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THE

PROGrE DINGS

OF THE

Peay SOCIETY

OF

Nisa Sears “Woes

FOR THE YEAR

1958

VOL. LXXXIII.

WITH SIX PLATES.
392 Text-figures.

SYDNEY:
PRINTED AND PUBLISHED FOR THE SOCIETY BY

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

and
SOLD BY THE SOCIETY,
1959.

CONTENTS.

CONTENTS GE PROCEEDINGS, 1258

PART 1 (No. 386).
(Issued 24th March, 1958.)

Pages
Presidential Address, delivered at the Highty-third Annual General Meeting,
26th March, 1958, by Dr. Lilian R. Fraser:
Summanryvwot Years) "Activities! 29 2) i) 5 eee ce te Os eee 1- 5
Virus Diseases of Citrus in Australia .. .. .. .. .. .. .. .. 9-19
ISTE CEIONGS! kiss. os neieiny, ROR” ul behdan Meeewe fetes, ura Test ceaien Peed deseies genclset cameras ict cis met Lali 5
Batamce Sheets for the Year ending 28th February, 1958 .. .. .. .. .. 6— 8
Seed Coat Anatomy and Taxonomy in Eucalyptus. I. By E. Gauba and L. D.
Pryor. (Plate i; nineteen Text-figures. ) Rea MERI oe See ee as deo 20-32
A New Spécies of Aédes (Finlaya) from Northern Australia (Diptera, Culicidae).
By Elizabeth N. Marks and Ernest P. Hodgkin. (One Text-figure.) .. .. 33-39
A Summary of the Atopomelinae (Acarina, Listrophoridae). By R. Domrow.
(Nine Text-figures. ) Pe ee ae ee Me oka eS ee at es 40--54
Inheritance of Oil Characters in Hucalyptus. By L. D. Pryor and L. H. Bryant.
(Five Text-figures.) Sey a, Monee crsis! made,” Mote Re gietcy (a Atel ic amiga noe ies 55-64

The Status of Nitrogen in the Hawkesbury Sandstone Soils and their Plant
Communities in the Sydney District. II. The Distribution and Circulation
of Nitrogen. By Nola J. Hannon, Linnean Macleay Fellow of the Society
in Botany. (Two Text-figures.) Rn ERE ae a Ae ens re ein, eo 65-85

CONTENTS.

PART 2 (No. 387).
(Issued 26th September, 1958.)

Pollen and Pollination in the Hupomatiaceae. By A. T. Hotchkiss. (Plate ii;
twenty-six Text-figures. )

The Species of the Genus Hrodium L’Hér. endemic to Australia. (With a Key
to all the Taxa known to occur in Australia.) By R. C. Carolin. (Twenty-
eight Text-figures.)

Catalogue of Australian Mammals and their Recorded Internal Parasites. I-IV.
By M. Josephine Mackerras. (Communicated by Dr. I. M. Mackerras.)

Studies of Nitrogen-Fixing Bacteria. VII. Cytochromes of Azotobacteriaceae.
By F. J. Moss and Y. T. Tchan

Migration and Utilization of Reserve Substances during Flight in Aphis
craccivora Koch. By M. Casimir. (One Text-figure.)

A New Bird-Flea from Tasmania. By F. G. A. M. Smit. (Communicated by
Mr. D. J. Lee.) (Ten Text-figures.)

Widespread Natural Infection of Barberry by Puccinia graminis in Tasmania.
By I. A. Watson and N. H. Luig

Systematic Notes on Some Eastern Australian Members of the Papilionaceae.
By Joy Thompson

Somatic Hybridization in Puccinia graminis var. tritici. By I. A. Watson and
N. H. Luig

A Note on the Status of Aphodius tasmaniae Hope. By B. B. Given. (Com-
municated by Mr. C. E. Chadwick.)

Sir William Macleay Memorial Lecture. Timing in Human Evolution. By A. A.
Abbie. (Seven Text-figures. )

Some More Bark- and Timber-Beetles from Australia. 158. Contribution to the
Morphology and Taxonomy of the Scolytoidea. By Karl E. Schedl. (Com-
municated by Dr. A. J. Nicholson.)

Two New Species of Hemicycliophora (Nematoda : Tylenchida). By M. R. Sauer.
(Communicated by Mr. A. J. Bearup.) (Two Text-figures.)

A New Species of Frog of the Genus OCrinia Tschudi from South-Hastern
Australia. By Murray J. Littlejohn. (Communicated by Mr. S. J. Copland.)

92-100

.. 101-160

.. 161-164

. 165-172

.. 173-180

. 181-186

. 187-189

. 190-195

196

.. 197-213

. 214-216

. 217-221

222-226

(7389

CONTENTS.

PART 3 (No. 388).
(Issued 23rd March, 1959.)

Acarina from Australian Bats. By Robert Domrow. (Twenty-six Text-figures.)

A Turbidimetrie Method for Estimating the Number of Nematode Larvae in a
Suspension. By C. D. Blake. (Two Text-figures.)
The Oviposition Behaviour of Aédes australis (Erickson) (Diptera, Culicidae).
By A. K. O’Gower. ..
Palaeozoic Geology of the Cooleman Caves District, New South Wales. By
N. C. Stevens. (Plates iii-iv; one Text-figure.)

Melampsora lini (Pers.) Lév. Uredospore Longevity and Germination, By H. B.
Kerr. (Three text-figures.)
On some Pergine Sawflies reared by Mr. M. F. Leask (Hymenoptera, Pergidae).
By Robert B. Benson. (Communicated by Mr. K. EH. W. Salter.) (One
Text-figure. )
The Diptera of Katoomba. Part 2. Leptidae and Dolichopodidae. By G. H.
Hardy. (Nine Text-figures.)

Bat-infesting Ornithodoros (Ixodoidea-Argasidae) of the Oriental-Australian
Region. By L. J. Dumbleton. (Communicated by Dr. J. W. Evans.)

(Highteen Text-figures. )

Notes on Australian Thynninae. II. The Genera Dimorphothynnus, Rhagigaster
and Hirone. By B. B. Given. (Communicated by Dr. A. J. Nicholson.)
(One hundred and twenty-six Text-figures.)

Notes on Australian Thynninae. III. The Genus Thynnoides. By B. B. Given.
(Communicated by Dr. A. J. Nicholson.) (Wighty-eight Text-figures. )

Australasian Ceratopogonidae (Diptera, Nematocera). Part VIII. A New Genus
from Western Australia attacking Man. By Willis W. Wirth and David J.
Lee. (Five Text-figures. )

Mode of Inheritance of Resistance to Powdery Mildew in Barley and Evidence

for an Allelic Series Conditioning Reaction. By N. H. Luig, K. S.
McWhirter and EH. P. Baker.

Spores and Pollens from a Permian-Triassic Transition, N.S.W. By J. P. F.
Hennelly. (Plates v-vi; two Text-figures.)

Abstract of Proceedings

List of Members

List of Plates

List of New Genera, Species, Subspecies and Name ..

Index

Pages
227-240

. 241-244

. 245-250

. 251-258

. 259-287

.. 288-290

.. 291-302

. 303-308

.. 309-326

. 327-336

.. 337-339

.. 340-362

. 363-369
. 371-376
. 377-882

383
383

- 384-387

ANNUAL GENERAL MEETING.
26th Mancu, 1958.

The Highty-Third Annual General Meeting was held in the Society’s Rooms, Science
House, 157 Gloucester Street, Sydney, on Wednesday, 26th March, 1958.

Dr. Lilian R. Fraser, President, occupied the chair. é

The minutes of the Highty-Second Annual General Meeting, 27th March, 1957,
were read and confirmed.

PRESIDENTIAL ADDRESS.

It is first my pleasant duty to convey to the Society’s honorary officers, Dr. W. R.
Browne and Dr. A. B. Walkom, our deep appreciation of their work during the past
year. It is impossible to overemphasize the debt which we owe to these gentlemen
for their services in an honorary capacity, as secretary, treasurer and editor, their
invaluable experience in all branches of the Society’s functions and organization, and
their patience and good humour in dealing with the occasional problems which arise.
I must also express our appreciation of the services of the assistant secretary, Miss
Allpress, who watches over the needs of secretary and treasurer, Council members
and ordinary members alike, and ensures the smooth running of day-to-day activities.
I should also like to record my personal indebtedness to the Council for their support
during the past year.

Parts 1 and 2 of Volume 82 of the Society’s PROCEEDINGS were published in 1957,
and Part 3 in March, 1958. Volume 82 consists of 384 pages, 14 plates and 190
text-figures. A contribution of £50 towards the cost of publication of “Australian Tree
Frogs of the Genus Hyla’* was made by Mr. S. J. Copland. A leaflet, giving a short
account of the history, aims and objects of the Society has been issued for prospective
members and others who may be interested in the Society’s activities; forms for
application for admission as an Associate are also now available.

During the year 21 new members were added to the list, two members died,
three resigned and two were removed from the list under Rule VII. The numerical
strength of the Society at 28th February, 1958, was: Ordinary Members, 225; Life
Members, 32; Corresponding Members, 2; Associate Member, 1; total 260.

Members will note that during the year fluorescent lighting was installed in the
meeting room and the ceiling and walls were painted. It was decided by Council that
ordinary monthly meetings during 1958 should commence at 6 p.m., instead of 7.30 p.m.
This experiment was started during 1957 in the hope that members might find the
earlier meeting time more convenient than the latter. The changed time appears to
have resulted in rather larger attendances at most meetings.

Council resolved to establish a Sir William Macleay Memorial Lecture to be given
biennially in some branch of science covered by the Society’s activities, the text of the
lecture to be published in the Procrrpines. The first lecture has been fixed for
Thursday, 19th June, 1958, Professor A. A. Abbie, of Adelaide, having consented to
deliver it.

Lecturettes were given at the following monthly meetings: April: New Caledonia
and its Plants, by Dr. H. S. McKee; May: A Visit to Some American Universities
and Marine Biological Stations, by Miss Isobel I. Bennett; June: Some Aspects of
Plant Distribution in Southern Africa, by Dr. A. R. H. Martin; July: Some Aspects
of the Biology of the Bandicoot, by Mr. Gordon Lyne; October: Social Behaviour in
the Australian Bulldog Ants, by Mr. John Freeland. On 25th September, 1957,
commemoration was made of the 250th anniversary of the birth of Linnaeus. An
article on the botanical work of Linnaeus, written by Mr. L. A. S. Johnson, was,

PROCEEDINGS OF THE LINNEAN Society or NEw SoutH WALES, 1958, Vol. Ixxxiii, Part 1.

A

2 ANNUAL GENERAL MEETING.

in his unavoidable absence, read by Miss Mary D. Tindale, and Mr. G. P. Whitley gave
an illustrated talk on the life and zoological work of Linnaeus, exhibiting first
editions and other interesting “Linnaeana’”’. We wish to express our appreciation and
thanks to all lecturers for their interesting contributions to our meetings. Members
again showed keen interest in bringing notes and exhibits.

Library accessions from scientific societies and institutions were slightly fewer
than in previous years, the total being 1,858. Requests for library loans and the
demand for reprints continued to be made as frequently as previously. Mrs. F. C.
Blanchard, a daughter of Dr. N. A. Cobb who was a member of the Society from 1889.
to 1906, forwarded a reprint for the Society’s library of ‘Nathan A. Cobb, Botanist.
and Zoologist, a Pioneer Scientist in Australia’ (from “the Asa Gray Bulletin’, n. s.
Vol. 3, No. 2, pp. 205-272, 1957). A first edition of ‘“Lachesis Lapponica, or a Tour in
Lapland, now first published from the original manuscript journal of the celebrated
Linnaeus” (2 vols. in one), by J. EH. Smith, London, 1811, was presented to the
Society on the occasion of its celebration of the 250th anniversary of the birth of.
Linnaeus, by Mr. A. EH. Jobson. Mr. David S. Macmillan presented to the Society a
copy of his book “A Squatter Went to Sea. The Story of Sir William Macleay’s New
Guinea Expedition (1875) and his Life in Sydney”, in which due acknowledgement
was made of his use of extracts from Sir William Macleay’s diary and other sources.
on the “Chevert” expedition in the possession of the Society. In January, the library
was thoroughly cleaned and dusted. During the year steel shelving was installed in
the storeroom to accommodate the stock of PRocEEDINGS, which has been rearranged
and checked. ;

Council has decided that in future the price of the Macleay Memorial Volume
shall be £1. 1s., post free, instead of £2. 2s. as previously.

The Society, through its Council, has supported the efforts made to oppose the
renewal of grazing leases in the Snowy Mountains area in the interests of soil conserva-
tion and prevention of soil erosion. The Society was represented at conferences held.
during the year of bodies interested in conservation. A report supporting the reservation.
of the Deep Creek area was forwarded to the Cumberland County Council. The Society
has also taken up Corporate Membership of the National Parks Association of New
South Wales, Central Region. Following the receipt of representations from the
Joint Scientific Advisory Committee the Council adopted two resolutions regarding
the Kosciusko area, the first strongly deprecating any proposal that there should be
a return to snow-lease grazing and the second urging the Kosciusko State Park Trust.
to take immediate action in regard to the setting apart of a Primitive Area. The
resolutions were forwarded to appropriate State Ministers and to the Kosciusko State
Park Trust respectively.

The total net return from the Society’s one-third ownership of Science House:
for the year was £1,065. The rental determinations of the Fair Rents Board as from.
ist October, 1957, gave an increase of 19% to the rents in Science House, the Society’s.
rent of rooms and storeroom being now £813 as against the previous approximate
amount of £648.

It is my pleasure to offer congratulations to Dr. Germaine A. Joplin on the award
of a Carnegie Grant for study in the United States of America, and to Professor
J. M. Vincent on the award of the medal of the Australian Institute of Agricultural
Science.

Linnean Macleay Fellowships. :

In November, 1956, the Council reappointed Miss Nola J. Hannon and Mrs. Mary B..
Williams to Fellowships in Botany for 1957.

During the earlier months of 1957 Miss Hannon devoted her time to the compilation
of unrecorded observations and unpublished files of relevance to the status of nitrogen
in the Hawkesbury Sandstone ecosystems. This information covered a wide variety of
subject matter and was obtained from private firms and individuals and government
officials. By combining these data with those that had been collected during the
three-year tenure of the Linnean Macleay Fellowship, a general survey of nitrogen in
these communities was completed. A second paper in the series “The Status of

ANNUAL GENERAL MEETING. 3

Nitrogen in the Hawkesbury Sandstone Soils and their Plant Communities in the
Sydney District. The Distribution and Circulation of Nitrogen” has been accepted
for publication in the Procrepines and further papers are being prepared. Investiga-
tions of the native rhizobial population have been concerned with the effectiveness of
the symbiotic associations with Acacia suaveolens (Sm.) Willd., the cultural charac-
teristics and the nutritional requirements of isolates chiefly from the Sydney sandstone
soils.

During 1957, Mrs. Williams continued her study of the family of freshwater Algae,
the Characeae. More determinations of chromosome numbers were made. A provisional
count of 18 chromosomes was made on Nitella stuartii A. Br., the one member so far
collected of the section Anarthrodactylae. It was particularly interesting to find that
a member of this section had a chromosome number conforming to the basic number,
nine, as determined for other sections of the genus, although the uniformity of numbers
does not give any clues to possible phylogenetic relationships within the genus.
Within certain species, however, it has been established that the chromosome number
may vary according to locality, so that several “chromosome races” have been found,
forming polyploid series within these species. Further investigations were made into
the dormancy of Chara australis spores; treatments included pretreatment at 5°C.
for 140 days, pretreatment with cold concentrated sulphuric acid, and pretreatment
with infusions of rotting Chara plants. A low percentage germination was obtained
in some cases, but no method gave 100% breaking of dormancy. It seems likely
that dormancy in Chara spores is a complex phenomenon, which is not overcome by
single treatments.

In November, 1957, the Council reappointed Miss Nola Hannon and Mrs. Mary
Williams to Fellowships in Botany for 1958.

In consequence of her appointment to a lectureship in the N.S.W. University of
Technology the resignation of Miss Hannon from her Fellowship as from 11th April,
1958, has been accepted by Council. We congratulate her on her new appointment.
It is hoped that Parts III-VI of ‘The Status of Nitrogen in the Hawkesbury Sandstone
Soils and their Plant Communities in the Sydney District’, together with two other
papers, will be submitted shortly for publication in the PROCEEDINGS.

During 1958, Mrs. Williams, who will be working in the Department of Botany,
University of New England, instead of the University of Sydney, proposes to continue
her work on the Australian Characeae. Owing to the recent disastrous fire in the
Department of Botany, University of New England, in which Mrs. Williams lost much
material, her work has necessitated replanning, which is not yet complete. We wish
her every success in her research work.

Linnean Macleay Lectureship in Microbiology.

Dr. Y. T. Tchan, who was appointed Linnean Macleay Lecturer in Microbiology as
from ist August, 1955, has furnished reports from the date of his appointment to
3lst December, 1957. His activities may be summarized as: (1) Teaching: His first
year was mainly used for organizing lectures and practical classes. Complete new
courses dealing with microscopy, infectious bacteria and bacterial and protozoan
systematics were prepared for Science and Agriculture students. These courses were
then modified in the second year in the light of the previous year’s experience.
(2) Research: Some of the research has been carried out with financial assistance
from C.S.I.R.O. (Land Survey and Utilization Section). The survey of the distribution
of Beijerinckia in Northern Australia was continued and some physiological studies
of representatives of the genus were made. Studies of Northern Territory soils
(Katherine Station soils) were made. A complete bacteriological analysis of these
soils was completed. (3) Microscopy: In conjunction with the University Hxtension
Board, a post-graduate course on Light Microscopy was given. The lectures have been
attended by research workers of various biological disciplines. Also, in collaboration
with Dr. W. H. Steel, of C.S.I.R.O., a paper is in preparation for publication on the
theoretical implications of overlap in phase microscope image formation. (4) N-fization.
bacteria: In collaboration with Dr. F. Moss, Department of Bacteriology, Sydney

4 ANNUAL GENERAL MEETING.

University, a paper is under preparation on the subject of cytochromes of Azoto-
bacteriaceae. (5) Fertility test: Using the algal method the P availability could be
determined. Good correlations have been obtained with higher plants. (Part of this
programme was carried out with the collaboration of Mr. R. Hawkes.) (6) A new
project on the Northern Territory soils has been started. It is aimed to study the
N fixation algal population and its ecological conditions.

In the year 1956-57, more work on the soil algae was done. A new approach to
the soil fertility test with algae was made. Some very promising results were
obtained for nitrogen estimation by this means. The technique also seems applicable
for phosphorus estimation. Further work is needed before any conclusion can be
reached. A preliminary paper has been published (VI Congres internationale de la
Science du sol, 1956). An improved technique of fluorescence-phase microscopy has
been developed and a short paper has been accepted for publication in Nature. A new
species of Beijerinckia has been described in a paper published in Part 3 of the
Society’s PROCEEDINGS.

In connection with the Bacteriology Fund, it is reported that the property at
53 Margate Street, Ramsgate. N.S.W., has been sold for £4,250, £2,200 of which remains
under a 15-years’ mortgage bearing interest at 7% per annum, the mortgagors having
the right to repay the whole of the principal on three months’ notice and the additional
right of reducing the principal sum by £50 per annum or multiples of £50.

Obituaries.

It is recorded with regret that the following members died during the year:

Emeritus Professor Wiitt1Am Nort Benson, B.A., D.Sc., F.R.S., F.G.S., who had
been a member of the Society since 1907, died at Dunedin, New Zealand, on 20th August,
1957. He was born near London on 26th December, 1885, and educated at the Friends’
High School in Hobart and later at the University of Tasmania and the University
of Sydney, where he came under the influence of the inspiring personality and teaching
of Sir Edgeworth David. From 1914 to 1916 he held a Linnean Macleay Fellowship
in Geology. He contributed ten papers to the Society’s PROCEEDINGS; also one with
F. Chapman and one with W. S. Dun and W. R. Browne. These papers included his
classic study of the geology and petrology of the Great Serpentine Belt of New South
Wales He was Professor of Geology at the University of Otago, Dunedin, from 1917
to 1949. In Dunedin he turned his attention io problems of New Zealand geology as
well as its relation to the south-west Pacific as a whole.

During his life-tinie Professor Benson received many honours. He was a
foundation Fellow of the Australian and New Zealand Assosciation for the Advancement
of Science, and its Mueller Medallist (1951); Fellow of the New Zealand Institute,
later Royal Society of New Zealand (1926); Hector medallist (1933), Hutton medallist
(1944) and president (1945-47) of the same Society; Lyell medallist of the Geological
Society of London (1939); Fellow of the Royal Society (1941); Clarke medallist of
the Royal Society of New South Wales (1945); correspondent of the Geological Society
of America (1949); honorary D.Sc., University of New Zealand (1951), and honorary
member of the Mineralogical Society (1954).

Mr. Davin Grorcr STEAD died at Watson’s Bay, Sydney, on 2nd August, 1957. He
had been a member of the Society since 1898, thus being the oldest member at the
time of his death. He was a noted authority on fishes and an all-round naturalist.
For many years he was actively interested in the work of the Wild Life Preservation
Society and the Royal Zoological Society of New South Wales. He contributed two
papers (1898 and 1899) to the Society’s ProcrEEpINGsS and in his earlier years took a
keen interest in the Society's meetings and made presentations of books to the
Society’s library.

PRESIDENTIAL ADDRESS.
Virus Diseases of Citrus Trees in Australia.
Seven diseases caused by virus infection are known to affect citrus trees in
Australia. Two of these, scalybutt of Poncirus trifoliata rootstock, and stem pitting
disease of grapefruit, are of major importance; tristeza, a very important disease

ANNUAL GENERAL MEETING. 5

elsewhere, is not a problem in Australia, and psorosis, crinkly leaf, woody gall and
enation are minor or unimportant. The major symptoms, distribution, importance
and method of control are discussed. Brief reference is made to two virus diseases,
stubborn and xyloporosis, which are important overseas but not as yet definitely known
to be present in Australia. (For full text, see pp. 9-19.)

The Honorary Treasurer, Dr. A. B. Walkom, presented the balance sheets for the
year ended 28th February, 1958, 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: S. Smith-White, D.Sc.Agr.
Members of Council: A. J. Bearup, B.Sc.; F. V. Mercer, B.Sc., Ph.D.; S. Smith-
White, D.Sc.Agr.; EH. Le G. Troughton, C.M.Z.S., F.R.Z.S.; H. S. H. Wardlaw,
D.Se., F.R.A.C.I.; and A. R. Woodhill, D.Sc.Agr.
Auditor: S. J. Rayment, F.C.A. (Aust.).

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

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PRESIDENTIAL ADDRESS.

VIRUS DISEASES OF CITRUS IN AUSTRALIA.
By LiniaAn R. FRASER.

(Detivered 26th March, 1958.)

Studies of plant viruses are of two general types, those which seek to elucidate
the disease condition caused by a virus, its vector relationships, variation in host
reaction, influence of environment, virus strain relationships, etc., and those which
seek to investigate the virus itself as an entity by biochemical or biophysical means.
These fundamental studies are providing the foundations upon which some rational
system of virus taxonemy may in the future be based, and have already made it
possible to demonstrate or ecorfirm relationship or lack cf relationship in certain
virus groups. This type of study is so far only possible with easily handled, stable
and highly infectious viruses, and the diseases of woody perennial plants, for the
most part, do not come into this category. ‘The study of the citrus viruses is still
largely in the stage of disease delineation, and the identification of a virus is based
on the symptoms produced in inoculated seediings, and relationships are assumed on
the basis of general similarity of symptoms. The need for precise means of establishing
relationships is urgent.

There are a number of special disease problems of perennial vegetatively propagated
plants which are particularly important in citrus. The first is the accumulation of
virus diseases and their propagation and spread in vegetative plant parts. Varieties
of orange or grapefruit or mandarin, locally esteemed, are imported into other
countries, taking with them any virus which they may be carrying. The second is
the existence of symptomless carriers which makes the accumulation of viruses so
dangerous. Varieties and species of citrus vary very greatly in their reaction to
infection by a particular virus. In some cases an apparently quite healthy plant can
carry a virus which is damaging or lethal to other varieties or combinations of
varieties. A third problem is that of latency. Some viruses may take many years
to preduce recognizable symptoms even on susceptible varieties, and during the period
of latency such affected plants can be used as a source of propagation material
in ignorance, with conseauent spread of the disease.

The complete examination of a new virus proklem involves not only a study of
behaviour in the field, but a search for indicaters, i.e. varieties which, when inoculated,
will give ar unmistakable, specific and reasonably rapid reaction. Only when such
indicators are available is it possible to make surveys of suspect populations, to
index material for disease freedom sc that sources of disease-free material for
propagation can be built up and to screen introductions from foreign countries.
Confusion can occur if more than one virus is present, and the search for indicators
which will adequately separate different viruses has been beset by pitfalls arising
from this cause. The danger here is that if more than one virus is present the
reactions which develop on indicators may be attributed to one cause only rather
than several.

The identification of disease relationships has been based largely on symptom
comparison. Attempts have been made to apply the technique of cross protection,
but this without corroborative evidence is not entirely reliable. Symptom expression
even in a single strain can vary under differing environmental conditions and almost
every virus appears to exist in numbers of strains. In addition there is convergence
of symptom type by apparently distinct viruses. These points make it clear that

PROCEEDINGS OF THE LINNEAN Society oF NEw SoutTH WALES, 1958, Vol. Ixxiii, Part. 1.

10 VIRUS DISEASES OF CITRUS IN AUSTRALIA,

considerable caution must be exercised in the identification of viruses and that much
more exact information is required on all the diseases which are here discussed and
the range of variation which can occur. Technical difficulties also are associated
with the study of virus diseases of trees, which make progress in this field somewhat
slow. All citrus viruses so far known must be transferred by budding, grafting, or
by means of insects. No meehanical means of inoculation is available. All are
restricted to citrus, and citrus seedlings must be used as indicators.

Some of the diseases can take years to produce symptoms, necessitating considerable
space for trials, and making such investigations costly and time-consuming. The
process of testing and selecting varieties suitable for survey and indexing is slow,
and ensuring that the symptoms produced are of the virus under study, and are not
modified by the presence of other, perhaps latent, viruses, is also slow and sometimes
difficult.

Citrus in Australia: Native Species. 2

No species of the genus Citrus is native to Australia, but the closely related genera
of the Citrinae, Hremocitrus and Microcitrus, are represented by one and five species
respectively. The Australian desert lime Hremocitrus glauca is a shrubby species
native to arid western New South Wales and south-western Queensland, and Microcitrus
australis, M. australasica, M. inodora, M. maidenii and M. garrowayi occur, though
not commonly, in eastern Queensiand rainforest areas, the first two extending into
northern New South Wales. There is no evidence to suggest that any of the virus
conditions affecting citrus in Australia are endemic to this country. Three collections
of Hremocitrus glauca have been made from localities in western New South Wales
and seven of WMicrocitrus australasica from north-eastern New South Wales and
southern Queensland. All of these proved to be free of detectable virus infection.

Introduction of Citrus.

The introduction of citrus to Australia commenced with the first settlement of the
colony in New South Wales. Seeds and plants of a number of varieties were brought
by the first fleet and planted at Port Jackson in 1788 (Bowman, 1956), and in
subsequent years new and desirable varieties were introduced from all citrus-growing
countries. At first trees were usually grown from seed or layers but it was soon
found that trees budded on selected stocks were superior. By 1835 nurseries round
Sydney were producing thousands of such trees and the industry was flourishing.
There was at first no restriction on imports of plants and no examination for presence
of disease or pests, but since 1908 quarantine restrictions have slowed up the volume
of introductions and perhaps saved us from some troubles present commonly in
other lands.

Virus Diseases Known to Occur in New South Wales.

I will refer briefly to seven diseases shown to be virus in cause, which have been
found to be present in Australia. A number of other diseases, quite serious in their
effect, are almost certainly of virus origin but proof has not yet been obtained; these
will not be included. Finally, I shall very briefly refer to two virus diseases of
importance overseas which are not yet definitely known to be present here.

Psorosis.

This virus is bud transmissible and no insect vector is known. It is of very
minor importance in most citrus growing areas of Australia but is a major cause
of tree decline in the U.S.A. and Mediterranean countries and appears also to be common
in the Orient. The symptoms are scaling or exfoliation of bark in patches on limbs
and trunk, which does not begin to develop until the trees are eight to twenty years
old or older, and is associated with slow deterioration and death. A virus pattern of
flecks or oak-leaf markings occurs on immature ‘foliage in the spring and sometimes
in the autumn, and the disease can be diagnosed in affected trees by means of this
symptom before the scaling starts, and suspect trees can be indexed onto young
seedlings of orange and mandarin for the production of this symptom.

We have no record of the first entry of this disease into Australia, but locally
propagated trees now 45 and 50 years old are known to carry it.

BY LILIAN R. FRASER. 11

The common type of psorosis appears to be identical with that described as
psorosis A in California (Fawcett and Bitancourt, 1943) but the range of symptom
development is extreme. In some orchards some affected trees start to produce
‘Sealing at 12 years and almost all trees show this symptom by the age of 20 years.
A number of orchards are known where scaling was observed for the first time in a
few trees at the age of 25 or 26 years, but the great majority are still vigorous
and productive at 48 to 50 years, though the presence of leaf patterns in the spring
proves that they also are infected. At least one orchard is known where the trees
are more than 40 years of age and no bark symptoms have occurred in any of them,
though leaf patterns are produced. One case has been investigated of trees known
to have been propagated from a single bud source. Of 63 trees 35 years old only
three have well-developed bark symptoms but all show strong leaf pattern development
‘and are not otherwise affected. The reason for this variability is not known, but
the simplest explanation is that the virus exists as a number of strains of differing
virulence.

The relative unimportance of psorosis in Australia can be attributed in part to
the fortunate chance that few infected trees were imported in the early years of
eitrus growing, in part to the restrictions which in recent years have limited the
free importation of budwood, and in part to the activity of a bud-selection society in
New South Wales which was controlled by the nursery industry with assistance from
the Department of Agriculture and operated from 1935. Though certification was
largely on agronomic characters, the parent trees chosen had to be mature and of ~
good habit and no budwood was cut from trees showing symptoms of disease. Since
1953 a more rigid bud registration scheme has been in operation which guarantees
freedom from this disease. In Queensland since 1934 budwood from disease-free
‘sources has been provided for nurserymen by the Queensland Department of Agriculture.

No citrus species or varieties are known which are not susceptible to infection,
but lemons, Seville oranges, rough lemon and possibly others can carry the virus
without production of bark symptoms.

The original home of psorosis is not known, but may have been in Asia, where
it is believed to be common.

Lemon Crinkly Leaf.

This virus is bud transmissible and no vector is known. It affects lemons,
producing crinkled foliage and protuberances on the fruit rind. Sometimes the whole
tree may be affected but more often symptoms are restricted to a few or only one
limb. There is evidence of the existence of strains of differing severity (Fraser,
unpublished). Trees affected by a severe strain have a somewhat upright habit and
reduced vigour, other strains have a negligible effect on vigour and productiveness.
It has a somewhat restricted distribution in New South Wales and Victoria, and has
been recorded in Western Australia on trees imported from New South Wales. It
was described originally from California (Fawcett, 1936; Fawcett and Bitancourt,
1943), and was included in the psorosis group. No information is available relating
‘to its presence in other countries.

Seedlings of Hureka lemon are useful as indicators for crinkly leaf if no other
virus is present. ‘Six weeks after inoculation small star-like spots occur in immature
leaves, followed by crinkled growth as the leaf enlarges and matures. Sweet orange,
Mandarin and rough lemon seedlings inoculated with crinkly leaf develop a few
star-like spots in the young foliage and occasionally one or two crinkled areas, but
for all practical purposes are symptomless.

Enation.

This virus is transmitted by budding, and very readily by the tropical citrus
aphid Toxoptera citricidus and probably by other aphids also. It was described
originally from California (Wallace and Drake, 1953) and later from South Africa
(McClean, 1954) where it was found during surveys for tristeza virus. In Australia
-also its presence was detected during surveys for other diseases (Fraser, unpublished).
Small enations, or outgrowths on lower surfaces of the main veins, are produced on

12 VIRUS DISEASES OF CITRUS IN AUSTRALIA,

seedling rough lemon and, if no other virus is present, on grapefruit, Seville orange:
and acid limes. In sweet orange and mandarin there is no reaction, or at most a very
weak slight development on very young rapidly growing seedlings. It appears to be
widespread in New South Wales but of no discernible importance. It is likely that
strains of different virulence occur since some isolates produce many large enations.
on rough lemon seedlings and others few and small.

Woody Gall.

This condition has been under observation in New South Wales for some years:
and has been shown to be due to a bud transmissible virus (Fraser, unpublished). Its:
prevalence in several naturalized seedling communities of rough lemon suggested that.
it has an active vector. The symptoms are woody galls on branches older than one
season and on the main trunk and on the upper roots. These are smooth, covered
with normal bark and may be single rounded swellings or large-knobbed or cauliflower--
like structures. It is possible that strains of differing virulence occur since some:
affected trees have only a few minor swellings and in others the swellings become
large. Orange and mandarin trees on affected stocks show no gall development even
when substantial outgrowths are present on the stock below the bud union. No
significant reduction in vigour of the scion is associated with infection of rootstocks,,
but affected rough lemon seedling trees are lacking in vigour. It is common in many,
localities. Rough lemon seedlings have been used as indicators.

Scaly Butt of Poncirus trifoliata (Hxocortis).

This virus is bud transmissible and no insect vector is known (Benton e#¢ al., 1950,.
1952). It is carried symptomlessly in orange, mandarin, grapefruit and rough lemon,
but in Poncirus trifoliata and some of its hybrids the virus produces a bark-scaling
condition which is latent for two to eight years following inoculation. Infected trees
on susceptible rootstocks are considerably stunted. The spread of the disease is by
means of infected budwood. No citrus varieties have so far been found which can
be used in the seedling stage as indicators for demonstrating the presence of this.
virus. Consequently P. trifoliata itself or one of its hybrids must be used. P. trifoliata
is completely resistant to root rot caused by the fungus Phytophthora citrophthora and
since 1942 has been in considerable demand in New South Wales as a stock for oranges,
mandarins and grapefruit for use in areas where root rot is liable to occur. The:
presence of scaly butt virus carried symptomlessly in some scion clones therefore:
becomes of considerable importance.

Control of the disease has been effected by a bud registration scheme, guaranteeing”
freedom from virus. Since the scaly butt virus is symptomless in trees on stocks
other than P. trifoliata only trees on this stock are registered, and these must be
older than ten years, so that it is certain that the virus is not present.

From field observations it appears that several strains of scaly butt occur, differing
in severity and type of scaling, time of onset, and the degree of stunting which is
produced in the scion tree. In addition there is a type of stunting of trees on
P. trifoliata with which no scaling is associated and it is eseHme that this also may
be due to a strain cf the scaly butt virus.

In New South Wales the virus appears to be universal in Hureka lemons and
rather common in Washington and Thompson navel oranges, less so in Valencia oranges
and grapefruit, and very unusual in mandarins. It occurs in North (Fawcett and
Klotz, 1948) and South America (Knorr et al., 1951), but no information regarding
its possible presence in the Orient is available. Its time of introduction cannot be
stated and very little information can be obtained from current plantings, since prior
to 1940 the susceptible P. trifoliata stock was of very minor importance. However, a
few infected trees of Valencia and Washington navel orange propagated locally are
known, which are more than 50 years of age. <A similar disease of Rangpur limes.
which has been attributed to the same virus has heen described in South (Rosetti,.
1955; Moreira, 1955) and North America (Olsen and Shull, 1956; Reitz and Knorr,.
1957).

BY LILIAN R. FRASER. 13

Tristeza.

Tristeza is a virus disease which has its most serious effect on orange and
mandarin trees on Seville orange stock. Trees may wilt and die in a few months after
inoculation or may linger on for a number of years making very poor growth before
they finally die. Its principal vector is the tropical citrus aphid, Toxoptera citricidus,
but a number of other aphids can also transmit it with a rather low degree of
efficiency.

The disease affects chiefly orange and mandarin varieties on Seville rootstocks.
On their own roots, on rough lemon, P. trifoliata and a number of others, no symptoms
are produced—the varieties are symptomless carriers. A number of other varieties
used as rootstocks, including grapefruit, induce a tristeza effect in an affected scion,
but combinations such as grapefruit on Seville orange or lemon on Seville orange
do not decline to the same extent or in the same way, and seedling unbudded Seville
oranges do not suffer. Various theories have been put forward to explain this, but
none are entirely acceptable, or account for all the anomalies seen. The history of the
investigation which led to the recognition of this disease has been well reviewed by
Webber (1943), Wallace (1956), and Bennett and Costa (1949). It is presumed that it
is endemic in oriental countries, to which the tropical citrus aphid is also endemic,
but ne surveys have yet been made in India or China which might throw light on
its place of origin. The recognition of the virus condition is dependent on the use
of the susceptible stock and there seems to be no evidence that Seville orange is or
has ever been in use as a stock in the Orient.

Seville orange came into prominence as a stock first in Spain during the middle
of last century, when it was found to be very resistant to the fungus root rot caused
by species of Phytophthora (Fraser, 1949). It was widely used in Mediterranean
regions, where it is at present the major rootstock, and in South America and
California.

Wherever its place of origin may have been, it is clear that tristeza virus and
its vector must have become established in South Africa at an early stage (McClean,
1956; Webber, 1943).

One of the greatest epiphytotics of recent times occurred following the introduction
of the virus into Argentina about 1930. In the following decades 7,000,000 trees of
the susceptible sweet orange on Seville orange stock were destroyed in Argentina
and an equal number in Brazil (Costa, 1956). In California an outbreak which is still
continuing commenced in 19398 (Wallace, 1956).

Tristeza has been present in Australia, though unrecognized, for very many years,
possibly from the time of the first establishment of this erop. Seeds and plants of
citrus were obtained during the voyage of the first flest, from Rio de Janeiro, but
tristeza virus was not present in South America at that time. The ships also
touched at the Cape of Good Hope on this voyage, and though there is no specific
reference it is quite possible that further citrus trees or fruit would have been
obtained from the garden of the Hast India Company there. If introductions were
not made then, certainly they would have occurred during later voyages. Suttor
in 1843 (quoted by Bowman, 1956) mentioned having worked with success China
orange to Seville orange stock, though these may have been seedling trees and
therefore virus-free, and Shepherd in 1851 mentioned the production of Seville orange
as a stock in the nursery.

By 1870 heavy losses of citrus trees round Sydney had been sustained from
root rot. The resistance to root rot of Seville orange stock was known and its use
advocated to overcome the disease, and it is clear that if it had proved generally
successful it would have been widely used, but this was not so (Fraser, 1949).
Alderton (1884) mentions that Seville orange, though admittedly a hardy tree, was
not favoured as a rootstock because of the poor growth of trees worked onto it, and
Mackay (1875), though advocating the use of Seville orange on theoretical grounds,
pointed out that growers were not unanimous on the subject of the best stock. The
general lack of enthusiasm for this stock can be taken to indicate its unsatisfactory
behaviour even at that time, since in the absence of tristeza, Seville orange is an

14 VIRUS DISEASES OF CITRUS IN AUSTRALIA,

extremely vigorous and desirable stock. In 1890, giving evidence at an enquiry into
the citrus industry, Pye, a leading grower in the Parramatta district (mear Sydney),
specifically states that Seville orange stock should never be used as it had proved
a complete failure. Pye also made the first reference to the vector, the citrus aphid,
as being a common pest at that time. The introduction of a virus of this type into a
plant community cannot have its most devastating effect unless the vector is also:
present.

It seems reasonable to assume, therefore, that both the virus and its vector had
been introduced at an early stage and infection had spread through the citrus
population, undetected because trees were often on their own roots, seedlings or
layers. When working onto rootstocks became the practice, the general preference
for rough lemon stock is noted by all early writers.

There have been citrus nurseries round Sydney since the early days of citrus:
culture, and when new areas were opened up in other parts of Australia, trees from
these nurseries were in demand. Hence the virus was spread, unwittingly in
symptomless carrier trees, to new settlements in Queensland, Western Australia and
New Zealand. It is generally accepted that the epiphytotic which worked its way
through South America was started by the introduction in 1930 of infected trees.
on rough lemon stock from South Africa (Wallace et al., 1956). At this time also
several large shipments of orange trees were sent from Sydney to growers in
Argentina (Eyles, private communication) and these would undoubtedly have carried
tristeza, so that Australian trees may well have contributed to the South American
disaster. As McClean (1939) has emphasized, the introduction of the vector is as
important as that of the virus. This also could have come from New South Wales.

In the years following 1890 a very considerable expansion of citrus growing took
place in the very isolated district of Mildura on the lower Murray River, with the
introduction for the first time in Australia of irrigation for fruit production in an
arid area. The Chaffey Bros., irrigation engineers from California, who started this.
venture, introduced budwood of favoured citrus varieties from California and also
seed for stocks, the most favoured being the Seville orange, a variety now known
as bitter sweet, and the bitter rough being also used. These were propagated locally
and Seville orange stock was found to be extremely satisfactory and a number of
excellent orchards were established. Its resistance to the fungus Phytophthora
citrophthora enabled these trees to flourish where trees on rough lemon or sweet.
orange gradually deteriorated with root rot. Further propagations were made and
the stock enjoyed considerable local popularity. However, by 1942 some of the trees
were starting to decline and now no satisfactory orchards on this stock exist. This.
decline has been attributed to the introduction of tristeza virus in the Robertson
navel from California (McAlpine ef al., 1948). This, however, is unlikely. A tree of
Robertson navel propagated from the originally introduced budwood was located at
Curlwaa in south-western New South Wales in 1948 and this proved to be free of
tristeza. However, in addition to locally propagated trees, many young trees on rough
lemon or sweet orange stock were brought into the lower Murray district from Sydney
nurseries in the early days of the settlement and these would certainly have carried
the virus. The later spread of infection to trees on Seville orange stock most probably
was the result of increased activity of aphids.

This behaviour of Seville orange as a stock completely satisfactory in one district
and as complete a failure in others was a challenge and a mystery to horticulturalists
until the virus nature of the disease was established.

Although a very great deal of information has been obtained about tristeza, the
full story of its relationships has not yet been told. Further comments on tristeza
virus distribution and reaction and seedling indicators follow the discussion of
stem pitting.

Stem Pitting of Grapefruit.

This disease is readily transmitted by budding and Toxoptera citricidus is an
efficient vector. It was described first in South Africa (Oberholzer, 1949) and later

BY LILIAN R FRASER. 15

found to be almost universal in Australia. The symptoms in the most severe form
of the disease include pitting of the trunk and branches with deep furrows, often
somewhat contorted (small furrows or pits can occasionally be seen on young twigs if
the bark is removed), reduced vigour, poor growth and fruit which are reduced in
size, flattened or lopsided and thick-skinned. Symptom expression is, however,
extremely variable, sometimes one growth feature predominating, sometimes another,
so that in the field trees are encountered which have deeply pitted contorted trunks
but with reasonably good fruit, others with a high proportion of very poor fruit
and little or no visible pitting. Many variations and combinations of symptoms
occur. The test seedling used for indexing for the presence of this virus is the West
Indian or Key lime (McClean, 1950). On this the virus produces small pits or furrows
on the xylem surface and flecks or cleared areas in the veins of the leaves. The
identification of the virus causing symptoms in lime with that causing the disease
of grapefruit is based on the fundamental similarity of the symptoms and on the fact
that inoculation from limes back to virus-free grapefruit seedlings reproduces
symptoms of stem pitting and fruit distortion in them. Using the West Indian lime
as an indicator, surveys of New South Wales citrus have shown the stem pitting virus
to be far more widespread than the visible symptoms would suggest. Many completely
symptomless grapefruit produced when inoculated on lime seedlings a mild type of
pitting and occasional vein flecks. It is assumed from this that there are strains of
stem pitting virus so mild that no field symptoms sre produced by them even in the
most susceptible varieties. Also, on the basis of the lime reaction, it has been
determined that the stem pitting virus is present in New South Wales in all citrus
of all varieties except Poncirus trifoliate and some of its hybrids, with the exception
of a number of orange and grapefruit trees in south-western New South Wales
propagated from the original Chaffey introduction. In this area, however, the disease
is spreading and its progress is being watched in a number of grapefruit groves.

Most citrus varieties carry the disease quite symptomlessly. In Australia symptoms
of pitting are only produced on grapefruit and rarely on smooth Seville orange and
Bengal citron.

The severity of symptoms produced in indicator seedlings of West Indian lime
varies considerably. Buds from symptomless grapefruit produce, in all cases indexed,
the mildest reaction encountered on lime. In general the most severe reactions are
obtained from trees showing the severest pitting and stunting, but the correlation is
not exact. It is possible that this could be explained by the presence of a number
of different strains in the one tree. :

No direct control measures are possible for a disease of this nature. The
perennial host, the widely distributed virus and the ubiquitous vector make protection
from infection impossible and it is only a matter of time before all grapefruit trees
in New South Wales will be infected. The question therefore arises, can grapefruit
be successfully produced in Australia in the future? Two possible means are being
investigated: (1) the location of a grapefruit variety which is not damaged by the
presence of the virus, i.e., a tolerant type, and (2) mild strain protection. In many
viruses the presence of one strain delays or prevents the establishment of a second
strain of the same virus. Whether this is practicable for a perennial crop constantly
subjected to reinfection can be determined only by field experiment. The possibility
of mild strain protection for a perennial crop was investigated by Posnette (1947) in
the case of cocoa swollen shoot virus but he reported only partial success with the
method.

Relationship between Tristeza and Stem Pitting.

Proof of the virus nature of tristeza was first obtained by using as test plants
budded trees of sweet orange on Seville orange rootstocks (Fawcett and Wallace,
1946; Meneghini, 1946). Later, Costa et al. (1950), using a range of seedling and
budded trees, obtained on lime the reaction of vein flecking and pitting described
by McClean (1950) as a reaction to infection by stem pitting virus. They suggested
that this reaction was also that of tristeza, and as a corollary that tristeza and stem

16 VIRUS DISEASES OF CITRUS IN AUSTRALIA,

pitting are different effects of the one virus. This has been generally accepted, and
the lime reaction has been used to index citrus for the presence of tristeza in many
parts of the world.

During the course of a survey of citrus in New South Wales, using a range of
seedling varieties as indicators, a type of reaction was found which appeared to be
quite distinct from the vein clearing, pitting reaction on lime. It was obtained with
buds from oranges, mandarins, rough lemon, Rangpur lime and some tangelos and
calamondin. When these were inoculated to Seville orange, grapefruit, lemon and citron
seedlings, there was produced very severe reduction of growth, yellowing and, in
the scion, a yellow tristeza-like effect. This was considered to be a distinct virus
from that causing the lime reaction and it was named seedling yellows (Fraser,
1952). The lime reaction virus was present as well in all trees indexed and it has
not yet been possible to separate yellows from this virus. The same reaction had
been seen by Costa et al. (1949) but not considered to be different from the lime
reaction. Mature trees of grapefruit, lemon, Seville orange and some other varieties
have never been found to be infected with the yellows virus and appear to be
susceptible to infection only as very young seedlings. As pointed out by McClean and
Van der Plank (1955) the yellows reaction is undoubtedly a tristeza reaction. This
at once brings into question the status of the stem pitting and vein flecking reactions
of lime. A number of theories have been put forward.

Costa (1956), Knorr and Price (1957), and others regard the West Indian lime
reaction as a symptom of tristeza virus and yellows as an associated symptom.
McClean and Van der Plank (1955) and McClean (1956) put forward the view that
tristeza disease has two components, yellows and stem pitting, and that these are most
probably related strains of one virus. Wallace (1957) holds somewhat similar views.

Results of experimental work in New South Wales favour the view that the
seedling reaction of yellows and that of stem pitting and vein flecking are caused by
distinct viruses. Virus isolates of stem pitting alone do not cause a tristeza type of
decline when inoculated into grafted seedlings composed of sweet orange top and
Seville orange rootstock. The most severe strains, obtained from grapefruit trees
showing strongly developed symptoms of pitting, stunting or fruit distortion, reduce
the vigour of growth of such indicator trees but do not cause the characteristic
yellowing and deterioration of the scion. The mildest strains of stem pitting do
not reduce vigour, and strains of intermediate virulence have an intermediate effect
on vigour of indicator trees.

When, however, buds containing the yellows-producing virus are used to inoculate
indicator trees of sweet orange on Seville orange rootstocks, the scion ceases to
grow, turns yellow and ultimately dies. These reactions appear to be distinct, and the
direct reactions of seedling indicators also appear to be distinct and without inter-
grading forms to suggest they are related.

It is considered, therefore, that the lime reaction of vein clearing and xylem
pitting is a seedling reaction to stem pitting virus and should not be used without
corroboration as a test for the presence of tristeza disease in a citrus community.

The use of the word virus complex by McClean (1956) and McClean and Van der
Plank (1955) carries the implication that both components contribute to the severe
tristeza reaction. There is, however, no evidence available which can cast any light
on this question and the solution must await the separation of yellows from admixture
with stem pitting.

Viruses Not Known to be Present in Australia.

Stubborn.—This disease was described first by Fawcett (1946) in California. Grape-
fruit and orange varieties, particularly Washington navel, are affected. Trees lack
vigour, fruit is small and sometimes misshapen, often with the rind at the distal
end thinner than that at the stalk end.: Symptoms are somewhat variable; not all
the fruit on an affected tree show symptoms each year. The foliage is somewhat
chlorotic, growth is short and affected with die-back and the development of multiple
buds gives a bushy appearance to the trees. The disease is widespread also in

BY LILIAN R. FRASER. 17

Morocco and possibly other Mediterranean citrus growing countries. No satisfactory
indicator is known and the disease may take a considerable time to develop.

Xyloporosis.—The virus disease first observed in Palestine in 1928 (Reichert and
Perlberger, 1931) affects particularly the sweet lime, both as an unworked seedling
and as a stock for sweet orange and other varieties (Reichert et al., 1953).

The symptoms on sweet lime are as follows: Pits develop on the xylem surface
of the wood of the stock. These are rounded or conoid in shape and have corresponding
pegs on the bark inner surface. A brown discoloration occurs in the bark. As the
disease progresses the wood becomes impregnated with gum, bark cankers develop
and the tree ultimately dies.

Growth of sweet orange on this stock is poor, with considerable dieback, leaves
are small, upright and yellowish. Xyloporosis also occurs in South America (Fawcett,
1937; Knorr et al., 1952) and is common and widespread in Florida, where it affects
principally the Orlando tangelo, mandarins and kumquats, but is symptomlessly
carried in grapefruit and orange varieties (Childs, 1952).

Xyloporosis and stubborn disease have nothing in common and there seems to be
no possibility that they are related, yet different aspects of the grapefruit stem
pitting disease have something in common with each.

The problem of separating and identifying these different viruses should all be
present in the citrus community offers great difficulty particularly in the case of
stubborn, for which no satisfactory indicator has yet been found.

Future Outlook.

The means which have to be adopted to safeguard the citrus crops of Australia
from inroads by virus disease are of several kinds.

1. Where the virus has no known insect vector and is spread by the use of infected
budwood it is necessary to ensure that material used in propagation is virus-free.
The diseases which fall into this category are psorosis and scaly butt. The principal
bark symptoms of psorosis may not become apparent for many years, but the
associated foliage symptom is evident from the earliest stage. Seedlings inoculated
with psorosis-infected budwood develop these symptoms and so a means is available
of checking the health of budwood sources. Symptoms of scaly butt and associated
stunting of P. trifoliata take two to eight years to develop, possibly longer in some
cases. Trees on this stock, more than eight years of age, of good vigour and stock
character, can be assumed to be free of scaly butt virus, and can safely be used as a
source of budwood. There are some varieties, however, of which no suitable trees on
P. trifoliata are known, and in order to obtain scaly butt free budwood the only course
as yet available is to index a number of selections onto P. trifoliata and grow the
trees until they are old enough to produce symptoms.

Lemon crinkly leaf appears also to fall into the category of diseases which can
be eliminated by the use of virus-free budwood taken from trees free of symptoms,
and if necessary indexed onto seedling lemons.

2. Where an active insect vector and a reservoir of virus are present no purpose
is served by the use of virus-free propagation material. The presence of the virus
must be accepted and its effects minimized if possible. In the case of tristeza, control
is achieved by the use of non-susceptible scion-stock combinations. .In the ease of stem
pitting, where the most susceptible variety is also a favoured commercial one for
which no substitute is at present available, the only hope seems to lie in the
possibility of mild strain protection.

The enation and woody gall viruses would appear to be of such slight importance
that no action against them would be justified.

The introduction of citrus varieties from overseas poses a difficult problem for the
pathologist. To ensure that no new disease is brought into the country all intro-
ductions should be tested for the presence of all known viruses. This would take a
considerable time and would involve more expense than is justified for the results
obtained, and might still permit tbe entry of undescribed or unknown viruses. For
in addition to those which are known, there are the host of disorders which are not

B

18 VIRUS DISEASES OF CITRUS IN AUSTRALIA,

yet proved to be, but almost certainly are, virus in nature. Hach may be unimportant
from a world standpoint, but is perhaps Jocally important, or potentially so if
disregarded.

There are, however, ways of eliminating virus infections from plant material.
With one exception (Xyloporosis, Childs, 1956) citrus viruses are not known to be
seed-borne, so that seed can safely be introduced. Most citrus varieties produce a
proportion of nucellar seeds, so that by careful selection of seedlings, types closely
approximating to the parent can be obtained. There are, however, some varieties
which produce no nucellar seed, and others with particular agronomic qualities which
are not transmitted to their nucellar offspring. For such varieties the possibility of
elimination of virus infection by heat therapy holds some promise. Grant (1957) has
recently shown that psorosis virus can be eliminated by growing an infected plant
at a continuously high temperature for three months or more and it is possible that
further investigations will show that other viruses may similarly be eliminated,
perhaps paving the way for the development of a standard treatment which could
be given to imported material to eliminate the hazard of importing disease as well.
A third and perhaps even more promising line of attack is based on the fact that
the embryonic tissue of the growing point is not invaded by virus. It has already
been shown that for some plants propagations from growing tips will eliminate virus
disease. This possibility should certainly be explored for citrus viruses also.

Literature Cited.

ALDERTON, G. E., 1884.—Treatise and Handbook of Orange Culture. Auckland, New Zealand.
BENNETT, C. W., and Costa, A. S., 1949.—Tristeza Disease of Citrus. Journ. Agricultural

Research, 78: 207-237.

BENTON, R. J.. BowMAN, F. T., FrAspr, L., and Keppy, R. G., 1950.—Scaly Butt and Stunting
of Citrus. Dept. Agriculture N.S.W., Science Bulletin No. 79.

—— ; , 1952.—Significance of Trifoliata for Citrus Rootstock
Problems. Report of the 13th International Horticultural Congress, Vol. 2: 1235-40.
BowMan, F. T., 1956.—Citrus-Growing in Australia. Sydney.

CHILbs, J. F. L., 1952.—Cachexia Disease, its Bud Transmission and Relation to Xyloporosis

and to Tristeza. Phytopath., 42: 265-268.

, 1956.—Transmission H!xperiments and Nyloperosis-Cachexia Relations in Florida.

Plant Disease Reporter, 40: 143-145.

Costa, A. S., 1956.—Present Status of the Tristeza Disease of Citrus in South America.

F.A.O. Plant Protection Bulletin, 1V: 97-105.

Costa, A. S., GRANT, T. J.. and Moreira, S., 1949.—Investigacoes sobre a tristeza des citrus,

Il. Bragantia, 9: 59-80.

—- ; ——, , 1950.—A Possible Relationship between Tristeza and the Stem-

pitting Disease of Grapefruit in Africa. Calif. Citrog., 35: 504, 526-528.

Fawcett, H. S., 1936.—Cilrus Diseases and their Control. New York.
, 1937.—Contacts with the Citrus Industry and other Observations in South America.

Calif. Citrograph., 22: 552-5538, 571-572, 575.

, 1946.—Stubborn Disease of Citrus, a Virosis. Phytopath., 36: 675-677.
Fawcett, H. S., and Birancourt, A. A., 19438.—Comparative Symptomatology of Psorosis

Varieties on Citrus in California. Phytopathology, 33: 837-864.

Fawcert, H. S., and Kuorz, L. V., 1948.—Bark Shelling of Trifoliate Orange. Calif.

Citrograph., 33: 230.

Fawcert, H. S., and WaAuLacn, J. M., 1946.—Hvidence of the Virus Nature of Citrus Quick

Decline. Calif. Citrograph., 32: 50, 88-89.

FrAseR, L. R., 1949.—A Gummosis Disease of Citrus in Relation to its Environment. Proc.

LINN. Soc. N.S.W., 74: 5-18.

, 1952.—Seedling Yellows, an Unreported Disease of Citrus. Agr. Gaz. N.S.W.,

63: 125-131.

Grant, T. J., 1957.—Effect of Heat Treatments on Tristeza and Psorosis Viruses in Citrus.

Plant Disease Reporter, 41: 2382-234.

Knorr, L. C., and BENATENA, H. N., 1952.—Xyloporosis en Mandarino Comun de Concordia.

Idia, 5: 19-20.

Knorr, L. C., DuCHARME, EH. P., and BAnri, A., 1951.—Exocortis in the Citrus Groves of

Argentina. JIdia, 4: 8-12.

Knorr, L. C.. DuCHARME, EH. P., and Buspy, J. N., 1954.—Discovery of Exocortis in Florida

Citrus. Plant Disease Reporter, 38: 12-138.

Kxnorr, L. C., and Price, W. C., 1957.—Is Stem Pitting of Grapefruit a Threat to the Florida

Grower. Citrus Industry, 38: 11-14.

BY LILIAN R. FRASER. 19

McALPIN, D. M., Parsal, P. S., Ropprts, R., and Hopr, R. H., 1948.—‘*Bud Union Decline”
Disease of Citrus. Journ. Dept. Agric. Victoria, 46: 25-31.

McCLeAN, A. P. D., 1950.—Virus Infections of Citrus in South -Africa III. Stem Pitting of
Grapefruit. Farming in South Africa, 25: 289-296.

—————, 1954._Citrus Vein Mnation Virus. S. Afr. Jour. Sci., 50: 147-151.

, 1956.—Tristeza and Stem-pitting Diseases of Citrus in South Africa. F.4A.0. Plant

Protection Bulletin, 4: 88-94.

——, 1957.—Virus Infections of Citrus Trees. Jbid., 5: 133-141.

McCLEAN, A. P. D., and VAN DER PLANK, J. H., 1955.—The Role of Seedling Yellows and Stem
Pitting in Tristeza of Citrus. Phytopath., 45: 222-224.

Mackay, A., 1875.—The Australian Agriculturist.

MENEGHINI, M., 1946.—Sobre e natureza e transmissibilidade “da doenca’’ ‘‘tristeza’” dos
citrus. Biologico, 12: 285-287.
Moreira, S., 1955.—A molestia “Exocortis’’ e o cavalo de limoeiro cravo. Rev. Agric.,

Piracicaba, 30: 99-112.

OBPRHOLZER, P. C. J., MatTHews, I., and STipMin, S. F., 1949.—The Decline of Grapefruit
Trees in South Africa. A Preliminary Report on So-called Stem-pitting. Union S. Afr.,
Dept. Agr. Sci. Bulletin, 297.

OLSEN, H. O., and SHuLL, A. V., 1956.—Hxocortis and Xyloporosis—Bud Transmission Virus
Diseases of Rangpur and other Mandarin-Lime MRootstocks. Plant Disease Reporter,
40: 939-946.

Posnette, A. F., 1947.—Virus Diseases of Cacao in West Africa. Ann. Appl. Biol., 34: 388-402.

Pys, T., 1890.— Bull. No. 1, N.S.W. Dept. Agriculture.

REICHERT, I., and PERLBERGER, J., 1931.—Xyloporosis, the New Citrus Disease. Hadar,
7: 163-167, 172, 183-202.

REICHERT, I., YORPFE, i., and BENTAL, A., 1953.—Shamouti Orange on Various Rootstocks and
its Relations to Xyloporosis. Palestine Journ. Botany, Rehovot series, 68: 163-183.
Reitz, H. J., and Knorr, L. C., 1957.—Occurrence of Rangpur Lime Disease in Florida and

its Concurrence with Exocortis. Plant Disease Reporter, 41: 235-240.
Rossetti, V., 1955.—A doenca do lomoeiro eravo nos laranjais de Sao Paulo. Biologica,

21: 1-8.
WALLACH, J. M., 1957.—Tristeza and Seedling Yellows of Citrus. Plant Disease Reporter.
41: 394-397.

—, 1956.—Tristeza Disease of Citrus, with Special Reference to its Situation in the
United States. F.A.O. Plant Protection Bulletin, 4: 77-87.

WALLACE, J. M., and DRAKE, R. J., 1951.—Newly Discovered Symptoms of Quick Decline and
Related Diseases. Citrus Leaves.

————_, ——_—,, 1953.—New Virus Found in Citrus. Calif. Citrograph., 38: 180.

WALLACE, J. M., OBERHOLZER, P. C. J.. and HormMryer, J. D. V., 1956.—Distribution of Viruses
of Tristeza and other Diseases of Citrus in Propagated Material. Plant Disease Reporter,
40: 3-10.

WEBBER, H. J., 1943.—The “Tristeza’’ Disease of Sour-orange Rootstock. Amer. Soc. Hort.
Sci., 43: 160-168.

BB

20

SEED COAT ANATOMY AND TAXONOMY IN HUCALYPTUS. L.
By EH. GAaurga and L. D. Pryor.
(Plate i; nineteen Text-figures.)
[Read 26th March, 1958.]

Synopsis.

Anatomical and histochemical investigations of the seed coat of Blakely’s two Renantherous
Sections of Hucalyptus aimed at revising these groups show for the bulk of the species a
remarkable consistency. The seed develops from an anatropous, bitegmic, crassi-nucellate
ovule. The salient testa characters are: the outer integument is several layered, with its
outer epidermis built up by sclereids, its inner epidermis crystal-bearing but without forming
a typical “crystal epithelium”; the inner integument persistent, two-layered and suberized.
An elaborately ramified vascular system extends from the basal hilum to the distal chalaza.

However, three of Blakely’s series, Ochroxylon, Steatoxylon and Myrtiformes, deviate
considerably from each of these testa features and vascularization pattern and probably are
not correctly placed in the Renantherae.

INTRODUCTION.

Since Bentham’s (1866) original treatment the taxonomy of Hucalyptus has been
based on various characters of the anther, such as shape, mode of dehiscence,
attachment to the filament and relative position of the gland. It has been obvious
more recently that some of the groups formed on this basis are not homogeneous.
It has now been found that the anatomy of the seed coat, together with some
morphological ovule and seed characters, promises to give valuable data for a regrouping
of some species.

The value of the study of seed coat anatomy is not confined to its application
to taxonomy, since it is likely also that seed coat structure and seed behaviour are
related.

Blakely’s “Key to the Hucalypts” is the most recent classification of the genus,
and undoubtedly the most satisfactory yet published. In such a large genus, however,
it is to be expected that even in so excellent a work there will be anomalies in the
arrangement of some of the species, and at different times some of these have been
suggested by investigations carried out in other ways. For example, Blake (1953)
has shown on general morphological grounds how the species belonging to the
Clavigerae as understood by him form a natural group, but in Blakely’s ‘‘Key” they are
partly separated. It has also been suggested (on the grounds of cotyledon shape)
that some species having bisected cotyledons and separated by Blakely in his “Key”
would more appropriately be placed together in the same taxon (Pryor, 1956).

Amongst all of Blakely’s groups, however. the combined group Renantherae and
Renantherae-Normales, containing about 100 species, is one which possesses a high
degree of homogeneity, and it seems clear that this group, largely as defined by
Blakely, might well be constituted a subgenus.

It was already known at the time this investigation began that in morphological
characters the seed of the great majority of species within this composite group had
features in common but at the same time these were to be seen seldom in species
placed by Blakely in other sections. It seemed likely, therefore, that this group would
be essentially a natural unit. It followed, therefore, that if a similar uniformity were
found in seed coat anatomy this would conform with this view. Further, if among
the species thus placed together by Blakely occasional exceptions to the common
‘pattern of seed coat structure were found, it might be deduced that these were
incorrectly included in the group. At this point, therefore, it appeared that an
examination of seed coat structure would be profitable and the study was commenced.

PROCEEDINGS OF THE LINNEAN Society oF NEw SoutTH WaArTKES, 1958, Vol. Ixxxiii, Part 1.

BY E. GAUBA AND L. D. PRYOR. 2

The seeds used in this investigation were mostly the result of our personal
collection, but some which were either rare or difficult to collect were obtained from
different State Forestry Departments, Botanic Gardens and Herbaria, to which we are
indebted for generous assistance. In a few cases seed from commercial sources has
been used.

The microscopic sections were made by hand without prior preparation, and the
drawings with a camera lucida.

Blakely (1955) has been followed in nomenclature and systematic grouping.

The purely anatomical aspects of the investigation considerably extend our
knowledge in this field of research on Hucalyptus. They have been linked with
previous publications on the subject, not all of which have been fully correct. ‘There
has been no previous reference in the literature to the taxonomic implications of
seed coat anatomy in Hucalyptus, and the conclusions in this regard are arrived at
entirely from the present work.

In the following, “testa” is taken to mean the seed coat derived from both seed
integuments. Published work on the anatomical structure of the Hucalyptus testa has.
up to the present been quite limited. In an extensive work, Petit (1908) investigated
the structure of 14 genera of Myrtaceae, including 59 species in the genus Hucalyptus.
However, he gave emphasis principally to the anatomical structure of the fruit wall,
both torus and pericarp, rather than to the seed coat. Only in 10 species was the testa
also examined, and of these four species alone were illustrated. One of these,
Hucalyptus macrocarpa (Fig. 22, page 51) is erroneous, illustrating, most probably, the
testa of one of the species of the Renantheraet The remaining nine species all belong
to the Section Macrantherae, sc that Petit’s work does not give a balanced presentation
of the distinct diversity of testa structure in the genus Hucalyptus. Raphe, chalaza
and the course and structure of the conducting tissue were not investigated. Nor
were the wall substances microchemically tested, so that the very interesting presence
of an inner cuticle and of suberized tissues in the seed coat (with their physiological
implications) escaped him.

Netolitzky (1926) gives a detailed and penetrating summary of seed structure for
any work dealing with seed anatomy. The author records all facts known at that
time and adds the results of his own critical studies. In particular with regard to
Hucalyptus he reports generally the results of Petit’s work and reproduces also,
amending the annotation, three of his figures, H. calophylla, E. globulus and #.
“macrocarpa”’. Harada (1956) has examined in no great detail ten Hucalyptus species,
but inaccuracies of observation and conclusion impair the value of the publication.’

In the fundamental works of Ferdinand von Mueller (1879) and J. H. Maiden
(1929), figures and descriptions of seed are presented purely from the morphological
viewpoint. Maiden gives a classification of seed according to colour, size, surface
structure, position of hilum and similar features. Some of these characters are better
understood, as will be seen later, if interpreted anatomically.

In addition we shall refer to other specific relevant points in the literature as
they arise.

Finally it may be said that, no matter how important the role of anatomy may
be in providing clues for taxonemic relationships, we are conscious of the fact that
anatomical features alone (and from one part of the plant only) cannot lead to final
conclusions until they are supported by evidence, especially morphological, from other
parts. However, characters like testa structure and the vascularization pattern of the
seed are features fixed by deep-seated factors of inheritance and scarcely affected by
environmental factors. Therefore they may provide valuable facts for the establishment
of taxonomic affinities.

1 This view results from the study of seed collected ourselves from H. macrocarpa growing
naturally near Wagin, Western Australia.

2Harada appears to be unaware of the precise delimitation of the two integuments.
Several times he presents only the outer epidermis as the outer integument, and designates
in text and figures the subepidermal parenchyma of the outer integument as the inner
integument. In surface sections he speaks of a “network of strips’ without realizing that
these are cross-sections of anticlinal epidermal cell walls.

22 SEED COAT ANATOMY AND TAXONOMY IN EUCALYPTUS. I,

SECTION RENANTHERAE-NORMALES.

The Outer Integument of the Ripe Seed.\—This is composed of several layers. The
outer epidermis is made up of a solid layer of sclereids with secondary, lamellated
thickenings on the outer and radial walls, the latter traversed by corresponding
simple or ramiform pit canals. ‘The excessively centripetally thickened walls leave
a very small lumen. They are mostly lignified, and this very strongly so in H. triantha,
HE. laevopinea, EH. macrorrhyncha, E. alpina and others, but no phloroglucin reaction
was obtained in H. haemastoma, indicating the absence of lignin. The size of the
epidermal cells is variable in one and the same seed. At the edges and ribs and around
the hilum the height of these epidermal cells can be many times the breadth
(macrosclereids). On the flat side of the seed they are more or less isodiametrical
or even frequently elongated parallel to the surface of the seed. They are relatively
small-in #. stellulata (Pl. i, fig. 4) and #. salicifolia, and by comparison very large
in #£. planchoniana, EH. laevopinea (Pl. i, fig. 1) and others. The outer cuticle is
missing, or at the most just discernible as fragmentary remnants as in EZ. oreades,
HL. kybeanensis, H. stellulata and #. piperita. The inner epidermis is made up of
small, thin-walled tabular cells. The number of layers of parenchymatous cells between
the two epidermal layers changes not only with the species but also with the position
on the seed at which the examination is made, that is to say, at ribs and corners, and
particularly along the raphe there is always a iarger number than on the flat side
of the seed. There are a few layers only in #H. sparsifolia, B. salicifolia and EH. oreades
(two to three), and they are numerous in H. planchoniana (six te eight).

lr Monoclinic crystals of primary calcium exalate monohydrate (identical with the
mineral whewellite) occur dispersed in the cells of the outer integument (excluding
the outer epidermis). They are most plentiful in the raphe parenchyma just over
the hilum, where they are arranged in vertical files accompanying the vascular bundle
(Text-fig. 4). They are often also more numerous in the ribs. There is special
significance in their localization in the inner epidermis.* In some species (H. piperita,
#H. pilularis, EH. planchoniana) they are missing or extremely rare on the raphe-free
sides. In #H. stellulata they occur only in the lower part of the seed. In #H. salicifolia
their occurrence is patchy. They are small and scarce in #. oreades but scattered
over the whole raphe-free sides. They are abundant in #. macrorrhyncha (Text-fig. 5)
and #. haemastoma (Text-fig. 1). In #H. sparsifolia nearly every epidermal cell accom-
modates a large, well-developed crystal. In H. muelleriana many of the crystal-bearing
cells have irregular, uneven wall thickenings which partly surround the crystals or
enclose them completely (Text-fig. 2). Furthermore, some of these crystals in the
hilar region have an additional cellulose membrane as an envelope (Text-fig. 3).

This discontinuous presence or complete absence of calcium oxalate crystals in
gererally unmodified cells of the inner epidermis of the outer integument is likely
to have taxonomic significance. As will be indicated later, this epidermis in other
sections of the genus is formed of cells strongly thickened at the base, each carrying
one or more crystals. In the Angiosperms the occurrence of this “crystal epithelium”’
is widely distributed, especially in the more primitive families, and is generally
considered in the phylogenetic sense as a primitive character. Within the Renantherae
the inner epidermis has, to a greater or lesser extent, lost its character as a “crystal
epithelium’’.

