphysati, cylindracei, 8-spori, apice rotundati, subsessiles vel brevissime stipitati,
usque ad 120 x 18u, evanescenti. Sporae oblique vel transverse 1-seriatae, ellipsoideae,
hyalinae, leves, continuae, 17-20 x 138-15yu, episporio 1-5u er.

Hab. in foliis Banksiae marginatae, Normanville, South Australia, WARI 2064, leg.
G. Samuel, Jan. 1924 (typus); Kuitpo, S. Austr., Dec. 1921, G. Samuel.

Ascomata hypophyllous, often amongst the colonies of Parodiella banksiae, from
which they are easily distinguished by their much larger size, in small close groups up
to 1 mm. diam., black, subglobose, smooth, sometimes slightly papillate above, up to
470u high and diam., opening at the apex by a minute round pore; often laterally connate

-or some of the larger individuals containing 2 loculi.

118 AUSTRALIAN FUNGI, II,

The mycelium is of limited extent in the leaf mesophyll, consisting of much
branched, septate, hyaline hyphae, intercellular, without haustoria. In the centre of
each infection an internal hyaline stroma is formed, reaching from the base of the
palisade tissue to the lower epidermis, composed of small pseudoparenchyma which
eventually darkens beneath the epidermis, this being burst as the stroma expands to
form an erumpent black tubercle containing one or sometimes two ascigerous loculi.
Similar tubercles arise in close proximity from the same stroma till a group of 5-10
ascomata is formed. Each loculus is surrounded by a wall of 4-10 layers of much
compressed hyaline to yellow-brown parenchyma, passing outside into the stroma tissue,
but often distinct from the latter around the base and sides, this inner layer being
5-15u thick. The middle layers of the tissue of the ascostroma are plectenchymic like
the hypostroma, of which they are a continuation, and may often be much compressed
or even absent, then the inner loculus wall passing direct into the black parenchyma of
larger cells forming the external stroma wall; the whole wall from outside to loculus
cavity is about 50u thick around the sides. The outermost layers are of thick-walled
rounded cells, opaque dark brown to black, about 104 diam., the external surface
somewhat roughened with outgrowths, or quite smooth and shining black.

The loculus, when single, is somewhat flask-shaped, with short apical neck, about
300u high and 270u diam., the sides and base lined with numerous asci directed towards
the apical pore; the neck is short conoid, about 1004 long and wide at the base, con-
tracting to the apical pore, which is about 30u diam., lined with delicate parallel
periphyses directed upwards, but with few free ends. The young loculus is filled with a
loose tissue of hyaline hyphae connected above and below with the innermost wall layer;
amongst these the asci develop, and their remains in the mature loculus simulate
septate paraphyses about 1-5u thick. The ostiolar cavity appears to be of lysigenous
origin. Asci numerous, cylindric, 8-spored, rounded at the apex, sessile or with very
short wide basal stipe, thin-walled, about 120 x 18u, disappearing early to leave the
spores free inside the loculus. Spores obliquely or even transversely 1-seriate, ellipsoid.
hyaline, smooth, continuous, 17-20 x 13-154, the epispore 1-54 thick. Intact asci were
rare in both collections examined.

(105) PSEUDOSPHAERIA SAMUELI Hansf., n. sp.

Maculae amphigenae, irregulariter rotundato-dendriticae, leniter rugulosae, fulvo- vel
atro-brunneae, subzonatae. Mycelium sub epidermide stromatice aggregatum, ex hyphis
hyalinis 1-24 cr., septatis, intertextis compositum, in mesophyllum hyphis singulis
intercellularibus profunde penetrantibus, haustoriis nullis. Ascomata in stromate
subcuticulare orta, usque ad 120u diam. et 60y alt., apice poro rotundato circa 104 diam.,
pertusa, glabra, nigra, 1-(2)-loculata; paries olivaceo-brunneus, plectenchymaticus,
deorsum circa 8-10 cr., sursum obscurior vel opace niger et incrassatus usque ad
15-204; intus hyphis hyalinis intertextus primo repleta. Asci modice numerosi, lateralo-
basales, apicem loculi versus directi, aparaphysati, cylindracei vel saccati, sursum late
rotundati, deorsum breviter stipitati, usque ad 100 x 35, 8-spori, tenuiter tunicati.
Sporae subparallelae, cylindraceae utrinque rotundatae, hyalinae, 3-septatae, haud
constrictae, eguttulatae, 60-65 x 6-7.

Hab. in foliis Hucalypti obliquae, Cape Jervis Peninsula, South Australia, Jan. 1924,
WARI 2069, leg. G. Samuel; Balhannah, S. Austr., May 1922, WARI 2068, leg. G. Samuel;
in foliis #. rostratae, Noarlunga, S. Austr., May 1924, WARI 2072, leg. G. Samuel.

On Hucalyptus obliqua the leafspots are amphigenous, irregularly rounded to
dendritic, slightly rugulose-elevated, buff to dark brown, indistinctly zonate, with rather
indefinite rings of black ascomata; on H. rostrata the spots are only 3 mm. diam., more
condensed, with the ascomata closely crowded and not circinately scattered. The
mycelium is entirely within the host leaf, and aggregates beneath the epidermis as a
thin stroma of fine hyaline septate interwoven hyphae, penetrating deep into the
mesophyll as isolated hyphae, which are intercellular and without haustoria, sometimes
reaching the opposite epidermis; the subepidermal stroma is up to 25m thick, the
individual hyphae 1-5-3 wide, in many places parallel and closely adnate.

BY C. G. HANSFORD. 119

The ascomata arise as thickened darker areas within the stroma, each surrounded
by an olivaceous brown wall of compressed plectenchyma, about 8-10u thick below the
loculus and thickened around the sides and upper part to 15-20u, where it is much
darker to subopaque or black. The loculus is flattened globose, up to 120u diam. and
60u high, at first filled with a tissue of interwoven hyaline septate hyphae, the remains
of which form a lining to the wall of the mature ascoma, the rest being replaced by the
developing asci. Sometimes a single ascoma may contain at first two separate loculi,
separated by a central column of tissue similar to that of:the wall; this may eventually
break down and the loculi become confluent. The ascoma opens at maturity by a
lysigenous apical pore about 104 diam. Around the upper part of the ascoma the stroma
turns dark brown and is confluent with the apex of the loculus wall, peripherally
becoming distinct and merging into the subepidermal stroma; traces of the epidermal
cells remain above the dark stroma mass, which can hardly be said to constitute an
epidermal clypeus.

The asci develop around the base of the loculus, the older ones more peripheral,
and enlarge at the expense of the original plectenchymic centrum, eventually more or
less completely filling the loculus; they are fairly numerous, aparaphysate, cylindric to
saceate, widely rounded and not thickened above, contracted below into a very short
stipe, up to 100 x 35u, 8-spored, thin-walled and apparently uni-tunicate. Spores parallel
in ascus, cylindric with rounded ends, often slightly bent, hyaline, smooth, transversely
3-septate, not constricted, not guttulate, 60-65 x 6—7u.

(106) PARODIELLA BANKSIAE Sacc. & Bizz., Syll. Fung., 9:410, 1891.

This species appears to be fairly common on species of Banksia in Tasmania,
Victoria, New South Wales and South Australia. It forms indistinct black patches on
the lower surface of the leaf, often obscured by the tomentum and then with only the
groups of ascomata showing through. The mycelium penetrates through the stomata
and ramifies in the intercellular spaces of the mesophyll, rarely penetrating high up in
the palisade tissue, the internal hyphae being hyaline, septate, 3-5u thick, not forming
haustoria. The external mycelium spreads between the leaf-hairs over the surface, and
consists of brown, much branched, septate hyphae 4—5u thick, without hyphopodia or
stomopodia, aggregated in places to form small stromata and attached to the bases of
the ascomata. The latter are closely aggregated and often sub-confluent at the sides,
black, smooth above, obpiriform to globose, 100-1604 diam., and high, completely
superficial amongst the leaf-hairs and attached only by mycelial hyphae, glabrous,
rather hard in texture. The wall consists of several layers of dark brown, angular
parenchyma, thick-walled in the outer layers, thinner inside, the cells up to 12 x 8u,
the whole wall of the mature ascoma about 254 thick at the apex and sides, thicker
below; enclosing at first a hyaline parenchymatous centrum, which is absorbed as the
asci develop, leaving finally only a few loose scattered cells amongst and above the asci.
Asci basal, erect, aparaphysate, fairly numerous, sessile on a hyaline tissue of delicate,
interwoven, branched hyphae at the base of the loculus (? ascogenous hyphae), elongate
clavulate, rounded and slightly thickened at the apex, without definite internal apical
canal, not obviously bi-tunicate, 8-spored, up to 70 x 14u. Spores obliquely 2-seriate,
dark brown, 1-septate, slightly constricted, smooth, 14-15 x 4-5u, the upper cell slightly
broader than the lower. There is no ostiole and the ascomata open by irregular fracture
of the apex when mature.

(107) PHYSALOSPORA ACACIAE Hansf., n. sp.

Mycelium in cortice ramulorum perennans et gallas efformans; perithecia in cortice
immersa, globosa, atra, usque ad 150u diam., dense stipata et pycnidiis Diplodiae
intermixta. Asci clavata, subsessiles, apice late rotundati incrassatique (—7/), recti vel
curvuli, 8-spori, usque ad 90 x 28u; paraphyses filiformes, hyalinae, simplices, continuae,
ascos aequantes. Sporae irregulariter positae, ellipsoideae, obtusae, hyalinae, leves,
continuae, subinde lenissime inaequilaterae, haud guttulatae, 20-24 x 9-10u, episporio
tenui.

120 AUSTRALIAN FUNGI, II,

Hab. in ramilis Acaciae baileyanae, Pennant Hills, New South Wales, Aug. 1958,
L. Fraser (WARI 3568).

The fungus invades the cortex of young twigs, causing at first greyish areas with
a dark margin, these later becoming slightly elevated, developing in later seasons into
galls and causing distortion of the twigs; the gall tissue consists mainly of the
hypertrophied cortex with intercellular hyaline hyphae without haustoria. The perithecia
are completely sunken in the gall tissue with only the apical pore reaching the surface;
globose, black, glabrous, up to 150u diam., closely crowded and mixed with pycnidia of
Diplodia type. Asci clavate, subsessile, widely rounded and thickened up to Tu
around the apex, with a shallow wide internal apical canal, straight or slightly bent,
8-spored. Spores irregularly arranged in the ascus, ellipsoid with obtuse ends, straight
or slightly inequilateral in side view, hyaline, smooth, continuous, thin-walled, not
guttulate, 20-24 x 9-10u. Paraphyses equalling the asci, filiform, hyaline, continuous,
simple.

(108) PHYSALOSPORA XANTHORRHOEAE Hansf., n. sp.

Perithecia numerosa, dispersa vel lineare 2-3-aggregata, subepidermalia, depresse
globosa, atra, glabra, submembranacea vel coriacea, 100-1804 diam., circa 130 alt.,
apice poro pertusa; paries atrobrunneus, 2—4-stratosus, intus hyalinus. Asci basales,
elongato-clavati vel subcylindracei, apice late rotundati incrassatique (-—10u), deorsum
in stipitem brevem attenuati, 8-spori, usque ad 120 x 20-25u. Sporae oblique 2-seriatae,
hyalinae, ellipsoideae vel clavulatae, continuae, leves, 22-25 x 7-9u. Paraphyses
numerosae, filiformes, ascos aequantes, 2-2-5u cr., continuae, rectae vel flexuosae,
simplices.

Pyenidia (Macrophoma) peritheciis consimilia et commixta, usque ad 200" diam.;
sporophoras breviter cylindracea, pycnosporae cylindraceae utrinque rotundatae, hyalinae,
continuae, leves, 15-19 x 3-4u, singula et acrogenea.

Hab. in caule emortuo Xanthorrhoeae semiglaucaeé, Meningie, South Australia, July
1953, L. D. Williams 43.

The perithecia occur mixed with the Macrophoma pycnidia on the dead flowering
scape of the host, numerous, scattered singly or in linear groups of 2-3, subepidermal,
with a minute pore opening to the surface, flattened globose, black, smooth, glabrous,
leathery-membranous, 100-1804 diam. and about 130 high, easily separable from the
host tissue and from the covering epidermis. Perithecial wall of several layers of dark
olivaceous parenchyma, lined with compressed hyaline layers. Asci in a basal hymenium,
erect, elongate clavate to subcylindric, broadly rounded and thickened at the apex to
10u, with central internal canal, attenuate below into a short flexuous stipe, 8-spored, up
to 120 x 20-254. Paraphyses numerous, filiform, equalling the asci and about 2-2-5
thick, continuous, straight or flexuous, not swollen at the apex, hyaline. Spores obliquely
2-seriate, hyaline, smooth, ellipsoid to slightly clavulate, the ends broadly rounded,
continuous, 22-25 x 7—9u.

Macrophoma pycnidia similar to the perithecia, but often up to 200u diam.; the wall
lined with short cylindric simple conidiophores abstricting single conidia from their
apices; pycnospores cylindric with rounded ends, hyaline, continuous, smooth, 15-19 x
3—4u.

(109) LrprosPHAERIA HARDENBERGIAE Hansf., n. sp.

Maculae 1-3 mm. diam., amphigenae, griseae, non secedentes, in centro arescentes,
margine atrobrunneae, primo zono flavo 1-1-5 mm. cr. circumdatae, subinde confluentes
irregularesque. Perithecia amphigenea, immersa, nigra, levia, glabra, globosa, usque
ad 180u diam.; paries membranaceus, parenchymaticus, 3—4-stratosus, strato exteriore
ex cellulis polygonalibus atrobrunneus, circa 124 diam. composito, stratis interioribus
subhyalinis, fortiter compressibus. Asci numerosi, basales, fasciculati, cylindraceo-
clavati, subsessiles, usque ad 70 x 12-15u, 8-spori. Paraphyses hyalinae, filiformes,
continuae, asci subaequantes. Sporae 1-—2-seriatae, olivaceae, fusoideae utrinque
rotundatae, curvulae, leves, 5-septatae, haud constrictae, 16-20 x 4-5-6y.

BY C. G. HANSFORD. 121

Hab. in foliis Hardenbergiae violaceae, Port Lincoln, July 1952, N. T. Flentje.

Leaf-spots amphigenous, at first surrounded by a yellow zone 1-1-5 mm. wide, then
enlarging and with a dark brown margin, the centre drying out to grey-white, not
usually secedent, rounded in outline, 1-3 mm. diam. or confluent and irregular. Perithecia
amphigenous, immersed in the mesophyll, black, smooth, glabrous, globose, up to 180
diam.; perithecial wall membranous, parenchymatous, consisting of an outer layer of
dark polygonal cells about 124 diam., enclosing two or three layers of subhyaline, much
compressed cells. Asci numerous, in a single large basal fascicle, the inner ones
straight, the outer incurved and directed towards the apical pore of the perithecium,
cylindric-clavate, subsessile, rounded at the apex, not noticeably thickened, 8-spored,
up to 70 xX 12-15u. Spores olivaceous-brown, fusoid with rounded ends, slightly bent,
smooth, 5-septate, not constricted, 16-20 x 4-5—-6u, 1—-2-seriate in the ascus. Paraphyses
hyaline, filiform, continuous, equalling the asci.

Associated with Septoria hardenbergiae Sacc. and Pleospora herbarum Rabh.

(110) LrprosSPHAERIA MiIcHoTIZ (Westd.) Sacc., Fungi ital. t. 279; Sace. Syll. Fung.,

2:58, 1883.

On dead culms of Bromus gussonii, Meningie, South Australia, L. D. Williams
(WARI 3525, p.p.).

This specimen corresponds closely with the revised description given by H. Muller
in Sydowia, 4:253, 1950.

(111) LreproSPHAERIA RESTIONIS Hansf., n. sp.

Perithecia in corticem immersa, subaggregata, atra, ampullacea vel subglobosa, usque
ad 180u diam., et 200u alt., ostiolo haud prominente, obtuse conoideo praedita. Paries
perithecii pluristratosus, extus atrobrunneus, intus hyalinus, in massam parenchymaticam
hyalinam centralem transeuns. Asci numerosi, erecti, cylindracei, apice rotundati
inerassatique (—5u), sessiles, 8-spori, 70-80 x 15-194. Paraphyses hyalinae, furcatae,
apice leniter clavulatae, continuae vel indistincte septatae, deorsum circa 1m crassae.
Sporae oblique 1—2-seriatae, ex hyalino brunnescentes, fusoideae utrinque rotundatae,
3-septatae, leniter constrictae, leves, 16-19 x 7—8u, cellula subapicale majore.

Hab. in caulibus Restionis australis, Pipers Bridge, Kosciusko, New South Wales,
A. Costin 63, April 1947.

Perithecia completely immersed in the cortex with only the obtusely conoid ostiole
reaching the surface and not protruding, closely scattered and sometimes sub-connate, -
on greyish-white areas of the stem, where the killed epidermis easily separates from
the cortex; black, flask-shaped to subglobose, up to 2004 high and about 180u diam.
Wall of perithecium of several layers of dark brown parenchyma, passing inwards to
the delicate hyaline ground tissue, which at first completely fills the interior and is
replaced later almost entirely by the developing asci, leaving only a few, hyaline,
branched and anastomosing “paraphyses”, often attached to the “wall” at both ends, or
with their apices slightly clavulate and free, indistinctly septate, about lu thick below.
Asci numerous, basal, erect, maturing in succession and then collapsing, cylindric with
rounded apex thickened up to 5H when immature, sessile, 8-spored, 70-80 x 15-19u.
Spores obliquely 1—2-seriate, becoming brown when fully mature, fusoid with rounded
ends, 3-septate, constricted at the middle septum and less so at the others, the upper half
slightly wider than the lower, with the subapical cell the largest, epispore smooth,
16-19 x 7-8uy.

Mixed with the perithecia are conidial pycnidia, smaller than the perithecia and
with thinner walls, lined with hyaline parenchyma; pycnospores borne direct on short
sterigmata formed in the innermost wall layer, cylindric with rounded ends, hyaline,
single and terminal on the sterigmata, smooth, continuous, 4-5 x lu.

(112) Lrnocarpon sStTIpArE Hansf., n. sp.
Perithecia dispersa vel laxe gregaria, nigra, plus minusve globosa, glabra, 300-500u
diam., superne ostiolo obtuse cylindraceo, usque ad 150u alt. et 100u diam. ornata;

J

122 AUSTRALIAN FUNGI, II,

paries pluristratosus, atrobrunneus, ex cellulis 4-15 x 4-10” compositus, circa 30m crassus,
intus subhyalinus. Asci numerosi, cylindracei, apice rotundati et incrassati (-5y),
deorsum in pedicello attenuati, 8-spori, usque ad 180 x 10-144. Sporae parallelae, rectae,
hyalinae vel in massa dilute olivaceae, filiformes, apice rotundatae, deorsum lenissime
attenuatae, basi rotundatae, 100-140 x 3-4u, 11-13-septatae, haud constrictae, leves.

Hab. in culmis emortuis Stipae spec. indet., Meningie, South Australia, WARI 3488,
leg. L. D. Williams.

Perithecia scattered or loosely gregarious, black, more or less globose, immersed,
with projecting short cylindric obtuse ostiole up to 1504 long by about 1004 diam.; body
300-500 diam.; wall of several layers of angular dark brown cells, 4-15 x 4-10, the
whole wall about 30u thick, lined with paler to hyaline compressed cells. The ostiole is
lined with short, upward directed, hyaline periphyses. Asci numerous, cylindric with
rounded apex which is thickened to 4-5u, leaving a rather long, narrow central canal in
immature asci, this disappearing more or less completely at maturity, narrowed below
into a stalk portion about 3u thick and 20-30. long, 8-spored, the complete ascus up to
180 x 10-144. Paraphyses numerous, filiform, delicate, hyaline, septate, about 1» thick,
apparently simple, not expanded at the apex. Spores parallel and straight in the ascus,
filiform, hyaline individually, pale olivaceous in mass, rounded at the apex, very slightly
attenuate downwards to the rounded base, 100-140 x 3—4u, 11—-13-septate, the septa very
thin, not constricted, smooth, becoming more or less bent to curved after discharge. No
conidial stage seen.

(113) LiINocARPON WILLIAMSII Hansf., n. sp.

Perithecia immersa, dispersa, nigra, globosa vel lateraliter subcompressa, usque ad
180u diam., ostiolo prominulo, breviter obtuse conoideo, poro pertuso, glabro. Paries
perithecii membranaceus, lentus, atrobrunneus, ex cellulis polygonalibus 8-124 diam.,
compositus, 2-—3-stratosus. Asci cylindracei, apice rotundati, deorsum in stipitem
filiformem attenuati, 8-spori, usque ad 180 x 8—-10u. Sporae parallelae, rectae, filiformes,
hyalinae, leves, rectae vel flexuosae, indistincte tenuiter 5—7-septatae, usque ad 130yu
longae, apice 2u crassae et rotundatae, deorsum lenissime attenuati, basi rotundati et lu
crassae, in massa dilutissime olivaceae. Paraphyses numerosae, filiformes, hyalinae,
simplices, indistincte septatae, lu cr., ascos subaequantes.

Hab. in culmis emortuos Graminearum spec. indet., Meningie, South Australia,
WARI 3542, leg. L. D. Williams.

Perithecia immersed, scattered, black, globose to somewhat flattened laterally, up to
180u diam., with slightly erumpent, short, obtusely conoid ostiole pierced by a pore and
lined with very short, hyaline, upward-directed periphyses. The perithecia are quite
glabrous, and have a membranaceous, tough, dark brown wall, consisting of 2-3 layers
of flattened, polygonal cells, 8-124 diam., irregularly arranged save in the ostiolar region
where they are more regularly longitudinal. Asci numerous, cylindric with rounded
apex, attenuate below into a narrow elongate (-—30u) stipe, 8-spored, the wall very thin
save in immature asci, where it is thickened at the apex to leave a very narrow internal
canal, which disappears completely at maturity, up to 180 x 8-10u. Spores parallel and
straight in the ascus, becoming more or less curved after discharge, filiform, gradually
attenuate from the rounded apex, 2u wide, to the base (lu wide), hyaline, smooth,
indistinctly and thinly 5—7-septate, up to 130u long, when fully mature appearing faintly
olivaceous in mass. Paraphyses numerous, filiform, hyaline, simple, indistinctly septate,
lu thick, equalling or somewhat shorter than the asci, and closely resembling the spores.

(114) MycoSPHAERELLA SUAEDAE-AUSTRALIS Hansf., n. sp.

Perithecia in caulibus dispersa, immersa, nigra, punctiformia, circa 1504 diam.,
glabra, sursum in ostiolum brevum papillatum nigrum glabrum attenuata, haud
prominula; paries perithecii tenuiter membranaceus, 1—2-stratosus, ex cellulis atro-
brunneis angulosis compositus, deorsum pallidior. Asci fasciculati, aparaphysati,
plerumque incurvati, subcylindracei, apice incrassati (-5u), rotundati, deorsum in

BY C. G@. HANSFORD. 123

stipitem brevem attenuati, usque ad 60 x 13, 8-spori. Sporae pluriseriatae, parallelae,
ellipsoideae utrinque rotundatae, hyalinae, leves, 1-septatae, haud constrictae, 18-20 x
3-3 bu.

Pyenidia (SEPTORIA SUADAE-AUSTRALIS Hansf., sp. n.): Pyenidia laxe vel subdense
dispersa, nigra, punctiformia, immersa, glabra, usque ad 150u diam., apice poro lato
pertusa; paries pycnidii tenuiter membranaceus, ex cellulis angulosis brunneus com-
positus. Pycnosporae filiformes, rectae vel curvulae, hyalinae, 3-septatae, leves, haud
constrictae, 45-58 x 3u, utrinque lenissime attenuatae.

Hab. in caulibus emortuos Suaedae australis, Meningie, South Australia, WARI
3490, leg. L. D. Williams. '

Neither stage of this fungus causes spots on the dead stems of the host. The
perithecia are scattered, immersed, black, punctiform, about 1504 diam., with slightly
papillate, blunt ostiole reaching the surface of the epidermis and pierced by a pore,
glabrous; wall thinly membranous, of 1-2 layers of dark brown angular cells, paler
below. Asci fasciculate, aparaphysate, mostly incurved, subcylindric with broadly
rounded apex thickened to 54 when immature, attenuate below into short nodose stipe,
up to 60 x 13u, 8-spored. Spores multiseriate and parallel in ascus, overlapping, narrowly
ellipsoid with slightly attenuate rounded ends, hyaline, smooth, 1-septate, not constricted,
18-20 x 3-33u.

Pyenidial stage: Spots none; pycnidia loosely to rather closely scattered on the
dead stems, black, punctiform, immersed save for the apex which penetrates the
epidermis and has a rather wide pore, glabrous, up to 150u diam. Pyecnidial wall thinly
membranous, of brown angular cells, darker around the apex, passing to subhyaline
deeper within the host tissue. Pycnospores filiform, straight or bent, hyaline, smooth,
3-septate, not constricted, 45-58 x 3u, very slightly attenuate to each rounded end, often
slightly more towards the apex. Sporophores not seen.

The pycnidia occur mixed with the perithecia.

(115) PLEOSPORA HERBARUM (Pers. ex Fr.) Rabh., in Herb. Myc. 2: no. 547.

On dead scape, of Allium cepa, Meningie, South Australia, L. D. Williams, WARI
3539.

This is on the type host of the species; the spores are transversely 7-septate, with
usually two longitudinal septa in the larger cells, and measure 28-35 x 15-17u. Other
South Australian collections more or less closely matching the above are:

On dead wood of Limonum sp., Meningie, L. D. Williams, WARI 3546; on dead stem
Dianthus sp., l.c., WARI 3495; on dead stem Lupinus paludosus, l.c. WARI 3497; all
collected by L. D. Williams.

(116) PLEOSPORA SCHIRROIDES Sacc., Michelia 2:68, 1880 (prox.).
On dead leaves Stipa sp., Meningie, S. Australia, L. D. Williams, WARI 3480.
Spores transversely 3-septate, with 1 longitudinal septum in one or both central
cells, 18-22 x 7—9u.

(117) PLEoSpoRA vULGARIS Niessl, Verh. Naturf. Ver. Brunn, 3:27, 1864.
On dead stem Geranium sp., Meningie, South Australia, L. D. Williams, WARI 3538.
This specimen has 5-septate and longitudinally 1-septate spores, measuring 16-18 x
7-8.

(118) PSEUDOSPHAERELLA PERSOONIAE (P. Henn.) Hansf., comb. n.

= Mycosphaerella persooniae P. Henn., Hedwigia, 42:81, 1903. = Sphaeretla
persooniae (P. Henn.) Sacc., Syll. Fung., 17:639, 1905.

The conidial stage was described by Hennings, l.c., 42:87, 1903, as Dichomera
persooniae.

Specimens examined: On Persoonia salicina, Nowra, New South Wales, G. Samuel,
February 1922, WARI 2030; Pennant Hills, N.S.W., L. Fraser, July 1953, WARI 3862.

The fruiting structures of the fungus are developed on epiphyllous dark spots up to
8 mm. diam., scattered over the host leaf; the colour of these spots is mainly due to a

124 AUSTRALIAN FUNGI, II,

subcuticular plate of 1-2 layers of dark brown, erect, rectangular cells, up to 20u deep,
extending in places between the epidermal cells to reach the palisade tissue, above which
it breaks up into single hyaline hyphae, without haustoria. This subcuticular mycelial
plate is connected around the developing fructifications, directly through the cuticle by
fine dark filaments, with an external mycelium on the leaf surface, of pale brown,
septate, exhyphopodiate, crooked, much branched and reticulate hyphae 3-54 wide.

The conidial stage can be construed either as a widely open pycnidium or as an
acervulus; it develops first as a local thickening of the dark subcuticular mycelial
plate, then forming a flattened-globose loculus, which raises, and finally bursts the host
cuticle, to become widely open and surrounded by the remains of its “wall”; the latter
appears to be derived almost entirely from the subcuticular mycelium. The base of the
loculus consists of short erect cells of the stroma, which bud off single conidia from
their apices, and no sterigmata or conidiophores are differentiated. Conidia dark
brown, ovate-ellipsoid, rounded at the apex, very slightly truncate at the base, smooth
ov indistinctly roughened-granulose on the surface, 1-(3)-septate, not constricted, 12-19 x
6-7u. A few conidia are muriform aggregations of 3—4 cells, but these are regarded as
abnormal. The acervuli when single reach 260u diam., but are frequently more or less
confluent in small groups up to 1 mm. diam.

The ascomata develop in a manner similar to the pyenidia or acervuli, from local
thickenings of the subcuticular mycelial plate, which enlarge to burst through the
cuticle and then become globose, black and hard, up to 2004 diam., usually containing
a single loculus, and scattered unevenly over the surface of the leafspot, becoming very
brittle when fully mature, without a definite pore or ostiole. The outer wall is
parenchymatous, the cells black and with rather thickened walls on the exterior surface,
the inner layers passing gradually into a central mass of hyaline, delicate parenchyma.
In the base of this central ground tissue a rosette of asci develops, without paraphyses,
for a long time remaining surrounded by the residue of the ground tissue, and the
individual asci sometimes separated by strands of this, simulating septate paraphyses.
The whole group of asci can be forced out from slightly immature ascomata more or less
intact and enclosed within the hyaline ground tissue. Asci ellipsoid-saccate, rounded-
attenuate and thickened up to 4u at the apex, with some trace of a rather wide “internal
canal’, slightly constricted at the shortly nodose-stipitate base, straight or bent, 8-spored,
about 50 x 154 when fully mature. Spores 2—3-seriate and somewhat parallel within the
ascus, hyaline, ellipsoid with rounded ends, 1-septate, not constricted, the cells more or
less equal, 15-16 x 3-5—-4-5u.

(119) AScCOBOLUS GLABER Pers. in Romer’s Neues Mag. Bot., 1:115, 1794.
On cowdung, Meningie, L. D. Williams, WARI 3516.

(120) ASCOPHANUS CARNEUS (Pers.) Boud., Ann. Sci. Nat., 5:10:250, 1869.

On sterilized soil in greenhouse, WARI, July 1953. These specimens differ from
Seaver’s description (N. Amer. Cup-Fungi, 1:115, 1942) in that the spores remain
permanently quite smooth and no orange granules are contained in the paraphyses,
though these appear pink in mass.

Also collected on dung, Meningie, South Australia, June 1953, L. D. Williams 31.

(121) ASCOPHANUS GRANULIFORMIS (Crouan) Boud., Ann. Sci. Nat., 5:10:245, 1869.
On cowdung, Meningie, 8. Australia, L. D. Williams, WARI 3516.

(122) ASCOPHANUS OCHRACEUS (Crouan) Boud., Ann. Sci. Nat., 5:10:247, 1869.
On cowdung, Meningie, S. Australia, L. D. Williams, WARI 3517.

(123) GODRONIA DAVIESIAE Hansf., n. sp.

Apothecia dispersa vel 2-3-aggregata, per peridermidem erumpentia tum cortice
disrupto circumdata, sessilia, circa 5004 diam., extus nigra, levia, tenuiter cupulata;
hymenium applanatum, molliter carnosum, in sicco obdurescens, griseum. Asci cylindracei,
deorsum leniter attenuati, apice abrupte rotundati, 8-spori, usque ad 200 x 6-8u. Sporae

BY C. G. HANSFORD. 125

parallelae, in asco rectae, liberae varie curvatae, hyalinae, filiformes utrinque obtusae,
usque ad 140 x 1-5-2u, obsolete multiseptatae, segmentibus circa 3yu longis. Paraphyses
filiformes, numerosae, simplices vel in medio furcatae, ascos aequantes vel leniter
superantes, 1-5—2u crassae, verisimiliter continuae, hyalinae.

Hab. in ramulis emortuos Daviesiae brevifoliae, Meningie, South Australia, L. D.
Williams, WARI 3520. :

Apothecia on dead twigs, more or less scattered or in groups of 2-3, erumpent
through the bark and then surrounded by the broken cortex, sessile, about 500” diam.;
externally black, the hymenium grey and surrounded by the rather thick irregularly
shallow-cupulate exciple; hymenium flat, soft fleshy when moist, drying hard. Asci
eylindric, somewhat attenuate below into stalk portion which may be very short,
abruptly rounded at apex, 8-spored, up to 200 x 6—-8u. Spores parallel and straight in
ascus, becoming variously curved after discharge, hyaline, filiform with obtuse ends, up
to 140 x 1-5-2u, with many indistinct cross-septa, the contents being divided into
segments about 3u long. Paraphyses filiform, numerous, simple or once-furcate at or
near the middle, equalling or somewhat exceeding the asci, 1-5-2u thick, apparently
continuous.

(124) GORGONICEPS CLELANDIIT Hansf., n. sp.

Apothecia dispersa vel raro 2-aggregata, fiepinata vel brevissime stipitata, sursum
cupulata, carnosa, 1-2 mm. diam. et 1-5-2-5 mm. alt., extus flavida vel fulva, margine
albo-pruinosa, haud setosa, levia vel longitudinaliter subtiliter striato-rugulosa, deorsum
in stipitem brevem obconicam attenuata. Hymenium concavum, margine incurvato
circumdatum, flavum vel aureum, leve. Asci leniter clavulati, usque ad 130 x 6—8y, apice
rotundati, inoperculati, deorsum attenuati, 8-spori. Paraphyses numerosae, filiformes,
simplices, verisimiliter continuae, apice haud incrassatae, ascos aequantes. Sporae
parallelae, filiformes, 45-55 x 2u, utrinque lenissime attenuatae, continuae, hyalinae,
intus granulosae.

Hab. in ligno emortuo, Dorrigo, New South Wales, Jan. 1918, J. B. Cleland, WARI
3125.

Apothecia scattered, or rarely 2-aggregate, turbinate or very shortly stipitate,
cupulate above, fleshy, firm, 1-2 mm. diam. and 1-5-2-5 mm. high, the exterior pallid
yellow to yellow-brown, becoming white-pruinose around the margin, but not hairy or
setose, smooth or finely longitudinally striate to rugulose, contracted below into a short
Obconic stem portion, without evident mycelium around the base. Hymenium concave,
surrounded by the even, rather thick, incurved margin, shallowly cupulate to almost
plane in old specimens, from yellow to bright orange, smooth. Asci narrowly clavate,
up to 130 x 6—-8u, rounded at the apex, inoperculate, 8-spored. Paraphyses numerous,
filiform, simple, doubtfully septate, not enlarged at the apex, equalling the asci; the
orange colour of the hymenium is due to the pale yellow coloration of the paraphyses
and asci as seen under the microscope, this colour extending throughout the tissues of
the apothecium. Spores parallel and often spirally twisted inside young asci, filiform,
45-55 x 2u, Slightly attenuate to lu» at the rounded ends, with finely granular contents,
apparently not becoming septate, quite hyaline.

(125) HELOTIUM CYATHOIDEUM (Bull.) Karst., Bot. Soc. Fauna & Fl. Fenn., 11:237, 1871.
On dead wood, Blue Mts., New South Wales, J. B. Cleland, May 1915, WARI 2736.

(126) HuMARINA ARANEOSA (Bull. ex Fuckel) Seaver, N. Amer. Cup-Fungi, 1:130, 1942.
On sandy soil, National Park, New South Wales, J. B. Cleland, WARI 2755.
Apothecia scattered, stipitate, expanded above and discoid to shallowly cupulate

with even margin, sometimes slightly distorted, orange-red outside, the hymenium

smooth and orange-red, to 7 mm. diam. and 5 mm. high, the stem portion 3-4 x 1 mm.,

dilated above and slightly bulbous at the base in the sand, concolorous. Asci cylindric,

to 180 x 10-11, attenuate below into a long stipe portion, 8-spored, operculate. Spores
obliquely 1-seriate, long remaining hyaline but finally becoming brownish, smooth,

126 AUSTRALIAN FUNGI, II,

ellipsoid with the basal end rather more acute than the apex, 15-19 x 6-5-7-5y.
Paraphyses filiform, hyaline, equalling the asci, 2u thick, not expanded above, straight,
septate. The whole texture is firmly fleshy.

(127) HuMARINA SEMIIMMERSA (Karst.) Seaver, N. Amer. Cup-Fungi, 1:131, 1928.

On rotting leaves and soil, Mt. Lofty, South Australia, July 1921, J. B. Cleland,
WARI 2540.

Apothecia gregarious or scattered, sessile or subsessile, to 10 mm. diam., expanded
and shallowly cupulate to almost discoid, pale yellow externally, subfloccose, soft fishy,
with somewhat irregular pale margin. Hymenium dark honey-coloured, concave to
almost plane; asci cylindric, up to 250 x 15-20u, 8-spored, operculate, with long stalk
portion which is more or less suddenly contracted into a short foot at the base. Spores
1-seriate, oblique, straight to almost transverse, occasionally 2-seriate in upper part of
ascus, ellipsoid, hyaline, quite smooth, 18-20 x 9-llyw. Paraphyses slender, septate,
hyaline, slightly yellowish and expanded to 3-4u at the apex.

(128) HuMARINA CONVEXULA (Pers.) Seaver, N. Amer. Cup-Fungi, 1:128, 1942.
On damp soil, R. Tug, Central Australia, Mayo Expedition 1079, June 1953.

(129) LACHNELLA CLELANDII Hansf., n. sp.

Apothecia stipitata, circa 9 mm. alt., apice dilatata, discoidea vel subcupulata,
usque ad 5 mm. diam., stipite circa 7 x 0-8 mm.; extus brunnea, setosa; setae usque ad
1000 x 8u, pluriseptatae, atrobrunneae, rectae, simplices, acutae, subrigidae. Hymenium
subcupulatum vel planum, margine tenue regulare circumdatum, leve, luteum, ceraceum.
Asci cylindracei, inoperculati, breviter stipitati, apice rotundati, 8-spori, usque ad 150 x
9-11u. Sporae l-seriatae, plus minusve obliquae, ellipsoideae, utrinque attenuato-
. rotundatae, continuae, hyalinae, leves, 15-19 x 5-7.

Hab. in terra, National Park, South Australia, July 1925, J. B. Cleland, WARI 2542.

Apothecia stipitate, about 9 mm. in total height, with expanded, shallowly cupulate
or subdiscoid apex, up to 5 mm. diam., rather suddenly contracted into a stem about
7 x 0-8 mm.; externally brown and clothed with setae which continue down the stem
to soil level. Setae dark brown, up to 1000 x 8u, acute, simple, stiff, many-septate,
suberect; beneath the setae the stem appears loosely white-striate. Hymenium shallowly
cupulate to almost plane, surrounded by a very thin, regular margin composed of the
thin exterior brown layer of the apothecium, slightly incurved around the hymenium,
which is smooth, yellow-brown and waxy. The upper part of the apothecium is soft
fleshy-gelatinous, the subhymenium being composed of very pale yellowish, contorted
and much interwoven, septate hyphae. Asci cylindric, inoperculate, contracted below
into rather short stipe, 8-spored, 150 x 9-llu. Spores 1-seriate, usually slightly oblique
and overlapping, narrowly ellipsoid with attenuate-rounded ends, continuous, quite
hyaline, permanently smooth, 15-19 x 5-T7u.

A second collection from the same locality, also by Cleland, May 1926, is preserved
at the WARI as no. 2545, growing on wood.

(130) LAMPROSPORA TETRASPORA Hansf., n. sp.

Apothecia gregaria, usque ad 5 mm. diam., cupulata vel subdiscoidea, flavida,
carnosa, margine crassa, regulare; extus levia, concolorata. Asci cylindracei, breviter
stipitati, usque ad 180 x 17-19u, operculati, (2)-4-(5)-spori. Sporae 1-seriatae, globosae,
hyalinae, leves, 15-174 diam., grosse 1-guttulatae. Paraphyses numerosae, filiformes,
indistinecte septatae, circa 2u cr., apice haud incrassatae.

Hab. in terra, Milson Island, Hawkesbury R., New South Wales, May 1915, J. B.
Cleland, WARI 2737.

Apothecia gregarious, sessile, attached by hyphae in the centre of the base only, up —
to 5 mm. diam., deep yellowish when moistened after drying (possibly orange when
fresh), fleshy, rather tough, shallowly cupulate to subdiscoid-expanded, the margin thick,
even, fertile. The exterior is smooth and concolorous with the hymenium. Asci cylindric,

BY C. G. HANSFORD. 127

narrowed below into a short foot, up to 180 x 17-19, operculate, mostly 4-spored when
mature, a few with 2 or 5 spores, but forming 8 spores at first. Spores 1-seriate, globose
to slightly ellipsoid, hyaline, filled with a single large oil globule, quite smooth, rather
thin-walled, 15-17% diam. Paraphyses numerous, filiform, doubtfully septate, not enlarged
at apex, about 2u thick, exceeding the young asci, but the mature asci slightly protruding
from amongst them.

(181) LAMPROSPORA LEIOCARPA (Curr.) Seaver, Mycologia, 6:21, 1914.

On soil, Mt. Lofty, South Australia, J. B. Cleland, WARI 3137.

Apothecia gregarious or scattered, becoming shallowly cupulate, regular or irregular,
2-3 cm. diam., finally almost discoid, brown to almost black, smooth or somewhat
roughened externally. Hymenium smooth or somewhat wrinkled, dark brown to almost
black. Asci cylindric, tapered below into a rather long stipe, 8-spored, up to 190 x 10-12y,
operculate. Spores 1-seriate, globose, hyaline, smooth, 8-10u diam. Paraphyses numerous,
often fasciculate and adherent at the apices, filiform, with globose apex swollen to 10y,
pale yellow-brown below, much darker at apex, septate.

(132) LaAmprRospora sp. near L. constellatio (B. & Br.) Seaver.

On sandy soil, Mt. Compass, Oct. 1924, J. B. Cleland, WARI 25438; ditto, Encounter
Bay, May 1924, J. B. Cleland, WARI 2550.

Apothecia scattered or gregarious, up to 5 mm. diam., coral red outside with darker
red hymenium, in old dried specimens the colours fading to uniform dull yellowish-buff,
discoid-expanded, very slightly concave when fully mature, about 8004 thick, without
distinct sterile margin, the edge rather thick and regular, fertile. Hymenium fleshy in
texture; asci cylindric, contracted below into rather short stipe, up to 170 x 20u, 8-spored,
very thin-walled, operculate. Spores 1-seriate, hyaline, permanently smooth, perfectly
globose, the wall about ly thick, 15-174 diam. Paraphyses filiform, hyaline, septate, not
thickened at apex, with granular contents, often bent to curved near the apex.

(1383) PATELLA SCUTELLATA (L.) Morgan, Journ. Mycol., 8:187, 1902.
On soil and wood, Katoomba, New South Wales, 1916, J. B. Cleland, WARI 3132.

(134) PATELLA sp. near P. albo-cincta (B. & C.) Seaver, N. Amer. Cup-Fungi, 1:168, 1942.

On soil, Warri, Oct. 1923, J. B. Cleland, WARI 2536.

Apothecia rather deeply cupulate, up to 10 mm. diam. and 12 mm. high, sessile,
indistinctly longitudinally plicate, firm fleshy, dark brown in dried specimens, the colour
dissolving on moistening and leaving the moist apothecium orange-brown both externally
and internally; the exterior clothed with hyaline hairs forming a short tomentum which
does not extend as bristles above the margin; the latter slightly incurved and sterile;
hairs up to 150 x 10-124 wide at base, often suddenly constricted above the base or near
the middle and thence gradually attenuate to the narrowly rounded apex (2u wide),
the wall 1-5u thick, smooth, 2—3-septate. Asci cylindric, up to 300 x 18-20u, thin-walled,
operculate, slightly narrowed below into stipe, 8-spored. Spores obliquely 1-seriate,
ellipsoid, 20-25 x 12-14u, continuous, the wall about lu thick, becoming finely verruculose
with minute blunt tubercles, quite hyaline.

(135) PATELLA sp. near P. theleboloides (Alb. & Schw.) Seaver, N. Amer. Cup-Fungi,

1:170, 1942.

On damp hay, Meningie, S. Australia, L. D. Williams, WARI 3492.

Apothecia scattered to closely gregarious, shallowly cupulate, yellow, sessile or
contracted below into the rudiment of a stem-like base, smooth, circular to somewhat
irregular in outline, up to 2 mm. diam. Exciple concolorous with hymenial disc or
slightly more brownish-yellow, smooth, with scattered erect hairs around the margin;
cells of outer layer rounded-angulose, thin-walled, yellowish, up to 40u diam., irregularly
arranged. Setae erect-spreading, simple, straight, pale yellow under microscope, yellow-
brown under lens, up to 300u long, gradually attenuate from the base 20 thick to the
simple acute apex, the wall 1—1-5y thick, smooth, 2—5-septate. Hymenium flat to concave,

128 AUSTRALIAN FUNGI, II,

yellow, the subhymenial tissue slightly more orange-yellow; asci cylindric, rounded at
apex, slightly attenuate below into a stipe, 8-spored, up to 230 x 12u, operculate.
Paraphyses filiform, hyaline, 24 wide, thinly septate, slightly swollen to 44 at apex,
contents hyaline. Spores 1-seriate, ellipsoid, hyaline, perfectly smooth, 14-16 x 9-1lp.

(136) PrzizA CLYPEATA Schw. (prox.)

On wood, Mt. Wilson, New South Wales, J. B. Cleland, June 1915, WARI 3141;
Hogans Brush, Gosford, N.S.W., J. B. Cleland, WARI 3140.

Apothecia single, discoid to shortly stipitate, becoming widely open with slightly
incurved margin, dark brown, drying almost black, 25-50 mm. diam. and 10-20 mm. high,
soft fleshy in texture, externally smooth or slightly rugose on under surface, centrally
attached to the wood. Hymenium dark brown to almost black when dry, the colour
being contained as granules in the paraphyses and soluble to some extent in warm
water; smooth, even; asci cylindric but tapered below into a long stipe portion, up to
about 400 x 15-17y, 8-spored, thin-walled, operculate. Spores 1-seriate, cymbiform with
rounded ends, hyaline, perfectly smooth with granular contents or 2—3-guttulate, 25-35 x
12-17u, the wall about lu thick. Paraphyses numerous, slightly exceeding the asci and
sometimes somewhat agglutinate at their swollen apices (5—-7u), simple, septate, with
brownish internal granules.

(187) PEZIZA SYLVESTRIS (Boud.) Sacc. & Trott, Syll. Fung., 22:612, 1913.
On humus under Pinus, Mt. Gambier, South Australia, Oct. 1953, H. Lower; not
present in the more open parts of the woods.

(188) Prziza sp. near P. brunneoatra Desm.

On soil, Mount Lofty, S. Australia, Sept. 1920, J. B. Cleland, WARI 3133.

Apothecia scattered, cupulate, subsessile, up to 10-12 mm. diam. and about 6 mm.
high, centrally attached by a short stipe-like portion about 1 mm. long and 2-3 mm.
diam., which is enlarged into a bulbous root immediately below the soil surface to
4-5 mm. diam., from which mycelium passes outwards into the soil. Apothecia glabrous,
externally brown, fleshy, soft, about 1 mm. thick; hymenium dark brown, smooth, the
margin finely crenulate and slightly paler. Asci cylindric, to 280 x 17-18, operculate,
8-spored, slightly contracted below into elongate stipe. Spores i1-seriate, ellipsoid,
continuous, hyaline, 22-25 x 12-154, becoming sparsely and finely verruculose when
mature. Paraphyses numerous, brown in mass, simple, clavate-enlarged at the apex to
Tu, covered there with yellow-brown granular matter, the contents also with fine
yellowish granules, septate. The contents of mature, shed spores are also sometimes
faintly yellow-granular.

(139) SARCOSPHAERA AMMOPHILA (Dur. & Lev.) Seaver, N. Amer. Cup-Fungi, 1:333, 1942.

In sand, Mt. Compass, South Australia, J. B. Cleland, Oct. 1924, WARI 3117.

This differs slightly from Seaver’s description: the spores become pale yellow-brown
when fully mature and have minute scattered tubercles on the surface, measure 14-16 x
9-104; paraphyses pale yellow-brown, darker towards the apex, fasciculate and clinging
together at the apex, septate.

Also collected by J. B. Cleland in sand at Encounter Bay, S. Australia, Sept. 1924,
WARI 3129; and at Narragin, Western Australia, 1926, WARI 3502.

BASIDIOMYCETES.
(140) PUuccINIA COCKAYNEI Cunn., Trans. N.Z. Institute, 54:679, 19238.
On Gentiana diemensis, Kosciusko, New South Wales, A. Costin 17, March 1947.

(141) PUCCINIA EUPHRASIANA Cunn., Trans. N.Z. Institute, 55:6, 1924.

On Huphrasia collina, Mt. Kosciusko summit, New South Wales, March 1947, A.
Costin 16.

This collection contains only the uredo-stage, and no paraphyses were found, so that
the determination is rather doubtful.

BY C. G. HANSFORD. 129

(143) PuccrntaA HreRAcIT (Schum.) Mart., Fl. Mosq., p. 226, 1912.
On Microseris scapigera, Kosciusko, New South Wales, March 1947, A. Costin 14.

(144) Puccrinta RUIZENSIS E. Mayor, Mem. Soc. Neuchatel, Sci. Nat., 5:486, 19138.
On Oreomyrrhus andicola, Kosciusko, New South Wales, Dec. 1946, A. Costin 5.

(145) PUccCINIA ARTHROCNEMI Hansf., n. sp.

II. Uredosori in caulibus et spicibus dispersi, discoidei, epidermide rupta cincti,
aurei, usque ad 0-75 mm. diam., rotundati vel elliptici, pulverulenti. Uredosporae
ellipsoideae, fulvae vel aureae, 29-40 x 21—27u, episporio 2—2-5y cr., laxe echinulato (—ly),
poris germinationibus 8-13, dispersis, leniter papillatis.

III. Teleutosori pauci, dispersi, usque ad 0-5 mm. diam., rotundati, leniter bullati,
atrobrunnei, epidermide rupta cincti. Teleutosporae ellipsoideae vel oblongae, utrinque
rotundatae vel apice subpapillatae vel truncatae, leves, sursum castaneae, deorsum
pallidiores, 44-60 x 29-40u; 1-septatae, non vel leniter constrictae; episporio 2—-2-5u er.,
apice incrassato usque ad 7u; pedicello persistente, hyalino vel apice dilute luteo,
50-110 x 7—-10p, collabascente.

X. Mesosporae paucae, teleutosporis commixtae, subglobosae vel ellipsoideae, 35-45 x
25-34u, apice leniter incrassatae.

Hab. in caulibus et spicibus Arthrocnemi halocnemoidis, Eucolo Gorge, Pimba, S.
Australia, Dec. 1953, J. B. Cleland, WARI 3699.

The uredosori are scattered on the stems and flowering parts, discoid, surrounded
by the broken epidermis, golden brown, up to 750u diam., rounded or elliptic, pulverulent.
Uredospores ellipsoid, yellow to golden brown, 29-40 x 21—27u, sparsely echinulate to 1p,
the epispore 2—24u thick, with 8-13, slightly papillate, paler to hyaline germ pores
scattered all over.

Teleutosori few, scattered, up to 0-5 mm. diam., rounded, somewhat bullate, dark
brown to black-brown, surrounded by the ruptured epidermis, becoming pulverulent.
Teleutospores eventually loosely pulverulent, mixed with a few mesospores but no
uredospores, ellipsoid to oblong, rounded at both ends, or the apex somewhat papillate
or flat and subtruncate, chestnut-brown above, somewhat paler towards the base, smooth,
or in some fluids appearing very finely and indistinetly granulose on the surface, not or
very slightly constricted at the septum, 44-60 x 29-40u; epispore 2-layered, the outer layer
lu thick, paler than the inner, which is 2—24u thick, but thickened around the apex up
to 7w. Germ pore of upper cell apical, of lower just below septum or half-way to the
base, and then slightly papillate. Pedicel persistent, hyaline, or with faint yellow tinge
at junction with spore, 50-110 x 7-10u, thin-walled and collapsing.

Mesospores few, similarly coloured to teleutospores and with similar double wall,
often slightly thickened around the rounded or flattened apiculate apex, 35-45 x 25-34.

(146) TILLETIA sp.

On Aira caespitosa, Myponga, South Australia, Jan. 1929, J. B. Cleland, WARI 2000.