8 Strictly, one ought to speak of the derivative of the outer integument, but to simplify
discussion we shall refer to the two parts forming the seed coat as inner and outer integument.

4Tsolation of the inner epidermis exposing larger areas is necessary to obtain correct
information about their distribution. It is also important to compare raphe-free sides above
the hilar region because crystals are always present in the vicinity of the bundle, hilum
and micropyle.

5 Blakely’s Series Pseudo-Stringybarks comprises three species: EH. pilularis, which we found
erystal-free, H. muelleriana, where about half of all epidermal cells are crystal-bearing, and
E. wardii. Of the last-named we had only a few seed fragments from the holotype for
examination, revealing about the same frequency of crystals in somewhat irregularly thickened
eells as in H. muelleriana, though in a lesser degree. The hilar region could not be investigated.

BY E. GAUBA AND L. D. PRYOR. 23

As a whole the outer integument represents the “pigment layer’, and the ripe
seed displays, due to this, a brown to black colouring. There are dark-coloured
compounds, giving tannin reaction,® impregnating the walls and filling as amorphous
deposits the lumina of the epidermal cells, the parenchyma and the chalaza cork
tissue (Text-fig. 18). They are probably closely related to the phlobaphenes (anhydrides
of tannin) of the bark. In the White Mahoganies (#. triantha and EH. carnea)
impregnation of the walls of the outer epidermis is weak or even absent. On thin
sections they appear ivory white and give a strong phloroglucin reaction which is
partly masked in other species, due to the brown colouring by tannin impregnation.

As the result of this study of the structure of Hucalyptus seed we consider the
raphe as part of the outer integument and not—as it still is described in some recent
literature—as the fusion of the funicle with the testa. The parenchyma and the
epidermis of the raphe have the same structure and contents as the rest of the
outer integument. Thus in the Renantherous seed coat the same sclereid epidermis
covers the raphe as well as the remainder of the seed. Where the inner wall of the
outer epidermis cells is mucilaginous, as in H. marginate, or where palisade-like thin-
walled epidermal cells filled with solid dark-coloured material occur in other systematic
groups (#. maculata, H. citriodora) we find them also unchanged over the raphe in
wall structure and zontents.

The hilum is a clearly seen sear surrounded by a raised rim of sclereids. There
is no protective layer covering the exposed surface, which is neither cutinized nor
suberized, nor covered by a cuticle. Thus the exposed cells wear away, the surface
breaks up and in the course of time the hilum becomes more or less hollowed and
air-filled. This may rather retard than facilitate the intake of water for germination.

From the hilum an amphicribral vascular system with helically thickened tracheids
extends in the expanded raphe parenchyma right up to the chalaza (which is not
externally visible) where it finally spreads out (Pl. i, figs. 3, 4, and Text-fig. 19, 1).
Integumentary buné@les, i.e. a system of strands branching off the raphe bundle and
traversing the outer integument, do not exist.

The Inner Integument—This occurs immediately below the outer integument. In
ripe seeds the median cuticle delimiting the two integuments is resorbed. In the
genuine Renantherae the inner integument is two-layered, being formed of both
epidermal layers alone. The cells are tabular and without intercellular spaces. In
contrast with species in Blakely’s two series, Steatoxylon and Myrtiformes, in all
remaining series of the Renantherae-Normales this integument has not been resorbed
in the course of ripening. It is suberized‘ throughout (PI. i, figs. 1, 3, 4), the walls
being not merely impregnated with fatty substances. If after previous treatment with
Eau de Javelle, tests are made with zinc chloroiodide, brown-coloured delicate suberin
lamellae separate themselves from the primary cellulose walls which appear blue-
coloured. The walis are in addition more or less impregnated with tannin-like material.

The shape and size of the micropylar end of the inner integument vary considerably
within the same species. There is either a biunt or pointed tip without any sign of
an aperture, or the apex is lengthened to form a cylindrical or conical tube, the
endostome, with a straight or somewhat funnel-like dilated rim. The capillary canal
is blocked by reddish-brown material which also fills the surrounding cells (Text-figs.
8, 9, 10 and 11). Thus there is little chance of an appreciable intake of water or
nutrients through the endostome to the embryo once the suberization is completed.*

®The microchemical test for tannins in the wide sense of the term (which comprises
very different substances) is generally limited to ferric salts staining green or blue. This
of course does not reveal much about the precise nature of these compounds.

7 Besides the fat-staining compounds which can give ambiguous results, Hoehnel’s cerin
acid reaction was used to test the suberin lamellae.

§The exostome is occluded, most of the species showing—sub lente—a faint short fissure
or ridge between hilum and the micropylar region which could be considered perhaps as the
exostome suture. However, this needs further investigation. But in view of the fact that
moisture is readily absorbed by the whole cuticle-free epidermis it is of secondary importance
for germination as to whether or not the intake of water is effective through hilum and
micropyle.

24 SEED COAT ANATOMY AND TAXONOMY IN EUCALYPTUS. I,

Netolitzky’s opinion is that the main role of cork tissues and cuticles in Angiosperm
seed coats lies in the prevention of emigration of crystalloids (formed by mobilization
of the colloids during germination) from the embryo.

8 3 10 11
Text-figs. 1-11.

1-7: Monoclinic crystals of caicium oxalate monohydrate, ca. 180x.

Fig. 1, 2, 5, 6, 7: crystal-bearing inner epidermis of the outer integument in surface
view. 1: Eucalyptus haemastoma; 2: H. muelleriana; 5: H. macrorrhyncha; 6: E.
brachyandra, the crystal epithelium is seen iying beneath the outer epidermis; 7: H. guwilfoylei,
the crystal epithelium is viewed from the inside of the seed. Through the gap (g) in the
erystal layer (in the chalaza region) the tracheary endings (¢) of the raphe bundle are seen.

Fig. 3: H. muelleriana, crystals with cellulose envelopes in thick-walled cells.

Fig. 4: #. muelleriana, crystals in short files accompanying the raphe bundle.

8-11: Some endostome forms, ca. 180x. 8: Bucaiyptus muelleriana; 9: EH. oreades;
10: #. salicifolia; 11: H. gigantea. The forms are variable within the same species.

At the inner limit of the inner integument (and therefore of the testa in general)
an inner cuticle is encountered which is rather conspicuously developed and forms
rib-like projections between the nucellus epidermal cells (Pl. i, fig. 1), so that in

BY E. GAUBA AND L. D. PRYOR. 25

surface view this cuticle simulates cellular structure. It is the product both of
the inner integument and nucellus where their surfaces are in contact, and therefore
of double origin. However, the formation of this cuticle is suppressed in the chalaza
region.

The chalaza is reckoned also as part of the testa. It is the parenchymatous tissue
where nucellus and integuments merge. It is the pathway for nutrients to the
integuments and through the gap in the inner cuticle to the embryo and its associated
tissue. Once the supply to the embryo is completed this gap is closed by suberization
of neighbouring chalaza tissue and by filling the cork cells with dark-coloured solid
material (PI. i, figs. 3 and 4). The suberization is of the type of the inner integument,
therefore there is an inner suberin lamella alongside the cellulose wall. It must be
noted, however, that this “chalaza cork” is not of uniform origin, because not only
do the cells of the chalaza sensu stricto undergo suberization, but partly the neigh-

14

Text-figs. 12-18.
12-13: Diagrams of germinating seed. 12: Hucalyptus dives; 13: EH. microcorys — cd,
clinging disc; ch, chalaza cork; cr, crystal layer; h, hilum; hy, hypocotyl; ic, inner cuticle;
m, micropylar region; 7, radicula; vb, vascular (raphe) bundle.

14-17: Chaff structure. 14: H. iaevopinea, transy. section; 15: H. microcorys, transv.
section of chaff with ventral hilum; 16: EH. microcorys, transv. sec. of chaff with basal hilum:
17: E. guilfoylei, tang. sect. of outer epidermis of.chaff with ventral hilum showing the
blind pits in the outer wall.

18: EH. fraxinoides, transv. sect. of the cuter integument, the phlcbaphene-bearing cells
(pigment layer) shaded. All sections ca. 180x.

bouring tissue of the nucellus is also involved—often, indeed, the latter forms the
major portion of the whole “chalaza cork’. Furthermore, the suberization extends
also into the raphe parenchyma and adjoins the suberized inner integument (PI. i,
fig. 4), forming finally (in surface view) a large circular, oval or elliptical, dark-
coloured disc above the smaller cuticular gap. With some care it is possible to
extract the inner integument with the adherent chalaza cork revealing its size and
shape.

It is beyond the limits of the present study to make an extended histochemical
and structural investigation of the embryo and its supporting tissues and the alterations
they undergo during the maturation of the seed. It seems, however, that nucellus
and endosperm also in some circumstances can throw light on taxonomic relationships.

26 SEED COAT ANATOMY AND TAXONOMY IN EUCALYPTUS. I,

Therefore a few remarks about the final stage of these parts as seen in the mature
seed are justified.

In the resting seed there is a more or less broad remnant of the nucellus tissue,
especially in the chalaza region, the cells of which are empty and largely obliterated
so that without application of swelling media the cellular structure in general cannot
be distinguished (PI. i, figs. 1, 3, 4). The endosperm is resorbed, except in the micro-
pylar region, where an intact layer one cell thick and rich in proteins is retained
(2. muelleriana, H. alpina, H. sparsifolia, H. planchoniana, EH. oreades and others).
Testing the germination residue of H. planchoniana with Millon’s or Raspail’s reagent
reveals that these proteins have not been used during the germination. They represent
a reserve surplus.

Finally, the embryo is itself covered by a cuticle which, however, does not at this
stage form a tough skin but rather a semisolid envelope on which oil drops from the
embryo dislocated by sectioning remain attached. In the resting seed, therefore, it
has not reached its final consistency and it separates easily from the embryo epidermis.
The seed being exalbuminous (without endosperm) the food reserves for germination
are stored in the large embryo (cotyledons and hypocotyl) which fills completely the
cavity enclosed by the seed coat. Storage substances are fats (in the form of oil
droplets) and proteins (aleuron grains) .®

With the exception of the two series Steatoxylon and Myrtiformes the testa
structure described above is uniform in principle in all the series and subseries of the
Renantherae-Normales, almost all of which have been investigated.° In these groups
the following species were examined: H. pilularis, H. muelleriana, EL. umobra, HE. triantha,
E. laevopinea, EH. macrorrhyncha, H. alpina, EF. sparsifolia, E. obliqua, E. gigantea,
EH. planchoniana, E. oreades, HE. fraxinoides, H. kybeanensis, HE. pauciflora, E. stellulata,
EH. salicifolia, H. dives, H. tasmanica, HE. piperita and H. haemastoma. ‘There is little
doubt in assuming that Blakely’s categories, series and subseries, are essentially
homogeneous. On this basis, therefore, the material examined satisfactorily represents
94 species.

In summary, the following points are distinct and characterize the testa of the
Renantherae-Normales:

1. The outer integument consists of several layers, its outer epidermis is formed
of sclereids, its inner epidermis does not form a specific crystal layer.
2. The inner integument persists, is two-layered and suberized.
Whether this structure is exclusively confined to this group of Hucalyptus and may be
used as an index of relationship or whether it appears also in other systematic groups
as the result of convergent development will not be fully apparent until the investi-
gation of all the remaining sections in the genus is complete. For the moment we are
able to point out that, for example, the testa of #. patens and EH. diversifolia (Sect.
Renantheroidae) and that of H. jacksoni (Sect. Macrantherae) are in all histological
and morphological details precisely that of the pattern in the Renantherae-Normales.4
Relating structure with function of the seed coat, it is obvious that the closely-
packed epidermal sclereids confer high mechanical protection, while the biological
function of the tannin deposits in the outer integument no doubt lies first of all in
the chemical protection against decay. The strongly reduced permeability resulting
from the suberized tissues (inner integument, chalaza) and the inner cuticle, all
enclosing the embryo, indicates that in the mature seed the embryo has attained a
far-reaching physiological independence from its coat.

9 We have examined the aleuron grains in the embryo of H. marginata. ‘They include
numerous globoids and one druse (cluster crystal) of calcium oxalate each. Treatment with
picric acid and eosin also reveals a protein crystalloid (stained bright yellow) in the amorphous
protein substrate (stained red). Thus they represent the most highly differentiated form of
aleuron grains.

10Only the Subseries Seminudae has not been examined. The four species belonging to
this group are considered to be hybrids and authentic seed could not be obtained.

11Jt seems unlikely because of basic differences in testa structure that EH. jacksoni and
E. diversicolor are closely related. This is contrary to Blakely’s view since he places them
in the same Subseries, Inclusae.

BY E. GAUBA AND L. D. PRYOR. 27

In some respects the testa as a whole may be compared with the (outer) bark.
The suberized inner integument is a dead tissue and excretion of oxalate crystals
and phlobaphenes into the outer integument excludes its cells from metabolic activities.

The series Steatoxylon and Myrtiformes, both of which Blakely places in his
Renantherae-Normales, present a quite different type of testa.

E. microcorys, of the monotypic series Steatoxylon, has flat, bifacial somewhat
elliptical seeds. The coat is much reduced in thickness. The inner integument is in
the course of the ripening process resorbed and is missing altogether in the mature
seed (PI. i, fig. 6). Only in the chalazal region, in the corner between the crystal
layer and the inner cuticle, do a few non-suberized cells still appear. The outer
epidermis is formed of thin-walled cells whose outer wall is often sunken (concave)
or even torn and whose anticlinal somewhat wavy walis are partly separated from
each other so that the seed surface appears somewhat shaggy. The inner epidermis
forms a typical crystal layer in which each cell has a thickened basal wall and contains
one large crystal enveloped in a cellulose membrane and some smaller ones. All cell
walls of the integument give the tannin reaction with ferric salts. The major portion
of the nucellus remnant is obliterated. Traces of endesperm are still seen, especially
in the chalaza region, where, as with the nucellus remnant which also occurs there,
cellular structure can be perceived. Here one can also see that suberized cells of the
nucellus remnants contribute in a very high degree to the formation of the ‘chalaza
cork: (Pl i} fiz. 5).

E. microcorys is distinguished therefore from the normal Renantherous type
described above in the following ways:

1. The outer integument, except the raphe side, is only two-layered. The
outer epidermis is composed of thin-walled cells which are often torn and
separated from each other. The inner epidermis forms a typical crystal
layer.

2. The inner integument is resorbed.

The other exception appears in the series Myrtiformes, of which Z£. deglupta,
#H. brachyandra (Text-fig. 6 and Pl. i, fig. 9) and H. raveretiana were examined. The
structure of the testa is in principle like that of H. microcorys except that the epidermal
cells are of lesser depth, more elongated, with straight anticlinal walls, remaining
attached to one another, and the outer wall is not torn. The inner periclinal walls
of the crystal cells are only moderately thicker than the outer walls. The crystals
are not enveloped in a cellulose membrane.

SECTION RENANTHERAE.

Blakely has erected a section Renantherae containing two series which are not in
the section Renantherae-Normales, i.e. the Occidentaies and the Ochroxylon. Of the
five species of the Occidentales we were able to examine H#. marginata, HE. staerii and
H. sepulcralis, which agree with one another well and are characterized by the
following features:

The outer epidermis has a cuticle-free, thick, strongly lignified outer wall which
on the inside has a thick mucilaginous lamella extending along the side walls almost
to the thin base wall of the cells (Pl. i, fig. 2). This mucilaginous layer is strongly
impregnated with tannin material, swells in water displaying lamellations, and
contracts in alcohol.”

The parenchyma of the outer integument is strongly developed and filled with
tannin deposits. In EH. staerii crystals are scattered over the whole inner epidermis,
while in #. marginata and E. sepulcralis they are confined to the hilar region.

The inner integument is suberized and in general two-layered, in H. staerii in some
places three- or four-layered. The suberin lamellae display a fine granulation which

2 Staining to test the nature of this mucilage is masked by tannin impregnation. This
can be removed by Eau de Javelle, whereupon chliorciodide of zine gives a dark blue colour
and cuprammonia a light blue. This points to cellulose mucilage. On the other hand,
ammoniacal ruthenium sesquichloride stains deep red, indicating the presence of pectin
mucilage. It seems, therefore, that both kinds are present, if staining alone is a sufficient
proof. There is no specific microchemical reagent for pectin.

28 SEED COAT ANATOMY AND TAXONOMY IN EUCALYPTUS. I,

was also observed in some of the Renantherae-Normales species, though to a lesser
degree.

The nucellus remnant for the most part has lost cellular structure, but here and
there it can be distinguished (PI. i, fig. 2).

This type of testa appears as a variant of the type of the Renantherae-Normales.
The mucilaginous wall component of the epidermal cells does not represent a significant
structural variation, since in an alcohol preparation the outline of the total thickening
(cellulose plus mucilage layer) is exactly the same as that of the epidermal sclereids
of the Renantherae-Normales. It is irrelevant to this study—and therefore has not
been investigated—as to whether this mucilage as such has been deposited upon the
cellulose walls or whether it is due to subsequent partial conversion of the walls
into mucilage.

The structure of H. guilfoylei, of the monotypic series Ochroxylon, is quite different
Cea, i, ile, 110). The outer integument of the flat seed is, viewed from the
raphe-free side, formed from the two epidermal layers only. The cuticle-free outer
epidermis is made up of thin-walled cells with walls impregnated with brown material
which often adheres aiso to the wall in the form of irregular lumps which do not
give any tannin reaction with FeCl;. The inner epidermis is a typical crystal epithelium.
The inner integument is resorbed. The structure has therefore nothing in common
with the large bulk of the Renantherae and resembles much more in general
#. microcorys, with which it also has in common the cellulose envelopes of the crystals.

To sum up, we can therefore say that on the basis of testa structure the three
series, Ochroxylon (#H. guilfoylei), Steatoxylon (H. microcorys) and Myrtiformes (£.
deglupta, H. brachyandra and EH. raveretiana), do not conform to the combined group
Renantherae and Renantherae-Normales. This evidence conforms with Maiden’s view
of the relationship of HE. microcorys. He says (1, 262): “This species appears to stand
by itself amongst the Renantherae to a greater extent than any other members of
that group.” To place correctly HE. microcorys and the other species mentioned many
more factors must be considered. This will be done at the conclusion of the anatomical
study of the seed of the remainder of the genus.

There is also still another difference which separates these three series from the
remainder of the Renantherae. This is the close placing of hilum and chalaza and
consequently a different pattern of attachment of the seed and its vascularization.

The seed of the Renantherae is of somewhat irregular form, but it always displays
about four or five raised ribs which spread out from a clearly marked basal hilum,
that is at or near the base of the seed. As this type of seed originates from an
anatropous ovule, the hilum lies in the vicinity of the micropyle through which the
hypocotyl emerges. Therefore the germinating seed shows the hypocotyl close to the
hilum at the base of the seed which is more or less perpendicularly attached to the
‘placenta, while the chalaza occupies the other end of the seed (Text-fig. 12). The
vascular bundle diverging from the placenta as a single collateral strand into the
outer integument ramifies sooner or later in numerous more or less arcuate ascending,
not anastomosing, amphicribral branches which fan out over the chalaza, where they
terminate (Text-fig. 19, 1). In transverse section they are seen arranged in a flat are
facing the chalaza (PI. i, figs. 3, 4).

In the three series which do not conform to the general Renantherous pattern
the seeds are bifacial with a ventral hilum, that is in the middle or perhaps a iittle
towards the upper end of the one side, and close to the chalazal region, or even
within it (as in #. microcorys).“ Thus the germinating seed shows the hypocotyl at
the lower (micropylar) end and the hilum quite distant from it near the middle of
the seed which is attached like a snield to the placenta (Text-fig. 13). This leads to

18 Discussing the ventral hilum of the Corymbosae, Maiden (VII, 95) mentions that after
removal of the testa the scar beneath (this means probably on the inside) ‘is often larger
and more distinct and definite in shape to that observed on the testa’. This “‘scar beneath”
is of course the chalaza cork and has nothing to do with the scar of the hilum. The
occurrence of hilum and chalaza face to face, or nearly so, is characteristic for seed with
a ventral hilum, as in EH. microcorys.

BY E. GAUBA AND L. D. PRYOR. 29

the conclusion that such a seed originates from another type of ovule, the hemitropous.
But further investigation must be undertaken when suitable material for ontogenetic
study is available to see. whether this structure is not perhaps the result of change
by growth of an ovule also originally anatropous. Goebel (1933) has illustrated by
different examples a change by growth before or after fertilization resulting in an
alteration of ovule shape, for example one originally anatropous becomes campylotropous
(Geraniaceae) or an atropous ovule changes to hemitropous (TVorenia asiatica).

Through whatever phases these ovules may pass during their development the
investigation of flower buds of H. microcorys just before anthesis shows that they are
already of hemitropous shape (PI. i, fig. 7). The simultaneous occurrence of atropous
ovules (Pl. i, fig. 8) will be discussed in connection with the chaff structure.

The phylogenetic and taxonomic importance of the ovule shape and structure has
been emphasized often by many authors (see Warming, 1913), though in higher groups
(families, ete.) different types may occur. In the Myrtaceae, for instance, the bitegmic
anatropous ovule is dominant, but campylotropous ones occur too, and Jambosa
caryophyllus is quoted as having only one integument. Thus it is not surprising that
a genus as large as Hucalyptus has more than one ovule type.

Due to the ventral position of the hilum and in contrast with the elaborately
ramified raphe bundle of the Renantherae, the conducting tissue of these three
series is greatly reduced in extent and ramification, and its xylem built up by short,
wide, irregularly shaped and sometimes branched tracheids with truncate ends, while
in the Renantherae the tracheids are slender and long (Text-fig. 19, 1-6).

It is a well established fact that vessels in vegetative parts have undergone
phylogenetic modifications culminating in the formation of short, wide vessel members
with truncate ends (Bailey, 1944). Tracheids likewise have become shorter, though
in a lesser degree. But too little is known about their structure and evolutionary
trends in raphe bundles. Kuehn (1928) has investigated the course of intraseminal
(= integumentary) bundles of the Angiosperms. Though this vascularization type does
not occur in our species—there are only raphe bundles—her results are of great
interest: the intraseminal vascular system has arisen independently in unrelated
families, primitive and high ranking, and some families (Oleaceae, Leguminosae,
Compositae, etc.) comprise genera with and without such bundles. This leads to the
conclusion that they are of little phylogenetic significance. However, this does not
mean that the vascularization pattern is of no taxonomic value for generic or
infrageneric grouping.

Kuehn has not examined the Myrtaceae. Concerning the structure of intraseminal
bundles she remarks only that they are built up by helically or annularly thickened
tracheids. To our knowledge no other evidence in connection with taxonomic evaluation
of seed vascularization has been published and its significance in the genus Hucalyptus
will show up when all sections have been investigated. It already seems likely,
however, that the two vascularization patterns are closely connected with the ovule
type.

Finally. a few remarks on the nature and structure of the chaff seem appropriate.

We have examined these sterile bodies in H. muelleriana, E. laevopinea, H. alpina,
#. obliqua, E. planchoniana, EH. oreades and E. fraxinoides. They are in principle in
perfect confermity in all respects. The outer epidermis is similar to that of seed
but encloses a tissue which may be best termed sclerotic parenchyma. On thin sections
it has some resemblance to angular collenchyma in so far as the wall thickenings
are apparently only in the cell corners while the side walls appear thin due to the
very wide pits (Text-fig. 14). All cells are strongly lignified and filled with phlobaphenes
staining inky black with ferric chloride. A disorganized vascular bundle can be
located embedded in this tissue.

The chaff of HE. microcorys is of a strikingly different structure. While the seed
epidermis is formed by thin-walled cells (Pl. i, fig. 5), the chaff epidermis is exactly
of the Leguminosae type, that is to say, built up by palisade-like sclereids many times
deeper than wide (Text-figs. 15, 16). The bodies forming this chaff are of very

I,

SEED COAT ANATOMY AND TAXONOMY IN EUCALYPTUS.

30

if).

Text-fig.
1: Eucalyptus dives;

3: E. brachyandra ;

qNiCcrocerys ;
(ch, chalaza region; h, hilum. )

Bate

b)
z

130x.
5: H. deglupta;

EH. gwilfoylei, tang.
scalariform thickenings on the outer wa

raphe bundles, ca.

4: H. raveretiana;

1-6:

E. gwilfoylet.

67:

sect, of the outer epidermis o

Gr

f chaff with basal hilum showing

ca. 400x).

(photomicrograph,

ll

BY E. GAUBA AND L. D. PRYOR. 31

different size and shape, but they can be placed in two groups: small, narrow ones
with a basal hilum, and larger, flat ones, often glossy, with a ventral hilum.

To trace their origin an ontogenetical investigation is needed. We could examine
oniy flower buds just befere anthesis and this revealed the presence of two kinds of
ovules in the ovary. Firstly, the fertile, at this stage already hemitropous ovules
which have two distinct two-layered integuments with three cuticulae: an outer
covering the surface, a median between the two integuments and an inner one
delimiting the inner integument from the nucellus (Pl. i, fig. 7).1* These ovules
develop after fertilization into seeds Icsing during maturation the outer and median
cuticle and the inner integument.

Besides these fertile ovules there are many sterile, atropous ovules of prismatic
shape, with basal hilum and distal micropyle. They are built up by a parenchymatous
tissue with a procambial strand in the axis. At its apical end this parenchyma encloses
a cavity leading into the micropyle and lined with a cuticle. The outline of this cavity
varies with the section and is seen as either empty or containing a small, more or
less isolated group of parenchyma cells (Pl. i, fig. 8). If we interpret this cuticle
as the inner one, then the outer parenchyma represents a modified integumentary
tissue and that within the cavity a modified nucellar tissue.

It is probably correct to assume that these sterile ovules mature finally into chaff
with a basal hilum. Thus this chaff is already predetermined befcre anthesis. In
conformity with the ovules from which it originates it shows a poor histological
differentiation, the tissue consisting mainly of the epidermis and the parenchyma, both
strongly sclerified and lignified (Text-fig. 16).

The chaff with a ventral hilum develops probably from fertile but not fertilized
ovules. The hemitropous shape not only conforms with this suggestion, but also the
richer tissue differentiation does too: outer epidermis, crystal layer, inner cuticle and
an obliterated nucellus remnant, but of course no embryo (‘Text-fig. 15).

H. guilfoylei displays two similar kinds of chaff, of which we mention only the
different architecture of their epidermal cells. All walls, except the inner periclinal
ones, have conspicuous thickenings which are pitted even on the outer walls (blind
pits) in chaff with a ventral hilum (Text-fig. 17), while in chaff with a basal hilum
they are scalariform or scalariform-reticulate (Text-fig. 19, 7). As mentioned earlier,
the seed epidermis of this species is thin walled (Text-fig. 10).

Thus, not only the seed coat but the chaff structure too deserves consideration
and may contribute valuable information for taxonomic problems.

References.

Baitpy, I. W., 1944.—The Development of Vessels in Angiosperms and its Significance in
Morphological Research. Amer. Jour. Bot., 31: 421-428.

BENTHAM, G., 1866.—Flora Australiensis, Vol. 3.

BLAKE, S. T., 1953.—Botanical Contributions of the Northern Australia Regional Survey, I,
Studies on Northern Australian Species of Eucalyptus. Aust. Jowrn. Bot., I, No. 2: 185-352.

BLAKELY, W. F., 1955.—A Key to the Eucalypts (2nd edition).

GOEBEL, K., 1933.—Organographie der Pflanzen, 3 (3rd edition).

HarapA, M., 1956.—Anatomical Characteristics of the Seed Integument observed on 10 species
of Eucalyptus. Reports of the Kyushu University Forests, No. 6: 1-14.

KUHN, G., 1928.—Beitrage zur Kenntnis der intraseminalen Leitbiindel bei den Angiospermen.
Bot. Jahrb., 61: 325-379.

MAIDEN. J. H., 1929.—A Critical Revision cf the tfenus Hucalyptus, 7.

MULLER, F. v., 1879-1884.—EHucalyptographia.

NETOLITZKY, F., 1926.—Anatomie der Angiospermen Samen. In: K. Linsbauer, Handbuch der
Pflanzenanatomie, 10: 14.

Petit, L. A., 1908.—Recherches sur la Structure anatomiaue du Fruit et de la Graine des
Myrtacées (Dissertation, Paris).

Pryor, L. D., 1956.—An Fil Hybrid between Hucalyptus pulverulenta and EH. caesia. Proc.
Linn. Soc. N.S.W., 81, Part 2: 97-100.

WARMING, H., 1913.—Observations sur la Valeur systématique de lOvule. Mindeskrift for
J. Steenstrup (Copenhagen).

147The outer cuticle is a single membrane, the two others are of double nature, as can be
seen where the two integumelts ‘separate from each other (Pl. i, fig. 7) or the nucellus
Separates from the testa (Pl. i, fig. 8).

32 SEED COAT ANATOMY AND TAXONOMY IN EUCALYPTUS. I.

BXPLANATION OF PLATH I.
Cork tissue and cuticles brown.

1: Hucalyptus laevopinea, part of transy. sect. of seed. 2: H. marginata, part of tranv.
sect. of seed. The mucilage layers of the epidermis (mu) swollen by water absorption.
Nucellus (7) partly crushed. 3: EH. tasmanica, part of transv. sect. through the raphe-chalaza
region at the level of the cuticula gap (the inner cuticle, ic, ending blind in the chalaza cork).
4: H. stellulata, part of tranv. sect. through the raphe-chalaza region, somewhat below the
cuticula gap (the inner cuticle, ic, is running through). The tracheids (t) of the amphicribral
raphe bundle fan out over the suberized chalaza (che). 5: H. microcorys, longit.-radial sect.
through the raphe-chalaza region and the cuticula gap showing the course of the raphe
bundle (rb) and the gap in the crystal layer (cr) and inner cuticle (ic). 6: H. microcorys,
part of transy. sect. of seed. 7: EH. microcorys, diagram and micropyle of the hemitropous
ovule showing the three cuticulae (oc, mec, ic). 8: H. microcorys, leng. sect. of a sterile
atropous ovule with the single outer and the double inner cuticle. 9: H. brachyandra, part
of tranmsy. sect. of seed. 10: H. guilfoylei, part of transy. sect. of seed. All figures (except
the diagram) ca. 180x. Figs. 7 and 8 microtome sections.

ch, chalaza ; che, chalaza cork; cr, crystal layer; e, embryo; ec, embryo cuticle;
en, endosperm; ic, inner cuticle; ie, inner epidermis; ‘w#, inner integument; mc, median
euticle; mu, mucilage; n, nucellus; nr, nucellus remnant; oc, outer cuticle; oe, outer epidermis ;
oi, outer integument; pl, placenta; ps, procambial strand; rb, raphe bundle; ¢t, tracheids.

33

A NEW SPECIES OF AEDES (FINLAYA) FROM NORTHERN AUSTRALIA
(DIPTERA, CULICIDAE).

By EvizagetH N. Marxs, National Mosquito Control Committee, Department of
Entomology, University of Queensland, and Ernest P. Hopexin, Department of
Zoology, University of Western Australia.

(One Text-figure.)

[Read 26th March, 1958.]

Synopsis.
Both sexes, larva and pupa of Aédes (Finlaya) Obritteni, n. sp., are described from
northern Western Australia. It occurs also in Northern Territory. The affinities of A. britteni
are discussed and it is placed in a new subgroup of Group D of the subgenus Finlaya.

For many years only two species of the subgenus Finlaya, Aédes notoscriptus
(Skuse) and Aédes purpureus (Theobald), were known from northern Western
Australia. Hodgkin and Britten (1955) recorded a third, undescribed, species which
is here described as Aédes britteni, n. sp. It has since been found also in Northern
Territory by Mr. A. K. O’Gower, to whom we are indebted for the opportunity to
study his specimens.

AEDES (FINLAYA) BRITTENT, N. Sp.

Distinctive Characters.

Adult: This is the only Australian Finlaya with hind tarsal segment V white with
a narrow basal dark band; other distinguishing characters are dark-scaled proboscis
and wings; absence of scutal pattern; scutellar scales ali broad, white; small patches
of broad white scales on pleuron, including paratergite; hind tarsal segment IV
all dark. ’

Larva: Antenna long, swollen near base; head setae 5 and 6 single or bifurcate,
14 short, stout, single; lateral comb a single row of fringed scales; distal margin of
saddle without spines.

Description of Adult. a

Female—wWing length 3:8 mm. Head: Integument orange-brown, clothed mesially
with rather sparse narrow-curved creamy scales, in front of which is a continuous
band of narrow-curved black scales, terminating laterally in a small patch of fiat black
scales, below which are flat creamy scales. There is a wide ocular border of flat
silvery-white scales, discontinuous at vertex, but a few narrow silvery scales extend
between the eyes. Upright forked scales long, black, numerous. A pair of strong
black vertical bristles with two pairs of smaller bristles above them and a row of
five strong mesially-directed ocular bristles, with finer bristles laterally. Torus dark
brown with some small dark setae mesially; flagellar segments of antenna black with
short silvery clothing hairs and sparse black verticillate hairs; first flagellar segment
paler on basal half and with flat black scales mesially. Clypeus dark brown. Palps
and proboscis black scaled; palp 0:2 length of proboscis, which is 1:2 length of fore
femur. lLabella dark brown.

Thorax: Integument bright orange-brown darkened beneath the silver scale patches.
Scutum clothed with rather sparse fine narrow-curved dark brown scales. Bristles
strong, black, about 13-16 acrostichal, 12-14 dorsocentral, 9-10 prescutellar, a group
above wing roots, scattered bristles along lateral margin of scutum, and one on fossa.
Scutellum darker brown, all lobes densely clothed with broad flat silvery-white scales;
4-6 long black bristles to each lobe in addition to shorter bristles.

-PROCEEDINGS OF THE LINNEAN SocineTY OF NEW SOUTH WALES, 1958, Vol. Ixxxiii, Part 1.

34 A NEW SPECIES OF AEDES (FINLAYA) FROM NORTHERN AUSTRALIA,

There is a patch of broad silvery-white scales on apn, propleuron and paratergite,
a small patch on posterior margin of ppn below the bristles, on upper stp, on lower
posterior margin of stp, and on upper msp. Bristles black; apn with 3-4 very strong
bristles above, numerous others below; 5 long and 1-2 shorter propleural; 4-5 strong
ppn; 3-4 postspiracular, one upper stp and 2-3 long bristles along posterior margin
of stp in addition to finer bristles; 8-9 prealar; 8-10 lighter brown upper and no
lower msp bristles.

Legs: Black scaled with purplish-blue reflections and banded tarsi. Coxae and
trochanters yellowish-brown; fore coxa with mixed dark and pale scales, mid and
hind with patches of silvery scales, hind also with some dark scales below; trochanters
dark scaled, mid and hind with pale scales posteriorly. Fore femur with pale scales
dorsally on basal 1/4, and extending as a tapering streak posteriorly for 2/3-3/4 length
and with a small silvery-white kneespot; tibia dark except for a few white scales
dorsally at base; tarsal segments I and II with basal white bands covering 1/5-1/4 I
and 1/3 II, I also with a small white apical dorsal patch, and one or two white scales
at apex of II and a few scales or small patch at base of III. Mid-femur with creamy
scales showing anteriorly as a small basal patch, extending on basal 1/4-1/3 dorsally
and as a tapering stripe on basal 1/2-3/5 posteriorly, and with white kneespot; tibia
as on fore leg; tarsal segments I and II with basal white bands covering 1/5-1/4 I,
1/3 II, basal patch 1/5-1/3 III, and a few pale scales also at apex of I, II and V.
Hind femur with white kneespot and with small dorsal basal patch of creamy scales
which has a few pale scales below it anteriorly, and extends as a tapering streak
on basal 1/3 posteriorly. Tibia entirely dark or with a few white scales at hbase;
tarsal segments I-III with basal white bands covering 1/5 I, 1/4-1/3 II and 1/3 III,
I and II also with small patches of white scales apically, IV all dark, V with basal 1/5
dark, and a ventral line of darkish scales, remainder white. Claws equal, those of fore
and mid legs toothed, hind simple.

Wings: Black scaled, outstanding scales all long and narrow. Cell R2 1:5-1:6 times
length of its stem; cell M1 0:8 times length of its stem, its base slightly proximal to
that of cell R2; r-m twice its own length distal to base of M3+4. Halteres pale with
dark scales on knob and a few dark and pale scales on upper side of stem.

Abdomen: Mainly black scaled with purplish-blue reflections; ‘numerous black
bristles along apical and lateral margins of tergites II-VIII and scattered over tergite I
and the sternites. Tergite I dark scaled mesially, with silvery-white scaled lateral
margin; tergite II with a pair of submedian basal patches of pale-refilecting scales,
contiguous with large lateral basal patches of silvery-white scales; III-VI with a
broad basal patch of pale scales, covering 1/2—2/3 length of tergite, indented in mid-line
on III-V, and with small patches of silvery-white scales slightly removed from basal
and lateral margins; VII dark with large lateral white patches, VIII short, dark scaled.
Sternites dark scaled, VIII large, exserted, integument dark basally, yellowish-brown
apically, clothed with fine hairs and bristles, bare of scales. Cerci short, dark.

Described from the holotype and one paratype female. The holotype has the
scutum and scutellum slightly rubbed.

Four females from Roper River Mission show the fcllowing differences from the
foregoing description: Wing length 3:6-4:-7 mm. Some small dark scales mesially on
torus (possibly obscured by shrinkage in holotype and paratype); 18-25 acrostichal,
11-13 prescutellar bristles; 7-8 long bristles on lateral lobe of scutellum; 2-4 ppn,
1-5 postspiracular, 1-2 upper sty and 7-16 prealar bristles. Fore leg: Basal band
1/4-1/3 tarsal segment II; there may be no pale scales at apex of I and II or base of III.
Mid leg: Pale scaling at base of femur may be reduced dorsally; basal band 1/4-1/3
tarsal segment II; there may be no pale scales at apex of I, II and V. Hind leg:
Posterior stripe may extend to mid length of femur; basal white bands 1/5-1/4 tarsal
segment II, 1/4 III; basal 1/4 V dark; no pale scales at apex of I and II. Wings:
Cell R2 1:4-2:0 times length of its stem; cell M1 0-7-1:0 times length of its stem, their
bases may be level; r-m 2-8 times its own length distal to base of M3+4. Abdomen:
Median patches of abdominal tergites may be indented on III—VI or lack indentatio
on IV-VI. :

BY ELIZABETH N. MARKS AND E. P. HODGKIN. 35

Male.—Resembles the female except as follows: Wing length 2:9-3:-2 mm. Head:
White scales between the eyes may be broad; lateral patch of flat black scales may
be absent. Torus large, dark, with fine short setae on mesial aspect; flagellar segments
of antenna brown with dense dark verticillate hairs lying mainly in an almost vertical
plane, first segment with some flat dark scales distally; two apical segments elongate,
dark, with silvery clothing hairs. Palps about equal in length to proboscis, dark scaled
with a small white basal band on segment III, sometimes also on II, and wide white
basal bands covering 1/5-1/4 IV and 1/2 V; segments IV and V down turned, without
dense hairs but with short dark bristles, about 6 at apex of III, 6 or 7 at apex of IV
and 4-6 at tip of V, as well as 12-18 along lower side of IV and numerous finer bristles
along V.

Thorax: 10-13 acrostichal, 9-16 dorsocentral, 4-9 prescutellar bristles. On mid
lobe of scutellum the white scales may be divided inte two patches by a narrow
median unscaled line; apn with 2-3 strong bristles above; 5-7 upper msp bristles.

Legs: Mid coxa may have a few dark scales below; hind coxa may have white
scales only; fore trochanter may be pale scaled posteriorly. Fore leg: The pale
posterior streak on femur may extend almost to apex; tibia may be dark basally;
basal bands covering 1/5-1/3 tarsal segment I, 1/4-2/5 II. Mid leg: Femur with a
streak on basal 1/3—2/3 posteriorly; tibia may be dark basally; basal bands covering
1/5-1/3 tarsal segment I, 1/4-1/2 II, III sometimes and V usually all dark. Hind
leg: Femur may have a complete ring of pale scales at base, or be dark to base
anteriorly with streak on basal 1/4-1/3 posteriorly; basal white bands covering 1/5-1/4
tarsal segment J, 1/5-1/3 III. On fore and mid legs, tarsal segment IV is 1/2 length
of V; claws (Fig. 1, a, 0) large, unequal, pilose basally, the anterior long with a
strong blunt tooth near mid length and usually with a slender pointed tooth arising
laterally at base (the strong tooth reduced on one mid claw in one specimen), posterior
claw shorter, with pointed subbasal tooth. Hind claws equal, simple.

Wings: Cell R2 1-2-1:5 times length of its stem, cell M1 0-6—0-8 times length of
its stem. Halteres with dark scales on knob and running down dorsal side of stem,
sometimes with a few pale scales on knob and stem.

Abdomen: Tergite II may have a pair of small submedian basal patches of pale-
reflecting scales, III-VI with median basal paie patches 1/4-3/4 length of III and IV,
1/3-3/4 length of V and VI, indented in mid-line on III and frequently also on IV-VI;
VII dark mesially or with median basal pale patch 1/3 its length; true tergite VIII
dark scaled or with a few silvery scales laterally. Sternites dark scaled, IIJ-VII may
have silvery lateral patches at mid length, VII may also have a median basal patch;
VIII with large median basal patch of silvery scales.

6 Terminalia (Fig. 1, c): Coxite densely clothed with setae and with large patches
of silvery scales laterally at base and dark scales towards apex; cylindrical, about
four times as long as broad at base, with a membranous area along its inner aspect.
The slightly developed basal lobe bears a dense patch of about 60 medium length setae,
those in the outer row appearing somewhat stouter; extending distally from this
patch, along the tergal side of the membranous area are about six rows of short fine
setae, which become longer near apex of coxite. Extending along the sternal side of
the membranous area on distal half of coxite are three rows of medium length fine
straight setae; sternal to these again, running the length of the coxite and likewise
directed mesially are 3-4 rows of longer stouter finely striated setae, the more distal
of which are the longest. There are numerous scattered long setae on the sternal,
lateral and outer tergal aspects of coxite, densest towards apex; there are also a few
shorter setae sternally at base. Style 2/5 length of coxite, strongly curved, slightly
tapering, non-pilose, with 1—2 short preapical setae; terminal appendage 1/7-1/6 length
of style, stout, apparently grooved with rounded tip. Harpago about 1/3 length of
coxite, stout basally, more slender on apical half, pilose except near apex, with a row
of 7-9 slender setae mesially on basal half; just beyond these tergally arise two longer
setae, reaching to apex of harpago; appendage about 4/5 length of harpago, broadening
on basal third, then tapering to a slender pointed tip, fairly evenly sclerotized except
for membranous broadening near base. Paraproct with single tooth. Phallosome simple,

36 A NEW SPECIES OF AEDES (TINLAYA) FROM NORTHERN AUSTRALIA,

elongate, tapering distally. Lobes of IXth tergite with 6-10 stout setae; IXth sternite
with 15-16 short setae. :

Larva (Fig. 1, d-k)—Nomenclature of setae as in Belkin (1950). Length about
8-10 mm. Cuticle dark. Setae in general rather short.

h ee

7
; m

Fig. 1—Aédes britteni, n. sp. a-c. Male: a. Anterior and b. posterior claw of foreleg.
ec. Terminalia (tergal view; blade of left harpago omitted).

d-k. Larva: d. Head. e. Prothoracic setae 1-3. f. Base of mesopleural and g. base of
metapleural setae (ventral view). h. Terminal segments. i. Lateral comb tooth. j and k.
Pecten teeth, 7 from apex and k from middle of pecten.

l-m. Pupa: Jl. Cephalothorax (head capsule separated). m. Metathorax and abdomen
(dorsal setae on left, ventral on right).

BY ELIZABETH N. MARKS AND E. P. HODGKIN. 37

Head: 0-8-0:9 times as long as broad. Antennae 0-6—-0-8 length of head, slightly
curved, broadening just above base (greatest breadth is 0:14-0:18 length of antenna),
then tapering to terminal. third which is parallel-sided; with sparse fine spicules; seta 1
arising at about mid-length, stout, plumose, single or bifurcate near tip, 0:3-0:5 length
of antenna; terminal and subterminal setae arising close together, 2 moderately long,
3 short, 4 shorter than 2, 5 broad with pointed tip, 6 short. Head seta 1 single,
slender, curved, simple; base of 7 slightly behind base of antenna, 6 level with base
of antenna and about 0-6 distance from 7 to midline, 5 slightly posterior to 7 and
lateral to 6, 4 mesial to and about midway between 5 and 6; 4 short, fine, 7-15-branched;
5 and 6 about 0-5-0-6 length of antenna, stout, plumose, single or rarely bifurcate
beyond mid-length; 7 plumose, 6—10-branched; 8 long, single; 9 3—-6-branched; 10 2-6-
branched, rarely single; 11 5-10-branched, plumose; 12 single to trifid; 13 fine, 5-10-
branched; 14 short, stout, single; 15 short, single. Setae of mouthbrushes apparently
all simple in 15 specimens; some of the more mesial setae finely pectinate in seven
specimens, including skin of holotype. (The occurrence of this type of dimorphism in
some species was first brought to our attention by Mr. P. F. Mattingly.) Mentum
triangular with median pointed teoth and 9-12 pointed lateral teeth, of fairly even
length, but the more lateral ones stouter.

Thorax: Prothoracic setae 1, 2 and 3 without sclerotized bases, 1 the longest, single
or rarely bifid, finely plumose; 2 single, simple, about 0:6 length of 1; 3 fine, 5-10-
branched, about ‘0:2 length of 1; 14 short, single or bifid. The bases of the meso- and
metathoracic pleural setae bear very large curved pointed spines.

Abdomen: Seta 6 on segments I-VI in length about 0:5 width of segment; on
I and II 2—5-branched, stout, plumose, arising from a large sclerotized base; on III-VI
bifid (rarely trifid on III), slender, finely frayed. Seta 7 on I slightly shorter and finer
than 6, 2-3-branched, piumose, arising from same base as 6; on II about 0-5 length of
6, 2-8-branched, plumose, arising from a separate sclerotized base. Highth segment:
Lateral comb a single curved row of 13-20 broad, apically rounded, coarsely fringed
scales (in one specimen one scale was posterior to the row; in one specimen, one comb
tooth was a small spine); seta 1 2-4-branched, finely frayed; setae 2 and 4 single,
simple; seta 3 5-9-branched, plumose, arising from a small sclerotized base; seta 5
2—4-branched, frayed. Siphon slightly tapering, with small acus; index 2-2-2-9; pecten
extending over basal 0:-4-0-5 of siphon, of 15-23 close-set dark spines with 2-4 fairly
even-sized denticles near base; the spines gradually increase in size from the base of
the siphon, the distal ones very long, with inconspicuous denticles; seta 1 arising
at 0-6-0:7 length of siphon, 2-3-branched, finely frayed, about 0-3 length of siphon;
seta 8 single to trifid. Anal segment: Saddle covering about dorsal 0-9 of segment,
without apical spines; seta 1 single, simple, about 0:6 length of saddle; seta 2 11-20-
branched, about equal in length to saddle; seta 3 single, about 2:5 times length of 2;
seta 4 of 11 14-22-branched tufts arising from a grid and 1 smaller precratal tuft;
anal papillae pointed, subequal, the lower 0:7 or more length of upper which are almost
equal in length to saddle.

Described from four skins, correlated with the holotype, allotype and two paratype
males, and 18 morphotype larvae.

Pupa (Fig. 1, 1, m)—The nomenclature of the setae follows Belkin (1952, 1958).
The setae are simple, unless stated; their lengths are indicated in the figure.

Cephalothorax: Trumpet evenly pigmented, 2-7-3-2 times as long as greatest width,
with oblique opening; ratio of meatus to whole 1 : 1-5-1:7, apical notch shallow. Setae 1,
2 and 3 single; seta 4 single to trifid; seta 5 single; setae 6 and 7 2~3-branched; seta
8 2-5-branched; seta 9 single; seta 10 single or bifid; seta 11 single; seta 12 single
to trifid.

Abdomen: Segment I. Seta 1 strongly developed, with about 10 primary branches
each subdividing into 2—4 simple or sparsely plumose branches. Seta 2 single or bifid;
seta 3 single, may be inconspicuously frayed; seta 4 single or bifid; seta 5 2—3-branched;
seta 6 single to trifid; setae 7 and 10 single. Segment II. Seta 1 single to trifid, may
be inconspicuously frayed; setae 2 and 3 single; seta 4 2-4-branched; seta 5 3-5-
branched; seta 6 single or bifid; seta 7 single; seta 10 single or bifid, Segment III.

Cc

38 A NEW SPECIES OF AEDES (FINLAYA) FROM NORTHERN AUSTRALIA,

Seta 1 single to trifid; setae 2 and 3 single; seta 4 single or bifid; seta 5 3—4-branched;
seta 6 single to tetrafid; seta 7 single; seta 8 2-3-branched; seta 10 single to trifid;
setae 11 and 12 single. Segment IV. Seta 1 single to trifid; setae 2, 4 and 5 single;
setae 3 and 6 single to tetrafid; seta 7 single; seta 8 single or bifid; seta 10
2—3-branched; seta 11 single; seta 12 single or bifid. Segment V. Setae 1 and 3 single
or bifid; seta 2 single; seta 4 2-5-branched; seta 5 single; seta 6 single to pentafid;
seta 7 single; seta 8 single or bifid; seta 10 2-3-branched; setae 11 and 12 single.
Segment VI. Setae 1, 2, 3 and 5 single; setae 4 and 6 single or bifid; seta 7 single;
seta 8 2—4-branched; setae 10, 11 and 12 single. Segment VII. Setae 1, 3 and 5 single .
or bifid; setae 2 and 4 single; seta 6 4-5-branched; seta 7 stout, single; seta 8 single
to trifid; setae 10, 11 and 12 single. Segment VIII. Seta 5 single; seta 7 stout, single,
slightly frayed. Paddles broad with bluntly pointed apex; index 1:2-1:5; midrib
moderately and buttress slightiy developed; fine denticles along margin; seta 1 single.

Described from three pupal skins, one correlated with the allotype and two with
paratype males.

Types: Holotype female, Kalumburu (Drysdale River Mission), 14° 25’ §.,
126° 37’ H., Western Australia, 14.111. 1954 (0830-7), EH. P. Hodgkin. Allotype male
(0830-9), one paratype female (0830-8), four paratype males (0830-5, -6, —10, —11),
and 18 morphotype larvae, same data as holotype. The holotype has a correlated larval
skin, and the allotype and two paratypes (0830-5, -10) have correlated larval and
pupal skins.

Holotype, allotype, one paratype male and six morphotype larvae in University of
Queensland collection; paratype male’ (with correlated skins) and female and six
morphotype larvae in University of Western Australia collection; one paratype male
and two morphotype larvae in C.S.I.R.O. Division of Entomology collection, Canberra;
one paratype male (with correlated skins) and two morphotype larvae in British
Museum (Natural History); two morphotype larvae in School of Public Health and
Tropical Medicine, Sydney.

This species is named after Mr. H. J. Britten, of the Department of Public Health,
Western Australia, whose collections have added considerably to knowledge of mosquitoes
of that State. ;

Biology: The type series was bred from larvae collected from a rot hole in the
fork of a baobab tree (Adansonia gregortui). Hodgkin and Britten (1955) noted that
larvae of A. purpureus which were found in the same treehole were preying on the
larvae of A. britteni. The mean annual rainfall at Kalumburu is 37 inches, almost
all of it falling from November to April. Mr. O’Gower found adult females occurred
not infrequently in collections at the Roper River Mission, where the mean annual
rainfall of 28 inches has a similar seasonal distribution.

Distribution: WESTERN AUSTRALIA: Kalumburu (the type series). NORTHERN TERRI-
TORY: Roper River Mission, 14° 8’ S., 134° 8’ H. (17, 21, 22, 23.iv.1957, A. K. O’Gower).

Discussion: Though it lacks the characteristic pattern of lines on the scutum,
A. britteni belongs to Group D (aureostriatus-group) of the subgenus Finlaya, in which
Knight and Marks (1952) recognized eight subgroups.

It shows relationships to A. quasirubithorua (Theobald) and A. keefei King and
Hoogstraal (subgroup IV, quasirubithorax) and also to A. candidoscutellwum Marks
(subgroup V, candidoscutellum), species which occur in north-eastern Australia and
New Guinea. Males of both subgroups IV and V differ from A. britteni in having long
hairs on the distal segments of the palp instead of short bristles.

A. britteni resembles A. candidoscutellum (which sometimes lacks a scutal pattern)
in having broad white scales on the scutellum and in small patches on the pleuron,
but in A. candidoscutellum the paratergite is bare (usually ppn also). The male
terminalia of A. britteni bear a fairly close resemblance to those of species in sub-
group IV, but differ in the short stout appendage of the style and the long setae on
the harpago; the resemblance is less close to A. candidoscuiellum which, however, has
long setae on the harpago. The larva of A. britteni resembles those of both subgroups
in the shape of the head, the long antennae, the form and position of head setae 5 and 6.
The shape of the antennae and the form of seta 14 in A. britteni are distinctive. The

BY ELIZABETH N. MARKS AND E. P. HODGKIN. 39

lateral comb is similar in A. britteni, A. quasirubithoraxy and A. keefei, but the two
latter species have distinct fine spines on the distal margin of the saddle.

The foregoing differences indicate that A. britteni does not belong to either sub-
group IV or V; nor does it fall into any of the other six subgroups of Group D.
Another species from eastern Australia, A. wasselli Marks, which belongs to Group D
and is known from females only, could not be placed in a subgroup by Knight and
Marks (1952). A. wasselli has a scutal pattern and subspiracular scale patch; other-
wise the thoracic scaling quite closely resembles A. britteni and the two species appear
likely to be nearly related. The following definition (in the terms of Knight and
Marks, 1952) of a new subgroup to include A. britteni has therefore been widened to
allow the tentative inclusion of A. wasselli also.

Group D (aureostriatus-group: Hulecoeteomyia). Subgroup IX, britteni. Defini-
tion: Australasian. Male palpi with segments IV and V downturned, III-V with a
few strong apical hairs; short hairs only along IV and V (male of wasselli unknown).
Basistyle without a specialized scale tuft. Hind tarsi with basal pale bands on I-III,
I and II with or without apical pale patches or bands; V all white or with basal
dark band. No postspiracular scales. Paratergite scaled. Subspiracular scales present
or absent. Supplementary characters: Claspette filament blade-like. Female tori with
fine hairs medially. Scutal linear pattern distinct or wanting. No prealar scale patch.
Larval head hair 5 posterior to 6, 7 and 4 on a line between 5 and 6. Comb consisting
of a curving row of scales. Larval habitat: treeholes. (Larva of wasselli unknown.)

References.
BELKIN, J. N., 1950.—A revised nomenclature for the chaetotaxy of the mosquito larva
(Diptera: Culicidae). Amer. Midl. Nat., 44: 678-698.
, 1952.—The homology of the chaetotaxy cof immature mosquitoes and a revised
nomenclature for the chaetotaxy of the pupa (Diptera, Culicidae). Proc. ent. Soc. Wash,,
54: 115-130.
, 1953.—Corrected interpretations of some elements of the abdominal chaetotaxy of
the mosquito larva with pupa (Diptera, Culicidae). Proc. ent. Soe. Wash., 55: 318-324.
HopGkKIn, EH. P., and Britten, H. J., 1955.—A survey of the mosquito fauna of tropical
Western Australia. Rep. Comm. publ. Hlth. W. Awst., 1953: 97-107.
KnicHT, K. L., and Marks, EH. N., 1952.—An annotated checklist of the mosquitoes of the
subgenus Finlaya, genus Aédes. Proc. U.S. nat. Mis., 101: 513-574.

CC

40

A SUMMARY OF THE ATOPOMELINAE (ACARINA, LISTROPHORIDABE).
By Rogsrert Domrow, Queensland Institute of Medical Research, Brisbane.
(Nine Text-figures.)

[Read 30th April, 1958.]

Synopsis.

The known species of atopomeline fur-mites are listed, and keys are given to the genera
and to the species of the large and essentially African genus Listrophoroides.

Two new Australian species are described: Awstrochirus perkinsi, n. sp., from the koala
(Phascolarctos) and Atellana papilio, n. g., n. sp., from the brush-tailed possum (Trichosurus).

Cumpylochirus is shown to contain only the geno.ype, C. chelopus, from Pseudocheirus in
Tasmenia.

Neolabidocarpus, from Macropus in New Guinea, of which only the holotype nymph
appeais . be ex.ant, has been reexamined and shown to be an atopomeline and not a
labidocarpine genus. Its exact status remains uncertain. F

New combinations: Listrophoroides adherens (Trouessart, 1893) and Chirodiscoides
oryzomys (Radford, 1954).

New synonymy: Marquesania Wornersley, 1943 = Listrophoroides Hirst, 1923; Listro-
phoroides trdgdrdhi Radtord, 1940 = Marquesania expansa form queenslandica Womersley,
1943 = Listrophoroides expansus Ferris, 1932; Marquesania elongata Lawrence, 1951 = Listro-
phoroides mastomys Radford, 1940; Marquesania imbricata Lawrence, 1954 = Listrophoroides
lemniscomys Radford, 1940; Cricetomysia andréi Lawrence, 1956 = Campylochirus cheloprs
Trouessart, 1893.

A study of the atopomeline fur-mites which have accumulated recently at this
Institute has yielded two new species and given supplementary data on certain other
Australasian species. This led to an interest in the essentially African genus
Listrophoroides, and the subsequent receipt of an extensive collection of this genus
from Dr. C. D. Radford induced me te add a synopsis of the known genera and species
of the subfamily. Twelve genera and 35 species have been included in this group,
of which 10 and 29 respectively are here recognized. | Species not commented on in
the text are readily recognizable from the available descriptions.

This paper is not meant to be a full study of the subfamily, but simply to
elucidate the known genera and species, many of which have long been unrecognizable.
It is certain that new genera and species remain to be found, and the following
generic key is not designed to indicate relationships. For example, I suspect that
Tenrecobia will prove quite close to Listrophoroides.

Key to genera of Atopomelinae.

1. Clasping apparatus enlarged posteriorly, forming a pair of clam-like flaps between coxae II
and III. Female with anterior half of body covered dorsally with two shields and
posterior half covered by small, pointed papillae. Males with legs IV grossly enlarged,
extending far beyond end of abdomen, and as long as entire body; basal movable
SERGE SrA, WR) SUOMI SITHAGES oot anoscoosmooanDestnDUodesOu OS onesas Atopomelus.

Clasping apparatus not so enlarged posteriorly. Body of female, if covered with papillae,
with single dorsal shield (Austrochirus). Leg IV of male moderately enlarged, and,
even if reaching beyond end of abdomen, never more than half as long as body;

. Wal uowKe, Shoo Om lose “maenwelolkey GesMAeMe goconorevonsbenuouopeaudsunobosoUSmSaO rae

2. Posteapitular shield with narrow transverse frental lobe arched over the capitulum,

producing a hunched facies; clasping apparatus immediately followed by a transverse

band sOftStOURCEetrOLSE SDINESE leeNere eet emer ee erence tein ne dace trot nec eh er ak oo ae Atellana.
Postcapitular shield simple, and set behind the evenly tapering capitulum; no spines
behind AclaSpinig "App AACS eye lepers asic wusioawe tepleemedeye xowtausliomeyroiiel s\ sieve! tobe vos 2) stiees ca ieiccd cure OReRe RoR 3.

3. With transverse oval shield immediately behind longitudinal postcapitular shield ........
st ae oa) oa Gee tt ne S1G CO LL OICHOROED DVOTOIOTO OCOD Clo CIOS 8 crow cia bo orcnoteDra ete Tenrecobia.

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutTim WALES, 1958, Vol. Ixxxiii, Part 1.

BY ROBERT DOMROW. 41

4, Postdorsal shield not separated from middorsal shield ....................----se0.> 5.
Postdorsal shield (if present) widely separated from mid- or anterodorsal shield ..... CG

5. Abdomen of female with long, thin process arching forward over dorsum; leg IV of male
much’ enlarged and modified, including tarsus .........2..5...---.4-<«.. Campylochirus.
Abdomen of female without long terminal process; leg IV of male only slightly enlarged
and with normal tarsus ....... Peery eae ete Sate caverta Rent all SAE ROR CMT Col sree oteg SgeLsau sph oh a ssiier Susie) whanerse 6.

6. Postdorsal shield of female covering entire hysterosoma (except extreme apex). Male
with tarsus IV straight, bearing caruncle apically, and without anal suckers ........
EE at ed scare R Seo eST sek seat visto ey a ovleabe: Ausmevarobaus iat ech oMamohegey elisterainavecate Alters Listrophoroides.

Postdorsal shield of female truncate, occupying only anterior half of hysterosoma. Male
with tarsus IV hooked, bearing caruncle subdapically, and with large anal suckers
PR MoM a saci taster syiayr sitar su ahsbises fattayediei riot aige erg cs snicriamteistysu etnies vsecet er eh aneuey eye eavnaticl a engeauen's! wate . Chirodiscoides.
With two broad anterior dorsal shields, the hinder one deeply concave anteriorly to accept
convex posterior margin of the other; third dorsal shield separated from middorsal
SIAN! lonir loueOeVel loguovel Ore Biavonsillengeol pil osonuooacoccogeon0ddudooaudHdD Cytostethum.
With only single dorsal shield anteriorly (which may have lateral accessory lobes) ;
male sometimes with a postdorsal shield, which is widely separated from anterodorsal

=

STC ret cBopeticnes cates tex cliccay steuciomepematee el osislian sicohistis heya lietiet «! aribs etiyeicettenia aliguaties myer entaera: es: c6l s}iale/ aucyeite yee ve wih venire 8.

8. With compact, well developed striate clasping apparatus between coxae I and II. Male
HEM, [Une Ore loslavioel Coxe INS aooococoodcocvguedsoucesecuespobue Austrochirus.
Described as having coxae I and II widely separated medially, and lacking a striate
clasping apparatus; male genitalia between coxae III .................... Centetesia.

AUSTROCHIRUS Womersley.
AUSTROCHIRUS QUEENSLANDICUS Womersley, 1943 (genotype).

The type series of this species is recorded and labelled as from the phalangerid
Trichosurus vulpecula (Kerr), and I had considered the two unnamed species from
this host and from bandicoots listed in the Annual Report on the Health and
Medical Services of the State of Queensland for the year 1937-38 to be the same.
Subsequent collections, however, have shown that A. queenslandicus occurs only on
the bandicoots Thylacis obesulus (Shaw and Nodder) and Perameles nasuta Geoffroy,
and have yielded a new genus and species described below from Tvrichosurus. It
seems certain that mislabelling has occurred, and that the type host of A. queenslandicus
should be T. obesulus, not the phalangerid.

AUSTROCHIRUS ENOPLUS Domrow, 1956.
The holotype female and allotype male of this species are now in the Queensland
Museum, Brisbane.

AUSTROCHIRUS PERKINSI, 0. Sp.

Types: Holotype female and allotype male in Queensland Museum, Brisbane;
paratypes of both sexes in U.S.N.M. and B.M. (N.H.). All specimens from the koala,
Phascolarctos cinereus (Goldfuss), Lone Pine Sanctuary, Brisbane, 26.ix.1955, F. A.
Perkins coll. .

Female—Dorsum with single, small anterior (postcapitular) shield, which is
rather longer than wide, and with a distinct row of heavy punctae on either side.
Four setae, of which the external pair is longer, flank this shield. Remainder of
dorsum covered with closely annulated cuticle, the contours being as shown; without
any medial pattern as in A. enoplus. Dorsal setae as follows: two small setae mid-
dorsally, a transverse row of four similar setae further back, and five pairs of
longer marginal setae, the anterior pair of which is set above the two pairs of setae.
in front of coxae III. Length of body 425-4444. Venter: Capitulum with usual pair
of setae in posterolateral corners. Inner surface of coxae I and II hollowed and
striate; with single pair of setae between coxae I and If. With two pairs of longer
setae above coxae III. Apodemes of coxae IV stronger than those of III, and both
pairs separated by a short median longitudinal sclerotization. Four setae in a line
between coxae III, and four in a square between coxae IV. Anus longitudinal and
subterminal, with one shorter anterior and one longer posterior pair of adanal setae.
. Vertral cuticle similar to dorsal, with two pairs of setae posteriorly. Legs: Tarsi I
and If with caruncles and usual recurved seta dorsally. Apart from the tarsi, the only
movable segments of legs III and IV with setae are the basal and penultimate segments
of lez Ill. Tarsi III and IV with setal pattern similar to other species of genus.

A SUMMARY OF THE ATOPOMELINAE,

Male.—Dersum and anterior half of venter as in female. Length of body 438—467u.
Apodemes of legs III and IV T-shaped, those of IV being much the stronger. Genitalia
behind coxae IV, set in sclerotized ring with two lateral setae. Intromittent organ
well sclerotized, quite short and projecting backwardly. Anus flanked by two sclerotized
areas, each with a short seta anteriorly and minute sucker posteriorly. Apex of
abdomen slightly indented and flanked by five pairs of setae, the central pair being

very short. Legs III as in female. Leg IV swollen, with setae on tarsus as shown.
Caruncle IV much smaller than III.

Text-fig. 1.—Austrochirus perkinsi, 0. sp. Left, venter of female. Right, dorsum of
female. Inset above, dorsal shield of A. queenslandicus Womersley. Inset below, anus of
female A. enoplus Domrow in lateral view.

Text-fig. 2.—Austrochirus perkinsi, n. sp. Venter of male.

Remarks—Mr. P. J. O’Sullivan recorded a severe outbreak of mange due to
Notoedres cati (Hering) in the same sanctuary (Minutes of the Hntomological Society
of Queensland for December, 1949), but A. perkinsi is the first native mite to be
recorded from the koala. The new species is closely related to A. queenslandicus
Womersley,. both possessing a simple dorsal shield and annulated cuticle. They may
be separated on the shape and pattern of punctae on the dorsal shield, the sclerotization
of coxae III and IV, and the genitalia and posterior legs of the male. The other two

BY ROBERT DOMROW. 43

species of the genus, A. sminthopsis Womersley and A. enoplus Domrow, have lateral
accessory lobes to the dorsal shield and modified cuticular patterns, especially in
the former.

CAMPYLOCHIRUS Trouessart.
CAMPYLOCHIRUS CHELOPUS Trouessart, 1893 (genotype).

I have already (1956) redescribed this species, but Dr. R. F. Lawrence, who is at
present engaged on groups other than the Listrophoridae, has asked me to clarify the
position regarding one of his species. Ameng some listrophorid material sent to him
from the Trouessart collection in the Muséum National d’Histoire Naturelle in Paris
were specimens labelled as from an African rodent, Cricetomys gambianus
(Waterhouse), which he deseribed (Lawrence, 1956) as a new monotypic genus and
species, Cricetomysia andréi.

After this paper was published, he received a separate of my redescription
(1956) of the genotype of Campylochirus from the Tasmanian phalangerid, Psewdo-
cheirus convolutor (Oken), and began to suspect that the two species were at least
congeneric. Since then we have exchanged specimens, and these have proved to be
conspecific. There is no doubt that the syntypes of Cricetomysia andréi are also the
syntypes from which Trouessart made his original description of Campylochirus
chelopus in 1893, but how the labels became mixed is conjectural. Cricetomysia andréi
Lawr. is thus an objective synonym of Cumpylochirus chelopus Trt.

Three other specific names have been traditionally associated with this genus
(Radford, 1950), and may be conveniently considered here. Through the courtesy of
Dr. Mare André, I have been able to examine the syntypes of Campylochirus adherens
Trouessart, and these have proved to belong to the African genus Listrophoroides
Hirst. As the species is unrecognizable from the published data, it is redescribed below.

Ewing (1929) synonymized Chirodiscoides Hirst (monotypic for C. caviae Hirst)
with Campylochirus without stating any reasons. It is now clear that his assumptions
were wrong, and that he had not seen authentic material of the genotype of
Campylochirus. The two species here included in Chirodiscoides are distinct, and
possibly closer to Listrophoroides than Campylochirus.

The third name to be considered is Campylochirus latus, ascribed to Trouessart
(without date) by Radford. As a result of correspondence with Dr. Radford and a
search through the literature, this name may now be placed as a nomen nudum.
Campylochirus thus contains only the genotype, C. chelopus Trt.

ATELLANA, Nl. g..

Diagnosis —Atopomelinae with three dorsal shields (somewhat reduced in female) ;
posteapitular shield anteriorly with narrow, transverse frontal lobe. Coxae II and Ili
closely approximated, with numerous heavy retrorse spines at posterior margin of
clasping apparatus. Leg IV of male enlarged, with modified caruncle. Anal suckers
present in male. Nymph with postcapitular shield only; otherwise similar to female.
With two attenuate tracheal tubes in all stages. Genotype: A. papilio, n. sp.

The new. genus may readily be separated from all known atopomeline genera by
its characteristic dorsal shields (particularly the frontal lobe) and the spines behind
the clasping apparatus.

ATELLANA PAPILIO, N. Sp.

Types: Holotype male, allotype female and morphotype nymph in Queensland
Museum, Brisbane; paratype male in B.M. (N.H.). All specimens from fur on the
thighs and rump of the phalangerid Trichosurus vulpecula (Kerr), D’Aguilar Range,
S.E. Queensland, 1.iv.1957.

Maile—Dorsum: A frontal lobe, which appears narrow in dorsal view but more
extensive laterally, precedes the anterodorsal (postcapitular) shield, and masks the
capitulum, producing a characteristic hunched facies. Anterodorsal shield surrounded
by four setae, one pair of which is in the longitudinally striated marginal cuticle,
and the other in the posterolateral corners of the shield. Middorsal shield broad and

44 A SUMMARY OF THE ATOPOMELINAE,

with two pairs of posterolateral setae. Postdorsal shield almost entireiy divided
medially by band of transverse striations; with eight setae arranged 2.4.2. Length of
body 378-3884. Venter: Capitulum almost covered by frontal lobe of anterodorsal
shield, but of similar structure to A. perkinst above, as are the structure of legs I
and II and the clasping apparatus. Immediately behind coxae II a transverse row of
several strong retrorse spines. Two setae above coxae III and a further seta above
these. Internal apodemes of coxae III and IV very strongly sclerotized, in form of a
butterfly. With four setae in arc between coxae III and two on the triangular lobes
between coxae IV and genitalia, which have two minute setae posteriorly. Four setae
between genitalia and anus, which is flanked by two small suckers. Apex of abdomen
irregularly sclerotized, with four pairs of setae, of which one is very much stronger
than the other three. Leg III as in female, with short seta on penultimate segment.
Leg IV swollen, with caruncle weak and slender compared with that of tarsus III.

Text-fig. 3.—Atellana papilio, n. g., n. sp. Left, dorsum of male. Right, venter of male.

Female.—Frontal lobe and antercdorsal shield as in male. Length of body 448uy.
Mid- and postdorsal shields reduced, and without setation. Laterally with longitudinal,
and postdorsally with transverse annulations; setation as shown. Anus terminal, with
two internal sclerotizations and two pairs of adanal setae. Capitulum and legs I
and II as in male. Legs III and IV similar in size. Ventral surface not clearly
visible, but probably with four setae between coxae III and two between coxae IV.

Nynvph.—bDorsally only with frontal lobe and anterodorsal shield flanked by four
setae. Otherwise generally as in female adult. Body length 420u. The nymph
illustrated is somewhat distended, being ready to moult, and containing a full-grown
but weakly sclerotized male.

CytTostErHtm Domrow.
The holotypes of the five species of this genus which I described in these
PROCEEDINGS in 1956 have been transferred from this Institute to the collection of
the Queensland Museum, Brisbane.

NEOLABIDOCARPUS Gunther.
NEOLABIDOCARPUS BULOLOENSIS (Gunther, 1940).

Gunther recorded placing the “type specimen” of Labidocarpus buloloensis in
the School of Public Health and Tropical Medicine, Sydney, and later erected a new
monotypic genus (Neolabidocarpus) for this species, at the same time dividing
the Listrophoridae into four subfamilies, Neolabidocarpus being placed in the
Labidocarpinae.

BY ROBERT DOMROW. 45

Through the courtesy of Mr. D. J. Lee, I have been able to examine the sole
specimen of this species (from Thylogale coxenii Gray, Gunther det.) in the §.P.H.T.M.
It is not labelled as type, but certainly belongs to Gunther’s species. This specimen
is a late nymph and is figured and described below. It is a typical member not of
of the Labidocarpinae, but of the Atopomelinae, legs I and If not being greatly

Text-fig. 4.—Atellana papilio, n. g., n. sp. Lateral view of female.

Text-fig. 5.—Atellana papilio, n. g., n. sp. lateral view of preadult nymph, slightly
distended, enclosing an adult male.

Text-fig. 6.—wNeolabidocarpus buloloensis (Gunther). Lateral view of holotype nymph.

flattened, and possessing definite caruncles. The coxal apparatus, as shown in
Gunther’s figures, is also typical of the Atopomelinae. Since the holotype is a nymph,
and the remainder of the material was destroyed during the war (Gunther, in litt.),
it appears best to keep this genus and species apart until fresh adult material
proves them valid or otherwise.

46 A SUMMARY OF THE ATOPOMELINAE,

Redescription of holotype nymph.—With single anterodorsal shield flanked by two
pairs of setae. Remainder of dorsum covered by striations with contours and setation
as shown. Anus terminal, with two internal sclerotized bars and two pairs of adanal
setae (this type of anus is also present in the female of Awustrochirus enoplus, see
inset). Capitulum and legs I and II typical, with recurved seta dorsally on tarsi I
and II. Clasping organ not clear in detail, but typical of Atopomelinae; probably with
pair of setae between coxae I and II. With pair of setae above coxae III and a
further seta above these. Coxae III and IV also not clear, but of general atopomeline
facies. Legs III and IV with usual four movable segments, the setal pattern of the
(foreshortened) tarsi being typical of cther atopomeline genera. Body length 370u.

LISTROPHOROIDES Hirst.

Diagnosis.—Atopomelinae with three dorsal shields which cover entire dorsum,
apart from apex of hysterosoma. Anterior dorsal (postcapitular) shield longer than
broad, flanked by two small setae and lateral sclerotized zones which serve for the
attachment of legs i and II. Middorsal shield subquadrate, with four setae along
anterior margin. ostdorsal shield longer than broad, always with seta in each
anterior corner and es pairs of setae on dise of shield, though additional setae may
sometimes be present marginally. Capitulum with two basoventral setae. Legs I
and II incrassate, with one strong seta dorsally on fused apical segments, and provided
with caruncles. Clasping organ between coxae I and II always with two transversely
striate areas, between which are a pair of setae. Two attenuate tracheal tubes present.
Genitalia of female between coxae III and preceded by a sclerotized arc; with two
pairs of anterior setae, two pairs of suckers and one pair of posterior setae. Coxae III
with three setae. Legs III and IV not enlarged; with four movable segments.
Penultimate segment of leg III with small dorsal seta. Dorsobasal seta on tarsus IV
weak. Male genitalia between coxae III and IV; with one pair of anterior setae, two
pairs of suckers and two pairs of posterior setae. Penis usually short, but exceedingly
long in L. mastomys. Coxae III with three setae. Anus without suckers, but flanked
by two setae. Posterior body lobe variable, but typically with three pairs of stalked
setae, of which the median pair is the strongest. Leg IV somewhat enlarged, with
dorsobasal seta of tarsus very strong and elongate. Tarsi IV also with two inner
apical sclerotized points and termina! caruncle. Genotype: L. aethiopicus Hirst by
monotypy.

The genotype has recently been redescribed and refigured (Lawrence, 1956), thus
putting this genus on a firm basis. A second species, Ll. expansus, was described by
Ferris (1932), who stressed the form of the anterior legs and the lack of spurs on
coxae III. However, the former character is only of specific value, and the spurs
described by Hirst for the genotype are artefacts (Lawrence, loc. cit.). Thus
Marquesania Womersley, 1943, monotypic for Ferris’s species, becomes a synonym of
Listrophoroides. :

Apart from LL. oryzomys, an American species which has been transferred to
Chirodiscoides below, there are now fifteen names referable to this genus. A close
study of these species has revealed a rather tangled situation, hence the full diagnosis
above. BHleven species (nine African, one Ceylonese and one semi-cosmopolitan) are
here recognized as valid.

Key to species of Listrophoroides,
1. Body very broad, terminating in two or four exceedingly long setae; postdorsal shield

reduced ini females On ws 2 thy Sr SClae rere cteteelateiencneteicenenc ir) ened iiellelcieiokceaiei onaitel ieee naan 2.
Body quite slender, never terminating in long flagellate setae; postdorsal shield not reduced
oa FESOTENIE, (Orn, IMENTAL) (EOE Thy CONGO) 5 ocanccocobacconongobonbounUnoo DO OON 3

bo

Female with apex of hysteroscma lacking pronounced conical process; postdorsal shield -
evenly sclerotized. Posterior body lobe of male with three pairs of stalked setae, the
MIECCIAN MD AnD CLUS sev CTVAES UL Ol) Same leicusns il MencneiennAaielsien Meek enenetenien ai eckeitaiiisiele nes .. bathyergians.

Female with apex of hysterosoma with large conical process; postdorsal shield wiih two
heavily sclerotized areas laterally. Apex of abdomen of male with subquadrate shield
bearine- ar flacellatessetan in) ealchy postemorNconner =). eee leineiene eerie eumpti.

BY ROBERT DOMROW. 47

3. Dorsl shields with very regular pattern of scales like those of fish or snakes; penis very
NO See rarnet trae tee) Sate yet cae cleanness odie foley cated SANDS anadodts. a uMuayfetlted ye oop Stroy yultetint aqcemalkaye) fete) las mastomys.
Dorsal Saige yr Bimo@WMEeIp WEAN S TOSS! Were SOOKE scococccsccnuvcosunccucuuaudad 4.

4. Dorsal shields heavily sclerotized and roughly pitted to produce a sponge-like effect;
venter of hysterosoma of female entirely covered by coarse, pointed papillae .. africanus.
Cuticle of dorsal shields otherwise; venter of hysterosoma normally striate and typically
WHTAOIE TORN ONNEK SS TG cose Gime oln G.rora 6 Oro OR oie ice GIceee SRS icin Pact hOncE Cha icity Decne Cec ecu eee oa SP De
IDersal siglals wiliin chisbaer Iliteene SibeenelOMs Gocoscoodsooe0cccuducocccauuunguGonuad 6.
IDOREA Saiglels ay MnO. ine ATe GUSIENHIOINS:. Gaoogoeo soup oo pldod bcoo no Uden oO eclol Old. o Dore 9.
6. Striations of dorsal shields irregular, and not evenly spaced and transverse; middorsal

oO

Sinielél jowoackee joOsteiclomhy Waren enmesrelorbhy SoscoscagcobobuvavocduonoungpdoonDOUDE Ts
Striations of dorsal shields evenly spaced and transverse; middorsal shield narrower
WOSKCIIORI NBM Mee ANCE IOM iy Weewey stare cens: seers skeen sate. cys cat les tobi oriay oleh ewe al ichcivel evista sce Je coy et etlavievie) site ceevauate) 8

7. Postcapitular shield flanked by two pairs of strong setae; body setae strong; venter of
hysterosoma of female without tubercles; coxae III of male simple; posterior body lobe
WERIRIK? COIR CIO” » Bee a isn rorercrcle a. oeo cCRC CHE Rey Re Een ete econ “aren Nica: CRC ten Farce oS Seo mee adherens.

Posteapitular shiela flanked by two pairs of very weak setae; body setae weak; venter or
hysterosoma of female with several drop-shaped tubercles posteromedially; coxae IIl
of male produced inwardly to form two sclerotized processes; posterior body lobe
StrOMahy Seleroiniwecl eimGl welll clemineg! ssoococasaoococtcoudnodnaouroeeoded lemniscomys.

8. Striae on dorsal shields very distinct; discrete and crescentie anteriorly and laterally,
but transverse medially and posteriorly; striations of clasping organ not reaching
Ievitereail GCkSSs Cie COpeyey IL elvaVOl TOL 5 Beach oicenl den jelolo Srtea, Ome loiclO oCrOrcucricaa Ohare Saaioio te Ginn leggadilla.

Striae on dorsal shields weaker (sometimes lacking on middorsal shield), and evenly
transverse; striations of clasping organ reaching extreme lateral margins of coxae
Flite ARO CHES rare ert acdyr sh sBenspstarichtrcptcyatiem eto isaieuicuie) skein selenite shee feet oo aus euslarious) sPeledsue el ebeake) Schesicansystecats expansus.

9. Lateral margins of hysterosoma serrate; female with posterior margin of middorsal
shield even, and venter and dorsal apex of hysterosoma non-striate; male with expanded
posterior body lobe with four short and two long stalked setae ........... aethiopicus.

Lateral margins of hysterosoma smooth; female with posterior margin of middorsal shield
armed with stout median spine; venter of hysterosoma with longitudinal striae, and
apex dorsally with transverse striae; male with posterior body lobe not expanded and
vglulameet 0 Ollcgek OUI el OMG) WSCLA CE aie intua cies nucde ave cate mc mete ieee yr er saan esos Ge a womersleyi.

LISTROPHOROIDES ADHERENS (Trouessart, 1893), n. comb.

Description of female—A slender, well-sclerotized species; body length 378—-3892u.
Anterodorsal (postcapitular) shield slightly longer than broad, flanked laterally by
two pairs of setae above insertions of legs I. Middorsal shield subquadrate, with
irregular scale-like markings and without setae. Postdorsal shield longer than wide,
with rather more distinct scale-like markings, and an anterior and posterior transverse
row of four slender setae. Apex of hysterosoma triangular, with minute apical point;
not covered by postdorsal shield, but with two slender setae. Venter: Capitulum with
two basal setae. Genitalia placed between coxae III, with usual three pairs of small
setae and two pairs of minute suckers. Posteroventral margins of hysterosoma covered
by lateral lobes of postdorsal shield. Medially with fine longitudinal striae, but
without tubercles; with two seta posteriorly. Anus longitudinal and subterminal,
flanked by three pairs of slender setae, the posterior pair being much the shortest.
A tracheal system similar to that of L. expansus figured below is present. Legs:
Legs I and II typical of subfamily. Inner surfaces of coxae I and II hollowed and
striate, with a pair of small setae between the two striate zones. Coxae I] with
stronger seta on posteroventral margin (this seta probably represents the outer pair
of the six seta normally present flanking the postcapitular shield and along the
anterior margin of the middorsal shield). Coxae III with single anterior seta and
flanked by a larger and a smaller seta. Coxae IV without setae. Legs III and IV
with usual four movable segments; penultimate segment of leg III apparently without
dorsal seta.

Description of male—As in female dorsally and anteriorly, but somewhat smaller;
length of body 350-360u. Genitalia set between coxae IV, with usual three pairs of
small setae and two pairs of minute suckers. Anus longitudinal and subterminal,
with pair of small adanal setae anteriorly. Posterior body lobe simple, with extremely
shallow posteromedian lobe; with two pairs of long slender ventrolateral setae and
one pair of small terminal setae; also with two setae arising dorsally, being the outer
pair of the posterior row of four setae on postdorsal shield. Legs III (including coxae)

48 A SUMMARY OF THE ATOPOMELINAE,

as in female. Legs IV somewhat enlarged, with dorsobasal seta of tarsus very strong
and much elongated. Tarsus IV with two minute ventroapical spurs. Caruncle
terminal.

Remarks.—Trouessart (1893) placed this African species in the genus Campy-
lochirus, but it is unrecognizable from the available descriptions. (The specific name
was spelt adhaerens by Trouessart in 1917, but this is regarded as an erroneous

B

Text-fig. 7.—Listrophoroides adherens (Trouessart). Left, venter of female. Right,
dorsum of female.

Text-fig. 8.—Listrophoroides adherens (Trouessart). Venter of male. Inset at right,

middorsal shield of Listrophoroides ajricanus Radford; below, outline of posterior margin of
mid@orsal shield of female of Listrophoroides womersleyi (lawrence).

subsequent spelling.) Through the courtesy of Dr. Marc André I have been able to
reexamine Trouessart’s ten syntypes from Anomdalurus fraseri erythronotus Milne
Edwards from the Congo, Dybowsky coll. The specimens have been remounted
successfully on two slides which have been returned to the Muséum National d’Histoire
Naturelle in Paris. One slide (with the original labels) contains the lectoholotype

BY ROBERT DOMROW. 49

female and the lectoallotype male, and the other six paratype females and two
paratype males. The species is a typical member of the genus Listrophoroides, and
may be separated from the other species of the genus by the above key. Lawrence
(1956) did not include it in his revision of Listrophoroides, although he had examined
the material.

LISTROPHOROIDES AETHIOPICUS Hirst, 1923.

This species has been discussed above, but it should be noted that in Hirst’s
figure the third (posterior) pair of genital suckers are artefacts, and that the second
pair of postgenital setae are not depicted. In Lawrence’s figure (1956) the adanal
setae are lacking. I attach no significance to the minor variation in the posterior
body lobe and dorsobasal seta of tarsus IV in the male.

LISTROPHOROIDES AFRICANUS Radford, 1944 (emend.).

This species was described from the same host and locality as L. mastomys
Radford. The slide I have examined is labelied as containing three female L. mastomys,
but really contains one female L. mastomys and a pair of L. africanus. Vertrally
the male may be recognized by the strongly sclerotized, fused coxal plates of legs III
and IV, which are extended inwardly, though not meeting medially, behind coxae IV
to flank the genitalia posteriorly. The clasping apparatus is striate. The most striking
character of this species is, however, the texture of the dorsal shields. These are
heavily sclerotized, and with typical fine punctae. However, this punctation is overlaid
by very numerous, much larger and deeper pits of variable size and shape, which are
evenly spread over the entire surface, producing a rough, areolate and almost sponge-
like appearance. The female is in lateral view and freshly moulted, but has similar
dorsal shields to the male. The venter of the hysterosoma is entirely covered with
strong, outstanding tuberculate processes. Lawrence (1956) in his key says that in
L. mastomys the female has the “ventral surface roughened with large sharp
granules”, and probably examined this same slide. However, he has associated the
sexes wrongly, since in the female in question the dorsal shields have the same
characteristic texture as the male of L. africanus. As the name of the genus is of
masculine gender, the termination of the specific name has been amended.

LISTROPHOROIDES BATHYERGIANS Radford, 1939.

This characteristic species and L. zumpti form a distinct group found only on
bathyergid rodents, the other species being typically from Muridae. These two species
may be recognized by their broad bodies and the possession of long terminal flagellate
setae. Lawrence (1956) says that both lack striae on the clasping apparatus between
coxae I and II, but this is incorrect. Under oil immersion, typical striae are present
both in Radford’s types and in specimens of L. zumpti with collection data as in the
type series. This character appears to be constant throughout the subfamily, but
should be checked in Centetesia Lawrence.

LISTROPHOROIDES DASYMys Radford, 1942.

This species, which certainly belongs to Listrophoroides, is known only from a
single male. As the description contains no detail of value in determining its specific
status, and the figure is semidiagrammatic and inaccurate, I have left this form as a
species inquirenda. It will undoubtedly prove to have striae between coxae I and II,
two pairs of small genital suckers, and 4-segmented legs III and IV. The description
calls for a long seta on leg II, but in the figure it is leg I that has a long seta. The
dorsal surface is not described.

LISTROPHOROIDES EXPANSUS Ferris, 1932.

Synonymy.—Womersley (1943) described and figured Marquesania expansa form
queenslandica from rats from S.H. Queensland on the basis of the lack of striations
on coxae I and II, and the absence of a “tooth” on leg I. However, he was mistaken
on both points, and the form is here regarded as a synonym of Ferris’s species.
Coxae I and II in all stages are obviously striate as illustrated. An oval, punctate
sternal area is always present between the striations of coxae II. The “tooth” on

50 A SUMMARY OF THE ATOPOMELINAF,

leg I requires further explanation. In dorsal and particularly in ventral view, leg I
appears to have a sclerotized process, but this is due to observing a narrow hyaline
lobe from end to end. In dorsolateral view the hollowed inner surface of this lobe
may be seen to clasp the shaft of the hair of the host, the free segments of legs I
and II passing right around the hair. The mode of attachment is similar to that
figured by Lawrence (1954) for Tenrecobia pauliani (the original spelling “pauliana”’
is in contravention of Article 14, and thus subject to automatic correction) from
Madagasear. Some other details of Womersley’s description also need correction. The
anterior dorsal (posteapitular) shield is not hroadly transverse, but decidedly longi-

150»

Text-fig. 9.—Listrophoroides expansus Ferris. weft, dorsum of male preadult nymph.
Centre, dorsum of female preadult nymph. Right. venter of female preadult nymph. Below,
dorsum of penultimate nymphal stage showing suture line.

tudinal, with distinct lateral margins and setae arranged as figured for the nymph.
The areas to the side of this shield serve for the attachment of legs I and II.
The inverted Y containing the genitalia of the adult male is heavily sclerotized
cuticle rather than a definite structure.

A second synonym of this species is Listrophoroides trdgdrdhi Radford, 1940, the
striking similarity between the published descriptions being confirmed by the study of
specimens kindly lent to me by Dr. Radford. Of the available figures, those of
Ferris are the best.

Distribution.—This species is apparently almost cosmopolitan on Rattus rattus
(Linné) and R. norvegicus (Berkenhout), and has probably spread onto native rodents
from these two species. It has been recorded from Sierra Leone, Uganda, Ceylon

-

BY ROBERT DOMROW. 51

and the Maldive and Marquesas Is. It is also common on R. assimilis (Gould) in
S.H. Queensland, and may now be recorded from North Queensland as follows:
R. rattus, Sundown, 23.viii.1956, and Innisfail, 9.x.1956, and &. assimilis, Bartle Frere,
13.11.1957. Recent material from R. gestroi gestroi (Thomas), Porebada Village, Port
Moresby, Papua, 19.xi.1956, comprises preadult nymphs of both sexes. Hach nymph
is still enclosed in the skin of the penultimate nymphal stage, which shows a distinct
central longitudinal suture from the anterior edge of the middorsal shield to the
apex of the hysterosoma, which may be still intact or gaping widely. In one specimen
the final nymph came out of its old skin during mounting procedure. Womersley
only described the adults, and although the present nymphs are quite pale, they are
adequate for illustration and are described below.

Description of preadult male nymph.—Length 350-362y. Anterodorsal (post-
capitular) shield longitudinal, slightly wider posteriorly, and flanked by two small
setae. Suture between postcapitular and middorsal shield well marked, with four setae
arranged along it. Middorsal shield subquadrate, with few scale-like markings. Third
(postdorsal) shield covering remainder of dorsum, with about eight setae as figured.
Apex of hysterosoma without well-developed accessory lobe of adult, but with six
fairly strong setae. Venter: Genitalia small and poorly developed, situated between
coxae III and IV, and without any inverted Y sclerotization. Legs as in preadult
female nymph, but leg IV slightiy thicker.

Description of preadult female nymph—Length 374-3854. The structure of the
anterior two pairs of legs is described above. Anterior half of body as in preadult
male nymph. Postdorsal shield constricted medially, with transverse markings, and
flanked by lateral cuticular areas with longitudinal striations; with six setae.
Venter: Two zones of striae between coxae I and II, the former with small seta near
inner posterior margin, and latter extending to extreme lateral margins of body.
Radford’s interpretation of these normal striae as a “toothed semicircular process” is
incorrect. Oval, punctate sternal area between coxae II. Genitalia between coxae III,
with setae and minute suckers arranged about an inverted Y. Anus subterminal,
flanked by about six small setae. Legs III and IV not greatly enlarged. With distinct
tracheal system on either side, consisting of a spherical atrium between coxae I and II
opening into a single trachea, which becomes thicker along its course, and thins
again posteriorly. A similar tracheal system was illustrated by Hirst (1921) for
Listrophoroides aethiopicus.

Description of penultimate nymphal stage—Length in distended condition 409—432u.
Similar to preadult female nymph anteriorly and ventrally. Dorsum with postcapitular
shield and weakly defined middorsal shield. Remainder of body covered with striations
which are longitudinal midlaterally and transverse posteriorly. With central suture-
line along dorsum, through: which the preadult nymph emerges. Legs III and IV
slightly thinner than figured for preadult nymphs. The earlier, rather similar nymphal
stage with six legs described by Ferris has also been seen. ;

LISTROPHOROIDES LEGGADILLA Radford, 1947.

This form was originally described as a full species from Ceylon, but may prove
to be a variant of the cosmopolitan L. expansus. Both forms are of identical facies
and setation, particularly as regards the shape of the dorsal shields and their texture,
and the posterior body lobe and the form and setation of the legs in the male. The
postdorsal shield is narrower than shown by Radford, and the ventrolateral hystero-
somal shields, if present, very weakly defined and with margins indiscernible. The
transverse markings on the dorsal shields are much more distinct than in L. expansus,
but are not crescentic as depicted by Radford, except laterally and anteriorly. The
discal markings, especially posteriorly, are transverse and evenly spaced as in
L. expansus. The Y-shaped sclerotization around the male genitalia is not well defined,
and the striae on the clasping apparatus do not reach the extreme lateral margin of
the coxae as in L. expansus, but finish evenly, well in from the edge of the coxae, along
a longitudinal line as figured by Radford.

52 A SUMMARY OF THE ATOPOMELINAE,

LISTROPHOROIDES LEMNISCOMYS Radford, 1940.

Both this species and Marquesania imbricata Lawrence, 1954, were described from
Lemniscomys in Uganda and Zululand respectively. Comparison of both sexes of
Lawrence’s species from the type series and of Radford’s from the type host has
revealed complete identity in cuticular pattern and fine detail, including the two -
characteristic crescentic marks at the anterior edge of the middorsal shield, the
large bases to the gnathosomal setae, the posteromedian ventral tubercles of the
female and the posterior body lobe of the male. WM. imbricata is therefore considered
a synonym of L. lemniscomys.

LISTROPHOROIDES MASTOMYS Radford, 1940.

This species may be readily recognized by its evenly scaled cuticle, which is a
reminiscent of snake or fish skin. Some rather strong longitudinal lines are present
laterally on the middorsal shield. Specimens of both sexes from the type series of
this species agree in cuticular pattern and in all fine detail with the full description
of Marquesania elongata Lawrence, 1951, and possess in the male both the characteristic
posterior body lobe and the ventrolateral, inwardly directed points behind coxae IV.
Lawrence (1951, 1956) is in error in saying coxal striae are absent. The enormously
long penis is characteristic of the species, but apparently could not be seen by
Lawrence, whose only specimen was obscured by a hair. The two species are here
considered identical. The females referred to this species in Lawrence’s key (1956)
are really L. africanus (q@.v.), which was originally collected from the same host and
locality.

LISTROPHOROIDES WOMERSLEYI (Lawrence, 1951).

This species may be immediately recognized by the characteristic pattern of
cuticular striae at the apex of the hysterosoma of the female, and (at least in the
female) by a distinct pointed median process (not shown in Lawrence’s figure) on
the posterior margin of the middorsal shield. Two nymphs examined show a
longitudinal middorsal suture and dorsal shields similar to those figured above for
L. expansus:

LISTROPHOROIDES ZUMPTI Lawrence, 1956.
This species has been discussed under L. bathyergians.

CHIRODISCOIDES Hirst.
This genus is monotypic for C. caviae Hirst, 1917, the widespread parasite of
guinea-pigs, which originally came from South America. The species is well illustrated
in Hirst (1922) and Lawrence (1956).

CHIRODISCOIDES ORYzZOMYS (Radford, 1954), n. comb.

This American species was originally described as a Listrophoroides, but has the
following important generic characters in common with C. caviae: Males with four
setae between coxae III and IV; genitalia flanked posteriorly by two setae and four
minute suckers; anus surrounded by two setae and two large suckers; posterior body
lobe well developed; leg III normal; leg IV enlarged; tarsus IV hooked distally, with
two inner apical sclerotized points and subapical external caruncle. Both sexes with
three dorsal shields, the middorsal being rather narrow. Females with postdorsal
shield truncate, leaving posterior half of hysterosoma covered only by annulated
cuticle. Radford’s species has therefore been reassigned as a second species of
Chirodiscoides. Since his figure is incorrect in several details, the following
supplementary data are given.

Male—Clasping apparatus striate between coxae II as well as coxae I. Dorsum
with three shields as follows: Postcapitular shield deeply convex posteriorly, extending
back to level of coxae II; flanked posteriorly by four setae. Middorsal shield very
narrow medially, reaching back only to level of posterior edge of basal movable
segment of leg II; straight posteriorly and concave anteriorly to accept postcapitular
shield. Postdorsal shield covering remainder of hysterosoma and deeply cleft
posteriorly. Penis slender and of moderate length, running forward and then turning
abruptly backwards. Posterior body lobe with strong median cleft, the inner posterior

angles of the two lateral processes being turned inwardly;
ventrally and four stalked setae laterally on each process.

BY ROBERT DOMROW.

53

With strong, rod-like,

transverse coxal apodemes between legs III and IV, that of IV being the thickest;

united medially by strong longitudinal strut
similar pattern is illustrated for Austrochirus perkinsi above).

and enclosing four setae (a rather
Legs III and IV with

Synopsis of the Subfamily Atopomelinae (Listrophoridae).
(Genotypes listed first, followed by other species in alphabetical order.)

Genera and Species.

Host.

Locality.

ATELLANA, D.g.
A. papilio, n.sp... at
ATOPOMELUS Trouessart, 1917.
A. locusta Trouessart, 1917
AUSTROCHIRUS Womersley, 1943.

A. queenslandicus Womersley, 1943 ..

A. enoplus Domrow, 1956

A. perkinsi. n.sp. Re oh

A. sminthopsis Womersley, 1954
CAMPYLOCHIRUS Trouessart, 1893.

C. chelopus Trouessart, 1893. ..

C. latus Be 55
CENTETESIA Lawrence, 1954.

C. tiptont Lawrence, 1954

C. tessellata Lawrence, 1954
CHIRODISCOIDES Hirst, 1917.

C. caviae Hirst, 1917

C. oryzomys (Radford, 1954)
CRICETOMYSIA Lawrence, 1956.

C. andréi Lawrence, 1956

CYTOSTETHUM Domrow, 1956.
C. promeces Domrow, 1956
C. charactum Domrow, 1956
C. nanophyes Domrow. 1956 ..
C. pseudocharactum Domrow, 1956
C. trachypyx Domrow, 1956
LISTROPHOROIDES Hirst, 1923.
. aethiopicus Hirst, 1923
. adherens (Trouessart, 1893)
. africanus Radford, 1944
bathyergians Radford, 1939
dasymys Radford, 1942
. elongatus (Lawrence, 1951)
expansus Ferris, 1932
imbricatus (Lawrence, 1954)
. leggadilla Radford, 1947
lemniscomys Radford, 1940..
mastomys Radford, 1940

. trdgardhi Radford. 1940
. womersleyt (Lawrence, 1951)
. zumpti Lawrence, 1956
MARQUESANIA Womersley, 1943.
NEOLABIDOCARPUS Gunther, 1942.
N. buloloensis (Gunther, 1940)
TENRECOBIA Lawrence, 1954.
T. pauliant Lawrence, 1954

SESESESESESESESESESE SSE iSns|

. queenslandicus (Womersley, 1943). .

Trichosurus.
Neotetracus.
Thylacis.
Hydromys.
Phascolarctos.

Sminthopsis.

Pseudocheirus.

Hemicentetes.
Hemicentetes.

Cavia.
Oryzomys.

Cricetomys (!).

Potorous.
Potorous.
Potorous.
Potorous.
Potorous.

Cricetomys.
Anomalurus.
Mastomys.
Bathyergus.
Dasymys.
Aethomys.
Muridae.
Lemniscomys.
Leggadilla.
Lemniscomys.
Mastomys.
Rattus.
Muridae.
Otomys.
Georychus.

Thylogale.

Ericulus.

Queensland.

China.

Queensland. Occasionally on Perameles.
Queensland.

Queensland.

South Australia.

Tasmania.
Nomen nudum.

Madagascar.
Madagascar.

Widespread.

U.S.A. New combination.

Synonym of Campylochirus.

Africa (!). Synonym of Campylochirus
chelopus.

Queensland.
Queensland.
Queensland. Also Tasmania.
Queensland.
Queensland.

Africa.

Congo. New combination.

Sierra Leone. emend. °

South Africa.

Uganda. Species inquirenda.

Natal. Synonym of LZ. mastomys.
Widespread.

Zululand. Synonym of LZ. lemniscomys.
Ceylon.

Uganda.

Sierra Leone.

Queensland. Synonym of L. expansus.
Widespread. Synonym of L. expansus.
South Africa.

South Africa.

Synonym of Listrophoroides.

New Guinea. Species inquirenda.

Madagascar. emend.

usual four movable segments (the long medial segment of leg IV in Radford’s figure

is weakly divided centrally).

Penultimate segment of leg III with small dorsal seta.

Tarsus IV hooked distally, with twe inner apical sclerotized points as in JListro-
phoroides; caruncle set dorsally and subapically.

Female.—Anterior half of body as in male.
to midway between level of coxae IV and apex of hysterosoma;

Postdorsal shield truncate, extending

quite concave

with two normal setae

54 A SUMMARY OF THE ATOPOMELINAE,

posteriorly. Apex of hysterosoma simple. Genitalia between coxae III. Legs III
and IV of normal size; tarsus III with enlarged dorsobasal seta.

Acknowledgements.
In addition to those mentioned in the text, I am grateful to Drs. I. M. Mackerras .
and HE. H. Derrick for reading my manuscript, to Dr. K. H. L. Key for advice on
some nomenclatural problems. and to Miss EH. Wood for her careful typing. .

References.

Domrow, R., 1956a.—Notes on Ausiralian fur-mites (Listrophoridae, Atopomelinae), with
description of a new genus. Proc. LINN. Soc. N.S.W., 80: 191-200.

——§—, 1956).—The genera Campylochirus Trouessart and Austrochirus Womersley in
Australia (Acarina, Listrophoridae). Proc. Linn. Soc. N.S.W., 80: 234-239.

Domrow, R., and SmitH, D. J. W., 1956.—Acarina from five hundred native mammals from
Queensland. Proc. LINN. Soc. N.S.W., £0: 201-206.

EwiIne, H. W., 1929—A manual of external parasites. Bailliere, Tindal! & Cox, London.
Page 44. i

HERRIS, G. F., 1932.—Hctoparasites of Marquesan rats. Bull. Bishop Mus., Honolulu, 98:
117-127.

GUNTHER, C. H. M., 1940.—A listrophorid parasite of the wallaby from New Guinea. Proc.
Linn. Soc. N.S.W., 65: 3538-354.

, 1942.—Notes on the Listrophoridae (Acarina, Sarcoptoidea). Proc. Linn. Soc.

N.S.W., 67: 109-110.

Hirst, S., 1917.—On three new parasitic Acari. Ann. Mag. nat. Hist., (8) 20: 431-434.
————, 1921.—Notes on parasitic Acari. A. On the presence of a system of tracheal tubes
in the families Sarcoptidae and Listrophoridae. J. Quweckett micr. Cl., 14: 229-232.

, 1922.—Mites injurious to domestic animals. Brit. Mus. (Nat. Hist.), Hconomic
Series No. 13, 107 pp.
, 1923.—On some new or little known species of Acari. Proc. zool. Soc. Lond.,
971-1000.
LAVOIPIERRE, M., 1946.—New records of Acari from Southern Africa and the Belgian Congo.
J. ent. Soc. S. Afr., 9: 78-81.
LAWRENCE, R. F., 1951.—New fur mites from South African mammals. Ann. Natal Mus.,
12: 91-133.
, 1964a.—Two new fur-mites from rodents. J. ent. Soc. S. Afr., 17: 38-46.
—, 1954b.—Studies on the listrophorid mites (Sarcoptiformes) of Centetidae from
Madagascar. Mém. Inst. Sci. Madagascar, 9A: 129-149.
— , 1956.—Studies on South African fur-mitcs (Trombidiformes and Sarcoptiformes).
Ann. Natal Mus., 13: 337-375.
RADFORD, C. D., 1939.—Notes on some new species of parasitic mites. Parasitology, 31: 243-257.
, 1940.—Notes on scme new species of parasitic mites. Parasitology, 32: 91-104.
, 1942.—New parasitic mites (Acarina). Parasitology, 34: 295-307.
1944._New parasitic mites (Acarina) from rodents. Parasitology, 35: 161-166.
, 1947.—Parasitic mites from snakes and rodents (Acarina Cheyletidae, Listro-
phoridae and Laelaptidae). Proc. zool. Soc. London., 117: 228-240.
, 1950.—The mites (Acarina) parasitic on mammals, birds and reptiles. Parasitology,

40: 366-394.
, 1954.—Three new species of fur mites (Acarina: Listrophoridae). Riv. Parassit.,

15: 593-599.
TROUESSART, HE. L., 1893.—Note sur les sarcoptides pilicoles (Listrophorinae). C. R. Soe.

Biol., Paris, (9) 5: 698-700.
, 1917.—Troisi¢me note sur les sarcoptides pilicoles et description de genres nouveaux.
Bull. Soc. zool. Fr., 42: 151-158.
WOMERSLEY, H., 1943.—Australian species of Listrophoridae Canest. (Acarina) with notes on
the new genera. Trans. roy. Soc. S. Aust., 67: 10-19.
, 1954.--Two new species of ectoparasitic mites from pouched mice, Sminthopsis,
from Australia. Rec. S. Aust. Mus., 11: 117-120.

5D

INHERITANCE OF OIL CHARACTERS IN EUCALYPTUS.

By L. D. Pryor, Parks and Gardens Section, Department of the Interior, Canberra,
and L. H. Bryant, Division of Wood Technology, Forestry Commission
of New South Wales.

(Five Text-figures.)
[Read 30th April, 1958.]

Synopsis.

Examination of segregates from EH. cinerea x EH. Macarthuri and E. pauciflora x E.
Robertsoni or E. dives, together with Fl hybrids between #. Maideni x EH. rubida, shows
that recombination between oil yield and components on the one hand and morphological
characters on the other occurs. Yield is sometimes determineaG in a far-reaching way in
accordance with that of one parent. The variance of physical and chemical constants of
oils derived from segregating hybrids is much greater than that of those for the parents.
Oil constituents may be found in much greater quantity in some segregates than in either
parent.

It has been evident that oil characters of Eucalypts, especially yield and the
nature of their various constituents, are determined to a large extent by heredity.
The review of the genus by Baker and Smith (1920) and their attempt to develop
its taxonomy by the consideration of oil composition shows the high degree of oil
specificity which species have sometimes been found to possess. On the other hand,
something is also known about the variation within species largely as a result of
the recognition of “physiological forms” by Penfold and Morrison (1927) in EH. dives.
Such forms are more or less similar morphologically, differing mainly in the chemical
nature of their oils. These have been found since also in many other species of
Eucalypts as well as in other genera of the Myrtaceae.

A study by Willis (1951) of families planted in one locality, made up of several
progenies from different oil varieties of H. dives from different localities, showed that
within this species the characters of the parents were largely repeated in the progeny
too. Bryant (1950) has summarized unpublished work carried out with Smith-White
indicating that wide variations in oil yield occur within a number of commercially
important species. Major constituents of the oils were also shown to vary within
the species and also even within a single tree but there has been little study of the
variation in minor components.

The biological significance of essential oils in plants has not yet been clearly
established, although different theories have been advocated. Two main ideas may
be mentioned. The first, held by James (1953) and others, suggests that in plants
generally, constituents such as alkaloids or essential oils are produced as a by-product
of metabolism and have no adaptive significance. The second, demonstrated in some
plants by Dethier (1941) and held by Barber (1955) for Eucalypts (and oil or alkaloid
bearing plants generally), is that the varying characters of the oils indicate part of
a system which is of distinct adaptive value by determining the palatability or
resistance to insect pests. It has not yet been established for Eucalyptus that
palatability to insects is influenced by the chemical nature of the essential oil, but
if this is so one could imagine through time a sequence of changes both in oil
composition and insect variation, leading to a series of alternating changes in closely
adaptive responses in both plant and insect. Dethier (1941), using pure compounds
found in the oils of some species of the Rutaceae and Umbelliferae, was able to induce
Papilio larvae to eat filter paper treated with them and so indicate positive attraction
by them for these insects. In Hucalyptus strong preferences by leaf-eating insects are

PROCEEDINGS OF THE LINNEAN Society OF NEw SoutH WALES, 1958, Vol. lxxxiii, Part 1.

56 INHERITANCE OF OIL CHARACTERS IN EUCALYPTUS,

already known, and in one or two combinations there is evidence that this is an
inherited character (Pryor, 1953). It is possible that the major oil constituents in
Eucalyptus, while of great consequence in a similar system in some earlier evolutionary
period, may now be of little importance, and the real significance as far as insect
attack is concerned may rest with quantitatively minor constituents.

There is considerable interest, therefore, both from the point of view of evolutionary
genetics and in understanding physiological forms, in gaining knowledge of the mode
of inheritance of oils in Hucalyptus. The study might also provide basic information
which could lead to very important results in applied work, such as in the control
of insects feeding on Hucalyptus on the one hand and in the Hucalyptus oil industry
on the other. Investigation can proceed some distance by raising progeny from open
pollinated natural hybrids which give access to segregating groups of individuals
containing recombinants derived from pairs of parent species. This will permit some
deductions concerning the inheritance of oil characters.

Method.

Combinations of parent species were selected which showed some strongly
contrasting characters both in oils and morphology. Open pollinated progeny raised
from naturally occurring hybrids have been examined at about the age of six years.
Progeny of the parents of the same age, raised at the same time, have been compared.
The offspring of parents (supposedly the “pure” species) are not necessarily of
precisely the same stock which led to the production of the hybrids, and there is a
potential source of error here which imposes some limitations on the interpretation
of the data, and may account for some minor anomalies which have been found.

The crude oil samples were obtained by steam distilling 10 lb. of green leaves
and small twigs collected in the same way and in the same position from each of the
trees. Approximate weighing to a little more than 10 lb. with a spring balance was
earried out in the field, and a more precise weighing to reduce the sample to
10 lb. + 3 02z., was made in the laboratory. The leaves were steam distilled to obtain
the first oil sample. The physical and chemical analyses were carried out by one of
us (Bryant). Two sets of data were compiled. Firstly, the constants usually calculated
for essential oils were established, particularly specific gravity, refractive index,
optical rotation and saponification number. Secondly, by a method of circular
chromatography (Bryant, 1955) using glass coated with inagnesol, an assessment of
most of the oil components was made. The oils were developed with n.hexane
containing 15 parts by volume of ethyl acetate and then sprayed with concentrated
H.SO, or examined under ultra-violet light. Most of the oil constituents reacted with
the H.SO, to give characteristic colours. Cymene and the pinenes did not react
satisfactorily and, although they are known to occur to some extent in these oils,
their proportion was not assessed. To assess recombination between oil features and
other characters, measurements of selected leaf features were made as well. In some,
the leaf shape as indicated by the length-breadth measurement was used, and in
others the angle which the primary veins made with the midrib. Transformation
of the length-breadth figures to a logarithmic scale results in reducing the variance
of the data for the parents in each case to more nearly the same level, and makes
the relationship clearer between the various groups and intermediates.

A small amount of material was also available from Fil hybrids produced by
manipulation, and this, though limited in extent, gives some additional indications
of the inheritance pattern.

E. cinerea x H. Macarthuri.

The hybrid H. cinerea x H. Macarthuri is found naturally from time to time
where these two species meet in the field. Two segregating progenies were raised
from two separate hybrid trees of this combination found at Paddy’s River on the
Hume Highway near Marulan, N.S.W., from which open pollinated seed was collected.
The parent trees differ widely in morphological and oil characters. Table 1 shows
the principal physical differences between the two species. Table 2 and Figures la
and 1b show the range of variation of oil and morphological characters within the

BY L. D. PRYOR AND L. H. BRYANT. ; 57

parent species and in the segregates. These latter show marked segregation and
recombination of beth characters, and contain individuals which closely resemble
either parent together with a series of intermediates between them.

~ TABLE 1.
— E. cinerea. E. Macarthuri.

Juvenile leaves .. | Orbicular. Lanceolate.

Sessile. Sessile.

Glaucous. Green.
Mature leaves .. | Sessile. Petiolate.

Opposite. Alternate.

Glaucous. - Green.
Oil yield se .. | High (about 1:75% vol./wt.). Low (about 0:25%).
Main constituents .. | High cineole (about 40%). Cineole nil.

Geranyl acetate (nil). Geranyl acetate (about 50%).

In the hybrid progeny 50/755a there are more segregates approaching FH. cinerea
than #£. Macarthuri. In 50/755 they are approximately evenly distributed between
the parental limits. It will be noticed that the quantity of oil produced in both
hybrid progenies is within the range of the low-yielding parent (#H. Macarthuri),
and there is not a single exception to this (Fig. la). This suggests, therefore, that

°. Sine
nN ® @ e oF
a ae
e
; 2
aS x
s
S
; a x
ad
& fy come § *
x ® =
Q 2 ae AG)
z 8 Ss x x
s 8 5S 8
i :
a
= ae ;
N x sep i fe
> S x ® 0 x J se
x 5 x ey 4
mx 4
a
-2 = 7] 2 3 7 F o
Leaf log “/a Lear Log. */a8
la &. cinerea - £ Macarthur 1h 4&ctherea - £. Macarthur
s e s e

Fig. la.—Yield of oil in ml. per 10 Ib. of green leaf against juvenile leaf shape is shown.
The “x’s” indicate hybrids. It will be seen that the leaf shape in the hybrid progeny ranges
between either parent, but that the oil yield is entirely within the range of the H. Macarthuri
parent.

Fig. 16.—Saponification number in relation to leaf shape shows in this particular progeny
a tendency for the #. cinerea characteristic of low saponification number to persist in the
hybrid progeny indicated by the ‘“x’s’, but there are some recombinants which have
#H. cinerea leaf shape but saponification number equal to that of H. Macarthuri and one
which is the reverse.

the factors determining quantity of yield in the H. Macarthuri parent are dominant
in this combination, and they determine low yield. On the other hand, it will be seen
that the amount of geranyl acetate in the oil assessed by saponification number,
when compared in relation to the morphological character of leaf shape, shows distinct
evidence of recombination. Hybrids 23 and 34, while having a leaf shape which

D

58 ‘ INHERITANCE OF OIL CHARACTERS IN EUCALYPTUS,

approaches that of HE. cinerea, have a saponification number which is closely comparable
with that of H. Macarthuri, whereas, on the other hand, No. 16 has a leaf shape
which is near the centre of the range of #H. Macarthuri, but has a much lower
saponification number. In the data as a whole there is low correlation between leaf’
shape and saponification number, which suggests rather free recombination between
these two characters (see Fig. 1b). Both hybrid progenies are somewhat limited,
and a considerably larger population would be desirable to assess more accurately
the pattern of recombination, and to determine whether there is any tendency for
characters to recur in association in any degree, perhaps suggesting linkage.

In Table 2 the presence of the various constituents of the oils (excluding terpenes.
such as the pinenes already mentioned which do not react readily with concentrated
H.SO,) has been assessed by the chromatographic method mentioned above. It will
be observed that cineole and geranyl acetate, which could not be detected in the
parent #. Macarthuri and EH. cinerea oils respectively, are both present together in
the oils of some of the hybrid individuals. In no case is there a hybrid oil where
both these constituents are absent. Geraniol, present in small quantities in all

TABLE 2.
Oil Constituents (Hacept Cymene and Pinenes) of cinerea, Macarthuri and Hybrids.

Fluo-
Geranyl Hudes- Sesqui- rescent
— No. Acetate. Geraniol. | Cineole. | Hudesmol. mene. terpene. Com-
ponent.

| 5258 3

Mx
x

x XX

E. Macarthuri. 64

XS 2S 08 OS os OK BK OX OS OK OS OK OS OK CK 2K OS

x
x

oS ON OS 26 OS 3K OK OK OS OK OK OS OE OK RK OS OK OK OK OR
x

for)
or
x OS OS BS BK RS OS OS MOK OS OK OK OX OS OK OK OK OK OOS
KR KX KOK KKK OS OK OK OK
OS Oh ES es 2S ES OS OS OST BS BS OS POON ES BS OS OM OM
BS oS OS OS OS PS OS OS BS OS os O6 O46 OM BS 24 PS es OS OX
|
os Ps OS Oh OS PS OS ES OS OS BS ES PS OS eS OS OS OS OS eS
“S 2S OS OS 8% OS OX OK OS EK OS OS OK OK OK OK SOS OS
DK Oh 2G BS OS BK BS OS OS OM OS OS OS 2S OS OS OS OM OS OX
PS Px 25 PS BS Om OS BX ES OK OC OS OS OK BS OS OS PS OS OS
x
x

x

nS
pia
|

| IxX|x x

ws
o> oO
||
! fl

nN
co
|

| Ix |

E. cinerea. 51 -

(o2)
oi
|
|

par
S 0
oOo @
lll
|

|

|
xxxxxxXxXxXxXxXXXXXXXXXX*X
xxxxXxXxXxXxXxXXxXXXxXXXXXX*X
De aE OX KEE XK KX XXX
xxxXxxXxxXxXxXxXxXXXXXXXXXXX*X
x
Ibs allel lel bal bs 4] bal bal bal bal allt) bal -alle-allb-a) b-ball a
x
x
|

x x Ixlx Pix x x |

x
x
x
Se | Se
x

t BY L. D. PRYOR AND L. H. BRYANT. — 59

TABLE 2.—Continued.
Oil Constituents (Except Cymene and Pinenes) of cinerea, Macarthuri and Hybrids.—Continued.

Fluo-
Geranyl Eudes- Sesqui- rescent
— No. Acetate. | Geraniol. Cineole. | Eudesmol. mene. terpene. Com-
ponent.
1 x 3 38 x x SS x KO
3 — _— x x SESE x 36 3K 3K
4 x _ — x 5k BS OK x x x SKK
5 x Xx x _ x 3K »< 3K <x x SK OK
8 = x x x xX XX xX XX x XX
11 = x x x x XX xX X X 36
12 = - <x x xX XX xX XX x XX
13 _— x <x S< 3K ax «XxX K 3K RK
14 32 34 _ xk OK x BS 3< 3K x 3K —
15 _ Dx 3K MK x SS x Xx x Ke OK
16 x 3K = — x 3K x 3K x 3X paex
17 xox x x x 3K x XK x Xx 3 3K OK
18 = x x x XX x xX x XxX
Hybrids. 21 _ x 3K x x xxx x SK KX
22 _— =— x x x x xox _
23 x 3K «KX — 34 3K OK x x — KKK
24 x x Xo KK x x6 OK x 3K OK x OX
25 Not recorded x ORS x KK 36K
26 x ax x 3K OK x6 3K 3K 3K ex x de
27 x x 3K XK x 3 x a <e
29 — x xe x KK KX xx
32 _ x x SKK x x XK SKK OK x SE
33 XK OK x x 3 3K S< x ae 363K SE
34 oS 3K SK _ KK 4 34 x
35 _ ax x x x6 Sax XK x 3K
36 — x aex >aex 36 36 OK x eX ox
38 — - dK OK OK 32K xX xX 34K

#. Macarthuri oils, is present but in still smaller quantities in only 25% of the
E. cinerea parent oils and in 60% of the hybrids. In 22% of the hybrids it is present
in greater quantity than in any of the parent H. Macarthuri oils.

The sesquiterpene alcohol, eudesmol, which is present to the extent of about 15%
in all the parent E. Macarthuri oils, is largely absent from the EZ. cinerea parents but.
is present in all the hybrids, sometimes in quantities as large as the EH. Macarthurt
parents. Eudesmene, though present, in small amounts in all the oils of the
EH. Macarthuri parents, is ‘present at the most as a trace in the E. cinerea parent but
present in all the hybrids, over 50% showing considerable amounts.

The unidentified. sesquiterpene is present in almost all oils. It is in greater
amount in Z#. cinerea than in E. Macarthuri and still greater than either in the hybrids.
The fluorescent compound is present in about half the oils of both parents, whilst
in the hybrids 93% contain larger individual amounts than either of the parents.

With regard to the specific gravity, refractive index and optical rotation of the
oils, the parents show relative uniformity within themselves, but the hybrids display
wide variation in these properties. In this respect hybrid 29 with its low specific
gravity and high laevo-rotation’is of interest. This could be due to the presence of
substantial. quantities of l-pinene but was not further investigated. The means of the
physico-chemical constants of the hybrids lie between those of the parents excepting
in the case of the refractive index (Fig. 4). This is explained by the larger
quantities of sesquiterpene present in the hybrid oils.

It is clear from these data that recombination of oil constituents is a feature of
the hybrids of this cross. Gershtein (1951) studied the oils of a number of Eucalypts
and their hybrids. The only chemical constituent assessed was cineole which did
not vary greatly in parents or hybrids. The physico-chemical constants published by
him, however, indicate that the hybrids contain similar constituents to those found.
in the parents. Mirov (1932) in Pinus turpentines has described a P. ponderosa x P.

60 INHERITANCE OF OIL CHARACTERS IN EUCALYPTUS,

i

&

NY

: i

a S

g 8

:

Ss N
RY
~“
:

' Vein Angle b Yein Angle
2a 2 é
£, paucttlora - £.aves £.peucthora ~ &.Robertsoni
e o e 6

Fig. 2a.—Intermediate position of the hybrids in yield in relation to vein angle is
illustrated, although there is a preponderance in the progeny of individuals which are
identical with the #. pauciflora parent and none actually quite reaches the range of the
E. dives parent.

Fig. 2b.—E. pauciflora x E. Robertsoni. Hybrids are indicated by x. Again many of
the hybrid individuals are intermediate, but some are identical with either parent. It will
be noticed, however, that the recombinants occur in one way only, an individual with a
low vein angle sometimes has a high oil yield instead of the normal low yield characteristic
of the HE. pauciflora parent, but the reverse, i.e. a high vein angle and low oil yield, does
not appear in the material examined.

elective dex

Spectlic gravity

S S
“3 = ® 3 J
Lop. +18. of Leak x x
3 £.Matdent - £. rubids g g S
: . y 8.
x = Hybrid

Fig. 3.—The Fi hybrid H. Maideni x EH. rubida is compared with the “pure” parents in
oil yield. It is clear that the F1’s are entirely in this respect identical with the H. Maideni
parent.

Fig. 4.—The means of various physico-chemical characters of the H. Macarthuri x E.
cinerea oils and the hybrids. Three separate conditions are illustrated and in yield the
hybrids are almost identical with the low-yielding parent. In specific gravity and leaf shape
the hybrids are approximately intermediate between the parents. In refractive index and
optical rotation the characters of the hybrids taken as a group considerably exceed those
of either of the parents.

BY L. D. PRYOR AND L. H. BRYANT. 61

Jeffreyi hybrid containing terpenes inherited from the ponderosa parent and heptane
from the Jeffreyi parent. Work by Snegirev (1936) on Fl and F2 hybrids of two
species of Ocimum shows a similar situation in inheritance of oil constituents in the
Fl hybrids. However, F2 hybrids in this case on the basis of specific gravity,
refractive index and optical rotation data he considered showed “that the composition
of the oil in these hybrids had undergone profound changes’’.

49/102

pueda

180 166 193 191 190 188 182

49/203

279 (

224 204 219 2i.
49/201
254
19
49/806
222

Fig. 5.—Typical examples of the leaf venation of the parents are shown: No. 254,
#H. pauciflora; No. 279, H. dives; and No. 119, EH. Robertsoni. All are intermediate leaves
taken from trees of the same age on about the third pair of leaves on a shoot about four
feet from the ground. No. 49/102 indicates the range of venation pattern in the hybrid
progeny, H. paucifiora x H. Robertsoni, and No. 49/203 in H#. pauciflora x EH. dives.

5

205

Although no evidence of new compounds (i.e. compounds not present in the
parents) was found in the oils of the hybrids, H. cinerea x EH. Macarthuri, the
considerable increase in the content of eudesmene, unidentified sesquiterpene and
fluorescent component is of interest, since they exceed substantially the amounts of
these materials present in either parent.

#. pauciflora x E. Robertsoni and EL. paucifiora x H. dives.

No analysis has been made of the oil components in these combinations, but they
afford a good opportunity to examine the inheritance of oil yield in relation to
morphological characters. Figure 2 shows the leaf character which has been compared
with oil yield, and the various yields obtained with the assessment of the vein angle

62 INHERITANCE OF OTL CHARACTERS IN EUCALYPTUS,

of the different individuals (Fig. 5). In the progenies 49/104 and 49/102 there is
very distinct segregation, and a wide range of variation between the two parents, and
the same is true of 49/203 in relation to H. dives. Unlike the previous hybrid
combination the character, oil yield, segregates and recombines freely. In each hybrid
progeny there are some individuals of the same order as either parent with a full
range of intermediates. There is also distinct recombination between the character
of leaf vein angle and oil yield, as illustrated by Figure 2.

TABLE 3.
Component
Outer Just in
: Fluorescent Sesqui- Inner Component Front of
Sample. Cineole. Eudesmol. | Component. terpene Sesqui- Just Behind Inner
( Eudesmene. terpene. Eudesmol. Sesqui-
: 3 terpene.
=: 2 (Sa ial |; ee ae fem a aT
MXM Self 267 | xxxx [<i = x x x x
MxM BO || 8M SEK SK «6 il — x x x x
MxM 302 xX XX X xX X = x x x ox
MxM 303 xx XX x Xx = x x x x
MxM 304 xX XX XOX _ x x x x
Mx R (F1) 297 KK OK x — x x x x
MxR . 298 x KK IK — x x x x
MxR 299 SoS OS 3S x — Xx x x x
MxR 300 xX X X X = x x xX x x
MxR 264 x XxX = x x x x
RxR 309 xa <x <4 x 3K KK x 3< Daas
RxR 310 x XX <x ax x x< ax x
RxR 313 xX Xx eax X< SEK 3K <x x
RxR 311 x XX x x x X al. oo XX X . *& x
RxR 312 x xX xX x x a oe “KK ees x

Note.—All oils examined gave fluorescence in P. cymene zone of similar intensity. This component would be
expected to be present. Pinene is probably also present in all species, but its Rf being similar to that of the outer
sesquiterpene (eudesmene) results in cortfusion with this.component on reacting with conc. H.SQ,.

M=Maideni. R=rubida. if ye
F1 Hybrids. We \ /

The hybrid studied principally in relation to the parents is H. Maideni x E. rubida.
The oil constituents in the’ parénts are rather similar. Quantitatively, however,
EH. rubida is a very low yielding species giving about 0:1% oil, whereas H. Maideni
is reasonably high, giving about 1:-4% oil. The various components of the oil have
been assessed chromatographically, as shown in Table 3. The most striking differences
exist in the “fluorescent component”, the sesquiterpene and the component “near
eudesmol’’. In the case of the fluorescent component and the component near eudesmol,
the Fl hybrid is like the #. Maideni parent. In the case of the sesquiterpene, it
appears to be intermediate, although the data are not fully consistent. There are no
differences between the cineol and eudesmol contents in the parents and none in the
hybrid. A similar examination has been made of FH. cinerea x EH. Blakelyi F1 hybrids,
but the differences between the parents are not sufficiently striking to derive any
conclusions from the limited material available. In the H. Maideni x EH. rubida the
same kind of inheritance pattern is displayed with regard to yield as in the E. cinerea
x H. Macarthuri combination, but on this occasion it is reversed, and in the Fl the
yield is identical with that of the high yielding parent (Fig. 3). Gershtein’s (1951)
figures for the Fl EH. Maideni x EH. viminalis show a similar position. At the same
time there is a tendency for the constitution of the H. Maideni parent to be followed
in the oil combinations; this suggests that the determinants of the yield and certain
of the constituents derived from H#. Maideni in this F1 hybrid are displaying dominance,
perhaps like the determinant of the red pigment of the Rutgers tomato (Tomes ef dl.,
1952).

BY L. D. PRYOR AND L. H. BRYANT. 63

Discussion.

While the material available does not permit an exhaustive analysis of the
pattern of inheritance, it indicates some interesting aspects. It seems that the yield
may be determined by one parent in a very far-reaching way, as is shown by the
E. cinerea x HE. Macarthuri segregating populations and also in the #. Maideni x EH.
rubida F1 hybrid. The same may be true of some of the different components, as is
indicated in the fluorescent component in the H. Maideni x H. rubida F1. On the other
hand, in some combinations, as the #. paucifiora x H. Robertsoni and E. paucifiora x EL.
dives, there is very distinct segregation and recombination of yield, as there is also
for geranyl acetate in the H. Macarthuri x E. cinerea combination. In the EH. cinerea
x H. Macarthuri and the #H. pauciflora x H. Robertsoni and EL. pauciflora x EH. dives
there is evidence of recombination of various characters. In the first place it is
between oil constituent and leaf shape, and in the second place between yield and
angle of leaf vein. The morphological characters within these combinations have
been selected for ease of measurement. There are several characters less easy to
measure, as for example leaf thickness in the H. pauciflora x LE. Robertsoni combination,
which by inspection seem to follow the same pattern.

A characteristic of segregating progeny derived from hybrids is that the variance
of both morphological and oil characters is much greater than that of the parent
species. If the facts disclosed so far have a more general application, they suggest
that a recombinant between high yield and desirable oil constituents may be found
if suitable segregating populations are produced.

The pattern of variation assessed by Willis et al. (1951) in the progeny from
presumed forms of E#. maculata shows close analogy with the segregating hybrid
populations described above. It is suggested that the origin of these variants is by
hybridization between #H. citriodora and E. maculata. The most closely occurring
H. citriodora trees, which in this case could be one parent, are believed to be about
40 miles distant from the above forms which occur in a H. maculata stand. Isolated
hybrid individuals have been found at this and greater distances and may arise either
from long-range outerossing, for example by birds, or as the result of species
population movements associated with climatic change. it is not difficult to imagine
the origin of such individuals if they are hybrid, and it is suggested, therefore, that
this is the explanation for them.

Conclusions.

It is clear that oil characters, both in yield and make-up, are strongly inherited
in Hucalyptus, although the determination of the precise pattern will need a good
deal more study. So long as evidence is still lacking that Hucalyptus oil is either
an attractant: or repellent to insects the function of oil in this regard can only be
presumed. From the point of view of the Hucalyptus oil industry, three main lines
for developing improved forms are clearly indicated. First and most obviously is
the raising of plants from seed from selected stands with known desirable oil
characters, as already initiated by Willis; secondly, raising segregating populations
from selected naturally-occurring F1 hybrids with a view to obtaining superior
recombinants; thirdly, producing experimentally F1 hybrids between parents which
have separate desirable characters which, if combined together, would give individuals
superior to either parent. It is of particular interest that in some combinations the
oils display characters which transcend in magnitude those of either of the parents.
This suggests that the biochemical processes resulting in oil formation may be, as in
other plants, subject to separate genetic control at each of a number of steps, and
that at times in hybrids some steps may be characteristic of one parent leading to the
formation of a precursor which is then subject to genetically determined influences
derived from the other parent somewhat in the manner suggested for the carotenoid
pigment system of tomato (Tomes et al., 1952). Such a process could lead to the
formation of compounds in greater quantity than those present in either parent species
or even to compounds present in neither. Very much more study and experiment would
be necessary to predict reasonably when and how this sort of change would occur.

64 INHERITANCE OF OIL CHARACTERS IN EUCALYPTUS.

However, the likelihood that it is present offers very considerable possibilities in a
breeding programme for improved oil production.

It suggests considerable flexibility in evolutionary aspects in the genus if oil is
significant in this regard.

Acknowledgements.
The generous provision of facilities to make the initial distillations by Professor
A. H. Ennor, of the John Curtin School of Medical Research, Australian National
University, is greatly appreciated, and sincere thanks are due to Mr. O. Ruzicka for
preparing the drawings.

References. '

BAKER, R. T., and SmiTH, H. G., 1920.—A Research on the Eucalypts especialiy in regard to
their Essential Oils. Sydney, Government Printer.

BARBER, H. N., 1955.—The Natural History of Natural Selection. Awust. Jowr. Sci., A.N.Z.A.A.S.
Report, 3, Section M.

Bryant, L. H., 1950.—Technical Notes, For. Comm. N.S.W., Div. of Wood Technology, Vol. 4
(Special Issue): 6-10.

—, 1955.—Circular Chromatecgraphy of Terpenes on Absorbent Coated Giass. Nature,
WHS HSO-

DETHIER, V. G., 1941.—Chemical Factors Determining the Choice of Food Plants by Papilio
Larvae. Amer. Nat., 75: 61-73.

GERSHTBIN, L. A., 1951.—Characteristics of Ethereal Oil in some Eucalyptus Hybrids.
Biochemistry, U.S.S.R., 937-940.

' JAMES, W. O., 1953.—Alkaloids in Plants. Endeavour, 12: 76.

Mirov, N. T., 1932.—A Note on Jeffrey and Western Yellow Pine. Jour. For., 30: 93-94.

PENFOLD, A. R., and Morrison, F. R., 1927.—The Occurrence cf a Number of Varieties of
E. dives as determined by Chemical Analyses of the Hssential Oils, Part I. Jour. & Proc.
Roy. Soc. N.S.W., 61: 54-67.

PENFOLD, A. R., McKrRN, H. H. G., Wiuuis, J., 1953.—Studies of the Physiological Forms
of the Myrtaceae, Part 6, An Examination of the Progeny obtained from #H. citriodora
Hook, var. A.

Pryor, L. D., 1953.—Variable Resistance to Leaf-eating Insects in some Eucalypts. Proce.
LINN. Soc. N.S.W., Vol. 77, Parts 5-6: 364-368.

SNEGIREV, D. P., 1936.—Bull. Applied Bot. Gen. Pl. Breeding, U.S.S.R., I11: 15.

Tomes, M. L., QUACKENBUSH, F. W., NELSON, O. E., JR., and Betty NortTH, 1952.—The
Inheritance of Carotenoid Pigment Systems in the Tomato. Genetics, Vol. 38, No. 2:
117-127.

Wiis, J., 1951.—The Jnheritance of Oil Characters in H. dives. M.Se. Thesis, Sydney
University.

65

THE STATUS OF NITROGEN IN THE HAWKESBURY SANDSTONE SOILS AND
THEIR PLANT COMMUNITIHS IN THE SYDNEY DISTRICT. II.

THE DISTRIBUTION AND CIRCULATION OF NITROGEN.
By Nora J. HANNON, Linnean Macleay Fellow of the Society in Botany.
(Two Text-figures.)
[Read 30th April, 1958.]*

Synopsis.

The nitrogen content of a typical Hawkesbury Sandstone ecosystem is estimated to be
approximately 2,400 lb. per acre. It is fractionated in the rooting medium, living plant tissue
and plant litter in the ratio of 200: 35:1. On the basis of estimates of nitrogen release:
from the soil organic matter complex and measurements of litter returns (700 lb. per acre
per annum), it appears that an annual cycle of 2-3 lb. nitrogen per acre occurs. These:
values for Hucalyptus forest on Hawkesbury Sandstone are shown to be low in comparison
with other types of forest. The litter accumulated on the forest floor occupies only 0:5%
of the nitrogen capital of the sandstone ecosystem and proportionately greater amounts of
nitrogen are contained in living plant tissue than are found in pine forests.

INTRODUCTION.

An earlier paper (Hannon, 1956) described the concentration values of nitrogen in
the parent rock, the soil and the plant material of the naturally occurring communities
on Hawkesbury Sandstone in the Sydney district. In the present instance, an attempt
will be made to convert these concentration values into absolute terms to allow the
estimation and fractionation of the nitrogen capital of the ecosystem. This conversion
is necessary to provide a concrete expression of the nitrogen content of the communities
and to allow comparison with other ecosystems.

Hstimates of this nature are relatively simple in pot trials where the initial
quantities of the various component fractions may be calculated, the growth rate
accurately measured, the losses accounted for and final computations compared with
the initial. Even in field trials, a certain acreage is sown uniformly with a single
variety at a given time. The sandstone communities, however, are very complex.
Although heaths and scrubs of long-lived shrubs and relatively uniform structure do
occur, the most widely spread formation is the Hucalyptus forest. Like the scrubs, the
forests are characterized by a great variety in their species composition, but possess
both shrub and tree layers. These communities are composed of individuals of
different physiological age and of diverse genetic constitution. Stands of the same
formation frequently display appreciable variation, especially in relation to aspect
‘effects or the stage of regeneration from fire. Difficulties, which will be mentioned
later, also apply to the study of the rooting medium and the litter fraction. All of
these factors complicate a study of nutrient distribution, which, to be complete,
requires detailed and quantitative analyses of typical stands. In the absence of these
observations, only approximations and estimates can be made. These will, however,
provide useful information.

THE DISTRIBUTION OF NITROGEN.

The physiography of the sandstone-covered areas allows a division into three
major ecological niches—the undissected portions of the plateau surface, the gully
slopes, and the gully floor. The plateau surface typically bears heath, scrub, tree-scrub
or low scrub forest. These are the least luxuriant of the sandstone formations and are
the communities in which exposure effects play a significant role. At the other extreme,
the tallest forests, containing a more mesomorphic understorey, are found only on the

* Manuscript received 18th November, 1957.

PROCEEDINGS OF THE LINNEAN Soctety or New SoutH WaAtgEs, 1958, Vol. Ixxxiii, Part 1.

66 NITROGEN IN HAWKESBURY SANDSTONE SOILS AND THEIR PLANT COMMUNITIES, II,

gully floor. Here not only are exposure effects eliminated, but plant nutrient and water
supply are increased by accumulation from the surrounding ridges and slopes. In
both of these niches the soil cover is better developed than on the slopes—on the
plateau, because erosive forces have not been so active, and on the gully floor because
accumulation has occurred from topographically higher areas.

It is thus obvious that, on the plateau, exposure effects complicate the existence
of a direct relationship between plant development and soil nutrient levels. When the
distribution of nutrients is under consideration, it is logical to select an example
where soil nutrient supply rather than any other factor of the environment is limiting
plant development. On the other hand, the gully floors occupy a very small percentage
of the total area where sandstone outcrops avd cannot therefore be taken as typical
of the sandstone areas as a whole.

The gully slopes are usually occupied by tall scrub forest. This is a very widely
spread formation occurring chiefly on slopes where conditions of shelter and soil
moisture are favourable. It is restricted to the lower portions of the sunny-north-facing
slopes, but may extend to the top of the gully on south-facing slopes. It also occupies
the restricted areas on the upland plateau where exposure is not too rigorous.

The plateau, composed chiefly of Hawkesbury Sandstone, which encircles the Sydney
Plains, is very dissected, particularly to the west and north of Sydney. To the south,
on the Nepean Ramp, dissection is not so marked as in the other areas, but occurs
extensively. The physiography thus presents areas of very considerable size which
are topographically suited to tall scrub forest development.

An illustration of the distribution of nitrogen amongst the three major divisions
of the ecosystem—the rooting medium, the undecomposed plant litter and the plants—
will therefore be given by considering a hypothetical stand of the poorer quality of
tall scrub forest formation. This stand would be described as “average” with respect
to its plant and litter cover, its soil depth and the nitrogen concentrations of its
mineral and organic components. No information is available to permit the inclusion
of the animal life of these communities.

Methods of Estimation.

Part of the data used in making this calculation has been previously presented
(Hannon, 1956—Tables 3-8, 14-18). Part is given in Tables i-iii of the Appendix
(pp. 83-84, below) and details concerning the estimation of soil and plant mass will now
be discussed.

A visual method of weight estimation of plant cover was employed, and attention
was confined mainly to the shoot systems. After estimation, shrub species were
frequently cut at ground level and the estimates checked on a _ spring-balance.
Experience in the estimation of tree mass was gained in areas of private property
where the trees were being felled and weighed to be sold for pulp and firewood.
Corrections were applied to convert these values to an oven-dry basis.

The figure for litter mass was obtained by direct measurement of the total litter
collected from a number of areas. A correction was applied to these values to allow
for the fact that outcropping rock surfaces, tree bases and shrub cover prevent the
accumulation of litter over the entire surface in these communities.

Soil depth, the occurrence of rock floaters and the rooting habits of the plant
cover have been examined in excavation works made by the Metropolitan Water,
Sewerage and Drainage Board in areas of Hawkesbury Sandstone. Even on the
plateau surface, soil depth generally does not exceed 24”-30” and rock floaters
commonly occur, mixed with the soil, above the underlying layer of almost solid rock.
The soils on the gully slopes are very skeletal. Rock boulders outcrop over most of the
surface and, apart from very isolated pockets which may extend to 5-6 feet, soil
depth is very shallow. Usually pockets barely exceed 12”. Inspection at the ground
surface level and by using an iron bar to test soil depth indicates that soil occupies
only a very small percentage of the rooting medium. However, when large areas
have been excavated with the assistance of heavy drilling and blasting equipment, it
is obvious that greater amounts of soil are to be found underlying the floaters than

BY NOLA J. HANNON. 67

casual inspection would have suggested. Close examination of this material, however,
shows that small rock fragments and pulverized rock which have originated in the
drilling and blasting operations are contributing to this fraction. The soil cover is
variable from one patch to another, but usually occupies no more than 40% of the
rooting medium volume.

The rooting habits of the trees occupying these areas are remarkable. Main tap
roots are not found and usually a division into three or four major roots occurs at the
stem base. These spread laterally along the rock surface to a point where jointing
occurs. The roots penetrate downwards through fissures—instances have been observed
where the root is flattened to only half an inch in thickness and is of a width of
12 inches to allow its passage through the rock layers. The interweaving of the roots
through the rock crevices must provide a very firm anchorage. It is obvious that
penetration by tree roots freyuently is a factor of importance in the fracture of the
sandstone. The great bulk of the feeding root zone is found in the surface layers but
a few large roots may extend to depths of up to ten feet or more to allow for
Support and the absorption of water. For the present purpose, a soil depth of 20 inches
will be chosen, since only occasional roots are found beyond this depth and their
absorptive surfaces are restricted to their extremities; in addition, soil nitrogen levels
are very low below this depth. The region occupied by the feeding roots is of the
greatest relevance to the present consideration of the soil—plant—litter system.

On the basis of these measurements and observations, the nitrogen capital of a
typical stand of tall scrub forest on Hawkesbury Sandstone has been estimated as
follews:

Calculation of the Nitrogen Content of a Hawkesbury Sandstone Ecosystem.
1. The Rooting Medium (60% rock+40% soil).