Sori in ovaries, remaining enclosed by the outer membrane of the host, which is
one cell thick, becoming black and powdery, irregularly dehiscent. Spores dark brown,
more or less globose, 18-234 diam., surrounded by a subhyaline wrinkled-reticulate
epispore, the meshes of which are up to 2u high, so that external spore dimensions are
21-28 x 21-234, mostly 22-274 diam. The reticulations of the exospore are irregular in
shape and size, even in fully mature spores, varying from almost hexagonal and about
34 diam., to elongate or ramose, up to 15 x 2u, with divisions between them about 2u
thick.

Hx descr. this is not unlike 7. flectens Lagerh., but which is limited to leaves; there
is no infection on the leaves of the present specimen. Germination of the spores could
not be obtained, so that even the generic determination is open to doubt, pending further
collections for comparison.

130 AUSTRALIAN FUNGI, II,

(147) IvaAgAHYA HORNSEYI Hansf., n. sp.

Receptaculum capitatum, stipitatum; stipes 15-20 x 38 ecm., cylindraceus, basi
attenuatus, extus lacteo-flavidula vel dilutissime puniceus, intus albus, cellulosus, cellulis
usque ad 5 mm. diam., concavus, contextus circa 10 mm. cr., apice dilatatus et revolutus,
usque ad 10 cm. diam., et pileus ferens. Pileus circa 3-5 mm. cr., discoideus, albus,
cellulosus, cellulis 1-3-stratosis, usque ad 2 mm. diam., sursum membrana pallide

Figures 1, 2. Itajahya hornseyi.

1, Vertical section through unexpanded plant, x1. 2, Vertical section through expanded
plant (x4/,), showing on right the gleba prior to dehiscence, on left, the gleba after spore
dispersal.

a, Exoperidium; 6b, mesoperidium, mucilaginous; ec, endoperidial membrane; d, stipe,
showing cellular structure; e, central cavity of stipe; f, pileus, with cellular structure; g, outer
membrane of gleba; h, inner membrane of gleba, bearing the tramal plates (Kk); j, expanded
stem apex, in Fig. 1 slightly off centre.

brunnea viscidula, facile secedente vestitus. Gleba ex apice dilatato stipitis dependens,
annulata, circa 5-8 mm. cr. et usque ad 50 mm. alt., foetida, primo nigra, in expanso
viride demum grisea, deliquescens et laminas imbricatas tramatis relinquens. Basidia
8-spora. Sporae sessiles, subhyalinae, ellipsoideae, leves, continuae, 5 x 2—2-5u, episporio
tenui, massam mucilaginosam, virideo-griseam efformantes.

Hab. in terra, Cadell, R. Murray, South Australia, April 1954, E. Hornsey, WARI
Silas

The fungus originates from the usual type of “egg” of the Phallaceae, buried just
beneath the surface of the sandy soil habitat, smooth, subglobose to obovate, brownish

BY C. G. HANSFORD. 131

to umber in colour, membranous, 4-5 cm. diam. and 4-7 cm. long, with a long basal
rhizomorph extending from the centre of the base deeper in the soil; in the only
specimen seen this rhizomorph was brown, 15 cm. long by 4 mm. diam., smooth, rather
tough, flexible, composed of parallel hyphae with little differentiation between the
surface layer and interior, save that it is white internally. In longitudinal section
(Fig. 1) the peridium is seen to consist of three layers, the outer a rather thick
membrane of closely agglutinated hyphae, the meso-peridium soon degenerating into a
watery mucilage, with some trace of areolae formed by the remains of the original
hyphae, and an inner, white, smooth membrane more or less closely adherent to the
receptacle inside, composed of more or less closely agglutinated hyaline hyphae, many
of which are swollen at the septa, very thin-walled. The receptacle prior to expansion
consists of a central slightly ellipsoid, hollow stem, somewhat expanded and recurved at
the apex, narrowed at the base, of minutely cellular structure, and covered over the
apex with a rounded, thin, separate discoid pileus. Around the edges of the expanded
stem apex depends the annular gleba, enclosed in a thin membranous sheath, consisting
of numerous, more or less horizontal, folded thin tramal plates, on which the basidia are
formed, traversing the solid black spore mass. At maturity (Fig. 2) the stem of the
receptacle elongates within 2—4 hours, bursting the peridium by an irregularly circular
fissure around the edge of the pileus, carrying the upper portion of the peridium. into
the air aS a membranous covering over the pileus; this covering is quite loose and has a
rather ragged edge, soon becoming detached from the pileus itself, especially in rain
storms. The surface of the pileus is smooth and creamy white, and when expanded it
consists of 1-3 layers of small cells, similar to those of the expanded stem. The pileus
‘is held to the stem apex merely by soft mucilage, and is easily detached as a disc,
somewhat incurved over the edge of the expanded stem-apex. The expanded stem is up
to 20 ecm. long by about 3 cm. diam., cylindric above the slightly attenuated and thinner
base, traversed by a central canal about 1 cm. diam., smooth inside, the tissue of cellular
structure, quite soft, the cells up to 5 mm. diam., in several layers. The apex is expanded
into a recurved disc portion, the flat, free edge bearing the annular gleba, which hangs
down for a distance of up to 50 mm. around the stem, from which it is quite free, and at
first is enclosed within a delicate surface membrane. On the inner surface next the
stem this membrane is somewhat thicker and tougher, and bears the more or less
horizontal, imbricate, white tramal plates; the latter remain more or less intact after
the dispersal of the spore mass. The outer part of the covering membrane splits
irregularly, often near the attachment to the stem apex, and may for a time hang down
around the base of the exposed gleba, but normally disappears rapidly. The exposed
spore mass is green in colour, soon darkening to almost black, and in the fresh state
bears numerous droplets of water. The basidia bear star-like groups of usually 8 spores
each; the spores individually are subhyaline to faintly grey in colour, sessile, ellipsoid,
smooth, thin-walled, 5 x 2—2-5u, the epispore slightly mucilaginous. The basidia them-
selves are rather ephemeral, collapsing immediately after spore formation, but the
clusters of spores tend to hang together on the apex of the collapsed basidium. The
tramal plates are at first irregularly folded, but expand and separate as spore discharge
occurs, their somewhat loose and ragged edges finally giving a very rough appearance to
the surface of the gleba. The latter has a very objectionable odour of rotting cheese,
quite characteristic of this species, and very different from the foetid smell of other
Phallaceae.

This is the first record of this genus in Australasia, it having previously been
recorded from South America, Egypt and Palestine.

FUNGI IMPERFECTI.
(148) PHYLLOSTICTA ALYXIAE Hansf., n. sp.
Maculae nullae; pycnidia epiphylla, rarius etiam hypophylla, dispersa, nigra,
subglobosa vel lenticularia, usque ad 300u diam. et 200u alt., primo subepidermalia,
demum epidermidem elevatam circulariter disrumpentia et late aperta. Paries pycenidii

132 AUSTRALIAN FUNGI, I,

parenchymaticus, circa 30-404 ecr., atrobrunneus, intus hyalinus. Sporophora stipata,
erecta, hyalina vel flavidula, continua, simplicia, 15-20 x 2-3-5u. Sporae acrogeneae,
singulae, cylindraceae, apice rotundatae, basi lenissime truncato-apiculatae, plerumque
rectae, hyalinae, continuae, leves, eguttulatae, 15-17 x 3-4y.

Hab. in foliis subvivis Alyxiae buxifoliae, Sandringham, Victoria, Aug. 1900 (? leg.
D. McAlpine), WARI 3729 ex Herb. Dept. Agr. Victoria.

Leafspots none; the pycnidia are scattered evenly over the dying or dead tissues of
the host leaf, usually epiphyllous but sometimes amphigenous, black, globose to lenticular,
up to 300u diam. and 200u high, showing through the elevated epidermis as shining
black spots, eventually bursting the epidermis, often circularly, and becoming widely
open like an acervulus. Wall of several layers of subopaque dark brown parenchyma,
30-404 thick all round the loculus, lined with lighter and thin-walled parenchyma,
opening at the apex by an irregularly rounded pore, and later more widely. Conidiophores
forming a close palisade on the base and lower part of the sides of the pyenidium, erect,
hyaline to yellowish, continuous, simple, 15-204 long, forming spores singly at the apex.
Spores cylindric, hyaline, continuous, smooth, thin-walled, not guttulate, the apex
broadly rounded, the base slightly truncate-apiculate, usually straight, 15-17 x 3-4u.

(149) PHYLLOSTICTA CONFUSA Bubak in Tranzschel & Serebr., Mycoth. Ross. no. 330, 1912,

et in Hedwigia, 57:339, 1916.

On Atriplex elachophyllus, Ooldea, South Australia, Aug. 1922, G. Samuel, WARI
1996.

This specimen differs from the original description in not causing definite leafspots,
the whole leaf being turned yellow, and the pycnidia scattered over it. Pycnidia
epiphyllous, immersed, black, punctiform, membranous, to 200u diam., glabrous, flattened-
globose, about 120u high; wall of several layers of brown, angular cells enclosing an
inner layer of hyaline cells 8-10 x 2—4u, projecting as a palisade into the loculus and
forming conidia direct on their apices; the whole wall 20-30 thick, thicker around the
apex (—40u) and pierced by a round pore 10-154 diam. Pycnospores exceedingly
numerous, filling the whole loculus, hyaline, bacterioid, slimy, adherent in loose masses, ©
straight with rounded ends, continuous, 2—4 x 1-1-5u, eguttulate, smooth, extruded as a
white tendril or mass.

(150) AscocHyTA sp. near A. graminicola Sace.

On dead leaves Stipa sp., Meningie, South Australia, L. D. Williams, WARI 3489 p.p.

Pyenidia seattered, not on leafspots, punctiform, black, depressed-globose, 80-904
diam., with apical pore, glabrous; wall membranous, of a single layer of brown, angular
cells with some traces of linear arrangement, darker around the apex, lighter below,
5-84 diam. Conidiophores not seen. Conidia subfusoid, usually straight, widest in the
middle, slightly attenuate towards both rounded ends, hyaline, smooth, 1-septate in
middle, not constricted, 8-13 x 2-5-3-5u; older spores tend to become greenish-hyaline
and the contents sometimes divided into three parts, thus appearing indistinctly 2-septate.
These spores are smaller than those described by Grove (British Coelomycetes, 1:3438,
1935) as 10-15 x 3—-4u.

(151) AScocHYTA HETERODENDRI Hansf., n. sp.

Maculae amphigenae, usque ad 3 mm. diam., primo brunneae atro-marginatae,
demum cinerescentes. Pycnidia nigra, immersa vel suberumpentia, punctiformia, plus
minusve globosa, glabra, levia, usque ad 140u diam., dispersa vel 2—3-aggregata, sursum
leniter papillata; paries membranaceus, pluristratosus, dilute olivaceus, intus hyalinus.
Sporophora erecta, stipata, simplicia, continua, usque ad 12 x 3yu, hyalina. Conidia
“acrogenea, hyalina, cylindracea, apice rotundata, basi lenissime attenuato-truncata,
1-septata, haud constricta, levia, recta vel curvula, 10-15 x 2-5-8u.

Hab. in foliis Heterodendri alnifolii, Ooldea, Aug. 1922, G. Samuel, WARI 1987.

Leafspots amphigenous, to 3 mm. diam., at first brown with darker marginal line,
finally becoming cinereous and slightly rugulose with the suberumpent pycnidia.

BY C. G. HANSFORD. 133

Pyenidia black, punctiform, subepidermal, more or less globose, glabrous, smooth, up to
140u diam., single or 2-3-aggregate and subconfluent; wall little differentiated from the
surrounding mycelial hyphae save by slightly darker colour, membranaceous, of several
layers of pale olivaceous cells, lined with other hyaline layers, the innermost bearing a
close palisade of conidiophores. The pycnidia are surrounded outside by a loose
plectenchymic stroma of hyaline hyphae, extending downwards between the palisade
cells and then dissolving into loose hyphae in the mesophyll. Conidiophores erect,
closely crowded, simple, continuous, up to 12 x 3u, forming single apical conidia. Conidia
hyaline, cylindric, straight or bent, rounded at the apex, slightly attenuate-truncate at
the base, 1-septate, not constricted, smooth, 10-15 x 2-5—-38uy.

(152) ASCOoCHYTA VERBENAE Siem., Bull. Mus. Caucasie, Tiflis, 12, extr. p. 4, 1919.

On dying stem of Verbena sp., Meningie, South Australia, L. D. Williams 23.

Pyenidia subepidermal, with punctiform ostioles, globose, black, membranous,
smooth, up to 90u diam. Wall parenchymatous, of a single layer of brown angular cells,
darker around the apical pore, which is 15-20% diam. Pycnospores single and acrogenous
on short conidiophores lining the loculus; oblong with obtuse ends, straight or slightly
bent, hyaline, thin-walled, becoming 1-septate and sometimes slightly constricted in the
middle, 8-12 x 3-3-5u.

(153) CoNIOTHYRIUM SP.

On dead leaf Stipa sp., Meningie, South Australia, L. D. Williams, WARI 3489 p.p.

Pyenidia seattered, not on leafspots, minute, punctiform, immersed, black, globose
to somewhat flattened, about 804 diam. and 60—70u high, the ostiole scarcely protruding,
merely reaching the leaf surface and pierced by a round pore about 15y diam.; wall
smooth, membranaceous, glabrous, parenchymatous, olivacecus, darker brown around
the apex, about 5u thick, consisting of angular cells in 1-2 layers, 7-94 diam., lined with
a hyaline inner layer, which forms the conidia on the inner surface; conidiophores not
evident. Conidia soon becoming chestnut-brown, oblong with rounded ends, continuous,
smooth, 6-8 x 5-6u, the wall about 1p thick, contents clear, not guttulate.

This appears to be close to C. fuegiana Speg., described on rotting leaves and culms
of Poa spp., South America, but no authentic material of this has been available for
comparison.

(154) STAGONOSPORA ACACIAE Hansf., n. sp.

Pyenidia primo singulariter dispersa, demum laxe aggregata, in areolas pallidibus
indeterminatas, plerumque epiphylla, immersa, atra, levia, membranacea, usque ad 400
diam. et 250u alt.; paries sursum opace atrobrunneus, pluristratosus, parenchymaticus,
deorsum pallidior et tenuior, cellulis rotundato-angulosis, circa 10 x 8u, apice poro circa
154 diam. pertusus. Sporophora stipata, simplicia, hyalina, continua, circa 15 x 3—4u.
Pycnosporae singulariter acrogeneae, rectae vel curvulae, utrinque rotundatae, leves,
2-3-septatae, haud constrictae, 18-24 x 3-5-4-5uw.

Hab. in foliis Acaciae elatae, Burragorang Valley, New South Wales, L. Fraser,
Feb. 1953.

Pyecnidia at first single, then loosely grouped, on indefinite pale areas of the leaf,
amphigenous, mostly epiphyllous, completely immersed in the leaf, with the ostiole and
terminal pore opening through the epidermis, becoming slightly raised above the level
of the leaf when fully mature. The pycnidia occupy the upper half of the leaf thickness
When epiphyllous, and force aside the remains of the palisade tissue, remaining quite
separate; there is a faint reddish tinge in the contents of the conidiophore layer and
the young conidia, not apparent when these are mounted separately. Pycnidia black
around the apex, smooth, glabrous, thin-walled, up to 400u diam. and 250u high; the wall
in the upper part opaque dark brown, of several layers of parenchyma, becoming paler
to hyaline, and thinner below; the wall cells rounded-polygonal, about 10 x 5u; opening
by an apical pore about 154 diam. The loculus in the lower part is filled with a rather
loose fibrous tissue, which on its plane or somewhat folded upper surface bears a palisade

134 AUSTRALIAN FUNGI, II,

of erect, short, simple, continuous, hyaline sporophores, about 15 x 3-4u, projecting into
the loculus and abstricting single spores from their free ends. Conidia hyaline, cylindric
or sometimes slightly clavulate, straight or slightly bent, the ends rounded, smooth,
2-3-septate when mature, not constricted, 18-24 x 3-5-4-5y.

(155) STAGONOSPORA ATRIPLICIS Lind., Danish Fungi, p. 444.

On Chenopodium murale, Meningie, June 1953, L. D. Williams 17.

Leafspots scattered, yellow-brown, becoming almost white, circular to rather
irregular, smooth, with definite margin and indefinite zones outside, up to 3 mm. diam.,
or sometimes confluent and larger; on lower surface similar, but the centre rather
smaller and more indistinct. Pyenidia, loosely scattered or in crowded groups,
epiphyllous, immersed, black, 70-150u diam., globose, smooth, glabrous, the ostiole not
projecting beyond the leaf surface; wall surrounded by loose mycelial hyphae, of 1-2
layers of pale brown parenchyma 5-8 diam., darker around the circular apical pore
which is about 15u diam. Pyecnospores issuing in white masses, not tendrils, cylindric,
obtuse at both ends, from almost straight to slightly flexuous, hyaline, smooth, thin-
walled, at first pluri-guttulate, then with 1-3 septa, 15-30 x 3-5u, not constricted; the
largest spore seen was 48 x 5u with 5 septa.

The Ascochyta form of this was collected in the same locality on Atriplex muelleri,
L. D. Williams 22.

Pyecnospores 15-22 x 4-5u, pale yellowish in mass, cylindric with rounded ends,
l-septate and rarely slightly constricted, hyaline.

(156) STAGONOSPORA GREVILLEAE Hansf., n. sp. :

Maculae usque ad 15 x 1 mm., griseae, leves, atro-marginatae. Pycnidia dispersa,
nigra, depresse-globosa, usque ad 400u diam., subepidermalia, glabra; paries usque ad 15u
er., pluristratosus, atrobrunneus, in maturitate irregulater et lateque aperta.
Pycnosporae sessiles, cylindraceo-fusoideae utrinque rotundatae, plerumque rectae,
hyalinae, leves, (4)-7-(8)-septatae, haud constrictae, 40-50 x 8—-9uw.

Hab. in foliis Grevilleae juncifoliae, Ooldea, Aug. 1922, G. Samuel, WARI 2078.

Leafspots up to 15 mm. long and 1 mm. wide, usually along the margins of the linear
leaves, grey, smooth, with narrow dark brown marginal line. Pyecnidia epiphyllous,
scattered, black, flattened-globose, up to 4004 diam., formed in or below the epidermis
and covered by the thick cuticle and remains of the epidermal cells, the latter filled with
the black pseudoparenchyma of the pycnidial wall, which is continued downwards around
the sides and base of the loculus as a multi-layered wall up to 15 thick, the outside
smooth and glabrous. There appears to be no definite apical pore and the developing
pycnidium raises the cuticle and epidermal remains, finally throwing these off, often by
a peripheral fissure and then opening irregularly and widely. The spores appear to be
formed directly on the inner face of the wall of the base of the pycnidium, with no
elongate sterigmata or sporophores; spores cylindric-fusoid with rounded ends, usually
straight, hyaline, smooth, transversely (4)-7-(8)-septate, 40-50 x 8—9u, the epispore
two-layered; the outer layer thin and smooth, the inner about lu thick and more
refringent, similar to the cross-septa, and often constricted slightly at these.

The pycnidial construction verges towards Leptostromataceae.

(157) HENDERSONIA AUSTRALIENSIS Hansf., n. sp.

Pycnidia dispersa vel subaggregata, haud connata, nigra, punctiformia, immersa,
80-120u diam., 50-60 alt., depresso-globosa, glabra; ostiolum epidermidem folii
perforans, obtuse conoideum, pertusum; paries aurantiaco-brunneus, 2-4 stratosus,
parenchymaticus, cellulis plus minusve compressis, intus hyalinus, conidiiferus.
Conidiophora bacillaria, hyalina, recta, simplicia, continua, usque ad 15 x lu. Conidia
acrogenea, singula, late fusoidea, castanea, utrinque attenuata, apice rotundata, basi
leniter truncata, 15-18 x 6-5-9u, transverse 3-septata, haud constricta, episporio dense
atro-verruculoso, in medio 1-5—2u cr., utroque tenuiore.

BY C. G. HANSFORD. 5

Hab. in foliis emortuis Stipae spec. indet., Meningie, South Australia, L. D. Williams,
WARI 3489 p.p.

Pyenidia closely scattered or few-aggregate but not confluent, not on leafspots,
black, punctiform, immersed with the obtusely conoid ostiole piercing the epidermis but
searcely protruding, 80-1204 diam., 50-604 high, flattened-globose, glabrous; wall golden-
brown, of several layers of flattened, rather thin-walled, angular parenchyma, lined by
one or two layers of hyaline thin-walled cells, on the inner surface bearing erect, straight,
simple, continuous, hyaline conidiophores, up to 15 x lw. Conidia apical, single, wide
fusoid, attenuate towards both ends, the apex rounded, the base subtruncate, chestnut-
brown when mature, transversely 3-septate, not constricted, 15-18 x 6-5—9u, the epispore
closely dark-verruculose, 1-5—2u thick in the middle of the spore, thinner towards each
end.

(158) HENDERSONIA EUCALYPTORUM Hansf., n. sp.

Maculae amphigenae, usque ad 1-5 mm. diam., dilute griseae, linea purpurea conctae,
rotundatae, numerosae, interdum confluentes. Pycnidia amphigenea, subepidermalia,
punctiformia, nigra, globosa, leves, usque ad 120u diam., laxe dispersa; paries atro-
brunneus, membranaceus, strato externo ex cellulis polygonalibus 8-10 diam. composito,
strato interno ex cellulis hyalinis conidiiferis composito; apice poro rotundato 10u diam.
pertusus. Pycnosporae olivaceae, cylindraceae, rectae vel curvulae, basi subtruncatae,
apice attenuato-rotundatae, transverse 3-septatae haud constrictae, 40-48 x« 5-6u;
episporio circa lu cr., dense subtiliterque atro-granuloso.

Hab. in foliis juvenilibus Hucalypti leucoryli, Clare, South Australia, Aug. 1922,
T. Osborn, WARI 2007.

Leafspots amphigenous, showing on both sides of the leaf, up to 1-5 mm. diam., pale
grey, surrounded by a purple marginal line, numerous and sometimes confluent, rounded.
Mycelium of subhyaline, septate, ramose hyphae, penetrating the whole thickness of the
leaf. Pyenidia 2-5 on one side of each leafspot, amphigenous, subepidermal, minute,
black, globose, smooth, up to 120u diam.; wall dark brown, membranous, of an outer
layer of polygonal cells 8-10u diam., enclosing an inner layer of hyaline cells which
form the spores direct on their inner surface; apical pore not projecting, about 10.
diam. Pycnospores olivaceous, cylindric, straight or slightly bent, the base subtruncate,
the apex attenuate-rounded, transversely 3-septate, not constricted, 40-48 x 5-6u;
epispore about lp thick, closely and finely dark-granulose on the surface.

(159) HENDERSONIA FRASERI Hansf., n. sp.

Maculae amphigenae, rotundatae, usque ad 2 mm. diam. vel in confluendo irregulares,
centro arescentes, albidae, margine atro-rufae. Pycnidia amphigenea, immersa, dispersa,
nigra, globosa, usque ad 120u diam., glabra; paries tenuis, deorsum subhyalinus, sursum
2—3-stratosus, atrobrunneus, apice poro rotundato 5-10u diam. pertusus. Pycnosporae in
sterigmatibus minutis singulariter acrogenae, ellipsoideae vel lenissime clavulatae, apice
rotundatae, basi subtruncatae, atrobrunneae, leves, 3-septatae, haud constrictae, 25-28 x
6-94, rectae vel curvulae, episporio circa 1-5yu er.

Hab. in foliis Hucalypti spec. indet., Molong, New South Wales, L. Fraser, Aug.
1950; in foliis H. polyanthemis, Kosciusko, N.S.W., A. Costin 201, July 1949.

Leafspots amphigenous, rounded, to 2 mm. diam. or confluent and irregular, the
centre drying out to whitish and surrounded by a dark red border. Pyenidia amphigenous,
totally immersed and scarcely visible, the ostiolar pore barely reaching the leaf surface,
few in each leafspot, scattered, black, globose, to 120u diam., glabrous; wall thin and
subhyaline in the lower part, where it is somewhat indistinct, in the upper half composed
of 2-3 layers of black parenchyma, pierced at the apex by a pore 5-10u diam. Pyecnospores
formed on minute apiculate projections from the cells of the lining layer of the wall,
single and terminal, ellipsoid to somewhat clavulate, becoming dark brown, the apex
rounded, the base with a slightly truncate hilum, becoming transversely 3-septate, not
constricted, smooth, 23-28 x 6—-9u, straight or slightly bent, the wall about 1-5 thick.

136 AUSTRALIAN FUNGI, II,

The internal mycelium consists of subhyaline intercellular septate hyphae, penetrating
the whole mesophyll and killing the cells, which contain a reddish-brown amorphous
substance.

(160) HrnpERSONIA LONGISTRIATA Hansf., n. sp.

Pyenidia dispersa vel 2—3-aggregata et subconnata, depresso-globosa vel ljenticularia,
nigra, punctiformia, usque ad 1504 diam., apice primo poro minuto pertusa, demum late
aperta. Paries lente membranaceus, atrobrunneus, 2-stratosus, parenchymaticus, cellulis
5-9 x 5u, polygonalibus. Conidiophora non visa. Pyecnosporae cylindraceae, apice
rotundatae, basi truncatae, castaneae vel aureo-brunneae, 3-septatae, subinde lenissime
constrictae, 13-19 x 4-5u; episporio circa lu cr., extus longitudinaliter striato-sulcato.
Pyenidia primo immersa, demum epidermide fisso subsuperficialia, glabra.

Hab. in culmis emortuos Graminearum spec. indet., Meningie, South Australia, L. D.
Williams, WARI 3540.

Pyenidia at first completely immersed, but breaking through the raised epidermis
and becoming almost superficial when mature, scattered or in small close groups and
then almost connate, usually circular in outline, up to 1504 diam., in section flattened-
globose to lenticular, black, punctiform, with minute apical pore, at length splitting
more widely open. Wall tough membranous, dark brown, composed of two layers of
thin-walled polygonal cells, 5-9 x 5u, enclosing a hyaline layer of thin-walled, compressed
cells. Conidiophores not evident. Conidia cylindric with rounded apex and truncate
base, becoming chestnut-brown to golden-brown, 3-septate, sometimes slightly constricted
at all septa, 13-19 x 4-5u; epispore about ly» thick, with about 8 very shallow longitudinal
ridges the whole length of the spore.

(161) HENDERSONIA PRUNI Died., Arypt.-Fl. Brandenb., 9:658, 1914.

On old prunings of Prunus armeniaca, Nuriootpa, South Australia, Hansford, WARI
3512.

Pyenidia scattered, covered by the swollen epidermis and becoming slightly
erumpent at the apex, with apical round pore, globose to somewhat flattened, up to 450u
diam., black, glabrous. Wall of dark brown angular parenchyma in 2-3 layers, tough
membranous when moist, lined with 2-3 layers of hyaline cells, the innermost forming
conidia on very short sterigmata, the conidia appearing almost sessile. Conidia oblong-
cylindric to somewhat ovate, the ends rounded, becoming yellow-brown and then darker,
at first 1-septate, then 2—3-septate, not usually constricted, quite smooth, 10-14 x 4—5u.

(162) HENDERSONIA TRIODIAE Hansf., n. sp.

Pyenidia immersa, seriata, globosa vel subirregularia, discreta, usque ad 150 diam.
et 100u alt.; paries levis, pluristratosus, parenchymaticus, usque ad 15 er., brunneus,
ostiolo erumpente leniter papillato, perforato; sub epidermide folii orta, cellulis
epidermidis illis nigris opacis fungi impletis. Sporophora hyalina, cylindracea, simplicia,
continua, circa 7 x 2-34. Sporae singulae, acrogeneae, hyalinae demum dilute olivaceae,
fusoideae, rectae vel curculae, apice attenuato-rotundatae, basi rotundatae vel sub-
truncatae, leves, 3—5-septatae, haud constrictae, 20-35 x 5—Tyu.

Hab. in foliis Triodiae irritantis, Meningie, South Australia, L. D. Williams, June
I1G)5y33,

Pycnidia immersed, forming lines along the leaf, globose to somewhat irregular,
separate, up to 150u diam. and 1004 high, developed below the epidermis, which is filled
with a black, opaque, clypeal tissue. Wall smooth, of several layers of compressed brown
parenchyma, up to 15m thick, with slightly papillate, erumpent ostiole, with apical pore.
Locule lined with short simple conidiophores, hyaline, cylindric, continuous, up to
7 x 2-3u, each bearing a single apical spore. Spores hyaline, becoming pale olivaceous,
fusoid, straight or slightly bent, rounded-attenuate at the apex, rounded or subtruncate
at the base, smooth, thin-walled, transversely 3—5-septate, not constricted, 20-35 x 5—7wu.

BY C. G. HANSFORD. 137

(163) CAMAROSPORIUM MyopPorR! Hansf., n. sp.

Maculae amphigenae, usque ad 12 mm. diam., aridae, cinereae, atro-marginatae, leves.
Pyenidia dense circinatim dispersa, immersa, globosa, glabra, usque ad 200u diam., nigra;
paries pycnidii crasse membranaceus, atrobrunneus, subopacus, 3—5-stratosus, ex cellulis
polygonalibus 104 diam. compositus, intus hyalinus et conidiiferus. Sporophora nulla.
Pyecnosporae atrobrunneae, subglobosae, ellipsoideae vel clavulatae, transverse
1—4-septatae, longitudinaliter aut oblique 1-septatae, haud constrictae, 10-18 x 5-10u,
apice late rotundatae, basi lenissime apiculatae vel truncatae, leves.

Hab. in foliis Myopori platycarpi, Henley Beach, South Australia, July 1922, G.
Samuel, WARI 2008.

Leafspots showing on both sides of the leaf, up to 12 mm. diam., dry cinereous with
raised black-line margin, smooth, with closely scattered or indefinitely circinate pycnidia.
Pycnidia immersed, black, punctiform, globose, glabrous, up to 200u diam., with apical
round pore about 25u diam., slightly projecting owing to shrinkage of host tissue; wall
rather thickly membranous, dark brown, subopaque, of 3-5 layers of polygonal dark
brown cells about 104 diam., lined with a hyaline conidiiferous layer. Sporophores none.
Pyenospores dark brown, smooth, from subglobose to ellipsoid, or clavate, with 1-4
transverse septa and one longitudinal or oblique septum in one or more cells, not
constricted, 10-18 x 5-10u, the apex broadly rounded, the base very slightly apiculate or
subtruncate.

(164) CAMAROSPORIUM SALICORNIAE Hansf., n. sp.

Pycnidia subepidermalia, dense dispersa, atra, globosa, 100-140u diam. et alt., levia,
membranacea, apice poro rotundato 10-154 diam. pertusa; paries pycnidii 5-10u er.,
translucente melleus, 2-stratosus, ex cellulis angulosis 5-124 diam., fortiter compressibus
compositus, apice atrobrunneus. Pycnosporae subglobosae vel late ovatae, olivaceae,
17-25 x 12-15u, leves, transverse 3-(4)-septatae et longitudinaliter 1—-2-septatae, subinde
leniter constrictae.

Hab. in ramulis subvivis Salicorniae australis, Eucolo Gorge, Pimba, South
Australia, Dec. 1953, J. B. Cleland, WARI 3700.

The pyenidia are closely scattered on the older parts of the stems, not causing
apparent damage, black, globose, 100-1404 diam. and high, with the apical pore
penetrating the dead epidermis, but without clypeus or stroma, smooth, glabrous,
membranous. Pycnidial wall 5-10u thick, pale translucent honey-brown, darker around
the apex, consisting of 2 layers of angular parenchyma 5-124 diam., much flattened,
enclosing an indistinct layer of hyaline cells on which the spores are apparently borne
direct. Spores at first subglobose, becoming widely ovate, olivaceous-brown, 17-25 x
12-154, smooth, transversely 3, or more rarely 4-septate, with most cells 1—-2-longitudinally
septate, sometimes the older spores having somewhat swollen cells and then slightly
constricted at the septa.

The same fungus also occurred in small quantity on Arthrocnemum halocnemoides,
growing close by.

(165) CAMAROSPORIUM SAMUELI Hansf., n. sp.

Pycnidia amphigenea et caulicola, dispersa, nigra, punctiformia, globosa, usque ad
140u diam., glabra, haud prominentia; paries pycnidii membranaceus, 10-124 ecr.,
2-3-stratosus, extus parenchymaticus, ex cellulis polygonalibus compressibus, fulvis vel
dilute olivaceis compositus, apice poro rotundato 10-18 diam. pertusus. Pycnosporae
atrobrunneae, ellipsoideae, utrinque late rotundatae, leves, transverse 5—7-septatae et
longitudinaliter 2-septatae, in medio leniter constrictae, 20-25 x 12-14uy.

Hab. in foliis subvivis et in caulibus Salsolae kali, Henley Beach, South Australia,
July 1922, G. Samuel, WARI 2005.

Not causing leafspots, the pycnidia scattered over the old leaves and also on
indefinite greyish areas of the stems; on the leaves amphigenous, punctiform, black,
globose, up to 140u diam., glabrous, not projecting from the host tissue; pyenidial wall
membranous, pale olivaceous to yellowish (the black colour of the mature pycnidia

IK

138 AUSTRALIAN FUNGI, II,

being due to the dark spores filling the loculus), 10-124 thick, of 1-2 outer layers of
flattened polygonal cells 8-104 diam., enclosing an inner hyaline layer of conidiiferous
cells; apical pore round, 10-184 diam., piercing the epidermis. Spores formed directly
on the lining cells of the pyenidium, ellipsoid, at first pale olivaceous, becoming dark
brown, smooth, transversely 5—7-septate when mature and usually with 2 longitudinal
septa in the larger cells, often slightly constricted at the middle septum and then looking
exactly like ascospores of Pleospora, 20-25 x 12-14.

(166) CAMAROSPORIUM SP. INDET.

On old stems of Suaeda australis, Meningie, South Australia, L. D. Williams, WARI
3490, p.p.

Pyenidia scattered or loosely gregarious, immersed, globose to depressed, about 150
diam., black, punctiform, with papillate, erumpent ostiole pierced by a round pore. Wall
thinly membranous, of dark brown angular cells, paler towards the base, in 1-2 layers.
Conidiophores not evident. Spores mostly ovate to subpiriform, honey-brown, smooth,
transversely 3-septate, with 1 longitudinal septum in the two central cells, 12-15 x 6—8n.

(167) SrprorRitA CARPHOLOBI Hansf., n. sp.

Maculae cinereae vel griseae, indeterminatae, orbiculares. Pyecnidia epiphylla,
punctiformia, dense irregulariterque dispersa, immersa, nigra, circa 80» diam.;
membranacea, glabra; paries parenchymaticus, 1—2-stratosus, cellulis 6-10 x 4-Ty,
sursum atrobrunneus, deorsum pallidior vel subhyalinus. Sporophora non visa. Sporae
hyalinae, filiformes vel cylindraceae, rectae vel curvulae, utrinque obtuse rotundatae vel
basi subtruncatae, leves, plerumque 2-septatae, 30-50 x 3u.

Hab. in foliis subvivis Carpholobi aequilateri, Meningie, South Australia, L. D.
Williams, WARI 3527, p.p.

Leafspots ashen to grey, indefinite, rounded, with numerous pycnidia on upper
surface, which soon extend into the dying tissues around. Pycnidia punctiform, closely
and irregularly scattered, immersed, appearing black, about 80u diam.; under microscope
greenish-brown, membranous, darker around the apical pore, smooth; wall of
parenchymatous cells irregularly arranged and angulose, 6-10 x 4—7y, in 1-2 layers, paler
to subhyaline towards the base of the pycnidium. Sporophores not seen. Spores hyaline,
filiform to cylindric, straight or slightly bent, the ends obtusely rounded or the base
subtruncate, smooth, mostly 2-septate, 30-50 x 3u, very slightly attenuate towards the
ends.

(168) SEPTORIA GERANIT Rob. & Desm., Ann. Sci. Nat., 20:93, 1853.

On leaves of Geranium pilosum, Kosciusko, New South Wales, A. Costin 174.

Leafspots indefinite, with reddish surrounding area, drying out to pale brownish in
the centre, irregular and often confluent. Pycnidia amphigenous, mostly hypophyllous,
black, at first completely immersed, then by enlargement elevating the epidermis and
becoming almost superficial, though still covered by the epidermis, which is joined to
the pyenidial wall around the upper part. Pycnidia black, soft membranous-fleshy, up ta
200u diam. and high, with terminal pore about 30-404 diam.; wall parenchymatous,
about 10 thick in lower part, thicker around apex and passing at the sides into
subhyaline to pale olivaceous mycelial hyphae 3u thick, branched and interwoven,
septate, intercellular in mesophyll; wall of 2-4 layers of olivaceous parenchyma, soft,
lined with a close palisade of simple fusiform conidiophores 10-18 x 3u, slightly attenuate
towards both ends, continuous, straight or bent, hyaline, forming single conidia at the
apex. Conidia hyaline, filiform with rounded ends, 30-50 x 1-1-5, smooth, bent to much
curved, indistinctly 3-septate.

(169) SEPTORIA GERBERAE Syd., Ann. Mycol., Berlin, 10:48, 1912.
On Gerbera jamesonii, Meningie, S. Australia, June 1953, L. D. Williams 48.

(170) SEPTORIA PELARGONIT Syd., l.c. 10:443, 1912.
On Pelargonium australe, Meningie, S. Australia, June 1953, L. D. Williams 16.

BY C. G. HANSFORD. 139

(171) Srprorta SttyBi Pass., Atti Soc. Critt. [tal., 2:34.

On Silybum marianum, Meningie, S. Australia, June 1953, L. D. Williams 26.

Leafspots more or less circular, often demarcated by a dark line and with traces of
zonation, brownish-grey, becoming pallid, smooth, shining, often raised above the general
level of the leaf, up to 7 mm. diam., though usually smaller, numerous, rarely confluent..
Pycnidia in mesophyll with the ostioles opening on upper surface, loosely scattered,
black, globose, smooth, 50-100u diam.; wall of meandering brown hyphae 4—5u wide,
much branched and agglutinate into a single layer, thin and leathery-membranous, not
parenchymatous; ostiole very slightly papillate with a narrow round pore. Sporophores
lining the loculus, very short, hyaline, simple, each producing a single apical spore.
Spores extruded in a long white tendril, hyaline, filiform, up to 65 x 1-5u, with numerous
minute vacuoles but no septa, straight or slightly bent, smooth, thin-walled, the ends
attenuate to a narrow apex and slightly wider base.

(172) SEPTORIA SP. INDET.

On Wahlenbergia gracilenta, Keith, South Australia, L. D. Williams, WARI 3695.

Leafspots often terminal, indefinite, whitish when dry, up to 5 mm. long, becoming-
greyish with formation of numerous pycnidia. Pycnidia closely scattered, immersed,
globose, black, about 100u diam., smooth, with apical round ostiole about 10u diam.; wall
of one or two layers of dark angular parenchyma with some traces on the exterior of
formation from interwoven hyphae, the whole 5—8u thick, bearing on the inner surface a
rather close palisade of simple, continuous sporophores about 10u long, each forming a
single spore at its apex. Spores filiform, hyaline, greenish in mass, straight or slightly
bent, the ends obtusely rounded, continuous or indistinctly 3-septate, smooth, 30-52 x
1-5-2yu.

This does not agree well with descriptions of species previously recorded on
Campanulaceae, but no material of these was available for comparison.

(173) DINEMASPORIUM GRAMINUM Ley., var. STRIGULOSUM Karst., Hedwigia 1884, p. 21.
On dead leaves Phragmites sp., Meningie, S. Australia, L. D. Williams, WARI 3494,
det. E. W. Mason.

(174) PresTaALoTIA QUERCINA Guba, Mycologia, 24:380, 1932.

On old leaves of Quercus sp., Adelaide, March 1952, isolated in culture by A. Kerr.

Petri-dish cultures produce a white mycelial growth with scattered acervuli, which
are frequently coalescent into black spore masses up to 4 mm. diam.; from single
acervuli the spores sometimes issue in long twisted black tendrils. Spores 5-celled,
fusoid-clavulate, tapering to the base, very slightly constricted at the septa, mostly
24-28 long; the three median cells olivaceous, equally coloured, 15-184 long; terminal
cells hyaline, the apical cell conoid with a crest of 3 divergent setae 10-15 x lu; the
basal cell conoid and with a terminal seta 5-7u long.

(175) PESTALOTIA BANKSIAE Hansf., n. sp.

Gallae in fructibus productae, usque ad 7 cm. diam., durae, brunneae, tuberculatae,
intus lignosae et hyphis hyalinis saepe aggregatis permeatae. Fungus culto mycelium
album et acervuli numerosi ferens. Acervuli usque ad 1 mm. diam., nigri. Conidia
fusoidea, 4-septata, cellulis mediis atro-olivaceis, leniter constrictis, levibus vel subtiliter
granulosis, 14-20 x 6—7u, cellulis terminalibus hyalinis, obtuse conoideis, 4-5u longis;
setae apicales 3, divergentes, hyalinae, solidae, usque ad 20 x 1u: seta basale usque ad
10 x lu; corpora tota conidiorum 22-29u longa.

Hab. in fructibus Banksiae marginatae, Keith, South Australia, Dec. UGH, Ie IR
Harris, WARI 3684.

Infected fruit-spikes show nearly every individual fruit greatly enlarged, and reach
7 cm. diam., to form a rough-tuberculate, woody, brown gall; internally the tissues are
brown and woody, permeated by loose hyphae and masses or small sheets of white

140 AUSTRALIAN FUNGI, II,

mycelium. In agar culture the fungus forms a white woolly aerial mycelium, with
numerous black acervuli, mostly around the colony margin, which reach 1 mm. diam.
Conidia fusoid, usually straight, with a basal setiform pedicel up to 10u long, and three
divergent apical setae up to 20 x lu; the body of the conidium is 22-294 long by 6—7u
wide in the widest part, consisting of terminal small conoid hyaline cells each 4—6u long,
with three, uniformly dark olivaceous, central cells, measuring together 14-20 x 6-T7yn,
slightly constricted at the septa, the exterior smooth or indistinctly granulose. This
was the only fungus isolated from the galled fruits, but has not yet been proved to be
the cause of the disease.

(176) MoNOCHAETIA MUEHLENBECKIAE McAlpine in Herb., n. sp.

Maculae amphigenae, in epiphyllo rotundatae, concentrice zonatae, usque ad 10 mm.
diam. vel confluentes, linea atrobrunnea circumdatae, intus alternatim atro- et fulvo-
brunneo zonatae, haud secedentes, in hypophyllo minus distinctae. Acervuli amphigeni,
plerumque epiphylli, dispersi, atro-brunnei vel subnigri, usque ad 1004 diam., primo
velati, deémum epidermidem irregulare disrumpenti. Sporae fusoideae, 21-26 x 2-5-5-5yu,
transverse 4-septatae, utrinque hyalinae, cellulis mediis dilute olivaceis; cellula apicali
conoidea, in setam terminaliam rectam vel curvulam, usque ad 10 x lu producta; cellula
basali obtuse conoidea, seta recta 4-7u longa excentrica praedita.

Hab. in foliis Muehlenbeckiae adpressae, Victoria (typus in Herb. Waite Institute,
2088).

The leafspots show on both sides of the leaf, on the upper surface rounded, con-
centrically zoned alternately with dark- and buff-brown, surrounded by a dark brown
marginal line, up to 10 mm. diam., drying out but not secedent; on the lower surface
the spots are less distinct. Acervuli amphigenous, mostly epiphyllous, evenly scattered
over the centre of the leafspot, dark brown to almost black, up to 100u diam., covered,
then bursting the epidermis irregularly. Spores slightly bent, fusoid, 21-26 x 4-5—-5-5y,
4-septate, the end cells hyaline and conoid, the three middle cells pale olivaceous, smooth,
and not constricted at the septa; apical cell produced into a terminal straight or slightly
bent hyaline seta up to 10 x 1u; basal cell with an excentric straight seta 4—7u long.

(177) MoNocHAETIA LOMATIE McAlpine in Herb., n. sp.

Maculae 2-30 mm. diam., rotundatae vel confluentes, linea atrobrunnea circumdatae,
luteae vel griseo-brunneae, haud secedentes, haud zonatae, arescentes. Acervuli
amphigeni, nigri, plus minusve rotundati, usque ad 3004 diam., epidermidem rumpenti.
Sporae late fusoideae, saepe curvulae, 15-20 x 5-5-7-0u, 4-septatae, cellulis mediis
eastaneis, tunicatis (1-5), extus verruculosis, cellulis terminalibus hyalinis, conoideis,
minutis, 2-34 long. et lat.; seta basali 5-154 longa, recta vel curvula, centrali; seta
apicali usque ad 33 longa, recta vel curvula.

Hab. in foliis Lomatiae ilicifoliae, Victoria, WARI 3728, ex Herb. Dept. Agr. Victoria.

Leafspots 2-30 mm. diam., round or confluent, each surrounded by a dark red-brown
line, the centre uniformly yellow to greyish-brown, drying out but not secedent.
Acervuli amphigenous, black, more or less circular, up to 300u diam., at first covered,
then bursting the epidermis. Spores broadly fusoid, often slightly bent, 15-20 x 5-5-7,
4-septate, not constricted, the terminal cells hyaline, obtusely conoid, 2-34 long and
wide; the central cells uniformly dark chestnut-brown, thick-walled (1-5u), the surface
coarsely and bluntly verruculose. Basal seta 5-154 long, apparently central; apical seta
a prolongation of the apical cell, up to 33 long, both straight or bent.

(178) MoNnocHABTIA UNICORNIS (Cke. & Ell.) Sacc., Syll Fung., 18:485, 1906.
On Cupressus sp., Adelaide, Jan. 1952, N. T. Flentje.

(179) PAPULARIA ARUNDINIS (Corda) Fr., Summa Veg. Scand., p. 509, 1849.
On old stems of Bamboo, Victoria, WARI 3727, ex Herb. Dept. Agr. Victoria; on old
culms Phragmites sp., Meningie, S. Australia, WARI 3494, L. D. Williams.

BY C. G. HANSFORD. 141

(180) TETRAPLOA ARISTATA B. & Br., Ann. Nat. Hist., 2:5:459, 1850.

On dead culm of Bromus gussonii, Meningie, South Australia, WARI 3525 p.p.,
L. D. Williams.

The conidia are about 35 x 184, composed of four lines of four cells each, slightly
constricted at the septa, at the apex each line being produced into a beak up to 80 long,
paler brown towards its obtuse apex 3u wide, slightly wider at the base, 2—4-septate,
smooth; the body of the spore is closely verruculose in the lower half. The conidia are
single and acrogenous on very short erect branches of the mycelium, sometimes almost
sessile on the repent hyphae.

(181) MOoNOSPORELLA SETOSA (B. & C.) Hughes, Canad. Journ. Bot., 51:654, 1953.
On wood, Hermitage, S. Australia, Oct. 1922, G. Samuel, WARI 2096.

(182) STEMPHYLIUM LANUGINOSUM Harz., Bull. Soc. Imper. Moscou, 44:182, 1871.
On dead haulms Solanum tuberosum, Mt. Gambier, South Australia, April 1917,
G. Samuel, WARI 1828, p.p.

(183) CErRCOSPORA LORANTHI McAlpine, Proc. Linn. Soc. N.S.W., 28:96, 19038.

On Loranthus pendulus, Black Swamp, S. Australia, June 1914, T. G. Osborn, WARI
1995.

Leafspots amphigenous, mostly epiphyllous, rounded, up to 4 mm. diam., raised,
grey, turning olivaceous with almost a continuous layer of densely fasciculate, erumpent
conidiophores. Internal mycelium penetrating the whole mesophyll and causing some
hypertrophy, not extending much beyond the edges of the visible spots and then only as
single intercellular hyphae; in the tissues of the leafspots aggregated to form sheets or
masses of pseudoparenchyma between the browned host cells; hyphae hyaline, not
forming haustoria. The mycelium aggregates beneath the epidermis to form a stroma
of olivaceous pseudoparenchyma, which extends outwards through the epidermis, at first
in separated masses between the host cells, but soon becoming laterally confluent and
up to 60 thick, crushing the epidermal cells, which are thrown off. The external
surface of this stroma is covered with a close palisade of short olivaceous conidiophores,
erect, simple, continuous, up to 25 x 4-5u, producing conidia singly at the apex. The
very slightly prominent scar left by the conidium becomes lateral by continued growth
of the conidiophore, so that old conidiophores are slightly geniculate and roughened by
successive conidial scars in the upper part, about 24 apart. Conidia versiform, from
ellipsoid to obclavate or elongate, 0-3-septate, not constricted at the septa, pale
olivaceous, the base internally rounded but externally with slightly prominent flat hilum,
slightly attenuated towards the rounded apex, thin-walled, smooth, 15-60 x 3-5—-4-5u, the
apex 2-5-3-0u wide.

142

NOTES ON AUSTRALIAN BEETLES IN THE TRIBE BOLBOCERATINI FORMERLY
IN THE GENUS BOLBOCERAS.

By Henry F. Howpen, Department of Zoology and Entomology, University of Tennessee,
Knoxville, Tennessee.

(Communicated by Dr. P. B. Carne.)
[Read 28th July, 1954.]

Synopsis.

The name Bolboceras could not be applied to any Australian species of the subfamily
Bolboceratini. Consequently Boucomont’s subgeneric names Blackburnium, Bolborhachium, and
Bolbapium are elevated to generic rank and a key is given to the Australian genera in the
subfamily. Also new names are proposed for two synonyms: Blackburnium quadriarmigerum
new name for Bolboceras armigerum Macleay and Blackburniwm bifoveatum new name for
Bolboceras cornigerum Macleay.

In a recent publication Cartwright (1953, Proc. U. S. N. M., 103:95-120) has shown
that the name Bolboceras should be applied to a small genus of beetles represented by
ten North American and one European species. Cartwright (1953, p. 101) presents the
reasons as follows:

“The genus Bolboceras was erected by Kirby in 1818 for the eight species
Scarabaeus mobilicornis Linnaeus [should read Fabricius], S. mobilicornis var.
testaceus Fabricius, S. quadridens Fabricius, S. farctus Fabricius, S. lazarus
Fabricius, S. cyclops Olivier, S. cephus Fabricius, and Bolboceras australasiae
Kirby. Technically no genotype was designated although he wrote, ‘My details
of Bolboceras were taken from B. quadridens’. Nearly all of the species mentioned
have since been moved to other genera. Curtis, 1829 (British Ent., vol. 1, pt. 1,
p. 74), selected the species Scarabaeus mobilicornis Fabricius [= armiger Scopoli]
‘as the type of Bolboceras. Therefore, since our species of Odontaeus are
congeneric with mobilicornis, they now take the generic name Bolboceras and it
becomes necessary to find an available name for those species we have formerly
placed in Bolboceras.”

Several names formerly placed in synonymy and a number of subgeneric names are
available. Most of these names are applicable to European and American forms which
are quite distinct from the Australian species. Of these only three seem easily referable
to the Australian forms. Blackburnium, Bolborhachiuwm and Bolbapiuwm were proposed
as subgenera under Bolboceras by Boucomont (1910, Ann. Soc. ent. Fr., 79:339-340) and
are herein slightly modified and raised to generic rank.

With some redefinition of generic characteristics, most of the Australian species
can be included in the genus Blackburnium. Boucomont’s description, translated from
the French, is as follows (1910, p. 339): First joint of the antennal club with a denuded
area not clearly delimited; anterior edge of the prothorax bearing behind the eyes two
little round and deep foveae; prosternal cavities carinate at the external edge; anterior
lobe of the mesosternal plate tectiform or more exactly cariniform, raised in front like
the prow of a boat; posterior lobe terminated in an acute angle with an abrupt edge and
vertical slant; first stria of elytra, as in most of the Australian species, reaching the
base by going around the scutellum from which it is separated by a fine carina; base of
elytra carinate.

From the material the writer has examined it seems doubtful if several of the above
characters should be given generic weight. Deep foveae can be found behind the eyes in
some species, indications of them in other species, and in many cases they are com-

BY HENRY F. HOWDEN. 143

pletely lacking. It is also doubtful if the sexual characteristics or carinate prosternal
cavities are good generic characteristics. However, further study of more adequate
collections than the writer has available may warrant not only the use of the above
characters but the establishment of a number of new genera.