Volume per Nitrogen Nitrogen
Depth of Sample. Acre Surface. Density. Mass. Concentration. Content.
(in.) : (cu. ft.) (Ib./cu. ft.) (Ib.) (p.p.m.) (Ib.)
Rock :
0-20 a 3 43,560 130-8 5,697,648 180 1025-6
‘Soil :
O- 8 a a 11,616 87-2 1,012,915 500 506-5
8-20 ae ar 17,424 93-5 1,629,144 300 488-7
Rock +soil of ee — — — — 2021
‘2. The Latter Fraction.
Mass per Nitrogen Nitrogen
Fraction. - Acre Surface. Concentration. Content.
(Ib.) (p.p.m.) (Ib.)
Total litter 5,600 2,000 11
3. The Plant Cover.
Mass per acre of plant material: 70 tons.
Composition: 5% living shoot and root tissue.
95% woody tissue.
Mass per Nitrogen Nitrogen
Fraction. Acre Surface. Concentration. Content.
(Ib.) (p.p.m.) (Ib.)
Living tissue 7,840 7,000 54-9
Woody tissue 148,960 2,000 297-9
Total 156,800 =

353,

Total nitrogen content of ecosystem =2,385 lb./acre.

The calculation shows that in this stand the nitrogen capital approximates 2,400 lb.
‘per acre and that the proportion of the nitrogen contained in the rock + soil, plants
and litter is in the ratio of approximately 200:35:1. Even though the presence of
the mineral fraction reduces the ,concentration of nitrogen in the rock and soil to a
level far lower than that of the organic material of the plant and litter, the very
great mass of rock and soil in comparison to the other fractions renders it the
chief reserve of nitrogen (and other nutrients) in these communities.

68 NITROGEN IN HAWKESBURY SANDSTONE SOILS AND THEIR PLANT COMMUNITIES, II,

THE CIRCULATION OF NITROGEN.

The foregoing fractionation illustrates the static distribution of nitrogen in the
ecosystem. The dynamic aspect—that of the interchange of nitrogen between the
three major fractions of the rooting medium, plants and litter — requires consideration
also. This cycle may be complicated by internal cycles within each of the three
fractions, e.g. the withdrawal of nutrients into a plant from senescent leaves, but the
fundamental principles of nutrient circulation may be summarized in broad outline
by the following items:

1. The release of nitrogen from the soil organic matter complex resulting in the
formation of an “available” form of nitrogen.

2. The absorption, translocation and synthesis of available nitrogen by plants,.
resulting in plant growth.

3. The maturity and senescence of plant organs, resulting in litter formation.

4. The partial decomposition of litter, resulting in its incorporation into the soil.
organic matter complex.

Little detailed information is available on these aspects of nutrient circulation in
Hawkesbury Sandstone communities, but certain data have been collected and will
permit discussion in generalized terms.

The Release of Nitrogen from the Soil Organic Matter Complex.
Discussion of Data.

On the basis of data presented previously (Hannon, 1956—Table 13), it may be
calculated that over a period of one year available nitrogen is released from the sand-
stone soil organic matter reserve in a concentration of approximately 100 parts per
million of soil. This value was obtained by bio-assay under conditions where
temperature, water and the nitrogen-free mineral supply were close to optimal. After
correction for growth due to the seed reserves, plant growth showed an approximate:
three-fold increase in the presence of a nitrogen-free mineral solution over that where:
only water had been added. This may be used as an approximate indication of the
extent to which the mineral supply stimulated the nitrification rate. Even this latter
value should be reduced since, under field conditions, water supply is very spasmodic
and must limit the activity of the microorganisms engaged in organic matter
conversions. Under field conditions an annual release of nitrogen in available form.
in a concentration of approximately 25 p.p.m. of soil might thus be anticipated.

Field observations show that a plant whose shoot system occupies one square metre:
of surface may be assumed to bear a root system, the extremities of which penetrate:
a volume of 3 x 3 x 0:25 cubic metres of soil and rock. With an apparent density of
1-4 g. per c.c., the 40% of this volume occupied by soil would weigh 1,260,000 g. On
the basis of the above-mentioned assumptions, from 1,260,000 g., 31:5 g. available
nitrogen would be released each year.

However, the propertion of soil within the absorptive range of the roots will be.
of greater relevance than the volume through which the roots extend. In addition,
the form in which the nitrogen becomes available, particularly whether it is freely
mobile as is nitrate or adsorbed as is ammonia, must be considered. The sparsity of
nitrate in the soils other than in swamps, and the consistent presence of ammonia
would indicate that the latter is the more likely form. Furthermore, the possibility
cannot be overlooked that forms of nitrogen and of other nutrients not considered
“available” for the growth of mesomorphic species may be acceptable to sclerophyllous.
vegetation under natural conditions when more easily assimilable sources are not
available. Since absorption of nutrients occurs only in regions close to the actively
growing root tips which, in woody perennial species, form a small percentage of the
total root mass, it is very probable that absorption may occur from only 1% of the
total volume penetrated by the roots. The volume,through which the roots pass is
shared with the root systems of adjoining shrubs. Certain quantities of available
nitrogen would also be required by soil-inhabiting microorganisms and some may be:
lost from the ecosystem by various factors which will be discussed later.

BY NOLA J. HANNON. 69

In view of these considerations, 0:315 g. available nitrogen may be absorbed by
the shrub during one year. Knowledge of the concentration of nitrogen in plant
tissue enables the calculation of the mass of tissue produced by the incorporation of
this quantity of nitrogen into new growth. On the basis of an average value of
4,000 p.p.m. nitrogen for the tissues of the root, stem and leaf, a mass of approximately
79 g. per square metre of tissue could be developed from 0-315 g. of available nitrogen.
On the basis of this figure, an amount of 700 lk. per acre (0:31 ton per acre) is
suggested as the annual production of organic matter. An unknown fraction of this
quantity would be diverted to the extension of the root system. Jenny (1941) quotes
the work of German investigators who have recorded that in forests the weight of the
underground mass of vegetation is from 20% to 30% of the total mass; 10% is often
quoted in the literature. On the basis of the very few studies where roots were
sampled, Rennie (1955) notes that of the total nutrients taken up by trees, 13% was
the average quantity present in the roots.

TABLE 1.
(Taken from Jenny, 1941.*)

Organic Matter Production by Vegetation.
(Roots not included unless stated.)

Annual
Type of Vegetation. Production. Remarks.
(Tons/acre.)
Alpine meadows a if <0-22-0-40 Clipped quadrats, Poland, air-dry (includes
: moss, lichens).
Short-grass prairie bes ae 0-71 Clipped quadrats, Colorado, air-dry.
Mixed tall-grass prairie ae 2-22 Clipped quadrats, Nebraska, air-dry.
Tall-grass prairies 2s ae 0-85-1-73 Annual prairie hay figures, Middle West.
Average forest (leaves, wood,
twigs) .. 5 5 aa 2-67 German forests, air-dry.
Beech :
ee < * a fay Central Europe, air-dry.
Pine:
ee x je ae Central Europe, air-dry.
White-pine needles ae Re 2-09 Connecticut, 1934, oven-dry.
Beech—birch—aspen forest 2-85 Leaf litter only, Germany, air-dry.
Tropical primeval forest (leaves, :
trunks, roots) .. aa a 11:1 Java, estimate, moist.
Tropical legumes are we 24-4 Dutch East Indies, moist.
Tropical savannas ae a 13:3 Estimate, moist ?
Monsoon forest .. Ne sie 22-2 Estimate, moist ?
Tropical rain forest eee it 45-90 Estimate, moist ?
Eucalyptus scrub forest me 0-31 Estimate, oven-dry, roots included.

—

* An estimate for a Hawkesbury Sandstone community is included for comparison.

Plant cover is not uniform or even continuous in the sandstone communities. It
is very probable, however, that the areal extent of bare ground is counterbalanced by
the interlacing of the shoot systems. This and the other limitations of this calculation
are realized; many assumptions have had to be made. Nevertheless, each of the
assumptions has been carefully considered in light of the known facts and observations.
Pending fuller investigation of nutrient—and in particular, nitrogen — circulation
in these communities, the calculation is of value in setting an approximate level of
the quantities involved, and emphasizing the gaps in our present state of knowledge.
It is of interest to compare this estimate with the figures compiled by Jenny (1941)
of the annual organic matter production by varying types of vegetative cover. These
values are shown in Table 1. It is seen that organic matter production in Hawkesbury
Sandstone communities is only small by comparison on these standards.

70 NITROGEN IN HAWKESBURY SANDSTONE SOILS AND THEIR PLANT COMMUNITIES, II,

Plant Growth Rate.

Observations of the growth rate of sandstone species are restricted to a few
measurements of shoot expansion in the shrub stratum over a period of one year.
Variation in the expansion of different shoots on the same individual and between
different individuals of the same species and similar age was very marked. Con-
sequently these values do not form a satisfactery basis for a measurement or an
estimate of the annual production of plant tissue, especially since no attention has
been paid to the dominants of the widespread forest communities—Eucalyptus and
Angophora trees.

TABLE 2.
Litter Fall in Forest Communities on Hawkesbury Sandstone.
Collections made in low scrub forest, Warrah Sanctuary.

Total Weight
Period and Dry Weight of Litter Total Area Total Leaf Total Total
Season of of Litter at at Hach of Sample. Litter. Eucalyptus Angophora
Sampling. Each Site. Collection. Leaf Litter. Leaf Litter.
(Month.) (g./sq. yd.) (g.) (sq. yd.) (g.) (g.) (g.)
(a) 19:5
4 (b) 24-0 2) Bey” 58-5 4 33:0 13-7 19-3
(July— (ec) 1:6
November) (d) 13-2 co
(a) 10-48
2 (6) 7:37 23-5 4 21-2 11:9 9-3
(November— (C) eit
January) EM tie OB ihar 4-07
Gey agaaay) 13-46
3 (6) 9-02 33-1 4 20-5 10-9 9-6
(January- (c) 6:45
April) Wie): Ree ee 4-21
AGO. 6-90
4 (b) 6°33 24-0 4 10-8 4-6 6:2
(April— (ec) 1:80
August) LOMO 8-97
ieee ee 8-15
(6) 3°73
1 (c) 14-48 37-2 6 30:9 6-1 24-8
(August-— (d) 0-91
September) (e) 4:15
Gp) He's
1 (e) 15-0
@ontiontner (f) 20-4 eS 35°5 2 26°8 10:8 16-0
October)

Litter Fall.

In a community which is in equilibrium with its ehvironment, the amount of
growth produced per annum must be approximately equivalent to the annual litter
fall. On theoretical grounds a net gain or net loss does not occur in a mature
community. Consideration of this fact somewhat reduces the shortcomings of the
foregoing data on organic matter production in these communities, both in the
theoretical calculation and in the measurements of the expansion of the shoot systems.
under field conditions.

Presentation of Data.

Litter collections have therefore been made in sandstone communities and the
results are presented in Tables 2, 3 and 4.

BY NOLA J. HANNON. ale

Hach site was chosen as being typical of the structure and floristics of the stand,
but variation in the litter collections from plot to plot is very marked; this may
reflect the influence of the direction of the prevailing winds. However, the data
clearly demonstrate that the amount of litter falling in these Hucalyptus forests is
uniformly low. Similar results were obtained with all the types of collecting agencies
used — wooden trays with a fine wire base, supported two inches above ground level,
hessian pegged flat onto the ground surface, and the ground surface cleared of litter.

The large-leaved tree species — Eucalyptus and Angophora— provide practically
all of the litter in the form of leaves and twigs. The contribution from shrubs in
areas other than directly beneath them is negligible. Apart from certain of the larger
shrubs, only seldom is any accumulation of leaf litter found at their base. The micro-
phyllous leaves of most shrub species would be difficult to detect when scattered
amongst the general litter accumulated on the ground surface, and were not found

TABLE 3.
Intier Fall in Forest Communities on Hawkesbury Sandstone.
Collections made in tall scrub forest, Kuring-gai Chase.

: Total Weight
Period and Dry Weight of Litter Total Area Total Leaf Total Total
Season of of Litter at at Hach of Sample. Litter. Eucalyptus Angophora
Sampling. Hach Site. Collection. Leaf Litter. Leaf Litter.
(Month.) (g./sq. yd.) (g.) (sq. yd.) (g.) (g.) (g.)
(a) 7-6
2 (6) 1-77 11-5 4 3:6 1:3 2°3
(March—May) (ec) 1:14 (
(d@) 1-01)
(a) 5-87)
2 (6) 3:16 21-3 4 2-9 1-4 1:5
(May—July) (c) |
(d) 8-66
(a) 31-91)
4 (0) 23-87 90-7 4 36-0 3-2 32-8
(Suly— (ec) 15-25
November) (d@) 19-65
(a) 4-37)
4 (6) 6-84 24-2 4A 6-1 1:2 4-9
(November— (ec) 7:19
March) (d) 5:78

in any abundance on hessian placed around the stem base and underlying the canopy
of individual shrubs. ‘The litter collected here consisted mainly of defoliated fine
twigs. The quantities of litter collected beneath shrubs are given in Table 4 and
are seen to be quite appreciable. The values shown in Table 4 are not representative
of the whole shrub stratum, since bulky species, as Banksia serrata and Banksia
ericifolia, while common, are not typical of much of the shrub stratum. JSpacris
pulchella on the other hand is less vigorous than the average shrub cover. On
the basis of the figures shown in Table 4 and acquaintance with the sandstone
communities, an average value of approximately 10-15 g. per sq. yd. (140 Ib. per acre)
for the total yearly fall would be suggested from the shrub layer. This estimate
appears low in light of the values shown in Table 4, where the figures refer to
monthly collections, but inspection of the growth form of many shrub species and
observations of their growth habit indicate that major growth bursts and litter fall
occur but once each year. Evidence to be presented later shows that leaf fall from
trees is at its peak at the season when the shrub litter was collected. A further factor.
which will reduce the values in Table 4 is that the shrub stratum is not continuous.

72 NITROGEN IN HAWKESBURY SANDSTONE SOILS AND THEIR PLANT COMMUNITIES, II,

A considerable proportion of bare space occurs between shrubs, particularly in the
forest formations and, in addition, part of the area is occupied by outcropping rock
and the trunks of the trees.
TABLE 4.
Intter Fall from Shrub Stratum.

Finely woven hessian was pegged onto the ground surface beneath the shrub canopy in a low scrub
forest stand in Warrah Sanctuary.

Period and Total Dry
Composition of Season of Weight of Remarks.
Shrub Stratum. Sampling. Litter.
(month.) (g./sq. yd.)
1 13-72
(July—Aug.)
Banksia serrata 10 ft. 1 57-64 Mostly bark and twigs from
tall ; Boronia ledifolia ; (Aug.—Sept.) B. serrata.
Dillwynia ericifolia ; 1 30-60 Mostly bark and twigs from
Tsopogon anemonifolius (Sept.—Oct.) B. serrata.
1 8-39
(Oct.—Nov.)
1 5-65
(July—Aug.)
5 a et 1 27-86
STU erie OM: Eon (Aug.—Sept.) ] Almost entirely twigs from
teretifolia, 3 ft. 6 in. See
inten, 1 10-06 B. ericifolia.
(Sept.—Oct.)
1 2-84
(Oct.-Nov.)
1 0:07
(July—Aug.)
Epacris pulchella cluster 1 1:14 |
of bushes averaging (Aug.—Sept.) ( Fine twigs bearing few leaves.
30 in. 1 3°64
(Sept.—Oct.) J
1 1-00
(Oct.—Nov.)

Inspection of Table 5 suggests that leaf material may form a greater fraction of
the total litter fall in low scrub forest than in the taller forest. However, since the
samples were not collected simultaneously in both stands, it is not possible to reach a

TABLE 5.
Intier Fall in Eucalyptus—Angophora Forest Communities.
Average Litter Litter Composed
Formation. Season of Sampling. Fall. of Leaves.

(g./sq. yd./month.) (%)

July—November 3°65 56°3

November—January 3:00 90-2

Low scrub forest January—April a 2-80 62-0
April-August Me oti 1-50 45-0

August-September 6-20 83-0

September—October 17-30 75°5

March—May 1-50 31:3

Tall scrub forest May-July 2-65 13-6
July—November 5-70 40-0

November—March 1-50 25-0

definite conclusion on this matter. Table 5 does illustrate, however, that litter fall
is never very heavy in these forest communities —an average figure would not exceed
six grammes per square yard per month. The areas which are not available for

-]

(es)

BY NOLA J. HANNON.

tree litter accumulation are the majority of rock surfaces (i.e. all other than those
which are horizontally placed) and the areas occupied by the tree trunks and the
low-growing shrubs. In view of these considerations, only 65-70% of the total area
is available for the accumulation of tree litter. The annual accumulation on an acre
basis would therefore amount to approximately 500 lb.

It is interesting to compare the litter fall from Hucalyptus—Angophora dominants
with that collected in an almost pure stand of Casuarina littoralis forest which also
occurs in localized patches on Hawkesbury Sandstone. Inspection of the morphological
character of Casuarina cladodes suggests that the canopy is gradually replaced during
each year. The quantities of litter collected in a Casuarina stand are shown in Table 6,
where an average fall of approximately 200 grammes per square yard per month is
indicated, at least over a period of seven months.

Table 5 shows that where the litter was collected at monthly intervals from
August to September and particularly from September to October in the low scrub
forest, the litter fall was heavier than at other seasons. The collection of litter at

TABLE 6.
Litter Fall in Casuarina littoralis Stand.
Period of Average Litter
Sampling. Season of Sampling. Litter Fall. Fall.
(Month.) (g./sa. yd.) (g./sq. yd./month.)
1 August-September .. 7" (a) 228) S51)
: ()) 33h
4 September—January .. ste (a) 1560 342
(b) 1176
2 January—March aye aes (a2) 266 185
(b) 473

relatively long and irregular intervals —such as the four-month periods from July to
November in both low and tall scrub forests—is likely to mask the occurrence of
a heavy fall during a relatively short period within that interval. Subsequently,
Turner (1954) made a preliminary and more detailed study over a period of six
months of litter fall in the Sydney district in rainforest and wet sclerophyll forest
occurring on Narrabeen shales and dry sclerophyll forest on Hawkesbury Sandstone.
In all three formations, Turner found that leaf fall was heaviest in October. His
graphs for leaf fall are reproduced (Text-figure 1) and show the small quantities of
litter falling in the sandstone community as contrasted with the others. Turner found
“no appreciable and definite trends” with season in twig falls. His values for the
leaf, twig and total litter fall in the sandstone community studied are given in
Table 7, and show general agreement with those given in Table 5.

In addition to leaf and twig returns, flower, fruit and seed litter must also be
considered. For an accurate estimate, samples need to be collected throughout the year;
but the majority of species flower in August and the fruit is close to maturity in
November. Samples were therefore collected from the living plants in November and
consist of the remains of the flowers and the fruits, some of which had matured and
opened, while others were still green. The results are shown in Table 8.

It is difficult to express this fraction as an annual addition on an area basis,
since many of the shrub species form large woody fruits which may be retained,
even after dehiscence, for several years. Firing is frequently responsible for the
opening of fruit which has long been mature but dormant. Other species form
abundant fruit of a leathery nature. In addition, the seed of different species shows
great variation, but a considerable mass of seed may be formed on both small and
large seeded varieties. A quantity of about 60 lb. per acre per annum is suggested
for the flower, fruit and seed litter from the shrub stratum. The woody fruits of the
trees were found in the litter trays and are included in the leaf and twig litter
fraction.

E

74 NITROGEN IN HAWKESBURY SANDSTONE SOILS AND THEIR PLANT COMMUNITIES, II,

Discussion of Data.
Detailed investigations of litter fall collected over a period of several years are
required before figures for bulk fall and nutrient returns may be recorded with
certainty. Kittredge (1948) gives several examples of variation in annual leaf fall

40

30

a 25 (pl ie

20

moe 2pm

10

g-/s4. 4
/* weeks)

Text-fig. 1—Taken from Turner (1954). Leaf fall (g./sq. yd./4 weeks) in rainforest,
wet sclerophyll and dry sclerophyll formations in the Sydney district.
----: rainforest ; rainforest ;
: dry sclerophyll forest on Hawkesbury Sandstone.
M, J, J, A, S, O, N: period May-November during which samples were collected.

wet sclerophyll forest;

in different years in the same stand and includes an instance in Scotch pine forest
in Germany. where, in the records of seven years’ falls, the maximum fall was three
times the minimum.

TABLE 7.
(Taken from Turner, 1954.)
Intier Fall in Dry Sclerophyll Forest.

Leaf Litter. Twig Litter. Total Litter.
Date of (g./sa. yd.) (g./sa. yd.) (g./sq. yd.)
Sampling. SE —————————————— ee | eee eee
Two Weeks. One Month. Two Weeks. One Month. One Month.
May 28 0-95 ; 4-10 ; ;
June = 1 ace Lek 5-10 f e230 ltd
June 25 0-40 0-55 i }
July 9 0-60 f ee 0:95 f ee 28
July 23 0°85 | Sho 0-65 | : f
August 6 11085 | B20 Pail jf age a0
August 20 0-60 | 1:20 | : ;
September 3 1-10 { ine DOTS Sf a oe oa.
September 17 1-35 0-85 |
October 1 2-30 f ans 3-35 goat RES
October 15 2°90 | 1°65 | ;
October 29 fe 5-05 f CoPe 2-90 f 298 12580
November 12 cy — j — \ : ;
November 26 "2 — } ee — f te’ BeBe
Total =. | 93-05 30-40 53°45

BY NOLA J. HANNON. 75.

Information is not yet available for Hawkesbury Sandstone communities as to the
extent of the effect of wind and rain and other climatic influences on litter fall and on
its nutrient content. Until this is obtained, litter fall results over any short period
must be considered unique, rather than typical, since they are the product of the
interaction of a particular set of environmental conditions on particular individuals.
While bearing this limitation in mind, and noting the agreement of Turner’s data
(Table 7) with that given in Table 5, a figure may be suggested tentatively to represent
the annual litter return in these forest communities. No heavy litter falls occurred
in the pericd of the year over which Turner's collections did not extend (December
to mid-May inclusive) (Table 5), and therefore a value of approximately 700 lb. per
acre would represent the annual litter return from bark, leaf, twig, flower, fruit and
seed from trees and shrubs. This figure includes only the usual litter fractions: it
must be noted, however, that death of shrubs, caused by insect attack, and death of

TABLE 8.
Potential Litter from Flower, Fruit and Seed Sources in Shrub
Stratum.

Samples collected from living plants in November, when most
of the fruit is mature.

©

Blower, Fruit
Formation. and Seed.
(g./sq. yd.)

Low scrub forest mi? Fire vs (6) 2:9

Tall scrub forest - aS % (c) 27-

large limbs of trees is of quite normal, though spasmodic, occurrence and is not
included in the above-mentioned figure. It is of interest to note that the annual litter
fall and organic matter production figures, each of which have been calculated from
different data and observations, coincide exactly (0-31 ton per acre). This agreement
strengthens the possibility that the assumptions approach the actual values; perfect
agreement is not anticipated, since so many factors are involved and therefore no
special significance is attached to it.

Table 5 shows that the mass occupied by bark, fruit and twigs (100 — leaf litter %)
is very appreciable, especially in the tall scrub forest. Turner (1954) estimated that
leaf mass in the stand of low scrub forest which he examined occupied only 43% of
the total fall. He found that twig fall was relatively much more important in the
sclerophyll communities, and especially in dry sclerophyll forest, than rainforest.
stands — both as regards the dry weight returned and also the nutrient return.

Reference to Tables 2 and 3 shows that of the leaf fall, Angophora forms at least
approximately 50% and usually much more of the total mass. It therefore was
analysed separately from the remainder for nitrogen. These analytical data together
with that of twigs, of a mixed leaf sample, and of mature fruit and seed were
presented in Tables 15, 17 and 18 (Hannon, 1956).

Of the total annual litter fall of approximately 700 lb. per acre per annum,
assuming that 250 lb. represents leaf litter averaging 5,000 p.p.m. nitrogen, and that
450 lb. represent twig, fruit, seed and bark fall from trees and shrubs, averaging
3,000. p.p.m. nitrogen, an annual cycle of only 2-6 lb. nitrogen per acre might be
anticipated. It will be shown that the annual return of litter in Hawkesbury Sandstone
communities is below the average of figures given in the literature; in consequence
this value for the annual nitrogen return is also low.

EE

76 NITROGEN IN HAWKESBURY SANDSTONE SOILS AND THEIR PLANT COMMUNITIES, II,

Hstimates of the total vegetative cover in low scrub forest are, according to the
topographic position of the stand, 10-40 tons per acre of dry matter; 60-120 tons per
acre is the yield in tall scrub forest. In light of these figures, the value for the
annual litter production, 0-31 ton per acre, represents only a small fraction of the
total plant mass and an exceedingly small fraction of the total nitrogen content of
the ecosystem. Leaf-bearing twigs of diameter up to approximately one inch, which
include the fractions most commonly found in the fallen litter, have been separated
from the crowns of trees felled in areas of sandstone which are being cleared. The
observed annual litter fall occupies 5-10% of the mass ef tiis fraction.

Inspection of the morphological character of several shrub species indicates that
the usual life span of leaves would be 3-4 years. Casuarina appears to replace its
foliage annually. Jacobs’ (1936) investigations indicate that the life of Hucalyptus
species leaves is short by comparison with other evergreens. Of the species studied,
only Hucalyptus haemastoma occurs on Hawkesbury Sandstone and its leaves are
considered to have an average life span of not longer than 18 months. Some leaves
lived 2-3 years, a few longer, but the average was surprisingly short. These results
were obtained from trees growing at Canberra, where environmental conditions are
markedly different from those of the sandstone communities. Saplings of Hucalyptus
piperita, another sandstone species, showed a similar life span. Jacobs records that
the life span is affected by position in the crown, bursts of growth, and flowering,
insect attack and species. Jacobs has shown that if general conditions are equal,
leaves fall from more rapidly growing crowns more quickly than from slow growing
ones. He remarks: ‘The tendency for some species to hold leaves longer than others
is probably partly a function of growth rate, and partly a genetical character.”

It is considered that the average life span of 18 months for Hucalyptus haemastoma
leaves as was found in Canberra cannot apply to this species when growing in the
Sydney habitat. On an 18 months’ life span, approximately 60% of the canopy would
be renewed annually. It is difficult to reconcile this estimate with observed leaf fall
for species of Hucalyptus and Angophora in the Sydney district. The above-mentioned
value of 10% of canopy probably is closer to the correct value for individuals in
these communities.

No information strictly comparable with sandstone forests is available. The
only other work that has been done in Hucalyptus forests is in Canberra, Western
Australia and Victoria. Jacobs (1936) estimated that the alpine ash (H. gigantea
Hook.) contained 5,800 lb. per acre of oven-dry weight of foliage in the tree canopy
and that 15-20 tons of litter (leaf litter and branch shed) is dropped in ten years
by the canopy of a fully stocked stand.

In virgin jarrah forests (#. marginata) of 105-115 feet height, 50-60% of the
total litter fall occurs between January and March and leaves form approximately
50% of the total of approximately 2,100 lb. per acre. Hatch (1955) estimates a return
of 9:20 lb. nitrogen per acre per annum, of which 5-23 lb. is contributed by leaf fall.

In a 220-year-old mature Eucalyptus regnans forest of trees 250 feet in height,
Ashton (private communication) collected an average of 7,208 lb. dry weight of litter
per acre per annum over a four-year period; this amount contained 53-5 lb. nitrogen.
Leaf fall from H. regnans alene contributes 22-2 lb. nitrogen.

The Western Australian and Victorian work is in close agreement as to the main
season of leaf fall. Both sites are further south than Sydney, where the maximum
litter fall apparently occurs in October. Very great differences in the quantity of
total litter fall occur between the different forests, but this is a reflection of the
differences in their quality.

In hardwood forests of Southern Carolina Piedmont, Metz (1952) records that
litter returns were 4,000—5,000 lb. per acre for the single year during which collections
were made. Chandler (1941, 1943) records 2,425-3,020 lb. per acre of litter in the
deciduous hardwood forest stands of North America and this includes a return of
16:6 lb. nitrogen per acre. In the evergreen coniferous forests, 2,463 lb. per acre of
litter containing 23:6 lb. nitrogen were recorded.

BY NOLA J. HANNON. 17

Handley (1954) and Scott (1955) have provided extensive reviews of the literature
on this subject. By comparison with all other forests, the litter fall in the sandstone
communities is very low.

Litter Decomposition.

Jacobs (1936, 1955) remarks that Hucalyptus leaf litter disintegrates “fairly quickly
by world standards”. He points out that the rate of disintegration varies with climate,
being faster in warm, moist localities and slower in cold or dry areas. It is of interest
to note Kittredge’s (1948) comment also: “Most of the species of Hucalyptus introduced
into the United States form little or no forest floor, and the question may be raised
whether these species have the beneficial effect on the soil that is attributed to most
other genera.” ;

Presentation of Data.

Despite the low rate of litter fall, the litter layer on the forest floor is always
well developed. In the sandstone communities the litter consists mostly of whole
leaves. It is only just above ground level that the leaves are broken into smaller
pieces. This is suggestive of chewing action by animals rather than enzymatic
decomposition by fungi and bacteria. The depth of litter is variable, not only between
different formations but also within single stands. It usually is of only one inch
depth, but in localized pockets it may accumulate to four inches. The mass of litter
from the forest floor is shown in Table 9 and provides evidence suggesting a slow
rate of litter decomposition. The samples were collected in typical stands and

TABLE 9.
The Mass of the Forest Floor.
Date of Mass of Forest
Formation. Sampling. Floor.
(g./sq. yd.)

28th November .. 508
778
4th March As 1580
Low scrub forest .. oS 704
987
20th September .. 884
670
4th March the 1892
: 1476
Tall scrub forest - 853
20th September .. 1516
1008

represent quite average samples of the general litter cover. The samples were
dried and sieved to remove sand grains, especially from the lower layers
where the leaf material is broken into fragments. The values in Table 9
represent the total litter on the ground (leaves, twigs and bark) and show no definite
trend with season. As has already been mentioned, although leaf fall rose in October,
the twigs showed no definite seasonal pattern. The total litter fall does not appear
to exhibit a cyclic nature as it does in Western Australia, Victoria or in overseas
deciduous forests. More critical investigations of the litter mass on forest floors in
various seasons would be of interest, but it appears that since only a small fraction
of the total is decomposed each year, variation in sampling may mask the incidence
of seasonal effects.

Discussion of Data.

Owing to the limited areas where litter may accumulate, the values in Table 9
require correction for conversion to an acreage basis. However, taking a figure of
1,000 grammes per square yard as an average value of the quantity of accumulated
litter and assuming a rate of 70 grammes per square yard per annum (based on
figures given in the preceding section), accumulation of litter has obviously proceeded

78 NITROGEN IN HAWKESBURY SANDSTONE SOILS AND THEIR PLANT COMMUNITIES, II,

over a period of 14 years. Periodic occurrences of adverse environmental conditions,
such as severe drought, may be responsible for exceptionally heavy litter falls and
contribute largely to the quantities of litter found on the forest floor. During occasional
gales, twigs with living leaves still attached, are frequently torn from tree tops. Very
heavy falls of scorched leaves have been observed immediately after bush fires. Such
occurrences would reduce the estimate of 14 years as the period during which litter
accumulates. Even allowing for these occurrences, however, and even were some
nutrients liberated when the litter eventually reached the ground surface and decom-
position began, certain amounts of nutrients must be locked within the litter for long
periods. Specht (1953) finds that in sclerophyllous heath vegetation in South
Australia, nitrogen and phosphorus are not liberated until decomposition is complete.

By studying the mass of the forest floor in several stands of H. marginata which
had been protected from fire for varying periods, Hatch (1955) records that after
twenty years’ protection, which was the maximum period studied, the forest floor was
still accumulating and the equilibrium between litter fall and litter decomposition

TABLE 10.
(Taken from Jenny et al., 1948.)*
Weights of Forest Floors in the United States.
(Mostly on oven-dry basis.)

Weights of Forest Floor.
Number of (Ib./acre.)
Locality. Vegetation. Samples. ||

Range. Average.

Minnesota and Wisconsin Basswood, maple, aspen, 16 25,050-59,787 39,556
birch.

Connecticut ae .. | Hardwoods. 5 7,000—94,000 60,244
Fern Canyon (California) Chapparral. 101 10,000—47,000 33,200
Ohio, Indiana Ae .. | Sassafras, black locust. ? 6,800-10,200 8,500
Ashville (North Carolina) | Pine, oak. 2 6,300 7,900 7,100
South Appalachians .. | Hardwood. 5 3,146—-16,359 10,435
Victoria, Australia .. | Eucalyptus regnans. = = 20,070
Western Australia .. | E. marginata. —_— i = 20,160
Sydney, N.S.W. .. .. | Hucalyptus—Angophora. — = 5.600

* Values for three types of Australian Hucalyptus forest are included for comparison.

was not yet attained. At this time of 20 years’ protection, 44-64 tons of litter had
accumulated per acre. He suggests 44 tons from a canopy cover of 35% (a stand of
55-65 feet in height) and 64 tons from 50% cover. Where the canopy was complete,
9 tons had accumulated after 20 years’ protection.

Ashton (private communication) collected 20,070 lb. per acre in the total litter
fraction of the mature H. regnans forest floor. The ratio of the total floor to the annual
litter fall was only 2:54, indicating a rapid decomposition of the litter.

It is of interest to compare the mass of the sandstone forest floors (average of
5,600 lb. per acre) and the figures of Hatch and Ashton given above with the data
summarized by Jenny, Bingham and Padilla-Saravia (1948) and given in Table 10.
Scott (1955) has provided an extensive review of the literature on this subject also.

Kittredge (1948) notes that the total forest floor accumulation as compared with
the annual litter fall may vary between a ratio of 1 or less in the tropics, where litter
is decomposed very rapidly, and 60, as is found in the birch-sugar maple—spruce forest
in northern New Hampshire. In this area, the mass of the total forest floor has a
value of 120 metric tons per acre. The ratio varies commonly betweeh 3 and 15,
and is essentially a measure of the rate of the decomposition of litter. Where
equilibrium between accumulation and decomposition has heen attained, the ratio also
represents the number of years for the establishment of that equilibrium. Jenny,
Gessel and Bingham (1949) have applied certain refinements to these latter principles
in their comparative study of the decomposition rates of organic matter in temperate
and tropical regions.

BY NOLA J. HANNON. a)

The agencies which are active in litter decomposition in the sandstone communities
other than Casuarina stands have received no attention. It is to be expected that the
low moisture and nutrient levels would limit the activity of microcrganisms, including
the animal population, just as it does the development of the plant cover. Clark (1950)
has paid attention to the role of the animal populations in Casuarina decomposition,
and several workers—Fenton, Fraser (1948), Anderson (1948), Coogan (1949), Simons
(1950), Carne (1950)—under the leadership of Burges, have examined the fungi and
bacteria which are engaged in Casuarina decomposition. Clark (1950) found that the
fauna is very similar to that of pine forests in Europe, both in respect of species and
numbers of animals present. It has already been pointed out that Casuarina grows
in almost pure stands where the heavy litter layer allows greater retention of moisture
than in Hucalyptus forests. The faunal population of the latter may resemble tnat of
the humus layers of regenerating jarrah forest which have been investigated by
MeNamara (1955). He reports a similarity in numbers and composition of the insect
fauna in these forests with those in Danish beechwoods and North American hardwood
forests, but the absence of the high proportion of earthworms.

GENERAL DISCUSSION.

The evidence presented indicates that organic matter production and litter and
nitrogen returns in the Hucalyptus forests on Hawkesbury Sandstone are small by
comparison with other forests. In addition, the forest floor is not so well developed
as in many other types of forests; although decomposition is relatively slow in the
sandstone forests, Hucalyptus forest is generally considered to show a rapid rate of
litter decomposition by world standards. The evergreen, broad-leaved and sclerophyllous
Eucalyptus forests are thus of a different nature from those found in other areas.

It would be of interest to compare the ratios of the nutrient content of soil: plant:
litter in Hucalyptus forests with the hardwood, coniferous and deciduous types of
forests. Such a comparison would clarify whether the differences in the nutrient
circulation mentioned above are a reflection of a smaller nutrient capital in the
ecosystem or a different proportion between the three fractions. It is possible that a
greater percentage of the capital is “mobile” in other forest. types, thus allowing the
exercise of a less rigid economy.

The quantities of nutrients contained in ecosystems have received no attention
until recently. Estimates of the absolute nutrient content had been made on only
separate aspects of the ecosystem such as the litter or the soil. There is much to
recommend the simultaneous examination of all three fractions of the ecosystem,
since such an approach involves the synthesis of otherwise isolated observations and
emphasizes the functioning of the ecosystem as the entity it is. The employment of
expression in absolute rather than relative terms also offers several advantages. Not
only are such values mentally digestible, but they avoid the deception which
concentration values frequently present.

In connection with the afforestation programme of the Calluna moors, Rennie
(1955) has made an outstanding and valuable contribution involving this approach.
He examined the original records of analytical data on the composition of the organs
of individual mature temperate forest trees and in the thinnings. By combining these
data with records of the density of wood, growth rate of trees and yield per acre
of forest stands, he estimates and fractionates the calcium, potassium and phosphorus
content of pine, other conifer and hardwood forests into the component plant organs
in the main timber stand and the thinnings. He notes that only seldom have nitrogen
analyses been recorded, but includes those that are available. By this means he
assesses the nutrient demands of pine and birch stands and compares these with that
‘of Calluna. An evaluation of the ability of Calluwna soils to provide for the nutrient
demands of the pine or birch is thus possible.

Table 11 is based on Rennie’s compilation of data relating to nitrogen. The
available information allowed the calculation of the nutrient content of the plant cover
and soil, but the litter fraction had to be ignored in some instances. The estimated
nitrogen content of the fractions of the Hawkesbury Sandstone ecosystem is included

NITROGEN IN HAWKESBURY SANDSTONE SOILS AND THEIR PLANT COMMUNITIES, II,

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for comparison. Inspection of this table shows that the nitrogen content of the
Hawkesbury Sandstone ecosystem is lower than any of the others listed.

Another feature of. considerable interest in the Eucalyptus forest is the plant
nitrogen content : rock and soil nitrogen content ratio value. It is in excess of all
values other than the pine-afforested moor community, which would appear to be
incapable of survival, if dependent on only the naturally occurring nutrient supply
of the Calluna moor soil. Comparison with the Scots Pine Forest stand would be of
greater significance, since the latter is a naturally occurring and somewhat similar
formation. It appears that the Hucalyptus forest plant cover is making a greater
demand on the soil reserve of nitrogen than does the pine forest. However, Rennie
does not include a value for the nitrogen content of the litter fraction in the Scots
Pine Forest. Since these forests have such a well-developed litter layer, which contains
considerable quantities of nutrients, it is illogical to exclude it from the calculation.
Presumably, its value would approximate that given for the pine afforested moor; in
this instance, the ratio of plant + litter N: soil N would become very similar in the
pine and Hucalyptus forests. The chief difference thus lies in the proportion of
nutrient immobilized in the litter layer. The litter fraction of the sandstone ecosystem
occupies only a minor fraction in comparison with the Calluna moor stands and
especially the pine forests. When nutrient balance is under consideration, the logic
of Rennie’s grouping of the fractions of the ecosystem into plant + litter as opposed
to soil is open to question. The litter fraction forms part of the reserve upon which
the plants draw for their nutrient requirements. Considerable proportions of the
root systems are located in the litter layer of pine forests. On this basis the ratios
of plant nitrogen content : soil + litter nitrogen content values are:

Eucalyptus forest Sot tebe eh OTA
Pine sforest: aap 052 005.0 2) 350-102

The abovementioned suggestion that the eucalypt forest plant cover makes a greater
demand on the nitrogen reserves of the ecosystem than does the pine forest would
therefore appear to be a valid conclusion, especially since more than 50% of the rooting
medium nitrogen content is bound within undecomposed parent material; elimination
of the nitrogen content of the rock increases the ratio of plant: “true soil’ to 0-354.
This character possibly is the key to the differences in nutrient circulation previously
noted between Hucalyptus and other forests.

This character of the Eucalyptus forest as contrasted with other ecosystems may
also be reflected in other nutrient elements such as phosphorus, potassium and calcium.
Evidence to be presented later suggests that the demand made by the plant cover on
the soil for calcium is even greater than that for nitrogen. This is a matter urgently
in need of investigation at the present time, when Australian ecologists are emphasizing
the possibility that the sclerophyllous character of the native vegetation may be
determined by low levels of nutrient supply.

Rennie (1955) has shown that the quantities of all three nutrients, calcium,
potassium and phosphorus, taken up by forest growth decrease in the order: hardwood,
coniferous other than pine, and pine forest. “All three types of forest take up
considerably less potassium and phosphorus compared with agricultural crops, but
the amounts of calcium are not greatly less; moreover, as all except very young
forest growth differs from agriculture in taking up calcium and not potassium, in
greater amount, the importance of calcium to forest growth appears to have been
markedly under-estimated.” It is of interest to note that in her studies of litter
decomposition in Casuarina stands on Hawkesbury Sandstone, Fraser (1948) has
suggested the occurrence of a vigorous calcium cycle in these communities. In
investigations of the composition of eucalypt forest litter, Turner (1954) has also
noted that the calcium and potassium content of the leaf litter in dry sclerophyll
Hucalyptus forest communities, when expressed on a concentration basis, exceeds that
in the wet sclerophyll forest stand studied which occurred on the fertile chocolate loam
of the Narrabeen series. In these areas the nutrient and water supply are at very
favourable levels for plant growth. Since litter fall in wet sclerophyll forest
communities exceeds by a factor of 2:6 that of the dry sclerophyll, the absolute

82 NITROGEN IN HAWKESBURY SANDSTONE SOILS AND THEIR PLANT COMMUNITIES, II,

amounts of all nutrients in the total litter fall are less in dry than wet sclerophyll
forest. Both concentration and absolute values for phosphorus in the litter were much
lower in dry than wet sclerophyll communities.

Features of the circulation of nitrogen in the Hawkesbury Sandstone ecosystem
which have been discussed above are summarized in Text-figure 2. Both the mass
of each fraction and its nitrogen concentration have been plotted quantitatively, the
mass of each fraction being proportional to the area it covers, and the nitrogen
concentration being proportional to the density of the stippling. The diagram thus
gives a graphic representation of the very small proportion of nitrogen present in
concentrated form in the ecosystem, since the great bulk is locked within the soil

Text-fig. 2.—The fractionation of nitrogen in a Hawkesbury Sandstone ecosystem.
A: Undecomposed parent material; B: Soil; C: Tree stratum; D: Shrub stratum; EH: Litter
accumulated on forest floor; F: Annual litter fall.

The mass of each fraction is proportional to the area it covers, and its nitrogen
concentration to the density of the stippling; consequently it has not been possible to
represent accurately the relative volume of each fraction, eg., tree height is grossly
underestimated in comparison to soil depth. Refer to pp. 66-67 for details of the calculation.

and woody tissue; it also shows the very small proportion that is “‘mobile’ — that is
the proportion involved in the annual circulation of nitrogen within the community.

While thus a closed cycle of nutrient movement will exist in the Hawkesbury
Sandstone communities as in all virgin communities, and while on theoretical grounds
a mature community exhibits no net gain or net loss, a realization of 100% efficiency
is most improbabie, particularly when so many transfers of the nutrients from one
form to another and from one phase of the ecosystem to another are involved. A
state of equilibrium exists in which processes tending towards the impoverishment of
the nutrient status, if not completely counterbalanced by nutrient release from the
weathering of the parent rock or by external accessions, at least proceed extremely
slowly. The sources of loss and gain of nitrogen in these communities require
examination and these studies will be reported later.

APPENDIX.
THE DENSITY OF HAWKESBURY SANDSTONE AND THE APPARENT DENSITY OF
SANDSTONE SOILS.
Methods.

Weighed rock fragments were coated with a thin layer of paraffin wax and
immersed in a known volume of water in a measuring cylinder. A glass rod was
used to remove all air bubbles from the rock surface and the volume of displaced
water measured. Measurements of the rock pieces are shown in Table i.

No special sampling tool designed to provide for apparent density measurements
of soil samples by standard methods was available. Tins used for the storage of

BY NOLA J. HANNON. 83

particular types of foodstuffs and consequently of uniform dimensions were therefore
opened at one end and punched with a small hole to allow for an air vent at the other.

A pit was dug and the tins were placed horizontally at various depths down the
exposed profile. A wooden block was used to cover the tins and hammered carefully
so that the tins were driven into the soil profile just to their full depth. By this
method, little, if any, compaction of soil occurred. Soil blocks containing the tins
filled with soil were then dug out. The soil was cut level with the top of the tin,
earefully removed, dried out and weighed. Density values were calculated by
comparing the values of the dried mass of soil with the volume contained by the tin.
Values of samples from various depths in several formations are shown in Table ii.

TABLE i.
Mass, Volume and Density Values of Fragments of Hawkesbury Sandstone.

Sample Mass. Volume. Density.
Number. (g.) (c.c.) (g./¢.c.)

ul 18-8 8-5 2-21

2 36-1 17-5 2-06

3 36:5 18-0 2-03

4 69-1 31-0 2-23

5 105-3 50-0 Zeal!

6 167-0 | 79-0 Zatt

ANALYSIS OF Woobdy TISSUE OF HAWKEFSBURY SANDSTONE SPECIES.

Details of the nitrogen and carbon content of the bark-free wood of several species
are shown in Table iii. As in the leaf and seed tissue values previously recorded,
the wood of the legume species has a much higher nitrogen content than the
non-legumes. Although the living tissue in woed forms only a small proportion by
mass, it contains proteins and sugars, whereas the lignified cells are empty. It would

TABLE ii.
Mass and Apparent Density of Hawkesbury Sandstone Soil.
Samples were collected at various depths in profiles located in several formations.
Tins of a volume of 230 c.c. were used throughout the sampling.

Depth of Apparent
Formation. Sample. Mass. Density.
(in.) (g.) (g./c.c.)
Shrub swamp is ae Be 1 - 32 289 1-26
4i- 7 271 1-18
Sedge swamp ae ee os 0-3 294. 1-28
3-6 317 1-38
10 -13 332 1-44
18 -21 348 1-51
30 -33 396 1:72
Low scrub forest age in 0 —- 22 326 1-42
94-12 351 1-53
152-18 333 1-45
Tall scrub forest .. ie ans 0-3 309 1-34
16 -19 348 1-51
25 -28 334 1-45
33 -36 366 1-59

thus be a proportionately greater contributor to the nitrogen content of the wood
sample. Russell (1950) notes that the lignins of leguminous plants are usually richer
in their nitrogen content than non-legumes. Apparently this, together with higher
nitrogen levels in the living cells of legumes, is responsible for the legume carbon-
nitrogen ratio, which is markedly lower than that of the others. The two species
of highest carbon-nitrogen ratio, Banksia serrata and Hucalyptus paniculata, have

84 NITROGEN IN HAWKESBURY SANDSTONE SOILS AND THEIR PLANT COMMUNITIES, II,

density values of approximately twice that of the other species. It is likely, therefore,
that differences in the carbon and nitrogen values could be correlated with anatomical
detail, if it were available.

Records of the nitrogen content of wood are not numerous, but it appears that
the values shown in Table iii are typical. Ramann (1890), quoted by Lutz and
Chandler (1949), reported the nitrogen content of wood as 1,000—-5,000 p.p.m.; Kalnins
and Liepins (1938) found that Scotch pine wood contained 1,200-1,600 p.p.m. Converting
the protein values of sapwood given by Wise and John (1952) to nitrogen by the
conventional factor of 6:25, the content of the following species is:

Ash (Frazinus elatior L.) Seen. on2alOZ pap ameNe
Elm (Ulmus sativa Mill.) .. .. .. 2,768 p.p.m. N.
Scotch pine (Pinus sylvestris L.) .. 1,328 p.p.m. N.
TABLE iii.
The Concentration of Nitrogen and Carbon in Bark-Free Wood.
Total Organic Carbon :
— Nitrogen. Carbon. Nitrogen
(p.p.m.) (p.p.m.) Ratio.
PROTEACEAE
Banksia serrata .. es 1,024 472,446 461
MYRTACEAE
Angophora costata te 2,976 460,570 155
Eucalyptus paniculata Sm. 1,107 507,502 459
E. piperita .. is as 1,736 468,522 270
LEGUMINOSAE
Acacia baileyana ¥. Muell. 4,784 421,385 88
A longifolia (Andr.) Willd. 5,027 389,908 78
A. maidenii F. Muell. oe 7,913 440,000 56
CASUARINACEAE
Casuarina torulosa. . | 1,966 464,546 236

Acknowledgement.

Grateful acknowledgement is made to the following persons, whose interest and
co-operation have been of great assistance: Associate Professor J. M. Vincent; Mr. S. N.
Blow, of the Colonial Sugar Refining Company, Sydney; Mr. N. A. Powell, of Bilpin,
N.S.W.; Mr. L. Bryant and several officers of the Forestry Commission of New South
Wales; officers and employees of the Metropolitan Water, Sewerage and Drainage Board.

References.

ANDERSON, BEVERLY J., 1948.—The Role of Fungi in the Decomposition of Litter formed by a
Pure Stand of Casuarina littoralis Salisb. (suberosa Ott. and Dietr.). Honours Thesis,
Department of Botany, University of Sydney.

CARNE, PHILLIDA, 1950.—An Investigation of Organic Acids in the Green Needles and
Decomposing Residues of Casuarina littoralis Salish. (swberosa Ott. and Dietr.). Honours
Thesis, Department of Botany, University of Sydney.

CHANDLER, R. F., 1941.—Amount and Mineral Nutrient Content of Freshly Fallen Leaf Litter
in the Hardwood Forests of Central New York. J. Amer. Soc. Agron., 33: 859-871.

—, 1943—Amount and Mineral Nutrient Content of Freshly Fallen Needle Litter of
Some North-Hast Conifers. Proc. Soil Sci. Soc. Amer., 8: 409-412.

CLARK, D. P., 1950.—The Fauna of Casuarina Litter and the Soil. Honours Thesis, Depart-
ment of Zoology, University of Sydney.

CooGAN, THELMA M., 1949.—The Role of Certain Fungi in the Removal of Bases from
Casuarina littoralis Salisb. (swberosa Ott. and Dietr.) Litter during the Harly Stages of
Decomposition. Honowrs Thesis, Department of Botany, University of Sydney.

FRASER, JupDITH A., 1948.—The Chemical Analysis of Living Casuarina littoralis Salisb.
(suberosa Ott. and Dietr.) Needles and Decomposing Residue. Honours Thesis, Department
of Botany, University of Sydney. :

HANDLEY, W. R. C., 1954.—Mull and Mor Formation in Relation to Forest Soils. Bwil. For.
Comm., Lond., 23, 115 pp.

HANNON, Nota J., 1956.—The Status of Nitrogen in the Hawkesbury .Sandstone Soils and
their Plant Communities in the Sydney District. 1. The Significance and Level of
Nitrogen. Proc. Linn. Soc. N.S.W., 81, 2: 119-143.

PLATE I.

Proc. Linn. Soc. N.S.W., 1958.

e&)
ESS
(RS

Ore

=
EER Bs ae
Ce Vess7 ‘
oN SZ
{\ Re

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ve

Oe

eS
SS

ae
ZI

oe)
gc)
om,

SU,
Kore
SOK

SS
vs
Ss

ES

Seed coat anatomy in Hucaiyptus.

BY NOLA J. HANNON. 85

Hatcu, A. B., 1955.—The Influence of Plant Litter on the Jarrah Forest Soils of the
Dwellingup Region, West Australia. Leafl. For. Bur. Canberra, 70, 18 pp.

Jacops, M. R., 1936.—The Primary and Secondary Leaf Bearing Systems of the Eucalypts
and their Silvicultural Significance. Bull. For. Bur. Aust., 18: 1-78.

, 1955.—“Growth Habits of the EHucalypts.” Forestry and Timber Bureau. Common-
wealth Govt. Printer. 262 pp.

JHNNY, H., 1941.—“Factors of Soil Formation.” McGraw-Hill, New York. 281 pp.

JENNY, H., BINGHAM, F., and PADILLA-SARAVIA, B., 1948.—Nitrogen and Organic Matter
Contents of Equatorial Soils of Colombia, South America. Soil. Sci., 66: 173-186.

JENNY, H., GesseL, S. P., and BINGHAM, F., 1949.—Comparative Study of Decomposition
Rates of Organic Matter in Temperate and Tropical Regions. Soil. Sci., 68: 419-432.

KITTREDGE, J., JR., 1948.—“Forest Influences.” McGraw-Hill, New York. 394 pp.

Lutz, H. J.. and CHANDLER, R. F., gR., 1949.—‘Forest Soils.” John Wiley and Sons, New York.
4th printing. 514 pp.

McNamara, P. J., 1955. A Preliminary Investigation of the Fauna of Humus Layers in
the Jarrah Forest of Western Australia. Leafl. For. Bur., Canberra, 71. 16 pp.

Merz., L. J., 1952.—Weight and Nitrogen and Calcium Content of the Annual Litter Fall of
Forests in the South Carolina Piedmont. Proc. Soil Sci. Soc. Amer., 16: 38-41.

RENNIE, P. J., 1955.—The Uptake of Nutrients by Mature Forest Growth. Plant €& Soil,
7, 1: 49-95.

RUSSELL, H. J., 1950.—‘‘Soil Conditions and Plant Growth.” Longmans, Green and Co.,
London, 8th ed. 635 pp.

Scotr, D. R. M., 1955.—Amount and Chemical Composition of the Organic Matter contributed
by Overstory and Understory Vegetation to Forest Soil. Bull. Yale Univ. Forestry School,
62. 73 pp.

SIMONS, HILDA R., 1951.—A Study of Fungi Active in the Decomposition of Litter of
Casuarina littoralis Salisb. (suberosa Ott. and Dietr.). Honours Thesis, Botany Dept.,
University of Sydney.

SPECHT, R. L., 1953.—The Ecology of the Heath Vegetation in the Upper South Fast of
South Australia — A Preliminary Investigation. Ph.D. Thesis, University of Adelaide.

TURNER, J. C., 1954.—Some Aspects of the Nutrient Cycle in Rainforest in New South Wales.
Honours Thesis, Faculty of Agriculture, University of Sydney.

Wise, L. E., and JOHN, H. C., Ed. 1952— “Wood Chemistry” I. Am. Chem. Soc. Monograph
Series. Reinhold, N.Y. 1-688.

A

86

POLLEN AND POLLINATION IN THE EUPOMATIACEHAE.
By A. T. HorcuHxKiss, Department of Biology, University of Louisville.

(Plate ii; twenty-six Text-figures. )
[Read 28th May, 1958.]

Synopsis.

The formation of the microspores in Hupomatia laurina is described and illustrated.
Standard descriptions and illustrations of the pollen grains of both H. lawrina and EL. bennettii
are given. Pollination in H. bennettii is similar to that in H. laurina. Comparisons are made
with other ranalian plants.

INTRODUCTION.

More information is needed as a basis for understanding the structure and
relationships of the woody ranalian plants. As a beginning of a series of papers on
the Eupomatiaceae, the geographical distribution of the family has been described
(Hotchkiss, 1955). It is hoped that the present paper will be followed by additional
information on this interesting family as studies are completed on various anatomical
and morphological details. Materials were collected in New South Wales and
Queensland as indicated in the text.

MICROSPOROGENESIS.

The long, narrow, sunken sporangia are closely filled with microspore mother cells
within cell walls which vary from an angular, isodiametric to a rather elongate form
which persists through the tetrad stage (Text-figs. 1-22). During meiosis the cell wall
thickens and becomes gelatinous in appearance. In EHupomatia lauwrina, meiosis in the
microspore mother cells is normal and ten bivalents can be seen at metaphase I.
These usually separate in a synchronous anaphase (Text-figs. 1-2). Infrequently, at
anaphase I, however, there appears to be early separation of one bivalent, as shown
in Text-figures 3-5; there also may be a tardy separation of one or more bivalents
(Text-figs. 6-8).

After division I, a cleavage furrow begins to form as in Magnolia (Farr, 1918)
but stops with the formation of an internal ridge filled with wall material at the
beginning of division II (Text-figs. 9-10). At the end of the second division, the first
furrow continues its growth and at the same time additional furrows appear. Then,
simultaneously, all the furrows divide the mother cell into four compartments containing
the microspores (Text-figs. 11-16). At interkinesis there is a conspicuous spindle
fibre apparatus remaining between the two dyad cells (Text-figs. 9-10). At the
completion of meiosis II, spindle fibres are conspicuous between the nuclei of the
tetrad and remain so until the cytoplasmic connections between the microspores are
finally cut off by the advancing furrows (Text-figs. 11-16). From the more isodiametric
young tetrads with 4-6 spindles there develop the commoner tetrahedral, isobilateral,
rhombic and decussate tetrads enclosed in nearly spherical mother cell walls; from
the linear, young tetrads with spindle connections between only the closely adjacent
nuclei, there develop the rarer linear and T-shaped tetrads enclosed in elongate mother
cell walls (Text-figs. 17-22).

POLLEN.

The pollen grains of Hupomatia lawrina have been figured by Hamilton (1897),
figured and described by Erdtman (1952). The pollen grains of both species of

PROCEEDINGS OF THE LINNEAN SocieTy oF New SoutTH WALES, 1958, Vol. Ixxxiii, Part 2.

BY A. T. HOTCHKISS. 87

Eupomatia are here figured and described (following the system of Erdtman) with
additional details as follows:

Eupomatia laurina R. Br. (Text-fig. 23). Bulli Pass, New South Wales. Grains
nearly spherical, spheroidal prolate; zonisulculate with a distinct groove on either side
of the equatorial zone, rarely 3-sulculate; exine finely granular in the sulculus-zone;

& \ | | i?

Text-figs. 1-8.—Hupomatia laurina, meiosis. 1, Metaphase I; 2, Anaphase I; 3-5, Meta-
phase I with early separation of a bivalent; 6-8, Anaphase I with tardy separation of
bivalents. x 862.

polar axis slightly longer, from 44 to 50u, the equatorial diameter shorter, from
43 to 48u.

Eupomatia bennettii F. Muell. (Text-fig. 25). Bellingen, New South Wales. Grains:
nearly spherical, more or less rectangular in section; at first nearly isopolar, later:
one pole protruding; zonisulculate; exine finely granular in the sulculus-zone, granules:

88 POLLEN AND POLLINATION IN THE EUPOMATIACEAE,

finer than in #. lauwrina; polar axis slightly shorter, from 35 to 40u, the equatorial
diameter longer, from 37 to 44u.

In £. laurina an occasional tetrad remains coherent and thus indicates the polar-
equatorial relationship. The young pollen grain bulges in the equatorial zone because
of the folding of the thickened wall in a two-pleated manner. This results in the
formation of a broad equatorial ridge bounded on either side by furrows which extend
completely around the pollen grain in £#. laurina (zonisulculate—EHrdtman, 1952) and
nearly around in H#. bennettii. As the grain is expanded the flexible equatorial zone
straightens out although the grooves impressed on the protoplast remain; when the
grain is dried the walls are refolded and contract again (Text-fig. 26). Occasional
pollen grains may be seen germinating within the anther. In #H. lauwrina the pollen
tube emerges from one of the grooves (Text-fig. 24). What appears to be a beginning
pollen tube can often be seen growing from the apex of the pollen grain of H. bennettiw
{thus changing its isopolar shape) in Text-fig. 26.

Text-figs. 9-16—Hupomatia laurina, developing microspores. 9-10, Arrested cleavage
furrows at interkinesis; 11-16, Spindles and cleavage furrows developing after meiosis II.
'The lobing of the future microspores can be seen before the furrowing is completed but no
cell plates are present. x 937.

Text-figs. 17-22.—Hupomatia laurina. Tetrads after cleavage showing some arrangements
of microspores. x 937.

POLLINATION. -

Robert Brown (1814) discussed the unusual pollination mechanism in Hupomatia
laurina “by certain minute insects eating the petal-like filaments (staminodia), while
the antheriferous stamina ... remain untouched”. Bennett (1860) reported that Harvey
in 1834 had found a small brown Curculio on ZH. laurina in the Illawarra district of
New South Wales. Hamilton (1897) confirmed the presence of beetles in the flower of
E. laurina and described in detail the behaviour of this insect, Elleschodes hamiltoni
T. Blackburn. To this account there can be added that the overarching, inner, rather
fleshy staminodes are equipped with succulent glands composed of delicate secretory-
like cells. Although the glands may be the first objective of the feeding beetles, a
large portion of the staminode is often consumed. No nectar has been observed. From
flowers placed in the laboratory, beetles emerge in the early evening darkness and fly
about. During the day, in the laboratory or the field, they show little tendency to
leave the flowers or to fly at all. Besides the Illawarra district and Bulli Pass, beetles
of this species have been collected from the upper Williams River valley (Messmer
and Musgrave, 1944), and south-east Queensland.

From flowering plants of H. bennettii at Bellingen, New South Wales, collections
of similar beetles were made in September, 1954. These were kindly identified by

BY A. T. HOTCHKISS. &&

Mr. A. Musgrave, of the Australian Museum, Sydney, as Hlileschodes sp. In this
connection it may be noted that the flowering time of H#. bennettii is several months
removed from that of #. laurina, and that the latter species flowers and fruits heavily
every other year and apparently not at all in the intervening years.

Text-figs. 23-24.—Hupomatia laurina. 238, Mature pollens grain; 24, Germination of pollew
grain from one of the grooves. x1150.

Text-figs. 25-26.—Hupomatia bennettii. 25, Mature pollen grain; 26, Outline of partly
dried pollen grain. x1150.

DISCUSSION.

The irregularities in anaphase I seem to have little effect on the subsequent
maturation of the tetrads which almost invariably are filled with four microspores of
equal size. Farr (1916) reported that the primary spindle persists in Nicotiana and
that four new and identical spindles were formed after the second division. However,
in Magnolia, Farr (1918) states that the primary spindle completely disappears as
meiosis II begins and that in telophase II a large number of fibres again appear.
Whether or not in Hupomatia the fibres of the primary spindle disappear as in
Magnolia remains undetermined. In Nicotiana there is neither a cell-plate nor a furrow
formed between meiosis I and II; in Magnolia both these structures are initiated,.
although neither is completed before meiosis II (Farr, 1916, 1918). Hupomatia
resembles Magnolia in possessing the arrested cleavage furrow but it lacks the cell-
plate vestige. In Hupomatia the persistent shape of the microspore mother cell wall
is related to the arrangement of the tetrad contained therein. Farr (1916) states that
it is quite probable that “the restraining tension of the wall has very little to do
with initiating the tetrahedral arrangement, though it may play a large part in the

90 POLLEN AND POLLINATION IN THE EUPOMATIACEAE,

maintenance of such orientation”. The tetrads of H. lawrina include the range of
forms (with intergradations) as shown by Maheshwari (1950) for the angiosperms.
In the Degeneriaceae, simultaneous divisions of the microspore mother cells result in
tetragonal tetrads (Swamy, 1949), in the Anonaceae the divisions are successive in
Asimina triloba (Locke, 1936), and successive or nearly so in Anona (Juliano, 1935).

Erdtman (1952) points out that the pollen grains of Hupomatia are similar to the
2- and 3-sulculate type found in the Atherospermoideae, and the sulculate pollen of
the Calycanthaceae. Possession of this type of pollen at once establishes the position
of the EKupomatiaceae in the Ranales (sensu latu) and, with some other features,
associates this family with those woody families generally considered the least
specialized in the order (Bailey and Nast, 1943; Bailey and Swamy, 1949; Canright,
1953; Swamy and Bailey, 1949; Wodehouse, 1935). Sulculate pollen is derived from
the monocolpate (one-furrowed—Wodehouse, 1935) pollen and distinguished from the
tricolpate pollen found in the majority of dicotyledonous plants.

Entomophily and, more specifically, pollination by beetles, has been considered by
various workers to possess considerable phylogenetic significance. The flowers of
Eupomatia (Plate ii) correspond to ‘the simple level with a haplomorphic arrangement
of petals” (Leppik, 1957). The strong, fruity, penetrating odour, and the generally
creamy white colour also suggest this classification. The nutritive tissues which are
confined to the tips of the inner floral appendages in Calycanthus (Grant, 1950) are
scattered over the margins and inner surfaces of the over-arching inner staminodes in
EHupomatia. Diels, in a paper on Kaferblumen (1916), compared beetle pollination in
Hupomatia with that in Calycanthus, other ranalian plants, and in Hncephalartos. He
suggested that beetles may have been one of the earliest existent orders of insects to
be anthophilous. The same views have been expressed by Grant (1950), whose
investigations on the pollination mechanism in Calycanthus occidentalis reveal a
striking similarity between that plant and Hupomatia. Grant sums up the evidence
from living plants and fossils for the hypothesis that beetle pollination was the
original condition in flowering plants.

Acknowledgements.

Materials for this paper were collected and studied while the author was a
member of the Botany Department, University of Sydney. For aid in collecting, I
am much indebted to Mr. W. D. Francis and Mr. L. S. Smith, of the Brisbane
Herbarium; to Mr. L. Webb, Plant and Soils Regional Laboratory, C.S.I.R.O., Brisbane;
to Dr. G. Hewitt, Dr. M. Elliott and Mr. B. Rickerby, of Bellingen, New South Wales;
and especially to Dr. S. Smith-White, University of Sydney, who guided me in making
the first collection.

References.
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Pollen and stamens. Jour. Arnold Arb., 24: 340-346. r

, and Swamy, B. G. L., 1949——The morphology and relationships of Austrobaileya.

Jour. Arnold Arb., 30: 211-226.

BENNETT, G., 1860.—Gatherings of a Naturalist in Australasia, p. 364. London.
Brown, R., 1814.—General remarks, geographical and systematical, on the botany of Terra

Australis. Appen. III in M. A. Flinders, A Voyage to Terra Australis, p. 65-66.
CANRIGHT, J. E., 1953.—The comparative morphology and relationships of the Magnoliaceae.

II. Significance of the pollen. Phytomorph., 3: 355-365.

Diets, L., 1916.—Kaferblumen bei den Ranales und ihre Bedeuting ftir die Phylogenie der

Angiospermen. Ber. Dew. Bot. Ges., 34: 758-774.

ERDTMAN, G., 1952.—Pollen Morphology and Plant Taxonomy. Angiosperms. Almaqvist and

Wiksell, Stockholm.

Farr, C. H., 1916.—Cytokinesis of the pollen mother cells of certain dicotyledons. Mem. N.Y.

Bot. Gard., 6: 253-317.

, 1918.—Cell division by furrowing in Magnolia. Amer. Jowr. Bot., 5: 379-395.
GRANT, V., 1950.—The pollination of Calycanthus occidentalis. Amer. Jour. Bot., 37: 294-297.
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N.S.W., 22: 48-55.

Hotrcuxiss, A. T., 1955.—The geographical distribution of the Eupomatiaceae. Jour. Arnold

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JULIANO, J. B., 1935.—Morphological contribution on the genus Anona. Philippine Agr., 24:

528-541.

BY A. T. HOTCHKISS. 91

LepPIK, H. E., 1957.—KHvolutionary relationship between entomophilous plants and anthophilous
insects. Hvolution, 11: 466-481.

Locks, J. F., 1936.—Microsporogenesis and cytokinesis in Asimina triloba. Bot. Gaz., 98:
159-168.

MAHESHWARI, P., 1950.—An Introduction to the Embryology of Angiosperms. McGraw-Hill
Book Co., New York.

Messmer, P. R., and MuscGraAve, A., 1944.—A trip to Barrington Tops. Aus. Mus. Mag.,
8: 194-199.

Swamy, B. G. L., 1949.—Further contributions to the morphology of the Degeneriaceae. Jour.
Arnold Arb., 30: 10-38.

, and Batiuey, I. W., 1949.—The morphology and relationships of Cercidiphyllum.

Jour. Arnold Arb., 30: 187-210.

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

EXPLANATION OF PLATE II.

Hupomatia bennett EF. Muell.—The solitary, terminal flower. A few of the thin, narrow,
petal-like, outer staminodia are seen reflexed at the right. Above these the staminodia are
longer, broad and fleshy. The arching, very fleshy, inner staminodia bear the glands of
nutritive tissue. The fertile stamens are invisible beneath the outer staminodia. Plant
collected at Bellingen, New South Wales. x1-25. (Photograph by Woodward-Smith. )

92

THE SPECIES OF THE GENUS ZRODIUM L’HER. ENDEMIC TO AUSTRALIA.
(With a key to ali the taxa known to occur in Australia.)

By R. C. Caroitin, University of Sydney.

(Twenty-eight Text-figures. )
[Read 28th May, 1958.]

Synopsis.
The specimens previously referred to H. cygnorum Nees in Lehm. have been critically
examined. Two new species are described (H. aureum and E. crinitum) and one new

Subspecies (H. cygnorum ssp. glandulosum). 'These all appear to be endemic to the Australian
continent, although #. crinitum appears to have been reported as a “wool alien” in Yorkshire,
England. A key to all the taxa known to occur in Australia and Tasmania is provided.

ERODIUM AUREUM, SD. Nov.

Herbeae perennes vel annuae, nanae, pilis glandulosis simplicibusque. Folia
elongata-deltoidea 3—4-lobis. Calyx ellipticum pilis glandulosis simplicibusque obtegitur.
Filamenta staminorum anguste lanceolata aristis longis, purpurea pallide. Pili rufoaurei
trans mericarpium ponent.

Dwarf, suberect, short-lived perennial! or possibly annual herb. Stems 1-3, up to
20 cm. tall, covered with soft glandular and simple hairs. Leaves 1:0-3:0 cm. long,
0-7-1-5 em. wide, lobed with three principal lobes; sinuses never reaching main vein,
glandular and simple hairs scattered over surfaces; margins toothed; petioles slender
with scattered hairs; stipules and bracts acute. Flowers in umbels of (2)-3(5) rarely
solitary, pedicels covered with glandular and, fewer, simple hairs. Sepals 5, oblong-
lanceolate, 0:5-0:8 ecm. long, 0-2-0-3 cm. wide, covered with both long glandular hairs
and simple hairs, the latter often more numerous towards the apex, shortly mucronate,
ciliate and membranous towards margin, particularly where covered in bud. Petals 5,
blue-purple with reddish veins and bases, ovate, c. 0:8 cm. long and 0-6 cm. wide.
Stamens 5; filaments narrow lanceolate, less than 0-1 cm. wide at their widest part,
0-45 cm. long including long aristate apex, often purple; anthers red; staminodes 5,
similar to staminal filaments, about 2/3 times as long. Stigmata 5, yellow. Mericarps 5,
oblanceolate, c. 0-5 cm. long and 0-2 ecm. wide, covered with short golden-brown, simple
hairs lying diagonally across the long axis; beak c. 3-5 cm. long with two pits at the
base and often 1-2 folds beneath each pit.

Chromosome number: 2n = 20.

Range: Central Australia, drier parts of South and Western Australia, and
extending into western Queensland.

Ecological distribution: Generally in open situations.

Holotype: Palm Valley, G. Chippendale, 26.8.1956. In Herb. N.S.W.

Specimens examined.—SoutH AUSTRALIA: N. end of Mann Ranges, Cleland, 23.8.1954
(S.A.95641035); Ernabella, Musgrave Ranges, Cleland, 19.8.1933 (S.A.95641016) ; between
Ernabella and Echo Hill, Cleland, 20.8.1933 (S.A.95641019 and 95641021); Echo Hill
between Moorilyanna and Ernabella, Cleland, 7.8.1933 (S.A.95641020 and 95641022);
Highway between Moorilyanna and Ernabella, Cleland, 7.8.1933 (S.A.95641023); H.-W.
line Wynbring, Cleland, 20.8.26 (S.A.95641041); E-W. line Barton, Black, 22.9.20
(S.A.95641003); E.-W. line Pimba, Cleland, 26.8.389 (S.A.95641015); Gov. (S.A.) North-
West Exped., Dr. H. Basedow, 1903, nos. 437, 116 (N.S.W.); Arkaringa Creek, R. Helms,
17.5.1891 (N.S.W.). WESTERN AUSTRALIA: Gnamma Paddock, Glenorn Station, N. T.
Burbidge, August, 1938 (W.A.); Herb., O. H. Sargent, no. 1410, rich red loam near
York, W.A. (MEL). NortHERN TERRITORY: Macdonnel Ranges, Strehlow, 1932-3
(S.A.95640011). QUEENSLAND: Queensland border, Cooper’s Creek, Gregory South,
J. More (Q.001877); Warrego River district, F. M. Bailey (N.S.W.); Parroo River
district, E. Betche, 9.1900 (N.S.W.).

PROCEEDINGS OF THE LINNEAN SocrrEtTy oF NEw SoutH WALES, 1958, Vol. Ixxxiii, Part 2.

BY R. C. CAROLIN. 93

ERODIUM CRINITUM, sp. Nov.

Herbeae perennes suberectae pilis simplicibus albis subrigidis et glandulosis
brevissimis instruntur. Folia tri-loba profunde vel palmata. Pedunculi graciles
hirsuti. Sepala oblonga anguste vel lanceolata pilis confertis sublaxis instruntur.
Filamenta staminorum 1/3-2/3-plo longiore staminodium. Pili longi appressi ponentur
trans mericarpium.

Sprawling or suberect herbaceous perennial with thick fleshy rootstock. Stems
1-numerous, 5—50 cm. tall with long stiff white simple hairs scattered over the surface.
Leaves 1-5-4:0 cm. long, 1:5-3-0 cm. wide, deeply palmately dissected with three
principal lobes; median lobe 1:5—4:0 cm. long, 1:0—2:5 em. wide; sinuses reaching to
midrib, long white simple hairs scattered over surface; margin toothed; petioles
slender with scattered simple hairs especially in groove, and very short glandular
hairs; stipules and bracts acute. Flowers in umbels of (2)—4—(6), very rarely solitary,
pedicels covered with long simple hairs. Sepals 5, lanceolate or narrow oblong, 0-6—1-:0
cm. long, 0-2-0-3 cm. wide, hirsute with long suberect white simple hairs and very
short glandular hairs where exposed in bud, shortly mucronate, ciliate and membranous
towards margin especially where covered in bud. JPetals 5, blue-purple with yellow
or white veins and base, elongate-ovate, c. 1:0 ecm. long and 0:5 em. wide. Stamens 5;
filaments oblanceolate or narrow oblong, 0-4 cm. long, 0:15 em. wide, acuminate-aristate
with an awn ec. 0-1 cm. long; anthers yellow; staminodes 5, ovate-triangular, 0:15 em.
long, 0-15 cm. wide, almost obtuse, occasionally toothed, 1/3-1/2 as long as staminal
filament. Stigmata 5, yellow. Mericarps 5, oblanceolate-obovoid, 0-65 ecm. long and
0-15 cm. wide, covered with long white simple hairs appressed diagonally to long axis;
beak 4-0-7:0 cm. long with two pits at base and often two folds beneath each pit.

Chromosome number: 2n = 40.

Range: Extra-tropical southern and eastern Australia, extending to the Musgrave
Ranges, Queensland, and rarely found in Western Australia.

Ecological distribution: Generally in open situations.

Holotype: Butler’s Peak Range, Fowler’s Gap. R. Carolin, June, 1956. No. H5. In
Herb. N.S.W.

The species is quite variable, although the variants grade imperceptibly into one
another. For this reason no subspecific taxa are recognized. The specimens from
Western Australia, where the species seems to be distinctly uncommon, tend to have
virtually glabrous pedicels and larger staminodes than the type. Towards the east
coast the leaves tend to be more dissected and the lobes somewhat narrower than in
the type. Near Adelaide and in the Murray “desert” the mericarp hairs tend to be
divergent and closely resemble those of #. cygnorum Nees in Lehm. ssp. cygnorum.

Specimens examined.—WESTERN AUSTRALIA: Main Camp Kurrawang, J. H. Maiden,
Sept., 1909 (N.S.W.); Toodgay, W.A., Oldfield (MEL). Soutmw Ausrraria: Ernabella
Station, H. E. Lord, 17.4.1950 (MEL); between Youldeh and Charlotte Waters, E. Giles
(MEL); Lake Letty, Crocker, 8.8.1939 (S.A.95640004); between Blinman and Wirrealpa,
east of Lake Torrens, Cleland, 1.12.1930 (S.A.95641032); Gawler Ranges, Lincoln Gap,
Cleland, 24.9.1942 (S.A.95641017); Flinders Range, Quorn, Cleland, 3.10.1945
(S.A.95641013); Flinders Range, Dr. Murray, Howitt Exped. (MEL); Mt. Lyndhurst,
Max Koch, Aug., 1898, no. 196 pro parte (Q@.001868); S. Flinders Range, Gladstone,
Black, 9.9.1932 (S.A.95641009); Koonamore Vegetation Reserve, Paltridge, 17.9.1928
(S.A.95640008); Koonamore, S.H. part of Vegetation Reserve, Hichler, 14.8.1956, no.
12452 (S.A.95641046); Koonamore, N.W. part of Vegetation Reserve, Hichler, 15.8.1956,
no. 12499 (S.A. 95641047); Gammon Ranges, Arcoona Bluff, Hichler, 15.9.1956, no. 12635
(S.A.95641048); Caroona, Cleland, June, 1885 (S.A.95641039); between Wintinna and
Marble Creek, J. M. Becharraise (MEL); Deep Creek, 5 miles east of Barra, Cleland,
24.8.1946 (S.A.95641029); Marino, Cleland, 10.9.1955 (S.A.95641025); Karoonda, Black,
5.10.1915 (S.A.95641009); Nonning, Cleland, 25.8.1928 (S.A.95641018); Nonning, Dr.
Pulleine, May, 1931 (S.A.95641045); Adelaide University grounds, Cleland, 5.9.1934
(S.A.95641012); near Adelaide, F. v. Miiller, Sept. 16, 1848, and Aug., 1848 (MEL);
Brighton, Black, 9.1904 (N.S.W.); Encounter Bay, near Callistemon Swamp beyond
Bluff, Cleland, 19.8.1947 (S.A.95641030); Hills above Encounter Bay, Cleland, 12.9.1945

94 SPECIES OF ERODIUM ENDEMIC TO AUSTRALIA,

(S.A.95641038) ; Adelaide-Melbourne, Kinchina, Cleland, 8.11.1924 (S.A.95641040); Byre’s
Peninsula, 30 miles west of Kimba, Cleland, 14.11.1955 (S.A.95641024); towards
Spencer’s Gulf, Warburton (MEL); Kangaroo Island, Waterhouse (MEL). NorrHern
TrRRiIToRY: Ayers Rock, R. Carolin, 21.6.1956, no. 76. Vicror1A: Nandooringa Creek,
Vict. Exploring Exped., Dec. 24.1860 (MEL); Shire of Dimboola, F. M. Reader, 26.8.1900
and 15.10.1887 (MEL); Grassy places around Port Phillip (MEL); Campaspe, Bathouse
(MEL). New SoutH WALES: Tibooburra, O. EH. Conch, 8.1930 (N.S.W.); W. of Nyngan,

Text-figs. 1-7. H. crinitwm Carol.—1, Flower bud; 2, Stamen filament; 3, Staminode;
4-5, Leaves; 6, Mericarp; 7, Hairs.

Text-figs. 8-14. H. awrewm Carol.—8, 10, Stamen filaments; 9-11, Staminodes; 12, Flower bud;
13, Mericarp; 14, Leaf.

I. Pidgeon and J. Vickery, 19.8.1939 (N.S.W.); Nyngan, J. L. Boorman, 8.03 (N.S.W.);
Sunny Corner, J. L. Boorman, 11.99 (N.S.W.); Tilpa, G. Turner, 5.1911 (N.S.W.);
Thuddingra, G. W. Caldwell, 16.12.1954 (N.S.W.); Boppy Mountain, near Cobar,
J. L. Boorman, 7.03 (N.S.W.); Narromine, J. H. Maiden, 9.98 (N.S.W.); Narrabri,
E. Breakwell, 9.13 (N.S.W.); Toonale-Goonery, J. L. Boorman, 10.1912 (N.S.W.);
Marsdens, Messrs. Bloomfield, 10.12 (N.S.W.); Gilgandra, J, D. Simon, 10.1913 (N.S.W.) ;
Gunnedah, Narr, 7.3.1841 (N.S.W.); Ardlethan, R. H. Cambage, 30.9.16 (N.S.W.) ;
Jerilderie, Bishop Dwyer, 10.1920 (N.S.W.); Waldoorah, Baracuba, R. D. Hay, Sept.,
1903 (N.S.W.); Canundia, J. Garden, 7.10.1949 (N.S.W.); Walgett, A. B. Little, 10.99

BY R. C. CAROLIN. 95

(N.S.W.); Culgoa River, T. S. Webb, 8.1901 (N.S.W.); Bingara, Miss P. M. Blundell,
5.1911 (N.S.W.); Boggabri, R. H. Cambage, 10.1912, no. 3624 (N.S.W.); Mugerie,
Coonamble, C: Featherstonehaugh, 7.1913 (N.S.W.); Plains of the Condamine, Timbe,
Dr. Lohb (Syd. Univ.); Coolabah, R. W. Peacock, 9.98 (N.S.W.); Fifield, R. H. Cambage,
9.1908, no. 1951 (N.S.W.); Mookoo, near Garah, R. H. Cambage, 19.2.22 (N.S.W.);
Condobolin, EH. Breakwell, Aug., 1913 (N.S.W.); 4 miles reserve, Condobolin, N. C.
Beadle, 8.45 (Syd. Univ.); Yanco Experimental Farm, H. Wenholz, 2.1913 (N.S.W.);
Yanco Experimental Farm, H. Breakwell, Nov., 1913 (N.S.W.); Temora, Rev. J. Dwyer,
915 (N.S.W.); Wentworth, Mrs. Forde (MEL); Sandhills near the Darling River,
Dr. Beckler (MEL); River Darling, Vict. Exploring Exped. (MEL); Darling River,
Mr. Neilson (MEL); east of Broken Hill, N. C. Forde, 31.8.46 (N.S.W.); Broken Hill,
A. Morris, 5.7.1920, 2506/r0 (N.S.W.); Ivanhoe, EK. H. Crisp, Rod. Herb. no. 11167
(N.S.W.); Hay, J. J. Fletcher, Sept. 22, 1889 (N.S.W.); Henty, E. J. McBarron, 15.2.1949,
no. 3079 (N.S.W.); Griffith, S. A. Blakely and D. W. Shiress, 7.1928 (N.S.W.); Railway
enclosure, Culcairn, EH. J. McBarron, 2.9.1949, no. 3542 (N.S.W.); Narrandera, EH. J.
MecBarron, 6.11.48, no. 2480 (N.S.W.); New England, A. Norton (Q.001878) ; Tamworth,
R. H. Goode, 10.11.1954, no. 95a (N.S.W.); Belltrees, near Scone, H. L. White, Feb., 1920,
no. 47 (N.S.W.); Scone, EH. Breakwell, 8.13 (N.S.W.); Dubbo, J. L. Boorman, 8.03
(N.S.W.); South Mudgee, M. Tindale, 6.10.1953 (N.S.W.); Pennant Hills, L. Fraser,
1934 (N.S.W.); Concord West, O. D. Evans, 27.10.1942 (Syd. Univ.); Arundale, Consett
Davis, 4.2.41, Rod. Herb. no. 12064 (N.S.W.); Nowra, Rod. Herb. (N.S.W.); Bega,
EH. Breakwell, Dec., 1913 (N.S.W.). QUEENSLAND: Charleville, Dr. A. J. Turner, 9.02
(Q.001871); Charleville, EH. W. Bichae, Dec., 1916 (Q.001878); Roma, R. E. Soutter,
Oct., 1924 (Q.001872); Bungewagorai, R. EH. Soutter, June, 1913 (Q.001867); Boatman
Stn., S. L. Everist, 21.7.1947, no. 3108 (Q.001869); Boatman Stn., S. L. Everist, 25.4.1947
(Q.001890); Hermitage, near Warwick, H. S. Pink, Sept., 1947 (Q.001870); Yelarbou,
€. T. White, 9.1919 (Q.001879); Darling Downs, about 3 miles south-east of Blaxland,
L. S. Smith and S. L. Everist, 13.10.1940, no. 797 (Q.001888); Darling Downs,
Goondiwindi, C. T. White, 28.9.1944, no. 12608 (Q.001887); Miles S2, EH. H. Belson,
10.1930 (Q.001886); Northampton Downs, near Blackall, S. L. Everist, 26.8.1935, no.
1216 (Q.001884); Mitchell district, Blackall, S. L. Everist, 24.8.1955, no. 1228 (Q.001889).

ERopIuM cygNnorum Nees in Lehm., Plantae Preissianae, 1: 162 (1844).
Sprawling or suberect herbaceous perennial with a thick fleshy tap-root. Stems
i-numerous, 5-60 cm. tall, covered with stiff white simple hairs or soft glandular
hairs. Leaves 1:5-6:0 cm. long, 0:5—4:0 ecm. wide with three principal lobes and simple
or glandular hairs scattered over the surface; petioles slender, 2-13 cm. long; stipules
and bracts acute. Flowers in umbels of 2-6, rarely solitary; pedicels almost glabrous
or covered with soft glandular hairs. Sepals 5, oblong-lanceolate or oblong-elliptic,
0-5-1-3 cm. long, 0:2-0:-7 em. wide, covered with short white simple hairs or soft
glandular hairs, mucronate, ciliate and somewhat membranous towards the margin,
especially where covered in bud.- Petals 5, blue-purple with red, yellow or white veins
and base, ovate, 1:0-1:2 cm. long, 0:5-0-7 em. wide. Stamens 5; filaments oblanceolate
to lanceolate, 0-4 cm. long, 0:15 em. wide, occasionally toothed; anthers red or yellow;
staminodes 5, oblong-lanceolate to lanceolate, 0:25-0:35 cm. long, 0:1-0:15 em. wide.
Carpels 5, hairy; stigmata 5, red or green. Mericarps 5, obovoid to oblong, c. 1:0 ecm.
long, 0-15-0-2 cm. wide, covered with long simple divergent or erect hairs; beak

5-0-12-:0 cm. long, with two pits at base and 1-2 folds often beneath each pit.

Ssp. CYGNOGUM.
Herbeae perennes suberectae simplicibus pilis albis rigidis instruntur. Folia
' triloba profunde plerumque ad nervam primam. Pedunculi graciles basi glabri hirsuti

summis. Sepala oblonga vel oblongo-lanceolata pilis brevissimis rigidis confertis
appressis instruntur. Filamenta staminorum oblanceolata anguste acuminata, 11/2-plo
longiore staminodia. Staminodia lanceolata. Mericarpium pilis rigidis longis

divergentibus et interdum brevioribus aliquot obtegitur.
Stems 1-numerous, 5-35 cm. tall with stiff white simple hairs scattered over surface.
Leaves 1:5-4:0 cm. long, 0-5-3:0 cm. wide, deeply dissected with three principal lobes;

96 SPECIES OF FRODIUM ENDEMIC TO AUSTRALIA,

-median lobe 1:5-4:0 cm. long, 0:3-2:0 cm. wide; sinuses reaching to main vein, white
simple hairs scattered over surface; petioles slender, c. 2:0 cm. long with scattered
white simple hairs, especially in the groove, and very minute glandular hairs; stipules
and bracts acute. Flowers in umbels of (2)-3-(5) rarely solitary; pedicels almost
completely glabrous but with numbers of short white simple hairs immediately below
calyx. Sepals 5, oblong or oblong-lanceolate, (0-5)0-7-0-°9 ecm. long, (0-2)0-3-0-4 cm.

Text-figs. 15-20. H. cygnorum Nees in Lehm. ssp. cygnerum.—-15, Flower bud; 16, Mericarp ;
17, Leaf; 18, Hairs; 19, Staminode; 20, Stamen filament.

Text-figs. 21-26. H. cygnorum ssp. glandulosum Carol.—21, Flower bud; 22, Mericarp ;
23, Leaf; 24, Stamen filament; 25, Staminode; 26, Hairs.

wide, covered with short appressed white simple hairs, mucronate, ciliate and
membranous towards margin, especially where covered in bud. Petals blue-purple with
yellow or white veins and base, ovate, c. 1:0 cm. long and 0:5 ecm. wide. Stamens 5;
filaments oblanceolate or narrow oblong, 9-4 em. long, 0:15 em. wide, acuminate-aristate;
anthers yellow; staminodes lanceolate, 0:25 cm. long, 0-1 cm. wide, 2/3 times as long
as staminal filaments. Mericarp obovoid, (0:6)0:8-1:0 em. long, 0:15-0:-2 cm. wide,

BY R. C. CAROLIN. 97

covered (usually sparsely) with long stiff spreading hairs often with a complement of
shorter hairs, densely hairy at base where hairs are erect; beak 5:0-10-0 cm. long.

Chromosome number: 2n = 60.

Range: Mostly extra-tropical Western Australia as far east as Lake Gilles in
South Australia and extending northwards to Onslow in the tropics and Ayers Rock.

Ecological distribution: Generally in open situations and possibly showing a
preference for sandy soils.

Holotype: In depressis arenosis umbrosis ad flumen Cygnorum supra oppidulum
Perth, Augusto 1839. Preiss no. 1902.

It has been impossible to trace this specimen, although enquiries have been made
at the following European herbaria in addition to the Australian institutes (the
abbreviations are those recommended in the current “Index Herbariorum’’): B, BM,
FI, G, GB, GOET, HBG, K, KIEL, LE, NBV, P, U, UPS, W, WU. It was possibly
housed at Berlin-Dahlem and was destroyed during the 1939-46 war. The description
is hardly adequate to decide to which taxon the type belonged. ‘‘sepalo .. . tota
superficie pilis brevissimis” and the locality from which it was taken indicate that
the taxon described above is probably the one concerned. It is fairly certain that the
type is lost and due to the inadequacy of the type description it is necessary to erect
a neotype. Unfortunately none of the specimens from the Swan River which were
' examined were in any way complete and a specimen from another locality has had
to be selected.

Neotype: 3 miles S. of Morawa, C. A. Gardner, 26 Aug., 1945, no. 7526. In
Herb. W.A.

Specimens examined.—WESTERN AUSTRALIA: Lakeside, ex Herb., W. V. Fitzgerald,
Aug., 1898 (N.S.W.); S. of Onslow, C. A. Gardner, Aug., 1932, no. 3063 (W.A.);
Murchison River and north of Murchison, Oldfield, pro parte (MEL); north of
Murchison, Oldfield (MEL); Tamala Stn., Hamelin Pool, C. H. Roberts, Aug., 1937
(W.A.); Gascoyne River, near Jimba-Jimba, C. A. Gardner, 1941 (W.A.); Yorkakrine,
C. A. Gardner, Sept., 1920 (W.A.); Yorkakrine, near Tammin, C. A. Gardner, Aug.,
1920 (W.A.); Merredin, C. A. Gardner, Aug., 1920 (W.A.); 30 miles east of Meekathera,
C. A. Gardner, July, 1931, no. 2374 (W.A.); Yandanooka, R. Morrison, Sept., 1904
(W.A.); Goomalling, C. A. Gardner, Aug., 1920, no. 648 (W.A.); Sturt Meadows Stn.,
C. H. Brockway, Oct., 1943 (W.A.); Mt. Magnet, W. S. Macpherson, 1897 (MEL); west
from Middle Mount Barren (MEL); Stirling Ranges (MEL); Basalt Ridge north of
Stirling Ranges, F. v. Mueller (?) (MEL); Mount Morgan §., Dr. H. J. Mitchell,
8.1913 (N.S.W.); Comet Vale, J. T. Jackson, 12.1916 (N.S.W.); Laverton, Maiden,
1909 (N.S.W.); banks of the Salt River,Max, no. 94 (MEL); Tone River, Oldfield,
no. 122 (MEL); Glenorn Stn., N. T. Burbidge, Aug., 1938 (W.A.); Coolgardie, A. J.
Vaga, 1896 (N.S.W.); Kurrawang, J. Burton Cleland, 5.1907 (N.S.W.). SouTH
AUSTRALIA: Ooldea Soak, Cleland, 22.8.1939 (S.A.); Ooldea, EH. H. Ising, 13.9.1920,
no. 1612 (N.S.W.); Tarcoola, E. H. Ising, 5.9.1920, no. 1191 (Q.001873); Lake Gairdner,
Babb. Exped. (MEL); Lake Giles, S. Burkitt (MEL). NorTHERN TeErRRITORY: Ayers
Rock, Maggies Spring, G. Chippendale (N.S.W.).

Ssp. GLANDULOSUM, ssp. Nov.

Herbeae perennes pilis glandulosis confertis instruntur. Folia triloba autem
numquam ad nervam primam. Pedunculi sepalaque oblongo-ellipticae pilis glandulosis
obteguntur. Staminodia aeque filamentis staminorum. Pili longi appressi obtegunt
dense mericarpium.

Stems 1-numerous, 10-60 cm. tall, covered with soft giandular hairs. Leaves 2:0
em. long, 1:0-4:0 cm. wide, lobed with three crenate principal lobes, sinuses never
reaching midrib, hairs almost all glandular on both surfaces; petioles slender, 2-13 cm.
long, glandular hairy, except in groove which has long stiff simple hairs; stipules and
bracts acute. Flowers in umbels of 2-6, very rarely solitary; pedicels slender, glandular
hairy. Sepals 5, oblong-elliptic, (0-6)0-8-1:3 cm. long, (0-3)0:-4-0:-7 em. wide, covered
with soft glandular hairs and very few or no simple hairs, membranous and ciliate
towards margin, especially where covered in bud. Petals 5, blue-purple with red veins

98 SPECIES OF ERODIUM ENDEMIC TO AUSTRALIA,

and base, ovate, c. 1:2 em. long and 0:7 cm. wide. Stamens 5; filaments lanceolate
or narrow oblong, 0:4 cm. long, 0:15 cm. wide, acute, occasionally toothed; anthers red;
staminodes 5, oblong-lanceolate, 0:35 cm. long, 0:15 cm. wide. Stigmata 5, red. Mericarps
5, oblanceolate or oblong, c. 1:0 cm. long and 0-2 cm. wide, covered with long white-
brown simple hairs appressed for the most part parallel to the long axis; beak
6:0-10:0(12:0) cm. long.

Chromosome number: 2n = 60.

Range: Mostly drier parts of extra-tropical South Australia extending to just north
of Alice Springs, westwards to Tarcoola and eastwards to the Darling River.

Ecological distribution: Generally in open situations.

Holotype: Butler’s Peak Range near Fowler’s Gap. Damp col in the ranges, 1200 ft-
R. Carolin, June, 1956, no. H4. In Herb. N.S.W.

Text-fig. 27. Distribution of H. aurewm Carol. @® and #H. crinitum Carol. x.

Specimens examined: SoutH AuSTRALIA: Mt. Davis, border of S.A., N.T., and W.A.,
Mrs. A. T. Reid, 9.1955 (N.S.W.) (see notes); Anna Creek, William Creek, Cleland,
10.9.1980 (S.A.95641027); Warburton River, Cowarie Stn., Crocker, 24.7.1939
(S.A.95640003); Arcoona, Murray, 9.8.1927, no. 83 (S.A.95640002) ; Cootanoorinna Creek,
Helms, 7.5.1891 (S.A.95640010); Railway line, Alice Springs to Adelaide, Irrapatanna,
Cleland, 25.8.32 (S.A.95641033); Curdimurka, Knox College Exped. no. 4, 1.9.1950
(N.S.W.); Leigh Creek, Dr. R. S. Rogers, 9.10 (N.S.W.); Lake Eyre, Prof. Baldwin
Spencer, 9.03 (N.S.W.); Alice Springs to Marree, Melbourne Ward, Aug., 1947 (Syd.
Univ.); Curnamona Stn., M. Pinches, 1931 (S.A.95641031); Gammon Ranges, Arcoona
Pound, c. 6 miles east of Ouianduina Outstn., Hichler, no. 12637 (S.A.95641049) ;
Flinders Range, near the road Hawker-Blinman, c. 4 miles after Wilpena, Hichler,
11.9.1956, no. 125387 (S.A.95641050); Mt. Lyndhurst, Max Koch, June, 1899, no. 196,
pro parte (MEL); Koonamore, T. G. B. Osborn, 5.1928 (Syd. Univ.); N. of Fowler’s
Bay, E. Giles, 1875 (MEL). New SoutH WALES: Broken Hill district, A. Morris, 28.8.21
(N.S.W.); Broken Hill, I. Pidgeon and J. Vickery, 23.8.1939 (N.S.W.); Broken Hill-
Menindie road, R. J. Constable, 19.7.1955 (N.S.W.); Broken Hill, E. C. Andrews, 9.1918
(N.S.W.); Thompson Siding, Broken Hill, A. Morris, 19.8.1928, no. 2373 (Q.001883) ;
Yanco Glen, P. Brough and N. C. Beadle, 26.8.1939 (Syd. Univ.); Umberumberka,
L. A. S. Johnson, 29.8.1946 (N.S.W.).

BY R. C. CAROLIN. 99

These two subspecies appear to be geographic differentiates of one species despite
their very distinctive characters. In the regions where their distributions abut or
overlap, ie., west of Lake Eyre in particular, the distinction becomes a little blurred.
Most of the specimens from this region are referred to ssp. glandulosum Carol. but
the hairs on the mericarps resemble those of ssp. cygnorum very closely. The specimen
“Mt. Davies, Mrs. A. T. Reid, 9.1955 (N.S.W.)” in particular shows a mixture of the
characters of the two ssp. The indumentum is almost wholly glandular and the
pedicels are hairy, whereas the leaves are dissected to the midrib and the mericarp
hairs are divergent and scanty. The two ssp. have been crossed and apparently viable
seed has been produced but, unfortunately, the seeds died before flowering. The
specimen from Mt. Davies mentioned above has well-formed seeds on it and it would
appear that the hybrid is viable.

Text-fig. 28. Distribution of £H. cygnorum Nees in Lehm. ssp. cygnorum x and
ssp. glandulosum Carol. e.

All these taxa have been grown in Sydney. FH. crinitum Carol. alone produced
good growth and this species is found locally native. The others retained their
characteristics. Crosses were attempted between them but apart from the results
mentioned above no hybrid seeds were obtained. No intermediates between the species
have been observed in the herbarium specimens examined and, indeed, should they
occur they would in general be sterile, assuming that the chromosome numbers
reported here are constant. b

These taxa probably represent a fairly discrete unit within the genus. A relation-
ship with H. malacoides Willd. may exist; this species extends into South Africa
but it appears to be an introduction there and it is essentially a European species.
The taxonomic position, then, must remain indefinite for the time being.

The diploid species is found in the arid areas, which suggests that, if the group
has a common origin in Australia, the genus entered Australia from an arid region
at a time when parts of the continent at least were arid. The apparent absence of
any of the species described above from the more northerly part of the arid area
would indicate that the genus may have entered the continent when the north was
somewhat cooler or from another direction altogether. Long-range dispersal of arid
plants over more clement regions is not likely. The polyploid species, particularly the

100 SPECIES OF ERODIUM ENDEMIC TO AUSTRALIA.

tetraploid E. crinitwm Carol., are more tolerant to differing ecological conditions than
E. aureum Carol. and are rather more abundant in the damper southern areas of the
continent, although they do occur in the arid areas. The hexaploid H. cygnorum Nees
in Lehm., at least, evolved prior to the last fleristic separation of the south-west from
the rest of the continent when two subspecies differentiated on either side of the
barrier. The aridity of mid-Recent times may have been this barrier, as these species
eannot exist under extremely arid conditions.

Nomen DvusiuM.—EH. PERISTEROIDES Turez. in Bull. Soc. Imp. Nat. Mosc., 36: 592
(1863).

Holotype: In Nova Zeelandia vidi specimina plurima in collectionibus All.
Cunninghamii et Hv. Homei. ;

Knuth in “Das Pflanzenreich” cites this name as a synonym of H. cygnorum Nees
in Lehm. There is a good deal of confusion about this name. _The Manual of the
New Zealand Flora by Cheesman does not mention either as occurring in that country
and it does not appear to have been reported. Without a specimen definitely named
by Turczaninow it would seem impossible to draw any conclusions with regard to type
specimen(s). No such specimen has been found in Leningrad, Moscow, Kiev, Kew or
London. Dr. R. Melville has suggested that the country of origin cited for
E. peristeroides Turcz. is an error and that it is possible that the description applies
to a Pelargonium. The ‘“sepala acuta ... Petalo primo alba dein rosea, unguiculata
subspathulata ... ovarium glabrum” preciude this specimen from all the indigenous
or introduced Australian species of Hrodiwm and certainly indicate Pelargonium as
the source of the material.

Key to the Australian Species of Hrodium L’ilér. Including Introduced Taxa.

1. Leaves pinnate-compound.
2. Leafiets very deeply pinnately lobed (often cut to the main vein) ....................
NEM OHO- OOo tn ORO GG CuO LG HHO eL Ta OA SI BeOTEe a onc cua thee EH, cicutarium (L.) L’Hér. ex Ait.
2§. Leaflets toothed or lobed but rarely dissected more than halfway to the main vein
SWS CH eCR OR ERO TST mOea eres ioe CREA AD OKCY SME NCEA. ESSER BOOS EE CeuICRES E. moschatum (lL.) Ait. introd.
1§. Leaves palmate or ternate compound or lobed or pinnate lobed.
3. Leaves very slightly lobed, ovate; beak of mericarp 2-3 ecm. long ....................
hs Gt (ovseuts eftahrowtce/jevfeccaes udaaisevteine Monon savas a8 tousabeWepen aver aten Seavowene austenite wiapcueme nie R H. malacoides Willd. introd.
3§. Leaves deeply lobed, dissected or compound; beak of mericarp more than 3 cm. long.
4. Leaves pinnate-lobed, basal lobes about the same size as subsequent lobes; staminodes
less than half as wide as staminal filaments ...... EH. botrys (Cav.) Bertol. introd.
4§. Leaves palmate-ternate, rarely pinnately, lobed with basal lobes larger than subsequent
ones; if leaves pinnately lobed staminodes as wide as staminal filaments.
5. All calyx hairs simple except the most minute ones (invisible except under high
magnification) ; leaves deeply dissected.
6. Hairs of calyx scarcely appressed, long, staminal filaments 3-2 times as long as
staminodes; mericarp hairs appressed diagonally ............ EH. crinitum Carol.
6§. Hairs of calyx short, appressed; stamina] filaments one and a half times or equally
as long as staminodes; mericarp hairs divergent .................02. cee eeeeeeee
LH ead a ctral arte) eee nsee ems, Gh eb au weenie Ge SUM eas ent eroneles EH. cygnorum Nees in Lehm. ssp. cygnorum.
5§. Visible calyx hairs glandular often mixed with some simple hairs or all glandular;
leaves lobed.
7. Staminal filaments oblong, acuminate; sepals almost devoid of simple hairs; mericarp
hairs erect or slightly divergent ........... EH. cygnorum ssp. glandulosum Carol.
7§. Staminal filaments lanceolate, aristate; sepals covered with both simple and glandular
hairs (former often towards apex); mericarp hairs appressed diagonally .......
sho abe necypeenci hs oP 2 debian d rie Pe hey tank ee aut Bagh A eee 8 ok oe ae ere map aan ie EF. aureum Carol.

The Government Botanists of each State have helped considerably by the prompt
and protracted loan of specimens. Dr. R. Melville, of Royal Botanic Gardens, Kew, has
kindly sent me descriptions and notes which were unobtainable in Australia, and
Mrs. M. Thompson, of the National Herbarium of N.S.W., Sydney, has read the MS.
and made a number of suggestions. A grant from the Research Committee of the
University of Sydney enabled some of the material to be collected.

101

CATALOGUE OF AUSTRALIAN MAMMALS AND THEIR RHCORDED
INTERNAL PARASITHS. LIV.

Part I. MONOTREMES AND MARSUPIALS (pp. 101-125). Parr II. HuTHERIA (pp. 126-143).
Part III. IntRopUCED HEeRBIVORA AND THE DomMESTIC Pic (pp. 143-153).
Part IV. MAN (pp. 153-160).
By M. JosEPHINE MACKERRAS, Queensland Institute of Medical Research, Brisbane.
(Communicated by Dr. I. M. Mackerras.)
[Read 30th July, 1958.]

PART I. MONOTREMES AND MARSUPIALS.

Synopsis.
Part I contains the names of three monotremes and 158 marsupials. Parasites have been
recorded from all the monotremes and from 72 marsupials. They have been found in

representatives of all the families except the Notoryctidae, which have probably not been
examined.

Blood protozoa are known from two monotremes (Trypanosoma and Theileria) and
from six marsupials (Trypanosoma, Theileria, and Haemogregarina (sens. lat.)).

Two trematodes are known from the platypus and five from polyprotodont marsupials.
No trematodes have been recorded from diprotodonts except, occasionally, Fasciola hepatica
(an introduced parasite).

Two adult ces.iodes are known from echidnas, five from polyprotodonts, and twenty-one
from diprotodonts. The majority of the cestodes belong to the Anoplocephalidae. Hydatids
(introduced with domestic animals) occur in wallabies and kangaroos.

Nematodes are known from an echidna and from 55 marsupials. Of about 156 described
species nearly 80% belong to the Strongyloidea. No unequivocal record of an adult member
of the Ascaridoidea has been made.

One species of Acanthocephala occurs in bandicoots.

It is desirable at times to take stock of the present position and to summarize
what is known of any particular subject. In this way the limitations of our knowledge
become clearer and new lines of attack may appear. Many years ago the late Professor
T. Harvey Johnston began to list the parasites of Australian animals. His 1916
census embraced ali vertebrate and invertebrate hosts. Today, largely as a result of
his own and his colleagues’ efforts, such a comprehensive list would require many
more than the 34 pages of print it occupied in 1916.

In the following four paris an attempt has been made to bring part of the
information up to date in readily accessible form, by recording the protozoan, helminth,
and pentastomid parasites known to occur in mammals in Australia. It is hoped
that these lists will provide a useful stepping-off point for young parasitologists. It
is hoped, too, that the publication of the names of very numerous species, from which
no parasites seem to have been recorded, may draw attention to the large gaps in our
knowledge and stimulate workers in other fields of animal study to preserve all
parasites which they may find, and also to submit blood films and unwanted
careasses to parasitologists, whenever it may be feasible to do so. Many strange and
unique forms have already been found in our indigenous fauna, but undoubtedly many
more await discovery.

The classification and synonymy given by Iredale and Troughton (1934)* and by
Troughton (1954) have been used as a basis for Parts I and II, but some alterations

* A Check-list of the Mammals Recerded from Australia, by Tom Iredale and Hi. Le G.
Troughton, Australian Museum Memoir VI. Sydney, 1934.

i Furred Animals of Australia, by Ellis Troughton. Angus and Robertson, Sydney, 5th ed.,
1954, 376 pp.

PROCEEDINGS OF THE LINNEAN SocieTY OF NEw SouTH WALES, 1958, Vol. Ixxxiii, Part 2.

B

102 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

have been made. A few records from non-indigenous animals, chiefly from islands north
of Australia, are included, because the same parasites may occur in mainland species.

Although care has been taken to search the literature and to check all references,
errors and omissions will undoubtedly be found, and the author would be grateful
to have attention drawn to them. An attempt has been made to make the bibliography
to Part I as complete as possible, but this has not been feasible in Parts II to IV.
A selection has therefore been made of those articles dealing with host-parasite records
(with due regard for synonymy), life histories, or epidemiology.

ARRANGEMENT OF PARASITES.
Abbreviations Used.

Long lists of names are confusing, so the parasites are arranged in a definite
order and an indication of their systematic position is given by abbreviations placed
before the name. The Protozoa are arranged in the following way: Mastigophora (M.),
Sarecodina (Sa.), Sporozoa (Sp.), and Ciliata (C.).

All the Trematoda mentioned belong to the Digenea Prosostomata, the following
families being represented: Paramphistomatidae (Param.), Cathaemasiidae (Catha.),
Dicrocoeliidae (Dicro.), Diplostomatidae (Diplo.), Fasciolidae (Fasci.), Heterophyidae
(Heter.), Microphallidae (Micro.), Notocotylidae (Notoe.), Opisthorchiidae (Opist.),
Plagiorchiidae (Plagi.), Pronocephalidae (Prono.), Rhabdiopoeidae (Rhabd.), Schisto-
somatidae (Schis.), and Strigeidae (Strig.).

Cestoda: Order Cyclophyllidea (CY.), with five families: Anoplocephalidae (Ano.),
Davaineidae (Dav.), Dilepididae (Dil.), Hymenolepididae (Hym.), and Taeniidae
(Tae.); Order Pseudophyllidea (FS.) with one family: Diphyllobothriidae (Dip.);
Order Tetraphyllidea (TH.) with one family: Phyllobothriidae (Phy.).

Nematoda: Superfamilies Rhabdiasoidea (RH.), Trichuroidea (TR.), Strongyloidea
(ST.), Oxyuroidea (OX.), Ascaridoidea (AS.), Spiruroidea (SP.), and Filarioidea (FI.).
The families of the Strongyloidea are indicated as follows: Trichostrongylidae (Tri.),
Aneylostomidae (Anc.), Strongylidae (Str.), and Metastrongylidae (Met.).

For the sake of brevity, the ietters J. and M. are used for T. H. Johnston and
P. M. Mawson; and Y. and M. for W. Yorke and P. A. Maplestone.

The authority for each parasite record is given after the author’s name and date.
Numbers in brackets refer to the numbered list of references. The seauence is
chronological, so the numbers are frequently not in serial order.

PARASITES OF MONOTREMES AND MARSUPIALS.
Histery of Discovery.

The first parasites of a marsupial to attract attention were the tapeworm inhabiting
the bile ducts of the grey kangaroo and the large filarial worm which infests the
bursae and tendon sheaths around the knee joint, 2nd even the joint itself, of the
same animal. The tapeworm was briefly described as Taenia festiva by Rudolphi in
1819 from a specimen in the Vienna Museum.

Two specimens of the filaria were recorded in the 1830 catalogue of the Royal
College of Surgeons. It is possible that these specimens were part of the original
collection of John Hunter, but proof of this is lacking. They are mentioned by
Dr. George Bennett in a letter to Sir Richard Owen, dated Sydney, February 4th,
1833. Bennett said he was sending a collection of “Ascarides taken from the inner
part of the knee joint of both legs in a large male kangaroo (of the common species)”’.
The same author referred to them in his book (Bennett, 1834), remarking that the
worm was recorded in the Catalogue of the Royal College of Surgeons as Filaria
macropi Mmajoris.

Cobbold (1879) mentioned these worms, “. . . which have been indicated as
Filariae macropodis gigantei”’, continuing: “It would, in my opinion, be far better
to call the worm after its discoverer, Webster’s filaria (fF. Websteri).’ Efforts to trace
Webster’s original discovery have failed. Hven the man’s initials do not seem to be
recorded. It seems probable that the discovery was made in London at some time
between 1791, when it is known that kangaroos were sent to Hngland, and 1830.

BY M. JOSEPHINE MACKERRAS. 103

Froriep* evidently confused Webster and Bennett, because the quotation said to
come from “Webster’s Wanderings” is word for word from Bennett’s “Wanderings in
New South Wales, etc.” except that, where Benneit used the prenoun “I”, Froriep
wrote “Herr Webster”. This reference to Webster may therefore be disregarded.

Hisig (1869) described, but did not name, a filarial worm, which has not been
rediscovered, from the pericardium of a wallaby which died in the Heidelberg Zoo.
Leidy (1875) described Filaria spelaea from the coelome of a wallaby which died in
the Philadelphia Zoo. Krefft (1871) studied the tapeworms of lecal birds and mammals
in New South Wales, but the complexity of the group was not appreciated at the time,
and his descriptions are too vague for recognition. Fortunately, T. H. Johnston
(1912) was able to rehabilitate many of Krefft’s species of bird tapeworms, but those
recorded from marsupials remain unrecognizable.

Cobbold (1879) gave provisional names to a tapeworm from the echidna and
one from the native bear, as well as changing the name of the kangaroo filaria to
Filaria websteri. However, as he did not describe any of the forms mentioned, his
names have been regarded as nomina nuda, although there is no real doubt as to the
identity of F. websteri. Lumholtz (1884) recorded the occurrence of worms lying
between the skin and muscles of the tree kangaroo, and implied that they were common
in other marsupials. These parasites of the tree kangaroo have not been rediscovered.

Collections of indigenous parasites were made by medical men, veterinarians and
others, notably by J. P. Hill, N. A. Cobb, W. H. D. Le Souef, G. Masters, and L. Gallard
in New South Wales, J. and T. L. Bancroft and the explorer R. Semon in Queensland,
during the latter part of the nineteenth and early twentieth centuries. Systematic
work was at first undertaken in Hurope by professional parasitologists, to whom
collections were sent. Thus, von Linstow in Germany, Zschokke in Switzerland,
Nybelin in Sweden, Beddard, Baylis, Yorke, and Maplestone in Hngland, named many
species. Following the pioneering work of Leidy and Wisig, parasitologists in various
parts of the world have been interested in the entozoa of exotic animals dying in
their local zoological gardens. Among these, Ortlepp, Wood, and Cameron in Britain,
Canavan, Chandler, and Schwartz in U.S.A., and Monnig in South Africa, have made
important contributions to the study of the parasitic fauna of Australian marsupials.

Harly in the present century workers in Australia began to play an increasing
part in recording the parasites found and in describing new species. Among these are
J. B. Cleland, T. H. Johnston, S. J. Johnston, J. A. Gilruth, W. Nicoll, A. Breinl,
D. A. Welsh, and H. Priestley. T. H. Johnston and his colleagues in Adelaide made
the first really planned attack on the problem. Recently D. F. Sandars has revised
the trematodes and cestodes, and compared them with those of American marsupials.
Valuable check lists have been published from time to time, for example by Sweet
(1909) in Melbourne, T. H. Johnston (1909 to 1918) in Sydney and Brisbane, Johnston
and Cleland (1912, 1937) in Sydney and Adelaide, Oldham (1933) and Young (1939)
in London. Skrjabin et al. (1954) listed many nematodes of monotremes and
marsupials under their hosts (“Key to Parasitic Nematodes”, vol. 4, pp. 733-742).

Practically no work has been done on blood parasites. Trypanosomes, haemo-
eregarines and Theileria are known to occur, but no Plasmodiwm has yet been found.
Some remarkable trematodes and cestodes are known, but the nematodes have received
most attention, and’ one of the most striking features of the marsupial record is the
immense development of the subfamily Trichoneminae of the family Strongylidae,
14 genera} having been erected to accommodate these parasites.

* Froriep, L. F. von (1834): Notizen aus dem Gebiete der Natur-und Heilkunde, vol. 42,
p. 328.
+ These are: Cloacina v. Linstow, Coronostrongylus J. & M., Cyclostrongylus J. & M.,
Labiostrongylus Y. & M., Macropostrongylus Y. & M., Maplestonema J. & M., Parazoniolaimus
J. & M., Paramacropostrongylus J. & M., Papillostrongylus J. & M., Pharyngosirongylus
Y. & M., Phascolostrongylus Canavan, Potorostrongylus J. & M., Spirostrongylus Y. & M.,

and Zoniolaimus Cobb.

104 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

Hosts AND PARASITES.
Class MAMMALIA
Order MonoTREMATA
Family ORNITHORHYNCHIDAE
Genus ORNITHORHYNCHUS Blumenback, 1800
O. ANATINUS (Shaw and Nodder, 1799) (The platypus)

Protozoa
(M.) Trypanvsoma sp. (36)
(Sp.) Theileria sp. (36)
Trematoda

(Catha. ) Mehlisia ornithorhynchi (S. J. Johnston, 1901) (45)
(? Fam.) Moreauia mirabilis S. J. Johnston, 1915 (48)

Family TAcHYGLOSSIDAE
Genus TAcHYGLOSSUS Illiger, 1811. (Hchidnas)
T. ACULEATUS (Shaw and Nodder, 1792)
Protozoa
(Sp.) Theileria tachyglossi Priestley, 1915 (104)
Cestoda
(CY., Ano.) Linstowia echidnae (Thompson, 1893) (117), (135)
Cittotaenia tachyglosst T. H. Johnston, 1913 (53)
(CY.) Taenia phoptica Cobbolid, 1879 (a nomen nudum) (32)
Nematoda
(ST., Tri.) Nicollina tachyalossi (Baylis, 1930) (7), (8)
N. echidnae (Baylis, 1930) (7), (8)
(¥#T.) Dipetalonema sp. (subcutaneous) (90)
T. SETOSUS (Geoffroy, 1803)
Cestoda
(CY., Ano.) Linstowia echidnae (Thompson, 1893) (74)

Order MARSUPIALIA
Suborder POLYPROTODONTIA
Family DASYURIDAE
Subfamily PHASCOGALINAE
Genus ANTECHINUS Macleay, 1841. (Marsupial mice)

A. APICALIS (Gray, 1842) No records
A. BELLUS (Thomas, 1904) No records
A. FLAVIPES (Waterhouse, 1838)
Nematoda

(ST., Met.) Plectostrongylus fragilis Mackerras and Sandars, 1953 (91)
A. GODMANI (Thomas, 1923) No records
A. MACDONNELLENSIS (Spencer, 1896) No records
A. MACULATUS (Gould, 1851) No records
A. MIMULUS (Thomas, 1906) No records
A. minimus (Geoffroy, 1803) No records
A. SwAInsonit (Waterhouse, 1840) No records

Genus PLANIGALE Troughton, 1928. (Flat-skulled marsupial mice)

Pu. INGRAMI (Thomas, 1906) No records
PL. SUBTILISSIMA (Lonnberg, 1913) : “No records
Pu. TENUIROSTRIS Troughton, 1928 No records

Genus PHAScoGALE Temminck, 1824. (Brush-tailed marsupial rats)
PH. caLurA Gould, 1844 No records
PH. TAPOATAFA (Meyer, 1793)
syn. penicillata Shaw, 1800
Two parasites from “brush-tailed rats’ may belong here.

NDNANNN

ANNNNN

7:

BY M. JOSEPHINE MACKERRAS. 105

Nematoda
(SP.) Denticulospirura dentata J. & M., 1941 (69), (70)
Acanthocephala
Gigantorhynchus sp. (50)

Genus DaAsycrrcus Peters, 1875. (Crested-tailed marsupial mice)
BLYTHI (Waite, 1904) No records
CRISTICAUDA (Krefft, 1867)

Nematoda
(SP.) Physaloptera sp. (131)

Genus DAsyuROImES Spencer, 1896. (Marsupial rats)

. BYRNEI Spencer, 1896 No records

Genus SMINTHOPSIS Thomas, 1887. (Marsupial mice)

. CRASSICAUDATA (Gould, 1844) No records
. FROGGATTI (Ramsay, 1887) No records
. GRANULIPES Troughton, 1932 No records
. HIRTIPES Thomas, 1898 No records
. LARAPINTA Spencer, 1896
Nematoda
(SP.) Rictulariidae (undescribed sp. collected by Dr. D. F. Sandars)

. LEUCOPUS (Gray, 1842) No records
. LONGICAUDATA Spencer, 1909 No records
. LUMHOLTZI Iredale & Troughton, 1934 No records
. MURINA (Waterhouse, 1838) No records
. PSAMMOPHILA Spencer, 1895 No records
. STALKERI Thomas, 1906 No records

Genus ANTECHTNOMYS Krefft, 1867. (Jerboa marsupials)
LANIGER (Gould, 1856) No records
SPENCERI Thomas, 1906 No records

Subfamily DASYURINAE
Genus DaAsyurus Geoffroy, 1796. (Native cats)

QUOLL (Zimmermann, 1777)
syn. viverrinus Shaw, 1800

Protozoa
(Sp.) Haemogregarina dasyuri Welsh, Dalyell & Burfitt, 1909 (122), (123)

= Hepatozoon dasyuri (Welsh, Dalyell & Burfitt) Wenyon, 1926 (126)
Toxoplasma sp. (113)

Trematoda
(Dicro.) Brachylaemus dasyuri (S. J. Johnston, i913) (47), (110D)
(Catha.) Mehlisia acuminata S. J. Jchuston, 1913 (47)

Cestoda
(PS., Dip.) Sparganum (49)

Nematoda
(SP.) Echinonema cinctum (v. Linstow, 1898) (52) (1 specimen)
(FI.) Dipetalonema dasyuri J. & M., 1938 (58)

DASYURUS SP. (Cairns, N.Q.)

Nematoda

(SP.) Spirocerca heydoni Baylis, 1927 (5)
Genus DaAsyurtnus Matschie, 1916. (Native cat)

GEOFFROYII (Gould, 1841) No records

D.

S.

Genus SATANELLUS Pocock, 1926
HALLUCATUS (Gould, 1842), the little northern native cat
Protozoa
(Sp.) Sarcocystis sp. (90)

106 AUSTRALIAN. MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

Trematoda
(Dicro.) Zonorchis sp. (110)
Nematoda
(FT.) Dipetalonema capilliforme Baylis, 1934 (11)

Genus DAsyurops Matschie, 1916. (Native cats)
D. GRACILIS (Ramsay, i888) No records
D. MACULATUS (Kerr, 1792), the tiger cat
Trematoda
(Strig.) Pharyngostomoides, n. sp. (110E)
Cestoda
(CY., Tae.) Dasyurotaenia robusta Beddard, 1912 (110A)
(PS., Dib.) Diphyllobothrium (Spirometra) erinacei (Rudolphi, 1819) as sparganum
(108)
Nematoda
(ST., Tri.) Trichostrongylus (s.l.) sp. (64)
(SP.) Physaloptera sp. (90)
(F1.) Filaria (s.1.) sp. (58)

Genus SARCOPHILUS Geoffroy & Cuvier, 1837. (Tasmanian devil)
S. HARRISI* (Boitard, 1841)
Trematoda
(Strig.) Alaria sp. (25)
Fibricola sarcophila Sarndars, 1957 (110C)
Cestoda
(CY., Tae.) Anoplotaenia dasyuri Beddard, 1911 (16), (25), (110A)
Dasyurotaenia robusta Beddard, 1912 (17)

(PS.) Sparganum (90)

Nematoda
(ST., Tri.) Nicollina sarcophili Cameron, 1931 (25)
(SP.) Physaloptera sarcophili J. & M., 1940 (67)

Subfamily THyLACININAE
Genus TuHyLtAcinus Temminck, 1824. (Tasmanian wolf)
T. CYNOCEPHALUS (Harris, 1898)
Cestoda
. Dithyridium cynocephaliy Ransom, 1907 (105)

Family MyrmercosiipAr
Genus Myrmecosius Waterhouse, 1836. (Marsupial ant-eaters)
M. FASCIATUS Waterhouse, 1836

Nematoda
(ST., Tri.) Trichostrongylidae (undescribed sp. cellected by Mr. J. H. Calaby)
M. rurus Wood Jones, 1923 No records
Family NororyctTiIpAE
Genus Novrorycres Stirling, 1891. (Marsupial moles)
N. cAurRINUS Thomas, 1920 No records
N. TYPHLOPS (Stirling, 1889) No records
Family PrRAMELIDAE
Genus TuyLaAcis Illiger, 1811
Syn. Isoodon Desmarest, 1817. (Short-nosed bandicoots)
T. AURATUS (Ramsay, 1887) - No recoras
T. BARROWENSIS (Thomas, 1901) No records

* Kreis (1952) recorded several parasites from two marsupials, Sarcophilus harrisii and
Macropus giganteus. However, it is not clear which parasites came from which host.

7 Tapeworm cysts were found in an animal which died in the National Zoological Park,
Washington. They may have been acquired in captivity.

BY M. JOSEPHINE MACKERRAS. 107

T. mAcRouRUS (Gould, 1842)
Cestoda
(CY., Hym.) Hymenolepis peramelidarum Nybelin, 1917 (98) -
(CY., Ano.) Linstowia’ semoni (Zschokke, 1896) (98)

T. NAuTICUS (Thomas, 1922) No records
T. OBESULUS (Shaw and Nodder, 1797)
Protozoa
(M.) Trypanosoma sp. (86)

Giardia sp. (90)
Trichoinonas sp. (90)
(Sa.) Entamoeba sp. (90)
(Sp.) Himeria sp. (90)
Klossiella sp. (35), (86)
Haemogregarina peramelis Welsh and Dalyell, 1909 (86)
Theileria sp. (86)
Sarcocystis sp. (86)
Toxoplasma gondii (Nicolle and Manceaux, 1908) (103), (103A)
Recorded as Hncephalitozoon sp. in (86)
Trematoda
(Dicro. ) Brachylaemus dasyuri (S. J. Johnston, 1913) (46), (86), (110D)
Syn. Harmostomum simile S. J. Johnston, 1913
Zonorchis australiensis Sandars, 1958 (110E)
Recorded as Platynosomum sp. in (86)
Cestoda
(CY., Hym.) Hymenolepis peramelidarum Nybelin, 1917 (86), (110B)
(CY., Ano.) Linstowia semoni (Zschokke, 1896) (134), (135), (110B)

Nematoda
(RH.) Parastrongyloides sp. (90)
Strongyloides sp. (90)
(TR.) Trichuris peramelis Baylis, 1932 (9), (86)

Capillaria sp. (86)

(ST., Tri.) -Filarinema peramelis J. & M., 1938 (60), (86)
Trichostrongylidae — undescribed spp. (90)

(ST. Str.) Cloacina sp. (61)

(ST., Met.) Marsupostrongylus bronchialus Mackerras & Sandars, 1953 (91)
Filostrongylus peramelis Mackerras, 1955 (88)

(OX.) Subulura peramelis Baylis, 1930 (6), (86)
(SP.) Hchinonema cinctum (y. Linstow, 1898) (83), (86)
Physaloptera sp. (90)
(FI.) Dipetalonema johnstoni Mackerras, 1954 (86) (87)
Acanthocephala

Moniliformis semoni (vy. Linstow, 1898) (83), (86)
Larval forms (90)

Pentastomida (Phylum Arthropoda)
Larval forms (90)

T. PENINSULAE (Thomas, 1922) No records
T. TOROSUS (Ramsay, 1877)
Nematoda
(OX.) Subulura peramelis Baylis, 1930 (73)
(SP.) Echinonema cinctum (v. Linstow, 1898) (73)

Genus PERAMELES Geoffroy, 1804. (Long-nosed bandicoots)

P. BOUGAINVILLEI Quoy and Gaimard, 1824 No records
P. EREMIANA Spencer, 1897 No records
P. FascraTa Gray, 1841 a No records
P. GUNNI Gray, 1838 No records

108 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

IP

124

HAS

MyOSURA Wagner, 1841
Nematoda
(OX.) Subulura peramelis Baylis, 1930 (64)
NASuTA Geoffroy, 1804
Protozoa
(Sp.) Klossiella sp. (90)
Theileria sp. (90)
Haemogregarina peramelis Welsh and Dalyell, 1909 (124), (125)
= Hepatozoon peramelis (Welsh and Dalyell, 1909) Wenyon, 1926 (126)
Toxoplasma gondii (Nicolle and Manceaux, 1908) (103) (103A)
Trematoda
(Dicro.) Zonorchis australiensis Sandars, 1958 (110E)
Cestoda

(CY., Hym.) Hymenolepis peramelidarum Nybelin, 1917 (110B)
(CY., Ano.) Linstowia semoni (Zschokke, 1896) (49)
(CY., Dil.) Mirandula parva Sandars, 1956 (109)

Nematoda
(RH.) Parastrongyloides sp. (90)
(TR.) Trichuris peramelis Baylis, 1932 (63)

(ST., Tri.) Trichostrongylidae — undescribed spp. (90)

(ST., Met.) Filostrongylus peramelis Mackerras, 1955 (88)

(OX) Subulura peranelis Baylis, 19380 (68)

(SP.) Hchinonema cinctum (vy. Linstow, 1898) (67)
Physaloptera parvicollaris J. & M., 1240 (67)
Ph. peramelis J. & M., 1939 (68)

(FT.) Dipetalonema johnstoni Mackerras, 1954 (87)
Dipetalonema sp. (lung) (63), (67)
Acanthocephala

Moniliformis semoni (v. Linstow, 1898) (50), (56)

Genus THyLAcomys Blyth, 1840. (Rabbit bandicoots or bilbies)
Syn. Macrotis Reid, 1837 (preoccupied); Paragalia Gray, 1841; (or Peragale)

. LAGOTIS (Reid, 1837) No records
. LEUCURA (Thomas, 1887) No records
. LEUCURA MINOR (Spencer, 1897)
Nematoda
- (OX.) Subulura peragale J. & M., 1940 (67)
(SP.) Physaloptera peragale J. & M., 1940 (66)
Ph. thalacomys J. & M., 1940 (67)
Genus CHAEROPUS Ogilby, 1838. (Pig-footed bandicoots)
ECAUDATUS Ogilby, 1838 No records

C.

9

Oe

? Banpicoor (New Guinea)

(? EHchymipera sp. or Peroryctes sp.)

Nematoda
(SP.) Physaloptera papuensis J. & M., 1940 (67)
Suborder DiIPpRoTODONTIA
Family PHALANGERIDAE
Subfamily TARSIPEDINAE
Genus TARSIPES Gray, 1842. (Honey possum)
. SPENSERAE Gray, 1842 No records
Subfamily PHALANGERINAE
Genus ACROBATES Desmarest, 1818. (Pigmy glider)
. PYGMAEUS (Shaw, 1793) No records
Genus CrErcARTETUS Gloger, 1841. (Pigmy possums)
. CONCINNUS (Gould, 1845) : No records
. NANUS (Desmarest, 1818) No records

BY M. JOSEPHINE MACK2ERRAS.

Genus Hupromicra Mijoberg, 1916. (Pigmy possums)

EK. LErPIpDA (Thomas, 1888) No records
E. MACRURA Mjoberg, 1916 No records
Genus GYMNOBELIDEUS McCoy, 1867. (Possum)
G. LEADBEATERI McCoy, 1867 No records
Genus PrETAuRUS Shaw and Nodder, 1791. (Gliders)
P. AUSTRALIS Shaw and Nodder, 1791 No records
P. BREVICEPS Waterhouse, 1839
Protozoa
(Sp.) Haemogregarina petauri Welsh and Barling, 1909 (90)
P. NORFOLCENSIS Kerr, 1792 No records
P. sciureus Shaw, 1794
Protozoa
(Sp.) Haemogregarina petauri Welsh and Barling, 1909 (120), (121)
= Hepatozoon petauri (Welsh and Barling) Wenyon, 1926 (126)
(host given as Petaurus sp., probably sciureus)
Genus DaAcTyLopsILA Gray, 1858. (Striped possum)
D. picatTa Thomas, 1908 No records
Genus PsmUDOCHEIRUS Ogilby, 1837. (Ring-tail possums)
P. coNVOLUTOR (OKen, 1816) No records
P. HERBERTENSIS (Collett, 1884)

Cestoda
(CY., Ano.) Prototaenia aberrata (Nybelin, 1917) (98)
Prototaenia pseudochiri (Nybelin, 1917) (98)
P. LANIGINOSUS (Gould, 1858)
Protozoa
(Sp.) Haemogregarina sp. (90)
Cestoda ;
(CY., Ano.) ? Prototaenia sp. (110)
P. OCCIDENTALIS Thomas, 1888 No records
P. PEREGRINUS (Boddaert, 1785) No records

Genus PsEupocHtIRorps Matschie, 1915. (Striped ring-tail possum)
Ps. ARCHERI (Collett, 1884) No records

Genus PETROPSEUDES Thomas, 1923. (Rock ring-tail possum)
P. DAHLI (Collett, 1895) No records

Genus HEMIBELIDEUS Collett, 1884. (Bushy-tipped ring-tail possum)
H. LEMURODES Collett, 1884
Cestoda
(CY., Ano.) Parabertiella campanulata Nybelin, 1917 (98)
Prototaenia undulata (Nybelin, 1917) (98)
Prototaenia pellucida (Nybelin, 1917) (98)

Nematoda
(FI.) Microfilariae in blood (112).
Genus SCHCINOBATES Lesson, 1842
Syn. Petauroides Thomas, 1888. (Greater gliders)
S. voLANS (Kerr, 1792) No records
S. VOLANS INCANUS Thomas, 1923
Nematoda
(OX.) Austrocyuris finlaysoni J. & M., 1938 (60)

Passalurus parvus J. & M., 1938 (60)
Oxryuris (s.1.) acuticaudata J. & M., 1938 (60)

109

110 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

Genus Triciiosurus Lesson, 1828, (Brush-tailed possums)
T. cANINUS (Ogilby, 1836)

Nematoda
(ST., Tri.) Asymmetricostrongylus trichosuri J. & M., 1939 (62)
(19115) Dipetalonema trichosuri (Breinl, 1913) (67)
Filaria sp. (50), (58)
T. FULIGINOSUS (Ogilby, 1831) No records
T. VULPECULA (Kerr, 1792)
Protozoa
Entamoeba sp. (90)
Cestoda

Taenia phalangistae Krefft, 1871 (76) (not rediscovered)
Nematoda
(RH.) Parastrongyloides sp. (90)
(ST., Tri) Trichostrongylus colubriformis (Giles, 1892) (15)
T. rugatus Monnig, 1925 (15)
Trichostrongylidae (undescribed sp.) (90)

(OX.) Syphacia trichosuri J. & M., 1938 (60)
(SP.) Protospirura marsupialis Baylis, 1927 (5), (12), (60), (63)
(FI.) Dipetalonema trichosuri (Breinl, 1913) (23), (58)

Filaria dentifera v. Linstow, 1897 (81) (not rediscovered)
Microfilaria (sheathed) (21)

Genus Wyvuitpa Alexander, 1919. (Scaly-tailed possum)
W. SQUAMICAUDATA Alexander, 1919 No records

Genus Spimocuscus Gray, 1862. (Cuscus)
SP. NUDICAUDATUS (Gould, 1850) No records

Genus PHALANGER Storr, 1780. (Phalangers or cuscuses)
P. MACULATUS KRAMERI Schwartz, 1910 (Manus Is.)
Cestoda
(CY., Ano.) Prototaenia kapul (Baylis, 1934) (13)
P. ORIENTALIS PENINSULAE Tate, 1945 No records
P. uRSINUS (Temminck, 1827) (Celebes)
Cestoda
(CY., Ano.) Prototaenia edulis (Zschokke, 1899) (136), (137)
P. sarasinorum (Zschokke, 1899) (136), (187)
PHALANGISTA sp. (? Trichosurus sp. or Phalanger sp.), New Guinea
Cestoda
(CY., Ano.) Prototaenia rigida (Janicki, 1905) (44)

Family PIrAScoLArctTiIpAr
Genus PHAScOLARCTOS de Blainville, 1816. (Koala)
P. cINEKEUS Goldfuss, 1817
Cestoda
(CY., Ano.) Prototaenia obesa (ZschokkKe, 1896) (134), (185), (110B)
Taenia geophiloides Cobbold, 1879, is a nomen nudum (32)

Family VoMBATIDAE
Genus VomBatus Geoffroy, 1803
Syn. Phascolomis Geoffroy, 1803. (Wombats)
VY. HIRSUTUS (Perry, 1810)
Syn. mitchelli Owen, 1838

Protozoa
(Sp.) Toxoplasma wenyoni Coutelen, 1932 (33)

Cestoda
(CY., Ano.) Moniezia sp. (49)

BY M. JOSEPHINE MACKERRAS. 411

Nematoda
(ST., Str.) Oesophagostomum giltneri Schwartz, 1928 (111)
Phascolostrongylus turleyi Canavan, 1931 (27)
V. uRSINUS (Shaw, 1800)
Cestoda
(CY., Ano.) Progamotaenia diaphana (Zschokke, 1907) (138)

Genus LASIORHINUS Gray, 1863. (Hairy-nesed wombat)
L. LATIFRONS (Owen, 1845)

Protozoa
(Sp.) Tleocystis wombati Gilruth and Bull, 1912 (41)
= Globidium wombati (Gilruth and Bull, 1912) Wenyon, 1926 (126)
Nematoda

(ST., Str.) Macropostrongylus lasiorhini Mawson, 1955 (91A)
Phascolosirongylus stirtont Mawson, 1955 (91A)

Genus WoMBPATULA Iredale and Troughton, 1934. (Queensland wombat)
W. GILLESPIEI (de Vis, 1900) No records
“WOMBAT” (ex Philadelphia Zoo)
Cestoda
Taenia bipapillosa Leidy, 1875 (79)
This is practically a nomen nudum

Family MAcrRoPopIDAE
Subfamily HypsIpryMNODONTINAE
Genus HyPSIPRYMNODON Ramsay, 1876. (Musk rat-kangaroo)
H. MOSCHATUS Ramsay, 1876 No records

Subfamily PorTororInar
Genus Brerronera Gray, 1837. (Rat-kangaroo)

B. CUNICULA (Ogilby, 1838) No records
B. GAIMARDI (Desmarest, 1822) No records
B. LESUEURI (Quoy and Gaimard, 1824) No records
B. LESUEURI GRAYI Gould
Protozoa
(Sp.) Sarcocystis bettongiae Bourne, 1932 (22)

B. PENICILLATA Gray, 1837 : No records
Genus AEPYPRYMNUS Garrod, 1875. (Rat-kangaroo)
A. RUFESCENS (Gray, 1837) No records

Genus Pororous Desmarest, 1804. (Rat-kangaroos)

GILBERTI (Gould, 1841) No records
. PLATYOPS (Gould, 1844) No records
. TRIDACTYLUS (Kerr, 1792)

Protozoa
(Sp.) Theileria sp. (90)

Nematoda
(ST., Tri.) Austrostrongylus potoroo J. & M., 1949 (72)
(ST., Str.) Labiostrongylus eugenii (J. & M., 1940) (72)

Potorostronaylus finlaysoni J. & M., 1939 (64)

(OX.) Ozyuris (s.l.) potoreo J. & M., 1939 (64)
(UM IE)) Filaria (s.l.) sp. (58)

ty

Genus CALOopRYMNUS Thomas, 1888. (Rat-kangaroo)
C. CAMPESTRIS (Gould, 1843) No records

112 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

Subfamily MAcropopiInAr
Genus DrnpROLAGUS Miiller, 1839. (Tree kangaroos)
D. BENNETTIANUS de Vis, 1887
Nematoda
(FI.) Filaria sp. (50)
Dipetalonema spelaea (Leidy, 1875) (58)
D. LUMHOLTzI Collett, 1884

Nematoda
(FI.) Dipetalonema sp. ? roemeri (v. Linstow, 1905) (58)
D. 1nustus Schlegel and Miiller, 1839-44 (New Guinea)
Protozoa
(M.) Trichomonas guttula Kirby and Honigberg, 1950 (75)
Trematoda
Species not specified (liver and bile ducts) (43)
Nematoda
(FT.) Dipetalonema dendrolagi (Solomon, 1933) (114)

Genus LAGORCHESTES Gould, 1841. (Hare-wallabies)
L. CONSPICILLATUS Gould, 1842
Cestoda
(CY., Ano.) Progamotaenia lagorchestis (Lewis, 1914) (80)
Cittotaenia villosa Lewis, 1914 (80)
L. HIRSUTUS Gould, 1844

Nematoda
(ST., Str.) Labiostrongylus communis (J. & M., 1939) (67)
L. LEPORIDES (Gould, 1841) No records

Genus LAGosTRoPHUS Thomas, 1887. (Banded wallaby)
L. FASCIATUS (Péron and Lesueur, 1807) No records

Genus ONYCHOGALEA Gray, 1841. (Nail-tail wallabies)
O. FRAENATA (Gould, 1841)
Cestoda
(CY., Ano.) Progamotaenia bancrofti (T. H. Johnston. 1912) (52), (98)
Nematoda :
(ST., Str.) Labiostrongylus onychogale (J. & M., 1939) (61)
(FI.) Dipetalonema annulipapillatum J. & M., 1938 (58)
D. rarum J. & M., 19388 (58)
D. roemeri (v. Linstow, 1905)- (58)
Microfilariae in blood (101)
Filaria sp. (50)
O. LUNATA (Gould, 1841) No records
O. UNGUIFER (Gould, 1841)
Cestoda
(CY., Ano.) Progamotaenia festiva (Rudolphi, 1819) (98), (119)

Genus Prraporcas Thomas, 1904. (Little rock-wallaby)

P. coNcINNA (Gould, 1842) No records
Genus PE&TROGALE Gray, 1837. (Rock-wallabies)
P. pRAcHYOTIS Gould, 1841 No records
P. HACKETTI Thomas, 1905 No records
P. INORNATA Gould, 1842 No records
P. LATERALIS Gould, 1842
Nematoda

(ST., Str.) Cloacina elegans J. & M., 19388 (59)
C. ernabella J. & M., 1938 (59)
C. hydriformis J. & M., 1938 (59)

BY M. JOSEPHINE MACKERRAS. 113

(ST., Str.) C. longelatiata J. & M., 1939, new name for minor J. & M., 1938 (59)
C. macropodis J. & M., 1938 (59)
C. parva J. & M., 1938 (59)
C. petrogale J. & M., 1938 (59)
Labiostrongylus longispicularis Wood, 1929 (59)
L. petrogale J. & M., 1938 (59)
Pharyngostrongylus alpha J. & M., 1988 (59)
Ph. beta J. & M., 1938 (59)
P. LONGMANI Thomas, 1926 No records
P. PEARSONI Thomas, 1922
Nematoda
(ST., Str.) Cloacina petrogale J. & M., 1938 (66)
Labiostrongylus longispicularis Wood, 1929 (66)
Macropostrongylus pearsoni J. & M., 1940 (66)
Pharyngostrongylus alpha J. & M., 1938 (66)
Ph. beta J. & M., 1938 (66)
P. PENICILLATA (Griffith, Smith and Pidgeon, 1827)

Protozoa
(Sp. ) Sarcocystis mucosae (Blanchard, 1885) Minchin, 1903 (subintestinal) (93)
= Globidium mucosae (Blanchard, 1885) Wenyon, 1926 (126)
Cestoda
(CY., Ano.) Tvriplotaenia mirabilis* Boas, 1902 (20)
Nematoda

(ST., Str.) Cloacina robertsi J. & M., 1939 (61)
C. similis J. & M., 1939 (61)
Pharyngostrongylus alpha J. & M., 1938 (61)
Ph. zeta J. & M., 1939 (61)

(FI.) Dipetalonema spelaea (Leidy, 1875) (58), (63)
Dipetalonema sp. (63)
Microfilariae in blood (102)