'The species in Blackburnium as here defined have the first elytral stria curving
around the scutellum, reaching the base of the elytra; elytra with seven striae between
suture and humeral umbone; base of elytra margined; mesosternal coxae separated
by the mesosternal plate which is tectiform or slightly to conspicuously cariniform,
raised anteriorly like the prow of a boat, although in some cases this “prow” is almost
truncate. Pronotum with or without horns, but lacking a transverse carina in front
of the posterior pronotal margin.

Type of Blackburnium as designated by Boucomont (1910, p. 339): B. reichei
Guérin. The genus seems to be entirely Australian, including most of the Australian
species formerly placed in Bolboceras.

The second of Boucomont’s subgenera, Bolborhachium, here accorded generic status,
has the prothorax with an abrupt, carinate edge parallel to the base, the prothorax
before the carina abruptly declivous, often excavated; first elytral stria shortened, barely
reaching scutellum; elytral base margined; seven striae between suture and humeral
umbone; mesosternal plate anteriorly flat between the middle coxae and truncate in
front.

Type of Bolborhachium as designated by Boucomont (1910, p. 339): B. recticorne
Guérin. Less than ten species have been placed in this strictly Australian genus.

The third of Boucomont’s subgenera accorded generic rank is Bolbapium. As defined
by Boucomont the genus includes both Australian and South American species. Since
none of the Australian species have been examined, a translation of Boucomont’s
description is given unchanged (1910, p. 340): Elytra with five striae between the
humeral umbone and the suture, the first reaching the base; ocular cavities of the
prosternum large, with lateral edges at a right angle with a carina at the intersection of
the two planes; mesosternal plate large, convex, pear-shaped.

Most species with elytral base unmargined. However, several Australian species,
according to Boucomont, have a small basal carina and their present inclusion in this
genus is open to question.

To facilitate the placement of the Australian species in the tribe Bolboceratini a
brief key to the genera is included below.

il, Seutelinm wea, waeONy, EubaaosE IWbboeeNe soocoucscuodoccoeonosuue abo bonobooEG Stenaspidius
Scueellimeerlanswlac ss mOt linear eels ccs cles ee aN Saete roel Chote i loic Oo glo a on eas 2
2. Eyes entirely divided; colour uniform brown or black. North American and European
SIDOCHOS: 5.b:bud.ssg-digr olla ce gear ROO LORD anes oS ean ne ROO Okc tO cia race me Spe ene Bolboceras (formerly Odontaeus)
IDWES Mot EmiEieShy clinnicleol lon EG} GAMUIMS sooconoboguosnooob ane ooUoe OD OUCDdooOU SOK GdoODOL 3
3. Middle coxae subcontiguous, mesosternum linear between coxae ..............00000e0ee 4
Middle coxae separated one from the other by the anterior lobe of the mesosternal plate
Vl Chie Sap Td VETS MING ATs seein ay se tise mcei at ap ai tecltetie ay cies Mecbente mised oeshisiictiewre coy cals s @bsmancteke oy aulereh anaes al Raed. 4 Uekeos 5

4. Elytra with five dorsal striae between suture and humeral umbone; males with clypeus not
AC atlavgme | OTS AC Cunvaweare weeaete le peureitoe ao, cease aden onteteuian cree meee atoll nkechie touch cosh oraub it peuatsneuaulhBoueuarsnahedelie Hucanthus

Elytra with seven dorsal striae between suture and humeral umbone; males with mouth

IMfehIOMmwitvhe chy peusseneatlyaelonsateds ameaanceie ace es emis wen selene. Elaphastomus
5. Elytra with five striae between suture and humeral umbone .................. Bolbapium
Hlytra with seven striae between suture and humeral umbone ........................ 6

6. Mesosternal plate tectiform or cariniform, often raised in front like the prow of a boat;
sutural striae of elytra curving around scutellum to the base .............. Blackburnium
Mesosternal plate flat, truncate in front; sutural striae of elytra short, barely reaching
S CULE MI TIMeMenene new romeo teiicis et aictsnslc aerate <o: s)starere oneretenorobecolevcre aus <: eu db atausicls tacuaisher itis ounia Bolborhachium

Further study and more adequate material will undoubtedly show the necessity
for the establishment of additional genera or more adequate definition of the present
ones.

The names of several of the Australian species have been found preoccupied and
therefore should be renamed.

144 AUSTRALIAN BOLBOCERATINI FORMERLY IN THE GENUS BOLBOCERAS.

BLACKBURNIUM QUADRIARMIGERUM, new name.
For Bolboceras armigerum Macleay, 1873, Trans. Entomological Soc. N. 8S. Wales,
2:360. Not Bolboceras armiger Scopoli, 1772, Ann. Hist., 5:78.

BLACKBURNIUM BIFOVEATUM, new name.
For Bolboceras cornigerum Macleay, 1873, Trans. Entomological Soc. N. S. Wales,
2:363. Not Bolboceras cornigerus Melsheimer, 1844, Proc. Acad. Nat. Sci. Phila., 2:138.

The writer is indebted to Dr. P. B. Carne, Division of Entomology, Commonwealth
Scientific and Industrial Research Organization, Canberra, for his advice and for
numerous specimens.

ANTIRRHINUM RUST, PUCCINIA ANTIRRHINI D. & H., IN AUSTRALIA.
By J. WALKER, Biological Branch, New South Wales Department of Agriculture.
(Plate vii; two Text-figures.)

[Read 25th August, 1954.]

Synopsis.

The rust disease of Antirrhinwum spp. caused by the fungus Puccinia antirrhini D. & H. is
recorded from Australia. Ail plant parts except the petals and the roots were attacked and
severe damage was caused. A determination of the physiologic race of rust present showed
that it is similar to the virulent race known in America as race 2. Studies on uredospore
longevity indicated that these spores remained alive longest at low temperatures and relative
humidities and their longevity fell off as both temperature and humidity rose.

The origin and the spread of the disease in Australia are discussed. An American origin
for the disease is proposed and evidence is presented to show the importance of wind-borne
uredospores in the spread of this disease.

INTRODUCTION.

The rust disease of Antirrhinum spp. caused by the parasitic fungus Puccinia
antirrhini Dietel and Holway is the most destructive disease of these plants known.
Until October, 1952, it had not been recorded in Australia, but at that time a severe
outbreak occurred in New South Wales in the Sydney metropolitan area, and within
a short time it had spread throughout eastern New South Wales. Since then it has
been recorded from Victoria, Queensland, South Australia and Tasmania. In this
paper details of the work carried out on this disease since its first outbreak are given.

HISTORY OF THE DISEASE.

Antirrhinum rust was first recorded by Blasdale (1903), who, in 1895, had found it
doing considerable damage in Californian gardens. Specimens of the rust had been
collected in California before this date, however, and a specimen in Professor J. C.
Arthur’s collection from Santa Cruz is dated 1879 (Peltier, 1919). The fungus was
described as Puccinia antirrhini by Dietel and Holway (1899) from specimens submitted
by Blasdale.

In 1913 the disease was recorded from the eastern State of Illinois, and by 1919
(Peltier, 1919) had spread throughout the United States. In 1921 it was reported by
Dickson (1921) from Canada, and in 1922 Whetzel (1924) found it causing severe
damage to snapdragons in Bermuda.

Up to 1931 no new records of rust were made, but Lepik (1941) states that, in
that year, Viennot-Bourgin reported it in the north of France. Soon after, the disease
was reported in England. Green (1933), reporting the outbreak, stated that the rust
appeared at a number of localities simultaneously and that spread was rapid and
destructive. Spread to other EKuropean countries occurred rapidly, and four years
later it had been recorded from as far east ds Odessa (Lepik, 1941), Palestine (Rayss,
1937) and Egypt (Fikry, 1937). More recently it has been recorded in South Africa
(Bottomley, 1940), Norway (Jorstad, 1946) and Tanganyika (Wallace, 1952).

The first record of the disease in Australia was made during October, 1952, on
plants growing in a Sydney (N.S.W.) suburban nursery. It spread rapidly and within
eighteen months was common in eastern New South Wales and had been reported from
all other States of the Commonwealth with the exception of Western Australia. Details
of its spread are given in the “Hpidemiology” section below.

More recently, in December, 19538, rust was reported in New Zealand at Auckland.
So far, no further spread in that country has been noticed (Brien, 1954, personal
communication).

146 ANTIRRHINUM RUST IN AUSTRALIA,

SYMPTOMS.

Puccinia antirrhini is able to attack all above ground parts of the plant except the
petals. On the leaves, the first sign of infection is usually a small yellowish spot seen
on the under surface. This gradually enlarges to form a shiny, brown, slightly raised
blister, about 1 mm. in diameter, which finally breaks through the leaf epidermis as
a reddish-brown uredosorus. Quite often, later uredosori are produced in rings around
the initial one, giving a distinctive pattern (Plate vii, 1). Sometimes, corresponding to
the masses of uredosori on the underside, a chlorotic area can be seen on the upper
side of the leaf (Plate vii, 2). As the season progresses, the leaf pustules darken and
the leaves become covered with black teleutosori.

Stems and petioles are also attacked and uredosori have been observed on the
sepals of young buds (Plate vii, 3). On the stems the pustules consist largely of
teleutosori, which often completely encircle the stems. In later stages of infection
plants may be almost completely covered with rust pustules (Plate vii, 5).

Quite commonly on infected leaves necrotic areas are seen to develop, especially
where many uredosori are massed together on the underside of the leaf (Plate vii, 4).
This effect has been noted in America by Dimock and Baker (1951), who found that,
under semi-arid conditions, injury resulted almost entirely from drying out of the
rust-invaded tissues. They found, however, that where the humidity was high and
rainfall frequent, damage was caused by facultative parasites entering through the
rust pustules and advancing into previously healthy tissue. In New South Wales the
main cause of these necrotic areas seems to be drying out of the tissues during
hot weather.

An interesting observation was made by Fikry (1938) in Egypt, who found
teleutosori occurring on the roots of some heavily infected snapdragon plants. No
such occurrence has been observed during this work.

THE CAUSAL FUNGUS.

The uredospores of the Australian isolate of P. antirrhini (Text-fig. 1, A) are
roughly spherical to slightly elongated in shape, and measure 19-27u x 19-234 (mean of
20 measurements: 23u x 21u). The outer wall is finely echinulate, 14-34 thick, and
penetrated by 2-3 germ pores which are sometimes arranged in an equatorial plane
around the spore but often appear to occur irregularly. The spore wall is slightly
thicker at the point of attachment of the stalk.

The teleutospores are of two main types (Text-fig. 1, B). One type is short and
measures 30-424 x 21-274 (mean of 20 measurements: 36u x 23u). It has a blunt
apical cell, quite thick at the top and dark reddish-brown in colour. The other type
is longer, measuring 43-574 x 17-214 (mean of 20 measurements: 49u x 19”), and has
a more or less acute apical cell. This cell is lighter in colour than the apical cell of
the shorter type. Other workers have noticed these two teleutospore types in P. antirrhini
(Doidge, 1941).

Both types of teleutospore are constricted at the septum, tapering or somewhat
rounded towards the base, and with a long pedicel, up to 954 long. In both the top
cell is darker in colour than the basal cell. The germ pore in the top cell is apical and
in the basal cell obscure.

Locally produced uredospores germinate readily in 18 hours when dusted onto
water or a dilute aqueous extract of antirrhinum leaves and incubated at 10°C., but
so far it has not been found possible to germinate teleutospores. Some workers (Green,
1941; Mains, 1924) have succeeded in germinating them, however, but have been unable
to reinfect Antirrhinum or infect any other plant with them. It thus appears probable
that P. antirrhini is a heteroecious rust whose alternate host has not yet been found.
Mains (1924) discusses this in some detail. The alternate host is not necessary,

however, as the rust is able to persist by means of continual infection of Antirrhinum
with its uredospore stage.

BY J. WALKER. 147

Physiologic Specialization.

As with other rust fungi, various races of P. antirrhini have been found. Up till
1937 only one race of this rust was known, but in that year Yarwood (1937), working
with excised leaves of various snapdragon varieties, demonstrated in America the
existence of another race. This new race was known as race 2, the original being
race 1. Race 2 proved to be more virulent than race 1 and was able to attack all
those varieties that were hitherto resistant to the disease and, at the present time,
there are no commercial lines of snapdragon resistant to race 2. Apart from these two
races, it is quite possible that others also exist. Baker (1953, personal communication )
stated that work done in California indicated clearly that other races are present there.

— /t
= /
=f i
/
Ne ePollongons

B oe Banya | Sy c

Figure 1. Figure 2.

Text-fig. 1—(A) uredospores and (B) teleutospores of P. antirrhini.
Text-fig. 2—Map showing initial zones of spread of Antirrhinum rust in New South Wales.

As far as can be determined, race 2 of P. antirrhini has been reported to date only
in America and Southern Rhodesia. In other parts of the world where this rust
has been recorded, race 1 appears to be the race present and, in these areas, the
resistant varieties are still quite successful.

Tests were conducted to identify the race of rust present in New South Wales and
to test the reaction of a number of snapdragon varieties to inoculation with this race.

Method:

All plants to be tested were raised from seed in four-inch pots in a glasshouse,
hardened in a frame outside and then inoculated in the frame by dusting with heavily
rusted plants. Plants were kept damp, giving optimum conditions for rust infection.

Results:

Inoculation results are summarized in Table 1. All varieties tested were quite
susceptible to the local rust isolate and, although some variation in reaction did occur,
all were quite heavily rusted. In the table those varieties classed as ‘fully susceptible”
Showed profuse uredosorus development, with practically no chlorosis visible on the
upper side of the leaf. With those classed as “susceptible” some chlorosis was visible

148 ANTIRRHINUM RUST IN AUSTRALIA,

on the upper side of the leat and in some cases the uredosori were slightly smaller
than those on the “fully susceptible” types.

Of the varieties listed in Table 1, Watkins and Simpson’s Pink Freedom, Wisley
Bridesmaid, Wisley Cheerful and Wisley Golden Fleece were sent out by Mr. D. Green
of Wisley, England, and No. 61 by Dr. K. F. Baker of California. These five varieties
were strain-1 resistant types and, as is seen from the table, they were quite susceptible
to the local rust race. This indicates that the race of P. antirrhini present in Australia
is not race 1, but is similar to the virulent race 2.

Two varieties of Linaria, Excelsior Hybrid and Fairy Bouquet, were also inoculated
with rust. Both showed no sign of infection. In the United States, Blasdale (1903)
recorded P. antirrhini on two species of Linaria.

TABLE 1.
Results of Antirrhinum Rust Inoculations. List of Varieties Tested and Their Reaction.

Fully Susceptible. Susceptible.

Copper King.

Giant Ruffled Tetraploid.

Giant Skyscraper.

Grandiflorum Rust Proof de Luxe.

Campfire.

Defiance.

Grandiflorum Rust-proof de Luxe Mixture.
Queen Victoria.

Rust Resistant Autumn Glow Shades.
Rust Resistant Orange Shades.

Rust Resistant Tall Mixed.

Rust Resistant Yellow.

Grandiflorum Rust Resistant University

of California de Luxe Mixture.
Grandiflorum University of California.
Hybrids Mixed.

Snowflake. Luteum.

Tango. No. 61.

Tetraploid Rust Resistant Selected Superfine Rose Marie.
Mixed. Ruby.

Rust Proof de Luxe Mixture.
Rust Proof Good Mixed.
Rust Resistant Spotlight.
The Rose.

Yellow King.

Yellow Wonder.

University of California Mixed.

University of California Rust Resistant
Mixed.

Watkins’ and Simpson’s Pink Freedom.

Wisley Bridesmaid.

Wisley Cheerful.

Wisley Golden Fleece.

World Favourite.

UREDOSPORE LONGEVITY.

In the literature on the longevity of uredospores of rust fungi, one of the most
important points made is the variation in the life of the uredospores under varying
conditions of temperature and humidity. With Puccinia antirrhini the life of uredo-
spores has been given from as short as six to eight weeks (Green, 1941) to as long as
almost a year (Baker, 1953, personal communication). Because of the importance of
the length of life of this spore stage, tests have been carried out to determine the
longevity of locally produced spores.

Leaves bearing uredosori were collected from plants growing outside and, after
drying at room temperature for 24 hours, were stored in closed jars at three different
relative humidities: 25%, 50% and 75%. The humidity was controlled by sulphuric
acid/water mixtures. Jars at each humidity were incubated at each of five different
temperatures: 5°, 10°, 20°, 25° and 30°C., giving fifteen different treatments in all.
Spore samples were taken at regular intervals and their germination tested by dusting
onto the surface of a dilute aqueous extract of antirrhinum leaves in a Syracuse dish
and incubating at 10°C. The antirrhinum leaf extract was used in preference to plain
water as preliminary tests showed that much better germination was obtained with it.

Results of the germination tests are shown in Table 2. It is readily seen that the
temperature and relative humidity under which the spores are kept considerably
influence their longevity. Best germination occurred after storage at the lower
temperatures and relative humidities, and as each rose a falling off in the life of the

149

BY J. WALKER.

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150 ANTIRRHINUM RUST IN AUSTRALIA,

uredospores was noted. The longest period for which uredospores remained viable
during these tests was 116 days at 5° and 10°C. at relative humidities of 25% and 50%.
The shortest period was 21 days at 30°C. at all humidities.

The longevity of uredospores is of great importance in the epidemiology of rust
diseases. These spores are specially suited to transport by air currents, but they
must be able to remain viable under the conditions existing in the atmosphere if this
is to be an effective method of spreading the disease. Many factors are concerned
but here only temperature and humidity have been considered. The results show that,
at low temperatures and relative humidities up to 50%, uredospores of P. antirrhini
produced under local conditions are able to remain viable for about four months. Their
transport in cool air currents, probably at high altitudes (Gaumann, 1950), would thus
constitute a definite hazard to plants quite long distances away.

It is also apparent that uredospores could remain viable for some time under cool
conditions in dust in seed samples. Such samples are dry and, unless conditions of
very low humidity adversely affected uredospore longevity, as has been found with
some rusts (Gaumann, 1950), they would seem to provide an excellent means for rust
dispersal. Air transport of seed samples would make this even more effective.

Further discussion of wind and seed transport of uredospores is given under
“Hpidemiology”’.

EPIDEMIOLOGY.
In this section will be considered first the possible means by which the fungus
was introduced to Australia, and second its spread within the country.

1. Entry of the Disease.

The way in which Antirrhinum rust was introduced to Australia is not definitely
known. In seasons when locally grown snapdragon seed is in short supply, bulk lots
of seed are imported from Europe and America, and small quantities of new varieties
are constantly being introduced from both countries. The crop on which the first
infection was recorded was grown from Huropean seed. Before the identity of the race
present was established, it was therefore thought that the Australian outbreak had a
European origin. As race 2, however, has been recorded only in America and Southern
Rhodesia, it now seems that the original entry must have been from America. A
European origin for the Australian outbreak would seem to be disproved unless race 2
has become established fairly recently in some countries there. No record of snapdragon
seed being imported from Africa has been seen.

Examination of many seed samples showed that the majority of overseas seed
coming into New South Wales contained rust uredospores and sometimes teleutospores
as well. Of 185 seed samples examined, 139 (75%) were found to contain rust spores.
The results of these examinations, with the country of origin of the seed, are given in
Table 3. A rough estimate of the number of uredospores present in the seed samples,
made using a Bausch and Lomb mould counter, showed that, in most samples, there
were from 1-2 x 10° uredospores present per ounce of seed. However, it was not found
possible to germinate these spores or to obtain infection on susceptible seedlings with
them. Moreover, plants grown from this seed, and from seed dusted with fresh local
uredospores, always produced disease-free plants.

In favour of seed transmission are the facts that under dry and cool conditions
rust uredospores can remain viable for a considerable time and that air transport of
seed could allow seed to be introduced within a couple of months of harvesting. Under
these conditions there seems to be little reason against the production of the disease,
for example in a nursery, by the accidental dusting of spores on to snapdragon seedlings
when seed was being sown in adjacent beds.

The entry of the rust into other countries has sometimes been attributed to seed
transmission, e.g., Sweden (Palm, 1937), South Africa (Bottomley, 1940) and Italy
(Preti, 1935), but other workers, after various investigations, have rejected seed trans-
mission (Green, 1941; Hassebrauk, 1937). These latter workers, however, did not seem

BY J. WALKER. Li

to consider the possibility of viable spores, carried in seed, being dusted onto established
plants in a nursery area. Baker (1953, personal communication) considered this a
distinct possibility and definitely favoured seed transmission.

In the case of the Australian outbreak, evidence against seed introduction are the
facts that it was not found possible to germinate the uredospores found in seed samples
or obtain infection with them and that, if seed transmission were effective, one would
have expected the disease to have heen introduced some time ago, considering the large
number of uredospores in introduced seed.

Other methods by which this disease could be spread are on cuttings and seedlings,
’ by wind and by spores adhering to clothing. In view of the strict plant quarantine
regulations in force in New South Wales it is most unlikely that seedlings or cuttings
could have been imported. Wind entry of the disease is unlikely as, at this time, the
nearest known source of inoculum was in the Hawaiian Islands, more than 4,000 miles
from Sydney. The entry of the disease by spores adhering to clothing is doubtful.

It can thus be seen that the most likely method of entry of this disease was by
means of viable uredospores adhering to seed.

TABLE 3.

Results of Antirrhinum Seed Examinations.

Number of Samples. |
Total Number | Percentage

Country of Origin. | Ron ie | pion Examined. Containing
+P = | | Spores.

Holland = Re Ne 125 1) 4 144 | 87
New Zealand 4 24 | 28 14
United Kingdom a0 bata 4 2 6 67
Italy tes re ao ac | 2 —- 2 100
France .. xe ae 20 || 2 — 2 | 100
Germany 1 — 1 100
America il — 1 | 100
Tasmania = il | 1 | 0

| 139 | 46 | 185 75

+ =spores present.
—=spores not seen.

Note.—The figure of “‘ 100% infection ’’ given in the above table for seed samples coming from Italy, France,
Germany and America is a misleading one in view of the small number of samples examined from these countries.
Examination of further samples would probably reveal the presence of some without rust spores.

2. Spread of the Disease in New South Wales.

After the first outbreak of rust was found in a Sydney suburb it spread rapidly
throughout eastern New South Wales. Within four and a half months it had been
recorded 100 miles from its source and within a year from its first discovery was
widespread in eastern New South Wales.

From a study of the records it has been found that spread in New South Wales
occurred in two main ways. These were: (i) primary spread by wind dispersal of
uredospores and (ii) secondary spread by the carrying of the diseased seedlings and
cuttings from place to place.

(i) Spread by wind-borne uredospores.

Wind is one of the most important agents for the dispersal of plant pathogens.
With rust fungi, the uredospore stage is well adapted for wind spread and considerable
work has been done in tracing the spread by air currents of many of these organisms,
especially the cereal rusts. Lambert (1929) showed that evidence exists that uredospores
of wheat stem rust may be carried in air currents several hundred miles from their

152 ANTIRRHINUM RUST IN AUSTRALIA,

original source. Wallace (1932), studying the prevalence of various physiologic races
of wheat stem rust in the United States in various years, found striking evidence in
some years of a south to north migration of rust. This migration was by means of
wind-borne uredospores formed early in the season in Mexico and Texas, and carried
northward by southerly winds. In western Canada, Craigie (1945) found that the
initial inoculum for the production of wheat stem rust consisted largely, if not
entirely, of wind-borne spores that originated outside western Canada.

In New South Wales, strong evidence is presented that the initial spread of
snapdragon rust was by means of wind-borne uredospores. Text-figure 2 shows the.
nature of this spread. Zone 1 represents the initial outbreak area of the disease,
and Zones 2, 3 and 4 its spread 9 weeks, 15 weeks and 18 weeks after the first
outbreak. Text-figure 2 was prepared by marking in the furthest points from the
initial outbreak area at which the disease had been recorded at the end of 9, 15 and 18
weeks respectively from the time of the first outbreak and then joining these points
up to give the three zones shown.

TABLE 4.

Surface Wind Direction Records at Sydney Weather Bureau for the Period from
1/11/1952 to 28/2/1953.

; | |
Wind | November, December, | January, February,
Direction. 1952. 1952. 1953. 1953.
N. 1 0 0 0)
Ss. 2 8 9 4
E. | 7 1 6 10
Ww... 13 3 2 2
N.-E. 29 | 47 | 40 29
E.-S. 13 14 29 23
S.-W. 1 10 | 2 7
W.-N. 10 8 2 8
Calm 4 2 3 1
Total 90 93 93 84

It can be seen from Text-figure 2 that the main direction of spread of the rust
during this period was in a south-westerly direction and only after this did a northward
spread to Gosford and the north coast of New South Wales occur. Surface wind
direction records at Sydney Weather Bureau have been obtained for this period and
are summarized in Table 4. Records are taken three times a day, at 9 am., 3 p.m.
and 9 p.m., and the table shows the number of records of each wind direction for the
period from ist November, 1952, to 28th February, 1953. It can readily be seen that,
in each month, wind was recorded more from a north-easterly direction than from any
other; in fact, over the four-monthly period 46% of the records were from the north
to east quarter and the prevailing winds for this period are given as north-easterly.

A definite correlation between prevailing wind direction and the initial direction
ot rust spread in New South Wales is thus seen to exist, and the evidence is strongly
in favour of wind-borne uredospores as the primary means by which the disease
spread in this State.

From the map it is seen that during the first four months spread was mainly in a
south-westerly direction, and in 18 weeks the disease had been recorded 100 miles from
its initial focus. Considering the comparatively thin population of snapdragons
throughout the area enclosed by Zone 4, this spread is quite rapid and shows the
efficiency of the wind-borne uredospores in extending the range of the fungus. From
the ninth to the eighteenth week of spread, i.e. from the edge of Zone 2 to the edge of

BY J. WALKER. 5S

Zone 4, the rust spread over an area approximately 10 times greater than that which
it had covered during its first nine weeks. This indicates that an initial build-up of
uredospore inoculum occurred on snapdragons in the Sydney suburban area (Zone 1
and part of Zone 2) during the first nine weeks and later spread in epidemic proportions
over a much wider area. The rapid spread and widespread destruction had all the
marks of a progressive or spreading epidemic, characteristic of the invasion of a new
area by a pathogen (Gaumann, 1950). The severity of the epidemic was increased by
the fully susceptible nature of all the host varieties being grown.

(ii) Spread on diseased plant material.

Once the disease had become established over a fairly wide area by means of
wind-borne uredospores, secondary spread began to occur by the transport of diseased
seedlings and cuttings from this area to other parts of the State. Quite often the
transported plant material was only very lightly infected and the disease escaped
attention, but, when planted out, the disease developed and set up secondary foci of
infection from which further wind spread could occur. In this way it was introduced
to Cooma in southern New South Wales, about 270 miles from Sydney, and also to
other areas. This considerably assisted its spread in Australia. Soon after the Cooma
outbreak it was widespread in southern New South Wales.

This vegetative transmission of snapdragon rust hastened its spread by carrying
it long distances and allowing it to become established in widely scattered places in
a comparatively short time.

3. Spread of the Disease in Australia.

By the methods outlined above, snapdragon rust became widespread in eastern
New South Wales. Very soon after, spread to other States in Australia occurred and
the disease was reported from the southern State of Victoria. Later, in the spring of
1953, it was recorded from Queensland and, in January, 1954, from South Australia.
By March it had reached Tasmania, leaving Western Australia as the only State still
free of the disease at the time of writing.

The exact method of introduction of the disease to other States is not known but
most probably both wind and vegetative spread were concerned. It is known that
diseased seedlings were sold in some States and quite possibly they provided foci from
which further spread by wind occurred.

4. Outbreak of the Disease in New Zealand.

Towards the end of December, 1953, rusted snapdragons were observed in a private
garden at Auckland, New Zealand. Brien (1954, private communication) states that
no further spread from this area appears to have occurred.

Possible sources of infection include seed and wind-borne uredospores. The question
of seed transmission has been discussed in the case of the New South Wales outbreak
and similar conclusions were reached in New Zealand. It is possible, however, that
uredospores have been carried by wind from Australia to New Zealand and resulted
in the outbreak of the disease in that country. Quite often, dust from Australia is
deposited in New Zealand, having been carried across by north-west winds, and it
would seem quite possible that with the large bulk of uredospore inoculum available
in eastern Australia some uredospores would reach New Zealand in air currents. The
distance from Sydney to Auckland is about 1,400 miles, but it is not improbable that
spores could be carried this distance in air currents.

SUMMARY.

1. Antirrhinum rust, caused by the fungus Puccinia antirrhini D. & H. was first
recorded in Australia in October, 1952.

2. The race present is similar to or identical with that known in America as race 2

and no varieties have been seen that are resistant to it. Two varieties of Linaria
inoculated with the rust showed no sign of infection.

154 ANTIRRHINUM RUST IN AUSTRALIA,

3. Locally produced uredospores were found to remain alive for 116 days ‘under
conditions of low temperature (5° and 10°C.) and relative humidities (25% and 50%).
As temperature and humidity rose, the longevity of the spores decreased.

4. The entry of the rust disease is discussed and it appears that the most probable
method of entry was as spore dust in seed samples. At planting time this dust could
be blown to nearby plants and infect them. Several points against this hypothesis
are discussed.

5. An American origin for the Australian outbreak is tentatively proposed from
evidence based on the physiologic race of the rust fungus present.

6. The spread of the disease in Australia is seen to consist of three main phases:
(i) primary spread from its initial focus by means of wind-borne uredospores;
(ii) secondary spread in New South Wales on diseased plant material; (iii) spread
throughout Australia by wind-borne uredospores which were spread from secondary
foci of infection established by diseased planting material.

7. The recent outbreak of the disease in New Zealand is mentioned and a possible
Australian origin put forward for it.

Acknowledgements.

The author wishes to thank Dr. C. J. Magee, Chief Biologist, and Dr. Lilian Fraser,
of the Department of Agriculture, for advice and suggestions during this work. To
Dr. K. F. Baker, of California, and Mr. D. EK. Green, of Wisley, England, thanks are
given for seed of some of the snapdragon varieties used and for advice on the disease.
He is also indebted to Dr. T. H. Harrison for notes on the spread of the disease in
Australia, to R. M. Brien, of New Zealand, for informaticn on the occurrence of rust
in that country, to the Sydney Weather Bureau for wind records, and to Mr. J. Rengger
for Plate vii, 3 and 5.

References.
BLASDALE, W. C., 1903.—On a rust of the cultivated snapdragon. J. Mycol., 9: 81-82.
Botromuey, A. M., 1940.—A destructive Antirrhinum disease new to South Africa. S. Afr.
hort. J., 2: 17. Abs. in R.A.M., 19: 350, 1940.
CRAIGIE, J. H., 1945.—Epidemiology of stem rust in Western Canada. Sei. Agric., 25: 285-401.
Dickson, B. T., 1921.—Plant diseases of 1920-21. Thirteenth Ann. Rept. Quebec Society for
the Protection of Plants, pp. 66-67. Abs. in R.A.M., 1: 374-375, 1922.

DIETEL, P., and Hotway, H. W. D., 1899.—Hedwigia, 36: 298 (from Blasdale, 1903). ~
Dimock, A. W., and BAksr, K. F., 1951.—Effect of climate on disease development, injuriousness
and fungicidal control as exemplified by snapdragon rust. Phytopathology, 41: 536-552.

Doiwee, KH. M., 1941.—South African Rust Fungi. IV. Bothalia, 4: 229-236.

Fixry, A., 1937.—Hgypt: appearance of Antirrhinum rust in the country. Int. Bull. Pl. Prot.,
iis als

, 1938.—Rust pustules on roots of Antirrhinum. Ann. Bot., Lond., N.S., 2: 536-537.

GAUMANN, EH., 1950.—Principles of Plant Infection. Pp. 543. Crosby Lockwood and Son, Ltd.,
London.

GREEN, D. E., 1934.—Antirrhinum Rust. <A disease new to Great Britain. J. Rk. hort. Soc.,
DOR eel 9 = 1216) :

- , 1941.—Antirrhinum Rust. IV. Improvement of rust-resistant warieties. J. R. hort.
Soc., 66: 83-86.

HASSEBRAUK, K., 1937.—Zur Frage der Verbreitung des LOwenmaulrostes durch das Saatgut.
Blumen- wu. Pfl. Baw ver. Gartenwelt, 41: 261-262. Abs. in R.A.M., 16: 752, 1937.

JORSTAD, I., 1946.—Snapdragon rust and snapdragon mildew. Two newly introduced parasitic
fungi. Norg. Gart. Foren. Tidsskr., 36: 497-498. Abs. in R.A.M., 26: 398, 1947.

LAMBERT, E. B., 1929.—The relation of weather to the development of stem rust in the
Mississippi Valley. Phytopathology, 19: 1-71.

Lepik, H., 1941.—Spread of snapdragon rust in Hurope. IJnt. Bull. Pl. Prot., 15: 93m.

MAINS, E. B., 1924.—Notes on the life history of the snapdragon rust, Puccinia antirrhini Diet.
and Holw. Phytopathology, 14: 281-287. °

Pau, B. T., 1937.—FPuccinia antirrhini Diet. et Holw. pa Oland. Svensk bot. Tidskr., 31:
288-289. Abs. in R.A.M., 16: 679, 19387.

PELTIER, G. L., 1919.—Snapdragon rust. Univ. Ill. Agric. Hxup. Sta. Bull., 221: 535-548.

PrRet1I, G., 1985.—Una malattia dell’ Antirrhinim majus Ll. nuova per la micologia italiana
(Puccinia antirrhini Dietel e Holway). kit. P-dta Veg. 253 301-32. Abs. in R.A.M.,
15: 228, 1936.

PLATE Iv.

Proc. Linn. Soc. N.S.W., 1954.

Aeschiidiopsis flindersiensis (Woodward).

Proc, Linn. Soc, N.S.W., 1954, PLATE VY,

Bacteria from Macquarie Island.

Proc. LINN. Soc. N.S.W., 1954. PLATE VI.

Collenia frequens.

Proc. LINN. Soc. N.S.W., 1954.

PLATE VII.

. & H.

tirrhini

mia an

Antirrhinum Rust, Puec

BY J. WALKER. 155

Rayss, T., 1937.—Contributions a la connaissance des Urédinées de Palestine. Reprinted
from “Hommage au Professeur E. C. Teodoresco’’, Bucharest, 1937, pp. 13. Abs. in R.A.M.,
18: 204, 1939.

WALLACE, G. B., 1952.—1950 Annual Report of the Plant Pathologist, Lyamungu, Moshi. Rep.
Dep. Agric. Tanganyika, 1950, pp. 167-172. Abs. in R.A.M., 32:117, 1953.

WALLACE, J. M., 1932.—Physiologic specialisation as a factor in the epiphytology of Puccinia
graminis tritici. Phytopathology, 22:105-142.

WHETZEL, H. H., 1924.—Special Report. Repts. Board and Dept. of Agric. Bermuda for the
Year 1923, pp. 45-52.

Yarwoop, C. E., 1937.—Physiologic races of snapdragon rust. Phytopathology, 27: 113-115.

EXPLANATION OF PLATE VII.
1. Rust-infected leaves showing concentric ring development of uredosori. x 1.

2. Rust-infected leaves, the leaf on the left showing the chlorotic area developed on the
top surface and the one on the right showing uredosorus development on the under surface
of a similar leaf. x1.

3. Rust on flower head. x 3.

4. Rust-infected leaves, showing the necrotic area developing. Leaf on left shows the
top surface and the one on the right the under surface. x1.

5. Snapdragon plants showing heavy rust infection. x 3

ACARINE PARASITES OF TWO SPECIES OF RATTUS FROM BRISBANE,
AUSTRALIA.

By Rosert Domrow, Queensland Institute of Medical Research, Brisbane.
[Read 29th September, 1954.]

Synopsis.

The mites and ticks were identified from 51 rats, of which nine out of 18 R. rattus and
21 out of 33 R. norvegicus harboured parasites. Nine species belonging to six families were
found, bringing the total known from Australia to eleven.

The acarine parasites of R. rattus and R. norvegicus have been listed from several
countries, but never from Australia. Hirst (1914) recorded Bdellonyssus bacoti (Hirst)
from Australia, and Womersley (1937) recorded Laelaps echidninus Berlese and
L. nuttalli Hirst on both species of rats from several loealities. There are two records
from south-east Queensland, Haemaphysalis humerosa Warb. & Nutt. (Ixodidae) from
R. rattus at Ipswich and Laelaps anomalus Hirst from R. norvegicus at Brisbane
(Annual Report of the Health and Medical Services of the State of Queensland for
the year 1937-38). ,

Some of these species are discussed further below, together with additional material
collected from 51 wild rats (18 R. rattus and 33 R. norvegicus) captured in Brisbane
over the period August, 1950, to June, 1952. The specimens were mounted by Dr. E. H.
Derrick and Mr. J. H. Pope, of this Institute. Nine species of acarines from six families
were detected.

I wish to thank Drs. I. M. Mackerras and EK. H. Derrick, of this Institute, for their
kind advice during the preparation of this paper.

Genus CoOSMOLAELAPS Berlese, 1903 (Laelaptidae).
CoOSMOLAELAPS, sp. n. in ms. Womersley.
Two females of this species were taken on one specimen of R. norvegicus captured
at this Institute, Ist December, 1950.

Genus LAELAPsS Koch, 1839 (Laelaptidae).
LAELAPS NUTTALLI Hirst, 1915.

Excellent figures of this species are to be found in Hirst’s original paper and in
Womersley (1937). This species was by far the commonest encountered, being collected
on four R. rattus and 19 R. norvegicus in numbers varying from three to 62, wandering
about through the fur. However, all the mites present were not collected in every
case. There was a marked preponderance of females over males, the ratio being 5:1,
and most were gravid, each carrying a single, large egg, in which a larva was often
discernible. Nymphs were slightly more numerous than males. From four R. rattus,
34 99, one 4, February to May, 1951; from 19 R. norvegicus, 320 99, 64 gg, 87 nymphs,
August, 1950, to June, 1952.

LAELAPS ECHIDNINUS Berlese, 1887.

This species is adequately described and figured in Hirst (1913, 1922) and Baker
and Wharton (1952). Only four specimens, all females, were taken on both species of
rats. The presence of this species is of interest, since it transmits Hepatozoon muris
(Balfour), which is common among wild rats in Brisbane.

BY ROBERT DOMROW. 157

Genus GYMNOLAELAPS Berlese, 1916 (Laelaptidae).
GYMNOLAELAPS, sp. n. in ms. Womersley.
Forty females of this species were taken from three FR. rattus from Taringa,
Brisbane, 6th October, 1950.

Genus BpELLONYSSUS Fonseca, 1941 (Macronyssidae).
BDELLONYSSUS BACOTI (Hirst, 1913).

Adequate descriptions and figures of this species are to be found in the original
paper and in Hirst (1922). This species, represented by two females, was taken on a
single specimen of R. norvegicus (Brisbane, 10th August, 1951). They were both
blood-filled. However, it is more common than this record would suggest.

Genus Ixoprs Latreille, 1795 (Ixodidae).
IxODES TASMANI Neumann, 1899.
This native species was found as larvae on two R. rattus (Taringa, October, 1950).
All the seven larvae were attached on or near the ear margins. They were identified by
Dr. F. H. S. Roberts, of the C.S.I.R.O. Animal Health Laboratory, Yeerongpilly,
Brisbane.

Genus RApDFORDIA Ewing, 1938 (Myobiidae).

RADFORDIA ENSIFERA (Poppe, 1896).

Figures and descriptions are to be found in the original paper and Radford (1949).
In Radford’s fig. 27, four posterior dorsal median setae are shown. Actually two of
these are ventral, and Poppe, fig. 14, is correct. Skidmore (1934) found this species
causing injury to white rats. He states: “these rats were scratching themselves
about the head, nose and neck. Many had small, dry scabs on the head, about the
ears and upper sides of the head. Some had blood scabs due to bleeding caused by
severe scratching.” One R. norvegicus with this parasite had wart-like growths on the
ear-edge which may have been caused by scratching. The species was quite common,
being taken on two R. rattus (nine gg, one nymph) and six R. norvegicus (49 99,

14 Jd).

Genus NOoTOEDRES Railliet, 1893 (Sarcoptidae) .
NOTOEDRES MURIS (Mégnin, 1877).

This species is figured in Hirst (1922). Death may follow heavy infestation. This
mite was found on two specimens of R. norvegicus (30 99, mostly. gravid, and three
nymphs). One of the rats had itch-scales on the ear, shaped like small, hollow domes
with a hole in the top. On dissecting these open in lactophenol, Notoedres mites were
found inside, bearing out Hirst’s remark, “this mite especially affects the ears of the
host ... giving rise to characteristic crusts and warty excrescences’’.

Genus LEpIpOGLYPHUS Zakhvatkin, 1936 (Glycyphagidae).
LEPIDOGLYPHUS DESTRUCTOR (Schrank, 1781).

This species is a serious pest of grain products. One male was taken off R. rattus
from Taringa, Brisbane, 7th October, 1950. It is an accidental record.

In addition to this mite, two specimens probably of the family Acaridae were found
on one R. norvegicus captured at this Institute, 7th October, 1950. This rat was caught
in the animal feed store, where large amounts of farinaceous material are kept. These
mites again are not true parasites.

One R. norvegicus was of interest, harbouring four species of mites, Cosmolaelaps,
sp. n. in ms. Womersley, L. nuttalli, R. ensifera, and N. muris. None of the other rats
had more than two species present. The complete absence of Trombiculidae is perhaps
to be correlated with the city-dwelling habit of the rats examined.

References
BAKER, H. W., and WHARTON, G. W., 1952.—An Introduction to Acarology, New York, Macmillan.
465 pp.
Hirst, S., 1913.—On three new species of Gamasid mites found on rats. Bull. Ent. Res.,
4:119-124.

158 ACARINE PARASITES OF TWO SPECIES OF RATTUS.

Hirst, S., 1914.—On the parasitic Acari found on the species of rodents frequenting human

habitations in Egypt. Bull. Hnt. Res., 5: 215-219.
, 1915.—On some new Acarine parasites of rats. Bull. Ent. Res., 6: 183-190.

———,, 1922.—Mites Injurious to Domestic Animals. Brit. Mus. (Nat. Hist.) Economic
Series, No. 13.

Poppr, S. A., 1896.—Beitrag zur Kenntnis der Gattung Myobia von Heyden. Zool. Anz., 19: 341.

RADFORD, C. D., 1949.—A revision of the fur mites, Myobiidae, Acarina. Bull. Mus. Hist. nat.
Paris, 2e Serie, 21: 91-97. :

SKIDMORE, L. V., 1934.—Acariasis of the white rat (Rattus norvegicus form albinus). Canad.
Ent., 66: 110-115.

WoOMERSLEY, H., 1937.—Studies in Australian Acarina, Laelaptidae. 1. New records and species
of Laelaps and allied genera. Parasitology, 29: 530-538.

159

A SECOND SPECIES OF HOLOTHYRUS (ACARINA: HOLOTHYROIDHA)
FROM AUSTRALIA.

By Rosert DomMRow,
Queensland Institute of Medical Research, Brisbane.

(One Text-figure. )
[Read 29th September, 1954.]

Synopsis.
Holothyrus constrictus, n. sp., from South Queensland, is described and discussed. The
key of Womersley (1935) to the known world species is amended to include it.

The rare suborder Holothyroidea includes only the genus Holothyrus Gervais, 1842,
with seven described species, of which one is from Mauritius, three from Seychelles, two
from New Guinea, and one (H. australasiae Wom., 1935) from South Australia and the
North Island of New Zealand. An eighth species from South Queensland is described
in the present paper.

The history of the genus is fully discussed by Thon (1906), and summarized by
Womersley (1935). All the species are forest dwellers, living in humus and the like.
The Australian forms are the smallest known, not exceeding 3 mm. It is also of note
that the female genital shields of the Australian forms differ from those figured by
Thon, and copied by Tragardh (1938), the anterior and lateral shields being much more
definite and well developed, as shown by Womersley (Plate VIII), and in Figure 1A
below. The two Australian forms are also the only ones described as having distinct
processes on the femur, genu and tibia of leg II in the male. However, in all other
respects they are typical of the genus, and there seems no good reason to separate them.
Full generic details are given by Thon and Womersley, and minor details by Vitzthum
(1941) and Baker and Wharton (1952).

I wish to thank Mr. H. Womersley, of the South Australian Museum, Adelaide,
where this work was done, and Dr. I. M. Mackerras, of this Institute, for their kind
advice during the preparation of this paper.

Family HOLOTHYRIDAE Thorell, 1882.
Genus HoLorHyrus Gervais, 1842.
HOLOTHYRUS CONSTRICTUS, DL. Sp.

Types: The holotype female and allotype male are in the collection of the South
Australian Museum, Adelaide, and two paratype males in the collection of the Queens-
land Institute of Medical Research, Brisbane. All four specimens were collected in a
composite lot of soil and litter from Brookfield, Brisbane, Queensland, 21st May, 1949.
by Dr. E. H. Derrick.

Female.

A small, dark brown, heavily chitinized species; idiosoma length 2080 wu, breadth
1150 uw. Dorsal shield completely covering dorsum, and underlapping venter as shown;
raised into small, irregularly-spaced tubercles, each carrying a short seta 18 to 25 wu long,
which has one edge with a double row of short ciliations, while the other edge is smooth;
shield very finely granulated, with punctate, linear semi-circular, and peculiar eye-like
pores.

: Venter (Fig. 1A), sternal, metasternal, endopodal, ventral and anal shields fused
to form holoventral shield, 1488 » long, 720 u wide at greatest width (behind coxae IV),
markedly thickened around acetabula (representing the endopodal shields), and
especially so at the anterior sternal part of the shield; sternal section of shield concave
anteriorly, broadening to behind coxae IV, whence ventri-anal portion tapers slowly

160 SECOND SPECIES OF HOLOTHYRUS FROM AUSTRALIA,

to posterior of venter; at least two pairs of longer sternal setae and pores distinguishable
from the accessory setae. Genital shield (Fig. 1B) wider basally than high, 352 uw x 304 u,
with two anterior horns, behind which the shield constricts markedly, and then expands
broadly to its base, as figured. Lateral shield (Fig. 1A) longer than broad, 352 w x 192 uw,
narrower anteriorly, and with rounded ends, as figured. Median shield (Fig. 1A)
broader than long, 208 uw x 160 uw, tapering twice to posterior. Two anal valves (Fig. 1A)
roughly semi-circular, 96 uw x 46 uw, each with three or four simple, short setae. Sternal
and genital shields covered by thickly set, simple setae to 54 w long, posterior part of
yenter with simple setae on tubercles, except at extreme posterior where a few setae

Text-figure 1.—Holothyrus constrictus, n. sp.

A-E, female, F-M, male.—A, ventral view; B, genital shield; C, chelicera; D, ventral view
of gnathosoma; EH, dorsal shield of H. constrictus, n. sp., above, and H. australasiae Wom., below ;
F, leg I; G, leg II; H, leg III; I, leg IV; J, genital shield; K, pit behind coxa IV; L, palpal
tibia + tarsus; M, chelicera.

are ciliated like those on dorsum. Behind each coxa IV a sub-spherical pit (Fig. 1K)
with aperture guarded by heavy brush of fine setae. Cuticle between dorsal and holo-
ventral shield without setae. Tritosternum present, but small, with two weakly ciliated
laciniae.

Gnathosoma (Fig. 1D) enclosed in camerostome formed by anterior part of dorsal
shield. Palpi with five free segments, palpal tibia expanded medially, tibia + tarsus
176 « xX 78 uw, with 4-tined sensory seta at tip of tarsus, palpal claws indistinct. Chelicerae
(Fig. 1C) large, fixed finger with two main teeth, and finely serrate edge between teeth,
and with two setae dorsally at base, movable finger 264 uw, also with two teeth and

BY ROBERT DOMROW. 161

serrations. Legs fairly long and slender, closely tuberculate like dorsum (except on
coxae), all with two claws, caruncle and pad (caruncle reduced on leg I); leg I 1490 uy,
leg II 1170 uw, leg III 1180 yp, leg IV 1740 uw.

Male.

Somewhat smaller than female, heavily chitinized; idiosoma length 1760 u, breadth
960 uw. Dorsal shield of same structure as female, entirely covering dorsum, and under-
lapping venter by approximately 80 uw. Holoventral shield of similar composition to
female, 1360 u x 688 uw (behind coxae IV). Genital shield (Fig. 1J) quadrate with
rounded corners, broader than long, 168 w x 116 uw; genital aperture sub-circular, 168 uw
in diameter. Anal valves longer than broad, 80 wu x 42 uw, with three or four pairs
simple setae. Glands present behind coxae IV. Ventral cuticle ornamented posteriorly
with tubercles as in female. Tritosternum present, small (only very narrow space
between sternal margin and gnathosoma).

Gnathosoma and palpi as in female; palpal tibia + tarsus (Fig. 1L) 176 uw x 88 ux.
Chelicerae (Fig. 1M) very similar to female, but second tooth on movable finger slightly
stronger; length of movable finger 256 uw. Legs (Fig. 1F—-I) fairly long, slender, except
leg II; all with caruncle, two claws and pad (caruncle of leg I reduced). Femur II with
large process ornamented as general leg surface, femur III with similar, smaller process,
genu II and tibia II with simple, small process, as figured, coxae of all legs smooth;
leg I 1490 pw, leg II 1200 u, leg III 1280 uw, leg IV. 1810 wu.

Distribution—Known only from the type locality in South Queensland.

Taxonomic notes.

This species may be readily separated from the other known Australian species,
H. australasiae Wom., by the great difference in cuticular ornamentation (Fig. 1E),
and from the other species by its very small size and armature of leg II in the male.
In dH. constrictus, n. sp., the cuticle of the dorsal shield in both sexes has numerous
irregularly-spaced tubercles, each bearing a single, short seta, 18 u to 25 uw long. These
setae are stout, and have a double row of ciliations on one side only. Between the
tubercles are punctate, semi-circular and eye-like pores. The tubercles are present
ventrally on the posterior half, but only the setae on the extreme posterior tubercles
are ciliated. The setae on the anterior half of the venter are simple and not set on
tubercles, to 35 uw in the male and to 50 uw in the female. In H. australasiae Wom. the
tubercles are smaller and not set so closely together, and the setae are not set on them,
but directly in the cuticle. The setae are much longer, to 90 u, and simple. The ventral
surface is rather similar to the dorsal, with somewhat shorter setae. The general
cuticular surface in both species is very minutely granulated.

The key of Womersley may be amended to include the new species by inserting
the following caption:

4a. Leg II of male with brushes of setae on femur, genu and tibia in addition to processes;
female genital shield not constricted behind anterior horns; male 2:9 mm., female 2:8 mm.

(South Australia and New Zealand).................... H. australasiae Womersley, 1935.
Leg II of male without such brushes; female genital shield markedly constricted behind
anterior horns; male 1-76 mm., female 2 mm. (Queensland)......... H. constrictus, n. sp.

There have been two errors in recent literature concerning this genus which should
be noted. Radford (1950) in his checklist gives the reference to the genotype as
“Ann. Soc. ent. France, 2”, etc., but this should read ‘11’. Secondly, in Baker and
Wharton’s book (1952), Fig. 33 (page 39) is labelled as ‘“Holothyrus longipes Thorell,
1882. (After Hirst, 1922.)”. Hirst’s figure 77, page 92, is simply labelled “Holothyrus sp.
(After Thorell.)”. Vitzthum (1941, Abb. 478, S. 752) also reproduces the same figure by
Thorell, and labels it (correctly) ‘‘Holothyrus nitidissimus Thorell, 1882, nach Thorell’’.
Thorell described four forms of the genus in 1882 (Descrizione di alecuni Aracnidi
inferiori dell’ archipelago malese, in: Ann Mus. civ. Stor. nat Genova, 18), viz., H.
longipes, H. 1. var. ferrugineus, H. nitidissimus and H. scutifer. Of these, Thorell himself
said H. 1. var. ferrugineus was a nymphal form, and that ‘“nitidissimo haec forma

162 SECOND SPECIES OF HOLOTHYRUS FROM AUSTRALIA,

[scutifer] valde similis est, et forsitan alter sexus ejus”. Thon regards this to be true,
and only two of Thorell’s four forms are valid, H. longipes and H. nitidissimus. Thon
re-examined H. longipes and stated: “Die Fiisze sind sehr lang und auffallend schlank
wie bei keiner andern Art. Thorell hat den Artnamen sehr treffend gewahlt. Der 1. Fusz
ist der langste, unbedeutend langer als der letzte.’ He gives the ratio of body length:
length leg I: length leg IV as 17:41:40. The figure given by Baker and Wharton, then,
cannot be Holothyrus longipes Thorell, 1882, but is really Holothyrus nitidissimus
Thorell, 1882, as given by Vitzthum.

References

Baker, E. W., and WHARTON, G. W., 1952.—An Introduction to Acarology. New York,
Macmillan, 38-39.

Hirst, S., 1922.—Mites Injurious to Domestic Animals. Brit. Mus. (Nat. Hist.), Hconomic
Series, No. 13, p. 92.

RApDForRD, C. D., 1950.—Systematie Checklist of Mite Genera and Type Species. Secrétariat
général de 1’U.1.8.B., Série C (Section d’Entomologie), No. 1. Paris, p. 9.

THON, K., 1906.—Die atissere Morphologie und die Systematik der Holothyriden. Zool. Jahrb.
(Syst.), 238: 677-724.

TRAGARDH, I., 1938.—Further contributions towards the comparative morphology and classi-
fication of the Mesostigmata. Ent. Tidskr., 59: 123-158.

V1ITZTHUM, H. GRaF, 1941.—Acarina, in Bronn’s Klassen und Ordnungen des Tierreichs. Leipzig.
5, Abt. IV, Buch 5, p. 752.

WoOMERSLEY, H., 1935.—A Species of Acarina of the genus Holothyrus from Australia and New
Zealand. Ann. Mag. Nat. Hist. (10), 16: 154-157.

163

THE NATURE AND SIGNIFICANCE OF NON-RECIPROCAL FERTILITY IN
AEDES SCUTELLARIS AND OTHER MOSQUITOES.

By S. SMITH-WHITE (Dept. of Botany), and A. R. WoopHILL (Dept. of Zoology),
University of Sydney.

(Three Text-figures. )
[Read 29th September, 1954.]

Synopsis.

When crosses are made between two subspecies of Aédes scutellaris, the matings prove
fertile or infertile according to the direction in which the matings are made. In the mating
of subsp. scwtellaris (S) females with subsp. katherinensis (K) males, fully viable eggs are
produced. In the reciprocal mating, copulation, insemination and egg laying are normal, but
the eggs are totally inviable.

This non-reciprocal fertility shows strictly maternal inheritance. Backcrosses of F,
females to subsp. katherinensis males are viable and the B, progeny are of scutellaris mating
type. In successive backcrosses to katherinensis males, to the B, generation, the scutellaris
mating type is retained. Bex males, derived from repeated backcrosses to subspecies
katherinensis, are still incompatible with katherinensis females.

The genetic system determining the inheritance of mating type must depend either on
anomalous meiosis in oogenesis or on nucleus-independent cytoplasmic factors. There is no
critical evidence enabling a choice between these two hypotheses.

A: survey of the available data on non-reciprocal fertility between species, subspecies, and
races of Aédes and Culex suggests that it has had significance as a source of incipient
speciation in mosquitoes.

INTRODUCTION.

The natural history of mosquitoes has long held a place of importance in entomo-
logical research, and the systematics of the Culicidae has advanced to a stage where the
limits between species are often finely drawn on the bases of morphological, physiological
and ecological criteria. The application of the genetic concept of species, which
emphasizes intrinsic isolation, is likely to cause some reorientation of the taxonomy of
the group in the near future.

The isolating mechanisms which exist between closely allied species include a range
of types similar to those known in Drosophila. They may be genetic, mechanical,
ecological, physiological, or behaviouristic in nature. Such isolating mechanisms usually
operate in both reciprocal directions between males and females of the species concerned.
Differences in fertility between reciprocal crosses, however, are known to occur between
races, subspecies or species in several genera (Toumanoff, 1939, 1950; Downs and Baker,
1949; Bonnet, 1950; Perry, 1950; Woodhill, 1949, 1950; Marshall, 1938; Laven, 1951,
1953; Dobrotworsky and Drummond, 1953) and it is apparent that the phenomenon is
widespread in the Culicidae. It has introduced difficulties and complexities in the
appreciation of specific and subspecific categories, and it has a significant bearing on
problems of medical entomology. It is also significant to genetical and evolutionary
theory. Its mechanics must depend on a uniparental genetic system, and its role in the
origin of intrinsic species barriers is problematical.

THE MATERIAL.

Aédes scutellaris Walker belongs to a species-complex which includes a number of
species, subspecies and geographical races of doubtful rank. Many of the forms show
complete intrinsic isolation, and undoubtedly deserve specific status. A peculiar non-
reciprocal fertility was described by Woodhill (1949, 1950) in crosses between two
subspecies, A. scutellaris scutellaris Walker and A. scutellaris katherinensis Woodhill.
Complete fertility was found when subsp. scutellaris females and subsp. katherinensis
males were mated, but hybrid eggs from the reciprocal cross were totally inviable. This

164 NON-RECIPROCAL FERTILITY IN AEDES SCUTELLARIS AND OTHER MOSQUITOES,

mating incompatibility was found to extend to the backcrosses to katherinensis.
Smith-White (1950) drew attention to the possible genetic and evolutionary significance
of the phenomenon, and suggested a backcross program which might clarify its nature.

The original isolation of subsp. katherinensis was obtained from Katherine, in the
Northern Territory, in January, 1948, and consisted of a batch of eggs from an unknown
number of females. This isolation has been maintained in the laboratory under the
culture designation “K’. An isolation of subsp. scutellaris was obtained from New
cTuinea at about the same time, and has been maintained as culture ‘‘S”.

Q co Q oi

K? ie ise Ss x K?
F2 as
die ey ae

Ke cnaear aE el
Bix me
die ya

B B

K x Bp Bix x K
Box Box
die SS an.

ne x Bay Bo. x ie

B

Bay Bs
die yr

Ke x Bay Bere eX kK"

Bax Bax
die ee
a
B 4k

K* Xin x Ken
eg
B 5. B 5k
die ee
K° Dileep Bey ax K?
Box Box
die SUTViVve

Text-figure 1.—The Backcross Program.
S = A. scutellaris scutellaris; K = A. scutellaris katherinensis ; a and 8B represent maternally
inherited factors, affecting survival. They may be carried either in the cytoplasm or in
the nuclei.

In April, 1953, Mr. K. O’Gower obtained eggs from eleven females of subsp.
katherinensis at Batchelor, 126 miles north-west of Katherine. This new isolation,
designated culture ‘“B”, is morphologically identical with the Katherine material, and
shows an identical behaviour when crossed with subsp. scutellaris. In the cross § x B,
3200 F, eggs yielded 2495 larvae (77-9% hatch), but in the reciprocal cross B x S,
2514 eggs were totally inviable. A. scutellaris katherinensis possesses a wide
geographical distribution in northern Australia, and it can be distinguished from the
type subspecies by morphological criteria.

BY S. SMITH-WHITE AND A. R. WOODHILL. 165

Aédes scutellaris is a less satisfactory laboratory subject than is Culex molestus. It
requires warm temperatures and high humidity, which necessitate special culture rooms.
Blood-feeding is necessary before egg-laying, and matings are only successful when large
numbers of the two sexes are confined in the breeding cages. The eggs are laid singly,
and the progeny of individual females cannot easily be isolated from the mass mating
cages.

THE BREEDING PROGRAM.

The main part of the breeding program has consisted of a series of backcrosses to
test for permanence or breakdown of incompatibility with K females. Data on sex ratio
in cultures, and of F., segregation for a morphological character ‘‘white line’, are
reported. An inbreeding program with culture S was commenced for the isolation of
possible recessive genes, but has been abandoned temporarily.

All larval cultures were maintained in incubators at 27°C. and matings were made
in muslin cages measuring 10” x 10” x 12”, in a warm room at 27°C. and 75-80%
relative humidity.

The backcross data. Three series of backcrosses have been made. The first series
commenced with the F, hybrids produced by Woodhill (1950), and was continued to the
fifth backcross generation (Text-figure 1 and Table 1). In the Text-figure, and in Tables

TABLE 1.
The First Backcross Experiment.
Progeny Eggs | Percentage
Cross. Designation. | Eggs Laid. Hatched. Hatch.
fi
1 a 8" | er? = os 0
a 6 C2 |
2 S xk Ry | — — high
|
|
ee ot |
3 Fi a) B | 650 normal high
a a | (od |
4 S xF, Be | 1150 normal high
eo (8) a |
5 BF xK Bi | 4380 normal high
Oe, S| 6 | |
6 K xF By | 1220 0 | 0
o4 6 o4 | |
7 By xK Bo normal normal ca 90
8 Ke xB 5 | B » 2000 13 0-65
1k 2k | %
ce (6 a |
9 Boy SK Ba. | normal normal ca 90
10 meee ml | 4800: 2 0-25
2k | 3k
@ (6 a
11 Bo. xk Ba normal normal ca 90
6 a 6
12 K xB, Bay | 4200 58 1-38
od 6B io ;
13 B hie xk Bey | normal normal ca 90
6 6 |
14 K x By, Be. 4600 84 IP StSZy
6 _ 6 |
ify) Ba. xk B 4k | not recorded over 90
6 6 6 | |
16 K x Bay Bay | not recorded | over 90

ra
co
for)

NON-RECIPROCAL FERTILITY IN AEDES SCUTELLARIS AND OTHER MOSQUITOES,

1-3, the backcross formulae require brief explanation. The subscripts indicate the
generation, B,x being the third backcross to K after the F,, and the superscripts indicate
the “maternal line’ of each family, and are significant for any cytoplasmic or nuclear
factors having a strictly maternal inheritance. B,,¢ is the third backcross to K, tracing
its maternal ancestry back to the original S stock. BS has had the K maternal line

introduced.

TABLE 2.

The Second Series of Backecrossing.

Eggs Laid. Hatch.
Serial Progeny
No. Cross. Designation. Numbers Mated.
Total. % Total. %
Sa aes
a _@% |
1 | § _ selfed 5 @ 48 100 2022 42-1 1450 71:1
| ~)
Ot jen Keleselfed k 104 76 2104 20:2 1583 1°98
od od
3 Sisk Fy 56 62 2482 44-3 1499 60-4
Cree 6
4 Kelas) Fy 126 202 3390 27-0 (0) 0-0
| cane (C4 od i
Bg ae lee x F, Fy 135 205 3534 26-2 863/1254 68-7
| v
(4 4 4
6 F xs Bs 86 126 2006 23°3 320/573 55-8
|
C2 4 (od
0 Sie Fy B,. 164 150 9218 56-2 686/748 - 91-7
om 6 | or
3 Fy xK By 112 126 2589 23h 1647 63-6
9 Ke xe 2 | B P 91 103 1767 19-4 0 0-0
vest 1k :
10 B S ee : 2 2 0 70
M Qs 2, . 2 .
lk CK Bo. 82 68 189 23-1 134 7
11 a sein B 6 51 113 1408 27°6 0 0-0
aeell Dies |e :
12 B : e B a 5 7 27-2 1051 64-8
2 aK * k 3k 59 73 1618 2
13 Ke cB 7 B 6 88 56 1254 14°3 0 0-0
Bx 2k 2k 838 £5) G
14 B *: a B o 7 1831 23°8 1072 58-6
3k * he 77 91 83 7
115) asec - B » 92 92 1567 17:0 0 0-0
3k 4k i a
oe (6 4
16 B xk BL 71 98 1573 2292 1083 68:7
4k 5k
Le ses ‘ B ? 75 102 1564 20-8 0 0-0
ak 5k : ah
18 B re : 78 8 VAIL oF 989 60-5
Bk Be. 78 86 1654
19 ae B - F 19-9 0 0:0
cK xB Be. 69 68 1372

The first series of backcrosses show a slight breakdown in the inviability of K-line

hybrid eggs, in the B..8 generation, and there is a slight but very doubtfully significant
increase in breakdown in the later generations. Breakdown individuals possessed
complete fertility with the K parental stock in both reciprocal directions, but were not
tested against the 8 stock. These results parallel those obtained by Laven (1953). Since

BY S. SMITH-WHITE AND A. R. WOODHILL. 167

the successive backcrosses have increasing dosages of the K genotype, a more rapid
breakdown might have been expected.

The second series of backcrosses was carried out to collect additional data, to test
for the occurrence of breakdown, and to obtain breakdown individuals for a com-
patibility test with S stock. There was a complete absence of breakdown up to and
including the B,x.6 generation (Table 2). A third series of backcrosses was made solely
to check the occurrence of breakdown, and the result was also negative. It seems
probable that the breakdowns in the first experiment may have been due to an accidental
contamination of the F, x K mating with a K female. The complete inviability of eggs

TABLE 3.

Repeat Experiment to Test for “* Breakdown’’.
(Each mating of approx. 130 and 150.)

Seria: Progeny | | Percentage
No. Cross. Designation. | No. of Eggs. | Hatch. Hatch.
o8 ob | :
1 Ry xKk By normal normal | high
Bo B | |
2 Ke 310 1 B 1k 5600 | 0 | 0-0
Coe 1G} of |
3 Bi. xK Boy normal | normal high
a IG) : |
4 K xB lk B 2k 6894 0 | 0-0
|
a 88 a | | |
5 Boy xk By. normal | normal | high
6 me xB B | 6962 0 0-0
2k 3k =
446) a | ;
Cl Boy xk B 4k | normal normal | high
6 oe 6 | |
8 KelSaB} 3k B 4k | 5368 0 | 0-0
a 6 | |
9 B 4k xK B Bako normal normal | high
10 mT ap B » 5054 0 9-0
4k 5k

derived from K females and S maternal line males is maintained up to the Byxa
generation, and probably indefinitely. B..a individuals, predominantly K in genotype,
however, show full fertility and high egg viability when crossed to S stock in either
direction. The strictly maternal inheritance of some determinant, either on a chromo-
some or in the cytoplasm, is demanded.

Sex ratio. Data on sex ratios in S and K stocks, and in F, hybrids are presented
in Tables 4 and 5. In all cases there is a significant surplus of males, approximating to
a ratio of 1-25:1. The most probable cause of this inequality is differential mortality.
Males mature and emerge several days earlier than females, and the last larvae to
pupate in any culture are invariably female. In the larval and pupal cultures, the
death rate increases gradually with time, and there is no reason to assume other than a
normal genetic sex determination.

_ Segregation in the F,. The two subspecies differ in a small but distinctive character.
In K there is a line of white scales on the anterior surface of the femur of the mid-leg,
and this line is absent in the § race. In the F, there is a thin broken line, indicating an
absence of dominance. In the F, there is segregation for the character indicating a
monofactorial mendelian inheritance independent of sex (Table 6). Actually the genetic
‘control of the character is complex, and probably modifier genes influence its degree of

168 NON-RECIPROCAL FERTILITY IN AEDES SCUTELLARIS AND OTHER MOSQUITOES,

development. The significance of the segregation for the present purpose lies in the
independence of a major “line” gene and the sex chromosome. Normal meiotic segrega-
tion is indicated for two of the three chromosome pairs present in Aédes.

The inbreeding program. In the Aédes scutellaris group, mating seems to be con-
ditional on the formation of swarm flights of males, and multiple copulations may occur
for each female entering the swarm. Such a system seems adapted to the maintenance
of genetic heterogeneity.

TABLE 4.
Sex Ratios.
Rn |
Culture. Females. Males. Total. Sex Ratio.
|
S. 52/5B 176 330 EMG 4 ale ileey
52/11-B 92 214 | 306 1: 2-36
52/7-B 215 | 201 416 1:0-94
Total $ 483 | 745 1238 1:1-54
K. 52/1-B 39 45 84 1:1-15
52/4-B .. | 239 | 312 551 ibe Teel
52/10-B .. 105 | 84 189 1: 0-80
Total K 383 441 824 1:1-15
SxK (F,)* 546 | 672 1218 1:1-23

* See analysis in Table 5.

A need was felt for the isolation of genes with a marked phenotypic effect for use
as markers of chromosome segregation. An inbreeding program was commenced with
the laboratory stock 8, and based on the plan outlined by Spencer (1947). Mass matings
were made, and were given two blood feedings within three days. Gravid females were
then immediately isolated in 8” x 1” tubes, each with a pad of wet filter paper, and the

TABLE 5.
Sex Ratio in Sx K Progeny.
Nl
| Hatch. Adults Reared.
Egg Date | Number % Sex

Batch. Date Laid. Sown. | of Eggs. | Survival.| Ratio.

NOs | % fe) 3 Total.
iH 25-26/2/52 3/3/52 39 19 48-7 5 11 16 84:2 | 1:2-22
2 26-27 /2/52 38 139 96 69-1 24 31 95 57-3 ne Boe }9)
3 27-29/2/52 a | aloe 152 | 91-0 69 74 143 94-1 1:1-07
4 29/2-3/3/52 6 506 501 99-0 177 217 393 78-4 |1:1-22
5 3-4/3/52 10/3/52 86 Cee teil 2 2 4 57-1 1:1-00
6 4-7/3/52 pp 465 308 66-2 105 132 237 74-5 1:1-35
7 7-10/3/52 0 655 315 48-1 | 118 108 221 70-1 13 (0)2(8)55
8 10-17/3/52 | 17/3/52 425 201 | 47-3 60 105 165 70) }} al abo7a)

| |

Total .. sf ne SM ..| 2482 1499 | 60-4 546 672 1218 81-2 131-23

tubes were sealed with cotton plugs. Egg laying commenced and continued for 5 to 14
days without further blood feeding. From each family of progenies, sib mass matings
were made, and females isolated as before. On the assumption that females copulate
repeatedly, each female initially isolated would carry a sample of the sperm produced
by the available males. Should any initially isolated female be heterozygous for an
uncommon recessive gene ad, 50% of its progeny would be Aad. In the subsequent sib
mass mating, 25% of the sperm would be a. If three females are isolated from the sib
mating there would be an 824% chance that at least one would be Aad, and its progeny
should show 123% of homozygous recessives.

The results of this program are given in Tables 7A and 7B.
in the isolation of recessive genes.

BY S. SMITH-WHITE AND A. R. WOODHILL.

169

No success was achieved

Possibly the laboratory stock had lost genetic hetero-

geneity over four years and perhaps 25 generations of artificial culture. Probably the
mutant characters likely to be most frequent would be inconspicuous to casual inspection.
Nevertheless the results are of considerable interest.

TABLE 6.
F, Segregation for White Scales on Midfemur.
“ White.” | ** Intermed.” “ Black.” Total.
Females .. | 52 90 51 | 193
Males | 33 91 48 172
|

| |

|
Total 85 181 | 99 | 365

|

Total data: Fit to1:2:1 ratio. 7?=1-08. 2df. P=0-6.

Independence of sex linkage: /?=5-21. 5d.f. P>0-3.

Reference to Table 2 shows that, under the artificial breeding conditions of the
laboratory, there is a considerable difference in fecundity between S and K females.

There is no significant difference between self and cross matings,
fecundity of the K females appears to be dominant in the F,.

TABLE 7A.
Egg Laying of Isolated Females.

and the lower

Individual
Female.

52/11-
52/11-
52/11-
52/11-
52/11-
52/11-
52/11-
52/11-
52/11-
52/11-
52/11-
52/11-
52/11-
52/11-
52/11-
52/11-
52/11-
52/11-
52/11-
52/11-
52/11 -2¢
52/11-

De

BPE RE RP Rr ODN DD OP Ww
OR WH ©

Ne RPee
Seo nMDND

mw bv
wnwmre

| Means for Inbred Daughters.
|
Eggs Laid. | Hatch. Hatch
| Percentage. Percentage
| Eggs Laid. Hatch.
24 20 | 83-3 38°5 31-6
58 34 58-6 24-7 37-4
50 36 | 72-6 QZ, 59-5
29 19 65-5 117/283 33°3
85 59 69-4 29-7 39-3
96 | 50 | 52-1 32-0 8-7
56 | 34 H 60-7 46-2 33-0
50 | 30 60-0 26:8 66-7
30 19 63-3 1-0 0-0
35 30 | 85-7 19-0 67-5
29 | 24 82-7 56-6 54-1
32 26 | 81-2 38-5 51-9
80 60 | 75-0 13°33 22-6
42 37 88-1 18-5 5Y5)°'7/
11 0 0-0 = =
19 17 89-5 = =
31 | 0 0-0 = _
14 | 14 100-0 26-0 PRO
53 43 81-1 29-6 40-6
48 | 42 87:5 39-0 BZ iat
19 : 0 0-0 = =
139 131 94-2 16-3 39-1
1030 | 725 70-4 — 41-7
46-8 — = 24-7 =e

170 NON-RECIPROCAL FERTILITY IN AEDES SCUTELLARIS AND OTHER MOSQUITOES,

TABLE 7B.

Egg Laying of Inbred Females.

Eggs Hatch Eggs Hatch
Individual. Laid. Hatch. % Individual. Laid. Hatch. %G
52/11- 1-1 42 20 : 3 53 12
2 32 3 M. 56-0 — 54-1
3 35 20 52/11-13- 1 70 37
4 49 7 2 7 3
M. 39-5 — 31:6 M. 38-5 = 51:9
Splilo DES at iLL W 52/11-14- 1 . 0 _
2 25 20 2 6 1
3 4 0 3 26 17
4 59 10 4 38 0
M. 24-7 —_ 37-4 5 22 3
52/11- 3-1 45 30 6 0 aut
2 39 PC 7 1 0
3 7 0 M. 13-3 — 22-6
4 3 2 52/11-15- 1 17 17
5 61 37 2 41 0
6 8 1 3 1 0
M. UO —_— 59-5 4 0 =
52/11 4— i 23 8 5 0 Lee
2 0 0 6 21 21
3 17 4 7 0 —
4 29 11 8 1 0
M. W3 — 33°38 9 1 0
§2/11- 5-1 51 21 10 41 31
2 22 8 11 71 39
3 16 6 M. 18-5 = 59°7
M. 29-7 —— 39-3 52/11-19- 1 0 =
52/11- 6- 1 59 0 2 52 12
2 1 0 M. 26-0 = Besos
3 2 0 52/11-20- 1 0 —
4 1 0 2 39 29
5 97 14 3 89 25
M. 32-0 — 8-7 4 10 2
5§2/11- 7-1 40 0 5 0 —
2 44 0 M. 29-6 aS 40-6
3 43 38 52/11 -21— 1 59 35
4 33 26 2 58 26
5 53 0 3 0
6 34 7 M. 39-0 = o2-1
a 76 39 52/11 -23- 1 0 —
8 47 12 2 29 6
M. 416-3 — 54-1 3 0 —
52/11: 8- 1 58 48 4 28 19
2 0 — 5 49 24
3 0 — 6 49 5
4 38 Lost a 43 16
| 38 14 8 3 0
M. 6 8 — 66-7 9 10 5
§2/11- 9- 1 c =~ 10 9 4
2 il 0 11 16 12
3 33 0 12 18 9
4 0 — 13 2 0
M. 1:0 — 0-0 14 0 —_
52/11-10- 1 46 37 15 0 =
2 il 0 16 5 2
3 0 — M. 16-3 — 39-1
4 49 40
5 0 a SSS
6 18 0
M. 19-0 — 67-5 Total Ss 2422 1010 41-7
52/11-11- 1 | 62 35 General mean 24-7 — —
Dees 53 44

BY S. SMITH-WHITE AND A. R. WOODHILL. 171

Isolated females of race S show considerable variation in fecundity, but the mean,
46-8 eggs per female, is comparable with the figure obtained for S females in mass
culture (Table 74). The mean hatching, as a percentage of total eggs, was 70-4% in
the isolations, and 71-1% in the mass culture. Among the inbred daughters, many failed
to oviposit, and the means, eggs per female (24:7) and hatching (41:7%), were much
lower. The figures suggest a marked degree of inbreeding depression.

The time of action of lethality. In a considerable proportion of eggs derived from
K x S and B x S matings, sperm heads have been seen in the egg cytoplasm if the eggs
are crushed in aceto-orcein within half an hour of oviposition. Similar observation of
6-hour eggs indicates that some at least undergo early embryonic development.
Inviability is not due to any failure of sperm to penetrate the eggs, but rather to an
incompatibility between the sperm, or the hybrid embryo, and the egg cytoplasm.

In viable eggs of Aédes scutellaris, and in S x K hybrids, development is very rapid,
and an almost fully developed embryo is produced in 48 hours. In the inviable K x S
eggs, death ensues at an early stage, and after 24 hours they show a definite collapse.
Lethality is effective either before or during the cleavage divisions, or in the early
blastoderm—i.e., in the stage 1 of Hadorn’s (1948) classification of Drosophila lethals.
In contrast, lethality in Laven’s Culex molestus hybrids (Laven, 1953) is effective much
later, during late embryonic stages, or during or shortly after hatching. The Culex
molestus lethality is also characterized by a much lower penetrance.

DISCUSSION.

It is neither desirable nor possible to attempt an adequate discussion of the data
presented in the present paper without a consideration of the main features of non-
reciprocal fertility in other groups of the Culicidae. In particular, the results obtained
by Laven (l.c.) in Culex and by us in Aédes show such a close parallelism that they
must be dependent on similar genetic mechanisms.

1. Possible genetic mechanisms.

At the present stage, several possible genetic mechanisms must be considered, if
only for the summary rejection of some. Parthenogenesis and pseudogamy can be
dismissed on the basis of the F, segregations given in Table 6. Laven (1953) and
Toumanoff (1950) also give evidence eliminating parthenogenesis, but Downs and Baker
(1949), Bonnet (1950), and Perry (1950) find some support for its assumption.

Hypotheses of predetermination, of the Limnaea type, are incompetent to explain
the behaviour. A possible system of multiple incompatibility (s) genes affecting the
survival of S or of hybrid sperm in K egg cytoplasm, suggested by Smith-White (1950),
is denied by Laven’s and our own results. Such a system should give a uniformly
increasing rate of breakdown in the later backcross generations, the actual rate
depending on the number of s genes involved, and on their linkage relationships. In
our material, breakdown is absent or extremely rare, and in Laven’s there was a constant
rate of breakdown up to the 11th backcross generation. Both cases are characterized by
their permanence, and neither is affected by the building up of K or O* genoms, respec-
tively, in the backcrosses.

In a female-heterozygous sex system, a sex-linked incompatibility-lethal could
explain both the Culex and Aédes systems, with breakdown due to crossing over between
the lethal and sex genes. However, Gilchrist and Haldane (1947) have demonstrated a
male-heterozygous sex-system in Culex molestus, and this type of sex determination is
characteristic of the Nematocera (White, 1949).

Two kinds of genetic mechanisms remain. One is chromosomal, and is dependent
on the uniparental inheritance of chromosomes or of chromosome segments, and involves
anomalies of the meiotic cycle. The other is nucleus-independent and cytoplasmic.

Very peculiar meiotic cycles are known to exist in other families of the Nematocera.
In Sciara the chromosomes of paternal origin are eliminated during spermatogenesis

*O is the symbol used by Laven for his Oggelshausen strain of Culex molestus.
M

(2 NON-RECIPROCAL FERTILITY IN AEDES SCUTELLARIS AND OTHER MOSQUITOES,

(Metz, 1938). In Miastor and other genera of the Cecidomyidae, many chromosomes are
eliminated in spermatogenesis, and are transmitted only in the female line. In the
Culicidae, limited chromosomes are not present. In Culex (Callan and Montalenti,
1947) and in Aédes chiasmata are formed in all three bivalents in male meiosis.
Moreover, in Aédes scutellaris, the approximately normal sex ratio, and the sex
independent segregation of “line” provide genetic evidence for the normal meiotice
separation of two of the three chromosome bivalents.

0 Ole hs 0 of

DO or aka parents IS? CRAB Ke
biK)? +(S)° gametes +(S)° biKI?
2b Ey progeny +bF,-
die [sree
| 8 Y arents & 6
HIRI? oe EVE, parents 7S ea Geen bk
bik)’ a gametes +(F) bik)

biF)
+bB,"
Progeny +bB,
bbB” * j |
die pee

BOK xX ebBe . Dareniis +bBy X bbK®

bik)° gametes +(By bik)?
8)
+bB, F
Roe Z progeny +bB,
2
die survive
Etc. Etc

Text-figure 2.—Gene control of non-reciprocal fertility, dependent on the elimination of

a paternal chromosome segment in oogenesis.
The symbols kK, S, Fy, Bi, ete., refer to the wilimited gametic genoms. b and + are alleles
which condition the cytoplasm, and are carried on limited (maternally-inherited) chromosome

segments.
Note.—Sperm b(F,)® from a males are unadapted and hybrids derived from unadapted

sperm and b (kx) 8 eges die.

It is possible to devise chromosomal systems of permanent non-reciprocal fertility
on the basis of two assumptions. There must be a strictly polarized segregation of a
chromosome segment in oogenesis, with the elimination of the segment of paternal
origin in hybrids. It is not necessary to assume any similar elimination in spermato-
genesis, and the sperm wastage which would result from such behaviour is not evidenced.
In oogenesis, the paternal segment must be directed into the polar bodies. The second
necessary assumption is the existence of a cytoplasm-conditioning gene, carried on the
polarized chromatin, in one or other of the two species or races involved. An example of
this type of hypothesis is offered in Text-figure 2. Race K is there considered to be
homozygous for the cytoplasm-conditioning gene “bar” or “b’, carried on polarized

BY S. SMITH-WHITE AND A. R. WOODHILL. 173

ehromatin, which modifies the cytoplasm from a neutral « condition to a 6 condition,
intolerant of other than b-adapted sperm or hybrid embryos. Race § is inferred to be
homozygous for a neutral or + allele at the same locus. In this scheme, + must be
dominant to 0b, and there must be a lag of several cell generations in cytoplasmic
adaptation following a change of genotype. Consequently b sperm from b+ males are
initially unadapted.

The assumption of lag in the conditioning of the cytoplasm is necessary to allow
partial embryonic development. Exceptional breakdown is permitted by a slight
variability in the penetrance of the lethal effect following the fertilization of b(K)*/ eggs
by b(F,)¢ or similar unadapted sperm, thus allowing cytoplasmic adaptation in the
bb(B,) embryos. .

There is no cytological or genetical evidence to support the inference of meiotic
polarization which hypotheses of this type require, but if this inference is rejected, an
explanation based .on cytoplasmic inheritance independent of the genotype must be
invoked.

Cytoplasmic inheritance is known in many plants (cf. Caspari, 1948) and in some
animals. Laven (1953) has already drawn attention to the similarities between the
known cases of plasmagenic or genoid inheritance in Paramoecium and in Drosophila
and the inheritance of non-reciprocal fertility in mosquitoes, and it is unnecessary to
elaborate this comparison further. A plasmagenic hypothesis can be set up in simpler
form than the chromosomal scheme given in Text-figure 2, but is in fact very similar in
operation. It is only necessary to regard the cytoplasmic conditions @ and 6 as being
independent and _ self-perpetuating. Breakdown could occur either by a variable
penetrance of the lethal effect of the 6 plasmagene, or by the possibility that sperm could
occasionally carry a sufficient dosage of 6 to enable them to survive in 6 egg cytoplasm.

At present there is no evidence enabling a choice to be made between hypotheses
of uniparental inheritance of cytoplasm-conditioning genes and cytoplasmic inheritance.

2. The role of non-reciprocal fertility in speciation.

On present knowledge, the S and K races of Aédes scutellaris are geographically
isolated, and they are morphologically distinguishable. It is probable that the genetic
basis of their non-reciprocal fertility was able to develop because of their geographic
isolation. In Laven’s races of Culex molestus a geographical factor is also apparent.

In crosses between Aédes aegypti and A. albopictus, various authors have obtained
conflicting results. Downs and Baker (1949) and Bonnet (1950) found that fertile
hybrids could be obtained only by using A. aeyypti females. The reciprocal mating gave
effective insemination and egg laying, but the eggs were inviable. Both authors suggested
parthenogenesis as a possible explanation. Toumanoff (1939, 1950) obtained exactly the
opposite results, a fertile F, being obtained only when A. albopictus was the maternal
parent. Toumanoff’s strains of the two species were obtained independently of those of
Downs and Baker and Bonnet. Simmons, Saint John and Reynolds (1930) were unable
to cross the same two species in either direction, and were unable to obtain egg laying
in their matings. DeBuck (1942) obtained normal egg production in matings between
the two species in both directions, but the reciprocal F,s were both inviable. He found
that with A. albopictus as maternal parent hybrid eggs showed no embryonic develop-
ment, but in the reciprocal cross some embryonic development was usual. His results
indicate a reciprocal difference in the same direction as those of Downs and Baker and
Bonnet, but with a higher degree of incompatibility.

A. aegypti and A. albopictus must include geographical races differing in mating
type. It is suggested that the degree of isolation indicated in DeBuck’s results is
causally related to the partial isolation reported by Downs and Baker, and to the
absolute isolation found by Simmons et al. Partial, non-reciprocal isolation may
represent a stage in the origin of complete isolation, and may constitute a significant
factor in speciation.

174 NON-RECIPROCAL FERTILITY IN AEDES SCUTELLARIS AND OTHER MOSQUITOES,

In crosses between Aédes hebrideus and A. pernotatus Perry (1950) found that
viable eggs were only obtained when A. hebrideus was the maternal parent, but that only
a small proportion of the females of that species (3 out of 37) yielded viable progeny.
Hovanitz (1946) found considerable variation in the fertility of individual females of
Haemogogus sp. when mated to their own males, and we find a similar variability in
isolated females of A. scutellaris. It is possible that a species-population may be hetero-
geneous for sterility factors. Fixation of mating type in geographical isolates would
then be expected, and the conflict between the results of the various authors could be
understood.

In the Culex pipiens species complex an apparently confused picture of relationships
comes from an analysis of the available interfertility data. Marshall (1938) found that
races of C. molestus from Paris and London which were reciprocally intersterile, both

c
ATIGAN?

Text-figure 3.—Race relationships in the Culex pipiens complex.
Key:

<—— Full reciprocal compatibility.

—— Partial reciprocal compatibility.

Sie’

—— Non-reciprocal compatibility.
————Complete incompatibility.
F, Farid, 1949; L, Laven, 1951, 1953; M, Marshall, 1938; R, Roubaud, 1941; TV, Tate and
Vincent, 1936; W, Weyer, 1936.

gave viable eggs and fertile F, progeny when mated to males of a strain from Hayling
Island, and inviable eggs in the reciprocal crosses. Laven (l.c.) found that various
isolations of the same species could be grouped on a geographical basis. Isolations from
north-eastern Germany and western Hurope were intersterile in both directions, and
both were fertile with isolations from southern Germany only when the latter strains
were used as male parents. It is noteworthy that Laven’s isolations from London and
Paris were fully interfertile, whereas Marshall’s strains from the same localities were
reciprocally intersterile.

Tate and Vincent (1936) were able to cross C. molestus and C. pipiens in both
directions. Weyer (1936) found full fertility between C. molestus and C. fatigans but
was unable to cross C0. pipiens and C. fatigans. Farid (1949), however, found the last
two species to be fully and reciprocally interfertile, and Roubaud (1941) working with

oO

BY S. SMITH-WHITE AND A. R. WOODHILL. 1

the same species obtained results suggestive of a reciprocal difference (fatigans « pipiens,
300 eggs, 55 larvae; pipiens x fatigans, 850 eggs, 2 larvae).

It is again apparent that races in the ©. pipiens complex differ in factors affecting
interracial fertility, and that these differences have a geographical basis. However,
interracial fertility barriers are not conformable with the recognized species limits in
the group, and it would seem unlikely that they represent stages in speciation parallel
with those involved in the origin of ©. pipiens, C. molestus and C. fatigans. We are
inclined to agree with Mattingly (1951) that the whole group shoulc be recognized as a
single species, with many geographical subspecies and locality races.

The role of non-reciprocal fertility in the origin of total isolation and speciation is
problematical, but it is of particular interest as a possible evolutionary mechanism.
Perhaps intrinsic isolation and speciation may arise, on occasion, firstly in the cytoplasm,
and secondarily in the nuclear genom.

SUMMARY.

A study has been made of the inheritance of non-reciprocal fertility in two subspecies
of Aédes scutellaris, A. s. scutellaris and A. Ss. katherinensis.

Mating type shows a strict maternal inheritance. F, eggs from subsp. scutellaris (S)
females and subsp. katherinensis (K) males are normally viable, and are of:S mating
type. Backcrosses of F, females to K males are viable, and again, are of S mating type.
In successive backcrosses to K males, to the B, generation, the S mating type is retained.
All crosses involving K females yield inviable eggs, and it is not possible to test the
mode of inheritance of the K mating type.

The results obtained are essentially similar to those reported by Laven for mating
type inheritance in Culex, but the lethal effect in inviable eggs is earlier in operation,
and more severe, in Aédes than in Culex.

Meiosis in the male is apparently normal. The genetic system determining the
inheritance of mating type must depend either on anomalous meiosis in oogenesis, with
a polarized segregation of bivalents and the elimination of part of the paternal genom,
or on nucleus-independent cytoplasmic factors. There is no critical evidence enabling
a choice between the two types of hypothesis. The observed entry of sperm into eggs in
the inviable matings, partial embryonic development in the inviable Culex matings, and
F, segregation in viable S female x K male matings, allow the rejection of hypotheses
of parthenogenesis.

From a consideration of the available data on mating isolation between species and
races of Aédes and Culex, the type of non-reciprocal fertility described is considered to
be significant as a source of incipient speciation in mosquitoes.

Literature Cited.

BONNET, D. A., 1950.—The Hybridisation of Aédes aegypti Linn. and Aédes albopictus Skuse
in the Territory of Hawaii. Proc. Hawaiian Ent. Soc., 14: 35-39.

CASPARI, B., 1948.—Cytoplasmic Inheritance. Adv. in Genetics. Vol. 2, pp. 1-66. Ed. M. Demerec.
Academic Press Inc., N. York.

DeBuck, A., 1942.—Kreusungversuche mit Stegomyia fasciatus Fabricius und S. albopictus
Skuse. S. angew. Ent., 29: 309-312.

DoBROoTWORSKY, N. V., and DRUMMOND, F. H., 1953.—The Culex pipiens Group in South-Eastern
Australia. II. Proc. Linn. Soc. N.S.W., 78: 131-146.

Farip, M. A., 1949.—Relationships between Certain Populations of Culex pipiens L. and Culex
quinquefasciatus Say. in the United States. Amer. J. Hyg., 49: 83-100.

GILCHRIST, B. M., and HALDANE, J. B. S., 1947.—Sex Linkage and Sex Determination in a
Mosquito, Culex molestus. Hereditas, 33:175-190.

Hapborn, E., 1948.—Gene Action in Growth and Differentiation of Lethal Mutants of Drosophila.
In Growth Symposium, Soc. Exp. Biol., pp. 177-195. University Press, Cambridge.

HOovANITz, W., 1946.—Comparisons of Mating Behaviour, Growth Rate, and Factors Influencing
Egg Hatching in South American Haemogogus Mosquitoes. Physiological Zoology, 19: 35-53.

LAVEN, H., 1951.—Crossing Experiments with Culex Strains. Hvolution, 5: 370-375.

, 1953.—Reziprok Unterschiedliche krauzbarkeit von Stechmucken (Culicidae) und

ihre Deutung als Plasmatische vererbung.

MARSHALL, J. F., 1938.—The British Mosquitoes. British Musewm, London.

176 NON-RECIPROGAL FERTILITY IN AEDES SCUTELLARIS AND OTHER MOSQUITOES.

MAarTinGLy, P. G., 1951.—The Culex pipiens complex. Introduction. Vrans. Roy. Ent. Soc.
London, 102: 331-341.

Merz, C. W., 1938.—Chromosome Behaviour, Inheritance and Sex Determination in Sciara.
Amer. Nat., 72: 485-520.

Perry, W. J., 1950.—Biological and Crossbreeding Studies on Aédes hebrideus and Aédes
pernotatus. Ann. Ent. Soc. Amer., 43:123-136. i

RouBAuD, E., 1941.—Phenoménes e’amixie dans les intercroisements de Culicides du groupe
pipiens. CO. R. Acad. Sci. Paris, 212: 257-259.

Simmons, J. S., SAINT JOHN, J., REYNOLDS, —., 1930.—Transmission of Dengue Fever by Aédes
albopictus Skuse. Philippines J. Se., 41 (38) : 215-219.

SMITH-WHITE, S., 1950.—A note on non-reciprocal fertility in matings between subspecies
of mosquitoes. Proc. LINN. Soc. N.S.W., 75: 279-281.

SPENCER, W. P., 1947.—Mutations in wild populations of Drosophila. Advance in Genetics. Vol. 1,
pp. 359-402. Ed. M. Demerec. Academic Press Inc., N. York.

TATE, P., and VINCENT, M., 1936.—The Biology of Autogenous and Anautogenous Races of
Culex pipiens L. Parasitology, 28:115-145.

TOUMANOFF, C., 1939.—Les races géographiques de Stegomyia fasciata et St. albopictus et leur
intercroisements. Bul. Soc. Path. exot., 32: 505-509.

, 1950.—L’intercroisement de VAédes (Stegomyia) aegypti L. et Aédes (Stegomyia)
albopictus Skuse. Bull. Soc. Path. exot., 43: 234-240.

WeEYER, F., 1936.—Kreuzungversuche bei Stechmucken (Culex pipiens und Culex fatigans).
Arb. phys. angew. Ent. Berlin-Dahlem, 3: 202-207.

WHITE, M. J. D., 1949.—Cytological evidence on the phylogeny and classification of the Diptera.
Evolution, 3: 253-261.

WoopHILL, A. R., 1949.—A note on experimental crossing of Aédes (Stegomyia) scutellaris
sceutellaris Walker, and Aédes (Stegomyia) scutellaris katherinensis Woodhill. PROC.
LINN. Soc. N.S.W., 74: 224-226. i

, 1950.—Further notes on experimental crossings within the Aédes scutellaris group
of species (Diptera, Culicidae). Proc. LINN. Soc. N.S.W., 75: 251-253.

ILA

A NEW COPTOTERMES AND AHAMITERMES (ISOPTERA) FROM AUSTRALIA.
lsh 105 dio Geatce
(Communicated by Dr. A. J. Nicholson.)
(One Text-figure.)

[Read 29th September, 1954.]

Synopsis.
A new mound-building species of Coptotermes with which is associated a new species of
Ahamitermes has been collected in Western Australia. Descriptions of these two species are
given together with brief notes on their biology.

INTRODUCTION.

During the past two years officers of the Wildlife Survey Section of the
Commonwealth Scientific and Industrial Research Organization have carried out a
number of extensive survey trips in Western Australia. As a spare-time activity, one
of the officers, Mr. J. H. Calaby, has given special attention to the termite fauna and,
as a result, has collected some hundreds of series that have yielded considerable
information on the range of distribution of most of the species recorded from this
State by Hill (1942), established the presence of several species not previously known
to occur in Western Australia, and have included some hitherto undescribed species.

The termites described below were collected during a recent survey trip north of
Geraldton, and I am indebted to Mr. Calaby both for this interesting material and for
the excellent and detailed field notes that accompanied the collection.

It may seem a little strange that such a conspicuous and distinctive species as the
new Coptotermes described below.should not have been recorded previously, but this
may be due to its restriction to a relatively small area in a sparsely settled and
infrequently travelled region.

The association of a new species of Ahamitermes with the new species of
Coptotermes is not altogether unexpected, since this association between the two genera
occurs commonly on both sides of the continent. Indeed the specificity of this association
is such that it would have been a little surprising if the Ahamitermes found in the
mounds of such a distinctive species of Coptotermes had not been new.

The drawings of the soldier and alate heads were made with the aid of a camera
lucida.

COPTOTERMES BRUNNEUS, Sp. Nov.
Winged Adult (Fig. 1, A).

Medium sized, head, thorax, and tergites of abdomen very dark brown (almost
black), sternites of abdomen somewhat paler. Legs of varying shades of brown, the
distal portions of the tibiae and the tarsi yellowish. Basal segment of the antenna the
same colour as the head, the remaining segments, palps, postclypeus and labrum,
golden, brown variably suffused with darker brown. Wing stumps dark brown, the
basal portion of the media and cubitus and the wing membrane between the radial
sector and costal border brown, the remainder of the wing membrane pale fuscous.
Head, thorax, wing stumps, and abdomen hairy. Head narrowed in front of the eyes,
hemispherical behind. Antennae of 20 segments, third segment generally shortest and
narrowest of all; occasionally fourth segment shortest of all. Posteclypeus more than
three times as wide as long. Eyes moderately large; ocelli 0:10 to 0:14 mm. long and
separated by about half their length from the eyes. Fontanelle small and indistinct.

* Division of Entomology, C.S.1I.R.O., Canberra, A.C.T.

178 NEW COPTOTERMES AND ATLAMITERMES FROM AUSTRALIA,

Pronotum somewhat narrower than head and conspicuously wider than long, anterior
margin widely and shallowly concave, sides but little rounded, the postero-lateral
corners almost straight and the posterior margin slightly sinuate. Anterior wing stump
very large, almost twice as long as that of hindwing. Costal border, radial sector and
basal sections of media and cubitus well defined, the remainder of the veins weakly
developed. Wing membrane without hairs, densely covered with micrasters.

Text-figure 1.

A, Coptotermes brununeus, n. sp. Head and pronotum of winged adult. B, C. brwinews, n. sp.
Head and pronotum of soldier. C, Ahamitermes inclusus, n. sp. Head and pronotum of queen.
D, A. inclusus, n. sp. Head and pronotum of soldier.

(Camera lucida drawings by Miss B. J. Gemmell.) (A, B, x 10; C, D, x 16.)

Measurements (50 specimens).—Length, with wings, 10-50-13:00 mm.; length, with-
out wings, 6:20-8:00 mm.; head, to apex of labrum, long, 1:59-1:81 mm.; head, to
clypeofrontal suture, long, 1:08-1:21 mm.; head, wide, 1:41-1:54 mm.; eyes, maximum
diameter, 0:34-0:40 mm.; ocelli, long, 0:10-0:14 mm.; pronotum, long, 0:77-0:84 mm.;
pronotum, wide, 1:28-1:41 mm.; forewing, long, 8-20-8-90 mm.; forewing, wide,
2:20-2:50 mm.

Soldier (Fig. 1, B).

Large, somewhat variable in size, with a very large head. Head and pronotum
raw umber, meso- and metanotum progressively paler; abdominal tergites about the
same colour or a little paler than the metanotum, a pale median line extending from
the front of the pronotum to the second or third abdominal tergite. Femora somewhat
fuscous, tibiae pale brown. Head darkest anteriorly, basal segment of antenna the same
colour as the anterior portion of the head. second segment pale golden yellow, the
remaining segments as well as those of the maxillary palps variably suffused with
brown. lLabrum bright golden yellow, the central area suffused with brown, with a

ea ds Ao 179

hyaline tip. Mandibles dark ferruginous. Head and thorax with a moderate number
of stout reddish hairs, abdomen very hairy. Head longer than wide, very little rounded
on the sides, narrowing evenly to the anterior margin, generally widest at the posterior
third and broadly rounded behind. Antennae of 17-18 segments, third segment generally
shortest of all, occasionally third and fourth about equal in size and shorter than
remaining segments. Fontanelle of moderate size, distinct. Labrum longer than wide,
widest at basal third, tapering to the narrow, rounded, hyaline tip. Pronotum con-
spicuously narrower than the head, distinctly wider than long, the anterior margin
with a broad median concavity, the posterior margin widely and shallowly concave.
Abdomen without dark dorsal pattern.

Measurements (50 specimens) .—Total length, 5:50-7:50 mm.; head, with mandibles,
long, 2:75-3:30 mm.; head, without mandibles, long, 1:90-2:16 mm.; head, to fontanelle,
long, 1:72-1:96 mm.; head, maximum width, 1:56-1:76 mm.; head, minimum width,
1:02-1:17 mm.; gula, minimum width, 0-26—-0-42 mm.; fontanelle, wide, 0:14-0:20 mm.;
pronotum, wide, 1:08-1:32 mm.; pronotum, long, 0:57-0:68 mm.

Distribution.—Western Australia, 18 miles NNW of Galena,* 25.X.1953, J. H. Calaby,
workers, soldiers, and alates; 23 miles NNW of Galena, 25.X.1953, J. H. Calaby, workers
and soldiers; 51 miles NNW of Galena, 25.X.1953, J. H. Calaby, workers soldiers and
alates (type series); 45 miles NNW of Galena, 25.X.1953, J. H. Calaby, workers, soldiers
and alates; 59 miles NNW of Galena, 24.X.1953, J. H. Calaby, workers, soldiers and
reproductive nymphs; 66 miles NNW of Galena, 24.X.1953, J. H. Calaby, workers and
soldiers; 67 miles NNW of Galena, 24.X.1953, J. H. Calaby, workers and soldiers.

Biology.

This species is a mound builder and, so far, is Known only from a relatively small
area just north of the Murchison River in the direction of Shark Bay. The mounds
occur in sclerophyll woodland and mallee scrub, but not in mulga scrub, and are
generally situated close to a eucalypt tree or clump of mallee. They have a thick clay
outer wall, variously coloured red-brown, grey-brown, or yellow-brown, depending upon
the colour of the surrounding soil, and are present on the yellowish-brown and greyish-
brown soils, and less commonly on the red sandy soils, but not on the stony red soils
towards the Murchison River.

The average size of the mounds is about 5 feet high and 3-4 feet diameter at
ground level, whilst the largest was 8 feet high, 5 feet in diameter at ground level, and
extended below ground level to a depth of 4 feet. An unusual feature is the presence of
small mounds 6-9 inches in diameter and 2-6 inches high. Mounds of this small size
relative to the average size have not been recorded in any of the other Australian
species of Coptotermes.

The mound consists of two distinct regions, (i) an inner nest structure of rather
loose, coarse, woody material, honeycombed with galleries but without any “nursery”
area of thin-walled concentric cells, and (ii) a thick outer wall of clay. This outer
wall is generally separated from the woody nest region by a distinct gap of 1-3 inches,
and is traversed by only a few very large trunk galleries (up to 3 inches in diameter)
that lead up towards the summit of the mound, where there is an area of small flattish
galleries close to the surface. A few of these small flattish sub-surface galleries are
also found here and there on the sides of the mound. Occasionally the interior of the
mound gives the impression of a large cavity between the upper surface of the nest
region and the outer wall. In such instances there are no trunk galleries in the outer
wall itself, and this cavity is, in fact, a single very large trunk gallery communicating
with the summit of the mound. Alates concentrate in the flattish galleries referred to
above during the heat of the day, but appear to retreat to the central nest region at
other times.

Many mounds have a zone of extremely wet clay localized within the upper portion
of the outer wall. This zone, which may be a foot in diameter, is linked to the nest

* Galena is a small lead-mining town about a mile south of the Murchison River, on the
north-west coastal highway.

LSO NEW COPTOTERMES AND AHAMITERMES FROM AUSTRALIA,

region by the large trunk galleries, and may serve as a source of humidity or as a
reserve of readily available building material to re-seal the outer wall after the release
of alates.

Winged adults are present in the mounds in October.

Affinities.

The winged adult is much darker than either C. dreghorni Hill or C. lacteus
(Froggatt); in addition it has one more sezment in the antenna and the wing mem-
brane is devoid of hairs. The characteristic colour and large size of the soldier head
serve to distinguish it from all other Australian species of Coptotermes.

Types.—Holotype winged adult female and morphotype soldier and worker in the
Division of Entomology Museum, C.S.1I.R.O., Canberra.

AHAMITERMES INCLUSUS, Sp. Nov.
Winged Adult.

Not known.

Queen (Fig. 1, C).