P. ROTHSCHILDI Thomas, 1904 No records
P. WILKINSI Thomas, 1926 No records
P. XANTHOPUS Gray, 1855

Nematoda

(ST., Str.) Cloacina australis J. & M., 1938 (66)
C. communis J. & M., 1938 (66)
CC. curta J. & M., 1938 (66)
C. frequens J. & M., 1938 (66)
C. longelabiata J. & M., 1939 (66)
C. macropodis J. & M., 1938 (66)
Labiostrongylus longispicularis Wood, 1929 (66)
Pharyngostrongylus beta J. & M., 1938 (66)
(FI.) Dipetalonema sp. (58)
Filaria sp. (26)
PETROGALE SP.
Protozoa
(Sp.) Sarcocystis macropodis Gilruth and Bull, 1912 (41)
= Globidium sp. Wenyon, 1926 (126)

Genus Sretronix Lesson, 1842. (QuokkKa)

S. BRACHYURUS (Quoy and Gaimard, 1830)
Protozoa
(Sa.) Entamoeba sp. (90)

«This peculiar cestode is regarded by some helminthologists as a monster. It would be
very desirable to get more material from this host. ;

114 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

Cestoda
(CY., Ano.) Progamotaenia bancrofti (T. H. Johnston, 1912) (110B)
Progamotaenia sp. (110B)
Nematoda
(FI.) Microfilariae in blood (102)

Genus Dorcorsis Schlegel and Miller, 1842
D. VETERUM (Lesson and Garnot, 1826) (Japen Island)
Nematoda
(ST., Str.) Macropostrongylus dorcopsis Baylis, 1940 (14)

Genus THYLOGALE Gray, 1837. (Pademelons or little scrub wallabies)
T. BILLARDIERIZ (Desmarest, 1822)
Nematoda
(ST., Str.) Labiostrongylus uncinatus (J. & M., 1939) (64)
Pharyngostrongylus delta J. & M., 1938 (64)
Ph. epsilon J. & M., 1939 (64)
Zoniolaimus buccalis (J. & M., 1939) (64)
T. COXENII (Gray, 1866) No records
T. EUGENII (Desmarest, 1817)
Syn. derbianus (Gray, 1837)
Cestoda
(CY., Ano.) Progamotaenia festiva (Rudoiphi, 1819) (32), (119)
Nematoda
(ST., Tri.) Austrostrongylus thylogale J. & M., 1940 (66)
(ST., Str.) Cloacina curta J. & M., 1938 (70)
C. petrogale J. & M., 1938 (65), (70)
Labiostrongylus eugentt (J. & M., 1940) (65), (66), (70)
Spirostrongylus kartana Mawson, 1955 (91A)
(SP.) Physaloptera sp. (larval form) (65)
T. FLINDERSI Wood Jones, 1924
Nematoda
(ST., Str.) Cloacina macropodis J. & M., 1938 (66)
C. petrogale J. & M., 1938 (66)
Pharyngostrongylus beta J. & M., 1938 (66)
T. PARMA (Waterhouse, 1846)
Nematoda
(ST., Str.) Cloacina thetidis J. & M., 19389 (67)
Coronostrongylus coronatus J. & M., 1939 (67)
Parazoniolaimus collaris J. & M., 19389 (67)
Pharyngostrongylus alpha J. & M., 1938 (67)
Ph. delta J. & M., 1939 (67)
Ph. epsilon J. & M., 1939 (62)
Ph. gamma J. & M., 1939 (67)
Spirostrongylus parma (J. & M., 1939) (62), (91A)
Zoniolaimus buccalis (J. & M., 1939) (67)
T. sTigMATA Gould, 1860 No records
T. THETIS (Lesson, 1827)
Protozoa
(Sp.) Coccidium (Himeria) sp. (50)
Cestoda
(CY., Tae.) Hchinococcus granulosus (Batsch, 1786) as hydatid (49)
Nematoda
(ST., Anc.) Hypodontus thetidis J. & M., 1939 (62)
(ST., Str.) Globocephaloides thetidis J. & M., 1939 (62)
Cloacina bancroftorum J. & M., 1939 (62)
C. similis J. & M., 1939 (62)

BY M. JOSEPHINE MACKERRAS. 115

(Si. Sie) Cl WieHWiIS do Ke IML, ISEB) (Hz)
Coronostrongylus coronatus J. & M., 1939 (61)
Cyclostrongylus medioannuiatus J. & M., 1940 (67)
Labtostrongylus onychogale (J. & M., 1939) (67)
L. uncinatus J. & M., 1939 (65)
Pharyngostrongylus alpha J. & M., 1938 (62)
OCLC lanes eV 939) (G2)
Pi epsilon de is Nia 9389) (62)
Eh GWEtO Jen Nie 1939) \(62)
Ph. zeta J. & M., 1939 (61), (62)
Zoniolaimus australis (J. & M., 1939) (61)
ZL. ouccalis (J. & Me, 1939) (62)

(FT.) Dipetalonema sp. (67)

T. witcoxt (McCoy, 1866)
Cestoda
(CY., Tae.) Hchinococcus granulosus (Batsch, 1786) as hydatid (37)
(CY., Dav.) Calostaurus macropus (Ortlepp, 1922) (110B)
Nematoda

(ST., Str.) Cloacina macropodis J. & M., 1938 (61)
C. similis J. & M., 1939 (67)
C. thetidis J. & M., 1940 (67)
Coronostrongylus coronatus J. & M., 1939 (61)
Labiostrongylus communis (J. & M., 1939) (61)
Pharyngostrongylus epsilon J. & M., 1939 (61)
Zoniolaimus australis (J. & M., 1939) (61)
Z. buccalis (J. & M., 1939) (61)

(FI.) Dipetalonema thylogali Mackerras, 1954 (87)

Genus WALLABIA Trouessart, 1905. (Brush wallabies, or large wallabies)
The generic name Protemnodon Owen, 1873, based on
fossil material, is also used for this group cf wallabies
W. acizis (Gould, 1842)
Cestoda
(CY., Ano.) Progamotaenia festiva (Rudolphi, 1819) (98), (119)
Nematoda
(ST., Str.) Cloacina cornuta (Davey and Wood, 1938) (34)
C. digitata J. & M., 1940 (72)
C. longispiculata J. & M., 1939 (61)
C. macropodis J. & M., 1938 (61)
C. robertsi J. & M., 1939 (61)
Labiostrongylus insularis J. & M., 1949 (72)
L. labiostrongylus Y. & M., 1926 (10)
Macropostrongylus australis Y. & M., 1926 (10)
M. macropostrongylus Y. & M., 1926 (10), (61). (63)
M. yorkei Baylis, 1927 (10), (61), (63)
(FT.) D. annulipapillatum J. & M., 1938 (60}, (61)
D. roemeri (vy. Linstow, 1905) (72)
D. ? spelaea (Leidy, 1875) (11)
W. sBicotor (Desmarest, 1804) including subspecies mastersii, Krefft and apicalis
Gunther
Syn. walabatus Lesson and Garnot, 1827
Cestoda
(CY., Tae.) Hchinococcus granulosus (Batsch, 1786) as hydatid (49), (37)
(CY., Dil.) Bancroftiella tenuis T. H. Johnston, 1911 (50)
(CY., Ano.) Progamotaenia bancrofti (T. H. Johnston, 1912) (110B)
Nematoda
(ST., Tri.) Awustrostrongylus aggregatus J. & M., 1940 (67)

116 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

(ST., Str.) Globocephaloides thetidis J. & M., 1939 (67)
Cloacina gallardi J. & M., 1940 (67)
C. macropodis J. & M., 1938 (62)
C. similis J. & M., 1939 (61)
C. wallabiae J. & M., 1989 (62)
Cyclostrongylus dissimilis J. & M., 1939 (62)
Cy. wallabiae J. & M., 1939 (62)
Labiostrongylus clelandi (J. & M., 1989) (61), (62), (64)
L. communis (J. & M., 1939) (61)
L. ualabatus (J. & M., 1939) (62), (64)
L. uncinatus (J. & M., 1939) (61)
Macropostrongylus dissimilis J. & M., 1939 (62)
Maplestonema typicum J. & M., 1939 (62)
Parazoniolaimus collaris J. & M., 1939 (62), (64)
Pharyngostrongylus alpha J. & M., 1938 (62)
Ph. beta J. & M., 1928 (62)
Ph. epsilon J. & M., 1939 (62), (64)
Spirostrongylus gallardi (J. & M., 1942) (71), (914A)
Zoniolaimus brevicaudatus Cobb, 1898 (62)
Z. setifera Cobb, 1898 (62)
(FI.) Dipetalonema annulipapillatum J. & M., 1938 (58)
D. roemeri (v. Linstow, 1905) (58)
D. spelaea (Leidy, 1875) (49), (638)
W. BROWNI (Ramsay, 1877), New Britain, New Guinea
Nematoda
(ST., Str.) Cloacina dahli v. Linstow, 1898 (82)
W. BRuNIT (Schreber, 1778), Aru Island.
Cestoda
(CY., Dav.) Calostaurus macropus (Ortlepp, 1922) (100), (110B)
W. DORSALIS (Gray, 1837)
Cestoda
(CY., Tae.) Hcehinococcus granulosus (Batsch, 1786) hydatid (2), (87)
Nematoda
(ST., Tri.) <Austrostrongylus minutus J. & M., 1938 (60)
(ST., Str.) Globocephaloides affinis J. & M., 1939 (61)
G. wallabiae J. & M., 1939 (61)
Cloacina bancroftorum J. & M., 1939 (61)
C. burnettiana J. & M., 1939 (61)
C. digitata J. & M.. 1940 (67)
C. longispiculata J. & M., 1939 (61)
C. similis J. & M., 1940 (67)
Labiostrongylus longispicularis Wood, 1929 (67)
LL. uncinatus (J. & M., 1939) (61)
Papillostrongylus labiatus J. & M., 1939 (61)
Pharyngostrongylus delta J. & M., 19389 (61)
Ph. epsilon J. & M., 1939 (61)
Ph. gamma J. & M., 1939 (61)
Ph. theta J. & M., 1940 (67)
Ph. zeta J. & M., 1939 (61)
Zoniolaimus buccalis (J. & M., 1939) (61)
(FI.) Dipetalonema annulipapillatum J. & M., 1938 (58)
D. roemeri (v. Linstow, 1905) (58)
= Filaria websteri Cobbold (52)
W. ELEGANS (Lambert, 1807)
Syn. parryi Bennett, 1835
Protozoa
(Sp.) Coccidium (Himeria) sp. (50)

BY M. JOSEPHINE MACKERRAS. IAL

Cestoda
(CY., Tae) Echinococcus granulosus (Batsch, 1786) as hydatid (37)
‘Nematoda
(ST., Str.) Cloacina communis J. & M., 1988 (61)
Labiostrongylus bancrofti (J. & M., 1939) (61)
Macropostrongylus yorkei Baylis, 1927 (61)
Pharyngostrongylus brevis Canavan, 1931 (61)
Ph. gamma J. & M., 1939 (61)
Ph. macropodis Y. & M., 1926 (61)
Zoniolaimus buccalis (J. & M., 1939) (61)
(FI.) Dipetalonema roemeri (v. Linstow, 1905) (58)
= Filaria websteri Cobbold (52)
W. GREYI (Waterhouse, 1846) No records
W. irmA (Jourdan, 1837)
Nematoda
(ST., Str.) Cloacina curta J. & M., 1938 (65)
Labiostrongylus communis (J. & M., 1939) (64), (65)
Macropostrongylus irma J. & M., 1940 (65)
Pharyngostrongylus beta J. & M., 19388 (65)
W. RUFOGRISEA (Desmarest, 1817) (including Tasmanian subspecies frutica Ogilby, 1838)
Syn. ruficollis Desmarest, 1817; bennetti Waterhouse, 1838
Protozoa
(Sp.) Toxoplasma sp. (106)
Himeria macropodis Wenyon and Scott, 1925 (127), (118)
Tleocystis macropodis Gilruth and Bull, 1912 (127), (118)
Lymphocystis macropodis Gilruth and Bull, 1912 (127), (118)
Coccidia (116)
Trematoda
(Fasci.) Fasciola hepatica L., 1758 (50)
Cestoda
(CY., Ano.) Thysanotaenia incognita Meggitt, 1927 (92)
(CY., Tae.) Hchinococcus granulosus (Batsch, 1786) as hydatid (37)
Nematoda
(RH.) Strongyloides sp. (116)
(ST., Tri.) Asymmetricostrongylus asymmetricus (Cameron, 1926) (24)
As. dissimilis (Wood, 1931) (78)
Austrostrongylus macropodis Chandler, 1924 (29)
Au. wallabiae J. & M., 1939 (62)
(ST., Str.) Globocephaloides trifidospicularis Kung, 1948 (78)
Cloacina linstowi J. & M.,.1940 (67)
C. similis J. & M., 1939 (67)
C. thetidis J. & M., 1939 (67)
Coronostrongylus coronatus J. & M., 1939 (67)
Cyclostrongylus gallardi J. & M., 1940 (62)
Labiostrongylus communis (J. & M., 1939) (62), (64)
L. australis Kung, 1948 (78)
L. onychogale J. & M., 1939 (62)
Macropostrongylus .lesouefi J. & M., 1939 (62)
M. wallabiae J. & M., 1939 (62)
Pharyngostrongylus alpha J. & M., 1938 (62)
Ph. beta J. & M., 1938 (62)
Ph. brevis Canavan, 1931 (62)
Ph. delta J. & M., 1939 (62)
Ph. epsilon J. & M., 1939 (62)
Ph. eta J. & M., 1939 (62)
Ph. gamma J. & M., 1939 (62)
Ph. iota J. & M., 1939 (62)

118

(

(

W.

(

AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

ST., Str.) Ph. longibursaris Kung, 1948 . (78)
Ph. macropodis Y. & M., 1926 (62)
Ph. theta J. & M., 19389 (64)
Ph. zeta J. & M., 19389 (62), (64)
Zoniolaimus australis (J. & M., 1939) (62)
Z. buccalis (J. & M., 1939) (64)
Z. chaetophorus J. & M., 1949 (72)

New name for setifer J. & M., 1939 (62)

Z. cobbi Kung, 1948 (78)
Z. labiatus J. & M., 1939 (62), (64)

FI.) Dipetalonema roemeri (v. Linstow, 1905) (58)
Dipetalonema sp. (63)
Filaria sp. (38)

WELSBYI (Longman, 1922)

Nematoda

ST., Str.) Cloacina macropodis J. & M., 1938 (61)
Cloacina sp. (61)
Labiostrongylus insularis (J. & M., 1939) (61)
Macropostrongylus macropostrongylus Y. & M., 1926 (61)
M. yorkei Baylis, 1927 (61)

(FI.) Dipetalonema roemeri (v. Linstow, 1905) (58)
? WALLABY
Nematoda
(FI.). Dipetalonema sp. (encysted in liver) (65)
(AS.) Contracaecum erraticum J. & M., 1940 (65)

(The latter specimens were labelled “Ascaris from wallaby” in a
very old collection in the Australian Museum. Johnston and Mawson
(1940a) consider it is possible the label was misplaced, as the worms

M.

resemble forms from cormorants. No other ascarids have been

recorded in marsupials.)

Genus Macropus Shaw and Nodder, 1790. (Kangaroos)
Syn. Halmaturus Illiger, 1811
MAJOR* Shaw, 1800 (also referred to as giganteus Zimmermann, 1777)
Protozoa

(Sp.) Coccidium (EHimeria) sp. (50)

Trematoda

(Fasci.) Fasciola hepatica L., 1758 (49)

Cestoda

(CY., Tae.) Hchinococcus granulosus (Batsch, 1786) hydatid (49)
(CY., Ano.) Progamotaenia festiva (Rudolphi, 1819) (49), (110B)

Nematoda

(ST., Str.) Cloacina communis J. & M., 1938 (67)

C. expansa J. & M., 1939 (62)

C. magnipapillata J. & M., 1939 (62), (64)

C. obtusa J. & M., 1939 (62)

Cyclostrongylus clelandi J. & M., 1939 (62)
Labiostrongylus bipapillosus (J. & M., 1939) (61), (65)
L. longispicularis Wood, 1929 (62), (64)

L. kungi Mawson, 1955 (91A)

Pharyngostrongylus alpha J. & M., 1938 (62), (64), (73)
Ph. beta J. & M., 19388 (62), (64)

Ph. macropodis Y. & M., 1926 (61), (65)

(FL) Dipetalonema roemeri (v. Linstow, 1905) (4), (58), (64)

Dipetalonema sp. (58)
Filaria websteri Cobbold, 1879 (32), (39), (52)
Filaria macropi majoris (39)

* See footnote under Sarcophilus harrisii (p. 106).

BY M. JOSEPHINE MACKERRAS. 119

M. MELANOoPS Gould, 1842 (possibly only a colour phase of M. major)
Nematoda
(ST., Str.) Cloacina australis J. & M., 1938 (66)
communis J. & M., 1938 (66)
. curta J. & M., 1938 (66)
. frequens J. & M., 1938 (66)
. hydriformis J. & M., 1938 (66)
. longelabiata J. & M., 1939 (66)
. macropodis J. & M., 19388 (66)
. parva J. & M., 1938 (66)
. obtusa J. & M., 1939 (66)
. vestibulata J. & M., 1940 (66)
Labiostrongylus longispicularis Wood, 1929 (64)
Paramacropostrongylus typicus J. & M., 1940 (66)
Pharyngostrongylus alpha J. & M., 1938 (66)
Ph. beta J. & M., 1938 (66)
(FI.) Dipetalonema roemeri (v. Linstow, 1905) (58), (66)
M. ocypromus Gould, 1842

QQ Q qq eq 22 &

Nematoda
(ST., Str.) Cloacina curta J. & M., 1938 (65)
C. obtusa J. & M., 1939 (65)
Pharyngostrongylus beta J. & M., 1938 (65)

(FI.) Dipetalonema roemeri (v. Linstow, 1905) (72)
M. TASMANIENSIS Le Souef, 1923
Nematoda

(ST., Str.) Labiostrongylus longispicularis Wood, 1929 (67), (72)
M. FULIGINOSUS (Desmarest, 1817)

Nematoda
(ST., Str.) Labiostrongylus communis (J. & M., 1939) (65)
(FT.) Dipetalonema roemeri (v. Linstow, 1905) (67)

MACROPUS SP., presumably M. major

Nematoda
(ST., Anc.) Hypodontus macropi Monnig, 1929 (73)

Genus MrcaLeIA Gistel, 1848. (Red kangaroo)
M. gnura (Desmarest, 1822) (usually called Macropus rufus)

Protozoa
(Sp.) Toxoplasma sp. (42)
Cestoda
(CY., Ano.) Baeriella proterogyna Fuhrmann, 1932 (40)
Nematoda
(RH.) Strongyloides sp. (90)
(ST., Anc.) Hypodontus macropi Monnig, 1929 (96)
(ST., Tri.) Filarinema flagrifer Monnig, 1929 (97)
(ST., Str.) Cloacina hydriformis J. & M., 1938 (59)
C. inflata J. & M., 1938 (59)
C. Viebigi J. & M., 1938 (59)
C. longelabiata J. & M., 1939 (638)
New name for minor J. & M., 1938 (59)
C. longispiculata J. & M., 1939 (62)
C. magnipapillata J. & M., 1939 (62)
C. petrogale J. & M., 1988 (59)
Labiostrongylus longispicularis Wood, 1929 (59), (62), (65)
Pharyngostrongylus alpha J. & M., 1938 (62), (73)
Ph. australis (Moénnig, 1926) (94), (97), (129)
Ph. beta J. & M., 1938 (65)

120

AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

Genus OSPHRANTER Gould, 1842. (Wallaroos and Euros)

QO. ANTILOPINUS Gould, 1842

Nematoda
(FI.) Dipetalonema roemeri (v. Linstow, 1905) (84)
QO. ANTILOPINUS WOODWARDI (Thomas, 1901)
Cestoda
(CY., Ano.) Progamotaenia festiva (Rudolphi, 1819) (98)
Nematoda
(ST., Tri.) Asymmetricostrongylus australis (Wood, 1931) (130)

(ST., Str.)

A. dissimilis (Wood, 1931) (130)

Labiostrongylus longispicularis Wood, 1929 (128)
Macropostrongylus baylist Wood, 1931 (130)
Pharyngostrongylus australis (Monnig, 1926) (129)
Ph. woodwardi Wood, 1931 (130)

Q. BERNARDUS (Rothschild, 1904)

Nematoda
(ST., Str.) Pharyngostrongylus brevis Canavan, 1931 (27)
(0. ERUBESCENS (Sclater, 1870)
Nematoda
(ST., Str.) Cloacina australis J. & M., 1938 (59)

. communis J. & M., 1938 (59)
. curta J. & M., 1938 (59)
. dubia J. & M., 1938 (59)
. frequens J. & M., 1938 (59)
. longelabiata J. & M., 1939 (63)
New name for minor J. & M., 1938 (59)
C. macropodis J. & M., 1988 (59)
C. magna J. & M., 1938 (59)
C. parva J. & M., 1938 (59)
Labiostrongylus grandis J. & M., 1938 (59)
L. longispicularis Wood, 1929 (59)
L. macropodis J. & M., 1938 (59)

QRQaagaea

QO. ISABELLINUS Gould, 1842

Nematoda

(ST., Str.)

O. REGINAE (Schwartz, 1910)

Labiostrongylus longispicularis Wood, 1929 (59)

O. RopusTUS (Gould, 1841)
Protozoa

(M.)

Trichomonas guttula Kirby and Honigberg, 1950 (75)
Retortamonas mitrula Kirby and Honigberg, 1950 (75)
Monocercomonas sp. (75)

Jestoda

(CY., Tae.) Echinococcus granulosus (Batsch, 1786) hydatid (49)
Nematoda

(ST., Str.) Cloacina minor (Davey and Wood, 1938) (a4)

CFI.)

Labiostrongylus longispicularis Wood, 1929 (128), (65)
Macropostrongylus labiatus Davey and Wood, 1938 (34)
M. macrostoma Davey and Wood, 1988 (34)
Pharyngostrongylus alpha J. & M., 1938 (62)

Ph. beta J. & M., 1938 (67)

Ph. ornatus Davey and Wood, 1938 (34)

Dipetalonema robertsi J. & M., 1988 (60)

D. roemeri (v. Linstow, 1905) (58), (66)

D. tenue J. & M., 1988 (58)

Dipetalonema sp. (58)

No records

BY M. JOSEPHINE MACKERRAS. 121

Macropus sp. (unidentified kangaroos or wallabies)
Protozoa
(Sp.) Tleocystic macropodis Gilruth and Bull, 1912 (41)
= Globidium macropodis (Gilruth and Bull) Wenyon, 1926 (126)
Lymphocystic macropodis Gilruth and Bull, 1912 (41)
= Globidium sp. Wenyon, 1926 (126)
Toxoplasma sp. (106)
Cestoda
(CY., Ano.) Progamotaenia zschokkei (Janicki, 1905), New Guinea (44)
Bothriocephalus marginatus Krefft, 1871 (76)
Taenia fimbriata Krefft, 1871 (76)
Taenia mastersi Krefft, 1871 (76)
(Krefft’s species are unrecognizable)
Nematoda
(ST., Str.) Globocephaloides macropodis Y. & M., 1926 (132)
Labiostrongylus communis J. & M., 1939 (61)
L. labiostrongylus Y. & M., 1926 (132)
Macropostrongylus australis Y. & M., 1926 (132)
M. macropostrongylus Y. & M., 1926 (132)
M. yorkei Baylis, 1927 (5)
Pharyngostrongylus eta J. & M., 1939 (61)
Ph. macropodus Y. & M., 1926 (132)
Spirostrongylus spirostrongylus Y. & M., 1926 (132)
(FI.) Dipetalonema annulipapillatum J. & M., 1938 (60)
D. roemeri (vy. Linstow, 1905) (60)
Filaria macropodis gigantei (3)
F. macropi majoris (18)
F. websteri Cobbold, 1879 (32)
These three Filariae are synonymous with Dipetalonema roemeri
(v. Linstow, 1905)
MAacropus sp. (a wallaby from Millmerran, Q.)
Nematoda
(ST., Str.) Labiostrongylus uncinatus (J. & M., 1939) (61)
Pharyngostrongylus eta J. & M., 1939 (61)
“LE KANGAROO DES ROCHERS’’
Protozoa
(Sp.) Balbiania sp. (‘“‘Siegant dans le tissu conjonctif”’) (19)
(Minchin (1903) regarded Balbiania as a synonym of Sarcocystis)

Acknowledgements.

I would like to thank the Librarian of the Queensland Institute of Medical
Research, Mrs. M. Macgregor, for her unfailing help with the references. The list
could not have been compiled without her cooperation. I would also like to thank
the Director, Dr. I. M. Mackerras, for his encouragement and helpful criticism of
the MS., and Mrs. J. W. Phillips for her careful and accurate typing.

Thanks are also due to Mr. W. R. Le Fanu, Librarian of the Royal College
of Surgeons, for his kindness in endeavouring to trace the elusive Mr. Webster—the
discoverer of the kangaroo filaria. Gratitude is also expressed to Mr. G. Mack,
Director of the Queensland Museum, for his help with the names of animals.

References.

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BancrorT, T. L., 1893.—Entozoal parasites. Aust. Med. Gaz., 12: 258-60.

Bayuis, H. A., 1925.—Notes on some Australian parasitic nematodes. Ann. Mag. nat.
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5. Bayuis, H. A., 1927.—Some new parasitic nematodes from Australia. Ann. Mag. nat.

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EP GOS

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18.

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Australian kangaroos. Parasitology, 30: 258-66.

DERRICK, E. H., and SmitH, D. J. W., 1939.—Unpublished observations.

DUNCAN, C., 1950.—Personal communication.

Duriz, P. H., and Riek, R. F., 1952.—The role of the dingo and the wallaby in the
infestation of cattle with hydatids (Hchinococcus granulosus (Batsch, 1786) Rudolphi,
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Hisic, H., 1869.—Beschreibung einer Filaria aus Halmaturus. Z. wiss. Zool., 20: 99-102.

FLETCHER, J. J., 1883.—Notes and Exhibits. Proc. Linn. Soc. N.S.W., 8: 388.

FUHRMANN, O., 1932.—Les Ténias des oiseaux. Mem. Univ. Neuchatel, 8: 381 pp.

GinbruTH, J. A., and Buy, L. B., 1912.—Wnteritis associated with infection of the
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HacKkeL, D. B., KINNEY, T. D., and WENDT, W., 1953.—Pathologic lesions in captive
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43.

44,

BY M. JOSEPHINE MACKERRAS. 123

HAMERTON, A. E., 1931.—Report on the deaths occurring in the Society’s Gardens during
the year 1930. Proc. zool. Soc. Lond., 1931: 527-55.

JANICKI, C. VON, 1905.—Beutlercestoden des Niederlandischen Neu-Guinea Expedition.
Zugleich einiges Neue aus dem Geschlechtsleben der Cestoden. Zool. Anz., 29: 127-31.

JOHNSTON, S. J., 1901.—Contributions to a knowledge of Australian Entozoa. I. On a
new species of Distomum from the Platypus. Proc. LINN. Soc. N.S.W., 26: 334-8.

JOHNSTON, S. J., 1913¢a.—On some trematode parasites of marsupials and of a monotreme.
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JOHNSTON, S. J., 1913b.—On some Queensland Trematodes, with anatomical observations
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JOHNSTON, S. J., 1915.—On Moreauia. mirabilis gen. et sp. nov., a remarkable trematode
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JOHNSTON, T. H., 1909.—The entozoa of Monotremata and Australian Marsupialia. No. 1.
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JOHNSTON, T. H., 1911.—The entozoa of Monotremata and Australian Marsupialia. No. 2.
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JOHNSTON, T. H., 1912a@.—On a re-examination of Krefft’s types of Cestoda. Rec. Aust.
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JOHNSTON, T. H., 1912b.—Notes on some entozoa. Proc. roy. Soc. Qd., 24: 63-91.

JOHNSTON, T. H., 1913.—Cestoda and Acanthocephala. Rep. Aust. Inst. trop. Med.,
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JOHNSTON, T. H., 1914.—Some new Queensland endoparasites. Proc. roy. Soc. Qd.,
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JOHNSTON, T. H., 1916.—A census of the endoparasites recorded as occurring in Queens-
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JOHNSTON, T. H., and DELAND, HE. W., 1929.—Australian Acanthocephala. No. 1. Census
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JOHNSTON, T. H., and Hpmonps, S. J., 1952.—Australian Acanthocephala. No. 9. Trans.
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JOHNSTON, T. H., and Mawson, P. M., 1938a.—An account of some filarial parasites of
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JOHNSTON, T. H., and Mawson, P. M., 1938b.—-Strongyle nematodes from Central
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JOHNSTON, T. H., and Mawson, P. M., 1938¢.—Scme nematodes from Australian
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JOHNSTON, T. H., and Mawson, P. M., 1939a.—Strongyle nematodes from Queensland
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JOHNSTON, T. H., and MAwson, P. M., 1939b.—Strongylate nematodes from marsupials
in New South Wales. Proc. LINN. Soc. N.S.W., 64: 513-36.

JOHNSTON, T. H., and Mawson, P. M., 1939¢—Sundry nematodes from eastern Australian
marsupials. Trans. roy. Soc. S. Aust., 63: 204-9.

JOHNSTON, T. H., and Mawson, P. M., 1939d—Some nematodes from Victorian and
Western Australian marsupials. Trans. roy. Soc. S. Aust., 63: 307-10.

JOHNSTON, T. H., and Mawson, P. M., 1940a.—On a collection of nematodes from
Australian marsupials. Rec. Aust. Mus., 20: 360-6.

JOHNSTON, T. H., and Mawson, P. M., 1940b.—Nematodes from South Australian
marsupials. Trans. roy. Soc. S. Aust., 64: 95-100.

JOHNSTON, T. H., and MAwson, P. M., 1940¢—New and known nematodes from
Australian marsupials. Proc. LINN. Soc. N.S.W., 65: 468-76.

JOHNSTON, T. H., and MAwson, P. M., 1940d.—A key to the nematode parasites of
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JOHNSTON, T. H., and Mawson, P. M., 1941a— Some parasitic nematodes in the
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JOHNSTON, T. H., and Mawson, P. M., 1941b.—Some nematodes from Kangaroo Island,
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JOHNSTON, T. H., and MAwson, P. M., 1942.—The Gallard collection of parasitic
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JOHNSTON, T. H., and MAwson, P. M., 1949.—Some nematodes from Australian hosts,
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JOHNSTON, T. H., and MAwsoNn, P. M., 1952.—Some nematodes from Australian birds
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Kerr, T., 1935.—On Linstowia echidnae (Thompson, 1893) Zschokke, 1899: a cestode
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KirRBY, H., and HONIGBERG, B., 1950.—AIntestinal flagellates from a wallaroo, Macropus
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Kreis, H. A., 1952.—Helminthologische Untersuchungen in schweizerischen Tierpairken
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124

78.

Os

80.

81.
82.
83.
84.
85.
86.
87.
88.

90.
91.

91a.

92.
93.

94.
95.

96.

100.

109.

110.

AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

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LInstow, O. von, 1897.-—Zur Systematik des Nematoden nebst Beschreibung neuer Arten.
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LINSTOW, O. VON, 1898a.—Nemathelmirnthen gesammelt von Herrn Prof. Dr. F. Dahl im
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LINstow, O. von, 1898b.—Nemathelminthen yon Herrn Richard Semon in Australien
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LINStow, O. VON, 1905.—Helminthologische Beobachtungen. Arch. mikr. Anat., 66:
355-66.

LUMHOLTZ, C., 1884.—Notes upon some mammals recently discovered in Queensland.
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MACKERRAS, I. M., MACKERRAS, M. J., and SANDARS, D. F., 1953.—Parasites of the
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MACKERRAS, M. J., 1954.—Two new species of Dipetalonema (Nematoda: Filarioidea )
from Australian marsupials. Proc. roy. Soc. Qd., 64: 51-6.

MACKERRAS, M. J., 1955.—A new lung-worm from Australian marsupials (Nematoda:
Metastrongylidae). Proc. roy. Soc. Qd., 66: 77-81.

MACKERRAS, M. J.—Unpublished observations.

MACKERRAS, M. J., and SANDARS, D. F., 1953.—Two new metastrongyle lung-worms from
Australian marsupials. Prec. roy. Soc. Qd., 63: 71-6.

Mawson, P. M., 1955—Some parasites of Australian vertebrates. Trans. voy. Soe.
S. Aust., 78: 1-7.

MeceitT, F. F., 1927.—On cestodes collected in Burma. Parasitology, 19: 141-53.

MINCHIN, H. A., 1903.—‘“Sporozoa’”’ in Ray lLankester’s Treatise on Zoology, Part 1,
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MOnnNIG, H. O., 1926.—Three new helminths. Trans. roy. Soc. S. Afr., 13: 291-8.

Monniec, H. O., 1927.—On a new Physaloptera from an eagle and a trichostrongyle from
the cane rat, with notes on Polydelphis quadricornis and the genus Spirostrongylus.
Trans. roy. Soc. 8S. Afr., 14: 261-5.

Monniec, H. O., 1929a.—Hypodontis macropi, n. gen., n. sp., a hookworm of the kangaroo.
15th Ann. Rep. Dir. Vet. Services, Union of S. Afr.: 303-6.

MONNIG, H. O., 1929b.—Filarinema flagrifer, n. gen., n. sp., a trichostrongylid parasite
of the kangaroo. 15th Ann. Rep. Dir. Vet. Services, Union of S. Afr.: 307-10.

NYBELIN, O., 1917.—Australishe Cestodon. Results of D. EH. Mjoberg’s Swedish Scientific
Expedition to Australia, 1910-1913. K. Svenska Vetensk. Akad. Handl., 52: 1-48.

OLDHAM, J. N., 1933.—The helminth parasites of marsupials. J. Helminth., 11: 195-256.

ORTLEPP, R. J., 1922.—A new davaineid cestode, Raillietina (Paroniella) macropa, sp. n.,
from a wallaby. Ann. Mag. nat. Hist., (9) 9: 602-12.

PLimMeER, H., 1912.—On certain. blood parasites. J. R. micr. Soc., 1912: 1338-50.

PLiMMER, H., 1914.—Report on the deaths which occurred in the Zoological Gardens
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Pore, J. H., DERRICK, E. H., and Cook, I., 1957.—Toxoplasma in Queensland. I. Observa-
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. Pope, J. H., Bicks, V. A., and Coox, I., 1957.—Toxoplasma in Queensland. II. Natural

infections in bandicoots and rats. Aust. J. exp. Biol., 35: 481-90.
PRIESTLEY, H., 1915.—Theileria tachyglossi (n. sp.). A blood parasite of Tachyglossus
aculeatus. Ann. trop. Med. Parasit., 9: 233-8.
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of a marsupial wolf (Thylacinus cynocephalus). Trans. Amer. micr. Soc., 27: 31-2.
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mammals. Amer. J. Path., 27: 655-67.

RUDOLPHI, C. A., 1819.—Entozoorum synopsis. Berlin, 1819, 811 pp.

SANDARS, D. F., 1953.—A study of Diphyllobothriidae (Cestoda) from Australian hosts.
Proc. roy. Soc. Qd., 63: 65-70.

SANDARS, D. F., 1956.—Mirandula parva, n: gen., n. sp. (Cestoda, Dilepididae), from
the long-nosed bandicoot (Perameles nasuta Geoff.). J. Helminth., 30: 183-8.

SANDARS, D. F.—Personal communicaticn.

1104. SANDARS, D. F., 1957a.—Redescriptions of some cestodes from marsupials. ieee ILS

Taeniidae. Ann. trop. Med. Parasit., 51: 317-29.

BY M. JOSEPHINE MACKERRAS. 125

Sanpars, D. F., 1957b.—Redescriptions of some cestodes from marsupials. Part II:
Davaineidae, Hymenolepididae and Anoplocephalidae. Ann. trop. Med. Parasit., 51:
330-39.

SANDARS, D. F., 1957¢—A new Strigeid Trematode from an Australian marsupial.
J. Helminth., 31: 257-64.

SANDARS, D. F., 1957d.—On Brachylaemus (Trematoda) from marsupials. J. Helminth.,
31: 265-72.

SANDARS, D. F., 1958.—A pancreatic fluke, Zonorchis australiensis, n. sp. (Trematoda),
from Australian marsupiais. Ann. trop. Med. Parasit. (in press).

ScHWarR?Tz, B., 1928.—Two new nematodes of the family Strongylidae, parasitic in the
intestine of mammals. Proc. U.S. nat. Mus., 73: 1-5.

Scott, H. H., 1926.—Report on the deaths occurring in the Society’s Gardens during
the year 1925. Proc. zool. Soc. Lond., 1926: 231-44.

SEDDON, H. R., 1952.—Diseases of domestic animals in Australia. Part 4. Protozoan and
viral diseases. Serv. Publ. Dep. Hith. Aust. vet. Hyg., No. 8, 214 pp.

SoLomon, S. G., 193838.—A note on a new species of Breinlia (Wilariidae) from a tree
kangaroo. J. Helminth., 11: 101-4.

Sweet, G. (1909).—The endoparasites of Australian stock and native fauna. I Census.
Proc. roy. Soe. Vict., n.s., 21: 454-502.:

THUSCHER, E. V., and Sttnzi, H., 1956.—Uber parasitologische Kotuntersuchungen bei
Saugetiere des Zoologischen Gartens Zurich. Acta trov., Basel, 13: 262-9.

THOMPSON, D’A. W., 1893.—Note on a tapeworm from echidna (Taenia echidnae, n. sp.).
J. R. micr. Soc., 1893: 297.

TRIFFITT, M. J., 1926.—Some sporozoan parasites found in the intestinal wall of Bennett’s
wallaby (Macropus bennetti). Protozoology, 2: 31-46.

WARDLE, R. A., and McLeEop, J. A., 1952.—The zoology of tapeworms. Univ. Minnesota
Press, Minneapolis, 1952, 780 pp.

WELSH, D. A., and BARLING, J. E. V., 1909.—Haemogregarina petauri: a haemogregarine
of a marsupial flying squirrel. Aust. Med. Congr., 1908, 2: 329-33.

WELSH, D. A., and BaRuIneG, J. HE. V., 1910.—Haemogregarina petauri: a haemogregarine
of a marsupial flying squirrel. J. Path. Bact., 14: 536-41.

WELSH, D. A., DALYELL, E. J., and Burritt, M. B., 1909..—Haemogregarina dasyuri:
a preliminary note on an undescribed haemogregarine of the Australian native cat.
Aust. Med. Congr.. 1908, 2: 333-7.

WELSH, D. A., DAatyetu, E. J... and Burrirr, M. B., 1910.—Haemogregarina dasyuri: a
haemogregarine of the Australian native cat. J. Path. Bact., 14: 542-6.

WELSH, D. A., and DALYELL, E. J., 1909.—Haemogregarina peramelis: a free haemo-
gregarine of an Australian bandicoot. J. Univ. Syd. med. Soc., 2: 112-5. ,

WeELSH, D. A., and Datyvett, FE. J., 1910.—Haemogregarina peramelis: a free haemo-
greearine of an Australian bandicoot. J. Path. Bact., 14: 547-9.

WENYON, C. M., 1926.—Protozoology. MBailliére, Tindall and Cox, London, 1926. 2 vol.,
1563 pp.

WENYON, C. M., and Scorr, H. H., 1925.—(Demonstration of sections of intestine of
Bennett’s wallaby). Trans. roy. Soc. trop. Med. Hyg., 19: 7-8.

Woop, W. A., 1929a—On a new species of Labiostrongylus. Ann. Mag. nat. Hist.,
(10) 4: 550-1.

Woop, W. A., 1929b.—A note on Rugopharynsx australis (Monnig, 1926). Ann. Mag. nat.
Hist., (10) 4: 552-4.

Woop, W. A., 1931.—Some new parasitic nematodes from Western Australia. Rep. Inst.
Anim. Path. Univ. Camb., 1: 209-19.

Woop Jones, F., 1948.—The study of a Generalized Marsupial (Dasycercus cristicauda
Krefft). Trans. zool. Soc., 26: 409-501. ?

YORKE, W., and MapuEesTonn, P. A., 1926.——The nematode parasites of vertebrates.
J. and A. Churchill, London, 1926. 536 pp.

Younec, M. R., 1939.—Helminth parasites of Australia. Imp. Bur. Agric. Parasitol.
(Helminthology) England. Pp. 145.

ZSCHOKKE, F., 1896.—Die Taenien der aplacentalen Sdugethiere. Zool. Anz., 19: 481-2.

ZSCHOKKE, F., 1898a—Die Cestoden der Marsupialia und Monotremata. Denkschr.
med-naturw. Ges. Jena, 8: 358-80.

ZSCHOKKE, F., 1898b—Weitere Untersuchungen an Cestoden aplacentaler Saugethiere.
Zool. Anz., 21: 477-9.

ZSCHOKKE, F., 1899.—Neue Studien an Cestoden aplacentaler Sdugethiere. Z. wiss. Zool.,
65: 404-45.

ZSCHOKKE, F., 1907.—Moniezia diaphana, n. sp. Hin weiterer Beitrag zur Kenntnis der
Cestoden aplacentaler Sdugethiere. Zbl. Bakt., 44: 261-4.

126

PART II. EUTHERIA.

Synopsis.

Part II contains the names of 171 animals, eight of them introduced by man. There
are 85 native rats belonging to 17 genera; parasites have been recorded from 8 species
belonging to 4 genera. There are 49 native bats, including flying foxes, belonging to 24
genera; parasites have been recorded from 10 species belonging to 5 genera. EXxtra-
territorial records for 7 bats belonging to 4 genera are included. Other records are from
the dingo and certain marine mammals.

Blood protozoa are known to occur in three native rats (Trypanosoma, Hepatozoon, and
Bartonella), in flying foxes (Trypanosoma and Hepatocystis) and in insectivorous bats
(? Polychromophilus). Trematodes are known from a water rat (3 species), small bats
(several undescribed species), the dugong (10 species), and a seal (one species). Cestodes
are known from some rats and bats, and are common in carnivorous marine mammals.
Nematodes occur in all groups. Acanthocephala are recorded from two native rats and
several marine animals.

The parasites of the introduced animals are the same as those they harbour in other
parts of the world. The dingo has been found to be infested with many of the parasites
of the domestic dog.

This part deals with the native Eutherian mammals and the following introduced
animals: rabbit, hare, brown and black rats, mouse, dog, cat, and fox. The parasites
of some of the introduced animals were recorded by Johnston in a series of papers
and exhibits from 1909 to 1918. Some were recorded by Sweet (1909a, 19090), and the
helminths were brought together by Young (1939). The parasites of the dog and fox
have been dealt with by Pullar (1946), and Seddon (1950, 1952) listed parasites from
all this group except the mouse and rats.

The records made by Johnston and his colleagues from marine mammals in the
Antarctic and in the sub-Antarctic islands are included here, the hosts being occasional
visitors to our southern shores. Extra-territorial records for the dugong are included,
and also for some bats, because the Australian representatives of these wide-ranging
mammals may be hosts to the same or related parasites.

Relatively few records have been found for the indigenous Eutheria. This appears
to be due to the lack of examinations rather than to any deficiency in the parasitic
fauna, since a few species which have been carefully examined have been found to
harbour many parasites.

The explanation of the abbreviations used will be found at the beginning of
Part I (p. 102).

Hosts AND PARASITES
Order LAGOMORPHA
Family LErPorIDAE
Genus Lepus L., 1758
L. EUROPAEUS Pallas, 1778, the hare* (introduced)
Protozoa
(Sp.) EHimeria perforans (Leuckart, 1879) (98)
Trematoda
(Fasci.) Fasciola hepatica L., 1758 (20), (105), (97)
Cestoda
(CY., Tae.) Taenia pisiformis Bloch, 1780, as Cysticercus pisiformis (15), (109),
(45), (97)
Multiceps serialis (Gervais, 1847), as Coenurus serialis (109), (45), (97)

*'The species of hare which has gone wild in Australia is sometimes referred to as
Lepus timidus. However, Mr. G. Mack, Queensland Museum, considers that it should be
ealled ZL. europaeus.

BY M. JOSEPHINE MACKERRAS. 127

Genus OrycToLacus Lilljeborg, 1874
QO. CUNICULUS (L., 1758), the rabbit (introduced)

Protozoa
(Sp.) Himeria stiedae (Lindemann, 1865) (38), (39), (99), (45), (98), (79)

E. perforans (Leuckart, 1879) (99), (98), (79)
#. irresidua Kessel and Jankiewicz, 1931 (79)
#H. media Kessel, 1929 (79)
E. magna Pérard, 1928 (99), (98), (79)
Toxoplasma sp. (111), (98)
Trematoda
(Fasci.) Fasciola hepatica L., 1758 (20), (105), (97), (79)
Cestoda
(CY., Tae.) Taenia pisiformis Bloch, 1780, as Cysticercus pisiformis (109), (88),
(39), (103), (45), (93), (97), (79)
Multiceps serialis (Gervais, 1847) as Coenurus serialis (103), (38), (39),
(KO)! pe 25) ee 259 0 DD)
Echinococcus granulosus (Batsch, 1786) as hydatid (103), (38), (39),
(109), (45), (97)
(PS., Dip.) Diphyllobothrium (Spirometra) erinacei (Rudolphi, 1819) as sparganum
(experimental) (10)
Nematoda
(ST., Tri.) Trichostrongylus colubriformis (Giles, 1892) (90)
T. retortaeformis (Zeder, 1800) (90), (63), (79)
T. vitrinus Looss, 1905 (90)
Graphidium strigosum (Dujardin, 1845) (38), (109), (63), (45), (79)
Nematodirus spathiger (Railliet, 1896) (79)
Nematodirus sp. (79)
(OX.) Passalurus ambiguus (Rudolphi, 1819) (41), (45), (90), (63), (79)

Order RODENTIA
Family MurRIDAE
Subfamily HyDROMYINAE

Genus Hypromys Geoffroy, 1804. (Water rats)
H. cauRiInus Thomas, 1909 No records
H. CHRYSOGASTER Geoffroy, 1804
Trematoda
(Plagi.) Plagiorchis jaenschi Johnston and Angel, 1951 (51), (52)
(Micro.) Microphallus minutus Johnston, 1948 (51)
(Strig.) Fibricola minor Dubois, 1936 (27), (51)
Cestoda
(CY., Hym.) Hymenolepis diminuta (Rudolphi, 1819) (30)
(PS., Dip.) Diphyllobothrium (Spirometra) erinacei (Rudolphi, 1819) as sparganum*
(72) )
Nematoda
(TR.) Trichuris muris (Schrank, 1788) (30)
Capillaria sp.* (72)
(ST., Tri.) Trichostrongylidae, undescribed species* (72)
(ST., Anc.) Ancylostomidae, undescribed species (72)

(OX.) Ganguleterakis spumosa (Schneider, 1866) (30)
(AS.) : Neoascaris sp. (102)
(SP.) Cosmocephalus australiensis J. & M., 1952 (65)

Spirura (s.1.) sp. (65)
Protospirura muris (Gmelin, 1790) (30)
Gongylonema sp. (30)
*T am indebted to Mrs. D. G. Delamoir, Queensland Institute of Medical Research Field
Station, Innisfail, North Queensland, for the species marked with an asterisk from the
various native rats.

128 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

Acanthocephala

Pseudoporrorchis hydromuris* Hdmonds, 1957 (294)
Pentastomida (Phylum Arthropoda)

Linguatula sp.* (larvae) (72)

H. FULIGINOSUS Gould, 1853 No records
H. GROOTENSIS Troughton, 1935 No records
H. LAWNENSIS Troughton, 1923 No records
H. LonecmMAanrt Thomas, 1923 No records
H. MELICERTES Thomas, 1921 No records
H. moare Troughton, 1935 No records
Genus XEROMYS Thomas, 1889
X. MYOIDES Thomas, 1889 ; No records

Subfamily Murinar
Genus Rarrus Fischer, 1803. (Rats)
R. ASSIMILIS (Gould, 1858)—the allied rat
Protozoa
(M.) Trypanosoma lewisi (Kent, 1880) (72)
(Sp.) Bartonella muris Mayer, 1921 (72)
Hepatozoon muris (Balfour, 1906) (72)
Toxoplasma gondii (Nicolle and Manceaux, 1908) (83)
Cestoda
(CY., Hym.) Hymenolepis diminuta (Rudolphi, 1819) (95)
Hymenolepis australiensis Sandars, 1957 (95)
(CY., Dav.) Raillietina (Raillietina) celebensis (Janicki, 1902) (1), (95)
(CY., Dil.) Choanotaenia ratticola Sandars, 1957 (95)
(CY., Tae.) Taenia taeniaeformis (Batsch, 1786) as Cysticercus fasciolaris Rudolphi,

1808 (95)
Nematoda
(RH.) Strongyloides sp. (72)
(TR.) Capillaria sp. (72)

(ST., Tri.) Trichostrongylidae, sp. not identified (72)
(ST., Met.) Angiostrongylus cantonensis (Chen, 19385) (72)

(OX.) Unidentified species* (72)
(AS.) Neoascaris mackerrasae Sprent (101)
Amplicaecum sp. (larvae) (102)
(SP.) Physaloptera troughtoni J. & M., 1941 (60)
Acanthocephala
Unidentified species* (72)
R. coLuerttr1 Thomas, 1904 No records
R. conatus Thomas, 1923
Nematoda

(ST., Tri.) Nippostrongylus braziliense* (Travassos, 1914) (72)
Trichostrongylidae (2 unidentified species*) (72)
(ST., Met.) Angiostrongylus cantonensis* (Chen, 1935) (72)

Cestoda
Unidentified species* (72)
R. cutmorum (Thomas and Dollman, 1909) No records
R. Fruscrers (Waterhouse, 1839) No records
R. GREYII (Gray, 1841) No records
R. LEucopus (Gould, 1867) No records
R. LUTREOLUS (Gray, 1841) No records
R. MANIcCATUS (Gould, 1858) No records
R. MELVILLEUS Thomas, 1921 No records
R. sorpipus (Gould, 1858) No records
R. TUNNEYI (Thomas, 1904) No records
R. VELLEROSUS (Gould, 1847) No recerds

BY M. JOSEPHINE MACKERRAS. 129

R. VILLOSISSIMUS (Waite, 1897)
_ Protozoa
(M.) Trichomonas sp. (72)
Hexamita sp. (72)
Giardia sp. (72)

(Sa.) Entamoeba sp. (72)
(Sp.) Bartonella muris Mayer, 1921 (17)
Cestoda

Unidentified larvae in lungs (72)
R. norvecicus (Berkenhout, 1769), the brown rat (introduced)
Protozoa

(M.) Trypanosoma lewisi (Kent, 1880) (85), (88), (55), (45), (30)
Trichomonas muris (Grassi, 1879) (72)
Hexamita muris (Grassi, 1881) (72)

(Sa.) Entamoeba muris (Grassi, 1879) (72)

(Sp.) Hepatozoon muris (Balfour, 1906) (18), (38), (89), (55), (45)
EHimeria nieschulzi Dieben, 1924 (72)
Coccidia (30)
Sarcocystis muris (Blanchard, 1885) (38), (55), (45)
Bartonellé muris Mayer, 1921 (72)
Toxoplasma gondii (Nicolle and Manceaux, 1908) (83)

Cestoda
(CY., Tae.) Taenia taeniaeformis (Batsch, 1786) as Cysticercus fasciolaris Rudolphi,
1808 (68), (82), (88), (109), (45), (380)

(CY., Hym.) Hymenolepis diminuta (Rudolphi, 1819) (38), (39), (109), (45)
H. nana (v. Siebold, 1853) (39), (109), (45) :

(CY., Dav.) Raillietina (R.) celebensis (Janicki, 1902) recorded as Davainea sp. (45)

(PS., Dip.) Diphyllobothrium (Spirometra) erinacei (Rudolphi, 1819) as sparganum

(experimental) (10), (94)

Nematoda
(RH.) Strongyloides sp. (30)
(TR.) Trichuris muris (Schrank, 1788) (30)

Capillaria hepatica (Bancroft, 1893) (89), (109), (45), (80)
Trichosomoides crassicauda (Bellingham, 1840) (39), (109), (45)
(ST., Tri.) Nippostrongylus braziliense (Travassos, 1914) (45), (30), recorded as
Oesophagostomum sp. (39), (109)
(ST., Met.) Angiostrongylus cantonensis (Chen, 1935) (73)
(OX.) Syphacia obvelata (Rudolphi, 1802) (39), (109), (45)
; Aspiculuris tetraptera (Nitzsch, 1821) (46)
Ganguleterakis spumosa (Schneider, 1866) (389), (109), (45), (30)

(SP.) Protospirura muris (Gmelin, 1790) (39), (109), (45)
Gongylonema sp. (30)
Acanthocephala

Moniliformis dubius Meyer, 1932 (38), (39), (109), (45), (30), (57)
R. RATTUS (L., 1758), the black rat (introduced), including R. R. alexandrinus
Protozoa
(M.) Trypanosoma lewisi (Kent, 1880) (85), (388), (55), (45), (30)
(Sp.) Hepatozoon muris (Balfour, 1906) (38), (55), (45)
Coccidia (380)
Sarcocystis muris (Blanchard, 1885) (39), (55), (45)
Cestoda
(CY., Tae.) Taenia taeniaeformis (Batsch, 1786) as Cysticercus fasciolaris Rudolphi,
1808 (39), (109), (45)
(CY., Hym.) Hymenolepis diminuta (Rudolphi, 1819) (38), (39), (109), (45), (30)
H. nana (v. Siebold, 1853) (39), (109), (45), (30)

Nematoda
(RH.) Strongyloides sp. (72)

130 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

(TR.) Trichuris muris (Schrank, 1788) (389), (109), (45), (30)
Capillaria hepatica (Bancroft, 1893) (89), (109), (45), (30)
Trichosomoides crassicauda (Bellingham, 1840) (89), (45)

(ST., Tri.) Nippostrongylus braziliense (Travassos, 1914) (46)

(ST., Met.) Angiostrongylus.cantonensis (Chen, 19385) (73)

(OX.) Syphacia obvelata (Rudolphi, 1802) (389), (109), (45), (30)
Ganguleterakis spumosa (Schneider, 1866) (39), (109), (45)
(SP.) Protospirura muris (Gmelin, 1790) (89), (109), (45)
Gongylonema sp. (30) ;
Acanthocephala

Moniliformis dubius Meyer, 1932 (38), (39), (109), (45), (30), (57)

Rattus sp. (introduced)

M.

Ge acIlaclias a= Iac a

Protozoa
(M.) Trypanosoma lewisi (Kent, 1880) (19), as Haematomonas (3)
(Sp.) Hepatozoon muris (Balfour, 1906) as Leucocytozoon sp. (19)
Cestoda
(CY., Tae.) Taenia taeniaeformis (Batsch, 1786) as Cysticercus fasciolaris Rudolphi,
1808 (68), (82)
Nematoda
(TR.) Capillaria hepatica (Bancroft, 1898) (4)

Genus Mus L., 1758

MUSCULUS L., 1758, the house mouse (introduced )
Protozoa
(M.) Trichomonas muris (Grassi, 1879) (24)
Hexamita muris (Grassi, 1881) (72)
Giardia muris (Grassi, 1879) (24)
(Sa.) Entamoeba muris (Grassi, 1879) (24)
(Sp.) Klossiella muris Smith and Johnston, 1902 (24)

Himeria falciformis (Himer, 1870) (72)
Eperythrozoon coccoides Schilling, 1928 (24), (25)
Cryptosporidium muris Tyzzer, 1907 (24)
Cestoda ;
(CY., Tae.) Taenia taeniaeformis (Batsch, 1786) as Cysticercus fasciolaris Rudolphi,
1808 (82), (39), (109), (45)
(CY., Hym.) Hymenolepis diminuta (Rudolphi, 1819) (38), (39), (109), (45)
H. nana (v. Siebold, 1853) (39), (109), (45), (24)
(PS., Dip.) Diphyllobothrium (Spirometra) erinacei (Rudolphi, 1819) as sparganum
experimental (10), (94)
Nematoda
(TR.) Trichuris muris (Schrank, 1788) (39), (109), (45)
Capillaria hepatica (Bancroft, 18938) (39), (109), (45)
Trichosomoides crassicauda (Bellingham, 1840) (39)

(OX.) Syphacia obvelata (Rudolphi, 1802) (389), (109), (45), (24)
Aspiculuris tetraptera (Nitzsch, 1821) (45), (46), (24)
Ganguleterakis spumosa (Schneider, 1866) (39), (109), (45)

(SP.) Protospirura muris (Gmelin, 1790), (39), (109), (45), (60)
Gongylonema sp. (45), (30)

Genus PsEupoMys Gray, 1832. (Native mice)

. AUSTRALIS Gray, 1832 No records
. AURITUS Thomas, 1910 - No records
FIELDI Waite, 1896 No records
. HIGGINSI (Trouessart, 1899) No records
MINNIE Troughton, 1932 No records
. RAWLINNAE Troughton, 1932 No records

. SHORTRIDGEI (Thomas, 1907) No records

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BY M. JOSEPHINE MACKERRAS.

Genus THETomMys Thomas, 1910. (Native mice)

. FERCULINUS (Thomas, 1902) No records
. GOULDII (Waterhouse, 1839) No records
. GRACILICAUDATUS (Gould, 1845) No records
NANUS (Gould, 1858) No records
PRAECONIS Thomas, 1910 No records

Genus LEecgapIna Thomas, 1910. (Native mice)

BERNEYI Troughton, 1936 No records
. DELICATULA (Gould, 1842) No records
FIELDI (Waite, 1896) No records
FORRESTI (Thomas, 1906) No records
HERMANNSBURGENSIS (Waite, 1896) No records
MESSORIA Thomas, 1925 No records
PATRIA (Thomas and Dollman, 1909) No records
WAITEI Troughton, 1932 No records

Genus Gyomys Thomas, 1910. (Native mice)

ALBOCINEREUS (Gould, 1845) No records
APODEMOIDES Finlayson, 1932 No records
. DESERTOR Troughton, 1932 No records
FUMEUS Brazenor, 1934 No records
GLAUCUS Thomas, 1910 “ No records
NOVAEHOLLANDIAE (Waterhouse, 1843) No records
. OCCIDENTALIS Tate, 1951 No records
. PUMILUS Troughton, 1936 No records

Genus Mastacomys Thomas, 1882. (Broad-toothed rat)

. Fuscus Thomas, 1882 No records

Genus Laomys Thomas, 1909. (Thick-tailed rats)

. PEDUNCULATUS (Waite, 1896) No records
. WOODWARDI Thomas, 1909 ; No records

Genus ASCOPHARYNX Waite, 1900. (Kangaroo mice)

. CERVINUS (Gould. 1853) No records

Genus MESEMBRIOMYS Palmer, 1906. (Shaggy rabbit-rats)

. GOULDII (Gray, 1843) No records
. MACRURUS (Peters, 1876) No records

Genus ConiLturus Ogilby, 1838. (Rabbit-rats)

. ALBIPES (Lichtenstein, 1829) No records
. HEMILEUCURUS (Gray, 1858) No records
. PENICILLATUS (Gould, 1842) No records

Genus LEporRILLUS Thomas, 1906. (Stick-nest rats)

. APICALIS (Gould, 1853) No records
. CONDITOR (Sturt, 1848) No records
. JONESI Thomas, 1921 No records

Genus Notromys Lesson, 1842. (Kangaroo-mice)

. ALEXIS Thomas, 1922 No records
. AISTONI Brazenor, 1934 ; No records
. AQUILO Thomas, 1921 No records
. AMPLUS Brazenor, 1936 No records
. LONGICAUDATUS (Gould, 1844) No records
. MEGALOTIS Iredale and Troughton, 1934 No records

(Nomen novum for macrotis Thomas, 1921)

131

132 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

N. MITCHELLI (Ogilby, 1838) ' No records
N. morDAx Thomas, 1922 No records
N. RICHARDSONII (Gould, 1853) No records

Genus Zyzomys Thomas, 1909. (White-tailed rat)
Z. ARGURUS (Thomas, 1889) No records

Genus Metomys Thomas, 1922. (Scale-tailed tree rats)
M. AUSTRALIUS Thomas, 1924 No records
M. BANFIELDI (de Vis, 1907)
Nematoda
(SP.) Physaloptera banfieldi J. & M., 1941 (60)
M. cALLtoprs Finlayson, 1943 No records
M. CERVINIPES (Gould, 1852)
Nematoda
(FI.) Microfilaria in blood;
M. LimIcAuDA Troughton, 1935 No records
M. LirroraLtis (LOnnberg, 1916)
Protozoa
(Sp.) Bartonella muris* Mayer, 1921 (72)
Nematoda
(ST., Tri.) Trichostrongylidae, undescribed species* (72)
(ST., Met.) Angiostrongylus cantonensis® (Chen, 1935) (72)
(OX.) Species not identified* (72)
Cestoda
Species not identified* (72)
Pentastomida (Phylum Arthropoda)
Linguatula sp.* (larvae) (72)

M. meExticus (Thomas, 1913) No records
M. mixtus Troughton, 1935 No records
M. murInus (Thomas, 1913) No records
M. rusicoLaA Thomas, 1924 No records

Genus Uromys Peters, 1867. (Giant scale-tailed rats)
U. CAUDIMACULATUS (Krefft, 1867)

Nematoda
-(ST., Tri.) Trichostrongylidae, undescribed species*
U. ExILis Troughton and Le Souef, 1929 No records
U. SHERRINI Thomas, 1923 No records

Order SIRENIA
Family DUGONGIDAE
Genus Ducone Lacépéde, 1799
Syn. Halicore Illiger, 1811
D. puGcon (Miiller, 1776), the dugong—syn. australis Owen, 1847
Trematoda
(Param. ) Solenorchis travassosi Hilmy, 1949 (35)
S. gohari Hilmy, 1949 (35)
S. naguibmahfouzi Hilmy, 1949 (35)
S. baeri Hilmy, 1949 (35)
Indosolenorchis hirudinaceus Crusz, 1951 (22)
(Prono.) Taprobanella bicaudata Crusz and Fernand, 1954 (23)
Lankatrema mannarense Crusz and Fernand, 1954 (23)

Opisthotrema dujonis (Leuckart, 1874), syn. cochleare Fischer, 18838 (37)

Pulmonicola pulmonale (vy. Linstow, 1904) (71)
(Rhab. ) Rhabdiopoeus taylori S. J. Johnston, 1918 (37)
Nematoda
(AS.) Paradujardinia halicoris (Owen, 1833) (59)

+ See footnote under H. chrysogaster.

BY M. JOSEPHINE MACKERRAS. 133

Order CARNIVORA
Suborder FISSIPEDIA
Family FELIDAE
Genus Ferris L., 1758
F. catus L., 1758, the domestic cat (introduced)
Protozoa
(Sp.) Toxoplasma sp. (111), (98)
Isospora felis Wenyon, 1923 (11)
I. rivolta (Grassi, 1879) (11)
Trematoda
Four species known to occur in Brisbane cats (102)
Cestoda
(CY., Tae.) Taenia taeniaeformis (Batsch, 1786) (82), (15), (89), (89), (97)
(CY., Dil.) Dipylidium caninum (L., 1758) (82), (388), (39), (109), (89), (97)
(PS., Dip.) Diphyllobothrium (Spirometra) erinacet (Rudolphi, 1819), syn. Dibothrio-
cephalus felis (Creplin, 1825) (21), (109), (42), (97), (9), (10),
(94)
Nematoda
(ST., Ane.) Ancylostoma caninum (Ercolani, 1859) (42), (14), (89), (97)
A. braziliense (Gomez de Faria, 1910) (33), (97)
(ST., Met.) Aelurostrongylus abstvusus (Railliet, 1898) (31), (97)
(AS.) Toxocara cati (Schrank, 1788), syn. mystax Zeder, 1800 (68), (82), (89),
(97), (100)
Toxascaris leonina (v. Linstow, 1902) (only found once) (97)
(SP.) Gnathostoma spinigerum Owen, 1836 (33), (97)

Family CANIDAE
Genus Canis L., 1758
C. FAMILIARIS L., 1758, the domestic dog (introduced)
Protozoa
(Sp.) Isospora bigemina (Stiles, 1891) (72)
I. felis Wenyon, 1923 (11)
I. rivolta (Grassi, 1879) (11)
Coccidiosis (species not specified) (98)
? Hepatozoon sp. (16), (98)
Toxoplasma sp. (111), (98)
Trematoda
(Diplo. ) Alaria alata (Goeze, 1782) -(mot indigenous) (74), (97)
Cestoda
(CY., Dil.) Dipylidium caninum (L., 1758) (82), (15), (88), (389), (109), (92),
(89), (97)
(CY., Tae.) Echinococcus granulosus (Batsch, 1786) (106), (107), (108), (82), (39),
(109), (92), (93), (89), (97)
Multiceps serialis (Gervais, 1847) (21), (39), (109), (92), (97)
Taenia hydatigena Pallas, 1766, syn. marginata Batsch, 1786 (15), (38),
(109), (89), (97)
T. ovis (Cobbold, 1869) (109), (91), (88), (97)
T. pisiformis (Bloch, 1780), syn. serrata Goeze, 1782 (15), (21), (89),
(109), (92), (89), (97)
T. taeniaeformis (Batsch, 1786), syn. crassicollis Rudolphi, 1810 (92)
(PS., Dip.) Diphyllobothrium (Spirometra) erinaceit (Rudolphi, 1819) (88), (97),

(10)
Diphyllobothrium latum (L.. 1758) (32), (97)
Nematoda
(TR.) Trichuris vulpis (Frohlich, 1789) (92), (88), (97)

(ST., Ane.) Ancylostoma caninum (HErcolani, 1859) (82), (109), (80), (92), (89), (97)
A. braziliense (Gomez de Faria, 1910) (33), (97)
Uncinaria stenocenhala (Railliet, 1894) (109), (92), (88), (97)

134 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

(ST., Met.)

(AS.)

(SP.)
(FI.)

Angiostrongylus vasorum (Baillet, 1866) (91), (97)

Filaroides osleri (Cobbold, 1879) (109), (88), (97), (67)

Toxascaris leonina (v. Linstow, 1902), syn. limbata Railliet and Henry,
1911 (92), (89), (97)

Toxocara canis (Werner, 1782), syn. marginata Rudolphi, 1802 (82),
(39), (109), (92), (89), (97)

Spirocerca sanguinolenta Rudolphi, 1819 (not indigenous) (40), (109)

Dirofilaria immitis (eidy, 1856) (2). (5), (6), (88), (109), (14), (80),
(89), (97)

Pentastomida (Phylum Arthropoda)

Linguatula serrata (Frohlich, 1789) (86)

C. ANTARCTICUS Kerr, 1792, the dingo— syn. C. dingo Meyer, 1793

Protozoa

(Sp.)

Cestoda

(CY., Dil.)
(CY., Tae.)

(PS., Dip.)

Himeria canis Wenyon, 1923 (11)
Isospora rivolta Grassi, 1879 (11)

Dipylidium caninum (L., 1758) (109), (44), (97), (28)
Echinococcus granulosus (Batsch, 1786) (39), (109), (44), (97), (28)
Multiceps serialis (Gervais, 1847) (28)

Taenia hydatigena Pallas, 1766 (28)

T. pisiformis (Bloch, 1780) (28)

Diphyllobothrium (Spirometra) erinacet (Rudolphi, 1819) (88), (28)

Nematoda

(ST., Anc.)
(AS.)

Ancylostoma caninum (Hrcolani, 1859) (44), (97)
Toxascaris leonina (v. Linstow, 1902) (84)

Pentastomida (Phylum Arthropoda)

Linguatula dingophila Johnson, 1910 (36)
? = L. serrata (Frohlich, 1789) (44), (28)

Genus VuLres Skjoldebrand, 1777

V. VULPES (L., 1758), the fox (introduced)

Cestoda

(CY., Dil.)
(CY., Tae.)

(PS., Dip.)

Dipylidium caninum (L., 1758) (88), (97)

Multiceps serialis (Gervais, 1847) (39), (88), (97)

Taenia hydatigena Pallas, 1766 (88), (97)

T. ovis (Cobbold, 1869) (87), (97)

T. pisiformis (Bloch, 1780) (97)

Diphyllobothrium (Spirometra) erinacei (Rudolphi, 1819) (88), (97),
as sparganum (42)

Nematoda
(ST., Anc.) Uncinaria stenocephala (Railliet, 1894) (88), (97), (65)

(AS.)

(FI.)

Toxascaris leonina (v. Linstow, 1902) (88), (97)
Toxocara canis (Werner, 1782) (88), (97)
Dirofilaria immitis (Leidy, 1856) (97)

Suborder PINNIPEDIA
Family OTARIIDAE
Genus NroPpHOCcA Gray, 1866

N. CINEREA (Péron and Lesueur, 1816), the hair seal

Cestoda

(PS., Dip.) Diphyllobothrium (Cordicephalus) arctocephalinum Johnston, 1937 (48)
Nematoda

(AS.) Contracaecum osculatum (Rudolphi, 1802) (48), (59)

Acanthocephala
Corynosoma australe Johnston, 1937 (48), (54)

BY M. JOSEPHINE MACKERRAS. 135

Genus OTARIA Péron, 1816
O. FORSTERI Lesson, 1828

Nematoda
(AS.) Terranova piscium (Rudolphi, 1802) (66)
Contracaecum osculatum (Rudolphi, 1802) (76)
O. HOOKERI Gray, 1859
Nematoda
(AS.) Terranova piscium (Rudolphi, 1802) (66)

Genus GyPsopHocA Gray, 1866
G. DORIFERA (Wood Jones, 1925)
Acanthocephala

Corynosoma clavatum Goss, 1940 (54)
G. TASMANICA (Scott and Lord, 1926)

Cestoda

(PS., Dip.) Diphyllobothrium (Cordicephalus) arctocephalinum Johnston, 1937 (26).
Nematoda

(AS.) Contracaecum osculatum (Rudolphi, 1802) (59), (65)

Stomachus sp. (59), (65)

Family PHOCIDAE
Genus Lospopon Gray, 1844
L. CARCINOPHAGUS (Hombron and Jacquinot, 1842), the crab-eating seal
Nematoda
(AS.) Contracaecum osculatum (Rudolphi, 1802) (62)
C. radiatum (v. Linstow, 1907) (50), (62)

Genus HyprurGa Gistel, 1848
H. LEPTONYX (Blainville, 1820), the sea-leopard
Cestoda
(PS., Dip.) Diphyllobothrium (Cordicephalus) quadratum (vy. Linstow, 1893) (49)
D. (C.) scoticum (Rennie and Reid, 1912) (49)
Nematoda
(ST., Met.) - Parafilaroides kydrurgae Mawson, 1953 (76)
(AS.) Contracaecum osculatum (Rudolphi, 1802) (50), (€2), (76)
C. radiatum (v. Linstow, 1907) (62), (76)
C. stenocephalum (Railliet and Henry, 1907) (50)
Stomachus similis (Baird, 1853) (50), (59), (66), (76)
Terranova piscium (Rudolphi, 1802), (50), (66), (76)
Acanthocephala
Corynosoma bullosum (v. Linstow, 1892) (29)

Genus LEPTONYCHOTES Gill, 1872
L. WEDDELLII (Lesson, 1826), Weddell’s seal

Trematoda
(Notoc.) Ogmogaster antarctica Johnston, 1931 (47)
Cestoda
(PS., Dip.) Diphyllobothrium (Cordicephalus) lashleyi (Leiper and Atkinson, 1914)
(49)
D. mobile (Rennie and Reid, 1912) (49)
D. perfoliatum (Railliet and Henry, 1912) (49)
D. rufum (Leiper and Atkinson, 1914) (49)
D. wilsoni (Shipley, 1907) (49)
Nematoda
(AS.) Contracaecum osculatum (Rudolphi, 1802) (50), (62)

C. radiatum (v. Linstow, 1907) (50), (62)

C. rectangulum (vy. Linstow, 1907) (714)

C. stenocephalum (Railliet and Henry, 1907) (50)
Terranova piscium (Rudolphi, 1802) (50), (62)

136 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

(SP.) Physaloptera guiarti Garin, 1913 (112)
Acanthocephala '
Corynosoma antarcticum (Rennie, 1907) (53)

Genus MacroruHINUS Cuvier, 1826
M. prosposcipEus (Péron and Lesueur, 1807), the elephant seal.
Usually referred to as Mirounga leonina (L., 1758)
Cestoda
(PS., Dip.) Diphyllobothrium (Cordicephalus) tectum (v. Linstow, 1892} (49)
(TE., Phy.) Phyllobothrium sp. (49)
Nematoda
(ST., Anc.) Uncinaria hamiltoni Baylis, 1933 (62)
(AS.) Contracaecum osculatum (Rudolphi, 1802) (50), (62), (76)
C. radiatum (v. Linstow, 1907) (76)
Stomachus similis (Baird, 1853) (50), (62), (76)
Terranova piscium (Rudolphi, 1802) (50), (62), (66)
(FT.) Filaria (s.1.) sp. (76)
Acanthocephala
Corynosoma bullosum (v. Linstow, 1892) (29)

Order CETACEA
(Whales, porpoises and dolphins)
Only species from which parasites have been recorded in Australia are mentioned
in this list.