Head and thorax golden brown, tergites of abdomen somewhat paler, sternites of
abdomen, antennae and labrum light yellowish-hrown. Head, thorax, and tergites and
sternites of abdomen densely pubescent. Postclypeus more than half as long as wide,
the anterior margin straight, the posterior margin strongly convex. Fontanelle
lanceolate, with indications of anterior prolongation. Eyes moderately large and
prominent. Ocelli oval, separated by about half their long diameter from the eyes.
Antennae of 15 segments, third segment shortest and narrowest of all. Pronotum a/
little wider than the head, somewhat wider than long, the anterior margin widely and
deeply concave, the sides somewhat rounded at first and then narrowed evenly to the
shallowly concave posterior margin. Posterior margins of meso- and metanotum widely
and deeply notched. Tibial spurs 3:2:2.

Measurements.—Head to apex of labrum, long 1:06 mm.; head, to clypeofrontal
suture, long, 0-55 mm.; head, wide, 0-88 mm.; eyes, maximum diameter, 0:28 mm.;
ecelli, long, 0:09 mm.; pronotum, long, 0-82 mm.; pronotum, wide, 0:93 mm.; total length,
12-50 mm.; width of abdomen, 3-75 mm.

Soldier (Fig. 1, D).

Head light orange, mid-dorsal area paler, the entire head capsule occasionally
lightly suffused with brown. Basal third of mandibles light orange, distal two-thirds
pale ferruginous. Head almost parallel-sided or with sides very slightly concave.
Fontanelle distinct. Labrum a little wider than long, bluntly conical, the hyaline tip
usually with an obvious concavity. Mandibles relatively short and stout, about two-
thirds as long as the head capsule, generally with vestiges of teeth about the middle.
Post-clypeus strongly bilobed. Antennae short, extending for only about one-fifth of
their length beyond the tips of the mandibles, of 13 segments, third segment smallest
of all. Pronotum about twice as wide as long, a little narrower than the head, the
anterior margin convex, the posterior margin shallowly concave. Tibial spurs 3:2:2.

Measurements (30 specimens) .—Total length, 4:00-5:00 mm.; head, with mandibles,
long, 1:70-1:83 mm.; head, to clypeofrontal suture, long, 1:02-1:10 mm.; head, wide,
0:66-0:77 mm.; pronotum, long, 0:31-0:35 mm.; pronotum, wide, 0:60-0:69 mm.

Distribution.—Western Australia, 51 miles NNW of Galena, 25.X.1953, J. H. Calaby,
workers, soldiers, reproductive nymphs, and queen (type series); 66 miles NNW of
Galena, 24.X.1953, J. H. Calaby, workers and soldiers.

Biology.

This species is known from two series only, both of which were taken from
termitaria of Coptotermes brunneus, sp. nov. One series was collected from a concen-
tration of cells in the outer clay wall of the Coptotermes mound, the other from a
definite nest structure about a foot in diameter situated on top of the cellular portion
of a Coptotermes nest. This second series consisted of a queen, reproductive nymphs,

ie) 18S dig GMNr, 181

workers, soldiers and eggs. The Ahamitermes nest, which was densely populated,
consisted of a mass of flattish dark-coloured cells, and from this central nest area
galleries extended throughout the Coptotermes mound, frequently running alongside
Coptotermes galleries but always clearly separated from them by a thin wall.

Affinities.

Ahanitermes inclusus is distinguishable from A. hillii Nicholls, also from Western
Australia, by the generally smaller size and much paler colour of the queen. In the
soldier caste it is separated from A. hillii by its smaller size, parallel-sided head, the
short antennae and the relatively short, stout mandibles.

Types.—Holotype soldier, morphotype dealated adult female and worker in Division
of Entomology Museum, Canberra.

Reference
HIuu, G. F., 1942.---“'t'ermites (Isoptera) from the Australian Region’. Melbourne, 479 pp.

NOTES AND DESCRIPTIONS OF AUSTRALIAN CHLOROPIDAE (DIPTERA).

By CurtTis W. SAsprosky, Entomology Research Branch, Agricultural Research
Service, United States Department of Agriculture.

(Communicated by Mr. D. J. Lee.)
(Five Text-figures.)
[Read 27th October, 1954.]

In the last three parts of his long series of papers entitled “Notes on Australian
Diptera’, J. R. Malloch presented an extensive revision of the family Chloropidae in
Australia and Tasmania (Proc. Linn. Soc. N.S.W., 63: 334-356, 1938; 65: 261-288, 1940;
66: 41-64, 1941). Citations to that basic revision will be abbreviated here to date and
page. The following notes will clarify a few problems, as well as make known a few
new forms. Great as was the number of new species described by Malloch in this
family, I have no doubt that many more remain to be discovered in Australia, especially
in the western part of the continent. However, in these tiny flies the recognition of
new forms and their satisfactory separation from described forms will seldom be an
easy task, especially on the basis of published descriptions alone. In certain large
genera, in particular, one will almost surely have to have good series of specimens in at
least fairly good condition, except for unusually distinct species.

Subfamily CHLOROPINAE.
PACHYLOPHUS Loew.

Three species, all described by Malloch, have hitherto been known from Australia,
and a key to these was presented by Malloch (1927, Proc. LINN. Soc. N.S.W., 52: 428).
From relatively limited material seen thus far, I suspect that P. secundus Malloch is
only a dark colour form, or perhaps the fully matured form of P. luteus Malloch
(correction for lutea!). The third species, P. alienus Malloch, is very little different,
unless the stated formula of 1 + 1 for the notopleural bristles proves to be a consistent
feature.

The character of hairy mesopleura, used by Malloch for Pachylophus in his key to
the genera of Australian Chloropidae (1938, p. 334), is not characteristic of the genus
as a whole. The species P. rufescens, recorded here from Australia for the first time,
has the mesopleura bare, a feature which will distinguish it at once from the species
described by Malloch.

PACHYLOPHUS RUFESCENS (Meijere).

Myrmemorpha rufescens Meijere, 1904, Bijdr. Dierkunde, 17: 1138 (Java).
Pachylophus rufescens (Meijere) Becker, 1911, Ann. Mus. Nat. Hungarici, 9: 41.

I have before me eight specimens from Burpengary, Queensland (Dr. T. L. Bancroft,
seven labelled ‘5-12-99’), which appear to be this wide-ranging Oriental species. In
addition to the type series, I have seen the species from the Simla Hills in North
India, Coimbatore in South India, Calcutta, North-east Assam, Upper Burma, Tientsin,
China, and Sumatra, and there are published records also from Formosa and Ceylon.
The species is somewhat variable in colour, but generally the frontal triangle is entirely
black and the thorax reddish with indistinct stripes, the median one blackened on the
anterior slope of the mesonotum. The mesonotum and scutellum are brownish-grey
pollinose and somewhat dull, while the pleura are polished and entirely reddish. These
colour characters are quite different from those described by Malloch for the three
species already recorded from Australia, and will further distinguish P. rufescens.

a)

BY CURTIS W. SABROSKY. 18

Cutoroeps Meigen (= Oscinis Latreille).

There has been an unfortunate confusion in the use of the generic name Oscinis.
Oscinis of authors, especially of the last century, referred to the small, usually black
chloropids of the subfamily Oscinellinae, now called Oscinella, Oscinosoma, Conioscinella, |
etc. Even among those who recognize that Oscinis, in the strict sense, refers to certain
predominantly yellow and black striped chloropids of the typical subfamily Chloropinae,
some synonymize Oscinis with Chlorops and some maintain the two as distinct genera.
I follow the former course.

CHLOROPS SULCATA Becker.
Chlorops sulcata Becker, 1911, Ann. Mus. Nat. Hungarici, 9: 58 (New South Wales).
Oscinis federata Malloch (1938, pp. 338, 341) must be very close to Becker’s species,
from the descriptions of the two. I suggest that they are probably synonyms.

CHLOROPS ALBIFRONS Walker.

Chlorops albifrons Walker, 1849, List Dipterous Ins. British Mus., 4: 1121 (Adelaide,
South Australia). Oscinis canaliculata var. trisulcata Malloch, 1938, p. 345 (New South
Wales, Austral. Cap. Terr.). [New Synonymy. ]

I have seen the type of albifrons in the British Museum (Nat. Hist.), and it agrees
perfectly with Malloch’s description of trisulcata. Malloch’s two guesses as to the
identity of albifrons, either as a species with smooth ungrooved frontal triangle (1931,
Proc. LINN. Soc. N.S.W., 56: 73), or as equal to Chloromerus gracilis Malloch (cf. 1938,
p. 336),,are incorrect. Incidentally, this Australian form is quite a different species
from the Formosan Chlorops canaliculata Becker, of which I have seen the type series
in the Hungarian National Museum.

CHLOROPS AUSTRALIENSIS, Nl. Sp.

Yellow and black species with smooth frontal triangle, polished black mesonotal
stripes, black palpi, and predominantly black femora.

Male, female—Head yellow, frontal triangle dark reddish-brown to black, occiput
above with median black stripe of same width as triangle joining a transverse black
band between the eyes and above the foramen, palpus entirely black, median clypeal
plate brown on sides, dnd antenna reddish-yellow basally with distal three-fifths of
third segment and the arista black. Thorax with yellow ground colour, marked with
black as follows: three broad mesonotal stripes and two supra-alar vittulae, small spot
on each humerus, the metanotum, and large spots on meso-, ptero-, sterno-, and
hypopleuron, the upper half of pleuron otherwise rather suffused reddish; scutellum
brown to blackish on sides, sometimes leaving only a narrow median area and apex
yellow. Abdomen reddish-brown, fourth and fifth segments with distal yellow margins.
Legs with fore coxa laterally at base, all femora narrowly at base and apex, and hind
tibia centrally black; fore and mid tibiae obscurely brownish in some specimens, and
fore tarsus entirely and others distally somewhat browned. Halter with light brown
stalk and conspicuously whitish knob. Wings hyaline, veins brown, membrane brownish
tinted. Bristles and hairs black.

Head with front parallei-sided, approximately square, at vertex twice the width
of an eye and slightly more than half the width of the head, sparsely haired; ocellar
tubercle grey pruinose; frontal triangle polished and glabrous, at base barely more than
two-thirds the width of front and thus well separated from eyes, long and acuminate,
the sides slightly convex, converging strongly about two-thirds the length of triangle,
which then continues to anterior margin of front in a narrow tapering prolongation.
Head in profile with greatest length and height subequal, the front projecting slightly
before the eyes and face receding, greatest length of head 1:75 times the length of
lower margin of head; eye large and rounded, cheek one-fifth the height of an eye and
two-thirds the breadth of third antennal segment, the latter slightly longer than broad
and with concave upper margin. Arista short pubescent, base rather thick. Ocellar
bristles proclinate and divergent, much longer than postverticals, but shorter than
verticals; six pairs of orbital bristles, the upper two pairs longer than the others.

184 AUSTRALIAN CHLOROPIDAEF,

Mesonotum grey to brownish pollinose outside of the stripes, the lines of pollen
separating stripes and outlining them clearly, stripes polished but the shine interrupted
by numerous, coarse, piliferous punctures; metanotum heavily leaden-grey pollinose;
pleuron predominantly smooth and polished, grey pruinose on narrow postero-dorsal
corner of mesopleuron, the pteropleuron posteriorly and the hypopleuron. Notopleural
bristles 1 + 2, anterior somewhat weaker than others. Scutellum narrowly rounded
apically, not quite three-fourths as long as broad at base; apical scutellar bristles
longer than scutellum and inserted rather close together, with bases separated by a
distance barely greater than that between posterior ocelli; subapical pair rather short,
inserted as near to the apicals as the latter are from each other; disc of scutellum
moderately setose. Abdomen shining, thinly and finely pollinose. Legs slender, hind
tibia without sensory area. 2

Wing with second costal sector 1-7 times third sector; second vein almost straight
until its apical fifth or sixth, marginal cell relatively narrow, at its midpoint little
more than half the width of submarginal cell immediately opposite; third and fourth
veins widely divergent, the width of apical ceil measured between ends of third and
fourth veins 1:7 times its width opposite hind crossvein; ultimate section of fourth vein
a fine line, not arched, practically straight to margin of wing; penultimate section of
fourth vein 1-8 times length of penultimate section of third vein and equal to ultimate
section of fifth vein; small crossvein slightly beyond middle of discal cell, but proximad
apex of first vein.

Length, 2:5-2-75 mm. .

Holotype, female, Molonglo River, A.C.T., March 20, 1930 (L. F. Graham). Para-
types: female, same data as type; female, Illawarra, New South Wales (H. Petersen) ;
female, Blundell’s, A.C.T., Jan., 1930 (A. Tonnoir); male, Como, New South Wales,
Dec., 1923 (H. Petersen; swept from flowers); male, Mangalore, Tasmania, Nov. 15,
1911 (A. White). Type and two paratypes to be returned to collection of the School of
Public Health and Tropical Medicine, University of Sydney; two paratypes in U.S.
National Museum, Washington; paratype (Tasmania) in British Museum (Nat. Hist.).

The male from Como has the third and fourth veins less widely divergent, but
otherwise is essentially similar to the holotype.

The smooth unfurrowed triangle and black arista and palpus will distinguish this
species from most of the Chlorops previously described from Australia. In Malloch’s
key to Oscinis (1938, p. 338), it will pass to couplet 15 and is close to O. botanica
Malloch, but differs from the latter by having the penultimate section of fourth vein
subequal to ultimate section of fifth, and scutellum not entirely black.

DipLtotoxa Loew.

The genus Diplotoxa is weakly represented from the Australasian region, there being
known at present only D. tasmaniensis Malloch (1927) from Tasmania and three species
described by Malloch from New Zealand. It may be, however, that many more species
remain to be discovered when the fauna of tiny flies is more thoroughly known. In
the material before me there appear to be four new species, of which one from Victoria
is described below. Two specimens from Ohakune, New Zealand, are in poor condition
and will not be described, but it is possible to recognize from available characters that
at least one represents a new species, and perhaps the other also. Still another
specimen, from Illawarra, New South Wales (H. Petersen), may be new or it may be
only a variant of D. tasmaniensis. Description of species, if new, should await the
discovery of adequate and satisfactory material.

DIPLOTOXA VICTORIENSIS, Nl. Sp.
Tiny species, with frontal triangle smooth and polished black, and thorax entirely
black except for narrow yellow median stripe of scutellum.
Male.—Predominantly black, marked with yellow as follows: Front outside frontal
triangle, face, cheek except for narrow brown oral margin, palpus, prosternum, all coxae,
trochanters, ends of all femora broadly (approximately one-fourth at each end), tibiae

BY CURTIS W. SABROSKY. 185

chiefly, and all tarsi except brown distal segment; hind tibia indistinctly browned
centrally, the others faintly so; antenna with arista and dorsal two-fifths of third
segment black, the rest orange-yellow; scutellum with narrow orange median stripe,
to and including the apex; halter with large white knob and dark yellow stalk;
bristles black or brownish-black.

Head: Front slightly broader than long and twice the width of an eye; frontal
triangle at base nearly touching each eye, smooth and ungrooved, apparently entirely
polished but under high power with indistinctly bounded central area of fine sparse
pollen encompassing the ocellar tubercle and an approximately equivalent area anterior
to it: sides of triangle slightly convex posteriorly, then curving inward on anterior
one-third and becoming concave as the triangle tapers acuminately, apex at anterior
margin of front. Head in profile slightly higher than long, the eyes large and very
convex; cheek narrow, one-tenth the eye height and one-third breadth of third antennal
segment, the latter very slightly longer than broad, and with microscopically pubescent
arista; median clypeal plate polished black. Outer vertical bristles long and strong,
the others weak and scarcely distinguishable from hairs, including four pairs of short,
regularly spaced orbitals.

Thorax subshining, thinly dark grey pollinose except for polished black sterno-
pleuron, anteroventral half of mesopleuron, lower fourth of pteropleuron, and the
propleuron chiefly. Notopleural bristles 1 + 2, only the lower posterior bristle long and
evident, the others hair-like. Scutellum short and rounded apically, with a pair of
long apical bristles, widely separated at base, converging towards tips, the usual
subapical scutellars weak and but little stronger than hairs.

Abdomen shining, but apparently thinly pollinose on dorsum. Legs slender, hind
tibia without sensory area. Wing venation approximately as figured by Malloch (1938,
p. 340, fig. 1) for D. tasmaniensis, the crossveins separated by no more than length of
the anterior one.

Length of body, 1:25 mm.; of wing, 1-5 mm.

Holotype, male, Victoria, Australia (C. French). In the British Museum (Nat.
Hist.).

This species is in the same limited group of small species as D. tasmaniensis, but
the latter is a predominantly yellow species with three black mesonotal stripes. The
three described New Zealand species (cf. key by Malloch, 1931, Records Canterbury
Mus., 3: 416) are much larger (2-5 and 3 mm.) and differ from victoriensis in a number
of distinct characters.

As Malloch pointed out for tasmaniensis, the new species is also related to the
Palaearctic approximatonervis Zetterstedt. It may be noted that European monographers
have referred the latter to the genus Lasiosina. If true, that generic assignment must
also be made for these two Australian species. However, generic limits form a difficult
problem in the Chloropidae, and for the present I shall continue the use of the name
Diplotoxa for tasmaniensis and victoriensis.

THAUMATOMYIA Zenker (= Chloropisca Loew).

In my opinion, Thaumatomyia and Chloropisca cannot be maintained as distinct
genera, though some workers do so. The relation of the two was discussed in detail
by Sabrosky (1943, Canad. Ent., 75: 116-117). Malloch’s proposed restriction of
Chloropisca to species with hairs on the mesopleuron (1938, p. 353) does not affect the
problem of the use of the two names, as the type species of both have that character.
Malloch used the name Chloropisca throughout his work on the Australian fauna.

THAUMATOMYIA MALLOCHI, new name.
Chloropisca monticola Malloch, 1927, Proc. Linn. Soc. N.S.W., 52: 430. Preoccupied.
Primary homonym of Chloropisca monticola Becker, 1912, Ann. Mus. Nat. Hungarici
LOZ SO:

186 AUSTRALIAN CHLOROPIDAR,

PEMPHIGONOTUS Lamb.

One species of this genus has hitherto been known from Australia, the genotype
P. mirabilis Lamb, from Melville Island (type locality) and Darwin, the latter recorded
by Malloch (1938, p. 354). There are also two females in the U.S. National Museum
from Buderim Mt., Queensland, December, 1889.

An unusual male specimen from Darwin appears to represent an unnamed species.
According to published descriptions of mirabilis, strong sexual dimorphism is to be
expected in this genus, but the species before me differs from the published description
of the male of mirabilis, and differs from other known species of the genus in characters
not usually involved in sexual dimorphism.

I published notes and a tentative key to the genus in 1940 (Annals Mag. Nat. Hist.,
ser. 11, 6: 421-424). Most of the species, which are chiefly Oriental, are easily
distinguished from the Australian ones by having at least the third segment entirely
black. Only in mirabilis, in the small (3 mm.) P. ochrostoma (Becker) from New
Guinea, and in the species described here, are the antennae entirely yellow. The
present species is hear mirabilis in its large size and in the wing venation, with long
discal cell, long and diagonally placed hind crossvein, and fringe of very long hairs on
the first section of costa. However, the male of mirabilis is described as having the
middle femur and tibia “clothed with abundant long tangled hairs’, whereas the hairs
are short and appressed in the new species. Lamb also made no mention of any
peculiar development of the fore tibia or of the mesonotum centrally, such as are
found in peculiaris.

PEMPHIGONOTUS PECULIARIS, Nn. Sp.

Large orange-yellow species with entirely yellow antenna, black hind tibia, white
and black marked fore tibia, peculiarly depressed and pruinose median stripe on the
mesonotum, and unusually long hairs on first section of costa.

Male.—Predominantly shining and orange-yellow, the small ocellar tubercle polished
black, broad sublateral mesonotal stripes polished reddish, halteres ivory-white, wings
entirely brown; legs orange-yellow, except as follows: hind tibia black except for narrow
reddish stripes on extensor surface; mid tibia brown to blackish anteriorly and dorsally;
fore tibia white with broad black dorsal stripe and an expanded, flattened, velvet black
area antero-dorsally on basal third; all tarsi brown. MHairs in general light yellowish-
brown to dark brown, the long costal hairs black.

Head broad, its width 1:6 times its length, with broad shining front, depressed in
the present specimen though possibly not naturally so, still approximately as long as
broad and twice the width of an eye; frontal triangle not clearly defined, a narrow
glabrous darker stripe extending from ocellar tubercle to the anterior margin, with
the flat areas flanking it slightly more shining than remainder of front, the whole
shining flat area (if that constitutes the frontal triangle) at its base slightly more than
half (0:55) the width of front, with two irregular rows of hairs on its surface, merging
imperceptibly with those of front; eye bare; head in profile 1-1 times as high as long;
cheek broad, slightly more than one-third the height of conspicuously bulging eye and
three-fourths the breadth of large third antennal segment; front barely projecting
before eyes and face weakly receding; proboscis short, labella fleshy, third antennal
segment large, suborbicular, length and preadth subequal, but upper margin slightly
concave at insertion of slender bare arista; no bristles developed, the erect and cruciate
ocellars and post-verticals slightly longer than hairs but not usually distinct from them.

Thorax above (Text-fig. 1) with peculiar median depressed area, which is densely
pruinose and varying with the light from dull brown to shimmering silvery-brown, on
the median line a narrow groove which is wider anteriorly, the whole depressed area
bare of hairs; remainder of mesonotum, and the convex scutellum, covered with long
suberect hairs, those bordering the median depression especially long and appearing
almost as a row of pale bristles; mesopleuron sparsely pale-haired; usual thoracic
bristles not developed, or not easily distinguished from the long hairs. Abdomen short
and much broader than thorax and, like it, with numerous pale erect hairs.

BY CURTIS W. SABROSKY. 187

Fore leg as figured (Text-fig. 3), fore tibia broadened on basal third to form what
may be a sensory area, the anteroventral surface of tibia and anterior surface of femur
opposite the area conspicuously long-haired; mid and hind femora and tibiae with
usual short appressed hairs, those of mid leg not long-haired as described for male
mirabilis; hind tibia with large sensory area posterodorsally.

Wing as figured (Text-fig. 2), notable for the long costal hairs on basal portion,
ending abruptly at subcostal break, the long first costal sector, long and distally
broadened discal cell, and long and obliquely placed hind crossvein.

Length, 4-5 mm.; of wing 4-5 mm.

Holotype, male, Darwin, N.T., Australia, March, 1945 (B. Malkin). Type No. 62372
in the U.S. National Museum.

Ee)

Text-figures 1-5.

1, 2, 3, Pemphigonotus peculiaris: 1, mesonotum; 2, wing ; 3, anterior aspect of left femur
and tibia, fore leg. 4, Batrachomyia occidentalis, antenna. 5, Lasioplewra tonnoiri, profile of
head.

EHUTROPHA NOCTILUX (Walker).

Oscinis noctiluz Walker, 1859, Jour. Linn. Soc. (Zool.), 3: 126 (Aru Islands).
Chlorops nicobarensis Schiner, 1868, Reise der Novara, p. 244 (Nicobar Is.). Chlorops
fuscipennis Thomson, 1868, EHugenies Resa, p. 603 (Rossi Is.). Pseudoformosina nico-
barensis (Schiner) Malloch, 1938, Proc. Linn. Soc. N.S.W., 63: 355, genotype of Pseudo-
formosina Malloch. Hutropha noctilux (Walker) Sabrosky, 1940, Annals Mag. Nat. Hist.,
ser. 11, 6: 420. New generic combination, with nicobarensis and fuscipennis in
synonymy; Pseudoformosina a synonym of EHutropha. Pseudoformosina noctilux
(Walker) Malloch, 1941, Proc. Linn. Soc. N.S.W., 66: 64. Nicobarensis in synonymy.
Eutropha noctilux (Walker) Sabrosky, 1952, Verh. Naturf. Ges. Basel, 63 (1): 216.

N.

188 AUSTRALIAN CHLOROPIDAE,

A wide-ranging Oriental and East Indian species which Malloch mentioned in his
Australian revision as probably to be found in northern Australia. Malloch erected a
new genus for it, but after study of the genotype of Hutropha Loew I am convinced
that Pseudoformosina is a synonym. The above citations will bring the Australian
list up to date.

Subfamily OSCINELLINAE.
LASIOPLEURA Becker (= Parahippelates Becker).

A key to eighteen species and one variety from Australia and Tasmania was
published by Malloch (1940, pp. 268-270). A few notes and one new species from
New Zealand are added below. It should also be noted that there is a complex of
species, involving parva Malloch, pallipes Malloch, aequalis Becker, taylori Malloch, and
some possibly undescribed species, whose accurate determination is difficult without
fully matured specimens in good condition. 'The material before me is not sufficient
to do more than indicate that identifications in this part of the genus must be made
with unusual caution, and that some clarification needs to be done and perhaps new
species distinguished from very similar forms.

As far as I am aware, the new species described alow is the first species of
Lasiopleura to be recorded from New Zealand, although a number of species are
known from Australia and the Oriental region.

LASIOPLEURA PRUINOSA (Thomson), n. comb.

Oscinis pruinosa Thomson, 1868, Hugenies Resa, p. 606 (Sydney, New South Wales).
Parahippelates seticauda Malloch, 1928, Proc. Linn. Soc. N.S.W., 53: 301, 302 (Victoria).
[New Synonymy. ]

The generic reference is based on my examination of type material of pruinosa in
the Naturhistoriska Riksmuseum in Stockholm, and paratypes of seticauda which are
before me. Previous authors have listed Thomson’s species in Oscinella or Oscinosoma,
except for Duda (1934, Arbeiten morph. taxon. Ent. Berlin-Dahlem, 1: 44), who referred
it to Parahippelates. However, he admitted that the species was unknown to him, and
in his paper it will not key out to that name, but rather will pass directly to seticauda
Malloch.

LASIOPLEURA TONNOIRI, Nl. Sp.

Small black species with polished black frontal triangle, broad cheek (0-40 the eye
height), tiny hind tibial spur, and black legs.

Female.—Black, the anterior half of front yellow to orange-yellow, face and cheeks
whitish-yellow, almost silvery, palpus yellow, third antennal segment slightly marked
with orange basally below, and halter lemon-yellow; coxae and legs black, the knees
narrowly reddish and tarsi brown; bristles generally black (cf. note on variation) ;
vibrissal seta, the row of setae along lower margin of cheek, postocular rows, and all
hairs of head, body and legs pale whitish-yellow, glistening in the light at certain
angles; wing hyaline, veins brown.

Head in profile (Text-fig. 5) subquadrate, as long as high, the front sloping and
only slightly projecting beyond eye, the latter with long axis diagonal and 1:5 times
the short axis. Cheek broad, 0-40 times greatest vertical height of an eye and approxi-
mately as broad as height of third antennal segment; vibrissal angle approximately
90 degrees. Front moderately broad, at vertex 2:5 times width of an eye and nearly
0-60 times width of head, and approximately as long as broad; frontal triangle barely
more than half length of front and well separated from eyes at vertex, chiefly smooth
and polished black, the ocellar tubercle, basal corners and side margins bright grey
pruinose. Antennae relatively small, third segment subquadrate, slightly broader than
long; arista relatively short, with microscopic pubescence seen only with high power.
The usual cephalic bristles strongly developed: proclinate and divergent ocellars, erect
and cruciate postverticals, mesoclinate inner and lateroclinate outer verticals, and three
pairs of more or less proclinate orbitals.

BY CURTIS W. SABROSKY. 189

Thorax black, dark grey pruinose, including the metanotum and upper portion of
pleuron; the sternopleuron, except posterior half of narrow upper margin, the anterior
two-fifths of mesopleuron, and the anterior spiracle and area immediately behind and
below it polished black, without pruinosity; scutellum brownish to brown-grey; thoracic
bristles long and strong: two humeral, 1:+ 1 notopleural, 1 presutural, 1 supra-alar,
1 postalar, dorsocentrals 0 + 38, the anteriormost directly opposite the ends of the
incomplete mesonotal suture but apparently a postsutural bristle, 1 subapical and 1
apical scutellar; moderately strong prescutellar acrostichals may or may not be present;
dise of scutellum with two to six long pale hairs. Abdomen black, subshining, densely
grey pruinose.

Legs slender; posterodorsal sensory area on hind tibia short and narrow, less than
one-fifth length of tibia; hind tibial spur black and strong, easily distinguished from
hairs under high power, but short, only half the greatest diameter of tibia, and thus
relatively inconspicuous.

Wing: Second to fourth costal sectors (between apices of first to fourth veins) as
22:16:6:5; veins three and four slightly divergent, fourth vein ending at apex of wing;
penultimate section of fourth vein 1:4 times that of third vein and 0-60 times that of
fifth vein; small crossvein at about two-thirds length of discal cell; hind crossvein
straight, half the length of penultimate section of fourth vein, and at right angles to
penultimate sections of fourth and fifth veins.

Length, 1:75-2 mm.

Holotype and three paratypes, all females, Tahuna, New Zealand, August 27, 1922
(A. Tonnoir). Type and one paratype to be returned to the Canterbury Museum, two
paratypes in the U.S. National Museum.

This species is quite unlike any of the Australian species (cf. Malloch, 1940), which
have the frontal triangle pruinose. Its affinities are rather with several Oriental and
East Indian species, including L. meijeret Sabrosky (1952) (= ornatifrons Meijere,
preoccupied). In Duda’s 1934 key to the Oriental and Australian species, tonnoiri passes
to ornatifrons but differs by hyaline wing, black legs including fore coxa, and broad
cheek.

Variation: Some variation is apparently to be expected in the colour of the bristles,
most of which are black and strongly developed. In the type the postverticals. are
black, but they are yellow in the other three specimens. The type shows three black
orbital bristles on the right, but two black and one (anterior) yellow on the left,
whereas two paratypes show only the anterior pair of orbitals yellow, and the third
paratype has the upper two orbitals on the left and all orbitals on the right yellow.
The thoracic bristles are more constantly black, but even here an oceasional bristle
is pale.

BATRACHOMYIA Krefft.

Krefft, 1864, Trans. Ent. Soc. N.S.W., 1 (1863): 100. Skuse, 1889, Proc. Linn. Soc.
N.S.W., ser. 2, 4: 174.

In his papers on the Australian Chloropidae, Malloch regularly credited the genus to
Skuse (1889). However, in 1941 (Annals Ent. Soc. Amer., 34: 749), I pointed out that
the generic name was first proposed by Krefft (1864), who published on the life history
and figured the larva, pupa, host and a wing of the adult fly. Skuse (1889) was the
first to mention species in connection with the name Batrachomyia.

It is interesting to note the resemblance between this genus and Pemphigonotus
Lamb in the large and robust habitus (for chloropids), the hairy mesopleuron, unusually
long first vein, broad costal cell, and presence of a posterodorsal sensory area on the
hind tibia. Pemphigonotus, however, has the costa stopping at the third vein and
belongs to the Chloropinae. The lack of pubescence on the eyes in the latter genus is
another conspicuous feature separating it from Batrachomyia.

Malloch’s key (1940, p. 264) to his six Australian species may be modified in part.
as follows, to place Chlorops vicaria Walker and the new species described below, as
well as to indicate the position of the two species described by Skuse in 1889.

190 AUSTRALIAN CHLOROPIDAE,

Partial Key to Batrachomyia.

to

1. Thorax reddish-yellow, with shining black stripes on mesonotum .................000.
Thorax reddish-yellow, the mesonotum at most with a suggestion of reddish stripes.
Bb. nigritarsis Skuse (= varipes) and others.
(cf. Malloch, 1940, p. 264.)

2. Third antennal segment entirely black, somewhat triangular in outline, dorsal margin
longer than ventral and anterodorsally somewhat angulate ....................:+:. 3
Antenna yellow to orange-yellow except for black arista and sometimes a small spot at
its base; third antennal segment small, suborbicular, upper margin shorter than lower

and anterodorsally rounded and not angulate.
B. quadrilineata Skuse; B. major Mall.; B. strigipes Mall.

3. Palpus black; anterior ‘crossvein nearly opposite outer three-fourths of discal cell, the

penultimate section of fourth vein only slightly longer than that of third vein.
MEG ha Olchor at B. vicaria (Walker).

Palpus yellow; crossveins well separated, the anterior at or slightly before middle of

discal cell, and penultimate section of fourth vein thus four times the length of penulti-
mate. section cof ofitth: veins Wey esos ccsusiensiaccne un Hae aoc ee Ee ee B. occidentalis, n. sp.

BATRACHOMYIA NIGRITARSIS Skuse.

Batrachomyia nigritarsis Skuse, 1889, Proc. Linn. Soc. N.S.W., ser. 2, 4: 175-176,
Plate 10, figs. 2-5a (New South Wales). Batrachomyia varipes Mallozh, 1940, Proc. LInn.
Soc. N.S.W., 65: 264 (Victoria). Probable synonym.

The descriptions of varipes and nigritarsis are so similar that I have little doubt
of their identity. The combination of reddish-yellow thorax (without black stripes),
black antennae, and characteristically black-marked legs sets the species off from all
others known from Australia. All femora narrowly at apex, the fore and mid tibiae,
the hind tibia at base and apex, and all tarsi are black.

BATRACHOMYIA VICARIA (Walker), n. comb.

Chlorops vicaria Walker, 1849, List Dipterous Insects British Mus., 4: 1120
(Australia ).

Becker, in his 1911 monograph of the Indo-Australian Chloropidae, referred vicaria
to Scoliophthalmus, apparently because of Walker’s mention of the “nearly triangular”’
third antennal segment. It was thus listed by Malloch (1931) in his catalogue of the
Australian Chloropidae (Proc. Linn. Soc. N.S.W., 56: 75). However, the type in the
British Museum (Nat. Hist.), now lacking third antennal segments, shows that the
species is close to Batrachomyia. In wing venation, antenna, and general habitus, it is
similar to the new species described below, but I am informed by Dr. F. van Emden
that the type apparently lacks ordinary hairs on the mesopleuron, though it has dense
pile. With generic limits in this family still not well established, one cannot be sure
whether a new genus is appropriate or whether the generic characterization of
Batrachomyia should be altered. Biological information may some day shed light on
this. In the meantime, it is clear that vicaria is far removed from Scoliophthalmus,
and that in wing and general habitus it is suggestive of Batrachomyia. Accordingly I
refer it there until additional material makes possible further study of its position.

BATRACHOMYIA OCCIDENTALIS, N. Sp.

Yellow species with shining black mesonotal stripes, black and subtriangular third
antennal segment, and yellow palpus.

Female.—Yellow, marked with black as follows: Arista, large spot encompassing
the ocellar tubercle and equally broad area behind it, connecting with broad black band
across upper three-fifths of occiput, the band only narrowly separated from eyes; three
broad shining mesonotal stripes, the median full length from neck to scutellum, the
lateral distinctly divided at mesonotal suture into a slightly broader anterior portion
and a posterior portion; supra-alar vittulae, large humeral spot, metanotum, and spots
on meso- (anterodorsal spot and smaller one on lower margin), ptero-, sterno-, and
hypopleuron, all thoracic spots highly polished; abdomen brown, subshining; median
plate of clypeus brown to blackish; bristles black; hairs of front dark, but otherwise
hairs yellow to whitish-yellow; wing hyaline, veins brown.

BY CURTIS W. SABROSKY. 191

Head slightly collapsed, but front obviously broader than an eye, though only
three-fourths as broad as long; frontal triangle polished, apically rounded and
exceedingly short, barely showing around ocellar tubercle; third antennal segment large
(Text-fig. 4), 1:5 times as broad as long, dorsal margin longer than lower and giving
segment a subtriangular appearance; arista relatively short, its length to height of
third segment as 3 to 2, and barely longer than the whole antenna, microscopically
pubescent; bristles short, outer verticals and postverticals longest and strongest; ocellars
and postverticals erect, cruciate at tips; upper three-fifths of front with five pairs of
erect black orbital bristles, subequal in length to inner verticals, the anterior orbitals
hairlike and merging with frontal hairs, the latter numerous, short but conspicuous.

Thorax polished, without pollen or tomentum; mesonotum and scutellum with
numerous fine silky hairs, which, however, do not interrupt the polished appearance;
scutellum quite flattened on disc, almost as in Thaumatomyia (= Chloropisca), with
three pairs of black scutellar bristles grouped together near apex, the apical pair
slightly longer than the others.

Wing with first vein long, extending midway on wing; costal sectors one to four
as 40:35:18:10; costal and marginal cells broad, submarginal and first posterior cells
relatively narrow; first and second veins anteriorly concave, third only very slightly
so and almost straight on its outer half, and fourth anteriorly convex, the third and
fourth veins thus strongly divergent at apex of wing; ultimate section of fifth vein
slightly longer than penultimate section of fourth; other characters as in key.

Length of body 4 mm., of wing 4 mm.

Holotype, female, Dongarra, Western Australia, Oct. 4-10, 1935 (R. E. Turner).
Type in the British Museum (Nat. Hist.).

This species is noteworthy in having the scutellum so distinctly flattened. Another
species before me, either B. dubia Malloch or near it, shows a suggestion of flattening,
but such species as major and atricornis have the scutellum highly convex.

It is interesting to record this species from far western Australia. The previously
described species, except possibly vicaria. which was recorded only from “Australia”,
are from New South Wales, Victoria and Tasmania.

CAVICEPS DEFECTA (Becker).
Oscineila defecta Becker, 1911, Ann. Mus. Nat. Hungarici, 9: 163 (Java).

Malloch (1924, Proc. Linn. Soc. N.S.W., 49:356) certainly appeared to refer defecta
to his new genus Caviceps. but perhaps he did not intend to make a positive reference
inasmuch as he later (1927) mentioned C. flavipes as the only known species of the
genus. Still later (1940) he stated that he had said that defecta “probably belonged”’.
At any rate, I have seen the type in Amsterdam, and it unquestionably belongs to
Caviceps.

A male example of Caviceps from Dongarra, Western Australia, Aug. 23—-Sept. 5,
1935 (R. E. Turner) [British Museum (Nat. Hist.)] may or may not represent a new
Australian species. This fully matured specimen has darker legs than described for
either flavipes or defecta. the legs being yellow with brown to blackish marks as follows:
All coxae, a narrow and weak median band on mid femur, the median third to two-fifths.
of hind femur and hind tibia, and narrow median bands on fore and hind tibiae. The
concave face is black, as described for defecta. It is possible, in view of the close
similarity in other characters, that colour variation or difference in degree of maturity
of specimens may be involved, and I would not wish to describe this single individual
without supporting material.

SCOLIOPHTHALMUS Becker.

As noted under Batrachomyia, Chlorops vicaria Walker is not a Scoliophthalmus
and can be deleted from that genus in the Australian catalogue.

192 AUSTRALIAN CHLOROPIDAEK,

LIOSscINELLA Duda.

Malloch had little material from Western Australia, and from the few specimens
which I have seen thus far it appears that a number of new species remain to be
found there. In this large genus the species are often very close, and their proper
distinction will take careful study, and above all longer series than were available
to Malloch in most cases, in order to be sure of variation in colour and maturity, and
any sexual dimorphism. I am not satisfied with the generic assignments of Australian
species in this portion of the subfamily, but must postpone that more complex task
for the present.

LIOSCINELLA FLAVOAPICALIS (Malloch), n. comb.

Oscinosoma flavoapicalis Malloch, 1931, Rec. Canterbury Mus., 3: 411 (New Zealand).
Oscinosoma diversipes Malloch, 1931, l.c., p. 414 (New Zealand). Lioscinella nigropolita
Malloch, 1941, Proc. Linn. Soc. N.S.W., 66: 47 (New South Wales; Australian Capital
Terr.; “Darwin, N.T.’). [New Synonymy. |]

Malloch misinterpreted the labels on two paratypes of nigropolita and recorded them
as “N. Territory: Darwin (Palmerston)’’. In reality, the labels read “Palmerston Nth
[North] N.Z.’’. Although Malloch recognized no species of the genus as common to
Australia and New Zealand, this appears to be one. Both the holotype (New South
Wales) and allotype (Austral. Cap. Terr.) of nigropolita, and the two paratypes now
recognized as being from New Zealand, are before me, and I can find no evident
difference, nor did Malloch, as he included all in the same type series: Accordingly, the
prior name for the species is flavoapicalis Malloch. Oscinosoma diversipes, synonymized
by Miller (1950, Catalogue Diptera New Zealand Subregion, p. 110), is the slightly
smaller male of the same species. I may also note that the name flavoapicata used
three times by Malloch (1931, l.c., p. 414) is only a lapsus or typographical error for
flavoapicalis.

193

THE CULEX PIPIENS GROUP IN SOUTH-EASTERN AUSTRALIA. III.
AUTOGENY IN CULEX PIPIENS FORM MOLESTUS.

By N. V. Dosprotworsky, Georgina Sweet Fellow in Economic Entomology,
University of Melbourne.

(One Text-figure. )
[Read 27th October, 1954.]

Synopsis.

The number of eggs in rafts, laid autogenously, varies from seven up to one hundred and
thirty-five. High temperatures do not influence the frequency of autogeny, but reduce the
number of eggs laid and their viability. Females prefer polluted water for oviposition, but
do not avoid clean water entirely. The frequency of autogeny in wild populations of Victorian
molestus is high; it varies from 46-6 per cent. to 94:2 per cent.

Study of autogeny in C. pipiens L. led to the separation of the autogenous form
molestus from the anautogenous pipiens, s.str. (Roubaud and Gaschen 1932, Marshall
and Staley 1935, 1937, Jobling 1938) and initiated a wide discussion on the taxonomic
status of molestus. The separation of molestus and pipiens, s.str., is based mainly on
biological characters and the most important diagnostic character of molestus is its
autogeny.

It is now well established that autogeny is an inherited character and that it
depends on the ability of the larva to build up a large fat body which permits the
maturation of eggs without a blood meal. Moreover, in autogenous mosquitoes, formation
of eggs commences in virgin females immediately after emergence from the pupa. This
phenomenon has been observed in the three known autogenous species found in Victoria
(C. pipiens form molestus, Aédes concolor Taylor and Tripteroides tasmaniensis Stricl.),
and can be used as a basis for recognizing autogeny in forms which do not readily mate,
or Oviposit, in the laboratory.

NUMBER OF RAFTS
(Oo)
[e)

ID 2) 30 40 SO GO 70 80 GO 1CO IO 120 igo
NUMBER OF EGGS

Text-fig. 1—The number of the eggs in rafts laid autogenously.

Only one egg-raft can be produced autogenously; each later oviposition must be
preceded by a blood meal. In my experiments, the maximum number of rafts produced
by a single female was six; Laven (1951) states that this number may be exceeded. The
number of eggs usually decreases at each successive oviposition: in a group of five
females the autogenous raft contained an average of 101 eggs; the following five, each
deposited after a meal of human blood, contained an average of 84, 81, 73, 70, and 49
eggs respectively.

194 THE CULEX PIPIENS GROUP IN SOUTH-EASTERN AUSTRALIA. It,

The number of eggs in the autogenous raft is variable. In counts of 1152 rafts the
number ranged from 7 to 135; the mean was 57-7 (Text-fig. 1). The size of these rafts
is undoubtedly affected by the larval diet. When this is poor, the fat body is small and
there is, in consequence, a reduction in the number of eggs. Tate and Vincent (1936)
and Shute (1951) have observed that a poor larval diet also reduces the frequency of
autogeny. This would imply that there is a correlation between the frequency of
autogeny and the size of the egg-rafts. The data of Table 3 tend to support this view,
but statistical analysis has shown that the correlation is barely significant.

TABLE 1.

The Influence of Temperature on Autogeny.

| Number of Eggs.
Number of Percentage a
Females. Temperature. Autogenous. Viability.
Total. Average.
|
50 20° C. 80 2912 72-8 90:0%
50 31°C. 80 Fi 2275 56-9 58-8%

Raft size is also influenced by temperature. In one experiment, the adults reared
from a single batch of eggs were divided into two groups; one group was maintained at
20°C, the other at 31°C. The higher temperature did not affect the frequency of autogeny
but reduced the number of eggs laid and their viability (Table 1).

The preference shown by molestus for breeding in polluted water, rich in decaying
organic matter, is notorious, but it also breeds in clean water. Thus, in Cairo, Knight
and Malek (1951) found it in clear water ground pools and wells; it is found in similar
situations in Victoria. That this is due to a lack of more suitable breeding sites was

TABLE 2.
Autogeny and Siphonal Index in Wild Populations of molestus.

Number of Eggs
Number Per- in Raft. Siphonal Index.
Date of of Locality. Habitat. centage
Collection. Females. Auto-
genous. Min. | Max. | Mean.| Min. | Max. | Mean.
March—April ae 52 Yarram. Liquid 94-2 it 114 65-2 3°6 4-9 4-5
manure.
April He Ae 120 Rosanna. Liquid 55-3 10 46 P3303} Boy 4-5 4-2
manure.
March-April ahs 110 Spotswood. Liquid 86-4 20 121 85-5 3°83 4-6 4:3
manure.
March .. Beg 125 Melbourne. Liquid 76°8 31 100 56-0 3°6 4-5 4-2
manure.
February ae 110 Melbourne. Concrete 50-0 13 68 38-1 Bot 4-6 4-3
pit.
August-September 91 Ringwood. Quarry. 67-0 15 106 43-7 3°38 4-6 4°3
February—March 146 Pt. Lonsdale. —= 50-7 21 96 59-4 3:5 4-5 4-1
May-June ee 103 Moe. — 46-6 a 44 23-4 3:9 4-7 4-4
Average at ae =a es 65°8 = = 49-3 — — 4:3

BY N. V. DOBROTWORSKY. 195

shown by an experiment in which three containers (20” x 20”) were filled, the first with
a manure infusion, the second and third with clean water; the third container was
black, the others white. During May, 75 rafts were laid in the first container, two in
the second and one in the third. The ovipositing females evidently select sites which
provide optimum conditions for the larvae.

While molestus is described as an autogenous mosquito, many authors (Mathis 1940,
Knight 1951, Knight and Malek 1951, Mattingly 1952, 1953) have claimed that the
frequency of autogeny is low in natural populations; Mattingly has suggested that the
high frequency observed in laboratory colonies is a result of unconscious selection.
This, no doubt, is true of some strains of molestus, but in Victoria, as the following
observations show, the frequency is high in wild populations.

Larvae and pupae were collected from various natural breeding places and brought
into the laboratory where they completed their development in water from the original
sites. Hach day the emerged adults were removed to cages (1000 ce. inches) provided
with an oviposition dish and cotton wicks soaked in a sugar solution. The cages were
kept in a constant temperature room at 25°C. A count of the egg-rafts was continued for
twenty days following the introduction of the last adult. The results are shown in
Table 2. In two localities, Point Lonsdale and Moe, where the breeding sites were not
found, engorged females were collected from a chicken house and a bedroom respectively ;
the figures for these groups refer to an F, population reared from these females.

In the eight populations, the proportion of autogenous females ranged from 46-6
per cent. to 94-2 per cent., with an average of 65-8 per cent. This figure agrees with
that of Theodor (1953) who reported that in Israel 70 per cent. of ‘pipiens’ are
autogenous.

The siphonal index (Table 2) was uniform throughout the eight strains; no
differences in adult, or larval, morphology were observed.

Acknowledgement. For interest and assistance in the preparation of this paper, the
writer is indebted to Dr. F. H. Drummond. Zoology Department, University of Melbourne.

References

JOBLING, B., 1938.—On two Subspecies of Culex pipiens L. (Diptera). Trans. Roy. Ent. Soc.
Lond., 87: 193-216.

KNIGHT, K. L., 1951.—A Review of the Culex pipiens Complex in the Mediterranean Subregion
(Diptera, Culicidae). Trans. Roy. Ent. Soc. Lond., 102: 354-364.

KNIGHT, K. L., and Mauex, A. A., 1951.—A Morphological. and Biological Study of Culex
pipiens in the Cairo Area of Egypt. Bull. Soc. Fouad ler Ent., 35: 175-185.

LAVEN, H., 1951.—Untersuchungen und Deutungen zum Culex pipiens Komplex. Trans. Roy.
Ent. Soc. Lond., 102: 365-368.

MARSHALL, J. F., and Srauey, J., 1935.—Some Adult and Larval Characteristics of a British
‘Autogenous’ Strain of Culex pipiens L. Parasitology, 27: 501-506.

, 1937.—Some Notes Regarding the Morphological and Biological Differentiation of

Culex pipiens L. and Culex molestus Forskal (Diptera, Culicidae). Proc. Roy. Ent. Soc.
Lond., (A), 12: 17-26.

MATHIS, M., 1940.—Biologie d’une souche de Culex pipiens autogenicus Roubaud, 1933, au cours
dun élévage de vingt generations en série. Bull. Soc. Path. exot., 33: 201-207.

MATTINGLY, P. F., 1952.—The Problem of Biological Races in the Culex pipiens Complex.
Proc. Linn. Soc. Lond., 163: 53-55.

, 1953.—The Culex pipiens Complex. Trans. Ninth Int. Cong. Ent., 2: 285-287.

RouBAuUD, E., and GASCHEN, H., 1932.—Differentiation des races biologiques de Culex pipiens
L. par l’adaptation lavaire aux milieux ammoniacaux. Bull. Soc. Path. exot., 25: 1053-1058.

SHUTE, P. G., 1951.—Culex molestus. Trans. Roy. Ent. Soc. Lond., 102: 380-382.

TATE, P., and VINCENT, M., 1936.—The biology of autogenous and anautogenous races of Culex
pipiens L. (Diptera, Culicidae). Parasitology, 28: 115-145.

THEODOR, O., 1953.—Discussion: The Culex pipiens Complex. Trans. Ninth Int. Cong. Ent.,
2293.

196

AN Fl HYBRID BETWEEN HLUCALYPTUS CINEREA ¥F. MUELL. AND
EUCALYPTUS ROBUSTA SM. ;

By L. D. Pryor.
(One Text-figure.)
[Read 24th November, 1954.]

Synopsis.
The experimental production of a viable Fl between EH. robusta Sm. and H. cinerea
F. Muell., from the series Transversae and section Macrantherae (Normales) is recorded.
Inheritance of some juvenile characters is intermediate in the F1, while others resemble
either parent.

The systematic schemes proposed for various genera often give a good indication
of genetic relationships. This is true of Maiden’s scheme for the eucalypts which
Blakely (1934) developed and presented in the “Key to the Eucalypts”.

In a large genus such as this it would indeed be surprising if the first monographs
were entirely in accord with the relationships of the species as determined by genetic

E. cinerea. The Fl hybrid. E. robusta.
H. cinerea x E. robusta.

Although the F1 is taller than the two parents, it is too early to say whether there
is hybrid vigour or not.

study. Duffield (1952) has pointed out in connection with the genus Pinus that
established systematic treatment aids genetic study and, conversely, genetic study aids
systematic review. The evidence presented here helps to establish affinity between
two groups.

The primary subdivision of the genus in the work mentioned is made on the basis
of the shape and characters of the anthers. There is an amount of evidence (Pryor,
1951, 1953) to show that some such groups—as, for example, the Renantherae—coincide

BY L. D. PRYOR. 197

with a set of species in which the members are freely interbreeding but which are
apparently isolated from all other groups.

In the case of the Macrantherae, however, which contains a large number of species,
it is equally clear that several ot’,erwise rather distinct groups of species are placed
together on the basis of anther morphology. It is probable that some of these cannot
interbreed. One group of the Macrantherae, the Transversae, is well defined by a
number of characters. It is cut off rather sharply from the ‘“Macrantherae-Normales”’
_ group, which contains a set of species, many of which are known to hybridize between
themselves. It has been suggested, on rather slender grounds, that these two groups
can interbreed. Any data which would more rigorously support this view are of
interest because they would indicate affinity between them, and so help to reduce to a
more sound basis the grouping of the units at present lumped together somewhat
“artificially” in the Macrantherae.

The record, then. of the successful production by manipulated pollination of an FL
hybrid between H. robusta Sm.—which belongs to the series Transversae—and LH. cinerea
F. Muell—belonging to the section Macrantherae (Normales)—is of considerable
interest since it establishes experimentally that at least these two members of the
group can successfully interbreed to the extent of producing viable F1 progeny.

TABLE 1.
Juvenile Leaf Characters.
zy es el : bs ost
E. robusta. E. cinerea. | Fl, E. cinerea = E. robusta.
|
|
1. Stems square, somewhat | Terete. | Square, somewhat winged.
winged. |
2. Plants green. Glaucous. Somewhat glaucous.
3. Leaves petiolate. Sessile. Very shortly petiolate.
4. Leaves alternate. Opposite. Opposite.
5. Leaves lanceolate. | Orbicular. Intermediate (ovate-lanceo-
late).