Family PHYSETERIDAE
Genus Kocia Gray, 1846
K. BREVICEPS (Blainville, 1838), the pigmy sperm whale.

Cestoda
(TE., Phy.) Phyllobothrium delphini (Bose, 1802) (eysts in blubber) (58)
Nematoda
(AS.) Stomachus simplex (Rudeolphi, 1809) (58), (61)
Terranova kogiae (J. & M., 1939) (58)
(FI.) Orassicauda magna J. & M., 1939 (58)

Family PHOCOENIDAE
Genus GRAMPIDELPHIS Iredale and Troughton, 1933
G. ExILIS Iredale and Troughton, 1933, the grampus-dolphin
Nematoda
(FI.) Crassicauda grampicola J. & M., 1941 (59)

Genus GLOBICEPHALUS Hamilton, 1836
GL. VENTRICOSUS (Lacépéde, 1804), the blackfish, or pilot whale
Nematoda
(AS.) Stomachus oceanicus J. & M., 1951 (64)

Family DELPHINIDAE
Genus DELPHINUS L., 1758
D. pELPHIS L., 1758, the common dolphin—syn. forsteri Gray, 1846
Trematoda
Distoma sp. (68)
Cestoda
Tetrabothrius forsteri (Krefft, 1871) (68)
Nematoda
(ST., Met.) Halocercus delphini Baylis and Daubney, 1925 (112)
(AS.) Ascaris sp. (68)
Stomachus simplex (Rudolphi, 1809) (59)
(SP.) Echinocephalus wncinatus Molin, 1858 (probably ingested with prey) (59)

BY M. JOSEPHINE MACKERRAS.

Acanthocephala
Echinorhynchus sp. (68)
Corynosoma cetaceum Johnston & Best, 1942 (54)
Corynosoma sp. (56)

Genus LAGENORHYNCHUS, Gray, 1846
L. OBScURUS (Gray, 1828), the dolphin.
Nematoda
(AS. ) Stomachus simplex (Rudolphi, 1809) (61)

Genus TuRSIves Gervais, 1855
T. TRUNCATUS (Montague, 1815), the bottle-nosed dolphin
Nematoda

137

(ST., Met.) Halocercus lagenorhynchi Baylis and Daubney, 1925 (59)

Stenurus ovatus (v. Linstow, 1910) (112)
Acanthocephala

Corynosoma cetaceum Johnston and Best, 1942 (54)

Order CHIROPTERA
Family PTEROPODIDAE

Genus PrERopUS Brisson, 1762. (Flying foxes)
P. BRUNNEUS Dobson, 1878 No records

P. CONSPICILLATUS Gould, 1850
Protozoa
(Sp.) Hepatocystis pteropi* (Breinl, 1913) (12)
P. coLtinus Anderson, 1908 (New Hebrides)
Protozoa
(Sp. ) Hepatocystis pteropi (Breinl, 1913) (77)
P. GEDDIEI MacGillivray, 1860 (New Hebrides)
Protozoa
(Sp. ) Hepatocystis pteropi (Breinl, 1913) (77)
Nematoda :
(AS.) Toxocara pteropodis Baylis, 1936 (8)
P. GouLpIr Peters, 1867
Protozoa
(M.) Trypanosoma pteropi Breinl, 1913 (13)
(Sp. ) Hepatocystis pteropi (Breinl, 1913) (13), (12),
P. NEOHIBERNICUS Peters, 1876 (Bismarck Arch.)
Nematoda —
(FI.) Filaria hepatica v. Linstow, 1898 (70)
P. POLIOCEPHALUS Temminck, 1825
Protozoa
(Sp.) Hepatocystis pteropi (Breinl, 1913) (72)
Cestoda
(CY., Hym.) Hymenolepis sp. (48)
Nematoda
(FI.) Filaria sp. (body cavity) (43)
P. SCAPULATUS Peters, 1862
Protozoa
(Sp.) Hepatocystis nteropi (Breinl, 1913) (12), (69)

Genus Dossonta Palmer, 1898

(69), (75)

D. mMAcNA Thomas, 1905 No records

D. MOLUCCENSIS (Quoy and Gaimard, 1830) (New Guinea)
Protozoa
(Sp.) ? Hepatocystis sp. (75)

*'The first reference to this parasite was made by O’Brien (1909), who referred to the

presence of “enhaemamoebae”’ in the blood of a flying fox at Cairns.
to the quartan malaria parasite of man.

He noted its resemblance

138 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

Genus NycTIMENE W.A., 1797. (Tube-nosed fruit bats)
N. PAPUANUS Anderson, 1910 No records
N. ROBINSONI Thomas, 1904 No records

Family KIopOTIDAE
Genus Syconycrerts Matschie, 1899. (Blossom bats)
S. AUSTRALIS (Peters, 1867) No records

Genus ODONTONYCTERIS Jentink, 1902. (Northern blossom bats)
O. LAGOCHILUS PYGAMAEUS (Anderson, 1911) No records

Suborder MIcROCHIROPTERA
Family RHINOLOPHIDAE
Genus RHINOLOPHUS Gray, 1834. (Horseshoe bats)
R. MEGAPHYLLUS Gray, 1834 No records

Family HIPPOSIDERIDAE
Genus RHINONICTERIS Gray, 1847. (Horseshoe bats)
R. AURANTIUS (Gray, 1845) No records

Genus HIPpPOSIDEROS Gray, 1831. (Horseshoe bats)

H. ALBANENSIS Gray, 1866 No records
H. cERvINUS (Gould, 1854) No records
H. DIADEMA REGINAE Troughton, 1937 No records
H. SEMONI Matschie, 1903
Protozoa
(Sp.) ? Polychromophilus* sp. (72)
H. steEnotTiIs Thomas, 1913 No records
Family MreéaApERMIDAE
Genus MAcroprerMA Miller, 1906. (False vampire bats)
M. gigas (Dobson, 1880) No records
Family VESPERTILIONIDAE
Genus NyctopHiLtus Leach, 1821. (Long-eared bats)
N. BiIrAx Thomas, 1915 No records
N. DAEDALUS Thomas, 1915 No records
N. GEOFFROYI Leach, 1821
Cestoda
(CY., Ano.) Oochoristica nyctophili Hickman, 1954 (34)
N. TIMORIENSIS (Geoffroy, 1806) No records
N. WALKERI Thomas, 1892 No records

Genus VESPADELUS Iredale and Troughton, 1934. (Short-eared bats)
V. PUMILIS (Gray, 1841)
Protozoa
(Sp.) ? Polychromophilus sp. (72)
Trematoda: An unidentified species (72)
Nematoda: An unidentified species (72)

Genus PIPISTRELLUS Kaup, 1829. (Pipistrels)
P. ABRAMUS (Temminck, 1840) No records

Genus REGISTRELLUS Troughton, 1943
R. REGULUS (Thomas, 1906) No records

Genus FALSISTRELLUS Troughton, 1943
F.. TASMANIENSIS (Gould, 1858) No records

*'The malaria parasites of insectivorous bats in Australia have been tentatively assigned
to the genus Polychromophilus Dionisi, 1899, until more information has been obtained
about them.

BY M. JOSEPHINE MACKERRAS. 139

Genus CHALINOLOBUS Peters, 1866. (Lobe-lipped bats)

C. GouLpIr (Gray, 1841) No records
C. morio (Gray, 1841) No records
C. prcatus (Gould, 1852) No records
C. RoGERSI Thomas, 1909 No records
‘Genus Myotis Kaup, 1829
M. AUSTRALIS (Dobson, 1878) No records
M. macropus (Gould, 1855) No records
M. myoris (Bechstein, 1819) (Palestine)
Protozoa
(Sp.) “a haemosperidian” (78)
M. NATTERERII (Kuhl, 1819) (Palestine)
Protozoa
(Sp.) “a haemosporidian” (78)
Genus ScoTEANAx Troughton, 1943. (Broad-nosed bats)
S. RUPPELLII (Peters, 1866) No records
Genus ScoTOREPENS Troughton, 1943. (Broad-nosed bats)
S. BALSTONI (Thomas, 1906) No records
S. GREYII (Gould, 1858) No records
S. INFLUATUS (Thomas, 1924) No records
S. orion (Troughton, 1937) No records

Genus MINIopTERUS Bonaparte, 1837. (Bent-winged bats)
M. AUSTRALIS Tomes, 1858, New Hebrides

Protozoa
(Sp. ) Polychromophilus murinus Dionisi, 1899 (77)
M. BLEPOTIS (Temminck, 1840)
Protozoa
(Sp.) ? Polychromophilus sp. (72)
Trematoda
Two unidentified species (72)
Cestoda
(CY., Hym.) Hymenolepis miniopteri Sandars, 1957 (96)
Nematoda
(ST., Tri.) Anoplostrongylus heydoni Baylis, 1930 (7)
(FI.) Litomosa sp. (Heart) (72)
M. SCHREIBERSII (Natterer, 1819) (Italy to Malaya)
Protozoa
(M.) Trypanosoma vespertilionis Battaglia, 1904, Italy (110)
(Sp.) Polychromophilus melanipherus Dionisi, 1899, Italy (110)

“a haemosporidian” (Palestine) (78)
MINIOPTERUS sp.
Nematoda
(ST., Tri.) Nycteridostrongylus unicollis Baylis, 1930 (7)

Genus PuHoniscus Miller, 1905. (Dome-headed bat)
P. PAPUENSIS (Dobson, 1878) No records

Family HMBALLONURIDABR
Genus SAccoLaimus Temminck, 1841. (Free-tailed bats)

S. AUSTRALIS (Gould, 1854) No records
S. FLAVIVENTRIS (Peters, 1867) No records
S. GEORGIANUS (Thomas, 1915) No records
S. NUDICLUNIATUS (De Vis, 1905) No records

140 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

Family MonossipAaEr
Genus AuSTRONOMUS Iredale and Troughton, 1934. (Mastiff bats)
A. AUSTRALIS (Gray, 1839) No records

Genus Micronomus Iredale and Troughton, 1934

M. NORFOLKENSIS (Gray, 1839) No records
M. PLANICEPS (Peters, 1866) No records

Genus CHAEREPHON Dobson, 1874. (Northern mastiff bats)
C. cotonicus (Thomas, 1906) No records

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97. SEDDON, H. R., 1950.—Diseases of domestic animals in Australia. Part 1. Helminth
infestations. Serv. Publ. Dep. Hlth. Aust. vet. Hyg., No. 5, 223 pp

98. SEDDON, H. R., 1952.—Diseases of domestic animals in Australia. Part 4, Protozoan
and viral diseases. Serv. Publ. Dep. Hlth. Aust. vet. Hyg., No. 8, 214 pp.

99. SEDDON, H. R., and CARNE, H. R., 1927.--Incidence of coccidiosis in Australian rabbits
as determined by faecal examination. Vet. Res. Rep., 3: 33-42.

100. SprentT, J. F. A., 1956.—The life history and development of Toxocara cati (Schrank,
1788) in the domestic cat. Parasitology, 46: 54-78.

101. SprReENT, J. F. A., 1957.—A new species of Neoascaris from Rattus assimilis, with a
redefinition of the genus. Parasitology, 47: 356-60.

102. SprentT, J. F. A. (unpublished observations).

103. Sweet, G., 1909a.—The Endoparasites of stock and native fauna. Part I. Introduction,
and census of forms recorded up to date. Proc. roy. Soc. Vict., n.s., 21: 454-502.

104. Sweet, G., 1909b.—The Endoparasites of Australian stock and native fauna. Part II.
New and unrecorded species. Proc. roy. Soc. Vict., n.s., 21: 503-27.

105. Swept, G., SEDDON, H. R., and ROBERTSON, W. A. N., 1917.—Fluke in sheep. J. Dept.
Agric. Vict., 15: 705-10.

106. THomaAS, J. D., 1882.—On Hydatid disease in Australia. Trans. roy. Soc. S. Aust., 5: 120.

107. THomaAs, J. D., 1885a.—Notes upon the experimental breeding of Taenia echinococcus in
the dog from the echinococci of man. Proc. roy. Soc., 38: 449-57.

108. THomas, J. D., 1885b.—Note upon the frequent occurrence of YTaenia echinococcus in
the domestic dog in certain parts of Australia. Proc. roy. Soc., 38: 457-8.

109. TIDSWELL, F., 1910.—Parasites. Rep. Bur. Microbiol. N.S.W., 1909, 74-99.

110. WeENyon, C. M., 1925.—Protozoology. MBailliére, Tindall and Cox, London, 1926. 2 vol.
1563 pp.

111. WickHAm, N., and CARNB, H. R., 1950.—Toxoplasmosis in domestic animals in Australia.
Aust. vet. J., 26: 1-3.

112. YorKrE, W., and MAPpLESTONE, P. A., 1926.—The nematode parasites of vertebrates.
J. & A. Churchill, London, 1926, 536 pp.

113. Younc, M. R., 1939.—Helminth parasites of Australia. Imp. Bur. Agric. Parasitol.
(Helminthology), England, pp. 145.

PART III. INTRODUCED HERBIVORA AND THE DOMESTIC PIG.

Synopsis.

Part III contains the names of eleven introduced mammals, parasites being recorded
from six of them.

Important protozoa recorded from stock are the organisms causing tick-borne fevers
in cattle, Babesia argentina, B. bigemina and Anaplasma ‘marginale. Non-pathogenic
trypanosomes occur in sheep and cattle, Trichomonas foetus occurs in cattle, and Himeria
faurei in sheep and goats.

Only four species of trematodes have become established, namely, the liver fluke and
three flukes inhabiting the rumen of sheep and cattle.

Three species of adult cestodes occur in the horse, and four in ruminants. Hydatids
have been recorded from sheep, cattle, goats, pigs, horses, and camels. Larval stages of
other dog tapeworms occur in sheep, cattle, goats, and pigs.

Numerous species of nematodes are present in stock, most important being species of
Trichostrongylidae in sheep and cattle, and of Strongylidae in the horse.

One Acanthocephala is known from the pig.

Many Australian workers have studied the parasites of domestic animals in this
country, so that most of the species which were imported with their hosts are now
well known. The invaluable publications of Seddon (1950, 1952) form the basis of
this list, which could not have been compiled without them. W#arlier records were
listed by Sweet (1909a, 1909b), Johnston (1909a, 1909b, 1916) and Young (1939). The
reader is referred to these publications, particularly those of Seddon, for references to
the extensive literature on the parasites of stcck.

One of the reasons for publishing this account is to enable a comparison to be
Made between the parasites of the large herbivorous marsupials (i.e. kangaroos and

144 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

wallabies) on the one hand, and those of the introduced herbivora on the other.
Both groups have numerous parasites, particularly nematodes belonging to the
Strongyloidea. In the marsupials, the family Strongylidae is most abundantly
represented, greatly outnumbering the Trichostrongylidae. This occurs also in the
horse, in which 15 species of Strongylidae belonging to six genera are known in
Australia. About half the species belong to the subfamily Trichoneminae, which is
the most numerous also in the marsupials, but the genera. are distinct. In the
ruminants, on the other hand, the Trichostrongylidae predominate, being represented
by 23 species belonging to five genera in cattle and sheep in Australia.

It is interesting to find that, with a few exceptions, there has been no mixing
of the parasitic groups. The most important exceptions are the transference of
hydatids to marsupials, some of which have unfortunately proved good hosts, and
the occurrence of the liver fluke in some wallabies and kangaroos. Apparently no
parasites of marsupials have been found yet in domestic animals.

An interesting and instructive account: of the subject was given by Dr. F. H. S.
Roberts in his presidential address to Section L of the Australian and New Zealand
Association for the Advancement of Science at the meeting in Sydney in 1952.*

The explanation of the abbreviations used will be found at the beginning of
Part I (p. 102).

HOSTS AND PARASITES
Order ARTIODACTYLA
Family SuImDAE
Genus Sus L., 1758
S. scrora L., 1758, the domestic pig (introduced)

Protozoa
(Sa.) Entamoeba polecki Prowazek, 1912 (77)
(M.) Trichomonas sp. (114)
(Sp.) Coccidiosis (sp. not identified) (114)

Himeria debliecki Douwes, 1921 (77)
Sarcocystis meischeriana Kitihn, 1865 (64), (58)

(Cs) Balantidium sp., probably B. coli (Malmsten, 1857) (74), (114)
Trematoda
(Fasci.) Fasciola hepatica L., 1758 (108), (4117), (91), (113)
Cestoda
(CY., Tae.) Hchinococcus granulosus (Batsch, 1786) as hydatid (41), (14), (128).
(91), (113)
Taenia solium L., 1758, as Cysticercus cellulosae (Gmelin, 1790) (57),
(2), (113)

Taenia hydatigena Pallas, 1766, as Oysticercus tenwicollis (Rudolphi,
1810) (85), (14), (128), (117), (91), (118)
(PS., Dip.) Diphyllobothrium (Spirometra) erinacei (Rudolphi, 1819), as sparganum
(88), (118), (5), (49)

Nematoda
(RH.) Strongyloides ransomi Schwartz and Alicata, 1930 (67), (91), (113)
(TR.) Trichuris trichiura (l., 1771) (85), (58), (123), (67), (117), (91), (113)

(ST., Tri.) Trichostrongylus axei (Cobbold, 1879), rare (102), (118)
T. colubriformis (Giles, 1892), rare (102), (118)
Hyostrongylus rubidus (Hassall and Stiles, 1892) (67), (117), (91), (113)
(ST., Str.) Stephanurus dentatus Diesing, 1839 (80), (24), (25), (3), (128), (59),
(112), (91), (113)
Oesophagostomum dentatum (Rudolphi, 1803) (67), (117), (91), (113)
O. quadrispinulatum Marcone, 1901 (67), (91), (113)
(ST., Anc.) Ancylostoma duodenale (Dubini, 1843) (75), (117), (91), (118)
Necator americanus (Stiles, 1902) (1), (117), (91), (118)

* Roberts, F. H. S., 1953.—Host specificity of livestock parasites in Australia. Rep. Ausf.
Ass. Adv. Sci., 29: 247-57.

BY M. JOSEPHINE MACKERRAS. 145

(ST., Met.) Metastrongylus apri (Gmelin, 1790) (70), (85), (123), (67), (117), (91),

(113)
M. pudendotectus (Wostokow, 1905) (67), (117), (91), (113),
(AS.) Ascaris suum Goeze, 1782, syn. suilla Dujardin, 1845 (85), (14), (123),
(117), (67), (91), (93), (113). (Often referred to as lumbricoides L..,
1758)
(SP.) Ascarops strongylina (Rudolphi, 1819) (85), (123), (117), (91), (113)

Physocephalus sexalatus (Molin, 1860) (67), (117), (91), (113)
Gnathostomum hispidum (Fedschenko, 1872) (52), (117), (91), (113)
(only found once)
Acanthocephala
Macracanthorhynchus hirudinaceus (Pallas, 1781) (85), (57), (123),
(117), (91), (113)

Family CAMELIDAE
Genus CameEtus L., 1758
C. DROMEDARTIUS L., 1758, the Indian camel (introduced)
Protozoa
(M.) Trypanosoma evansi (Steel, 1885) (not indigenous) (15), (114)
Cestoda
(CY., Tae.) Hchinococcus granulosus (Batsch, 1786), as hydatid (15), (123), (113)
Nematoda
(TR.) Trichuris sp. (1138)
(FI.) Onchocerca fasciata (Railliet and Henry, 1919) (18), (100), (113)
Microfilariae in blood (15)
Dipetalonema evansi (Lewis, 1882) (123)

Family BovipDAE
Genus Bos L., 1758
B. taurus L., 1758, the ox (introduced)

Protozoa
(M.) Trypanosoma theileri Laveran, 1902 (124), (125), (114)
Trichomonas foetus Riedmuller, 1929 (34), (114)
(Sp.) Anaplasma centrale Theiler, 1910 (73), (114)

A. marginale Theiler, 1910 (71), (114)
Babesia argentina Ligniéres, 1901 (118), (72), (114) ;
B. bigemina (Smith and Kilborne, 1892) (86), (56), (14), (30), (114)
Theileria mutans (Theiler, 1906) (31), (32), (114) .
Bartonella bovis Donatien and Lestoquard, 1934 (82), (115)
EHperythrozoon wenyoni Adler and Hllenbogen, 1934 (83), (115)
Coccidiosis (sp. not identified) (114)
Himeria sp. (16)
Sarcocystis tenella Railliet, 1886 (64)
Sarcosporidiosis (30)
S. blanchardi Doflein, 1901 (114)
Trematoda* °

(Param.) Paramphistomum ichikawai Fukui, 1922 (36), (37)
Calicophoron calicophorum (Fischoeder, 1901) (36), (38)

; Ceylonocotyle streptocoelium (Fischoeder, 1901) (36), (37)

(Fasci.) Fasciola hepatica L., 1758 (89), (14), (57), (123), (91), (113)

* All flukes found in the rumen were referred to as Amphistoma conicum Rudolphi, 1809,
in the older literature (20), (85), (3), (14). This name proved to be a synonym of cervi
Schrank, 1790, and the parasites were then referred to as Paramphistomum cervi (123),
(120), (121), (60), (91). Later it was realized that at least three species were present,
and they were recorded as P. cervi (Schrank, 1790), P. explanatum (Creplin, 1847), and
P. cotylophorum (Fischoeder, 1901) (101), (113), (35). MWRecently, Durie (1951, 1953, 1956)
worked on the group, and fcund that the species present in Australia are those given here.

146 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

Cestoda
(CY., Ano.)

(CY., Tae.)

Moniezia benedeni (Moniez, 1879), syns. planissima Stiles and Hassall,
1893, and alba Perroncito, 1879 (1238), (91), (113), (104)

M. denticulata (Rudolphi, 1810) (14), (120), (128), (118)

M. expansa (Rudolphi, 1810), syn. trigonophora Stiles and Hassall, 1892
(85), (58), (128), (91), (118)

Helictometra giardi (Moniez, 1879), rare (45), (113)

Echinococcus granulosus (Batsch, 1786) as hydatid (89), (41), (14),
(123), (91), (118)

Taenia saginata Goeze, 1782, as Cysticercus bovis (Cobbold, 1861) (84),
(1138)

Taenia hydatigena Pallas, 1766, as Cysticercus tenuicollis Rudolphi, 1810
(85), (128), (91), (413)

Nematoda

(RH.)
(TR.)

(ST., Tri.)

Strongyloides papillosus (Wedl, 1856) (91), (118)

Trichuris globosa (v. Linstow, 1901) (96), (113)

T. ovis (Abildgaard, 1795) (58), (128), (91), (118).

T. parvispiculum Ortlepp, 1937 (96), (113)

Capillaria sp. (98), (118)

Cooperia curticei (Railliet, 1893) (101), (113)

C. mcemastersi Gordon, 1932 (45), (101), (1138)

C. oncophora (Railliet, 1898) (45), (101), (113)

C. pectinata Ransom, 1907 (91), (113)

C. punctata v. Linstow, 1907 (45), (91), (113)

C. spatulata Baylis, 1938 (101), (113)

Haemonchus contortus (Rudolphi, 1803) (4), (29), (128), (91), (118),
(105)

H. placei (Place, 1893) (105)

Nematodirus filicollis (Rudolphi, 1802) (101), (118)

“N. spathiger (Railliet, 1896) (101), (113)

(ST., Anc.)
(ST., Str.)

(ST., Met.)
(AS.)
(SP. )

(FI.)

Ostertagia circumcincta (Stadelmann, 1894) (29), (91), (1138)

O. occidentalis Ransom, 1907 (101), (113)

O. ostertagi (Stiles, 1892) (44), (91), (118)

Trichostrongylus axet (Cobbold, 1879), syn. extenuatus Railliet, 1898
(128), (10), (91), (4118)

T. colubriformis (Giles, 1892), syn. instabilis Railliet, Lege (91) (118)

T. longispicularis Gordon, 1933 (101), (113)

T. vitrinus Looss, 1900 (101), (113)

Bunostomum phlebotomum (Railliet, 1900) (111), (91), (118)

Bosicola radiatum* (Rudolphi, 1813) (38), (4), (14), (123), (101), (418)

Stephanurus dentatus Diesing, 1839 (91), (1138)

Dictyocaulus viviparus (Bloch, 1782) (14), (1238), (10), (91), (118)

Neoascaris vitulorum (Goeze, 1782) (68), (51)

Gongylonema pulchrum Molin, 1857 (121), (123), (1183)

G. verrucosum (Giles, 1892) (113)

Onchocerca gibsoni (Cleland and Johnston, 1910) (81), (42), (17), (18),
(65), (91), (113)

O. gutterosa Neumann, 1910 (62), (91), (118)

O. lienalis (Stiles, 1392) (62), (113)

Setaria cervi (Rudolphi, 1819) (not indigenous) (113)

Pentastomida (Phylum Arthropoda)

B. INDICUS L.,

Lingulatula serratum (Frohlich, 1789) (larvae) (66), (123)
1758, the zebu (introduced) No records

Genus BuBaALus Frisch, 1775

B. BUBALUS (L., 1758), the water buffalo (introduced) No records

* Confused

with Oesophagostomum columbianum by earlier workers.

BY “M. JOSEPHINE MACKERRAS. 147

Genus Ovis L., 1758
O. aries L., 1758, domestic sheep (introduced)

Protozoa*
(M.) Giardia sp. (127), (114)
Trypanosoma melophagium (Flu, 1908) (124), (125), (114)
(Sp.) Himeria faurei (Moussu and Marotel, 1901) (63)

Coccidiosis (sp. not identified) (114)
Sarcocystis tenella Railliet, 1886 (114)
S. (Balbiania) gigantea Railliet. 1886 (58), (64)
Globidium gilruthi (Chatton, 1910) (48), (114)
Toxoplasma sp. (128), (114)
Trematoda};
(Param. ) Paramphistomum ichikawai Fukui, 1922 (387)
Calicophoron calicophorum (Fischoeeder, 1901) (38)
Ceylonocotyle streptocoeliuwm (Fischoeder, 1901) (37)
(Fasci.) Fasciola hepatica L., 1758 (50), (85), (120), (59), (91), (4118)
(Dicro.) Dicrocoelium dendriticum (Rudolphi, 1819) (not indigenous) (27), (113)
Cestoda
(CY., Ano.) Moniezia benedeni (Moniez, 1879) (58), (123), (113)
M. expansa (Rudolphi, 1810) (26), (69), (85), (9), (58), (128), (91),
(113)
Helictometra giardi (Moniez, 1879) (22), (14), (58), (123), (45), (91),
(113)
(CY., Tae.) Hchinococcus granulosus (Batsch, 1786) as hydatid (55), (76), (85),
(41), (14), (91), (118)
Taenia hydatigena Pallas, 1766, as Cysticercus tenuicollis Rudolphi, 1810
(85), (19), (14), (1238), (91), (113)
T. ovis Cobbold, 1860, as Cysticercus ovis (33), (48), (101), (113)

Nematoda :
(RH.) Strongyloides papillosus (Wedl, 1856) (116), (91), (113)
(TR.) Trichuris globulosa (v. Linstow, 1901) (99), (113)

T. ovis (Abildgaard, 1795), syn. affinis Rudolphi, 1802 (6), (85), (119),
(58), (91), (118)

T. parvispiculum Ortlepp, 1937 (99), (113)

(ST., Tri.) Cooperia curticei (Railliet, 1893) (116), (78), (91), (118)

C: memastersi Gordon, 1932 (46), (113)

C. oncophora (Railliet, 1898) (45), (97), (113)

C. pectinata Ransom, 1907 (39), (97), (1183)

CO. punctata v. Linstow, 1907 (46), (97), (113)

Haemonchus contortus (Rudolphi, 1803) (26), (85), (21), (119), (123),
(91), (118), (105)

H. placei (Place, 1893) (105)

Nematodirus filicollis (Rudolphi, 1802) (123), (91), (118)

N. furcatus May, 1920 (99), (113)

N. spathiger (Railliet, 1896) (97). (118)

Ostertagia circumeincta (Stadelmann, 1894) (128), (91), (113)

O. mentulata (Railliet and Henry, 1909) (99), (113)

O. ostertagi (Stiles, 1892) (11), (116), (45), (91), (118)

O. trifurcata (Ransom, 1907) (45), (97), (113)

Trichostrongylus axei (Cobbold, 1879) (116), (91), (118)

T. colubriformis (Giles, 1892) (116), (91), (113)

T. falculatus Ransom, 1911 (97), (118)

* Commensal flagellates and ciliates are found in the rumen. Moir (1955) stated that
Six species were constantly present, but they were not identified.

7 See footnote under Trematodes of the ox. Rumen flukes in sheep were recorded as
A. conicum (70), (80), (25); as P. cervi (120); and as P. cervi, P. explanatum, and
Cotylophoron cotylophorum (118).

148 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

(ST., Tri.) TV. longispicularis Gordon, 1933 (46), (1138) °
T. probolurus (Railliet, 1896) (46), (97), (118)
T. rugatus Monnig, 1925 (45), (97), (113)
T. vitrinus Looss, 1905 (45), (97), (1138)
(ST., Anc.) Bunostomum trigonocephalum Rudolphi, 1808 (26), (21), (45), (99),
(113)
(ST., Str.) Chabertia ovina (Gmelin, 1790) (85), (123), (113)
Oesophagostomum columbianum Curtice, 1890 (4), (21), (119), (128),
(91), (1138)
O. venulosum (Rudolphi, 1809) (8), (97), (118)
(St., Met.) Dictyocaulus filaria (Rudolphi, 1809) (70), (26), (85), (8), (119), (128),

(113)
: Muellerius capillaris (Mueller, 1889) (109), (1138)
(FI.) Onchocerca gibsoni (Cleland and Johnston, 1910) (rare) (91), (113)

Genus Capra L., 1758
C. uircus L., 1758, the goat (introduced)
Protozoa
(Sp.) Himeria faurei (Moussu and Marotel, 1901) (53)
Trematoda
(F'asci. ) Fasciola hepatica L., 1758 (113)
Cestoda
(CY., Ano.) Moniezia expansa (Rudolphi, 1810) (91), (118)
(CY., Tae.) Echinococcus granulosus (Batsch, 1786) as hydatid (91), (118)
Taenia hydatigena Pallas, 1766, as Cysticercus tenuicollis Rudolphi, 1810
(57), (128), (113)

Nematoda
(RH.) Strongyloides papillosus (Wedl. 1856) (52), (53), (113)
(TR.) Trichuris ovis (Abildgaard, 1795) (53), (118)

(ST., Tri.) Cooperia curticei Railliet, 1893 (40), (113)
Haemonchus contortus (Rudolphi, 1803) (18), (538), (91), (118)
Nematodirus sp. (113)
Ostertagia circumcincta (Stadelmann, 1894) (40), (118)
Trichostrongylus axei (Cobbold, 1879) (53), (91), (118)
T. colubriformis (Giles, 1892) (52), (538), (91), (118)
T. probolurus (Railliet, 1896) (113)
T. rugatus Monnig, 1925 (113)
T. vitrinus Looss, 1905 (113)

(ST., Str.) Oesophagcstomum columbianum Curtice, 1890 (53), (91), (113)
O. venulosum ({Rudolphi, i809) (1138)

(ST., Met.) Muellerius capillaris (Muller, 1889) (61), (118)

Family CERVIDAE
Genus Crrvus L., 1758
C. ELEPHAS L., 1758, the red deer (introduced) No records

Order PERISSODACTYLA
Family EQUIDAE
Genus Equus L., 1758
E. CABALLUS L., 1758, the horse (introduced)
Trematoda
(Param.) Gastrodiscus aegyptiacus (Cobbold, 1876) (not indigenous) (123), (1138)
Cestoda
(CY., Ano.) Anoplocephala mugna (Abildgaard, 1789), syn. plicata Rudolphi, 1810
(3), (28), (58), (1238), (106), (91), (118)
A. perfoliata (Goeze, 1782) (85), (9), (28), (121), (58), (128), (106),
(91), (113)
Paranoplocephala mammillana (Mehlis, 1881) (28), (28), (58), (128),
(95), (118)

BY M. JOSEPHINE MACKERRAS. 149

(CY.. Tae.) Hchinococcus granulosus (Batsch, 1786) as hydatid (58), (113)

Nematoda

(ST., Tri.) Trichostrongylus axei (Cobbold, 1879) (47), (95), (118)
(ST., Str.) Oesophagodontus robustus (Giles, 1911) (106), (1138)

Strongylus edentatus (Looss, 1900) (121), (58), (128), (7), (91), (113)

S. equinus Mueller, 1780, syn. armatus Rudolphi, 1802 (85), (3), (28),
(123), (7), (91), (413)

S. vulgaris (Looss, 1900) (121), (123), (7), (91), (118)

Triodontophorus brevicauda Boulenger, 1916 (8), (1138)

T. minor (l.ooss, 1900) (8), (118)

T. serratus (Looss, 1900) (121). (106), (118)

T. tenwicollis Boulenger, 1916 (106), (113)

Triodontophorus sp. (123)

Gyalocephalus sp. (8), (1138)

Poteriostomum imparidentatum Quiel, 1919 (106), (118)

Trichonema aegyptiacum Railliet, 1923 (1138)

T. calicatum (Looss, 1900) (121), (1238), (1138)

T. longibursatum (Yorke and Macfie, 1918) (7), (118)

T. poculatum (Looss, 1900) (121), (128), (8), (118)

T. tetracanthum (Mehlis, 1831) (85), (28), (58), (128), (7), (91)

(ST., Met.) Dictyocaulus arnfieldi* (Cobbold, 1884) (85), (3), (123), (91), (118)
(OX.) Oxyuris equi (Schrank, 1788) (85), (38), (57), (123), (7), (91), (118)

Probstmayria vivipara (Probstmayr, 1865) (118)

(AS.) Parascaris equorum (Goeze, 1782), syn. megalocephala Cloquet, 1824

(85), (8), (28), (121), (58), (128), (91), (113)

(SP. ) Draschia megastoma (Rudolphi, 1819) (85), (3), (58), (128), (54), (12),

(91), (118)

Habronema microstoma (Schneider, 1866) (85), (58), (123), (54), (12),
(91), (113)

H. muscae (Carter, 1861) (54), (61), (12), (91), (4118)

(FI.) Onchocerca reticulata Diesing, 1841, syn. cervicalis Railliet and Henry,
1910 (8), (100), (113), (90)
BE. asinus L., 1758, the ass (introduced) No records
BH. CABALLUS x E. ASINUS, the mule (introduced) No records
References.
1. ALBISTON, H. E., 1922.—Nematodes in pigs. Med. J. Auwst., 1922, 2: 173.
2. ATKINSON, H., 1915.—Cysticercus cellulosae in the pig. Med. J. Aust., 1915, 1: 144-5.
3. Bancrort, T. L., 1893.—Notes on some diseases of stock in Queensland. Vet. J., 37:
327-30.
4. Barnes, A. W., 1898.—A new cattle disease. Vet. J., 47: 485-6.
5. Brarup, A. J., 1953.—Life history of a Spirometrid tapeworm, causing sparganosis in
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6. BENNETT, R., 1875.—Internal parasites of sheep. - Veterinarian, 48: 123-6.
7. BENNETTS, H. W., 1927.—The helminth parasites of Western Australian stock. Recorded
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8. BENNETTS, H. W., 1928.—Further contributions to the helminths of Western Australian
stock. J. roy. Soc. W. Aust., 14: 57-9.
9. Brown, A. A., 1902.—Animal parasites. J. Dep. Agric. Vict., 1: 698-700.
10. Bruce, G. S., 1912.—Disease in calves. Agric. Gaz. Tasm., 20: 270-2. (Original not
seen. )
11. Bruce, G. S., 1913.—Parasitic gastritis in sheep. Agric. Gaz. Tasm., 21:176. (Original
not seen.)
12. Buu, L. B., 1919.—A contribution to the study of habronemiasis: a clinical, pathological,
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13. CHARLTON, N. B., 1924.—Bowel parasites of Australia and her dependencies. Their
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14. CLELAND, J. B., 1907.—The diseases of animals and meat inspection in Western Australia.

E

J. Dep. Agric. W. Aust., 15: 84-94.

* Horse lung worms were recorded as Strongylus micrurus Mehlis by the earlier workers.

150

36.

37.

38.

39.

40.

41.
42.

43.

44,

45.

46.

ANT

48.

49.

AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

CLELAND, J. B., 1909.—Trypanosomiasis and other diseases in camels, with experiments
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CLELAND, J. B., 1913.—Note on the occurrence of Coccidiosis in house sparrows and in
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CLELAND, J. B., and JOHNSTON, T. H., 1910a.—Worm-nests in cattle due to Filaria
gibsoni (sp. nov.) —preliminary report. Agric. Gaz. N.S.W., 21: 173-4.

CLELAND, J. B., and JOHNSTON, T. H., 1910b.—Worm-nests in Australian cattle due to
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N.S.W., 44: 156-71.

CLUTTERBUCK, F., 1907.—A parasite of sheep. The Cysticercus tenuicollis: its progress
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Cops, N. A., 1898.—Extract from M.S. report on the parasites of stock. Agric. Gaz.
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Cops, N. A., 1902.—Probable occurrence of the tapeworm (Taenia ovilla) in Australian
sheep. Agric. Gaz. N.S.W., 13: 796.

Cops, N. A., 1905.—The tapeworms of Australia. Agric. Gaz. N.S.W., 16: 617-31.

CoBBOLD, T. S., 187la.—The new hog-parasite. Brit. med. J., 1871, 2: 394.

CoBBOLD, T. S., 1871b.—Report on Dr. Morris’ paper. Monthly Micr. J., 31: 245.

CoBBOLD, T. S., 1879.—Parasites, a treatise on the entozoa of man and animals, including
some account of the ectozoa. J. & A. Churchill, London, 1879, 508 pp.

Davin, T. W. E., 1900.—(Exhibit of New South Wales internal parasites.) J. roy. Soc.
N.S.W., 34: XX.

DESMOND, J., 1904.—Horse complaint on Yorke’s Peninsula. J. Agric. S. Aust., 7: 569-70.

Dopp, S., 1908.—Notes on the presence of two stomach worms in calves hitherto
unrecorded in Australia. Qd. agric. J., 21: 197-8.

Dopp, S., 1909.—Report of the principal veterinary surgeon and bacteriologist. Ann. Rep.
Dep. Agric. Stock Qd., 1908-9: 81-105. :

Dopp, S., 1910.—Diseases in stock. Ann. Rep. Dep. Agric. Stock Qd., 1909-10: 19-22.

Dopp, S., 1924.—The piroplasmoses of the Pacific Regions. Proc. Pan-Pacific Sci. Congr.,
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DRABBLE, J., 1934.—Measles (Cysticercus ovis) in sheep in New South Wales. Aust.
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DUMARESQ, J. A., 1948.—A note recording the presence of Trichomonas foetus infection
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Duribf, P. H., 1949.—A preliminary note on the life cycle of Paramphisiomum
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DourRIz£, P. H., 195i1.—The Paramphistomes (Trematoda) of Australian ruminants. Part I.
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Duri£, P. H., 1953.—The Paramphistomes (Trematoda) of Australian ruminants. II. The
life history of Ceylonocotyle streptocoelium (Fischoeder) Nasmark and Paramphi-
stomum ichikawai Fukui. Aust. J. Zool., 1: 193-222.

Durif, P. H., 1956.—The Paramphistomes (Trematoda) of Australian ruminants. III.
The life history of Calicophoron calicophorum (Fischoeder) Nasmark. Aust. Jd.
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HpGar, G., 1933.—Some observations on trichostrongylosis of young sheep. Auwst. vet. J.,
9: 149-54.

EpGar, G., 1936.—Fatal effects of heavy infestation with Cooperia curticei (Railliet,
1893) in goats. Aust. vet. J., 12: 58-61.

FEUERHEERDT, H., 1898.—Hydatids in stock. J. Agric. S. Aust., 1: 833-4.

GisBson, J., 1893.—Notes on certain ‘‘worm nests” or “worm knots” in the cellular tissue
of the brisket in cattle. Intercolon. med. Congr. Aust., 1892: 576-80.

GILRUTH, J. A., 1910.—Notes on a Protozoon parasite found in the mucous membrane
of the abomasum of a sheep. Proc. roy. Soc. Vict., n.s., 23: 19-20.

GILRUTH, J. A., and Sweet, G., 1910.—Gastritis due to trichostrongyle invasion. Cases
in adulz: cattle. Vet. J., 66: 418-21.

GORDON, H. McL., 1932.—Some helminth parasites reported from Australia for the first
time, with a description of Cooperia memastersi sp. nov. Aust. vet. J., 8: 2-12.

GoRDON, H. McL., 1933a.—Scome ovine trichostrongylids reported from Australia for
the first time, with a description of T'richostrongylus longispicularis sp. nov. from a
sheep. Aust. vet. J., 9: 34-7.

GorRDON, H. McL., 1933b.—A note on a case of Trichostrongylus infestation in a horse.
Aust. vet. J., 9: 68.

GorDON, H. McL., 1939.—A note on the experimental transmission of Cysticercus ovis.
Aust. vet. J., 15: 125-6.

GorRDON, H. McL., ForsytH, B. A., and ROBINSON, M., 1954.—Sparganosis in feral pigs
in New South Wales. Aust. vet. J., 30: 135-8.

50.

ol.

BY M. JOSEPHINE MACKERRAS. 152

Harrop, BH. A., 1870.—Remarks on the fluke (Fasciola hepatica). Pap. roy. Soc. Tasm.,
1869: 12-16.

Hart, B., 1951.—Recovery of the nematode Ascaris vitulorwm Goeze, 1782, from the
faeces of a calf. Awst. vet. J., 27: 208.

Heypon, G. M., 1929.—Creeping eruption or larva migrans in North Queensland and @
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Heypon, G. M., and GREEN, A. K., 1931—Some werm infestations of man in Australia.

Med. J. Aust., 1931, 1: 619-28.

Hiuu, G. F., 1918.—Relationship of insects tc parasitic diseases in stock. Part 1. The
life history of Habronema muscae, H. microstoma and H. megastoma. Proc. roy. Soe.
Vict., n.s., 31: 11-76.

Hupson, R. F., 1861.—On hydatids. Aust. med. J., 6: 75-88.

Hunt, J. S., and CoLuIns, W., 1896.—Report on tick - fever. Report of the Special
Commission of the Queensland Government to the U.S.A. Brisbane, 1896.
JOHNSTON, T. H., 1909a.—Notes on some Australian parasites. Agric. Gaz. N.S.W.,
20:.581-4. : - ;
JOHNSTON, T. H., 1909b.—Notes and exhibits of entozoa. Proc. LINN. Soc. N.S.W., 34:
117-8, 217-9, 417-8. ;
JOHNSTON, T. H., 1912.—Notes on some Entozoa. Proc. roy. Soc. Qd., 24: 63-91.

JOHNSTON, T. H., 1916.—A census of endoparasites recorded as occurring in Queensland
arranged under their hosts. Proc. roy. Soc. Qd., 28: 31-79.

JOHNSTON, T. H., 1918.—Notes on miscellaneous endoparasites. Proc. roy. Soc. Qa.,
30: 209-18.

JOHNSTON, T. H., 1921.—Onchocerciasis of Queensland cattle. Trans. roy. Soc. S. Aust.,
45: 231-47.

JOHNSTON, T. H., 1926.—Note on the occurrence of coccidiosis in South Australiam
sheep. Rep. Aust. Ass. Advance. Sci., 1924: 712-4.

JOHNSTON, T. H., and CLELAND, J. B., 1909.—Notes on some parasitic Protozoa. Proc.
LINN. Soc. N.S.W., 34: 501-13.

JOHNSTON, T. H., and CLELAND, J. B., 1910a—On the anatomy and possible mode of
transmission of Filaria (Cnchocerca) gibsoni. J. roy. Soc. N.S.W., 44: 171-89.

JOHNSTON, T. H., and CLELAND, J. B., 1910b.—Notes on the occurrence of Pentastomes
in Australian cattle. J. roy. Soc. N.S.W., 44: 315-8.

Kauzau, G. P., 1930—A survey of the helminth parasites of swine in New South
Wales. Aust. vet. J., 6: 51-6.

KeirH, R. K., 1951.—The occurrence of Ascaris vitulorum Goeze, 1782, in calves im
Australia. Aust. vet. J., 27: 129.

KENDALL, W. J., 1890.—Parasitic diseases of animals. The tape worms of sheep. Aust.
Vet. Live Stock J., 1: 193-8. 5

KRreEFrT, G., 1871.—On Australian Entozoa with descriptions of new species. Trans. ent.
Soc. N.S.W., 2: 206-32.

Lecce, J., 1933.—The cccurrence of Anaplasma marginale Theiler, 1910, in Northern
Queensland. Coun. Sci. industr. Res., Pamphlet No. 38, 31 pp.

Leee, J., 1935.—The occurrence of bovine babesiellosis in Northern Australia. Couwn-
Sci. industr. Res., Pamphlet No. 56, 48 pp.

Leee, J., 1936.—Anaplasmosis. Cross-immunity tests between Anaplasma centrale (South
Africa) and Anaplasma marginale (Australia). Aust. vet. J., 12: 230-3.

Lece, J., 1947.—Report of the Director, Division of Animal Industry, Ann. Rep. Dept.
Agric. Stock Qd., 1946-47: 42-53. :

Leee, J.. and RHEUBEN, J. A., 1921.—Note on the finding of Anchylostoma duodenale in
the intestine of the pig. Med. J. Aust., 1921, 2: 398.

MACKELLAR, C. K., 1883.—HExhibit of portions of the liver and lungs of a sheep with
large hydatid cysts. Proc. LINN. Soc. N.S.W., 8: 280-1.

MACKERRAS, M. J.—Unpublished observations.

McGratH, T. T., 1931.—Occurrence of Cooperia curticei in sheep in New South Wales.
Vet. Res. Rep. Dep. Agric. N.S.W., No. 6: 111-3.

Moir, R. J., 1955.—The seasonal variation in the ruminal microorganisms of grazing
sheep. Aust. J. agric. Res., 2: 322-30. ‘

Morris, W., 1871.—(Intestinal worms from Australia.) Monthly Micr. J., 6: 234-4.

Morris, W., 1881.—(Notes on an encysted Filaria found in the flesh of a bullock.)
J. roy. Soc. N.S.W., 14: 337.

MULHEARN, C. R., 1946.—A note on two blood parasites of cattle (Spirochaeta theileré
and Bartonella bovis) recorded for the first time in Australia. Aust. vet. J., 22: 118-9.

MULHEARN, C. R., 1949.—Personal communication.

PENFOLD, H. B., 1937.—The life history of Cysticercus bovis in the tissue of the ox.
Med. J. Aust., 1937, 1: 579-83. .

PERRIE, W., 1892.—Scientific Collections— Internal and External parasites of stock
(a prize-winning collection of Entozoa and Epizoa). Agric. Gaz. N.S.W., 3: 821-3.

100.
101.

102.

103.

104.

105.

106.

AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

PounpD, C. J., 1895.—Redwater disease in cattle in the Gulf district. Gov. Printer,
Brisbane. ,

Pounp, C. J., 1897.—Tick fever. Qd. agric. J., 1: 258.

PuLuaR, H. M., and MCLENNAN, G. C., 1949.—Sparganosis in a Victorian pig. Aust.
vet. J., 25: 302-4.

RALPH, T. S., 1865.—On the parasitic origin of pleuropneumonia in cattle; being the
report made to the Pleuropneumonia Commission. Awst. med. J., 10: 1-10.

RigkK, R. F., 1954-——A note on the occurrence of Onchocerca reticulata Diesing, 1841,
in the horse in Queensland. Aust. vet. J., 30: 178-81.

Rosperts, F. H. S., 1934a—Worm parasites of domestic animals in Queensland. Qd.
Agric. J., n.s., 41: 245-52.

Roperts, F. H. S., 1934b.—Parasites of the pig. Qd. Agric. J., n.s., 42: 208-21.

Roperts, F. H. S., 1934e—The large roundworm of pigs, Ascaris lumbricoides L., 1758.
Its life history in Queensland, economic importance and control. Bull. Dep. Agric.
Qd., No. 1, pp. 81.

Rosperts, F. H. S., 1934d.—The parasites of sheep. Qd. Agric. J., n.s., 42: 3387-59.

Roperts, F. H. S., 19384e—Parasites of the horse Qd. Agric. J., n.s., 42: 473-89.

Roperts, F. H. S., 1934f.—Parasites of cattle. Qd. Agric. J., n.s., 42: 674-89.

Roperts, F. H. S., 1935a—Helminth parasites of domesticated animals in Queensland.
Further records of occurrence. Qd. Agric. J., n-S., 44: 299-300.

RopBerts, F. H. S., 1935b.—The occurrence of Capillaria sp. in a calf. Aust. vet. Jd.,
11; 229.

Roperts, F. H. S., 1936.-—-The distribution of the gastro-intestinal parasites of sheep in
Queensland. Qd. Agric. J., n.s., 46: 30-7.

Roserts, F. H. S., 1938.—Onchocerciasis — Annotation. Aust. vet. J., 14: 32-5.

ROBERTS, F. H. S., 1989.—The gastro-intestinal helminths of cattle in Queensland: their
distribution and pathogenic importance. Proc. roy. Soc. Qd., 50: 46-54.

Roperts, F. H. S., 1940a—Notes on some helminths infesting domestic animals in
Queensland. Aust. vet. J., 16: 30-3.

Roperts, F. H. S., 19406b.—The incidence, prevalence and distribution of helminths
infesting the lungs and alimentary tract of the pig in Queensland. Aust. vet. J.,
16: 259-66. ;

Roperts, F. H. S., 1953.—Zygoribatula longiporosa Hammer (Oribatei: Acarina), an
intermediate host of Moniezia benedeni (Moniez) Anoplocephalidae: Cestoda in
Australia. Awst. J. Zool., 1: 239-41.

Roperts, FE. H. S., TURNER, H. N., and McKervetTtr, M., 1954.—On the specific distinctive-
ness of the ovine and bovine “strains” of Haemonchus contortus (Rudolphi) Cobb
(Nematoda: Trichostrongylidae). Aust. J. Zool., 2: 275-95.

Ross, I. C., 1926a.—Parasites of dogs and horses in New South Wales, from the
neighbourhood of Sydney. Rep. Aust. Ass. Advance. Sci., 1924: 693-7.

Ross, I. C., 1926b.—A survey of the incidence of Hchinococcus granulosus (Batsch) or
hydatid disease in New South Wales. Med. J. Auwst., 1926, 1: $6-103.

Ross, I. C., 1928.—Liver fluke disease in Australia: its treatment and prevention. Coun.
Sci. industr. Res., Pamphlet No. 5, 23 pp.

Ross, I. C., 19381. ame small lung worm of sheep in New South Wales. Aust. vet. J.,
7: 148-9.

Ross, I. C., and Gordon, H. McL., 1936.—The internal parasites of sheep: their treat-
ment and control. Angus and Robertson, Sydney, pp. xx + 238.

Ross, I. C., and KAuzau, G., 1931.—Monodontus phlebotomus and Oesophagostomum
venulosum. Two important parasites of cattle and sheep recorded from Australia
for the first time. Aust. vet. J., 7: 25-7.

Ross, I. C., and Kauzau, G., 1932.—The life cycle of Stephanurus dentatus Diesing,
1839: the kidney worm of pigs. Bull. Coun. sci. industr. Res. Aust., No. 58, 80 pp.

SEDDON, H. R., 1950.—Diseases of domestic animals in Australia. Part 1. Helminth
infestations. Serv. Publ. Dep. Hlth. Aust. vet. Hyg., No. 5, 223 pp.

SEDDON, H. R., 1952.—Diseases of domestic animals in Australia. Part 4. Protozoan and
viral diseases. Serv. Publ. Dep. Hlth. Aust. vet. Hyg., No. 8, 214 pp.

SEDDON, H. R., 1953.—Diseases of domestic animals in Australia. Part 5, vol. II.
Bacterial diseases. Serv. Publ. Dep. HIith. Aust. vet. Hyg., No. 10, 464 pp.

SEDDON, H. R., and McGratH, T. T., 1931.—Observations upon the conditions requisite
for the transmission of gastro-intestinal nematodes of sheep. Vet. Res. Rep. Dep.
Agric. N.S.W., No. 6: 40-57.

SHELTON, E. J., 1930.—Diseases of the pig. Qd. Agric. J., 34: 297-318.

Souns, J. C. F., 1918.—WMicrobabesia divergens in Nederlandsch-Indié. Nederl.-Ind. Blad.
u. Diergeneesk., 30: 385-459. (Original not seen.)

Stpwart, J. D., and Henry, M., 1908.—Experimental test of treatments for worms in
sheep at Glen Innes Experimental Farm. Agric. Gaz. N.S.W., 19: 981-5.

BY M. JOSEPHINE MACKERRAS. 153

120. Sweet, G., 1909a.—The Endoparasites of Australian Stock and Native Fauna. Part I.
Introduction, and census of forms recorded up to date. Proc. roy. Soc. Vict., n.s.,
21: 454-502.

121. Swher, G., 1909b.—The Endoparasites of Australian Stock and Native Fauna. Part II.
New and unrecorded species. Proc. roy. Soc. Vict., n.s., 21: 503-27.

122. Sweet, G., SEDDON, H. R., and RopBertson, W. A. N., 1917.—Fluke in Sheep. J. Dep.
Agric. Vict., 15: 705-10.

123. TIDSWELL, F., 1910.—Parasites. Rep. Bur. Microbiol. N.S.W., 1909: 74-99.

124. TURNER, A. W., and MURNANE, D., 1930a—Trypanosomes in the blood of Victorian
animals. 1. A preliminary note on the occurrence of Trypanosoma theileri in the
blood of cattle. 2. On the presence of Trypanosoma melophagium in the blood of
Victorian sheep, and its transmission by the sheep ‘‘tick’, Melophagus ovinus.
J. Coun. sci. industr. Res. Avust., 3: 120-2.

125. TurNpR, A. W., and MURNANE, D., 1939b.—On the presence of the non-pathogenic
Trypanosoma melophagium in the blood of Victorian sheep, and iis transmission by
Melophagus ovinus. Aust. J. exp. Biol. med. Sci., 7: 5-8.

126. TuRNER, A. W., and MURNANE, D., 1930c—A preliminary note on the occurrence of
Trypanosoma theileri in the blood of cattle in Victoria. Awst. J. exp. Biol. med. Sci..
02 Pahg

127. TURNER, A. W., and MURNANBS, D., 1932.—Giardia in sheep in Victoria, Australia. Aust.
J. exp. Biol. ned. Sci., 10: 53-6.

128. WickHAm, N., and CARN#, H. R., 1950.—Toxoplasmosis in domestic animals in Australia.
Aust. vet. J., 26: 1-3.

129. Youne, M. R., 1939—Helminth parasites of Australia. Imp. Bur. Agric. Parasitol.
(Helminthology) England, pp. 145,

PART IV. MAN.

Synopsis.

Highteen protozoa, five trematodes, nine cestodes, and thirteen nematodes have been
recorded from man in Australia. Several of these were certainly acquired outside this
continent (one protozoon, four trematodes, two cestodes, and three nematodes). Certain
others, while acquired locally, were accidental infections with parasites of other animals
(one trematode, five cestodes, and three nematodes). Others (including the malarial and
filarial parasites) are very rare.

Echinococcus granulosus is the most dangerous animal parasite of the white population
in the southern part of ‘Australia, and the hookworms of the native population in the north,

Schistosome dermatitis, creeping eruption, and visceral larva migrans are caused by
parasites of other animals, which are not adapted to complete their development in the
human host.

Australia is fortunately free from many harmful parasitic diseases, and others,
which were formerly important, are now becoming uncommon. Malaria belongs to
the latter group. It was once endemic in parts of Australia north of 19°S, and sharp
epidemics occurred from time to time in towns and mining camps. The available
evidence strongly suggests that the disease was imported, probably repeatedly, by
migration from the north; by Asiatics, by islanders from New Guinea and the
Solomons, and by Europeans entering from the north, particularly from New Guinea.
There is no evidence that the disease was endemic in the aboriginal people.

White (1867) and Dyson (1889) described severe fevers in North Queensland,
some of which were probably malarial. O’Brien (1908, 1909) recorded three types in
Cairns, benign, malignant and quartan, his cases being diagnosed by microscopic
examination. Epidemics occurred in Cairns from time to time. Breinl and Taylor
(1918) investigated one, in which Plasmodium vivax and P. falciparum were both
present. In 1942 there was an epidemic due to P. vivax alone. This epidemic was
investigated by Dr. G. A. M. Heydon, who showed that the vector was Anopheles
farauti Lay. This is the only occasion when the vector has been definitely proved.
Small outbreaks occurred during the war in army camps at Selheim and Canungra,

154 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

areas outside the distribution of An. farauti, and the vector was thought to be
An. annulipes Walk. Malaria has been acquired as far south as Victoria, but only
rarely.

Cilento (1924) and Ford (1950) have given good accounts of the history of
malaria in Australia. It seems to have disappeared as an endemic disease from
Queensland, except for small foci in some of the Torres Strait islands (Mackerras and
Sandars, 1954). There may still be endemic malaria in parts of the Northern Territory
and in the Kimberley District of Western Australia, but Black (1950) could find no
evidence of it in 1382 natives and 121 white people examined in these areas.

Filariasis is another diminishing disease. It was probably introduced into
Queensland in the middle of the last century, and probably by Chinese immigrants.
By the seventies it was a frequent cause of morbidity in Queensland, and it was
in Brisbane that the adult worms were discovered by J. Bancroft in 1876. The first
assessment of the frequency of the infection was made in 1910 by McLean, who found
that nearly 11% of 1200 admissions to the Brisbane General Hospital were infected.
When the staff of the Australian Hookworm Campaign carried out a survey for
malaria and filariasis in 1922-24, the infection rate in Brisbane was 5%, one of the
highest rates in the State. However, in 1938 Derrick examined 228 persons in the
Brisbane General Hospital without finding a single carrier. The disease has receded
everywhere in Queensland, only six cases having been notified in the last decade.
This recession was well advanced before any effective treatment became available,
and in spite of the fact that a good vector, Culex fatigans Wied., is still abundant
and widespread.

Ancylostomiasis has decreased greatly since the time of the Australian Hookworm
Campaign, 1919-24, when 8% of 202,582 persons examined in Australia were infected.
The great majority of these examinations were made in coastal Queensland, where
the overall rate was 9%. In some districts about 25% were infected at this time. In
a few high-rainfall areas of North Queensland there is still an infection rate of
about 1% in rural school children; elsewhere the disease has disappeared from the
white population. It is still widely disseminated among the aborigines, particularly in
Queensland and in the Northern Territory, although there has been a remarkable
improvement in the last few years in some settlements, for example at Mona Mona
Mission, Yarrabah and Palm Island.

Human schistosomes have been repeatedly introduced, particularly S. haematobiun,
which was brought back by many soldiers returning from the Boer War. Treatment
was ineffective, and they remained carriers for years. Nevertheless only two indigenous
cases were reported during ensuing years. They occurred in Western Australia in
1911 and 1912, and may have been infected from the same source (Nelson, 1912).
During the First World War Australian soldiers were infected in Egypt with S. mansoni
and S. haematobium, and during the last war some air force personnel, exposed to
risk in the Philippines, acquired infection with S. japonicum. More effective treatment,
however, was available. One case (S. haematobium) was reported in New South
Wales in 1924, but the source of infection could not be traced. There have been
no notifications since the last war, and it is safe to conclude that human schistosomes
have not become established anywhere in Australia.

Hydatid disease is undoubtedly the most serious helminthic disease present in
Australia. It was one of the first to attract attention, and the literature on the
subject is enormous. Graham (1937) observed that its incidence in Melbourne chiidren
had declined during the previous three decades. However, there does not seem to be
any cause for complacency, as the number of cases notified each year in Victoria, where
notification has been practised consistently, has not fallen since 1937. The infection
rate in rural dogs in New South Wales has not decreased in the last thirty years
(Gemmell, 1957), so that the risk of infection is evidently still considerable.

We have unfortunately no means of finding out what parasites the aborigines
harboured before they came in contact with white people. No parasites have been
found in them which were not already well known in other communities, and at the
present time they have the same array of species as the white people.

BY M. JOSEPHINE MACKERRAS. 155

The first observation of a parasite of man in Australia appears to have been
made by Surgeon P. Cunningham in 1829, in a book entitled “Two Years in New South
Wales”. He noted the presence of ‘teres’ (or roundworms) in children at Port
Jackson. The first collection of records and references to human parasites was made
by Georgina Sweet in 1908 (published in 1909). In 1909, T. H. Johnston began to
record carefully all the parasites recognized at the Bureau of Microbiology, Sydney
(Johnston, 1909a, 1909b, 1909c). In 1912, Johnston and Cleland gave an account of
the helminths of man in Australia, bringing their records up to date in 1937. Johnston
(1916) recorded all parasites found in Queensland. In 1939 Young listed the helminths
and gave many references to the subject.

The explanation of the abbreviations used will be found at the beginning of
Part I (p. 102).

Order PRIMATES
Family HoOMINIDAE
Genus Homo L., 1758
H. SAPIENS L., 1758—man (introduced)
Protozoa

(M. ) Trichomonas hominis (Davaine, 1860) (13)
T. vaginalis Donné, 1837 (67)
Giardia lamblia Stiles, 1915 (75), (113), (13)
Chilomastix mesnili (Wenyon, 1910) (75), (18)
Leishmania donovani (Laveran and Mesnil, 1903) (not indigenous)

(454), (424), (89)

(Sa.) Hntamoeba coli (Grassi, 1879) (59), (113), (18)

; E#. histolytica Schaudinn, 1903 (38), (52), (1084), (57), (118), (13)
Amoebiasis (clinical) (79)
H. gingivalis (Gros, 1849) (57)
Hndolimax nana (Wenyon and O’Connor, 1917) (13)
Dientamoeba fragilis Jepps and Dobell, 1918 (13), (20)
Iodamoeba butschlii (Prowazek, 1913) (75), (113), (32), (13)

(Sp.) Plasmodium vivax (Grassi and Feletti, 1890) (80), (81), (57), (60)
P. falciparum Welch, 1897 (81), (59), (60)
P. malariae (Laveran, 1881) (81), (59), (60)
Malaria (clinical) (111), (385)
Pneumocystis carinii Delanoé, 1912 (88)
Sarcocystis sp. (Seen in sections of heart at Anatomy Department,

University of Queensland. Subject was a healthy child killed in an

accident )
Toxoplasma sp. (90), (36), (82), (102)
(C.) Balantidium coli (Malmsten, 1857) (109)

Trematoda
(Fasci.) Fasciola hepatica* L., 1758 (1), (54)
(Opist.) Clonorchis sinensis (Cobbold, 1875) (not indigenous) (112), (53), (57),
(97), (98)
(Schis.) Schistosoma haematobium (Bilharz, 1852) (not indigenous except (77)
ena! (G0) )) (S3O)5 (Gs GO). ()5 Cas (GAs (SO), GN)
S. mansoni Sambon, 1907 (not indigenous) (61)
S. japonicum Katsurada, 1904 (not indigenous) (30)
Cestoda
(CY., Tae.) Taenia solium L., 1758 (not indigenous) (57), (104)
T. saginata Goeze, 1782 (56), (57), (78), (104), (85)
Echinococcus granulosus* (Batsch, 1786), as hydatid (34), (51), (107), ‘
(57), (33), (42), (40)
(CY., Hym.) Hymenolepis diminuta* (Rudolphi, 1819) (4), (21), (104), (14)
H. nana (v. Siebold, 1852) (109), (75), (66), (104)

* Species marked with an asterisk are parasites of other animals, and only accidentally
infected man.

156 AUSTRALIAN MAMMALS AND THEIR RECORDED INTERNAL PARASITES,

(CY., Dil.) Dipylidium caninum* (L., 1758) (14)
(CY., Dav.) Raillietina (Raillietina) celebensis* (Janicki, 1902) (5)
(PS., Dib.) Diphyllobothrium latum (L., 1758), syn. Dibothriocephalus parvus
Stephens, 1906 (not indigenous) (37) (57), (104), (94)
Sparganumy* (99), (69), (25), (95)

Nematoda
(RH.) Strongyloides stercoralis (Bavay, 1876) (2), (108), (57), (18), (78),
(66), (62), (104)
(TR.) Trichuris trichiura (., 1771) (48), (57), (109), (66)

Trichinella spiralis (Owen, 1835) (not indigenous, except (55)) (55),
(57), (83), (9)
(ST., Tri.) Trichostrongylus colubriformis* (Giles, 1892) (47), (92), (46)
T. axei* (Cobbold, 1879) (47)
Haemonchus contortus* (Rudolphi, 1803) (104), (47)
(ST., Anc.) Ancylostoma duodenale (Dubini, 1843) (49), (81), (57), (18), (78),
(66), (104), (8)
Necator americanus (Stiles, 1902) (70), (57), (78), (66), (104)

(OX.) Enterobius vermicularis (L., 1758) (26), (98), (57), (78), (66)

(AS.) Ascaris lumbricoides (L., 1758) (91), (68), (100), (56), (57), (109),
(104), (3)

(FI.) Wuchereria bancrofti (Cobbold, 1877) (27), (28), (29), (6), (7), (57),

(74), (17), (23), (105), (110), (48), (45)
Loa loa (Cobbold, 1864) (not indigenous) (106), (22)
Dracunculus medinensis (L., 1758) (not indigenous) (72), (84)

LESIONS DUE TO OTHER HELMINTHS
Larval stages of some helminths, which do not normally develop in man, may
gain entrance through the skin or alimentary canal and set up lesions of various.
kinds before they perish in the foreign host.

1. Dermatitis.

Two distinct types of lesions may be caused by the invasion by larval helminths
of the skin of a previously sensitized person.

(a) “Bathers’ itch” or “Schistosome dermatitis” is caused by forked-tail cercariae,
larval stages of various schistosomes, and is characterized by an itchy, papular rash.

In the swamps of the lower Murray River the lesions are probably due to Cercaria
parocellata Johnston and Simpson, 1939, emitted by the fresh-water snail Limnaea.
lessoni (63). C. parocellata is thought to be a larval Trichobilharzia, a group of blood
flukes, which live in the veins of the nasal mucosa and intestine of water birds.
Johnston (1941) thought that the black swan, Chenopis atrata (Latham), which is
known to harbour a schistosome, might be the definitive host, and recently Bearup.
(1957) has found schistosome eggs in the nasal mucus of the grey teal, Querquedula
gibberifrons (Miller) (12). Hither of these birds may prove to be the definitive host.
Macfarlane (1952) described the lesions produced experimentally by C. parocellata
from the Murray River, and reported similar lesions from the Wagin lakes in
Western Australia (71).

In the shallow estuaries and salt-water lagoons of the New South Wales coast.
the lesions may be caused by the cerecariae of Awustrobilharzia terrigalensis S. J.
Johnston, 1917, which are emitted by the common marine snail, Pyrazus australis
Quoy and Gaimard, and have been shown by Bearup (1955, 1956) to be capable of
producing the characteristic rash. The adults live in the portal system of the seagull,
* Larus novae-hollandiae Stephens (10), (11).

(bo) “Creeping eruption” or ‘cutaneous larva migrans” is caused by the infective
larvae of foreign hookworms, and is characteristically an itchy, serpiginous rash,
which advances as the larva burrows along below the stratum granulosum. WHeydon

+ Probably the larval stage of Diphyllobothrium (Spirometra) erinacei (Rudolphi, 1819),.
the adults being parasites of carnivores.

BY M. JOSEPHINE MACKERRAS. 157

(1929) showed that it could be caused by the infective larvae of cat and dog hook-
worms, Ancylostoma braziliense de Faria, 1910, and A. caninum (Ercolani, 1859),
particularly by the former (44).

2. Visceral larva migrans.

This is a chronic condition, characterized by enlargement of the liver and extreme

eosinophilia, accompanied sometimes by pneumonitis. It is caused by the migration
of the larvae of the dog ascarid, Toxacara canis (Werner, 1782), through the viscera
and especially through the liver. This disease is usually seen in small children who
have had close contact with puppies. Two presumptive cases were reported by
Blanch (1956) (16).

OV

15.

16.

17.

18.

19.

20.

21.

22.

23.

24.

265.

26.

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BY M. JOSEPHINE MACKERRAS. 159

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161

STUDIES OF N-FIXING BACTERIA. VII.
CYTOCHROMES OF AZOTOBACTERIACEAE.
By F. J. Moss and Y. T. TcHAn.
[Read 30th July, 1958.]

Synopsis.
Using a reflectance spectrophotometric method, the representatives of the Azotobacteriaceae
were studied. For comparison, three Rhizobia and one other, a Gram-positive bacterium,
were also studied and compared.

Azotobacter has the highest QO. (4000-5000) (P. W. Wilson, 1951). However,
Burke (Wilson, 1951) reported that the effect of pO, on Azotobacter is independent of
the N source and cannot indicate the nature of the mechanism of N fixation. Wilson
and Fred (Wilson, 1951) and Bond (1950) found that O, is connected with the fixation
process of N. (by legume-plant-Rhizobium symbiosis) but direct participation is not
necessarily implied. It is thus interesting to investigate the terminal respiratory
oxidation enzymes (cytochromes) of N-fixing bacteria. A more detailed study of the
cytochromes of Azotobacter was carried out by Groucher and Kocholaty (1957), who
reviewed the literature of the subject. Using a reflectance spectrophotometer, they
reported the absorption spectra of some Azotobacter.