Characters 2, 3 and 5 are intermediate; 1 is like H. robusta and 4 is like EH.
cinerea.

In spring, 1952, manipulated pollinations were made on bagged flowers of H. cinerea,
emasculated about a week before. The species used as the male parent in each case
were substantially distinct from #. cinerea, the combinations being chosen because they
represented supposedly wide crosses. Controls, not pollinated, and selfings were made
at the same time. The controls did not set seed, whereas the selfings set normally
and produced good seed. Of the cross pollinations, all failed except one, which was
notably successful, namely, H. robusta, Fruit developed normally in the four bags
used, producing a total of some thirty fruit. They were harvested in October, 1953,
and about seventy plants were raised. These by March, 1954, reached the seventh pair
of leaves and were vigorous. The progeny was strikingly uniform and had morpho-
logical characters in the juvenile state, some of which were intermediate between
those of the two parents, which in themselves are quite distinct, and some identical
with one or other parent. This is summarized in Table 1.

This hybrid combination is of considerable interest for two reasons. Firstly, no
other case is so far recorded of the experimental production of a hybrid between
species belonging one each to the systematic groups Macrantherae-Normales and the
Transversae.

Secondly, a similar combination has not yet been suggested, even by the less
critical method of progeny testing, from naturally occurring supposed hybrids, although
a few such mixtures are hinted at from morphological evidence collected in the field.
At the same time it is a combination which cannot occur naturally, as the two species
are separated by, a considerable geographical distance and occupy ecological sites which

198 Fl HYBRID BETWEEN EUCALYPTUS CINEREA AND E. ROBUSTA.

are widely divergent. HE. cinerea is a species of the Southern Tablelands of New South
Wales at elevations generally above 1,500 feet, and does not descend to the coastal
plain, whereas H. vobusta is a species of low-lying areas along the whole New South
Wales coast which are frequently subject to tidally induced flooding and the soil of
which is often somewhat saline.

While one may not proceed from this one example to the generalization that all
species of the Transversae and the Macrantherae-Normales can successfully hybridize,
the particular evidence makes the assumption that many of them may successfully
interbreed now much more probable than previously. In view of this, there are
important possibilities from the point of view of breeding programmes, aimed both at
research and practical ends, which warrant further experiment.

Many of the Transversae, which contains about 20 species, are excellent trees
(such as-#. saligna, E. resinifera, E. botryoides, E. propinqua and EH. diversicolor),
both from the point of view of timber and in cultural characters required in species
for rural planting. Most, however, are not very frost-resistant nor can they be planted
as a rule beyond the well-watered areas of the country. The prospect that characters
from this group may be brought together in hybrid combination with species within
Macrantherae-Normales (containing the series Microcarpae, Globulares, Semiunicolores,
Viminales, Argophyllae and Paniculatae), which amount to about seventy, is of
considerable interest. Many of these possess cold and drought resistant characters
which are missing from the Transversae. The combination already produced is of this
nature and, if ultimately it fiowers and is found to be fertile, it may give segregates
of considerable value in the F2 generation.

Following experience with other groups in the genus, it is likely by analogy that
since one hybrid combination has been produced, many others can be obtained between
pairs of species from the same groups, and some of these are likely to be economically
valuable.

References.
BLAKELY, W. F., 1934.—A Key to the Eucalypts. Sydney.
DUFFIELD, J. W., 1952.—Relationships and Species Hybridization in the Genus Pinus. Zeit-

schrift fiir Forstgenetik wu. Forstpflanzenziichtung, 1: 4, 93-97.

Pryor, L. D., 1951.—PrRoc. LINN. Soc. N.S.W., !xxvi: 140-148.
———. 1953.—PrRoc. LINN. Soc. N.S.W., Ixxvili: 43-48.

199

A NOTE ON THE OCCURRENCE OF “ANOMALOUS” KRASNOZEM IN THE
RICHMOND-TWEED REGION OF NEW SOUTH WALES.

By J. W. McGarity and D. N. MUNNS.
(Communicated by Di. W. R. Browne.)
(Plate viii, and One Text-figure.)
[Read 24th November, 1954.]

Syncpsis.

cy

The krasnozem soils lying to the west of the 55” rainfall isohyet in the Region, previously
described as “anomalous”, are not developed on basalt but on a highly sesquioxidic clay
material interstratified in basalt.

The interstratified material is described and some analytical data for this and associated
materials presented. The origin of the material is uncertain but the occurrence of some of
the contiguous red and “‘black’’ soils of the Region is thereby explained.

The red and “black” soils developing on Tertiary basalt in the Richmond-Tweed
region of New South Wales have been described by Hallsworth (1951) and classified
at the great soil group level. The pattern of distribution of these soils is such that
the red soils or krasnozems cover large areas of the basalt plateau to the east of
the 55” rainfall isohyet while the “black” soils, chocolate and praire, are found on the
more sharply dissected plateau remnants to the west. Hallsworth (1951, 1953) and
Nicholls (1952) consider that the climatic factor is the chief determinant in the present
distribution of the krasnozems, while -Teakle (1952, 1953) holds the view that they are
relict soils, some of which are derived from laterite.

The presence of small areas of krasnozems lying in lower rainfall areas to the
west of the 55” rainfall isohyet in this region, although observed by Hallsworth (1951),
has not been explained adequately. Hallsworth regarded these soils as “anomalous”
because they did not fit the climatic theory of distribution, and suggested that they
were relics of a previous wetter climate.

Recent observations made near Kyogle and Casino indicate that the ‘‘anomalous’”’
krasnozem soils are not derived from a normal basaltic parent material but from a
relatively easily weathered stratum within the main basalt mass.

The interstratified material, of which three distinct types are recognizable, often
forms extensive, roughly horizontal sheets from one to six feet thick, outcropping
at much the same level over distances of a half-mile or more. Several bands of this
material, separated by unweathered basalt, have been observed outcropping, on steep
scarps, indicating that the material is not uniquely associated with one flow or period
of activity (Fig. 1).

The material may be described as a compacted, unctuous clay which adheres to
the tongue on moistening and slakes in water with a faint crackling sound, but does
not show expansion properties.* Both red and grey types are found interstratified in
the basalt, but the former has the wider distribution. ‘A further type has a strong
vesicular structure.

The hardness of the red clay varies from slightly compacted to moderately indurated,
the latter variant being less commonly encountered, while colour ranges from bright
read to reddish brown. Typical examples are shown in Plate viii, figures 1 and 2,
underlying unweathered or partly weathered basalt. The red clay, at its junction with
the overlying basalt, generally shows an ashy grey layer from 3” to 3” in depth, which
is of fine particle size and often unconsolidated. This layer changes abruptly into the

*The term bentonite has not been used because of this last-named property. Bole, as
defined by Holmes (1920), closely resembles some of the material here described.

200 OCCURRENCE OF “ANOMALOUS” KRASNOZEM,

red compacted clay which is uniform with depth, brittle and crumbly, and which
fractures spheroidally. The fracture faces are lustrous and sometimes have a broken
black patina. Despite the ease of fracture, the hardened clay can be augered by hand
only with extreme difficulty or not at all. Sometimes the clay is slightly vesicular, the
vesicles being partly filled with a soft pale yellow, soapy material resembling kaolinite.

The red clay usually passes into a shallow layer of altered basalt, but in some
cases a pale grey to black friable layer up to one foot in depth is found between these
two zones.

Thompson and Beckmann (1953) have referred to similar red material near
Toowoomba as “baked clay”.

The grey clay is found less frequently and is generally in a more weathered state.
Apart from colour, the morphology closely resembles that of the red clay. Since the
grey type is not important in the genesis of the “anomalous” krasnozems, it is not.
considered further.

eS Nl

aes PID LS
oy ane — IOOO0'

BASALT

/
Pammmebta iy py mat ee OS

Text-fig. 1.—Three layers of outcropping red clay at Wyneden near Upper Eden Creek.

The remaining type referred to as vesicular clay appears to be a highly altered
type of scoriaceous basalt. In some exposures it is found grading into, and mixed with,
the red clay described above.

The red clays were examined mineralogically and chemically. They were found
to be free of primary minerals and could be dispersed with strong acid treatment into
particles of clay size. No nitrogen and only traces of carbon were found on analysis.
The content of Si, Fe, Ti and P for samples taken from the top and bottom of the
red layer is shown (Table 1) and compared with analyses of typical Krasnozems and
basalts. The ferruginous nature of the clay is of note.

The origin of the interstratified material is in doubt. The red material of a
similar nature observed by Thompson and Beckmann (loc. cit.) was associated with
laterite developed on basalt and buried beneath a later basalt flow. In the Kyogle
district there is no evidence of laterite formation on the basalt itself, although massive
laterites and companion horizons capped by basalt are found on the underlying Clarence
sediments.

As none of the beds shows evidence of sedimentation, it would appear that alteration
has taken place in situ. If the process were pedochemical, then the material represents
a highly weathered ferruginous, buried soil, consolidated by the overlying basalt flow.
Speculatively, the absence of primary minerals would indicate this could not have
been a chocolate or praire soil. The chemical data show the similarity of the red clay
and krasnozem soils on elimination of “loss on ignition” (Table 2).

Whatever the origin of this highly sesquioxidic clay material, its importance as a
factor in the genesis of some of the “anomalous” krasnozems cannot be overlooked.
It explains the occurrence of these red soils at Fairy Hill, Dome Mountain and
Rukenvale, where the “black” and red soils are intermingled in a complex pattern. In
these situations the parent material of the red soils is mainly the highly weathered,

BY J. W. MCGARITY AND D. N. MUNNS. 201

TABLE 1.
Analytical Data.
Percentage.
| Depth in |
Material. Locality. Position. Inches. Loss
| on Ig- Si. Fe. Ti. ipa
nition.
|
aie =a |
Red clay+ a .. | Fairy Hill. Top. 0-6 7:9 17:49 24-32 3:06 0-108
Bottom. 48 9-9 13-50 20-38 2-46 0-130
Red clay} eae .. | Backmede. Top. 0-6 9-5 19-78 17-60 2°13 0-123
Bottom. 24-36 9-4 20°14 12-92 2-04 0-105
Red-brown clayy+ .. | Backmede. Middle. 12-24 7-9 21-55 12-63 1-05 nd
Krasnozem a .. | Nashua. Top. 3- 6 19-0 14-01 13-23 3°48 0-222
Bottom. ‘60-72 11-4 15-74 11-83 2-40 0-272
Krasnozem a .. | Nashua. Top. 3-— 6 18-4 13-30 19-46 3-12 nd
Bottom. | 60-72 12-3 15:18 12-96 1:98 nd
Anomalous krasnozem Woodview. Top. 3- 6 13-8 14-15 14-70 3:18 0-282
Middle. 12-24 9-5 | 15°55 14-70 2-10 0-174
} |
Basaltt a Se ENE Saye | Average of 8 samples .. ae 21195 USD) 1-26
| Sf | 20-07 6-30 | 0-60
| Range ‘\ |—23-35 | —8-40 |—1-68
Basaltst and Doleritest | N.S.W. | Average of 17 samples .. Ae 21-46 | 8:04 1-26
| | JO, eae 6-30 | 0-60
| Panne ‘L |-23-85 |—11-90 |—2-64
|

* Analyses by Miss H. V. Rayner.
+ Interstratified material.
t David’s ‘‘ Geology of the Commonwealth of Australia ’’.

interstratified red clay with varying amounts of addition from surrounding higher
basalt. In less obvious cases where the original outcrop has been eroded and removed
(as on hilltops), the red soils still contain fragments of the hardened ciay in the
sand fraction, e.g. Woodview.

The view of Hallsworth, therefore, that these red soils are relict and represent
small areas left uncovered by later outpourings of.basalt, is not tenable. The patchy
distribution of these ‘‘anomalous” krasnozems merely reflects the patchy distribution
of the outcropping parent material. The varying degree of redness of the soils reflects
the variability of this material, and the effects of such factors as erosion and topography.
Where the influence of this material is comparatively weak and that of basaltic
colluvium strong, as on the sides of steep hills, soils with a reddish chocolate surface
and varying degree of subsoil redness may develop. This is particularly noticeable in
the dissected area between Kyogle and Nimbin.

Similar occurrences of interstratified material have been observed elsewhere in
New South Wales. Near Murrurundi in the Liverpool Range, and at Guyra in the

TABLE 2.
Comparison of Interstratified Clay and Krasnozem.
Composition as Percentage of Ignited Material.
Material.
Si. Fe. Mbt, IP,
Interstratified clay .. | Average 5. 20-14 19-13 2-34 0-127
Range. 14-87-23 -25 13 - 63-26-24 1-13-3-30 0-115-0-143
Krasnozem aa .. | Average 6. 16:50 16-56 3:10 0-270
Range. 15-75-17 -53 13-18-23 -04 2-22-4-14 0-191-0-321

202 OCCURRENCE OF “ANOMALOUS” KRASNOZEM.

New England Tableland, abnormally reddish soils amongst normal chocolate soils are
developed from similar red and vesicular clays apparently lying between basalt flows.
If such occurrences are widespread it may well be that some of the transitional
krasnozem — chocolate, and reddish chocolate —normal chocolate ‘catenas’, hitherto
regarded as the result of topographic effects (Hallsworth et al., 1952), may be effects
of parent material differences.

This study has been assisted by a University of Sydney research grant.

References.

HALLSWoORTH, E. G., 1951.—Aust. J. Agric. Res., 2: 411-28.

, 19538.—Aust. J. Sci., 15: 175-6.

HALLSWORTH, HE. G., COSTIN, A. B., GIBBONS, F. R., and ROBERTSON, GWEN K., 1952.—J. Soil
Sci., 3: 103-24.

Houmes, A., 1920.—The Nomenclature of Petrology. Thomas Murby. London.

NicHOLLs, K. D., 1952.—A.N.Z.A.A.S., Sydney Meeting, 1952.

TEAKLE, L. J. H., 1952.—Aust. J. Agric. Res., 3: 391-408

—-—, 1953—-Aust. J. Set., 152 176-7.

THOMPSON, C. H., and BECKMANN, G. G., 1953.—Australian Conference in Soil Science.
Summaries of Papers, Vol. 1, 1.19.1-5, Adelaide, 1953.

EXPLANATION OF PLATE VIII.
1. Fairy Hill. Chocolate soil and partly weathered basalt (A) overlying red clay,
(B) in a roadside cutting.
2. Fairy Hill. Close up, showing the ashy incoherent layer (C) at the junction of
basalt (A) and red clay (B).

203

SUPPLEMENTARY NOTES ON THE GENUS BRACHYCOME CASS.
DESCRIPTIONS OF FIVE NEW AUSTRALIAN SPECIES AND SOME NEW LOCALITY RECORDS.
By Gwenpba L. Davis, Department of Botany, University of New England, Armidale.
(20 Text-figures. )

[Read 27th October, 1954.]

Synopsis.

Descriptions of the five new species are accompanied by figures designed to illustrate habit,
vegetative features and fruits. Affinities and intraspecific variation are discussed in the text.

A number of new locality records are listed which extend the known range of several
species for a considerable distance.

INTRODUCTION.

Knowledge of the distribution and natural variation of genera and component
species is directly correlated with the activity of collectors, both amateur and pro-
fessional. Within recent years a large number of specimens of the genus Brachycome
Cass. have been received by the writer and, although the majority were referable to
well Known species, certain of them were quite distinct and are accordingly described as
new species.

The range of several species is now known to be more extensive than was previously
recorded (Davis, 1948, 1949), and these new records are now listed under the appropriate
species. The present location of each specimen cited is indicated as in previous papers
(Davis, 1948, 1949) with the following additions:

The Waite Institute, Adelaide (WAT).
C.S. & I.R.O., Canberra (©):
Parks and Gardens Herbarium, Canberra (GE)F

TAXONOMY.

CoMPpoSsITAE. Tribe ASTEROIDEA.
BRAacHycomMer Cass., Dict. Sci. Nat., xxxvii (1825): 471.
Subgenus HUBRACHYCOME G. L. Davis.
Superspecies LEPTOCARPA.

BRACHYCOME ULIGINOSA, Sp. Ov.
(Text-figures 1—4).

Holotype: Heathland at eastern foot of Black Range, Western Grampians Region,
Victoria, 2.11.1948, J. H. Willis (MEL). Paratypes: Two. Loc. cit. (MEL).

Herba perennis, glabra, 9-28 cm. alta; folia radicalia ad 6 cm. longa, 6 mm. lata,
oblanceolata, acuta, aut integra aut interdum 1-4 lobis linearibus, in basi in petiolum
gracilem fastigata; pedunculi robustiores, 1-3 linearibus phyllis; capitula 1-3, ligulis
circiter 2-5 cm. expansis, involucri phylla circiter 22, 4-5 mm. longa, 1-1-5 mm. lata,
subacuta-obtusa, linearia—anguste-ovata, integra; flores radii in speciem violacei,
circiter 36, 5-7-5 mm. longi, 1-1-5 mm. lati; receptaculum 5 mm. latum, 2-5 mm.
altum, hemisphaericum, non alte punctum; achaenia 1-8 mm. longa, 1-2 mm. lata, fusea,
ovata, compressa; corpus leve et glabrum, ala angusta, crassa, integra et pilis margine
minutis glandulosisque; pappus albis, simplicibus setis tam longis aut longioribus quam
incisura intra alas.

Glabrous perennials 9-28 cm. high. Leaves radical, clustered, up to 6 cm. long, 6 mm.
broad, oblanceolate, acute, entire or occasionally with 1-4 linear lobes; tapering
proximally into a slender petiole. Scape rather robust, with 1-3 linear bracts.
Inflorescences 1-8 to each plant, about 2-5 em. across the expanded rays. Involucral
bracts about 22, 4-5 mm. long, 1-1-5 mm. broad, subacute to obtuse, linear to narrow-
ovate, entire. Ray florets apparently mauve, about 36, 5-7-5 mm. long, 1-1-5 broad.

C0)

204 SUPPLEMENTARY NOTES ON THE GENUS BRACHYCOME,

veceptacle 5 mm. broad, 2-5 mm. high, hemispherical, shallowly pitted. Fruits 1-8 mm.
long, 1:2 mm. broad, dark golden-brown, oval, flat; the body oblong, smooth and
glabrous, sharply demarcated from the narrow, thick, entire wing which bears minute
white glandular hairs along the margin. Pappus of white simple bristles equal to or
slightly exceeding the notch between the wings.

Habitat: Swampy situations.

Range: Southern and western districts of Victoria. :

Distribution: Eltham, 21.8.1903, P. R. H. St. John (MEL); Brisbane Ranges, 17
miles S.W. of Bacchus Marsh, 30.10.1943, J. H. Willis (MEL); Moyston, 10.1881, D.
Sullivan (MEL); Mt. William Creek, 10.1879, D. Sullivan (MEL); Eastern foot of
Black Range, near ‘The Pass’, heathland swamp, 2.11.1948, J. H. Willis (MEL); Follett,
27.8.1905, F. M. Reader (MEL); Little Desert, 8 miles south of Lawloit and 2 miles east
of Catiabrin Springs, swampy tracks, 11.9.1949, J. H. Willis (MEL); Wimmera, 1893,
W. E. Matthews (MEL).

This species appears to be widely distributed in western Victoria, but due to the
close vegetative resemblance to B. scapigera (Sieb. ex Spring) D.C. may well have
been recorded as that species from other districts and even other States. Both species
bear similar leaves and the base of the plant in each instance is surrounded by the
fibrous remains of previous season’s leaves. The only vegetative feature which dis-
tinguishes B. uliginosa from B. scapigera is the tendency in the former species for
occasional leaves to bear a few linear lobes.

The fruits of B. uliginosa show certain similarities to both B. radicans Steetz ex
Lehmann and B. dissectifolia G. L. Davis, and specimens from Moyston and Mt. William
Creek were previously recorded as new records for the latter species (Davis, 1949). The
relationship, however, is closest to B. radicans, from which the fruits can be dis-
tinguished by their oval body, and non-inflated thick wing.

B. radicans is widely distributed in swampy situations in eastern Australia and it
is possible that both B. uliginosa of Victoria and B. dissectifolia of New England
originated as geographic subspecies of B. radicans.

BRACHYCOME RARA, SDP. NOV.
(Text-figures 5-8.)

Holotype: Floodwaters of the Wilson River, western Queensland, 9.1922, W.
MacGillivray. (WAI. 8118.) Paratypes: Two. Loc. cit. (WAI. 8140.)

Herba in speciem annua, circiter 10 cm. alta aut altior, a basi ramosissima et mic.
glandulosa; folia caulina ad 4 cm. longa, 1 cm. lata, lanceolata, sessilia, integra, acuta;
pedunculi axillares et terminales, graciles, aut nudi aut uno parvo filio; capitula
circiter 35, ligulis diametro 1-3 cm. expansis; involucri phylla 22, 3-5 em. longa, 1-1-1-5
mm. lata, oblanceolata, subacuta-obtusa, mic. glandulosa et fimbriato-ciliata; flores radii
circiter 24, 4 mm. longi, 1-2 mm. lati, probabiliter caerulei; receptaculum 2 mm. latum,
1-5 mm. altum, hemisphaericum; achaenia 1-1-2 mm. longa, 0-6-0-8 mm. lata, cuneata,
compressa, rubida; latus utrumque gibbus fuscum longum, utrimque longo sinu, habet;
margines crassae et teretes sunt; pappus mic. ora est.

An apparently annual plant about 10 cm. high or higher, much branched from the
base and microscopically glandular all over. Cauline leaves up to 4 cm. long, 1 cm.
broad, lanceolate, sessile, entire, acute. Peduncles axillary and terminal, slender, naked
or with a single small leaf. Inflorescences about 35, 1-3 em. diameter across the
expanded rays. Involucral bracts 22, 3-5 cm. long, 1-1-1-5 mm. broad, oblanceolate,
subacute to obtuse, microscopically glandular and torn ciliate. Ray florets about 24,
4 mm. long, 1-2 mm. broad, probably blue. Receptacle 2 mm. broad, 1-5 mm. high,
hemispherical. Fruits 1-1-2 mm. long, 0-6-0-8 mm. broad, cuneate, flattened, reddish-
brown, with a raised dark-brown protuberance running vertically on each face,
bordered on either side by a longitudinal fold; margins thick and rounded. Pappus
represented by a microscopic rim.

Range: Only known from the type locality.

Specimens examined: Type series only.

BY GWENDA L. DAVIS. 205

Text-figures 1-20.

1-4, B. uwliginosa.—1, Habit x 4; 2, Leaf variation x 4; 3, Fruit x 12; 4, Distribution.

5-8, B. rara.—5, Habit x 1; 6, 7, Fruit, front and side views x 12; 8, Distribution-

9-13, B. gracilis.—9, Habit x 4; 10, 11, 12, Fruit, side, front and back views, x 12;
13, Distribution.

14-17, B. riparia.—14, Habit x £; 15, 16, Variation in Fruits x 12; 17,. Distribution.

18-20, B. Eyrensis—18, Habit x 2; 19, Fruit x 12; 20, Distribution.

206 SUPPLEMENTARY NOTES ON THE GENUS BRACHYCOME,

The fruits of this species are unique in their possession of a dark projecting
longitudinal ridge on the centre of each face. No similar structure has been seen in any
other species, but the pad-like confluent tubercles on the fruits of B. Whitei G. L. Davis
suggest its origin from a structure of this nature. The thick margin-like wing can only
be seen to advantage in the mature fruits, following the development of the longitudinal
folds which demarcate the body.

Since this species has never been collected before, it is possible that it makes its
appearance only after heavy rain when the country is under flooded conditions and
collectors are unlikely to be in the area.

BRACHYCOME GRACILIS, Sp. Nov.
(Text-figures 9-13.)

Holotype: Mt. Major, Dacite outcrop near Dookie, Victoria, 28.10.1953, A. Tylee
(MEL). Paratypes: Five. Loc. cit.; in sandy loam at Mt. Major, 6.11.1949, A. S. Forgas
(MEL).

Herba annua, gracilis, pilis paucis-multis, ad 9 em. alta, a basi ramosa; folia
caulina 1-1-8 em. longa, pinnatipartita, in basi fastigata; partes 5, lineares, 2 mm.
longae, 1 mm. latae, acutae; pedunculi filiformes, terminales; capitula ligulis diametro
2 ecm. expansis, maturitate diametro 5-6 mm., hemisphaerica; involucri phylla 9-13,
2-5-3 mm. longa, 1-5 mm. lata, rhombata, acuta, mic. fimbriato-ciliata et glandulosa;
flores radii 8-12, ligulis albis, 4 mm. longis, 1-5 mm. latis; receptaculum 1mm. latum,
1 mm. altum, acute conicum, altius punctum; achaenia 1-5-1-7 mm. longa, 1 mm. lata,
nigra, cuneata, in ultima parti valde curvata; corpore ab crassis, rigidis marginibus,
quae modo sinuum supra pappi basim pro-currunt, inclare secto; pappus albus, con-
spicuus, non stellatus, setis simplicibus.

Slender sparsely septate-hairy annuals, up to 9 cm. high, branching from the base.
Leaves cauline, 1-1-8 em. long, pinnatipartite, tapering proximally; segments five, linear,
2 mm. long, 1 mm. broad, acute. Peduncles filiform, terminal. Inflorescences 1 em.
diameter across the expanded rays. Infructescences 5-6 mm. diameter, hemispherical.
Involucral bracts 9-13, 2-5-3 mm. long, 1-5 mm. broad, rhomboidal, acute, micro-
scopically torn-ciliate and glandular. Ray florets 8-12; the rays white, 4 mm. long,
1-5 mm. broad. Receptacle 1 mm. broad, 1 mm. high, steeply conical, rather deeply
pitted. Frwits 1-5-1-7 mm. long, 1 mm. broad, black, cuneate, strongly curved distally:
central body inconspicuously demarcated from the thick rigid margins which extend
upwards as curved folds above the point of insertion of the pappus. Pappus white,
conspicuous, not stellate, the bristles simple.

Range: Only known from the type locality.

Specimens examined: Type series only.

All specimens examined have the same slender appearance and are probably
ephemeral, the whole life cycle being completed during spring. Although this species
has only been collected from the type locality, it is possible that its actual distribution
is more extensive and that previously flowering plants have been referred to B.
goniocarpa Sond. et F. Muell., with which there is a close resemblance vegetatively and
in the young fruits. The mature fruits, however, are strongly curved through the
development of a thick wing and an affinity with B. campylocarpa J. M. Black is
indicated.

BRACHYCOME RIPARIA, Sp. NOY.
(Text-figures 14-17.)

Holotype: Snowy River, east of Butcher’s Ridge, “in crevices and amongst porphyry
rocks, below flood level’, 22.1.1954; N. A. Wakefield, No. 4804 (MEL). Paratypes: Three.
Loc. cit. (MEL.)

Herba perennis, ramosissima, aut erecta aut ascendens, ad 36 cm. alta, indumento
mic. glanduloso; folia multa, caulina, cuneata, inferiora ad 35 cm. longa, 7 dentibus
acutis et petiolo gracili, superiora minora et postremo sessilia; pedunculi plerumque
graciles, 1-2 foliis filiformibus; capitula plurima, ligulis diametro ad 1-5 cm. expansis;
involucri phylla 18-22, 4 mm. longa, 0-6—-0-8 mm. lata, linearia, acuminata, mic. fimbriato-

BY GWENDA L. DAVIS. 207

ciliata et glandulosa; flores radii circiter 25, ligulis 3 mm. longis, 0-8 mm. latis, aut
albis aut caeruleis; receptaculum 1-5-2 mm. latum, 1-5 mm. altum, conicum; achaenia
1-9-2-1 mm. longa, 1-1-1-4 mm. lata, oblonga-obovata, compressa; fulva; corpore
anguste-elliptico, ab latis in acuta segmenta inaequaliter pinnatisectis alis acute secto;
pappus simplicibus setis tam longis aut paulo longioribus quam incisura intra alas.

Much branched erect or ascending perennials, up to 36 cm. high, with a micro-
scopically glandular indumentum. Leaves numerous, cauline, cuneate; those towards the
base up to 3-5 cm. long, with 7 acute teeth, and tapering into the slender petiole. Upper
leaves smaller, and finally sessile. Peduncles usually slender, with 1 or 2 filiform
leaves. Inflorescences very numerous, up to 1:5 em. diameter across the expanded rays.
Involucral bracts 18-22, 4 mm. long, 0-6-0-8 mm. broad, linear, acuminate, micro-
scopically torn-ciliate and glandular. Ray florets about 25, the rays 3 mm. long, 0-8 mm.
broad, white or blue. Receptacle 1-5-2 mm. broad, 1-5 mm. high, conical. Fruits 1-9-2-1
mm. long, 1-1-1-4 mm. broad, oblong to obovate, flat, light brown; the body narrow-
elliptical, sharply demarcated from the broad wings which are very irregularly dissected
into acute lobes. Pappus of simple bristles equal to, or slightly exceeding, the notch
between the wings.

Range: Inland districts of eastern Victoria, along river banks.

Specimens examined: Victoria, Wangrabelle, Genoa River, “abundant amongst
granitic rocks, in crevices, about 400 ft.’’, 3.1948, N. Wakefield No. 2222 (MEL); Genoa
River gorge, 1.1947, N. A. Wakefield No. 3298 (MEL); Genoa. River, near Maramingo
Hill, “in crevices and amongst granitic rocks, below flood level”, 1.1.1954, N. A. Wakefield,
No. 4803 (MEL); Gelantipy, Snowy River, 22.1.1953, L. Hodge No. 4718 (MEL); Snowy
River, east of Butcher’s Ridge, “in crevices and amongst porphyry rocks, below flood
level’, 22.1.1954, N. A. Wakefield No. 4804 (MEL); gorge track of the Upper Snowy River
near Deddick, 27.1.1948, J. M. Béchervais (MEL); Betebolong, Snowy River, “growing in
erevices of granite rocks in the river bed’, 12.1945, N. A. Wakefield, No. 3299 (MEL).

This species is confined to eastern Victoria where “it extends for at least 20 or 30
miles along the Snowy and Genoa Rivers, and is never far away from the waterside
rocks”.* Mr. Leo Hodge has established some plants in his garden in East Gippsland,
one of which “produced a dense mass of stems, to form a tuft about 18 inches across’’.*
These were all transplanted by Mr. Hodge “from the sand high above the average rise of
the Snowy River where Acacia has grown tall. In such places it is quite spindly, but
higher up on the cliffs where the soil is better and there is less shade, it is quite bushy,
though not so much as those of my garden which are more vigorous and have more
flowers”.

Vegetatively B. riparia strongly resembles certain populations of both B. aculeata
(Labill.) Less. and B. marginata Benth., a resemblance which is also apparent in the
fruits. A notable feature of the fruits is the variation in the dissection of the wing, no
two fruits, even of the same capitulum, being identical in this respect. All, however, are
deeply and irregularly cleft into acute lobes. Originally regarded as an aberrant form
of one of these related species, the enthusiasm and interest of the two collectors
mentioned have produced sufficient material to show that its particular characters are
constant throughout its range, and to justify specific status.

Subgenus MrTABRACHYCOME G. L. Davis.
Superspecies IBERIDIFOLIA.
BRACHYCOME EYRENSIS, Sp. nov.
(Text-figures 18-20.)

Holotype: Figure-of-Hight Island, Recherche Archipelago, Western Australia,
7.11.1950, J. H. Willis (MEL). Paratypes: Three. Loc. cit. (MEL.)

Herba in speciem annua, 3-7-16 cm. alta, a basi in 2-7 primos caules ramosa, in
ultima parti mic. glandulosa; folia radicalia ad 2 em. longa, 4 mm. lata, petiolis et 3-5
altis acutis dentibus plerumque adsunt; folia caulina, inferiora ad 2 cm. longa, petiolis

* Personal communication of Mr. N. A. Wakefield, of Noble Park, Victoria.

208 SUPPLEMENTARY NOTES ON THE GENUS BRACHYCOME,

brevibus et 3-5 dentibus, superiora minora et plerumque integra, multa sunt; pedunculi
graciles axillares et terminales, aut nudi aut uno parvo folioso phyllo; capitula ex
magnitudine et ramorum numero 3-40; involucri phylla 9-10, exteriora 1-6-1-8 mm.
longa, 1 mm. lata, oblonga, integra, glandulosa, in ultima parti teretia; flores radii 25-40,
ligulis 1-5-2 mm. longis, 0-5 mm. latis; receptaculum 1 mm. latum, 0-5 mm. altum,
compressum, vix punctum; achaenia 1 mm. longa, 0-5 mm. lata, cuneata, compressa,
maturitate nigra, in media parti mic. canis tessellis et paucis crispis pilis interdum in
ultima parti; pappus abest.

Apparently annual plants 3-7-16 em. high, branching from the base into 2-7 primary
stems; microscopically glandular distally. Radical leaves usually present, up to 2 cm.
long, 4 mm. broad, petiolate, with 3-5 deep acute teeth. Cauline leaves numerous, the
lower shortly petiolate, up to 2 cm. long, 3—5-toothed; the uppermost smaller and often
entire. Peduncles slender, axillary, and terminal, naked or with a single small leaf-like
bract. Inflorescences 3-40 according to size and degree of branching of the plant.
Involucral bracts 9-10, the outer 1-6-1-8 mm. long, 1 mm. broad, oblong, entire,
glandular, rounded distally. Ray florets 25-40, the rays 1-5-2 mm. long, 0:-5 mm. broad,
“white or pale bluish”. Receptacle 1 mm. broad, 0-5 mm. high, flattened, hardly pitted.
Fruits 1 mm. long, 0-5 mm. broad, cuneate, flat, black at maturity, with microscopic grey
tessellations; a few curled hairs may be present distally, but a pappus is absent.

Range: South-eastern coastal belt of Western Australia in the Recherche-Israelite
Bay area. ;

Specimens exanined: Western Australia: Near Mt. Ragged, 1889, S. Brooke (MEL);
between Mt. Ragged and Victoria Spring, near Israelite Bay, 1886, S. Brooke (MEL);
Round Island, Recherche Archipelago, 18.11.1950, J. H. Willis (MEL); Figure-of-Hight
Island, Recherche Archipelago, 17.11.1950, J. H. Willis (MEL).

According to Willis (1953) “abundant on many islands, from Figure-of-Hight to
Goose (but not seen on the larger masses of Mondrain and Middle) is a noteworthy
variant of Brachycome pusilla. This decumbent annual has rather foliose branches,
broad strongly dissected leaves and short white or pale bluish rays”.

Specimens collected by Mr. Willis are similar in habit and vegetative features to
B. exilis Sond., but the fruits are closest to those of B. bellidioides Steetz. Their flattened
nature and complete absence of pappus are, however, distinctive features which justify
specific status.

The specific epithet was chosen at the suggestion of Mr. J. H. Willis, and refers to
the fact that this species appears to be confined to the Eyre Region of Teakle (1987),
which embraces the coastal sandheath between Albany and Israelite Bay.

New Locality Records.
Subgenus EHUBRACHYCOME G. L. Davis.
Superspecies TENUISCAPA.
BRACHYCOME oBovaTA G. L. Davis, Proc. Linn. Soc. N.S.W., Ixxiv (1949): 146.
New South Wales: Kosciusko, 7000 ft., swamp, 10.3.1953, E. Gauba (CP).
This species was previously recorded only from eastern Victoria.

BRACHYCOME STUARTIT Benth., Fl. Aust., iii (1866): 513.

New South Wales: Point Lookout, New England, 17.4.1937, K. Ingram (KI); Boyd
River, Blue Mts., wet mossy places, 4.3.1954, EH. Gauba (CP).

This species has now been recorded from as far west as Mudgee (Davis, 1949) and
south to the Blue Mountains, where Dr. Gauba reports having seen it also in the
Cudgegong Mountain District.

Superspecies LEPTOCARPA.
BrRACHYCOME DEBILIS Sond., Linnaea, xxv (1852): 477.
New South Wales: Between Cumnock and Beldry, 15.10.1947, C. W. E. Moore no.
867 (C).
Hitherto recorded in New South Wales only from western and south-western
districts.

BY. GWENDA L. DAVIS. \' 209

Superspecies BASALTICA. es,
BRACHYCOME NOVA-ANGLICA G. L. Davis, Proc Linn. Soc. N.S.W., Ixxiii cans) Jp
New South Wales: Between Grafton and Glen Innes, 25.4.1912, J. B. Clelan pce:

BRACHYCOME MULTIFIDA DC., Prod. V (1836): 302, Var. pILATATA Benth., Fl. Avust., iit
(1866) : 520.
New South Wales: Smoky Cape, Macleay River, 2.9.1941, K. Ingram (KI); South
West Rocks, 12.1946, K. Ingram (KI).
Previously not recorded north of the Manning River.

Superspecies ACULEATA.

BRACHYCOME MUELLEROIDES G. L. Davis, Proc. Linn. Soc. N.S.W., Ixxili (1948): 194.

New South Wales: Bulgandry Reserve, white rays, plentiful around lagoons, rooted
in mud or in water with Cotula and Centipeda spp., 14.10.1949, H. J. McBarron (NSW.
No. 10168); Fagan’s Reserve, 3 miles north of Walbundrie, white rays, plentiful on
margins of lagoons, in mud or water, mixed with Calotis sp., 14.10.1949, E. J. McBarron
(NSW. No. 10167).

These records link up the two original localities cited, Wagga and Nathalia, and the
ecological notes confirm the earlier suggestion (Davis, 1948) that this species occupies a
wet habitat among herbage. Until now, the colour of the ray florets has not been known.

Superspecies TESQUORUM.
BRACHYCOME BLACKII G. L. Davis, Proc. Linn. Soc. N.S.W., Ixxiii (1948): 207. i
Central Australia: Mt. Ultim, 1.9.1930, J. B. Cleland (JBC); Mt. Allen, Summit,
9.8.1936, J. B. Cleland (JBC).

Subgenus METABRACHYCOME G. L. Davis.
Superspecies IBERIDIFOLIA.
BRACHYCOME EXILIS Sond., Linnaea, xxv (1852): 449.
South Australia: Greenby Island, North Island, 20 miles from Eyre Peninsula,
12.1947, Adelaide Bush Walkers (JBC).

Superspecies TRACHYCARPA.
BRACHYCOME TRACHYCARPA F. Muell., Linnaea, xxv (1852): 339.
' Western Australia: Halfway between Mt. Ragged and Victoria Springs, 1886,
S. Brooke (MEL).
This is the first and only record of this species from Western Australia.

Superspecies SILPHIOSPERMA.
BRACHYCOME PERPUSILLA (Steetz) Benth. var. TENELLA (Turcz.) G. L. Davis, Proc. Linn.

Soc. N.S.W., Ixxiii (1948): 231.

New South Wales: Corner Reserve, 6 miles from Henty, fairly common in barer
grassland and red sandy loam, 6.9.1947, E. J. MecBarron (NSW. No. 10173); Bulgandry
Reserve, fairly common but little flowering in grassland, red sandy loam, 2.10.1949,
EK. J. McBarron (NSW. No. 10166); Walbundrie, common in damp depressions in open
paddocks, 17.9.1948, E. J. McBarron (NSW. No. 2014); Jindera Gap, common in gravel
soil 2.10.1949, E. J. McBarron (NSW. No. 10165).

These specimens extend the range of this species into southern New South Wales
where it appears to be relatively common in grassland.

Acknowledgements.

My thanks are extended to the Directors of State Herbaria and private collectors
mentioned in the text, for lending the specimens on which this paper is based.

In particular, I would like to acknowledge the assistance of Mr. J. H. Willis, of the
National Herbarium, Melbourne, and Mr. N. A. Wakefield, of Noble Park, Victoria, with
both of whom I have been in constant communication in connection with the new
species collected by them.

I am also indebted to Miss Greta Baddams, formerly of the University of New
England, for writing the Latin diagnoses of the new species.

Als) SUPPLEMENTARY NOTES ON THE GENUS BRACHYCOME.

References.
Davis, G L., 1948.—Revision of the genus Brachycome Cass. Part 1. Australian Species.
Proc. LINN. Soc. N.SiW., Lxxiii: 142-241.
1949.—Revision of the genus Brachycome Cass. Part 3. Description of three new
Australian species and some new locality records. Proc. LINN. Soc. N.S.W., Ixxiv: 145-152. .
TEAKLE, L. J. H., 1937.—Regional Classification of the Soils of Western Australia. Journ. Proc.
Roy. Soc. W.A., 24;146.
Wi.uis, J. H., 1953.—The Archipelago of the Recherche. Aust. Geog. Soc. Reports, No. 1: 21.

211

THE INFLUENCE OF THE PHYSICAL PROPERTIES OF A WATER CONTAINER
SURFACE UPON ITS SELECTION BY THE GRAVID FEMALES OF AEDES
SCUTELLARIS SCUTELLARIS (WALKER) FOR OVIPOSITION
(DIPTERA, CULICIDAE).

By A. K. O’GOwER.*
(One Text-figure.)

{Read 24th November, 1954.]

Synopsis.

Aédes scutellavis (Walker) belongs to the subgenus Stegomyia, the members of which
predominantly breed in tree holes and other similar water containers, thus the selection of
an oviposition site by the members of this group was thought to be guided more by visual
and tactile stimuli than by olfactory stimuli.

The physical properties of the water container surface were found to be important in
determining the attractiveness cf an oviposition site due to the visual and tactile stimulation
of its surface. These physical properties were the texture of the surface, the illuminance
reflected from it, and the amount of moisture or water present within the container.

When these properties were studied separately a rough textured surface was preferred
to a smooth one, a surface of low reflectance was preferred to one of high reflectance, and a
free water surface was preferred to a moist, porous surface.

When these physical properties of the water container surface were studied jointly, it
appeared that the gravid mosquitoes were first attracted to an oviposition site by its reflected
illuminance, but the decision whether to oviposit, or what proportion of the total egg batch
was to be deposited, was governed by the texture of the surface. A moist porous surface of
rough texture and low reflected illuminance was so attractive as an oviposition site that all
other single preferences for the various surfaces did not occur; it was therefore concluded
that such a surface, being similar to that of the preferred oviposition sites found in nature,
probably did determine the attractiveness of an oviposition site.

INTRODUCTION.

The selection of an oviposition site by gravid female mosquitoes has been found
to be influenced by (1) such chemical and physical properties of the water as salinity
(Woodhill, 1941), temperature (Hecht, 1930; Weyer and Hundertmark, 1941), organic
pollution (Buxton and Hopkins, 1927; Manefield, 1951); (2) the type and amount of
vegetation present in the water (Russell and Rao, 1942; Hess and Hall, 1945; Rozeboom
and Hess, 1944); (3) the illumination of the breeding site (Muirhead-Thomson, 1942) ;
and the physical properties of the surface of the water container, such as the illuminance
reflected from its surface (Jobling, 1935: Kennedy, 1942), the texture of the surface
and the water vapour pressure gradient.

In the majority of these oviposition investigations, gravid mosquitoes of different
species were offered choices between containers of water which varied in some property
which the author considered as important in limiting the larval distribution of the
species. Although much information has been gained in this manner about species
which breed in ground water, little has been learnt of the environmental factors which
influence the selection of an oviposition site by those species which breed in container
habitats.

As the larvae of the scutellaris group of mosquitoes are always found in small
accumulations of water, such as occur in tree holes, coconut shells, empty cans and
other similar containers (Farner and Bohart, 1945), and as Penn (1947) failed to
show any delimitation of the larval habitat of Aédes scutellavis due to organic pollution,
salinity, water flow, water temperature, pH, specific gravity or vegetation, it was
thought that the selection of an oviposition site by the gravid females of this species

* School of Public Health and Tropical Medicine, University Grounds, Sydney.

212 INFLUENCE OF PHYSICAL PROPERTIES OF SURFACE ON SELECTION FOR OVIPOSITION,

was probably influenced by the properties of the water container surface, rather than
by any properties of the water itself.

Thus the physical properties of the water container surface which were studied
in this investigation are the illuminance reflected from the surface, either high, medium
or low; the texture of the surface, either rough or smooth; and the gradient of the
water vapour pressure from either a free water surface or a moist porous surface, such
as is obtainable by wetting a filter paper.

EXPERIMENTAL TECHNIQUE AND RESULTS.

Because the adults of A. scutellaris readily feed, copulate and oviposit within
confined spaces, and as the larvae of this species are always found in small accumu-
lations of water, it was thought that the behaviour pattern involved in the selection
of an oviposition site by the gravid females of A. scutellaris in nature was similar
to that observed in colonies of this species in 10 inch x 10 inch x 12 inch cages, which
contained either filter papers folded into cones or Petri dishes as the water containers.
These cages were kept in a constant temperature and humidity room operating at
27° + 2°C., with a saturation deficiency of 10 2 mm. of mercury. Hach cage contained
approximately fifty adults of each sex, either two or three water containers whose
relative positions were altered daily, and some raisins for food. Human blood meals
were offered thrice weekly, and deaths were replaced by adults from a stock colony.

TABLE 1.
Percentage Reflectances of Filter Papers Used as Water Containers.

Wavelength in Millimicrons.

Filter Papers. | |
400 450 500 550 600 650 | 700

|
Black Smooth Dyed 4 4 3 3 4 4 5
Black Rough Dyed 4 4 3 3 3 4 6
Black Smooth igs ae 7 7 6 6 6 6 7
Grey Smooth ae ae 28 28 27 26 25 26 31
Grey Rough at is 22 23 Py) ill 21 23 26
White Smooth ss Bo 84 86 87 87 87 88 90
White Rough on 3.9 78 82 82 82 82 82 $3

When studying the texture and reflected illuminance of the water container surface
the containers were made by folding filter papers of 11 cm. diameter into cones and
placing them in beakers. Water was added to these until the beakers were full and
the water levels were half the height of the cones.

When studying the preference for either a free water surface or a moist, porous
surface, the containers were Petri dishes of 9 cm. diameter with paper taped around
the sides whose reflectance matched that of the dishes. Into one dish water was
poured and beneath it was placed a filter paper of the required reflectance, and in the
other dish was a water soaked pad of cotton wool with a filter paper of the required
texture and reflectance on top of it.

The filter papers* used to form the water containers had surfaces which were
(i) white and smooth; (ii) white and rough; (iii) grey and smooth; (iv) grey and
rough; and (v) black and smooth. White smooth papers and grey rough papers were
dyed black with a “Tintex Dye” as required.

The percentage reflectances of these filter papers used as water containers were
measured over the visual range of the spectrum by a General Hlectric spectrophoto-

* (i) Whatman No. 5; (ii) Postlip Mills No. 633C; (iii) Allnutt and Sons No. B.1;
(iv) Allnutt and Sons No. D.3; (v) Schleicher and Schiill No. 551.

BY A. K. O’GOWER. flees

meter* using magnesium carbonate as the surface of reference (Table 1). These
reflectances were expressed as ratios (column 3 of Tables 2, 3, 4,5 and 6) by assigning
a value of 100 to the paper of highest reflectance and expressing all the reflectances
of the other papers used in these experiments as ratios of it.

The times of oviposition upon black, grey and white filter paper water containers
were individually determined by counting the numbers of eggs deposited upon each ai
four-hourly intervals throughout four consecutive days.

- All the eggs deposited in water containers of low reflectances in a twenty-four
hour pericd were laid in daylight, with the majority being deposited between 9 a.m.
and 5 p.m.; however, the majority of the eggs deposited in a water container of high
reflectance were laid at dusk and to a lesser extent at dawn (Figure 1).

PERCENTAGES

SUMMED

MEAN

lam. Sam. Sam. lpm. Spm. Sem.
TIME

_ Yext-fig. 1—Time of oviposition by A. scutellaris. A, upon a black, water container
surface. B, upon a grey, water container surface. C, upon a white, water container surface.

The daily rate of oviposition and the mean total number of eggs deposited by single
female mosquitoes was individually determined for twenty females after each had fed
once. It was found that the eggs developed from a single blood meal were deposited
either in toto, over a period of two or three consecutive days, or over a period of four
days. The mean total number of eggs deposited by each female mosquito was 62.

1. The effect of the illuminance reflected from the surface of a water container upon
its attractiveness as an oviposition site.

The gravid mosquitoes were first given a choice between two water containers
whose surfaces were either black or grey, grey or white, and black or white; then
the choice was between three water containers whose surfaces were black, grey and
white.

From Table 2 it can be seen that a close correlation exists between the reflected
illuminance of a water container surface (columsn 3 of Tables 2, 3, 4, 5 and 6) and
its attractiveness as an oviposition site (column 6 of the same tables), provided the
textures of the surfaces are smooth (experiments 1 to 4 of Table 2). An example is
experiment 4, in which the percentage ratio of reflectances of the filter papers was
7:30:100, whilst the percentage ratio of eggs deposited upon each paper was 59:34:4.

. * These measurements were made by the Division of Physics, National Standards Labora-
tory, C.S.I.R.O., University Grounds, Sydney, and the author wishes to thank Dr. R. G.
Giovanelli for this service.

214 INFLUENCE OF PHYSICAL. PROPERTIES OF SURFACE ON SELECTION FOR OVIPOSITION,

TABLE 2.
The Relationship between the Reflectance of the Water Container Surface and Its Attractiveness as an
Oviposition Site to the Females of A. scutellaris.

Mean
Experiment | Number of | Water Container | Reflected | Number of Percentage
Number. | Replicates. | Surface. | Illuminance. | Eggs of Eggs
Deposited. Deposited.
|
1 | 7 | Black Smooth 7 | 5,593 70
Grey Smooth 30 2.384 30
2 7 | Grey Smooth | 30 | 3,783 91
White Smooth | 100 413 9
3 | 7 Black Smooth | 7 8,364 96
White Smooth 100 Patil 4
4 11 Black Smooth | 7 5,653 59
Grey Smooth | 3 | 3,504 37
White Smooth 100 391 4

2. The effect of the texture of a water container surface upon its attractiveness as an
oviposition site.

The gravid mosquitoes were given a choice, both in a normal night and day cycle
and in constant darkness, between two water containers whose surfaces were black
and smooth, and black and rough. Alternatively the mosquitoes were offered as an
Oviposition site a smooth, black, filter paper water container, from whose centre
radiated regularly spaced, black, cotton threads.

When the reflected illuminances of the water container surfaces were similar, the
texture of the surfaces determined their attractiveness (experiments 5 and 6 of
Table 3). Similarly the distribution of the eggs deposited upon a water container
surface was determined by the texture of that surface, as the majority of the deposited
eggs were laid along the black, radiating, cotton threads, whilst the remainder were
laid on the black, smooth paper between the threads (experiment 7 of Table 3).

TABLE 3.

The Relationship between the Texture of the Water Container Surface and Jts Attractiveness as an
Oviposition Site to the Females of A. scutellaris.

Mean
Experiment | Number of | Water Container Reflected. Number of Percentage
Number. Replicates. Surface. Tlluminance. Eggs of Eggs
Deposited. Deposited.
—— as | Sere — — ———
By 11 Black Rough 3 14,663 60
Black Smooth 3 9,750 40
67 | 5 Black Rough 3 3,068 61
Black Smooth 3 2,040 39
a 5 Black Smooth 3 1,494 40
Black Threads 3 2,253 60

* In normal night and day cycle.
+ In constant darkness.

9

3. The effect of the amount of moisture present within a water container upon its
attractiveness as an oviposition site.

The gravid mosquitoes were offered a choice between two Petri dishes as the water
containers for oviposition whose reflectances were similar; one dish contained water,
the other had a moist porous surface.

The container with a free water surface was slightly but distinctly preferred to a
moist, porous surface when both had equal reflected illuminances (experiment 8 of
Table 4).

ad Dee

BY A. K. O’GOWER.
TABLE 4.

The Relationship between the Amount of Water Present within the Container and Its Attractiveness as an
Oviposition Site to the Females of A. scutellaris.

| |
| | Mean
Experiment | Number of | Water Container Reflected | Number of Percentage
Number. Replicates. Surface. Illuminance. | Eggs of Eggs
| Deposited. | Deposited.
|
8 8 | Water Surface 100 | Bes BA
| Moist Porous | |
| Surface 100 | 6,521 | 46
| |

4. The effect of both the reflectance and the texture of a water container surface upon
its attractiveness as an oviposition site.

The gravid mosquitoes were given twc choices. The first was between a black,
smooth surface, a grey, rough surface and a white, smooth surface. The second choice
was between a black, smooth surface, a grey, smooth surface and a white, rough surface.

The attractiveness of water containers of unequal reflectance and varying textures
was primarily determined by the texture of the surface, and the reflected illuminance
became important only when it was so high that its unattractiveness masked any
difference in the texture of the surface (experiments 9 and 10 of Table 5).

TABLE 5.

The Relationship between the Reflectance and the Texture of the Water Container Surface upon Its
Attractiveness as an Oviposition Site to the Females of A. scutellaris.

| Mean
Experiment | Number of | Water Container Reflected | Number of Percentage
Number. Replicates. Surface. Tlluminance. Eggs of Eggs
| Deposited. Deposited.
9 @ Black Smooth 7 6,280 37
Grey Rough 24 10,111 60
White Smooth 100 410 3
10 5 Black Smooth Gi 7,856 63
Grey Smooth 30 4,622 31
White Rough 100 386 6

5. The effect of the reflectance and texture of the water container surface and the
amount of moisture present within the container upon its wttractiveness as an
oviposition site.

The gravid mosquitoes were given two choices between two water containers: the
first between a Petri dish with a free water surface with a black background, and a
Petri dish with a moist, smooth, grey, porous surface; the second choice was between
a Petri dish with a free water surface with a black background, and a Petri dish
with a moist, rough, grey, porous surface.

A water surface with a black background was preferred to a moist, grey, smooth,
porous surface, and this preference was greater than that due to either the difference
in reflectance (experiment 1 of Table 2) or the amount of moisture present (experiment
11 of Table 6). By altering only the texture of the grey, moist, porous surface from
smooth to rough, however, this above preference for the water surface of low reflectance
was completely reversed. Thus the moist, grey, rough, porous surface was decidedly
more attractive to the ovipositing females than the free water surface (experiment 12
of Table 6).

216 INFLUENCE OF PHYSICAL PROPERTIES OF SURFACE ON SELECTION FOR OVIPOSITION,

TABLE 6.
The Relationship between the Reflectance and the Texture of the Water Container Surface and the Amount of Water within
the Container upon Its Attractiveness as an Oviposition Site to the Females of A. scutellaris.
| | | Mean
Experiment | Number of | Water Container | Reflected | Number of | Percentage
Number. | Replicates. | Surface. | INuminance. | Eggs | of Eggs
Deposited. Deposited.
1 | 6 Water Surface with Black |
Background | i 6,522 83
Moist Grey Smooth Porous |
| Surface 24 | 1,298 17
12 5 | Water Surface with Black |
| Background | g 1,279 19
Moist Grey Rough Porous | | :
Surface 30 5,445 81
|

The minimum number of replicates for the calculations of the means of each experi-
ment was five, and. the determinations of significance, although not strictly necessary
because of the clear-cut preferences in all experiments, were made from determinations
of standard deviation differences. Tables of students’ “t’’ were used to read the
probabilities, which in every case were less than 0-001.

DISCUSSION.

Because the larvae of A. scutellaris, a typical member of the subgenus Stegomyia,
are always found in small accumulations of water, many of the factors of a larger
aquatic environment, which may influence ovipositing females of other species, such
as those found in swamps, rice fields and other similar large bodies of water, probably
would not influence the females of this species when selecting a tree hole for oviposition.
Therefore the behaviour pattern of oviposition by the females of A. scutellaris was
thought to be guided more by the properties of the water container surface than by the
properties of the water held in the container. That is, visual and tactile stimuli were
thought to be more important than olfactory stimuli in guiding the selection of an
Oviposition site; for example, a typical breeding site of this species is a tree hole or
a coconut husk, which has a rough texture, a moist, porous surface, a low reflected
illuminance and usually only a small water surface.

On this assumption filter papers of different texture and varying reflected
illuminances were used to form the water containers of the oviposition sites in the
experimental cages, and the only precautions thought necessary for the prevention of
any interaction of the variables introduced were: dyed filter papers only to be used
in the same experiments; textural experiments to be repeated in constant darkness to
eliminate possible visual stimulation; equal volumes of water to be maintained in
the water containers to prevent unequal evaporation affecting the water vapour
pressure gradients; and the sides of the Petri dishes to be taped with paper whose
reflected illuminance matched that of the dish.

Although both Jobling (1935) and Muirhead-Thomson (1942) have investigated the
influence of the environmental and the reflected illuminances upon the selection of an
Oviposition site by the gravid mosquitoes, little attention seems to have been given to
reflected illuminances intermediate between high and low. In the present investigation
the gravid mosquitoes of A. scutellaris were shown to have a preference for a breeding
site which was approximately inversely proportional to its reflected illuminance. By
studying the time of oviposition a relationship was shown to occur between the
environmental illumination and the illuminance reflected from the oviposition site, for
the bulk of the eggs of an egg batch was deposited upon surfaces of medium or low
reflected illuminance during the hours of brightest illumination, that is about midday;
however, the bulk of the eggs of an egg batch was depcsited upon surfaces of high

BY A. K. 0’GOWER. 217

reflected illuminance during the hours of lowest illumination, that is at dusk (see
Figure 1). This relationship explained why a water container surface of high
reflected illuminance was unattractive to ovipositing females.

The selection of an oviposition site by the grain weevils, Rhizopertha dominica and
Calandra granaria, has been shown by Crombie (1941) and Hastham and McCully
(19438) to depend mainly upon the texture of the grain surface. Similarly the gravid
females of A. scutellaris were influenced in their selection of an oviposition site by the
texture of its surface, and although this preference was not absolute, it was constant
for all conditions under which it was studied (see experiments 5, 6 and 7 of Table 3).

When both the texture and the reflected illuminance of the oviposition site were
studied, the preference for a rough textured surface of medium reflected illuminance
over a smooth textured surface of low reflected illuminance (experiment 9 of Table 5)
was the same as that for a rough texture over a smooth texture, both of equal reflected
illuminance (experiment 5 of Table 3). However, a rough textured, surface with a
high reflected illuminance was decidedly unattractive (experiment 10 of Table 5).
From these preferences it was assumed that the gravid mosquitoes first selected an
Oviposition site because of its reflected illuminance, but once there the texture of its
surface determined whether any eggs would be deposited or what proportion of the egg
batch would be deposited. As the gravid female mosquitoes were found to deposit their
eggs either in toto or over varying periods of days, both of these possibilities could
occur.

The preference for a water surface over a moist, porous surface of equal reflected
illuminance, although only slight, was decided and significant (experiment 8 of
Table 4). But by combining the water surface with a low reflected illuminance this
preference was increased until it was greater than that due to either the above preference
for the water surface, or the preference for a surface of lower reflected illuminance
(experiment 1 of Table 2). Thus it appeared that a summation of preferences occurred.

By altering the texture of the moist, porous surface from smooth to rough, and
comparing its attractiveness with that of a water surface of lower reflected illuminance,
the above preference for a water surface was completely reversed; that is, a moist,
porous surface of rough texture and medium reflected illuminance was more attractive
than a water surface of low reflected illuminance. It was therefore concluded that the
combination of a rough texture with a moist surface made the water container so
attractive to ovipositing females that all other preferences were ignored.

This investigation has therefore determined that the physical properties of a water
container surface are of decided importance in guiding the selection of an oviposition
site by a mosquito species which breeds in such water containers as tree holes. The
sequence of events in this selection would appear to be an initial attraction to a
water container due to its reflected illuminance, but once the texture of the surface
and its moisture content were determined by the gravid mosquito, the decision of the
attractiveness of such a site was made. From this it was concluded that a moist,
porous surface of rough texture and low reflected illuminance so closely resembled the
preferred oviposition site of the gravid females in nature that the selection of tree
holes, coconut shells and other similar water containers was governed by these physical
properties of the water container surface.

Acknowledgements.

Acknowledgement is made of the facilities kindly made available by Dr. A. R.
Woodhill with the permission of Professor P. D. F. Murray in the Department of
Zoology, University of Sydney, and of the encouragement and criticism given during
the execution and reporting of this work by Dr. A. R. Woodhill and Dr. L. C. Birch,
of the above Department, and by Mr. D. J. Lee and Dr. H. O. Lancaster, of the School
of Public Health and Tropical Medicine, University Grounds, Sydney.

Acknowledgement is due to the Director-General, Commonwealth Health Depart-
ment, for permission to publish this article.

218 INFLUENCE OF PHYSICAL PROPERTIES OF SURFACE ON SELECTION FOR OVIPOSITION.

References.

Buxton, P. A., and HOPKINS, G. H. H., 1927.—Researches in Polynesia and Melanesia. No. 1
of the Memoirs of The London School of Hygiene and Tropical Medicine, London, W.C.1,
260 pp.

CromsBin, A. C., 1941.—On oviposition, olfactory conditioning and host selection in Rhizopertha
dominica Fabr. (l1nsecta, Coleoptera.) J. exp. Biol., 18: 62-79.

HasrTHAm, L. E. S., and McCuuny, B., 1943.—The oviposition response of Calandra granaria
linn. J. exp: Biol, 2/0): 35-42)

Farner, D. S., and BoHart, R. M., 1945.—A preliminary revision of the scutellaris group of
the genus Aédes. Nav. med. Bull. (Wush.), 44; 37-58.

Hecut, O., 1930.—Ueber den W&armesinn der Stechmtichen bei der Hiablage. Riv. Malar.,
9: 706-724.

Hess, A. D., and Haun, T. F., 1945.—The relation of plants to malaria control on impounded
waters with a suggested classification. J. nat. Malar. Soc., 4: 20-46.

JOBLING, B., 1935.—The effect of light and darkness on oviposition in mosquitoes. Trans. roy.
Soc. trop. Med. Hyg., 29: 157-166.

KENNEDY, J. S., 1942.—On water finding and oviposition by captive mosquitoes. Bull. ent.
Res., 32: 279-301.

MANEFIELD, T., 1951.—Investigations of the preferences shown by Aédes (Stegomyia) aegypti
Linn. and Culex (Culex) fatigans Wied. for specific types of breeding water. Proc. LINN.
Soc. N.S.W., 76: 149-154.

PENN, G. H., 1947.—The larval development and ecology of Aédes (Stegomyia) scutellaris
(Walker, 1859) in New Guinea. J. Parasit., 33: 43-50.

RozEBoom, L. EH., and Huss, A. D., 1944.—The relation of the intersection line to the production
of Anopheles quadrimaculatus. J. nat. Malar. Soc., 3: 169-179.

RUSSELL, P. F., and Rao, T. R., 1942.—On relation of mechanical obstruction and shade to
Ovipositing of Anopheles culicifacies. J. exp. Zool., 91: 303-329.

THOMSON, R. M. C., 1942.—The control of Anopheles minimus by “Shade” and related methods.
Indian med. Gaz., 77: 675-676.

Weyer, F., and HUNDERTMARK, A., 1941.—Versuche tber die Vorzugstemperatur einiger
Anophelen bei der Hiablage. Riv. Malar., 20: 251-257.

WoopHILt, A. R., 1941.—The oviposition responses of three species of mosquitoes (Aédes
aegypti Linnaeus, Culex fatigans Wiedemann, Aédes concolor Taylor) in relation to the
salinity of the water. Proc. LINN. Soc. N.S.W., 66; 287-292.

219

NOTES ON THH MORPHOLOGY AND BIOLOGY OF SCAPTIA VICINA TAYL.
AND A NEW SPECIES OF SCAPTIA (DIPTERA, TABANIDAE).

By KATHLEEN M. I. PnGuisH, Department of Zoology, University of Sydney.
(Thirty-four Text-figures. )
[Read 27th October, 1954.]

Synopsis.

Larvae of Scaptia vicina (Tayl.) were found at Roseville, N.S.W. The larva and pupa are
described and figured. The type and paratypes of S. muscula, n. sp., were reared from larvae
found, in association with the larvae of a species of Myrmeleontidae, in the sand deposits which
occur beneath overhanging rocks in the sandstone areas near Sydney, N.S.W. Imago, larva
and pupa are described and figured.

INTRODUCTION.

Very little work has been done on the immature stages of any of the Pangoniinae.
The larva and pupa of the Nearctic Goniops have been described or mentioned by
McAtee, Malloch and Stone, and also by Schwardt and Walton, but these two papers
have not been seen by the writer. The immature stages of two Australian species have
been described in more detail, Scaptia auriflua (Don.) by Fuller (1936) and Hctenopsis
vulpecula var. angusta (Macq.) by English (1952). In the Tabanidae (Diptera) of
Australia (Mackerras, to be published), eighty-two described species of Pangoniinae are
listed from Australia, so it is apparent that a great deal remains to be discovered
about their early stages.

Feeding was a problem in rearing the Tabanid larvae described here, as the soil
was too dry for earthworms, which have been used successfully by other workers.
Something had to be provided that did not mind dry soil, would not crawl up the
side of the jar, would not decompose if left untouched or partly eaten, and would not
retaliate when attacked. Muscoid pupae filled all these requirements, but were not
readily taken by the larvae; various Lepidopterous pupae were accepted, but were not
always to be found. Dr. D. F. Waterhouse, of the Division of Entomology, C.S.I.R.O.,
suggested Hphestia, and provided a culture, which has been maintained for two years
and has proved most satisfactory.

ScAPTIA vICcINA Taylor (Text-figures 1-14).

This species was described under the name Hrephopsis vicina by Taylor (1918)
from one specimen, a male, from Wentworth Falls, N.S.W. It was transferred to
Scaptia by Ferguson (1926), within which it is being placed by Mackerras (paper to
be published) in the maculiventris group of species. In the writer’s collection are three
adults taken in the field in N.S.W., one female at Avalon in Feb., 1948, one male taken
at Woolwich in March, 1950, and a male taken at Roseville in March, 1953. It is also
known from Barrington Tops and the Dorrigo plateau at elevations of 4,000—5,000 feet.

Two larvae were found at Roseville, at one end of an area of about an acre,
surrounded by houses, where large trees had been left growing, and where at that time
weeds and decaying vegetation covered the ground. They were found on a steep slope
at the higher end of this area. One larva was found in August, 1952, when soil was
being turned over with a hand trowel; later it pupated, and a female emerged in
Feb., 1953. It was identified as S. vicina by Dr. I. M. Mackerras. The second larva
was found in March, 1953, under a raked-up heap of decaying leaves and grass. During
March and April it fed upon the Lepidopterous pupae supplied, but after that various
pupae and larvae were left untouched, except for a Tipulid larva which was sucked
out in August. In December the larva appeared to have decreased a little in size,

ip

220 MORPHOLOGY AND BIOLOGY OF SCAPTIA VICINA,

but this may have been caused by the soil in the jar becoming too dry and being left
dry for some weeks. It pupated in January, and a female with one wing slightly
deformed emerged in February. Eggs were not found.

Larva (Text-figs. 1-11).

The larva (Text-figs. 1-2) is large, stout, circular in cross-section, and, when fully
extended, it tapers very much anteriorly to the small head and very slightly to the
truncated posterior end. A full-grown larva, when extended, may be about 30 mm.
long and about 9 mm. across the widest part. When touched or otherwise disturbed
it will contract very much, and may then be about 17 mm. long and about 11 mm. wide,
and it appears to be short, cylindrical, and truncated at both ends. Contraction is
effected largely by the telescoping of the tapering anterior segments, with relatively
little shortening of the posterior segments.

The integument is coarsely striated all over, except on the anterior collar of the
prothorax, the small spiracular area, the anal tubercle, and very narrow areas between
the segments. Beneath the striated integument is a layer containing a pale brown
pigment in an irregular network pattern (Text-fig. 2). The pigment is much darker in
the middle line of both dorsal and ventral surfaces in the posterior half of each
segment, and the body colour is cream in the non-pigmented areas, so the living larva
appears mottled, with a central, dark, longitudinal, broken, double line on the dorsal
and ventral surfaces. Graber’s organ cannot be seen in the living larva, owing to the
pigment obscuring it. The pigment is not present in larval exuviae.

Head.—The head can be completely withdrawn, but the antennae, palps and tips
of the mandibles project when the larva is extended at all. The mandibles are large
and quite black, and the external openings of the mandibular canals can be seen
under the binocular magnifier with magnification x10.

The antennae (Text-fig. 3) are large. The basal segment is about 1 mm. long, with
a strong chitinous internal structure and a thick layer of integument, and it bears
two small, pointed, sensory processes on one side near the distal end. The second
segment is about 0:3 mm. long, with a chitinous internal structure similar to the
first segment, and a very thin integument. The third segment is bifid, with both
branches slender, tapering, pointed and about equal in length.

Mouth-parts.—The mandibles (Text-fig. 4) are large, almost square at the tip, with
very slight serrations on the lower edge, and so heavily chitinized that the mandibular
canal is not visible in a slide of the last larval exuviae mounted in balsam. The
maxillae (Text-fig. 4) are of clear chitin, with a wide basal portion and a long
narrow distal blade. The anterior edge of the base is armed with rows of long, slender,
pointed hairs, and the proximal half of the lower edge of the blade is similarly armed.
The maxillary palp is three-segmented; it has a short, thick, basal segment, a long
cylindrical second segment, and a long, slender, pointed third segment, which is nearly
half the length of the second segment.

The labrum (Text-fig. 5) is blade-like, about three times as long as high, mainly
of clear chitin, with some dark longitudinal bands of heavier chitin. ‘there are several
pairs of sensory organs placed midway along the down-curved dorsal surface; further
forward is a pair of strong setae; beyond these is a deep indentation; and the labrum
ends in a narrow, upturned, distal portion bearing a pair of slender setae. On the
latero-ventral surface is an extensive area covered with papillae and bordered dorsally
with long-pointed hairs or bristles. The labium (Text-fig. 5) ends in a thick, fleshy,
pointed, tongue-like structure covered with fine pointed hairs. It is attached to a
heavily chitinized pharynx support, and salivary ducts run back from it to the large
salivary pump. On its ventral surface is a pair of very small, forwardly directed
labial palps set in an area covered with hairs.

Except for minor differences, and the fact that it is much larger and more heavily
chitinized, the head capsule is very similar to that of S. awriflua (Don.) described by
Fuller (1936) and to the new species described in this paper.

BY KATHLEEN M. I. ENGLISH. 221

Thorax (Text-fig. 6)—The prothorax is encircled anteriorly with a wide collar, or
annulus, which narrows abruptly for attachment to the head capsule. It is covered
with more or less closely set scales, each armed on the posterior edge with backwardly
directed spines. On the anterior half of this prothoracic annulus, each scale bears a
few long, strong spines (Text-fig. 7); on its posterior half the scales are further apart,
and each bears numerous fine spines (Text-fig. 8). There is a difference in colour
in these two areas in the living larva, the anterior part being colourless and the
posterior part mottled light brown. The whole annulus can be withdrawn.

Each thoracic segment bears two groups of at least four long, strong, chitinous setae
on the ventral surface, one group on each side of the middle line. There are also some
hairs on each segment. The prothorax bears at least 14 hairs, 4 large and 4 small on
dorsal surface, 2 large on ventral, and 2 large on each lateral surface; the second and
third segments each bear 6 hairs. There is no pubescent annulus on the anterior border
of the second and third segments, but there are a few very small areas of brownish
rough integument on each.

The openings of the anterior spiracles were not found.

Abdomen.—On segments 1-7 are very small paired, dorsal and ventral pseudopodia,
and two lateral pseudopodia, one above the other, and behind each is a swelling, which
extends longitudinally almost the length of the segment and is very noticeable when
the larva is moving. lLaterally on each segment are three long hairs visible with a
magnification x20. Each segment also bears two small hairs on the dorsal surface and
two minute hairs on the ventral surface, visible only in slide mounts at a magnification
x60; on segments 3-5 at least there are in addition two smaller hairs. There is an
incomplete circlet of small areas of rough, brownish integument at the anterior border
of segments 1-8.

The eighth segment (Text-figs. 9-10) is abruptly truncated, and there is no siphon.
At the posterior end are two rounded dorsal tubercles, each with one long hair, on
each side is an elongated tubercle bearing two long hairs, and between these is the
flat spiracular area surrounding the posterior spiracle. The large anal tubercle, with
unstriated rugose integument, is situated on the ventral posterior surface of the
segment.

The posterior spiracle (Text-fig. 11) is of typicai Tabanid form. It lies in a vertical
slit in the spiracular area and protrudes very little, if at all. The spiracular area is.
roughly triangular in shape, with more or less vertical striations in the integument,
and it bears several hairs. On each side, just above the spiracle, is a pair so close
together that they look like one hair, while on each side below the spiracle are three
pointed hairs, one long and two short, and also one hair with a rounded tip: The
external opening of Graber’s organ is at the apex of the area above the spiracle.

Pupa (Text-figs. 12-14).

The head and thorax in the living pupa are of pale brown chitin, except the
spiracular tubercles which are dark brown; the chitin of the abdomen is very dark
brown.

The larger of the two pupal exuviae measures 30 mm. long and about 7 mm. across
the thorax. The head and thorax (Text-figs. 12-13) are armed with very fine, short
setae, which may be seen with magnification x10; they are placed directly on the
chitinous surface and are not raised on mound or tubercle. On the head are one pair
each of frontal and vertical setae, anterior and posterior orbital setae, and two pairs
of lateral orbital setae. The antennae are short, and they rise from low, broad,
antennal tubercles which almost meet in the centre line. Above is a low, broad, frontal
tubercle, and below is the carinate tubercle; together these form a slightly raised,
furrowed and ridged area. The sheath of the proboscis is very elongated (Text-fig. 13).
It is produced well beyond the sheaths of the maxillary palps, and extends to the
median junction of the sheaths of the first tibiae.

The thorax bears four pairs of very slender setae, a basal alar seta on each side,
and three pairs of setae on the dorsum, the anterior, posterior and median mesonotal

222 MORPHOLOGY AND BIOLOGY OF SCAPTIA VICINA,

External Pore of
Grabers Ogan7

i

vi

133
oy Sheath of

Ton
8 ALO Max. Pp

| f q Lentra/ ee cs
/} ii Seae. a i=

ve
Aymulus.

Posterior
10 1 Soiratle

4 Mawary Fenp
Maia.

Labiim |p

Text-figures 1-14. Scaptia vicina (Taylor).

1-11, Larva.—i, partly extended, dorsal view, x 3 approx.; 2, showing pigment pattern,
x 3 approx.; 3, antenna, x 25, approx.; 4, mandible and maxilla, x 25 approx.; 5, labrum and
labium, x 25 approx.; 6, thorax, lateral view, x 5 approx.; 7, scales with spines from anterior
part of annulus, x 250 approx.; 8, scale with spines from posterior part of annulus, x 250
-approx.; 9, 8th abdominal segment, lateral view, x 5 approx.; 10, 8th abdominal segment, end
view, x 10 approx.; 11, posterior spiracle and spiracular area, x 70 approx.

12-14, Pupa.—12, lateral view, x 3 approx.; 13, head and thorax, ventral view, x 3 approx. :
14, 8th abdominal segment, end view, x 8 approx.

BY KATHLEEN M. I. ENGLISH. 223

setae. These dorsal setae are very small and very pale, and not easy to distinguish,
as they are not raised on tubercles. Each thoracic spiracle is marked by a double
tubercle, consisting of a prominent, rounded mound with a slightly furrowed surface
placed adjacent to the medial aperture, and another mound with a furrowed sloping
surface edged by a narrow dark ridge. The rima of the spiracle is very small.

The metathorax bears three pairs of very slender setae.

Abdomen.—The abdominal spiracles are roughly circular in shape, flattened, and
very slightly elevated. The first segment bears two pairs of setae on the dorsum.
Segments 2-7 each bear a girdle of slender setae and heavy spines. On the dorsal
surface there are numerous short heavy spines and about eight long slender setae on
segment 2; the number of heavy spines is reduced in succeeding segments, until on
segment 7 there are four or six heavy spines and ten long setae. On the ventral
surface there are about twelve long, widely spaced setae on segment 2, and about
the same number close together on segment 7. Laterally there are about ten long
setae on each side of each segment. On segment 8 (Text-figs. 12-14) there is an
incomplete circlet of long setae, there being none on the dorsal surface; the setae
are very long dorsolaterally, and they decrease in length towards the ventral surface,
which is bare in the centre. The segment terminates in an aster of two long dorsal
projections, with a small knob placed at the base of each. An aster of this type was
described by Fuller (1936) for Scaptia auwriflua (Don.).

SCAPTIA MUSCULA. n. sp. (Text-figs. 15-34).

A grey, hairy, unadorned species, which belongs structurally to the violacea group
(Mackerras, unpublished), but because of its small size, rounded shape and generai
erey colour it cannot be confused with any other described member of the genus. The
flies rather resemble blowflies in general shape, with more or less rounded thorax and
abdomen. The sexes are very similar in colour and size; if anything, the males are
slightly larger than the females. Average length, excluding antennae, 10 mm., average
width across thorax at wing base 3-5 mm.; length of wing 9 mm. The eyes have
green and bronze lights in living specimens. '

The holotype 92, allotype g, two 9 and four ¢ paratypes were all reared from larvae
and one pupa collected in New South Wales within 60 miles of Sydney. The types are
in the Macleay Museum, University of Sydney.

The flies were submitted for identification to Dr. I. M. Mackerras, who considered
them to belong to a new species; he submitted them to Mr. H. Oldroyd, of the British
Museum (Natural History), who agreed with the determination.

Occurrence.

Four adults only of this species have been collected in the field, all in N.S.W.:
Woy Woy, 5 Oct., 1924, Nicholson, 2¢¢; Oatley, Oct., 1924, 1 g, in the coilection of
The School of Public Health and Tropical Medicine, University of Sydney; and
Chatswood, 21 Oct., 1952, 1 9, collected by the writer.

Larvae were found first by Mr. J. Burden, when collecting larvae of Myrmeleontidae
in the deposits of fine sand that occur beneath overhanging sandstone rocks. The pits
made by the ant-lion larvae are to be found even in very small sand patches, and even
where the overhang is very small or very low; but most of the Scaptia larvae have
been found where the overhanging ledge was five feet or more from the sandy floor,
though at Woodford larvae were found at the back of a small cave, where the roof
was iess than three feet high, but where there was quite an extensive area of sandy
floor. Six larvae were collected by Mr. Burden in 1949 and 1950, and 27 have since
been found by the writer; fourteen at Roseville, four at Woodford on the Blue Mts.,
and others in various localities in the vicinity of Sydney. Some of the larvae were
killed and preserved, some pupated, and eight adults emerged., Fifteen larvae still
remained alive in June, 1954, of which three had been collected in October, 1952. The
larvae had fed fairly readily on various Lepidopterous pupae and latterly on the
Ephestia pupae.

224 MORPHOLOGY AND BIOLOGY OF SCAPTIA VICINA,

19 PEE 16

17

Aedaegus.
21 \ Sipe.
t
Conifer”

Ventral
26 ix Se7ae

ee
a
ry

1
\iaal

if oo

32

Antenna a
AIepop en qty, yiay

Clee

Posterior
\ Sprracle

Text-figures 15-34. Scaptia muscula, n. sp.

15-21, Imago.—15, dorsal view, x 4 approx.; 16, head, lateral view, x 6 approx.; 17, frons,
<x 12 approx.; 18, antenna, x 20, approx.; 19, 2 maxillary palp, x 20 approx.; 20, & maxillary
palp, x 20 approx.; 21, ¢ hypopygium, x 20 approx.

22-31, Larva.—22, dorsal view, x 4 approx.; 23, antenna, x 45 approx.; 24, mandible and
miaxilla, x 45 approx.; 25, labrum and labium, x 45 approx.; 26, thorax, lateral view, x 8 approx. ;
27, armature of anterior part of annulus, x 250 approx.; 28, armature of posterior part of
annulus, x 250 approx.; 29, abdomen, segments 7 and 8, lateral view, x 8 approx.; 30, abdomen,
segment 8, end view, x 20 approx.; 31, posterior spiracle and spiracular area, x 150 approx.

32-34, Pupa.—32, lateral view, x 4 approx.; 33, head and thorax, ventral view, x 4 approx. ;
34, 8th segment of abdomen, end view, x 15 approx.

BY KATHLEEN M. I. ENGLISH. 225

After emerging, some of these Scaptia were kept alive up to six days in small jars,
in which they fed readily on moistened raisins. The jars. were kept covered with dark
material in an endeavour to keep the flies from battering their wings on the sides.
These flies at times emitted a high-pitched hum, quite audible a few feet away, and
apparently the sound was not due to vibration of the wings in any way.

Imago (Text-figs. 15-21).

Female (Text-figs. 15-16).—Head a little wider than thorax. Eyes covered with
light buff coloured hairs. Frons (Text-fig. 17) diverging slightly posteriorly and
anteriorly, and converging slightly near centre and at the anterior angle, with light
olive-grey pollinose covering and short light buff hairs. Subcallus moderately pro-
duced, with light olive-grey pollinose covering and without hairs; parafacials with
similar pollinose cover and long pale hairs. Beard of light buff and warm buff very
long hairs. Face prominent, with deep olive-grey pollinose covering, and with long
warm buif hairs at sides, sparse short hairs on upper centre and no hairs on lower
centre. Antennae (Text-fig. 18) length about half height of head; first segment,
viewed laterally, about as wide as long; second segment about same width, but very
short; both with capucine buff pollinose covering and long black and light buff hairs;
third segment subulate, 8-annulate, orange or vrange-buff in colour. Palpi about one-
fourth as long as shaft of proboscis; first segment mouse-grey, with long silvery and
tawny hairs; second segment (Text-fig. 19) as figured, flattened, orange-buff in colour,
with sparse pale hairs on lower edge and black hairs partly surrounding a lateral
bare area. Proboscis a little longer than height of head, moderately slender, with
expanded, only partly chitinized labella.

Thorax—Dorsum covered with deep olive-grey tomentum, with narrow light grey
stripes and tawny recumbent hairs, with light buff hairs above wing base. Pleural
hairs white, and much longer and thicker than on dorsum. Halteres chestnut-brown.

Abdomen a deep olive-grey pollinose covering, with shining orange-buff recumbent
hairs on tergites and shining light buff recumbent hairs on sternites. Cerci tawny.

Legs orange-buff, with long light buff hairs on femora, shorter hairs, mostly black
but some tawny, on tibiae, and short recumbent hairs on the tarsi, black on the dorsal
surface and tawny on the ventral surface.

Wings clear, with a smail but definite dresden-brown stigma between R, and R.;
cell R; open, but somewhat narrowed.

Male.—Very similar to female, except that it is more heavily clothed with longer
hairs, and there are slight differences in colour. Hyes contiguous, upper facets not
enlarged. Sides of face liver-brown in colour. The first segment of the palp (Text-
fig. 20) is bulbous, liver-brown in colour, and clothed with long silvery hairs; the
second segment is slender and cylindrical, liver-brown, with long silvery hairs, except
the tip, which is pale orange-yellow, slightly flattened, and bare except for some hairs
on the edges.

The hypopygium (Text-fig. 21) has a long, rather narrow aedeagus and a simple
pointed style.

The eight specimens examined are remarkably uniform in appearance, but there
are slight variations in the colour of the tomentum and in the length and colour of the
hairs. In two specimens, a male and a female, the dorsum of the thorax is much
darker, as the pollinose covering is lacking on a large part of it, and there are
numerous black upright hairs as well as tawny hairs. The absence of the pollinose
covering is probably not due to rubbing, as the hairy covering is intact.

Eggs were not found.

The colours in this description have been identified as accurately as possible with
the aid of Ridgeway’s Colour Chart.

Larva (Text-figs. 22-31).
The iarvae found varied in length, when contracted, from 7 mm. to about 16 mm.,
and up to 6 mm. in width. When the larva contracts, the head is completely withdrawn,
the thoracic segments are partially telescoped, and the larva appears to be truncated

226 MORPHOLOGY AND BIOLOGY OF SCAPTIA VICINA,

at both ends. When it extends, it tapers gradually anteriorly to the very small head,
and is widest at the fourth and fifth abdominal segments. The body of the larva appears
to be slightly flattened, as the large, elongated swellings of the lateral pseudopods make
it appear broader than high; but preserved specimens appear more cylindrical, because
the lateral swellings have contracted.

The integument has fine, longitudinal striations, is transparent and glass-like, and
there is a layer containing pigment beneath it. There is an irregular pattern of this
purplish-brown pigment (Text-fig. 22) on the dorsal and ventral surfaces of each
segment except the prothorax; it varies somewhat in different larvae, but the general
appearance is similar. The text-figure shows the pattern of a partly contracted larva,
when the lines of pigment appear broad; but, when the larva extends, the lines of
pigment lengthen and become thinner, and the unpigmented area is more extensive.
The pattern remains visible in carefully preserved spirit specimens, and may also be
preserved in slides, if the abdominal contents of the larva are removed but the muscles
attached to the body wall left undisturbed as much as possible. The pigmented layer
is not shed with the larval exuviae, but remains in the pupa, where it is distinct and
complete in the abdomen, and there are faint traces of it in the meso- and metathorax.
The integument is not transparent enough, when dry, for the eye-spots and Graber’s
organ to be seen. If, however, the larva is placed in water and the dust gently washed
off, then, whilst it is still wet, the black bodies of Graber’s organ can just be seen
and the eye-spots are large and conspicuous.

Head.—tThe head capsule is long and slender, up to nearly 4 miu. in length and
0-75 mm. broad.

The antennae (Text-fig. 23) appear to be three-segmented. The first segment is
0-47 mm. long, with a definite internal chitinous structure, and two small sense organs
on the anterior dorsal area; the second segment is short, 0-1 mm. in length, slender,
and with an internal chitinous structure similar to the first segment; it bears the bifid
terminal segment, which has both branches slender, tapering and pointed, and one
much longer than the other.

Mouth Parts—The mandibles (Text-fig. 24) are almost square at the tip, and only
very slightly serrated on the lower margin; they are heavily chitinized, but the
mandibular canal can be seen in slide mounts. The maxillae are of clear chitin, each
consisting of a long, narrow, blade-like distal part, and a wide basa! part armed on
the anterior margin with long, pointed hairs. The maxillary palp is three-segmented;
the first segment short; second long, with an internal chitinous structure similar to
that in the antenna; the third long, slender and tapering to a point.

The labrum (Text-fig. 25) consists mainly of clear chitin. On the dorsal surface
are several sensory pits, nearer the apex is a pair of strong setae, beyond these is a
deep indentation, and the labium ends in an upturned portion bearing a pair of slender
setae. The ventral surface is covered with papillae bordered laterally with long pointed
hairs or setae. The labium is a bi-lobed, tongue-like structure, densely covered with
hairs. It is attached to the heavily chitinized pharynx support; salivary ducts run
back to the large salivary pump; and there is a pair of labial palps on the ventral
surface. The labium is very similar to that of S. vicina Tay. deseribed in this paper
and S. auriflua (Don.) described by Fuller (1936).

Thorax (Text-fig. 26).—The prothorax is encircled anteriorly with a wide collar
or annulus armed with spines. The anterior part bears longitudinal bands of thickened
epithelium, with short backwardly directed spines (Text-fig. 27); these bands gradually
merge into separated scales which cover the posterior part, each scale (Text-fig. 28)
being armed on the posterior border with three to five short spines. The appearance
of scales is scarcely evident unless the exuviae are stained before mounting, but the
spines are to be seen in unstained mounts. Each thoracic segment bears two groups
of setae on the ventral surface, one on each side of the middle line, each group
consisting of one long, strong seta, which can be seen in preserved whole larvae with
magnification x10, and one or more very fine short setae, which can be seen only in

bo
bo
=)

BY KATHLEEN M. I. ENGLISH.

slide mounts with high power. There are also irregular rows or circlets of single
hairs on each thoracic segment: sixteen or eighteen small hairs in three rows on the
prothorax, about ten larger hairs in two rows on the mesothorax, and about eight large
hairs in two rows on the metathorax. These-hairs are mostly on the dorsal and lateral
surfaces, and some can be seen in spirit specimens with magnification x30.

Abdomen.—The integument is thickened and folded to form pseudopodia near the
anterior border of segments 1-7. On the dorsal surface of each segment are paired
pseudopods meeting in the centre and forming a low transverse ridge; on the ventral
surface is a slightly thicker pair, which do not meet in the centre; and on each side
is a small dorsolateral pseudopod and a larger, more prominent lateral one. There is
also a series of irregular folds or ridges in the integument, and behind the larger
pseudopod is a distinct swelling extending almost to the posterior edge of the segment;
it is this structure that gives the appearance of width to the live larva. On this
swelling are two long hairs, one above the other, which can be seen in spirit specimens
with a magnification x30, and these, together with four or six small hairs, form an
irregular circlet on each segment.

The eighth segment (Text-figs. 29 and 30) is abruptly truncated. It bears dorsally
two low, broad, pointed processes, with one long hair set below the apex on each;
laterally on each side is a long, low process bearing two long hairs; on the ventral
surface is the anus surrounded by the prominent folds of the anal tubercle; and on the
posterior surface is the posterior spiracle set in a flat almost triangular spiracular area.
The spiracle (Text-fig. 31) is of typical Tabanid form, and there is no siphon. There
are fine hairs on the spiracular areas visible only on slide mounts with high power,
ohne on each side just above the spiracle, and four on each side below it.

Pupa (Text-figs. 32-34).

Hight pupal exuviae were obtained from reared specimens. One female pupa is
approximately 18 mm. long and 4 mm. wide on thorax. The pupal skin is of very thin,
transparent chitin, which does not become dark in colour, and this permits the larval
pigment to be seen.

The head and thorax (Text-figs. 32-33) are armed with very small, slender setae,
which can be seen with a hand lens only with difficulty. The head bears six pairs of
small setae. The sheath of the proboscis is very long, and extends to the sheath of the
first pair of legs, as in S. vicina. The thorax bears a basal alar seta on each side, and
three pairs of very small setae on the dorsum, but the central pair may be hidden in
exuviae by the folding at the suture. The thoracic spiracle is on a definite mound, but
the C-shaped rima is so small that it is almost indistinguishable. The metathorax bears
three pairs of fine setae.

Abdomen (Text-fig. 32).—Abdominal segments 1-7 are divided by longitudinal lines
into dorsal, ventral and lateral regions. The first segment bears a small spiracle on
each lateral region, and two pairs of very small spines on the dorsum. Segments 2-7
bear a small spiracle on each lateral region, and a girdle of spines in a single row
about the middle of each segment. On the dorsal surface of segments 2-6 are six to
eight large spines flanked on each side by minute spines; on segment 7 there are only
four or five large spines, and there may be no minute spines. On the ventral surface
of each segment is a row of very small spines, and on segments 6 and 7 there are also
two or four larger spines. The spines on the lateral areas vary considerably in number
and size in different specimens. There may be four or six, some long, some short, and
they are more or less short on segment 2 and much longer on segment 7, but they do
not increase in size with any apparent regularity as they progress backwards. The last
segment (Text-fig. 34) bears an incomplete girdle of spines; the centre of the dorsum
is bare, and the dorso-lateral spines are more or less larger than the ventro-lateral ones.
In the male pupa there is a ventral row of small spines; in the female the centre of
the ventral surface is bare. The segment terminates in an aster of two tubercles each
bearing a single strong spine. The terminal spines are much longer than those figured
for S. aurifiua (Don.) by Fuller (1936).

bo
bo
co

MORPHOLOGY AND BIOLOGY OF SCAPTIA VICINA,

CONCLUSION.

Fuller (1936) gave some characters of the larvae and pupae of Scaptia and Goniops,
and suggested that they might prove to be distinguishing characters of the subfamily
Pangoniinae. The larval characters could be observed in living or spirit specimens.
Other characters, which can be seen in slide mounts of exuviae of larvae of Scaptia
and Hctenopsis, are now suggested as possibly distinguishing Australian Pangoniinae
at least:

Larva—Annulus of prothorax partly or wholly covered with scales armed with
backwardly directed spines. Maxilla with long, slender, blade-like distal part.

For the pupa, Fuller gave “The aster is reduced to two large projections at the
end of the terminal segment’. This character does not hold for Hctenopsis (English,
1952), so an addition is suggested:

Pupa—lf the terminal segment has six projections, then there are three pairs of
strong dorsal setae on the thorax.

These setae are strong in Hctenopsis. Some Scaptia pupae, if not all, also have three
pairs of dorsal setae, but they are so small and slender that they are difficult to detect,
and if, as so often happens, they are broken off, their position cannot be located, as
they have no basal tubercles.

On examining comparable characters of the available larvae and pupae of Australian
species of other groups, it was found that in the larvae (Tabanus two spp., Dasybasis
four spp., Lilaea one sp.) the annulus of the prothorax is densely covered all over
with short hairs or spines, and the maxilla is broad with a very short terminal portion;
in the pupae (Tabanus four spp., Dasybasis five spp.) the aster has six projections,
and there are two pairs of dorsal setae on the thorax. The aster is also described as
having six projections in Haematopota and Chrysops.

The larval habitat of S. muscula (clean sand under rock ledges, frequently very
dry) is very different from that of S. vicina and S. awriflua, and these differ from one
another, as the soil where S. vicina was found was too dry for Tipulid larvae, which
were numerous for at least part of the year in the soil where S. auriflua was found.

The two pupae described in this paper have the sheath of the proboscis very
elongated, but this does not occur in all Scaptia pupae. The writer was able to
examine pupae of two species of Scaptia from the collection of the Division of
Entomology, C.S.1.R.O., Canberra, and in these the sheath of the proboscis is short.
The length is presumably to be correlated with the length of the proboscis in the adult.

The material used in the preparation of this paper, i.e., the adult flies with pupal
exuviae, and slide mounts of larval exuviae of both species, together with slide mounts
of larvae and larvae in spirit of S. muscula, have been deposited in the Macleay Museum
at the University of Sydney.

Acknowledgements.

The writer is indebted to Professor P. D. F. Murray and Dr. A. R. Woodhill,
Department of Zoology, University of Sydney, who made available laboratory accom-
modation at the Department; to Dr. A. J. Nicholson, of the Division of Entomology,
C.S.1.R.O., Canberra, for the loan of specimens; to Dr. D. F. Waterhouse, of the same
Division, for advice and help on the problem of feeding larvae; and to Dr. I. M.
Mackerras, Queensland Institute of Medical Research, Brisbane, who identified the
adults and gave much helpful advice on the preparation of this paper.

References

MNGLISH, K. M. I., 1952.—Notes on the morphology and biology of Hctenopsis vulpecula Wied.
var. angusta Macq. (Diptera, Tabanidae, Pangoniinae). Proc. Linn. Soc. N.S.W.,
Ixxviil : 270-274.

FERGUSON, E. W., 1926.—Additional notes on the nomenclature of Australian Tabanidae. Bull.
Ent. Res., xvi: 293-306.

FULLER, M. E., 1936.—Notes on the biology of Scaptia auriflua Don. (Diptera, Tabanidae).
Proc. LINN. Soc. N.S.W., Ixi: 1-9.

McATEE, W. L., 1911.—Facts in the life history of Goniops chrysocoma (Dipt., Tabanidae).
Proc. Ent. Soc. Washington, xiii: 21-29.

MACKERRAS, I. M. (to be published).—The Tabanidae (Diptera) of Australia.

BY KATHLEEN M. I. ENGLISH. 229

MauwocuH, J. R., 1917.—A preliminary classification of Diptera exclusive of Pupipara, based on
larval and pupal characters, with keys to imagines in certain families. Part 1. Bull. Ill. Lab.
Nat. Hist., xii: 355-357.

RIDGEWAY, R., 1912.—Color Standards and Color Nomenclature. Washington.

ScHwarptT, H. H., 1934.—Biological Notes on Goniops chrysocoma (O.S.) (Dipt., Tabanidae).
J. Kansas Ent. Soc., McPherson, vii: 73-79.

STONE, A., 1930.—The bionomics of some Tabanidae (Diptera). Ann. Ent. Soc. America,
Xxiii: 261-304.

"‘PAYLOoR, FE. H., 1918.—Studies in Australian Tabanidae. Rec. Aust. Mus., xii: 57.

WALTON, W. R., 1908.—Notes on the egg and larva of Goniops chrysocoma O.S. Ent. News,
xix : 464-465.

A NEW GENUS AND SPECIES OF THE TRIBE LABENINI FROM AUSTRALIA
(PIMPLINAEH, ICHNEUMONIDAE).

By ArtHur W. Parrorr, Wakapuaka Road, Nelson, New Zealand.
(One Text-figure.)

[Read 24th November, 1954.]

Synopsis.

A new genus and species of ichneumon-wasp of the Sub-family Pimplinae are described.
The new genus, Neonotus, is placed in the Tribe Labenini as defined by Cushman and Rohwer
(1920). A generic diagnosis is given together with a key separating it from the closely allied
genus Certonotus Kriechbaumer. The type of this new genus, Neonotus, is N. chadwickii,
which was bred from a species of Buprestid, Hthon affine L. & G., by Mr. C. E. Chadwick,
Systematic Entomologist, Department of Agriculture, Sydney.

A small collection of reared Ichneumonidae’ from known hosts was recently
forwarded to me by Mr. C. E. Chadwick, Entomologist, N.S.W. Department of Agri-
culture, Sydney. Included in this collection were nine specimens of an undescribed
species, which represents a new genus closely allied to Certonotus Kriechbaumer. As
this new species was bred from the Buprestid beetle Hthon affine L. & G., it is deemed
advisable to give immediate diagnosis of this new form, so that it may be identified
and studied by local entomologists.

Tribe LABENINI.

The new genus falls into this tribe as defined by Cushman and Rohwer (1920)
in their revision of the Pimplinae. The Labenini is at present represented in Australia
by the genus Certonotus Kriechbaumer; the present genus is now added, to which
I have given the name Neonotus.

Genus NEONOTUS, gen. nov.

The following combination of characters will serve to distinguish this genus from
other genera of the Labenini:

Labium not exserted; malar space obsolete; antennae sub-clavate, mesonotum
elongated, not transversely rugulose or carinate, smooth and finely punctate, notauli
present; propodeum declivous posteriorly, aerolated; abdomen smooth and shining,
with a few large punctures scattered over the surface of the tergites; spiracles of first
abdominal tergite situated at, or slightly basad to, the middle of the segment; ovipositor
long, about as long as the abdomen; recurrent vein of forewing not strongly bent
inwards with the aerolet large, longer than high and sessile on the radius.

This new genus is allied to Certonotus but differs from that genus in many
important details. It may be separated as follows:

Malar space at least more than half width of mandibles at base, usually considerably wider ;

mesonotum transversely rugulose or carinate; scutellum planate ...... Certonotus Krb.
Malar space obsclete ; mesonotum smooth, finely punctate, without transverse carinae; scutellum
COMVEX lata Ns als gn mente o eikemedsokenes AUC eeise nuk MeO eae carer Reem ee Bee Ueasne nL Neonotus, gen. nov.

NEONOTUS CHADWICKII, Sp. nov.

Female: 10 mm. long; antennae 9 mm.; ovipositor 5 mm.

Colour black with creamy-white markings, head black with inner and outer orbits
white, interrupted dorsally; face white with median black area; mandibles, scape, and
antennae black, with segments 23 to 29 white; mesonotum’ medially brownish-black,
with the anterior lateral borders with a narrow white band; propleurae, mesopleurae,
metapleurae and sternum mainly dark brown with the sub-alar tubercles white;
scutellum black with apical third and metascutellum white; anterior and middle coxae

BY ARTHUR W. PARROTT. 231

mainly whitish, posterior coxae blackish with the apex white; dorsal and ventral
surfaces of femora and tibiae mainly whitish shaded with dark brown; middle legs
whitish, considerably clouded with brown, the apical tarsal joints and claws entirely
brown; posterior legs mainly dark brown, except trochanters dorsally, apex of femora,
tibial spurs and basal half of metatarsal joint which are whitish; abdomen black
with all segments widely margined posteriorly with white; ovipositor and sheaths
mainly dark brown or black.

Face slightly raised in centre and faintly rugose, with an obsolete, longitudinal
median furrow: clypeus has three or four large punctures and is clothed with long
whitish hairs; anterior margin gently rounded; internal orbits emarginate opposite
antennae and sub-parallel below; cheeks about half the diameter of eye in profile,
shining, almost impunctate, except for a few minute widely spaced punctures; mandibles
wide at base, narrowing towards apex, which is very narrow, with two fine teeth, the
upper slightly thicker and longer than lower, the entire margins of mandibles swollen;
antennae with 31 segments, first segment about twice as long as second, the following
segments decreasing slightly in length to about the eighteenth, the remaining segments
being subequal in length, about as wide as long, the apical segment elongated; upper
angles of pronotum not produced; mesonotum elongate, punctate, not transversely
rugose; notauli faintly marked, although the median lobe is somewhat prominent, with
an obsolete median depression; scutellum convex, declivous posteriorly without lateral
carinae; scutellum fovea deep and smooth; propodeum areolated, all areas plainly
indicated, basal area subquadrate, areolar area wider than long; spiracles elongate oval;
propleurae very finely punctate, mesopleurae and metapleurae similarly punctate and
clothed with short white pubescence; mesopleurae with a glabrous area adjacent to
posterior border with a short deep longitudinal groove; prepectal carina well developed
and sinuated; posterior coxae elongated, nearly twice as long as thick, and on dorsal
surface very finely and sparsely punctate; anterior tibiae subequal in length to femora
with outer side obliquely grooved on basal half; tibial spurs about half the length of
tibiae; spiracles of first abdominal tergite situated slightly before the middle of the
segment; abdomen elongated, compressed posteriorly from the fifth tergite; all abdominal
tergites smooth, with indications of longitudinal aciculations; ovipositor sheaths clothed
with short thick black pubescence. Forewing with nervulus interstitial with basal vein
and reclivous; first and second abscessae of cubitus subequal, slightly longer than
second abscessa of radius; recurrent vein distinctly bent inwards, with a long fenestra
in lower part of upper half; subdiscoidal originating below the middle of the transverse
portion of the discoideus; third abscessa of radius straight and longer than first
abscessa, which originates before the middle of the stigma; discocubital nearly straight
but somewhat faintly sinuated about the middle.

Posterior wings with abscissula shorter than the intercubitella; mediella curved
at base, longer than the first abscessa of cubitella; nervellus not broken and reclivous.
Fore and hind wings are illustrated in Figure 1.

Male: In all essentials agrees with the female but the apical fourth and fifth
segments and the face entirely white; the ocelli are slightly larger in the males and
the anterior tibiae are normally shaped.

Variation: The venation of the wings varies to some extent, as for instance the
relative lengths of the first and second intercubitus and the first and second abscessa
of cubitus. The number of white segments of the flagellum varies in different
individuals of both sexes.

Location of Types—The type material is in the collections indicated below. The
labels on the specimens refer to data in notes made by the collector.

Holotype 2 (d) HEmerged 20th March 1952. Mc.235. Department of Agriculture,
Sydney.

Allotype ¢ (h) Emerged 13th October 1952. Mc.343. Department of Agriculture,
Sydney.

232 NEW GENUS AND SPECIES OF LABENINI,

Paratypes: ¢ (e) Emerged September 1952. Mc.244. Division of Entomology,
C.S.LR.0., Canberra. ¢ (c) Emerged March 1952. Me.221. Australian Museum, Sydney.
3 (g) HEmerged September 1952. Mc.338. (Damaged.) Department of Agriculture,
Sydney. @ (i) Dead in gall, 21st March 1953. Mc.419. (Damaged.) Department of
Agriculture, Sydney. 9 (a) Emerged 3rd May 1951. Cawthron Institute, Nelson, New
Zealand. ¢ (vb) Emerged March 1952. (Head missing.) Cawthron Institute, Nelson,
New Zealand. 6 (f) Emerged March 1952. Mc.283. Cawthron Institute, Nelson, New
Zealand.

Text-fig. 1—Wing venation of Neonotus chadwickii Parrott.

Ali specimens were bred from galls of the beetle Hthon affine lL. & G. on Pultenaea
stipularis Sm. at Middlecove Point, Willoughby, Sydney.

Acknowledgement.

The author wishes to express his very sincere thanks to Mr. C. EH. Chadwick,
Systematic Entomologist, Department of Agriculture, Sydney, New South Wales, for
the opportunity of examining this interesting material. I have much pleasure in
naming this new species in honour of Mr. Chadwick.

Reference.

CUSHMAN, R. A., and RoHwer, S. A., 1920.—Holaretic Tribes of the Ichneumon-flies of the
subfamily Ichneumoninae (Pimplinae). Proc. U.S. Nat. Mus., 57, No. 23815: 379-396.

233

AUSTRALASIAN CERATOPOGONIDAE (DIPTERA, NEMATOCHRA).

Part VII2 Noves ON THE GENERA ALLUAUDOMYIA, CERATOPOGON, CULICOIDES
AND LASIOHELEA.

By Davip J. Ler, B.Sc., and Hric J. REYE,? M.B., B.S.
(Plate ix; 36 Text-figures.)
[Read 24th November, 1954.]

Synopsis.

The present paper records the occurrence of the genera Allwaudomyia and Ceratopogon
in Australia. Descriptions of seven new species of Culicoides are presented together with a
new name for one previously described and additional data concerning other species. A
discussion of the occurrence of Lasiecheleéa in Australia and the New Guinea area is also
included.

1. THE CERATOPOGON GROUP (Lee, 1948, p. 326).

No members of this group have previously been recorded from the Australian
mainland, and collecting up to the present does seem to indicate that the group is
rather poorly represented in this region. Two genera are now known to occur,
Alluaudomyia and Ceratopogon.

(a) Alluaudomyia.

Alluaudomyia Kieffer, J. J., 1913, Voyage Ch. Alluaud et R. Jeannel en Afrique
Orientale, Dipt., 1: 12; full synonymy in Wirth, W. W., 1952. Univ. of California Pub.
in Entomology, 9, No. 2: 194.

The generic diagnosis given by Wirth (1952) seems adequate for the Australian
species, which will be described elsewhere. Wirth’s diagnosis is ‘Eyes bare or hairy,
narrowly separated above. Antennae fifteen-segmented, last five segments elongated
(female). Wings with macrotrichia towards apex, often adorned with small black
spots; first anterior radial cell obliterated, second subequal to first, its apex with
veins thickened and ending at about half of wing length; media forked and petiolate.
Legs slender, without spines; claws of hind legs of female unequal, inner claws twice
as long as outer; empodium vestigial.”

This genus is known to occur in both Queensland and New South Wales.

(b) Ceratopogon.

Ceratopogon Meigen, J., 1803, Illiger’s Mag. Ins., 2: 261. = Helea Meigen, J., 1800,
Nouv. Class. des mouches a deux ailes, p. 18. For synonymy see Wirth, W. W., 1952,
loc. cit.

(Until the controversy concerning the Meigen names of 1800 and 1803 is resolved
we retain the name Ceratopogon since it is most familiar to Australian entomologists.)

The most recent generic diagnosis is that of Wirth (1952), from which the
following is largely taken. Body stout, hairs short and scanty. Eyes more or less
pubescent. Antennae with segments 3-10 rounded, 11-15 not very long; male antennae
with last three segments elongate. Humeral pits present. Legs rather slender, without

1 Continued from Vol. 77, p. 394, of these PROCEEDINGS.

2 Assisted by a research grant jointly contributed by the Commonwealth Science and
Industry Endowment Fund and the Commonwealth Scientific and Industrial Research
Organization.

234 AUSTRALASIAN CERATOPOGONIDAE. VII,

long bristles; fourth tarsal segment short but not cordiform; empodium very short or
absent; claws of female rather large, equal or slightly unequal. Wings broad, usually
milky, owing to absence of microtrichiae; macrotrichiae absent or confined to a few
at apical margin; costa reaching more or less beyond middle; two radial cells normally
present, about equal in size; cross-vein r—m slightly oblique; intercalary fork indistinct;
median fork with rather long stem or lower branch widely interrupted or absent; alula
and squame bare. Male genitalia not inverted; ninth tergite small; coxites swollen;
harpes separate.

The Australian species we have seen, from Sydney and northern New South Wales,
would fall into the genus Brachypogon (Kieffer, J. J., 1899. Bull. Soc. ent. France, p. 69;
genotype, Ceratopogon vitiosus Winnertz) now included in the synonymy of Ceratopogon
by most authors. :

We do not think that Ceratopogon is either common or widely represented in
Australia, as very few specimens have been included in our general collecting in
recent years.

2. THE GENUS CULICOIDES.
(a) New Species.
(i) Species in which the Distal Portion of the Second Radial Cell is Dark.
CULICOIDES CORONALIS, nN. sp.

Types: Holotype ? and three ? paratypes, all on slides in S.P.H.T.M.

Type Locality.—Prince of Wales I., North Queensland, 31:xii:1952, 2000-2100 hours,
on sandy beach (J. Menner).

Distinctive Characters—The indefiniteness of the wing pattern associated with
entirely dark radial cells means that C. coronalis could only be confused with
C. immaculatus, C. austropalpalis and C. subimmaculatus. Since the fourth segments
of the tarsi are not cordate it should not be misidentified as C. subimmaculatus. The
absence of a pale area over r—m, together with the form of the palp distinguishes the
present species from C. austropalpalis and from C. immaculatus, to which it comes
closest. It is distinct in the terminal segment of the palp and the interorbital space.

Description—From the type series. No pinned material available for coloration
details. Measurements from holotype and selected series comprising three paratypes.
(See Table 1.)

Female.

Legs uniformly light brown, halteres brownish. Wings (Plate ix, fig. 1) with no
pattern, only slightly pale areas below the radial cells and in intercalary fork.

Head: Eyes well separated (Text-fig. 1). Antennae (Text-fig. 17) with segments
3-10 cylindrical but not much longer than broad, 11-15 distinctly longer, giving definite
contrast between proximal and distal segments. Segment III of palp very distinctly
swollen on one side and with single round sensory pit in the distal half (Text-fig. 9),
segment IV very small, shorter than V. Mouthparts about equal in length to height
of head.

Thorax: Legs unmodified, tarsus IV subcylindrical, tibial comb of five spines.

Abdomen: Two fully-formed spherical spermathecae (Text-fig. 24). The chitinized
plate of the seventh abdominal segment in the form of a distally directed crown
(Text-fig. 32).

Mate.
This sex has not been taken.
Distribution.— QUEENSLAND: Only known from the type locality.

CULICOIDES WILLIWILLI, nN. sp.
Types: Holotype 9, allotype ¢ and six ? paratypes. All slide specimens in S.P.H.T.M.
except one paratype in each of C.S.I.R.O. and Q.I.M.R.
Type Locality.—Bundy, via Moree, New South Wales, 1:xi:1951, light trap (A. L.
Dyce). .

235

LEE AND ERIC J. REYE.

BY DAVID J.

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236 AUSDPRALASIAN CERATOPOGONIDAE. VII,

Distinctive Characters.—C. williwilli is, in most characters, close to C. nattaiensis,
but the wing is distinct from the latter and other species in the possession of two
rather boomerang-like elongated pale areas over M,,, and M., each partially framing a
marginal round pale spot.

Description.—From the type series and other specimens listed below. No pinned
material available for details of coloration. Measurements from holotype, and selected
series comprising four paratypes. Male measurements from allotype, one ¢ paratype
and a specimen from Texas. (See Table 1.)

Female.

Legs with preapical pale spots on the femora, subbasal ones on the tibiae. Halteres
pale. Wings (Plate ix, fig. 2) with very complex and distinctive pattern of strong
contrast. The large pale area in the intercalary fork is variable; it may enclose a
dark spot as in the holotype, or only partially enclose a dark spot running to the wing
margin, or be indented distally in various ways or even without any enclosed or
indented dark area.

Text-figures 1-$.—Interorbital space of various species (x 190 approx.): 1, C. coronalis.
2, C. williwilli. 3, C. nattaiensis. 4, C. bunrooensis. 5, C. moreensis. 6, OC. bundyensis.
7, C. waringi. 8, C. multimaculatus.

Figures 1, 3 and 5 are from holotypes, 2 is a specimen from Noonameena (425), 4 a

paratype (397), 6 a specimen from Texas (430), 7 a paratype (441), and 8 a specimen from
King Lake (411).
Text-figures 9-16.—Palp of various species (x190 approx.): 9, C. coronalis. lO, C:

williwilli. 11, C. nattaiensis. 12, C. bunrooensis. 13, C. moreensis. 14, C. bundyensis.
15, C. waringi. 16, C. multimaculatus.

Figures 9, 13 and 14 are from holotypes, 2 is a specimen from Texas (424), 11 a paratype
(436), 12 a paratype (397), 15 a paratype (441), and 16 a specimen from King Lake (411).

Head: The eyes are separated above and contiguous below (Text-fig. 2). Segments
3-10 of the antennae are cylindrical and not much longer than wide, 11-15 are
considerably more elongate (Text-fig. 18). Palp with third segment very much swollen,
a single round sensory pit on the distal half, IV and V very short (Text-fig. 10).
Mouthparts shorter than height of head.

Thorax: Legs unmodified, tarsus IV subcylindrical, tibial comb of four spines.

Abdomen: Two fully developed subspherical spermathecae, each with a distinct
duct and one very small spermatheca (Text-fig. 25).

BY DAVID J. LEE AND ERIC J. REYE. 237

Male.
Wing pattern similar to female, macrotrichia almost absent. Third segment of

palp divided like a boxing glove. In the genitalia the harpes are complex (Text-fig. 33)

and the aedeagus as in Text-fig. 34.
Distribution QUEENSLAND: Texas, 20:1:1952 and 26:iii: 1952, light trap (A. L.

Dyce). New SoutrH WatLes: Bundy (type series); Noonameena, 8:1:1952, light trap
(A. L. Dyce).

Text-figures 17-253.—Segments 9-15 of the antenna of various species (x 190 approx.):
17, C. coronalisi 18, C. williwilli. 19, C. nattaiensis. 20, C. bunrvooensis. 21, C. moreensis:
22, C. bundyensis. 23, C. waringi.*

Figures 19, 21 and 22 are from holotypes, 17 is from a paratype (392), 18 a paratype (416),
20 a paratype (397), and 23 a paratype (441).

Text-figures 24-31.—Spermathecae of various snecies (x 190 approx.): 24, CG. coronalis-
25, C. williwilli. 26, C. nattaiensis. 27, C. bunrooensis. 28, ©. moreensis. 29, C. bundyensis..
30, C. waringi. 31, C. multimaculatias.

Figures 24, 28 and 30 are from holotypes, 25 is from a paratype (415), 26 paratype (437).
27 a paratype (397), 29 is a Specimen from Texas (430), and 31 a specimen from King
Lake (411).

Text-figures 32-36.—(x190 aprrox.): 32, Terminal abdominal segments of ¢ C. coronalis.
33 and 34, Harpes and Aedeagus of C. williwilli. 35, Harpes of C. nattaiensis. 36, Harpes
of C. bundyensis.

Figure 32 is from the holotype, figures 33-36 are from the allotypes.

CULICOIDES NATTAIENSIS, Nl. sp.
Types: Holotype 9, allotype ¢ and one ? paratype. All slide specimens in S.P.H.T.M.
Type Locality—Stockyard Creek, Colo Vale, near Mittagong, New South Wales,
17:xi:1953, light trap (A. L. Dyce). This is near the headwaters of the Nattai River.

1The sensory pits illustrated on the antenna of this species are also to be found in other
species but were particularly obvious in C. coronalis. :
2 Only segments 10-12 are figured for this species.

238 AUSTRALASIAN CERATOPOGONIDAE. VII,

Distinctive Characters.—This species is closely allied to C. williwilli but lacks the
distinctive wing pattern of the latter. The wing pattern of (. nattaiensis is essentially
similar to that of C. robertsi. However, in C. nattaiensis the third segment of the palp
is considerably swollen (much smaller and not obviously expanded in C. robertsi).
A possibility of confusion may also exist with C. marmoratus, but again the third
segment of the palp is quite distinctive.

Description.—From the type series only. No pinned material available for details
of coloration. Measurements from holotype and selected series comprising two para-
types. Male measurements from allotype only. (See Table 1.)

Female.

Femora with preapical pale spots, tibiae with subbasal ones. Halteres pale. Wings
with well-contrasted pattern, as in Plate ix, fig. 3. Macrotrichia scanty, second radial
cell almost obliterated, its veins thickened.

Head: Hyes separated above, contiguous below (Text-fig. 3). Antennae with basal
segments almost globular, distal five cylindrical and decidedly longer, contrast well
marked (Text-fig. 19). Third segment of palp swollen with large round pit on distal
half, segments IV and V rather short (Text-fig. 11). Mouthparts not as long as
height of head.

Thorax: Legs unmodified, tarsus IV subcylindrical, tibial comb of four spines.

Abdomen: Two fully developed subspherical spermathecae each with a distinct
duct and one very small spermatheca (Text-fig. 26).

Male.

Similar to female on wing pattern and form of palp (cf. OC. williwilli). Genitalia
with complex harpes (Text-fig. 35).

Distribution: New SoutH Waters: Only known from the type locality.

CULICOIDES BUNROOENSIS, Nl. sp.

Types: Holotype 9, allotype ¢ and 10 9 paratypes. All types in S.P.H.T.M. except
four paratypes, one each in C.S.I.R.0., Q.1.M.R., B.M. and U.S.N.M. F

Type Locality—F¥or holotype and all paratypes, Texas, Queensland, light trap, dusk
to 2130 hours, 26:i111:1952 (A. I. Dyce); for allotype, Bundy, via Moree, New South
Wales, light trap, 1:xi:1951 (A. L. Dyce).

Distinctive Characters.—The wing pattern of this species has some basic resemblance
to OC. ornatus, from which it may be distinguished by the dumb-bell-shaped pale area in
the intercalary fork and the presence of pale areas over the extremities of M,, M.
and M,,,. There is also similarity in wing pattern to that cf OC. marmoratus from which
the pale areas at the extremities of the branches of the media are still distinctive and
the single sensory pit on the third segment of the pvalp is in contrast to the multiple
pit of C. marmoratus.

Description—From the type series. No pinned material available for coloration
details. Measurements from the holotype and selected series comprising six paratypes.
Male measurements from allotype only. (See Table 1.)

Female.

The slide specimens reveal that the legs have pale spots preapically on the
femora and also adjacent to the bases of the tibiae. The wings have a complex pattern
of moderately strong contrast (Plate ix, fig. 4) and the halteres are pale.

Head: Hyes rather closely approximated (Text-fig. 4). Antennae rather short,
with basal segments cylindrical (3-10) and distal five scarcely longer except the apical
one, and hence no obvious contrast between proximal and distal segments (Text-fig. 20).
The third segment of the palp (Text-fig. 12) is only slightly enlarged, with greatest
width at about two-thirds from the base with a single distinct sensory pit apically.
The mouthparts are scarcely as long as the height of the head.

Thoraz: Legs with no obvious modifications, tarsus IV subcylindrical, tibial comb
of four spines.

BY DAVID J. LEE AND ERIC J. REYE. 239

Abdomen: Two fully-formed subspherical spermathecae. each with a short duct and
two additional ducts, one of which may be slightly expanded to appear like a deflated
balloon (as in holotype). (See Text-fig. 27.)

Male.

This sex has not been taken.

Distribution QUEENSLAND: Yelarbon, turn off 4 mile on Texas Rd., suction trap,
1800-0045, 27:111:1952 (A. L. Dyce); Texas, 20:1:1952 (A. L. Dyce); Texas, 26:111:1952
(type series). New SoutH WaAtLsES: “Baronga’, Barwon R.—Boomi, suction trap,
19:xii:1951 (A. L. Dyce); Noonameena, light trap, 20:x:1952 (A. L. Dyce); Moree,
30:x:1951 (A. L. Dyce), mercury vapour light trap, 2130-dawn, 20:ii1:1952 (A. L. Dyce) ;
Bundy, via Moree, light trap, 1:xi:1951 (A. L. Dyce); Bundy Cr. crossing, 6:xii:1951,
light trap 0000-0200, 0200-0500, 7:xii:1951 (A. L. Dyce); Yagobie, light trap, 1915-2100,
Sexasil@Rl (CAG Ibe ADAWEE))e

(ii) Species in which the Distal Portion of the Second Radial Cell is Pale.

CULICOIDES MOREENSIS, N. Sp.

Types: Holotype 9 and six 9 paratypes, all slide mounted. Of these one paratype
in C.S.1L.R.O. and Q.1.M.R., the rest in S.P.H.T.M.

Type Locality—Moree, New South Wales. All specimens in type series taken biting
man, 1600-1800 hours, 25:xi:1951 (A. L. Dyce).

Distinctive Characters.—Confusion of the wing pattern of this species may arise
with C. austropalpalis and C. subimmaculatus, but from both of these it differs in having
the end of the second radial cell pale.

Description—From the type series. No pinned material has been available for
details of coloration. Measurements are from holotype and selected series comprising
six paratypes. (See Table 1.)

.

Female.

Legs with pale spots preapically on femora, subbasally on tibiae. MHalteres pale.
The wings (Plate ix, fig. 5) have a reduced rather inconspicuous pattern comprising two
major pale spots, one over r—m, the other at the tip of the second radial cell and just
including the apical portion of this cell. Rather more indefinite pale areas occur in
the cubital fork and below the distal portion of Cu,.

Head: Eyes separated (Text-fig. 5). Antennae (Text-fig. 21) with basal segments
(3-10) subcylindrical, broader basally, last five segments obviously longer than the
basal ones, so with moderate contrast between 3-10 and 11-15. Third segment of palp
swollen, broadest at about middle, outer margin straight, inner very convex. The
sensory pit is large, deep and opening on the distal half (Text-fig. 13). Mouthparts
scarcely equal in length to height of head.

Thorax: Legs unmodified, tarsus IV subcylindrical, tibial comb of four spines.

Abdomen: Two fully formed spermathecae and two accessory ducts (Text-fig. 28).

Male.

This sex is not yet known.

Distribution. QUEENSLAND: Texas, 28:xii:1951 (biting man). NrEw SourH WALES:
Moree, type series and 16:iv:1952, biting man (A. L. Dyce).

CULICOIDES BUNDYENSIS, Nl. sp.

Types: Holotype 2, allotype ¢ and one 9 paratype, all slide specimens in §.P.H.T.M.

Type Locality: Bundy, via Moree, New South Wales, 1:xi:1951, light trap (A. L.
Dyce).

Distinctive Characters.—Except for having the distal portion of the second radial
cell pale the wing pattern is reminiscent of C. ornatus. In the distribution of the
differential wing pattern C. bundyensis is closest to C. magnimaculatus but the pattern
is weaker and the pale areas in the centre of the wing are far more restricted.

Description.—From the type series and other specimens listed below. No pinned
material available for details of coloration. Measurements from holotype and selected

240 AUSTRALASTAN CERATOPOGONIDAR. VII,

series comprising one paratype and a specimen each from Texas and Moree. Male
measurements from allotype only. (See Table 1.)

Female.

Legs with pale subapical femoral spots and similar subbasal tibial ones. Wings
with definite pattern as in Plate ix, fig. 6.

Head: Eyes well separated (Text-fig. 6). Antennae with segments 3-10 cylindrical,
11-14 each longer than the basal segments but not markedly so, 15 distinctly longer
(Text-fig. 22). Third segment of palp expanded in distal half, on which is situated the
single large round sensory pit (Text-fig. 14). Mouthparts at least equal in length to
height of head.

Thorax: Legs unmodified, tarsus IV subcylindrical, tibial comb of four spines.

Abdomen: Two fully developed subspherical spermathecae, each with short duct
(Text-fig. 29).

Jlale.

Wing pattern similar to female but with almost no macrotrichiae. Male genitalia
with simple harpes (Text-fig 36).

Distribution.—QUEENSLAND: Texas, 20:1:1952, light trap (A. L. Dyce). Nrw Sourn
Wates: Moree, 30:x:1951 (A. L. Dyce); Bundy (type series).

CULICOIDES WARINGI, nN. sp.

Types: Holotype 9, allotype 4, five 9 and ten ¢ paratypes, all slide mounted. Of
these, one ¢ paratype in each of C.S.I.R.O., Q.I.M.R., B.M. and U.S.N.M., the rest in
S.P.H.T.M.

Type Locality.—Rottnest Island, near Fremantle (HE. J. Reye, 23: viii:1954, 1000 hrs.,
net, among Acacia on dune near jetty).

Distinctive Characters.—This species is closely related to C. magnimaculatus, from
which it may be distinguished by its larger size, the form of the palpal segments,
particularly the second and third, and the details of the spermathecae.

Description.—From the type series. Measurements from holotype and selected
series comprising five paratypes. Male measurements from six paratypes. (See Table 1.) —

Female.

Legs with dark kneespots. Wings with large pale areas as in Plate ix, fig. 7.
Halteres pale.

Head: Eyes widely separated (Text-fig. 7). Antennae with basal segments (3-10)
subcylindrical, narrowing distally, last five similar in shape but obviously longer
(Text-fig. 23). Third segment of palp expanded at middle, with large sensory pit on
distal half (Text-fig. 15); segment V longer than IV. Mouthparts scarcely as long as
height of head.

Thoraz: Legs unmodified, tarsus IV subcylindrical, tibial comb of four spines.

Abdomen: Two spherical spermathecae each with a distinct duct and a small
accessory duct (Text-fig. 30).

Male.

Similar to female except for usual sexual differences. Harpes long, slender and
curved.

Distribution— WESTERN AUSTRALIA: Only known from the type locality.

(b) A New Name and Additional Notes on Other Species.

CULICOIDES AUSTROPALPALIS, nom. Nov.

Synonymy: Culicoides palpalis Lee and Reye, 1953. Proc. Linn. Soc. N.S.W., 77
(1952): 380-1. Nee Culicoides palpalis Macfie, 1948. Ann. Trop. Med. & Parasit.,
42: 10; 18:

In describing C. palpalis we were not aware of the earlier use of the name palpalis
by Macfie for a species from Mexico. We are indebted to Dr. W. W. Wirth for drawing

BY DAVID J. LEE AND ERIC J. REYE. 241

our attention to this preoccupation. Accordingly we now propose the name austro-
palpalis for the species originally described by us as C. palpalis.

Previously the known range of this species covered the area from Cape York in
Northern Queensland to the vicinity of Sydney in New South Wales. We now know it
to occur further south still. It has been taken at Colo Vale, near Mittagong, New
South Wales, in light traps operated in November, 1953, and February, 1954, and at
Black Mt., Australian Capital Territory, in light traps operated in October, November
and December, 1953.

This species is occasionally taken in a sweep net but otherwise its presence is
only detected by the use of light traps in which it may be taken in large numbers in
favourable areas.

Apart from the variation in the wing pattern noted in the original description
we have since noticed a rare variation in a specimen from Bundy Cr., New South Wales
(7:xii:1951, A. L. Dyce); this is the presence of another large spermatheca in addition
to the two large and one small spermathecae normally present in this species.

CULICOIDES MAGNIMACULATUS Lee and Reye.

Lee, D. J., and Reye, E. J., 1953. Proc. Linn. Soc. N.S.W., 77 (1952): 388-9.

Further collections in New South Wales have confirmed the widespread occurrence,
often in large numbers, of this species. It is taken attracted to man, biting, in light
traps and by net sweeping. Its range into southern Queensland is extended west to
Longreach and it has been taken in numbers at Peel I. In the mountain country of
north-eastern New South Wales it has been taken at Mt. Warning (EH. N. Marks) and
also at Mulimbimby and on the Clarence River (B. McMillan). In the Australian
Capital Territory specimens have been examined from Canberra, Black Mt. (some on
horse, 18:1:1952, R. Mykytowyez) and from Tidbinbilla (some on cattle, 20:1:1953, R.
Mykytowycz). It has also been taken feeding on a rabbit (between Gravesend and
Terry Hie Hie, northern New South Wales, 28:1x:1952, A. L. Dyce). In Victoria it
is recorded from the Grampians (B. McMillan) and Gifford, Lake Denison and Tidal
River (G. W. Douglas). Its range in Tasmania has been extended to Port Davey
' (biting man and dog at dusk, 8:ii1:1954, E. N. Marks).

The Victorian and Tasmanian material shows a variation in the wing pattern which
appears to be present in increasing proportion the further south collections are made.
This variation is in the shape of the intercalary pale area which shows varying degrees
of indentation producing an effect resembling the three pale areas such as are found
in C. dycei, C. multimaculatus, C. memillani and C. marksi but running together at
their adjacent borders. These variants can be distinguished from C. marksi by its
possession of two pale areas in cell M, and three incompletely chitinized spermathecae,
from C. mcmillani and C. dycei by their lesser extent of pale areas on the wing generally
and by the form of their antennae, and from C. multimaculatus (with which the variants
seem most likely to be confused) by the form of the third segment of the palp.

From Fisher I., Bass Strait (15:iv:1954, R. Mykytowyez) comes an extreme variant
in which the intercalary pale area is completely divided into two very unequal portions,
the lesser being a small oval area towards the distal end of the lower branch of the
intercalary fork, the greater being bilobed and proximal in position.

One specimen is aberrant in its spermathecae (from Gifford, Victoria) there being
three large, subequal, subspherical spermathecae instead of two.

CULICOIDES IMMACULATUS Lee and Reye.

Lee, D. J., and Reye, HE. J., 1953. Proc. Linn. Soc. N.S.W., 77 (1952): 375.

The range of distribution of this northcrn species is extended in Torres Strait to
Goode I., Saibai I., Prince of Wales I. and Jacky Jacky Cr. on Cape York Peninsula.
It has also been taken on Low I. and Woody I. off the east coast of Queensland.

We have also examined pinned material from Mabuiag I. in Torres Strait which
shows this species to be entirely lacking in scutal pattern, the scutum being a fairly
uniform dark brown with the scutellum lighter brown.

242 AUSTRALASIAN CERATOPOGONIDAE. VII,

i CULICOIDES ORNATUS Taylor.

Taylor, F. H., 1911. Rept. Aust. Inst. Trop. Med., 73. Lee, D. J., and Reye, EH. J.,
1953. Proc. Linn. Soc. N.SW, 77 (1952): 381-2

This coastal species is now known to extend to Saibai I., Boigu I. and Prince of
Wales I. in Torres Strait, and to Woody I. in Hervey Bay, Green I. and Victoria Point
in Moreton Bay (all the above being Queensland localities) and the latter is now the
recorded southern limit for this species. One of the specimens from Saibai I. has three
subequal subspherical spermathecae instead of the two normally present in this species.
Specimens from Port Samson, near Roeburn (27:11:1954, E. P. Hodgkin) extend the
southern range of this species in Western Australia.

CULICOIDES MARMORATUS (Skuse).

Skuse, F. A., 1889. Proc. Linn. Soc. N.S.W., 4 (2nd series): 304-5 (Ceratopogon).
Macfie, J. W. S., 1989. Proc. Linn. Soc. N.S.W., 64: 556. Lee, D. J., and Reye, HE. J.,
19538. Proc. Linn. Soc. N.S.W., 77 (1952): 382-3.

The range of this coastal species is extended to Woody I., Fisherman I., and
Stradbroke I., in Queensland.

CULICOIDES SUBIMMACULATUS Lee and Reye.
Lee, D. J., and Reye, H. J., 1953. Proc. Linn. Soc. N.S.W., 77 (1952): 375-80.
This coastal species is now known to extend beyond its previously recorded range
to Port Douglas (Queensland) and Albany Passage (Torres Strait, North Queensland).
Off the east coast of Queensland new localities are Woody I. and Green I. In southern
Australia it has been taken at Lake Denison, Victoria and Yorke Peninsula, South
Australia.

_ CULICOIDES MOLESTUS (Skuse).

Skuse, F. A., 1889. Proc. Linn. Soc. N.S.W., 4 (2nd series): 305 (Ceratopogon).
Kieffer, J. J.. 1906. Chironomidae in Wytsman’s Genera Insectorun., fasc. 42: 54. Macfie,
J. W. S., 1939. Proc. Linn. Soc. N.S.W., 64: 556. Lee, D. J., and Reye, EB J., 1953.
Proc. Linn. Soc. NSW, 77 (1952): 382

The range of this coastal species is extended in Queensland by its capture at
Woody I, Maroochy R., Currigee and Southport, and in New South Wales at East Ballina.

CULICOIDES DYCEI Lee and Reye.
Lee, D. J., and Reye, E. J., 1953. Proc. Linn. Soc. N.S.W., 77 (1952): 390.
Previously known from southern Queensland and northern New South Wales,
specimens have since been captured at Colo Vale (near Mittagong, New South Wales)
and in the Australian Capital Territory.
The presence of this species is usually revealed by light trap collections, but it
has also been taken from rabbits and horses.

CULICOIDES MARKSI Lee and Reye.

Lee, D. J., and Reye, EH. J., 1953. Proc. Linn. Soc. N.S.W., 77 (1952): 392.

The distribution of this species is extended northwards to Magnetic I. and westward
to Noondoo in Queensland, and it has been taken at additional localities within its
known range at Greenslopes (biting man) and Yeerongpilly in Queensland, and
Gravesend and Colo Vale in New South Wales, and at Black Mt. and Tidbinbilla in
the Australian Capital Territory (the latter on horse and biting man).

CULICOIDES MULTIMACULATUS Taylor.

Taylor, F. H., 1918. Aust. Zoologist, 1: 169. Macfie, J. W. S., 1939. Proc. LINN. Soc.
N.S.W., 64: 556. Lee, D. J., and Reye, EH. J., 19538. Proc. Linn. Soc. N.S.W., 77 (1952):
390-1.

Additional information on this species has been provided by a long series of
specimens collected by B. McMillan at King Lake (Victoria), 1900-2000 hrs., 15:xii:1953.
A few of these were taken biting man, most were aspirated in flight. Further series
were collected by G. W. Douglas at Gifford, 1930 hrs., 10:ii1:1953, biting man and at

BY DAVID J. LEE AND ERIC J. REYE. 243

Lake Denison (north-east of Yarram), 1930 hrs., undated. Both these localities are
also in Victoria.

We are now able to illustrate the interorbital space (Text-fig. 8), the spermathecae
(Text-fig. 31) and correct a misinterpretation of the palpal sensory organ (Text-fig. 16).
In the badly mounted head of the holotype (previously the only Known specimen) the
palpal organ appeared to comprise three irregular pits; in the fresh material it is
obvious that these are fused into one large, very irregular pit, which is internally
divided by ridges.

As the holotype was not satisfactory for full measurement, measurements are now
given for a specimen from Lake Denison (461) and a series of six specimens from
Gifford and Lake Denison. (See Table 1.)

CULICOIDES ANTENNALIS Lee and Reye.
Lee, D. J., and Reye, H. J., 1958. Proc. Linn. Soc. N.S.W., 77 (1952): 386-7.
The range of this species is now known to extend to Colo Vale, near Mittagong,
New South Wales.

CULICOIDES BANCROFTI Lee and Reye.
Lee, D. J., and Reye, E. J., 1953. Proc. Linn. Soc. N.S.W., 77 (1952): 387-8.
The range of this species is extended to Colo Vale, near Mittagong, New South
Wales (taken February, April and November, 1953) and to Hall’s Gap in the Grampians,
Victoria (taken from calf, 1900-20000 hrs., 22—25:xii:1953, B. McMillan).

CULICOIDES ANGULARIS Lee and Reye.
Lee, D. J., and Reye, HE. J., 1953. Proc. Linn. Soc. N.S.W., 77 (1952): 384-5.
The range of this rather rare species is extended to Black Mt., Australian Capital
Territory (light trap, 2:xi:1953, A. L. Dyce).

CULICOIDES MCMILLANI Lee and Reye.

ee, D. J., and Reye, E. J., 1953. Proc. Linn. Soc. N.S.W., 77 (1952): 391.

The range of this species is extended north to Peel I., Queensland (taken in net,
11:x:1951) and south to Wilson’s Promontory, Victoria (biting man at Tidal R.,
22110: 1953) .

CULICOIDES PARVIMACULATUS Lee and Reye.

Lee, D. J., and Reye, HE. J., 1953. Proc. Linn. Soc. N.S.W., 77 (1952): 391-2.

The range of this species is extended south to Colo Vale, near Mittagong, New
South Wales.

CULICOIDES MAGNESIANUS Lee and Reye.

Lee, D. J., and Reye, H. J., 1953. Proc. Linn. Soc. N.S.W., 77 (1952): 385.

This species was previously only known from Magnetic I., the type locality.
Recently it has been found in the Torres Strait, where it has been taken in numbers
coming to light on Moa I. (M. J. Mackerras and EH. N. Marks) and biting man on
Horn I. (J. Menner). It has also been collected south of the type locality at Woody I.
in Hervey Bay, Fisherman I. and Peel I. in Moreton Bay (all localities in Queensland).

Among specimens from Moa I. were several which were found to have an additional
large subspherical spermatheca (three instead of two). This may represent full
expansion of the end of the isolated duct mentioned in the original description.

Having noted the close similarity of OC. magnesianus with the description of
C. cordiger Macfie from Negri Sembilan, Malaya (Ann. Trop. Med. & Parasit., 28: 193-4)
we asked Dr. Freeman at the British Museum to compare the two species. He replied
that he could “find no differences in proportions of antennal segments, colour and
proportions of legs, spermathecae, halteres or wings. They resemble each other in
appearance.”

It seems possible that the two species may be the same, but as in our experience
palpal differences may be critical it would seem wise to wait until the Malayan species
is more adequately known, and males compared, before committing magnesianus to
synonymy.

244 AUSTRALASIAN CERATOPOGONIDAE. VII,

3. THE GENUS LASIOHELEA.

Kiefter, J. J.. 1921. Arch. Inst. Pasteur de V Afrique du Nord, 1: 115. Kieffer, J. J.,
1925. Faune de France, II, Dipteres Chironomidae Ceratopogoninae: 49. Edwards, F. W.,
1922. Bull. ent. Res., 13: 166-7. Ingram, A., and Macfie, J. W. S., 1924. Ann. Trop. Med.
«& Parasit., 18: 377-92.

In an earlier paper (Lee, 1948) Lasiohelea was mentioned as a member of the
Forcipomyia group. Most authors place the genus as intermediate between Forcipomyia
and Atrichopogon and ascribe to it the venation of the latter associated with the wing
vestiture of the former. For example, Kieffer’s original definition (1921) was as follows:
“Alar pilosity as in Ceratopogon; [that is, Forcipomyia] “empodium as long as the
claws with short hairs; the rest as in Atrichopogon. Type Atrichopogon pilosipennis
Kieff.”. Later (1925) Kieffer expanded this to “Intermediate between Forcipomyia,
of which it has the wing pilosity, and Atrichopogon, of which it has the venation; costa
passing the middle, second [i.e. third] segment of the palp the longest, first, third and
fourth [i.e. second, fourth and fifth] short, except in L. silesiae n. nov. (Pilosipennis
Kieffer ¢ not 9, of which the fourth [i.e. fifth] is the longest. Type L. pilosipennis 9.”

Edwards (1922) and Ingram and Macfie (1924) discuss the characters of the genus
in greater detail but add little in the way of distinctive generic characters other than
a character of the wing fringe which Ingram and Macfie state to be composed of long
hairs between two rows of shorter oblique hairs, not as in Atrichopogon of practically
a single row of alternating long and short straight hairs.

In our limited experience with this genus we do not consider that there is ever
likely to be confusion between it and Atrichopogon, as the density of the macrotrichia
is always considerably greater in Lasiohelea. On the other hand, there is room for
confusion between Lasiohelea and both Forcipomyia and Dasyhelea. The latter, of
course, is distinct in the absence of an empodium, but the form and distribution of
macrotrichia, particularly in the tendency to develop bare lines on either side of the
wing veins, may be quite close to that of Dasyhelea. Similarly, the possession of short
blunt spines on the distal antennal segments is common to both Dasyhelea and
Lasiohelea. In Forcipomyia the distribution of macrotrichia is almost completely
uniform over the wing surface and the second radial cell is not elongated. Indeed the
most characteristic feature of Lasiohelea appears to be the elongation of the second
radial cell associated with a distinct upward curve of R,,; from r—m almost to the
wing margin.

In all, more than 50 species have been placed, at one time or another, in the genus
Lasiohelea, and at the present time about 47 are still ascribed to the genus. A few
of the better known of these are blood-sucking pests but a few others have been taken
attacking various other insects, although some at least of the latter do not appear to
be typical members of the genus. Despite this number of species most authors are
constrained to remark on the difficulty of differentiating species, and this also has
been our experience.

One species only has so far been described from Australia (L. townsvillensis) but
the genus is now known to occur widely in Queensland and New. South Wales and
is also known from Western Australia. We have also examined material from various
localities in New Guinea, and from New Britain and New Ireland. Our concern has
been to try to establish whether or not more than the one previously described species
is involved in these collections and whether or not any of the specimens before us
might be identical with the more notorious L. stimulans from Sumatra or L. lefanui
from the Ethiopian region. A review of the literature and examination of the material
available to us has convinced us that clarity on these points cannot be arrived at for
the time being and only by a comparative treatment of adequate material from many
sources can there be any hope of achieving these ends. Nevertheless, in view of the
possibility of Lasiohelea spp. being responsible for the transmission of certain diseases
of cattle and of the recent remarks by Muirhead-Thomson (1954) concerning Lasiohelea
as so-called “eye-flies” it is important that the specific status of the more prominent
members of the genus be clarified.

BY DAVID J. LEE AND ERIC J. REYE. 245

LASIOHELEA TOWNSVILLENSIS (Taylor).

Taylor, F. H., 1918. Aust. Zoologist, I, Pt. 6: 169 (Culicoides). Macfie, J. W. S.,
1939. Proc. Linn. Soc. N.S.W., 64: 558.

Type: There does not appear to be any specimen in existence actually designated
as type of this species. There are four slides in §.P.H.T.M., obviously of the material
originally examined by Taylor, including one which has been remounted by Macfie. As
most are fragmentary and none are particularly satisfactory as specimens we have
refrained from designating a neotype from this series.

Type Locality—Townsville, Queensland.

Distinctive Characters—Since this species was originally placed in the genus
Culicoides it was overlooked by Edwards (1922) when he said: “Of L. stimulans, the
Museum has a specimen from Deli, Sumatra . . ., several from Peradeniya, Ceylon .. .,
and a series of females, without doubt specifically identical with those from Ceylon,
from various localities in Queensland, some collected biting or labelled ‘troublesome
sandfly’ from Dr. T. L. Bancroft.”

Later, Edwards (1928) further remarks: ‘The species of Lasiohelea are so similar
in all parts of the tropics that it is difficult to suggest the exact affinities of this new
species [samoaensis]. It seems, however, to have more resemblance to L. stimulans .. .
than to L. townsvillensis (Taylor) of Queensland. From L. stimulans it differs in its
smaller size, shorter antennae and dark mesonotal pubescence. I have also seen examples
of undescribed species of Lasiohelea from New Ireland and New Britain, which seem
allied but distinct from this Samoan form.”

Macfie (1939) also remarks: “It [townsvillensis] resembles closely L. lefanui Carter,
an African species, and may indeed prove to be conspecific. Very similar if not identical
forms have been taken in Malaya and elsewhere.”

The close resemblance of L. townsvillensis to both L. stimulans and L. lefanui has
then been obvious to earlier authors. We have not seen L. lefanui but have had
available reputed L. stimulans from Java, which are very similar to Australian material.
In view of this, any distinctive characters which we might propose for L. townsvillensis
are of little more than generic significance.

However, townsvillensis is a Lasiohelea with only one spermatheca, bare eyes,
expanded third segment of the palp with a large sensory pit with a diameter of between
one-third and one-half the length of the segment and largely in the distal half of the
segment. The wing length approximates 1:0 mm. or very slightly less, R,,; terminates
well beyond the middle of the wing, and the tarsal ratio is about 2-0.

We have refrained from illustrating this species, as we feel that only comparative
figures of closely related species can now be of any real value. :

An examination of the male genitalia of specimens from Magnetic I. and Proserpine
has revealed no appreciable differences from the illustrations of the genitalia of
LL. nigeriae (Ingram and Macfie, 1924, p. 382) or L. stimulans (Macfie, 1934, p. 207).
Nevertheless a critical examination of carefully dissected material may yet reveal some
differences of diagnostic value.

Distribution.— QUEENSLAND: Bramston Beach, Inwoods Camp, N.Q., 9:ix:1949 (M.
J. Mackerras); Bramston Beach, Babinda, N.Q., 14:ix:1949 (hovering close to ground
in forest); The Boulders, Babinda, 11:ix:1949 (I. M. Mackerras); Harvey Cr., N.Q.,
15:1x:1949 (1. M. Mackerras); Magnetic I., 9:vii:1952 (EH. J. Reye, 0930 hrs., net,
Horseshoe Bay mangrove swamp and 1100 hrs., net, freshwater swamp); Townsville
(W. J. Young); Proserpine, 15:vii:1952 (EH. J. Reye, 0840 hrs., net, freshwater pool
near stockyard, 4 m. S.); Gin Gin, 5:vii:1952 (EK. J. Reye, 1530 hrs., biting man in
sunlight); Fraser I., 15:ii1:1949 (hovering around head, biting occasionally, rain forest) ;
Pialba, 27:11:1938 (biting in daylight); Hidsvold, 2:iv:1924 (T. L. Bancroft) ; Chinchilla
(J. Mann); Chinchilla, 13:xii:1949 (#. H. 8. Roberts, ex horse); Imbil, Brooloo forest,
29:vi:1938 (biting in daylight); Burpengary—Morayfield, 28:viii:1951 (1345 hrs., biting
man); Deception Bay, 28:viii:1951 (HE. J. Reye); Camp Mt., Brisbane, 19:vi:1938
(R. V. Smythe), 3:vii:1954 (HE. N. Marks); Cedar Cr., Upper Kedron, near Brisbane,

246 AUSPTPRALASIAN CERATOPOGONIDAE. VII.

30:iv:1950; Branch of Ithaca Cr., Mt. Cootha, 9:ix:1950 (J. Pope); Brisbane, 24:ii1:1941
(biting in daylight); Yeronga, 23:11:1954 (EK. J. Reye, 1200 hrs., hovering and biting
man); Blunder Cr., Oxley, 14:v:1950 (M. J. Mackerras); Sunnybank, 19:viii:1950;
Runeorn, 9:iv:1954 (W. Dowd, on goats); Creek near Boonah turnoff from Cunningham’s
Gap Rd., 23:iv:1950 (M. J. Mackerras); Victoria Pt., 11:ix:1954 (M. J. Mackerras) ;
Nunimbah Valley, 19:iii:1950 (M. J. Mackerras, in dense jungle cave, biting), 9: viii:1952
(M. J. Mackerras, biting man); Lamington, O’Reilly’s, 14:viii:1954 (B. McMillan,
biting); Currigee, 3:v:1953 (KE. J. Reye, 1700 hrs., net, mangrove shore line); Texas,
16:iv:1953 (A. L. Dyce, 1200 hrs. and 1600 hrs., biting man). Nrw Sournm Wates: Mt.
Warning, 10:vii:1954 (EH. N. Marks); Byron Bay, 27:vi:1954, 25:vii:1954 (B. McMillan,
biting); Lismore (P. H. Durie); Casino, 5:viii:1954 (B. McMillan, biting on ears);
Barrington Brush, 7:i11:1951 (B. McMillan, 1400 hrs., biting on legs, 1200 ft.) ; Williams.
R., 18:111:1952 (B. McMillan, some biting, 1500 ft.); Cooranbong, 26:11:1950, 1:vi:1951
(B. McMillan); Cowan Cr., 27:x:1949 (B. McMillan); Hornsby, 8:x:1950 (B. McMillan),
5:ix:1954 (H. J. Reye, 1545 hrs., net in gully); Cox R., 6:v:1954 (some biting on legs) ;
Springwood, 16:iv:1950 (B. McMillan, biting).

Apart from the above we have also seen a single specimen of Lasiohelea from the
north of Western Australia (Kulumburu, 11:i111:1954, E. P. Hodgkin) which we cannot
place in townsvillensis, as the third segment of the palp is longer than in the eastern
Australian form and the sensory organ is no more than one-quarter the length of the
segment.

From the New Guinea area the following material has been examined: Biak, D.N.G.,
—:1:1945 (W. R. Horsfall); Kanosia, Papua, 5:vi:1947 (R. H. Wharton, biting, 1400 hrs.,
in bush); Hilogo, Papua, 2:vi:1947 (D. J. Lee, on cattle); Samarai, Papua (F. J.
Williams); Keravat, New Britain, 11: viii:1953, 7:xii:1953 (G. S. Dun). None of this
are we prepared to identify as L. townsvillensis, but the only differences we can detect
are that the segments of the palps tend to be slightly smaller and the antennal ratio
slightly higher in the New Guinea material. Otherwise there is very close similarity
to L. townsvillensis.

It should also be noted that Muirhead-Thomson (1954) notes that Dr. Freeman has
advised that specimens of lL. lefanui var. squamipes have been received from New
Britain (in the British Museum).

References.
(Most systematic references are quoted in full in the text.)

EpWARDs, F. W., 1928.—insects of Samoa, Part vi. Diptera, fasc. 2. Nematocera: 51.
Leg, D. J., 1948.—Proc. LINN. Soc. N.S.W., 72 (1947): 313-331.

MUIRHEAD-THOMSON, R. C., 1954.—Ann. Trop. Med. & Parasit., 48; 121.

WIRTH, W. W., 1952.—Univ. Calif. Pub. in Hntom., Vol. 9, No. 2: 95-266.

EXPLANATION OF PLATE IX.

Wings of various species of Culicoides (x30 approx.): 1, C. coronalis. 2, C. williwilli.
3, C. nattaiensis. 4, C. bunrcoensis. 5, C. moreensis. 6, OC. bundyensis. 7, C. waringi.

Figures 2, 3, 4, 6 and 7 are from tke holotypes, 1 is from a paratype (392), and 5 from a
paratype (387).

i

Proc. LINN. Soc. N.S.W., 1954.

PLATE

“Anomalous”

krasnozem at Fairy Hill.

Vill.

TAU Hes Texe

Proc. LINN. Soc. N.S.W., 1954

icoidés.

ies of Cuwli

f spec

Wings o

a

(Issued 10th June, 1954.)

Vol. LXXIX. Nos. 371-372.
Parts 1-2.
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EXPLANATION OF PLATES II-III.

Plate ii.

1, Achnanthes brevipes Ag. var. intermedia Kutz. 73 x 20 pw. 10-11 striae/ 10 mw. x 1000.
2, A. brevipes var. intermedia. 58 x 20 w. 9-10 striae/ 10 w. x 1000. 38, Achnanthes lanceolata
(Breb.) Grun. 28 x 9 w. 12 costae/ 10 uw. x 2000. 4, Fragilaria (virescens?) Ralfs. 13-20 x By.
23 costae/ 10 uw. x 700. 5, Cymbella pusilla Grun. var.? 33 x 5-7 w. 16 costae/ 10 pw. x 700.
6, Diploneis smithii (Breb.) Cleve. 32 x 15 w. 12 striae/ 10 uw. x 2000. 7, Achnanthes subsalsa
Peterson. 18 x 8 uw. 14 costae/ 10 w. x 1000. 8, Diatomella Balfouriana Grey. Length 15 ph.
21 costae/ 10 w. x 1000. 9, Fragilaria Harrisonii (W. Smith) Grun. 21 x 13 uw. 10 costae/ 10 li, ¥e
700. 10, Hunotia trinacria Krasske. 16 x 2 uw. approx. 20 costae/ 10 mw. x 700. 11, Hunotia lunaris
Grun. 36 x 3 uw. 22-25 costae/ 10 w. x 700. 12, Hunotia alpina (Naeg.) Hust. 36 x 3 uw. 15 costae/
10 uw. x 700. 18, 14, Fragilaria capucina Desm. 34-53 x 3 uw. 17 costae/ 10 w. x 1000. 15, Fragilaria
virescens Ralfs. Length 40 uw. 16-18 costae/ 10m. x 700. 16, Gomphonema parvulum (Kutz.) Grun.
32 x 9 uw. 18-14 costae/ 10 mw. x 1000. 17, Gomphonema intricatum Kutz. 54 x 10 uw. 12 costae/
10 uw. x 1000. 18, Fragilariopsis antarctica Castr. 30 x 8 uw. 5-6 striae/ 10 w. x 1000. 19, ? Frus-
tulia minuta Rab. 25 x 4 uw. x 1000. 20a and b, Melosira granulata (Hh.) Ralfs. 20a, 11 x 7:5 ps.
x 2000. 206, 27 x 4 w. x 700. 21, Navicula (rostellata?) Kutz. 40 x 5 uw. 18 striae/ 10 yu.
x 1000. 22, Opephora Martyi Heribaud. Length 14 uw. 10 costae/ 10 uw. x 2000. 23, Pinnularia
appendiculata (Ag.) Cleve. 40 x 6 uw. 14 costae/ 10 uw x 700. 24, Pinnularia molaris Grun.
37 x 5 uw. 16 costae/ 10 w. x 1000. 25, P. molaris. 44 x 6 uw. 15 costae/ 10 uw. x 1000. 26, Pinnularia
divergentissima (Grun.) Cleve. 36 x 6 mw. 11-12 costae/ 10 w. x 700. 27, Pinnularia microstauron,
Eh, 48 x 8 w. 16 costae/ 10 w. x 1000. 28, Hantzschia amphioxys (Eh.) Grun. 75 x 10 yw. 23 costae/
10 uw. x 2000. 29, Pinnularia brevicostata Cleve. 78 x 15 uw. 9 costae/ 10 w. x 1000. 30, Pinnularia
cardinalis (Eh.) W. Smith. 74 x 12 w. 10 costae/ 10 mw. x 1000.

d Plate iii.

1, Pinnularia divergens W. Smith. 120 x 22 uw. 8-9 costae/ 10 mw. x 2000. 2, Pinnularia lata
(Breb.) W. Smith. 85 x 20 w. 4 costae/ 10 w. x 2000. 38a, b, c, Stawroneis acuta W. Smith.
1438 x 28 w. 15 striae/ 10 mw. 38a and 3c, x 1000. 3b, x 2000. 4, Pinnularia microstauron Eh.
60 x 12 w. 15 costae/ 10 w. x 1000. 5, Swrirella angustata Kutz. 74 x 14 w. x 1000. 6, ? Synedra
vaucheriae Kutz. 24 x 4 uw. 16 costae/ 10 w. x 1000. 7, Surirella bifrons Kutz. 100 x 28 uw. x 1000.
8, Stauroneis anceps Eh. var. hyalina Brun and Per. 68 x 13 uw. 20 striae/ 10 uw. x 1000.

PROCEEDINGS, LXXIX, PARTS 1-2, 1954

CONTENTS.

Pages.

Presidential Address, delivered at the Seventh-ninth Annual General Meeting,

31st March, 1954, by J. M. Vincent, B.Sc.Agr., Dip.Bact. The Root-Nodule
Bacteria of Pasture Legumes. (One Text-figure.) .. .. .. .. .. .. i-xxxii
OTS CEVOMS i552) Fico ae Week aT doe resend cal Clone 2 Payline cach ies 6 Ue IN toad aa ee ne ee a ae aT XXxii

Inheritance and the Genetic Relationship of Resistance Possessed by Two
Kenya Wheats to Races of Puccinia graminis tritici. By D. S. Athwal
andl. As WiatSOm i a1 ei sutra blew. hoe glee ete le see, cl ce Neloy SIH Lar Die 1-14

Macadamia ternifolia F. Muell. and a Related New Species. By L. A. S. Johnson 15-18

Experimental Crossing of Aédes (Stegomyia) pseudoscutellaris Theobald and
Aédes (Stegomyia) polynesiensis Marks (Diptera, Culicidae). By A. R.
WOO GIN ee ia Uneee eee eet sree wren ha cates, Ran. IRS Lk We A it NMA ale gsc ne a 19-20

Cytological Studies in the Myrtaceae. IV. The Subtribe Huchamaelaucinae.
By S. Smith-White. (Forty-one Text-figures.) .. .. .. .. .. «se. 21-28

A New Species of Hibbertia Andr. from Western Australia. By A. T. Hotchkiss.
(Blatesaitand mtourhy Next=fe unre sy) nein. cea ake alba) CAN a 7SHi es ssah SIR ee ec oe 29-33

A Comparative Account of the Terrestrial Diatoms of Macquarie Island. By
J. S. Bunt. (Plates ii-iii and twenty-three Text-figures. ) FR ReMi Ot tater Lt 34-57

A
WH LAEG 4

Em