The present study compares a wide range of species of Azotobacter and Beijerinckia.
The influence of the presence of NH,OH which affects the N fixation mechanism
(Wilson and Burris, 1947, 1953) is studied in relation to the cytochrome spectra.

Method.

(a) Organisms: Nearly all known species of Azotobacter and Beijerinckia were
included in the experiment. For comparison, the three groups of Rhizobium and
Serratia marcescens are also included. The classification of Azotobacteriaceae is based
on the systematic classification of one of us (Tchan, 1953) combined with Jensen’s
classification (Jensen, 1955). The cultures were grown on Winogradsky’s medium
With sucrose, agar, and Mo. For the Azotomonas insigne, 2:°5% of glucose was added to
the above medium. For medium containing ammonium 5 c.c. of 5% ammonium sulphate
was used for 100 c.c. of the medium. Cultures were first grown on the N-free medium
for 24 hours in a test tube. They were used for inoculation of larger quantities of
medium for experimental work. The organisms were harvested in sterile water after
48 to 72 hours of incubation at 25°C. Their purity was checked by microscopical
examination.

(0) Reflectance measurements: The technique used was described by Giovanelli
(1947). All the organisms were examined in the reduced state. When oxidized
conditions were used, the organisms were washed twice in Winogradsky’s medium
without sugar and then pure O, was bubbled through for several minutes and
examined immediately.

: Results and Discussion.
(a) General description of cytochrome spectra of Azotobacteriaceae.

Cytochrome absorption maxima from this and other experiments are shown in
the table.

The Sorét band is present in all organisms examined. Smith reported a band at
4400A for Azot. chroococcum. In the present experiment an absorption band at
4800-48404 was present in all members of the Azotobacteriaceae. This band was
particularly strong for Az. vinelandii and Az. agilis. However, this band may not be
related to the green fluorescent pigment of the above two species since Beij. derxii

PROCEEDINGS OF THE LINNEAN Society oF NEW SoutH WALES, 1958, Vol. Ixxxiii, Part 2.
EF

162 STUDIES OF N-FIXING BACTERIA. VII,

(Tchan, 1957) has a strong green fluorescence but a weak band at this wave-length.
The fluorescent pigments of Azotobacteriaceae have been recently studied by Johnstone
(1956). This band seems to be absent (or too weak to be detected) from the Rhizobium
(peanut). The 50504 band described by Gourcher and Kocholaty was not found.
However, one of us (F.M.) has observed the development of a band in this position
from Aerobacter-aerogenes when an aerated culture was stored at 4°C. overnight. It
is suggested that this band is not sufficiently consistent for the purposes of classification.

TABLE 1.
Sorét flavin 6 ac a aaa,
Azot. chroococcum—
(Smith) .. 3 ae ae 428 440 be te 530 se 552 560 590 628
After several GmemnerPara ©
(Smith) as Pee 427 —_— a6 x 522 de 552 ao 590 oe
(Negelein and Gersher) O° = = an an — Sh fs 563 a 632

(Goucher and Kocholaty) ae 422 hs she 505 520 ts 550 ts 590 625

Moss and Tchan ae a2 420 480 ve ste 524 ae 552-4 og 590 632
Azot. vinelandii—

Wilson and Wilson io ae 420 an Bie Sts 523 ae 551 ae ae 625

Fujita and Kodama es We ot “4 ae of 521 531 550 563 590 632

Goucher and Kocholaty nie 420 Rye us a 525 Le! 555 ote te 630

Moss and Tchan Pee a6 418 ae 480 ae 522 Sr 552 ne BS 632

+NH, ss es oe 412 oe 480 se 525 we 553 ae ts 648
Azot. beijerinckii .. see ne os ice a 527 2 553 me a 630

Goucher and TOCHOLat ya ae 422 he ah bac a Ae a bt, eye ad

Moss and Tchan x Ae 420 he 480 us 522 a 554 tye 589 632
(var. acidotolerans) ne ae 417 sh .480 aye 520 ie 554 ae: 589 632

+NH, he ie ave hie 416 as 480 on 523 ote 553 ae 590 633
Azot. macrocytogenes 0O—

Moss and Tchan eb Me 422 ae 480 ee 524 ae 557 noe 600 632
Azot. macrocytogenes 1—

Moss and Tchan arn ae 423 Pa 480 af 526 be 558 ans 596 632

+NH, se ee os 418 ay 480 bg 524 Lhe 558 ot 590 635
Azot. agilis— .
Goucher and Kocholaty ve 427 Dis as 505 A 530 Ak 560 590 625
Moss and Tchan a ys 427 a 480 fe 525 so, 6SEY ie 590 632

+NH, tye a Be 420 as 480 bes 534 Ay aie 560 590 636
Azomonas insigne—
Moss and Tchan aa kis 423 Ae 470 ae 522 ss 554 as 590 632
+NH, os af ie 418 vs 480 = 521 3 556 ts 590 632
Beij. indica \

Moss and Tchan iy ag 418 445 480 hs 518 55 551 Ls 604
Beij. indica var. alba—

Moss and Tchan aA aS 415 ES 480 a 518 ths 551 a 604
Beij. lacticogens—

Moss and Tchan Se 20 424 50 480 20 56 525 556 _ De 598 630
Beij. derxit a brs Ee 418 Ae 480 Soe. Ss 527 555 Ae 589 630
Rhizobium trifolai—

Moss and Tchan ine aS 422, ae 480 =e ae 525 555 30 600 640
Rhiz. meliloti—

Moss and Tchan 4 c 418 as 475 ae a: 525 555 a 601
Rhiz. cow-peat group Cent}

Moss and Tchan oe ze 418 ae ve a a 523 552, oC 604
Serratia marcescens—

Moss and Tchan Bs ea 422 Ae 480 a E% 530 ao 560 Se 632
Bot. bei). an we a 420 “ie 490 500 Pes 525 552 fe 590 631
mutant white nidasde Ke i 412 ai +t Bs Si 520 549 Re se 650

BY F. J. MOSS AND Y. T. TCHAN. 163

Beij. indica and its var. alba exhibited a band at 5180A but not the two other
species of the genus (Beij. derxii and Bei). lacticogenes). No other member of the
Azotobacteriaceae or Rhizobium possesses this band.

The absorption at 5520-55804 was obtained with all organisms under examination,
but we were unable to observe absorption at 5600—5630A; Beij. indica and its variant
alba present a band at 5510A. The absorption band at 5900A which Goucher and
Kocholaty considered as differential of the chroococcum-agilis group was observed by
us with Az. beijerinckii and its variants.

We were able to confirm the presence of a band at 5900A described by Fujita and
Kodama. However, the value of this band for classification of: Azotobacter has limited
value. Az. macrocytogenes and Beijerinckia and Rhizobium present a corresponding
absorption band at (5960-60404).

The absorption at 6300A was present in all those examined except the Rhiz.
meliloti, Rhizobium of the cow peat group, Beij. indica, and Beij. indica var. alba.

The Rhizobium meliloti has a similar cytochrome spectrum to Beij. indica var.
alba, but the cytochrome spectra of other Beijerinckia and Rhizobium are different.
The Beij. derxii and Beij. lacticogens have rather similar cytochrome spectra to
Azotobacter. Between mutants, variants of the same species, some variation of
cytochrome spectra was noticed.

Within the same species the recorded cytochrome spectra vary with the authors.
It is probable that the organism used and experimental conditions were not comparable.

Concerning the cytochrome spectra of Azotobacteriaceae, it is not possible at
present to group the organisms as suggested by Goucher and Kocholaty unless more
information is available. Our experiment does not support Derx’s hypothesis, which
suggests that Beijerinckia may be related to the Rhizobium.

(0) The influence of Ammonium on the cytochrome spectrum of Azotobacter.

Wilson (1951) has pointed out that ammonium is completely accepted as a
nitrogen source to the exclusion of the nitrogen fixation reaction. It is of particular
interest that the addition of NH, ion produces Azotobacter with a modified cytochrome
spectrum. All Azotobacter species cultivated in the presence of NH, showed a shift
of the Sorét band towards the shorter wave-length. The 6300A band was generally
shifted towards a longer wave-length. There are also modifications of the absorption
bands at other wave-lengths but they are not consistent. These modifications become
more interesting when the oxidized spectra of cytochromes are compared with those
from organisms grown in ammonia. When the organisms were washed and oxygenated,
some bands disappeared. A shift of the Sorét towards the shorter wave-length and
6300A towards the red was observed. Goucher and Kocholaty made a similar
observation.

Nevertheless this aspect of the problem should not be overlooked. If the N fixation
is realized by a reduction process of molecular N, to NH,, it is not impossible to
imagine that the cytochromes participate in electron transfer of the reduction reactions.
Recent work by Hamilton et al. (1957) showed that N, appears specifically to oxidize
flavins and cytochrome b.

Acknowledgement.

The authors express their thanks to Dr. Giovanelli, of C.S.1.R.O., for the use of
the recording spectrophotometer. We are indebted to Misses M. Cunningham, E. Jackson
and M. Baird for their technical assistance.

Bibliography.

Bonp, G., 1950.—The importance of the oxygen factor in nodule formation and function.
Ann. Botany, 15: 95-109.

FusiTa, A., and KopAMA, T., 1954. (See Goucher and Kocholaty.)

HAMILTON, P. B., SHuG, A. L., and WILSON, P. W., 1957.—Spectrophotometriec examination of
hydrogenase and nitrogenase in Soy bean Nodule and Azctobacter. Proc. Nat. Acad. Sci.
U.S., 43: 297-304.

GoucHsER, C. R., and KocHouaty, W., 1957.—A comparison of the Cytochrome structure and
radiation effects in Azotobacter. Arch. Bioch. and Biophysic., 68: 30-38.

GIOVANELLI, R. G., 1957.—The application of diffuse reflection spectrophotometry to chemica?
analysis. Aust. J. Hap. Bio. and Med., 35: 143-156.

164 STUDIES OF N-FIXING BACTERIA. VII.

JBENSEN, V., 1955.—The Azotobacter-flora of some Danish water course. Aaert. Botanisk
Tidsskrift, 52: 148-157. :

JOHNSTONE, D. B., 1956.—The use of a fluorimeter in the characterizations of fluorescing
substance elaborated by Azotobacter. App. Microbiol., 5: 103-106.

TCHAN, Y. T., 1953.—Studies of N-fixing bacteria. IV. Taxonomy of genus Azotobacter
(Beijerinck, 1901). Proc. Linn. Soc. N.S.W., 78: 85-89.

TOHAN, Y. T., 1957.—Studies of N-fixing bacteria. VI. A new species of N-fixing bacteria.
Proc. Linn. Soc. N.S.W., 82: 314-316.

WILSON, P. W., 1951.—Bacterial physiology. Acd. Press. N.Y. (Biological N-fixation, p. 474.)

WILSON, P. W., and Burris, R. H., 1947.—The mechanism of Biological Nitrogen fixation.
Bact. Review, 11: 41-78.

WILSON, P. W., and Burris, R. H., 1953.—Biological Nitrogen fixation. A reappraisal. Ann.
Review of Microbiol., 7: 415-432.

165,

MIGRATION AND UTILIZATION OF RESERVE SUBSTANCES DURING FLIGHT
IN APHIS CRACCIVORA KOCH.

By Max Casimir, Entomological Branch, Department of Agriculture,
New South Wales.

(One Text-figure.)
[Read 28th May, 1958.]

Synopsis.

A histological examination of A. craccivora alatae indicated that an important fuel reserve
in this species is glycogen. Deposits of fat were also considerable in young adults, although the
fat body cells contracted in size to a narrow layer next to the cuticle about one to two days
after the final ecdysis and there was an associated mobilization of part of the glycogen from
the abdomen to other parts of the body. Glycogen reserves decreased as the aphids aged.

Both glycogen and fat were utilized during starvation and aphids survived a maximum
of 4-5 days without food. In moribund aphids the glycogen reserves had almost completely.
disappeared but appreciable reserves of fat were still evident.

Aphids, suspended from a fine wire in the laboratory and stimulated by a slight air
current, vibrated their wings strongly for periods ranging from a few minutes to several
hours. However, aphids more than five days old could not be induced to ‘‘fly’’ and there
was a narrow age range at which the aphids would readily vibrate their wings and at which
the duration of flight was most prolonged. This occurred one to two days after the final
ecdysis. Glycogen was progressively reduced in all regions of the body as the duration of
flight increased.

Recently arrived alatae collected at Gosford and Sydney on the New South Wales coast
in September showed greatly reduced glycogen reserves compared with swarming aphids
taken a few days before at Moree, several hundred iiles to the north-west. From the
appearance of the fat body itself the aphids in each locality were approximately the same
age, namely, 0-2 days old. It was concluded that the aphids had depleted their glycogen
reserves during migration in upper wind currents from the inland breeding areas to the
central coast.

Swarming populations at Moree consisted of several aphid species including Aphis
craccivora Koch, Myzus persicae Sulz., Lipaphis erysimi Kalt., and Capitophorus elaeagni
Del Guer., and these species were represented in roughly the same proportions in the Gosford
eollections, M. persicae being the dominant species. However, collections in the Sydney
district showed a majority of A. craccivora with very few M. persicae.

INTRODUCTION.

Black aphids have been reported by the New South Wales Department of
Agriculture infesting broad beans and French beans since 1922 but it was not until
1950 that the species was definitely identified as Aphis craccivora Koch (synonymous
with A. leguminosae Theob. and A. medicaginis Koch) (Dyce, unpublished thesis,
University of Sydney). Johnson (1957) indicated the presence of extensive breeding
grounds of this species in north-western inland areas of New South Wales. From
observations in the field, correlations with wind data and other sources, he advanced
the theory that plagues of A. craccivora experienced in some years .on the central coast
of New South Wales resulted from long distance dispersal of the aphid away from
these inland breeding areas.

The appreciable reserves of glycogen present in many insects (Babers, 1941)
suggest the general use of this readily available material as an energy source during
flight. The duration of flight to exhaustion in Drosophila varies with the age of the
fly and with the total glycogen content of the insect (Williams et al., 1943). However,
it has been shown that the principal source of energy for flight in the Desert Locust,

PROCEEDINGS OF THE LINNEAN Society or NEw SouTH WALES, 1958, Vol. Ixxxiii, Part 2.

166 UTILIZATION OF RESERVE SUBSTANCES DURING FLIGHT IN APHIS,

Schistocerca gregaria Forsk., was fat, the remainder being glycogen (Weis-Fogh,
1952). Fat also provides energy for flight in another migrating insect, Hutettix
tenellus (Fulton and Romney, 1940). Rough determinations of the distances flown
were made from analysis of the chloroform extractive contents of these leafhoppers
as they moved along an extended migration route away from the breeding area.

The object of the present paper is to provide evidence for long-range dispersal by
histological examination of the distribution of glycogen or fat and their mobilization
during sustained fight in alate forms of A. craccivora in the laboratory. An analysis
of migrant aphids collected in the field is also made to measure differences in reserve
substances of aphids obtained from their breeding grounds in the north-west of New
South Wales and aphids collected in coastal areas of New South Wales, the latter
having presumably been transported by winds from the north-western districts
(Johnson, 1957).

METHOD OF EXAMINATION OF STORED RESERVES.

Alate forms of the aphid were used in all experiments, and broad beans in winter
and cow peas in summer proved suitable hosts in that the aphid populations increased
rapidly on them and produced large numbers of alatae as the plants began to wilt
2-3 weeks after germination. However, in ihe spring and early summer it was
necessary to use scarlet runner beans, medics and clovers to maintain colonies, and
these were not as suitable for production of alatae. Fourth instar nymphs with wing
buds were removed from stock colonies 24 hours before the adults were required
and a large proportion of these became adults the following day. All host plants
were grown in earthenware pots covered with perspex tubes and maintained in a
glasshouse.

(a) Glycogen.

Alate viviparous females were fixed in Carnoy’s solution and split in halves by a
median longitudinal cut with a fragment of a safety razor blade, following the method
described by Wigglesworth (1949) for Drosophila. The half insects were stained with
a saturated solution of light green in 90 per cent. alcohol and the glycogen revealed
by the iodine method (Wigglesworth, 1942) and with Best’s carmine (Carleton and
Leach, 1947). The two methods gave similar results, the glycogen appearing as
brilliant red granules with Best’s carmine and a mahogany colour with iodine.
Microtome sections of Carnoy-fixed material were cut in paraffin and stained with
Best’s carmine.

(b) Fat.

The usual method of fixing in Bouin’s fluid and staining with Sudan Black B
proved unsuitable with the half-aphids and fat was demonstrated by fixing aphids in
Baker’s formaldehyde-calcium (Baker, 1945) for five days or more. Half-aphids were
then prepared as above and washed in several changes of distilled water, transferred
to 50 per cent. glycerine, and mounted in glycerol jelly. Baker’s formaldehyde-calcium
fixes all tissues but leaves fat and, examined under the binocular microscope, it
appears as whitish globules in reflected light or bright shining yellow in transmitted
light, the rest of the insect tissue having a dark appearance. The aphids were dipped
in 50 per cent. alcohol before being immersed in the fixative to overcome difficulty in
wetting the insects. The method was useful for contrasting relatively large changes
in the appearance of fat reserves but small differences were not readily perceptible.

(c) Comparison of Reserves in Different Aphids.

The comparison was put on a roughly quantitative basis by giving each prepared
slide of stained half-aphids a serial number and comparing with an appropriate
control slide. A score system was used whereby separate parts of the body of each
aphid were allotted points with a possible maximum of ten in each case according
to the area and intensity of staining in the region. Thus, for example, glycogen in
the fat body of the abdomen, embryos, alimentary canal, thoracic muscles, head, etc.,
was compared in different slides and with a control slide and finally given score
ratings.

BY MAX CASIMIR. 167

STORED RESERVES IN ALATE APHIDS.
(a) Distribution of Glycogen and Fat.

The general distribution of glycogen in an adult viviparous female, approximately
one day old, is shown in Text-figure 1. The bulk of the glycogen is situated in the
fat body, especially that of the abdomen, although appreciable amounts occur along the
surface of the indirect flight muscles and their points of insertion in the wall of the
thorax. On the whole, glycogen in A. craccivora was not as conspicuous as the massive
deposits reported in Drosophila (Wigglesworth, 1949).

In the fat body, discrete fat droplets were particularly obvious in the caudal
region and at the base of the cornicles in the abdomen, in the prothorax, mesothorax
and head, and the scutellum of the thorax.

Text-fig. 1.—-The distribution of glycogen in an alate viviparous female Aphis craccivora,
0-1 days old, shown in a sagittal section. A, Glycogen between muscle bundles and insertions ;
B, Glycogen in the thoracic fat body; C, Fat body of abdomen with glyccgen; D, Hmbryos
with glycogen; EH, Rectum; F, Midgut; G, Fat body of head; H, Pharynx.

*(b) Changes in Reserves with Age.

From six to eight aphids, for each of the age groups 0, 1, 2, 3, 4, 7 and 14 days
old, were tested for glycogen and a similar number for fat. All aphids used were
removed from various stock broad-bean plants as nymphs and maintained on the one
plant throughout the experiment, aphids of the required age being taken from the
plant at intervals.

In the newly emerged adult what Wigglesworth (1949) termed the “larval fat
body” was still evident and filled a large part of the abdomen. The loosely packed
fat body cells contained very heavy deposits of fat and an intense staining reaction
for glycogen indicated large reserves of the latter. Glycogen was at a relatively low
level in the head and thorax but fat was here also present, in large amounts as
discrete droplets.

During the next 1-2 days the fat body cells in the abdomen contracted in size
to a narrow layer next to the cuticle with a consequent reduction in glycogen and
fat content. There was, at the same time, an increase in the glycogen content of the
fat body and muscles of the thorax, in the head and the alimentary canal. ‘This
glycogen level was maintained in all regions up to the fourth day but on the seventh
day there was a reduction in all regions except the head and alimentary canal, while
on the fourteenth day glycogen had been reduced in all parts of the body to about
two-thirds of that observed in a one-day-old aphid. The embryos and alimentary canal
maintained a fairly constant level of glycogen throughout the life of the aphid.

No decisive change in fat content was observed as age increased, after the initial
Major reduction in the size of the fat body cells.

(c) Changes in Reserves during Starvation.
One-day-old aphids were confined in a small perspex and organdie cage which was
placed over a dish of water to maintain an almost saturated atmosphere. The aphids

168 UTILIZATION OF RESERVE SUBSTANCES DURING FLIGHT IN APHIS,

immediately climbed to the top of the cage towards the light from a window near by
and after a brief period of activity remained motionless until death, which occurred
after a maximum of five days. Six aphids were placed in fixative at eight-hourly
intervals until death and stained half-aphids were compared with control slides of
aphids of the same age taken directly from the host plant.

A small drop in glycogen was observed after 24 hours but it was not until after
40 hours that glycogen was reduced significantly in the fat body. In the later stages
(90-120 hours) very little glycogen remained in the fat body but appreciable amounts
were retained in the thoracic muscles and alimentary canal. Similarly, fat was
reduced only slightly until about 40 hours without food, when there was a progressive
reduction until death, although, even in the moribund aphid, about half the fat
present in a recently-fed insect remained.

UTILIZATION OF RESERVES DURING FLIGHT.
(a) Flight in Suspended Aphids.

It is well known that if one of the higher Diptera is suspended in still air it will
readily vibrate its wings. However, some insects require further stimuli before they
will vibrate their wings and maintained flight has been induced in Locusta species
(Kennedy ez al., 1948) by suspending the insects in a current of air. The distribution
of stress within the body of the insect may be expected to be most natural when
the position of attachment is as close as possible to the wing bases (Holiick, 1941).

A fine phosphor-bronze wire with a diameter of 0:08 mm. was soldered to the end
of a long entomological pin from which the head had been removed. ‘The point of the
pin was inserted into a cork and clamped to a small retort stand. The aphid, lightly
anaesthetized with ether for 15 seconds, was placed on a sheet of cork on the stage
of a binocular microscope. A drop of water-colour paint made into a thick paste
Was painted on the tip of the bronze wire and the wire was then held horizontally
and symmetrically to the aphid so that the minute paint globule came in contact with
the mesothorax when the wire was gently lowered from above and behind the aphid.
The paint quickly dried and secured the aphid to the wire without interfering with
wing movement. It was essential that the paint be of the correct “gluey” consistency :
so that it would adhere to the wire in the form of a minute droplet. Aphids required
from 10 to 20 minutes to recover fully from the ether. A sharp puff of air then
generally resulted in the aphid elevating its wings to the ‘ready-to-fly”’ position.

In addition to the removal of the tarsi from contact with any object a further
stimulus of air movement was required before the aphids would vibrate their wings.
This was provided by an electric fan, blowing through a mosquito netting screen, so
that the resultant draught reaching the aphid was not more than 90 feet per
minute. Consistent wing movement was not possible at wind speeds greatly above this.
A sudden puff of wind from in front and below was then sufficient to induce the aphid
to vibrate its wings. Some aphids were very erratic in their response and constant
stimulation was required, while others “flew” strongly for hours in the steady air
draught. Flight to complete exhaustion was apparently not obtained.

An attempt was made to conduct the experiment at a constant temperature of
80°F. and relative humidity of 85 per cent. in a warm room but only a few aphids
responded under these conditions. The results are summarized in Table 1, where
duration of flight in the warm room as well as at room temperature is given for the
different age groups. A wide range of temperature and humidity conditions Was
encountered in the latter as the experiment was carried out over an extended period.
of time, temperature ranging between 50° and 90°F. during the period of testing.

(0) Changes in Stored Reserves during Flight.

Sixteen aphids, having “flown” for periods ranging from 20 minutes to 540 minutes,
were tested for glycogen by the carmine and iodine methods. A progressive reduction
of glycogen in all regions of the body except the embryos was observed as the
duration of flight increased. In the aphid that had flown continuously for nine hours
glycogen was negligible in the thorax but traces remained in the head and fat body
of the abdomen. Sixteen aphids were also tested for fat; they had flown for periods

BY MAX CASIMIR. 169

ranging from 7 minutes to 415 minutes and it was only in those aphids that had flown
for more than 20 minutes that a slight reduction in fat could be observed when
compared with a control slide. There did not appear to be a progressive reduction
of fat as the duration of flight increased, although comparisons were difficult as most
of the aphids were suspended for several hours at an age when a large reduction was
taking place in the size of the fat body cells and no direct conclusion could be
made concerning the importance of fat as a fuel during sustained flight.

TABLE 1.
Duration of Aphid Flight in Laboratory.

Aphid Age Number of Aphids Tested, with Duration of Flight
Groups in Minutes.
(in Days).

At Constant Temperature (80° F.) and Humidity (85%).

0-1 0, 0, 0, 0, 0, 0, 6, 0, 0.
1-2 0, 0, 0, 9, 0, 0, 18, 0, 78, 48, 0
2-3 0, 0, 0, 24, 0, 45, 21, 0, 62, 0, 0, 23.
3-4 23, 0, 0, 17, 0, 0
4-5 0, 0, 0, 0, 0, 0
6-7 0, 0, 0, 0
7-8 0, 0, 0, 0, 0, 0, 0, 0, 0, 0
At Room Temperature (Variable).
0-1 0, 0, 16, 0, 14, 0, 0, 12, 0, 0.
1-2 317, 547, 415, 307, 372, 292, 91, 224, 374, 96, 190, 74. 285, 273, 251.
2-3 0, 0, 0, 45, 0, 0, 0, 82, 0, 0.
3-4 124, 0, 0, 45, 0, 0, 0, 82, 0, 0.
4-5 0, 29, 0, 0, 0, 0, 0, 0.
6-7 0, 0, 0, 0, 0.
7-8 0, 0, 0, 0, 0.

OBSERVATIONS ON SwaARMING APHID POPULATIONS.

A visit was made to Moree in September, 1952, and aphids collected. On the
return trip observations were made at Gosford and again at Sydney a few days later.
Only alate aphids were considered in the field collections.

Moree is situated approximately 300 miles north-west of Sydney and it is an
area of extensive black soil plains upon which Burr medic, Medicago hispida var.
denticulata, thrives in the spring months. Burr medic is a favoured host of
A. craccivora. According to local reports there had been a big aphid increase in Moree
during August but after a week of rain the aphids almost vanished. There was
another increase in numbers during September, when heavy swarming occurred.
These swarms were particularly dense on 12th, 13th and 14th September in Moree
and several aphid species were thought to be involved. Observations were commenced
on 15th September, when there was evidence of previous huge colonies in the very
large numbers of cast skins in the old medic stands. The medics and other annual
Pasture plants were commencing to wilt and a mild swarm of aphids was observed
on the afternoons of 15th and 16th September. Samples were collected from various
host plants and from swarming aphids in the air. Wasp parasites and predatory
Coccinelid beetles were very common amongst the reduced aphid colonies.

It was observed that the main host of A. craccivora in the Moree district was
the widespread Burr medic. Other aphid species present were Myzus persicae Sulzer
which thrived on many host plants including the weeds Marshmallow, Malva parviflora
and milk thistle, Sonchus oleraceus, and the turnip aphid, Lipaphis erysimi Kalt.,
which was found on wild turnip, Brassica campestris. These weeds were unusually
abundant in 1952. Of the aphid species taken in the air, 62 per cent. were M. persicae,
22 per cent. A. craccivora, 14 per cent. I. erysimi and 38 per cent. Capitophorus
elaeagni Del Guer. from a total sample of 107 aphids collected by net.

170 UTILIZATION OF RESERVE SUBSTANCES DURING FLIGHT IN APHIS,

At Gosford on the central coast of New South Wales the first report of black
aphids on bean crops in 1952 was received by the Government Entomologist on
17th September. On 18th September several farms in the hbean-growing area were
visited; the aphid situation was similar on all farms, there being 3-4 A. craccivora
alatae per plant on about 40 per cent. of the bean plants in each crop. Aphids were
observed in the air and a sample consisted of 57 per cent. M. persicae, 25 per cent.
A. craccivora, 14 per cent. L. erysimi, and 3 per cent. C. elaeagni from a total of 173
aphids collected.

On 22nd September, at Sydney, 50 miles south of Gosford, very heavy aphid
Swarms were observed in several districts. Strong, dry winds kept the aphids
dispersed and after a cool southerly change about mid-day they disappeared. However,
in the late afternoon, when calm and warmer conditions prevailed, large numbers
were observed swarming above grass, etc., as they left unsuitable plants, and specimens
were collected. The latter consisted of 90 per cent. A. craccivora, 6 per cent. L. erysimi,
2 per cent. M. persicae and 2 per cent. C. elaeagni, from a total sample of 402 aphids.

STORED RESERVES IN FIELD—COLLECTED APHIDS.

Although several aphid species were collected, only A. craccivora were considered
in the glycogen and fat determinations. Specimens were placed in fixative when
collected and later tested for glycogen and fat in the laboratory. A slight variation
in the glycogen technique was necessary, as Carnoy’s solution fixes tissues in less than
one hour and Bouin’s fluid was used as an alternative so that the fixed aphids could
be stored for several days before staining.

At Moree 23 alatae were collected from French beans on 15th September. Both
glycogen and fat were at a high level in the fat body and thoracic muscles, being
comparable with that observed in 0—2-day-old aphids bred in the laboratory on cowpeas.
At Gosford 33 alatae were taken from French beans and 20 caught in flight on
18th September. The former had evidently recently settled on the host plants, as
no young were present. Glycogen was similar in both groups, being almost negligible
in the fat body of the abdomen and thorax and greatly reduced in the thoracic
muscles and head. No record of fat content was made. At Sydney 47 alatae were
taken in flight and 55 collected from Melilotus indica and Trifolium recumbens plants
on 22nd September. There were no aphids on these plants the previous day. Glycogen
in the aphids taken in flight was very much reduced and similar to that observed
in aphids collected at Gosford, although the level of glycogen in the fat body of the
abdomen was slightly higher. However, the fat body of the aphids appeared to be
extensive, particularly in the abdomen, and was similar to that observed in 0—2-day-old
aphids bred in the laboratory. The glycogen level in the aphids taken from the
legumes was slightly higher than in those taken in flight but was not comparable
with the large deposits in aphids collected at Moree. The fat body was greatly
reduced in size in all regions of the body.

A further 77 alatae were collected from a plot of WM. indica at Sydney University
on 9th October, 17 days later. These were not new arrivals, as no swarms occurred
in Sydney after 23rd September, nor could they have been produced as offspring of
previous migrants as alate viviparous females invariably give rise to apterous nymphs.
Glycogen in these insects was similar to that in aphids collected at Moree, although
the staining reaction was not quite so intense, indicating that, although well fed,
these aphids were probably somewhat older than those collected at Moree or at Sydney
previously.

DISCUSSION.

The age at which aphids flew most readily in the laboratory and at which the
duration of flight was most prolonged coincided with the maximum concentration
of glycogen in the body of the aphid. These deposits of glycogen in the fat body of
the thorax and abdomen were drawn on extensively by the aphids during starvation
and for vibrating the wings while suspended from a fine wire. The extensive use of
fat as a fuel during flight was not demonstrated, although the method of examination
for fat was not sufficiently precise to observe other than major changes in the

BY MAX CASIMIR. i7/iL

appearance of the fat body and no conclusion was drawn as to the importance of
fat as an energy source during flight. It was sufficient to know that the more readily-
observed glycogen reserves were progressively depleted during flight and could be
used as a pointer to any recent muscle activity in the aphids.

There was a very small range of ages at which aphids could be induced to fly
in the laboratory. A sharp peak of response was obtained in aphids 1-2 days old
(Table 1) and a period of settling, involving feeding and reproducing on a suitable
host plant, had an adverse effect on the ability of the aphids to vibrate their wings.
Johnson (1953) indicated that autolysis of the flight muscles occurred in several
aphid species and that flight was not possible for more than a few days after the
final ecdysis. Thus, not only was it necessary to stimulate special receptors by air
movement but it was apparent that the aphid must be in a particular physiological
‘state before flight could begin. Swarming in natural populations could only occur
when several conditions, such as the age of the aphids themselves, the state of the
host plants, and the related climatic conditions, combined to overcome the natural
resistance of aphids to any movement away from their food and shelter habitat.

It was only in very young adults that the fat body occupied extensive areas of
the abdomen, after which the celis were reduced in size to form a narrow layer
next to the cuticle, and aphids collected from swarming populations at Moree, Gosford
and Sydney in September all showed this extensive “larval fat body’ and were
apparently of approximately the same age.

The flying aphids collected at Gosford and at Sydney showed greatly depleted
glycogen reserves compared with aphids in the large swarms at Moree a few days
earlier and it is probable that these or similar swarms were transported by winds to
the New South Wales coast from their inland breeding areas several hundred miles
to the north-west. Migrants collected in Sydney after feeding for 2-3 weeks on a
favoured host had their glycogen reserves restored to the normal high level. The
aphids, because of their slow speed of flight and the presence of a mountain range
between their inland breeding grounds and the coastal plain, could not have travelled
unaided and were probably carried by upper prevailing winds as suggested by
B. Johnson (1957). However, C. G. Johnscn (1951) considered that once active flying
ceased the aphids might not remain aloft for long and, under natural conditions,
might continue to vibrate their wings and use up available glycogen reserves although
directed by prevailing winds.

The host plants of the several aphid species observed at Moree were annuals
which commenced to die at about the same time of the year and gave rise
simultaneously to conditions favourable for the production of alatae. Thus M. persicae,
L. erysimi and C. elaeagni, as well as A. craccivora. were observed swarming at
Moree and again at Gosford in roughly the same proportions, with M. persicae
predominating. UL. erysimi, C. elaeagni and A. craccivora and only a few WM. persicae
were observed at Sydney. The aphid arrivals at Gosford on 17-19th September may
have been the result of a single period of swarming in the Moree region while the
migrants observed at Sydney on 22nd September could have originated from a more
southerly area such as Gilgandra or Coolah, where the favoured hosts of M. persicae
may not be so widespread.

Long-range dispersal in favourable years by upper wind currents to the coastal
areas of New South Wales may be a feature of several small-bodied, flying insects,
particularly aphids, and the climatic conditions of the inland breeding areas allied
with the prevailing upper air currents could have an important bearing on the
likelihood of infestation by these insects on coastal crop and orchard areas, when even
a close study of the local coastal conditions would give no indication of a likely
infestation.

Acknowledgement.
This work was carried out in the Zoology Department of the University of Sydney
under the direction of Dr. A. R. Woodhill, whose guidance and helpful criticism are
gratefully acknowledged.

172 UTILIZATION OF RESERVE SUBSTANCES DURING FLIGHT IN APHIS.

References.
BABERS, F. H., 1941.—Glycogen in Prodenia eridania, with special reference to ingestion of
glucose. J. Agr. Res., 62: 509-530.
Baker, J. R., 1945.—Cytological Techniques. Methuen, London.
CARLETON, A. M., and LEACH, E. A., 1947.—Histological Techniques. Oxford Univ. Press.
FuLTON, R. A., and ROMNEY, V. E., 1940.—The Chloroform-soluble Components of Beet-
leafhoppers as an Indication of the Distance they Move in Spring. J. Agr. Res., 61: 737.
Houuick, F. J. S., 1941.—The Flight of the Dipterous Fly, Muscina stabulans Fall. Phil.
Trans. Roy. Soc. Lond., 230B: 357-390. -
JOHNSON, B., 1953.—Flight Muscle Autolysis and Reproduction in Aphids. Nature, 172: 813.
--, 1957.—Studies on the Dispersal by Upper Winds of Aphis craccivora Koch in New
South Wales. Proc. LINN. Soc. N.S.W., 82: 191-198.
JOHNSON, C. G., 1951.—The Study of Wind-borne Insect Populations. Science Progress, 39: 41.
KENNEDY, J. S., AINSWORTH, M., and Toms, B. A., 1948.—Laboratory Studies on the Spraying
of Locusts at Rest and in Flight. Anti-Locust Bull. 2.
PANTIN, C. F. A., 1948.—Wicroscopical Techniques for Zoologists. Cambridge Univ. Press.
Weis-FocH, T., 1952.—Fat Combustion and Metabolic Rate of Flying Locusts. Phil. Trans.
Roy. Soc. Lond., Series B. Biol. Sct., 237: 1-36.
WIGGLESWORTH, V. B., 1942.—Storage of Fat, Protein and Glycogen in Mosquito Larvae.
J. Exp. Biol., 19: 56-59.
, 1949.—Utilisation of Reserve Substances in DrosophiJa during Flight. J. Hap. Biol.,.
26: 150.
WILLIAMS, C. M., BARNESS, L. A., and SAwymER, W. H., 1943.—The Utilisation of Glycogen by
Flies during Flight and Some Aspects of Physiological Ageing of Drosophila. Biol. Built,
Marine Biol. Lab.. Woods Hole, Mass., 84: 2263-272.

173

A NEW BIRD-FLHA FROM TASMANIA.

By F. G. A. M. Smit, British Museum (Natural History), The Zoological Museum,
Tring, Herts.

(Communicated by Mr. D. J. Lee.)
(Ten Text-figures.)
[Read 30th July, 1958.]

Synopsis.

Three specimens of a new species of Hoogstraalia, a Pygiopsyllid genus of bird-fieas,
were found in the collection of the late Dr. HK. W. Ferguson; they had been bred from a
nest of the Tasmanian Scrub-tit, and the new species which they represent is described here
in comparison with the hitherto only Known member of the genus, Hoogstraalia turdella Traub.

INTRODUCTION.

When recently sorting out an accumulation of various documents relating to fleas,
in the Tring Museum, we found an envelope which contained several photographs of
a peculiar male flea, and on the outside of the envelope was a note (obviously written
a good many years ago) that the photographs had been received from .Dr. E. W.
Ferguson of Sydney. Since Dr. Ferguson’s active work on Australian fleas took place
mainly in the early twenties of this century, it seems probable that the photographs
may have been taken about 35 years ago. On examining the photographs I saw, to my
amazement, that the flea represented an undescribed species of the Pygiopsyllid genus
Hoogstraalia, erected by Traub in 1951 for his new species H. turdella, known only
from the male holotype, collected from a thrush in the Philippines.

Following advice kindly given me by Dr. G. M. Dunnet, I wrote to Mr. D. J. Lee
at the School of Public Health and Tropical Medicine in Sydney and asked him whether
he would try to trace the photographed specimen. Mr. Lee responded generously to
my request and I am most deeply indebted to him, not only for discovering the flea
in question but also for finding in Ferguson’s collection a male and a female of the
same species with the same data, and for obtaining permission to send me the three
specimens on loan for study and description.

The genus Hoogstraalia is a very peculiar one. As pointed out by Traub, it differs
from all other members of the Pygiopsyllidae by the presence of a genal ctenidium
and of a sinus in the dorso-anterior margin of the prosternosome for the reception of
one end of the first link-plate.

HOOGSTRAALIA VANDIEMENI, sp. nov. (Figs. 1-10).

Type material: Male holotype, female allotype and one male paratype bred from a
hest of the White-breasted Scrub-tit Acanthornis magna, collected on Mt. Wellington,
Tasmania. The specimens are in the E. W. Ferguson collection, now housed in the
School of Public Health and Tropical Medicine in Sydney.

Diagnosis: The new species is closely related to Hoogstraalia turdella Traub,
hitherto the only known representative of the genus. The male differs from that of
H. turdella by having a much more strongly convex frons, by the presence of fewer
Spines in the genal ctenidium (11 instead of 13) and more numerous setae on the nota
and terga, by the different outline of the posterior margin of tergum VIII, the fixed
process of the clasper is shorter and basally broader, the dorso-anterior margin of the
movable process convex instead of slightly concave and also with a somewhat different
chaetotaxy, the upper lobe of the bifid apical part of the distal arm of sternum IX is
shorter and the anal segment differently shaped. No comparison of females can be
given, since that sex of H. turdella is not yet known.

PROCEEDINGS OF THE LINNEAN SOCIETY OF New SoutH WALES, 1958, Vol. Ixxxiii, Part 2.

174 A NEW BIRD-FLEA FROM TASMANIA,

Description. Head (Figs. 1, 2): Fronto-clypeal margin strongly convex in the
male, less so in the female. Oral margin concave. Anterior wall of frons rather
narrow and of uniform width throughout. Submarginal frontal row consisting of

$

Text-figs. 1-4.—Hoogstraalia vandiemeni, sp. nov.: 1, Head, prothorax and fore coxa of
male (paratype). 2, Head of female (allotype). 3, Hind tibia of female (allctype). 4, Fifth
hind tarsal segment of female (allotype).

4

seven or eight slender setae. Ocular row of three setae, the long ocular seta placed
well in front of the eye. Between the submarginal and ocular rows one large seta and
a number of small and minute setae. The oblique genal ctenidium consists of 11
straight, fairly long and sharply pointed spines (in the female allotype there are 11 on

BY F. G. A. M. SMIT. 175

one side, 12 on the other); the uppermost of the genal spines is out of alignment with
the others and is situated behind the eye and just above the dorsal margin of the
genal process. The genal process has an obtuse apex. Hye large, not sinuate. Tentorial
rod fully developed, running from a point below the apex of the antennal clava to
just in front of the middle seta of the ocular row. The largest setae of the antennal
pedicellus reach to nearly the middle of the clava in the male, to near the apex of the
clava in the female. Antennal fossa closed, bordered dorsally by a number of thin setae.
Postantennal region of head with three rows of setae, the first consisting of four to
six, the second of five to seven, and the third of six setae. The occipital groove in
the male is rather shallow. The apex of the maxillary palp reaches to about the
middle of the fore coxa, while the tips of the epipharynx, of the finely serrated laciniae
and of the five-segmented labial palps reach to about two-thirds the length of the
fore coxa. In the female the interantennal suture is dorsally uninterrupted, but the
portion of the head anterior to the suture does not overlap the hinder portion as in
a true fracticipit head.

Thorax (Figs. 1, 5): Pronotum with two rows of setae, the first row consisting of
six to nine setae, the second row of seven setae each side; pronotal ctenidium with
31-33 long and slender spines. Prosternosome with a sinus for the reception of one
end of the first link-plate. Mesonotum with a main row of six setae each side, preceded
by numerous small setae which are scattered all over the surface; two pseudosetae
each side under the mesonotal collar near the dorsum. Mesepisternum much higher
than long, with one seta; mesepimeron with four or five setae in the male, in the
female with one seta on one side, three on the other. Metanotum with a main row of
five to six setae each side, preceded by a number of smaller setae which form about
three irregular rows; the large metepisternum with only one seta; metasternum with
a squamulum and with one large and one or two minute lateral setae, metepimeron
with 13-17 setae. *Pleural arch of metathorax well developed, as are also the pleural
rod and ridge.

Legs (Figs. 1, 3, 4): Fore coxa as in Figure 1. Mid coxa with an uninterrupted
oblique pale suture on the outer side, without setae on the inner side but on the
outer side with a patch of a dozen or more setae near the apical half of the anterior
margin. Hind coxa with a patch of about thirty setae adjoining the apical two-thirds
of the anterior margin, and a smaller group of small setae on the inner side near the
apical third of the anterior margin. Fore femur on the outer side with about ten
small lateral setae, several setae along the apical half of the dorsal margin and one
larger seta near the ventral margin at proximal fourth; on the inner side only one
small seta on the basal part. Mid femur with a ventral submarginal row of five to six
small setae and a larger one near the apex.. Hind femur with a ventral submarginal
row of five to eight setae, of which the apical one is the largest. Fore and mid tibiae
with six notches in the dorso-posterior margin, each notch bearing two setae (the
outer of the two setae in the penultimate notch often shifted a little away from the
notch onto the lateral surface). Hind tibia (Fig. 3) with seven notches in the dorso-
posterior margin, bearing 2, 2, 1, 2, 2, 2 (sometimes 1) and 2 setae respectively from
base to apex; on the outer lateral surface of the hind tibia about two dozen slender
setae. None of the tarsal setae reaches beyond the apex of the next segment. Fifth
tarsal segment (Fig. 4) of all legs with six pairs of lateral plantar setae, the three
apical pairs consisting of rather small setae, and four subapical plantar setae except
on the last segment of the fore tarsus which usually seems to bear only one such
seta (on the fifth tarsal segment of one fore leg of the female allotype there are four
subapical plantar setae); on the plantar surface there are also about 15 very small
setae.

Abdomen: Main row of setae on terga I-VII in the male with 4, 8, 8 (9), 8 (7),
8 (9), 7 and 7 setae respectively on each side; in the female with 4, 8, 8, 7 or 8,
7 or 8, 7 and 4 setae. These rows are preceded by numerous smaller setae which
may form three irregular rows. The most ventral seta of the main row on terga II-VII
is inserted below the level of the spiracle. Terga II-V in the male with 3 (2), 2 (3),
1 (2) and 1 (0) short marginal spinelets each side near the dorsum; in the female the

176 A NEW BIRD-FLEA FROM TASMANIA,

Text-figs. 5, 6—Hoogstraalia vandiemeni, sp. nov.: 5, Mesothorax, metathorax and
tergum I of male (paratype). 6, Terminal segments of male (holotype) with the omission
of clasper and sternum IX.

BY F. G A. M. SMIT. UTC

numbers of spinelets on each side of terga II—V are 3, 2, 1 and 1 (on one side of
tergum V of the only known female there is a spiniform seta instead of a normal
spinelet). Tergum VII with two antesensilial setae each side in both sexes; in the
male (Fig. 6) the upper seta is about half the length of the lower; in the female
(Fig. 10) the upper seta is about two-fifths the length of the lower. Margin of tergum
VII below the antesensilials markedly concave, especially in the female. Sternum II
(basal abdominal sternum) in both sexes with one ventro-marginal seta each side,
without setae on the lateral surface. Sterna III—VII in the male and III—VI in the
female with a main row of four setae each side, the rows preceded by a group of
smaller setae.

Modified abdominal segments and genitalia. Male (Figs. 6-8): Tergum VIII
(Fig. 6) small, bearing anterior to the fairly large spiracular fossa about a dozen
small setae, but no setae below the fossa. Sternum VIII (Fig. 6) large, the dorso-
posterior margin smoothly rounded except for a portion, which includes a short
ventro-posterior marginal row of about six thin setae, where the posterior margin is
straight or feebly concave, the very marked angle present in H. turdella above this row
of setae not developed in the present species. On the outer surface of sternum VIII
are numerous smallish setae. As in H. turdella, the ventral margin of sternum VIII
forms cephalad an elongate ellipsoid apodeme which protrudes deeply into the seventh
sternum; an intersegmental membranous process extends caudad from the apodeme,
dorsally joining up with the base of the distal arm of sternum IX. Fixed process of
clasper (Fig. 7) sugarloaf-shaped, with a number of small setae on the inner side
near the dorsal margin; its apex smoothly rounded and the sides much more divergent
than in H. turdella. Along the posterior margin of the large corpus of the clasper,
which projects strongly caudad, a row of about a dozen slender setae of which the
lower five are markedly larger than the rest. Manubrium long and narrow, more
strongly curved upwards than in H. turdella. Movable process of clasper (Fig. 7)
large, triangular, with convex dorso-anterior and posterior margins; along the former
Margin numerous minute setae and along the iatter margin a few small setae near
the obtuse apex and farther downwards along the margin seven or eight fairly long
ones. No setae on the outer lateral surface of the movable process, but on the inner
surface scattered minute setae are present. Proximal arm of sternum IX (Fig. 7)
greatly expanded apically, shaped like the head and neck of a curlew or ibis. Distal
arm of sternum IX divided into an anterior non-setose part and an apical portion
which is bifid in the apical half; the anterior (or dorsal) lobe of the bifid portion is
narrow and bears a number of minute setae along the dorsal margin and many long
or longish ones at and near the apex; the posterior (or ventral) lobe is nearly as
long as the anterior, but is much broader, with a rounded apex, and bears on each
side a row of six short spiniforms and proximad to these three large spiniforms
which are apically curved, the two most proximal especially so. Proximad to the three
large spiniforms are several small setae and a marginal dense patch of minute setae.
Sensilium with 18 trichobothria each side. Anal segment long and rather narrow
(see Fig. 6). Phallosome (Fig. 8) virtually as in H. turdella, described in detail by
Traub.

Female (Figs. 9, 10): Posterior margin of sternum VII (Fig. 10) with a deep and
narrow lateral sinus dividing an oblique and blunt short dorsal lobe from a more
rounded ventral lobe which extends farther caudad than the dorsal lobe, the whole
of the ventral lobe and most of the dorsal one strongly sclerotized; the sternum with
a main row of five setae each side, preceded by numerous smaller setae which are
not arranged in rows. Upper portion of tergum VIII without setae below the large
spiracular fossa but with a group of seven short setae anterior to it; on the lower
part of tergum VIII about a score of setae, of which the posterior and more dorsal
ones are the largest. Four normal-sized genital setae on the inner side of tergum VIII,
and anterior to these several minute ones. Sternum VIII not much sclerotized, with
a few minute setae at the blunt apex. Sensilium strongly convex, with 18 tricho-
bothria each side. Anal tergum with a number of slender setae; anal stylet four
times as long as broad, with one apical seta which is only a little longer than the

G

178 A NEW BIRD-FLEA FROM TASMANIA,

Text-figs. 7, 8.—Hoogstraalia vandiemeni, sp. nov.: 7, Clasper and sternum IX (holotype).
8, Phallosome (holotype).

stylet, and two short preapical setae. Anal sternum with a markedly straight ventral
margin, along which are placed four longish setae. Ductus bursae (Fig. 9) forming
a double bend; bursa copulatrix rather long and fairly broad; no ductus obturatus;
ductus spermathecae not very long, only just over twice as long as the bulga of the
spermatheca. Spermatheca (Fig. 9) with a trapezoid bulga, the dorso-posterior angle

BY F. G. A. M. SMIT. 179

Text-figs. 9, 10.—Hoogstraalia vandiemeni, sp. nov.: 9, Genital ducts and spermatheca
(allotype). 10, Terminalia and spermatheca (allotype).

180 A NEW BIRD-FLEA FROM TASMANIA

of which forms a distinct hump; the hilla rather short and strongly curved, the
portion of it which protrudes deeply into the bulga about twice as wide as the free
portion.

Length: g, 24-3 mm., 9, 3 mm.

Remarks: The only known specimen of Hoogstraalia turdella Traub was collected
from the body of a thrush (Turdus poliocephalus kelleri Mearns), taken at an altitude
of 7800 feet on Mt. McKinley on Mindanao, Philippine Islands. Traub (1951) suggested
on the basis of the large number of pronotal spines that H. turdella is a true bird-flea;
his suggestion has been confirmed by the discovery of the species described above as
H. vandiemeni, the specimens of which were bred from the nest of the White-breasted
Serub-tit (Acanthornis magna Gould), which occurs only in Tasmania. This bird, a
member of the family Muscicapidae (primitive insect eaters), subfamily Malurinae
(Australian Warblers), makes a large globular nest, with a side entrance, in thick
bush, from a few inches to nine feet or higher above the ground. Turdus poliocephalus
Kelleri, which according to Mayr and Amadon (1951) also belongs to the Muscicapidae
(subfamily Turdinae), makes a typically thrush-like open nest; this subspecies is
confined to Mindanao. It is of course by no means certain that the scrub-tit is the
only host of H. vandiemeni. Moreover, it is very probable that representatives of
Hoogstraalia will be found in birds’ nests in other parts of Australia (perhaps only
at particular altitudes) and in the islands to the north, up to the Philippines.

Traub mentions in the description of the genus Hoogstraalia, and also in that of
the species H. turdella, that the movable process bears spiniform setae, and in the
holotype of H. vandiemeni it would be extremely easy to think that this was their
position, since in this specimen the movable process and the distal arm of sternum IX
overlap just as in the holotype of H. turdella. But the fortunate chance that in the
paratype of H. vandiemeni the distal arm of sternum IX is slightly further removed
from the movable process has enabled me to ascertain that actually these spiniforms
are not on the movable process but on the distal arm of sternum IX. The resemblance
in the male terminalia of the two species is so close as to make it certain that the
three long spiniforms in H. turdella must also be placed on sternum IX, just proximad
of the shorter spiniforms on this structure.

To the generic description of Hoogstraalia should be added that the tentorial rod
is fully developed and that the female also has two antesensilial setae; the statement
about the shortness of the setae of the antennal pedicel should be omitted as it does
not apply to H. vandiemeni.

In spite of considerable morphological modifications which doubtless arose in
response to the nature of the host-associations of these fleas, the genus Hoogstraalia
shows clearly affinities with Acanthopsylla, a genus of fleas known only from Australia
and New Guinea.

; References.
Mayr, E., and Amapon, D., 1951.—A classification of recent birds. Amer. Mus. Novit., (1496):
1-42.
TRAUB, R., 1951.—Hoogstraalia turdella, a new genus and species of flea from the Philippines.
Proc. ent. Soc. Wash., 53: 97-104.

181

WIDESPREAD NATURAL INFECTION OF BARBERRY BY PUCCINIA GRAMINIS
- IN TASMANIA.

By I. A. Watson, The University of Sydney, and N. H. Luic, Department of
Agriculture of N.S.W., Sydney.

(One Text-figure.)
[Read 28th May, 1958.]

Synopsis.

Berberis vulgaris was found to be widespread throughout Tasmania and in the summer
of 1956-57 showed a high incidence of infection by P. graminis. Most of the rust recovered
resembled P. graminis secalis but P. graminis tritici was also recovered. The widespread
occurrence of Agropyron repens and its marked susceptibility to the variety secalis is con-
sidered to influence the prevalence of this latter throughout the island.

INTRODUCTION.

It has generally been accepted that the alternate host of Puccinia graminis Pers.
is unimportant in Australia as a source of inoculum each spring and as a medium
concerned in the production of new pathogenic strains of this organism. From time
to time changes have occurred in the prevalence and distribution of the strains of
Puccinia graminis tritici EB. and H. attacking wheat, but these have been attributed to
mutation in the first instance followed by a screening process resulting from the
widespread cultivation of wheat varieties having specific genes for resistance (Watson
and Waterhouse, 1949; Watson and Singh, 1952; Waterhouse, 1952). Barberries are
known to occur, however, and natural infection of them by P. graminis tritici has
been recorded on one occasion (Waterhouse, 1934). Workers concerned with investiga-
tions into the cereal rust problem in New South Wales have had several bushes on the
Central Tablelands under observation since 1933, and until 1957 no natural infection
had been observed on these bushes.

EVIDENCE FOR NATURAL INFECTION IN TASMANIA.

During 1956, under the terms of the Colombo Plan, Dr. R. Prasada, of India, was
engaged in research at the Faculty of Agriculture, The University of Sydney. In
December of that year he visited Hobart, Tasmania, and found within the heart of
that city a horticultural specimen of Berberis bearing the pycnidial stage of a rust
closely resembling Puccinia graminis. Following an examination of this material one
of us (f.A.W.) corresponded with Dr. G. C. Wade to ascertain the distribution of
barberries in Tasmania and the likelihood of their infection by P. graminis.
Subsequently more infected material bearing old aecidiospores typical of P. graminis
was forwarded for examination and Dr. Wade stated that barberries were probably
very common in certain areas of Tasmania.

During the latter part of February and early March, 1957, the senior author made
an extensive trip through Tasmania to survey the distribution of barberries and to
determine the extent to which they were infected. He was accompanied by Mr. Ian
Geard, of the Tasmanian Department of Agriculture, whose detailed knowledge of
the island was a big factor in the successful location of many bushes.

The map shown in Figure 1 makes clear the widespread distribution of B. vulgaris.
From the size of the bushes it is apparent that they have been established for
many years and in view of the association between large bushes and old buildings
it is likely that many of the former were planted more than fifty years ago. It is
certain that they are more abundant in Tasmania than in any other part of Australia
known to the authors. As shown in the map, a high percentage of the bushes examined
were infected.

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SouTH WALES, 1958, Vol. Ixxxiii, Part 2.

182 NATURAL INFECTION OF BARBERRY BY PUCCINIA GRAMINIS,

ASSOCIATION OF UreEDIAL Hosts With THE BARBERRY.

Throughout Tasmania Agropyron repens (Twitch grass) is widespread as a weed
both of cultivation and pasture land. Agropyron scabrum also occurs but less
abundantly and mainly on pasture land. Dactylis glomerata, Hordeum leporinum and
Lolium sp. are common in the more favoured areas. Rusted plants of four of those
grasses mentioned were found in association with infected barberries, the latter were
producing aecidiospores, and heavy uredial infection occurred on the grasses. Rye
grass (Lolium sp.), although found with barberry on many occasions, was not infected
by rust. However, in Launceston, a patch of rusted Rye grass was found well removed
from barberry. The organism responsible was P. graminis lolii.

Agropyron repens is the species that contributes most teleutospore material for
spring infection of barberries. This is inevitable since many of the bushes are to be
found in home gardens where, under proper care, they make attractive specimens.

A Infected bushes
_A Uninfected bushes.
Text-fig. 1.—Map of Tasmania showing the distrinution of Barberries found on the survey.

Should the gardens become neglected, however, even for short periods, A. repens
develops rapidly and spreads by a rhizomatous type of growth. In moist seasons,
such as 1955-56, the host plants are well placed for reciprocal infection to occur
when the rusted grass is growing up into the barberry. Despite the dry summer of
1956-57 evidence for a complete cycle of events was obtained. At Oatlands and Middle
Park old teleutospores formed during the 1955-56 season were found on Hordeum
leporinum, Agropyron scabrum and A. repens. The infected barberries growing in
association were producing aecidiospores resulting from the initial infection in the
spring of 1956. Uredospores on Hordeum leporinum and A. repens probably resulting
in part at least from the aecidiospore infections and teleutospores on this same host,
indicated that the cycle had been completed for another year.

GLASSHOUSE TESTS WITH AECIDIAL AND UREDIAL MATERIAL.

In view of the close association between the uredial and aecidial hosts throughout
the area surveyed, an attempt was made to relate the strains isolated from both hosts
at any one collecting point. This was not entirely successful, as many aecidial
collections proved non-viable. In Table 1 is shown a classified list of the localities
where barberries were found; it indicates whether infection was observed, and also
shows the incidence of rust,on the uredial host growing in association.

Since the variety of P. graminis responsible for the barberry infections was not
known, spores from the field collections of both aecidial and uredial material were
placed onto seedlings of wheat, oats, barley, rye and Lolium perenne. Two aecidial

BY I. A. WATSON AND N. H. LUIG. 183

and twelve uredial cultures were established on Black Winter Rye and on an
unnamed barley variety B125. This latter is used extensively for glasshouse tests,
since it is susceptible to both P. graminis tritici and P. graminis secalis HE. and H. and
reacts with a 2+ to 2: type of infection. Algerian oats were resistant to all 14
collections but susceptible to those taken on D. glomerata which in each case were of
P. graminis avenae HE. and H. Federation was resistant to all 14 collections with a type
2= and 2 reaction.

TABLE 1.

Location of Barberries in Tasmania and Evidence for Infection on Them and Associated Grasses.

Infection on Infection on
Locality. a Locality.

Barberry. Grass. Barberry. Grass.
Hobart (Botanic Gardens) + +1 Evandale a ne + —
Huonville (Longley Grove) + see Perth bs Aa ae ae +3
Kermandie : + — Perth e ae sE spose
Castle Forbes Bay =F — Longford .. rs Se + =
Franklin... Ss ans SE SF =F" Longford House ... : + -
Ranelagh = = Hagley .. 3a ae => spar oo =
Bagdad 3 pina Westbury + se 35°
North Bagdad - + — East Deloraine Se —
Kempton + - Deloraine Sis + =e
Oatlands ++ se abo & Penstock Lagoon + ia
Middle Park Be 1,4 Ouse = sb apts ¥
Tunbridge = _ Hayes + +1, 3
Campbell Town SF arta New Norfolk + +1
Youngtown + ai

+, light infection; ++, heavy infection; —, no infection.

1 Agropyron repens; * Dactylis glomerata: * Hordeum leporinum; * Agropyron scabrum.

The infection types on the cereals and grasses studied indicated that the various
collections were mostly of a rust that agreed closely with P. graminis secalis. It was
apparent, however, when a side-by-side comparison of the resulting cultures was made
on a number of selected wheat varieties, that at least four strains of the fungus could
be distinguished. Little Club was the variety most useful for the differentiation of
the strains as shown in Table 2. No attempt was made to compare these reactions

TABLE 2.

Reaction of Certain Cereals and Grasses to Four Strains of P. graminis secalis.®

Strain.
1 2 3 4
Wheat, Little Club .. ie ty) i2=,2= ) 2 24,3
Oats, Algerian Bt is 0 0 0 0
Barley, B125 .. a ne 2t Qt Bat 2t
Rye, Black Winter .. mee 3+ 3+ 3+ 3+
Rye grass, Z. perenne He 0 0 0 0

5 Since the completion of this paper collections of rust from infected barberries from
Tasmania have yielded strains of P. graminins tritict.

with those of overseas collections and the culture of P. graminis secalis isolated
previously in Australia (Waterhouse, 1957) was not available for comparison. Numbers
assigned to the strains have no relationship to similar numbers given to overseas
strains.

Table 3 shows the distribution of the strains throughout the island and it is
apparent that strains 2 and 3 are the most prevalent.

184 NATURAL INFECTION OF BARBERRY BY PUCCINIA GRAMINIS,

All of those listed in Table 2 were collected on grasses. The aecidial cultures
both proved to be strain 2 and came from “Quamby Plains’, Hagley and from Penshock
Lagoon. Uredial material collected at each of these two places failed to germinate.

Representative cultures of three of the four strains were used to inoculate more
than 40 wheat varieties but none proved as susceptible as Little Club. Strain 4,
which is the most virulent on Little Club, also gave high reactions on the average on
the other varieties but they could not be used for differentiation purposes.

Two of the collections made from A. repens proved to be mixtures of P. graminis
tritici and P. graminis secalis. Hach was taken in a home garden at Oatlands, where
barberries were heavily infected. The mixtures proved to be of P. graminis secdalis
strain 3 and P. graminis tritici strain 21Anz-1. These were the only occasions on which
a strain of wheat stem rust was found in association with barberry. 21Anz-1 was also
isolated from a commercial wheat field well removed from the alternate host at the
time of making this survey.

TABLE 3.

Distribution of Four Strains of P. graminis secalis in Tasmania.

Strain. Localities where Found.
1 Middle Park.
2 Hobart (Botanic Gardens), Campbell Town, Perth, New Norfolk, Ouse.
3 Hobart (Botanic Gardens), Huonville (Longley Grove), Oatlands,
Westbury.
4 Hagley (St. Mary’s Manse).

SIGNIFICANCE OF THE BARBERRY INFECTIONS.

The significance of susceptible barberries in promoting a diversity of pathogenic
strains of P. graminis is so well known that no further emphasis is needed. In
Australia the presence of relatively few strains of P. graminis tritici has been
attributed to the sparse distribution of susceptible species of barberry (Waterhouse,
1952). While this is still true for the wheat belt, the present results show that in
Tasmania susceptible barberries are widespread throughout the island and under
certain circumstances infection by P. graminis is very common. The climate on the
island is particularly favourable for infection. Teleutospores are produced under
relatively cool conditions and, unlike those produced in the mainland wheat belt, they
can retain their viability through the winter for a spring infection. On A. repens
teleutospores are produced right through the summer and those formed in late autumn
have not been exposed to high temperatures.

Certain areas of the mainland provide a similar environment, particularly the
tableland districts of New South Wales and the more elevated parts of southern
Victoria. However, susceptible perennial grasses comparable with A. repens are not
common in these areas and the teleutospores are seldom brought into close proximity
with susceptible bushes, although A. scabruwm was associated with barberries at
Yetholme in 1933 (Waterhouse, 1934). The most likely association would be between
Lolium perenne and the alternate host. In March, 1957, infected barberries were
found in New South Wales for the first time since 1933. These occurred at Meadow
Flat. Heavily rusted plants of Lolium perenne growing below were carrying both
uredospores and teleutospores of P. graminis lolii. This same variety probably caused
the infection on barberry but the aecidiospores failed to germinate.

Varieties other than P. graminis tritici have been mainly recovered in the col-
lections reported herein and this fact would appear to lessen the menace of barberries
in Tasmania. Waterhouse (1957) has already reported the presence of P. graminis
secalis on the mainland and it has not become important. Rye is not grown to any
extent in Australia and barley usually matures before the temperatures are high
enough for stem rust to become established. A. repens is so widespread on the island
and so susceptible to P. graminis secalis that it is inevitable that this latter should
predominate in this area. But A. scabrum and Hordeum leporinum are also commonly

BY I. A. WATSON AND N. H. LUIG. 185

associated with barberry and it is well known that these species are both susceptible
to the variety tritici (Waterhouse, 1929, 1952). With such congenial hosts available
there is always a possibility that infections with this variety will occur. It was, in
fact, recovered from grasses in two of the collections made and each proved to be
21Anz-1. Although little wheat is grown in Tasmania, strains from barberry with
new gene combinations and showing high survival ability would readily become
established. There is circumstantial evidence for spore movement across Bass Strait
so that once a strain predominated in Tasmania it would soon be carried to the
mainland.

There is also some evidence to suggest that in recent years barberries have become
significant in changing the dominant rust flora in eastern Australia. The isolation
of strain 21Anz-1 in 1954 from the south-eastern areas of the country and its rapid rise
to become the dominant strain in southern New South Wales, Victoria and Tasmania
in 1955 (Watson, 1955, 1957a) suggests that it first originated in the south. As it
differs so markedly from the prevalent strains that occurred between 1928 and 1953
it is unlikely that it arose by mutation. Apart from its ability to attack Celebration,
a variety not widely grown in the area, it shows no genes for virulence not present
in strains prevailing earlier. The phenomenal rate of spread suggests a manifestation
of heterosis resulting from a hybridization precess which has brought together several
dominant factors associated with the high survival ability shown by this strain. In
21Anz-1 certain of these have been for virulence and one at least for avirulence.
More detail will be given to this matter in another publication now in preparation.

The recovery of strain 21Anz-1 from two of the collections raised the question
of biotypes of Agropyron repens. tt is possible that certain clones are susceptible to
the variety tritici as well as to the variety secalis. Repeated inoculations with several
strains of the variety tritici revealed no susceptibility to any of them, but all were
susceptible to P. graminis secalis. The occurrence of plants of this species or any other
species with the double susceptibility to the variety secalis and the variety tritici
would present further possibilities for genetic recombination of virulence genes to
occur. While it has not yet. been demonstrated that dicaryons of P. graminis secalis
and P. graminis tritici will undergo somatic hybridization that possibility suggests
itself from the work of Watson (1957b). In these studies an avirulent strain of the
variety tritici, not unlike P. graminis secalis, hybridized readily on wheat seedlings
with a virulent strain of the variety tritici to give strains much more virulent than
either parent. Further experiments are being carried out on this matter and will
be published elsewhere, but already they emphasize that the degree of virulence that
can result from crossing relatively avirulent and genetically unknown dicaryons on
wheat plants cannot be predicted.

Whatever the mede of variability of P. graminis, barberries will always continue
to be a menace to cereal crops while they provide the efficient means to assemble and
reassemble particular gene combinations which are then free to mutate or undergo
somatic hybridization to even greater virulence. From this point of view the significance
of the infections in Tasmania should not be underestimated.

Acknowledgements.

The authors are pleased to acknowledge the help given at all times by their
colleague Dr. E. P. Baker. Financial help from the University Research Grant and
the Rural Credits Department of the Commonwealth Bank is also gratefully
acknowledged. Thanks are due to the members of the staff of the Tasmanian Depart-
ment of Agriculture, especially to Mr. Ian Geard, who traversed much of the barberry
area, and to Dr. G. C. Wade and Mr. A. H. Woodforde, who made transport available
for collecting.

References.

WATERHOUSE, W. L., 1929.—Australian Rust Studies. i. Proc. Linn. Soc. N.S.W., 54: 615-680.
, 1934.—Australian Rust Studies. iv. Proc. Linn. Soc. N.S.W., 59: 16-18.

, 1952.—Australian Rust Studies. ix. Proc. LINN. Soc. N.S.W., 77: 209-258.

, 1957.—Australian Rust Studies. xv. Proc. LINN. Soc. N.S.W., 82: 145-146.

186 NATURAL INFECTION OF BARBERRY BY PUCCINIA GRAMINIS.

Watson, I. A., 1955.—The Occurrence of Three New Wheat Stem Rusts in Australia. Proc.

LINN. Soc. N.S.W., 80: 186-190.

, 1957a.—Rust Investigations. Conf. of Cereal Breeders and Geneticists, Longerenong
Agric. College, Victoria. August, 1956. C.S.1.R.O., Melbourne.

, 1957b.—Further studies on the production of new races from mixtures of races of
Puccinia graminis var. tritici on wheat seedlings. Phytopath., 47: 510-512.

, and SINGH, D., 1952.—The future for rust resistant wheat in Australia. Journ.
Aust. Inst. Agric. Sci., 18: 190-197.

, and WATERHOUSE, W. L., 1949.—Australian Rust Studies. vii. Some recent observa-
tions on wheat stem rust in Australia. Proc. LINN. Soc. N.S.W., 74: 113-131, 2 pl.

187

SYSTEMATIC NOTES ON SOME HASTERN AUSTRALIAN MEMBERS
OF THE PAPILIONACEHAE.

By Joy THompson, National Herbarium, Sydney.
[Read 30th July, 1958.]

Synopsis.

A species of Oxylobium and a species of Pultenaea are described as new. A new name
is provided for a species of Pultenaea. Four new combinations are made, two for species of
Mirbelia, one for a species of Pultenaea and one for a variety of Dillwynia.

INTRODUCTION.
In the course of systematic studies on the Papilionaceae carried out at the
National Herbarium it has become evident that some taxa require naming or renaming.
The new names are now presented.

OxYLOBIuM Andar.
OXYLOBIUM ROBUSTUM, Sp. nov.

Frutex saepe 3 m. altus, caulibus pubescentibus. Folia plus minus opposita vel
verticillata, angusto-lanceolate vel linearia, apice pungente, saepe 7 cm. longa coriacea,
supra pilosa vel glabra, rarius reticulata et scabrida, infra laxe pubescentia vel
tomentosa, marginibus recurvibus. Stipulae 0. Flores plerumque 1 cm. longi, in racemos
breves terminales vel axillares dispositi. Bracteae et bracteolae deciduae. Calyx
sericeus, lobis acuminatis quam tubo longioribus. Ovarium sessile, ovules 10 vel
pluribus. Legumen plerumque 1 cm. longum, breviter stipitatum, brevi-oblongum,
valide compressum a latere, ad suturam contractum, oblique rostratum, pubescens,
semina estrophiolata.

A shrub, often 3 m. high, the stems pubescent. Leaves irregularly opposite or in
irregular whorls of three, narrow-lanceolate to linear, pungent-pointed, frequently 7 cm.
long, coriaceous, the upper surface pilose or glabrous, occasionally reticulate and
covered with small tubercles, the lower loosely pubescent or tomentose, the margins
recurved. Stipules absent. Flowers usually 1 em. long, in short terminal or axillary
racemes. Bracts and bracteoles deciduous. Calyx silky-pubescent, the lobes acuminate,
longer than the tube. Ovary sessile, the ovules 10 or more. Pod usually 1 cm. long,
shortly stipitate, short-oblong, strongly laterally compressed, contracted at the suture,
obliquely beaked, pubescent, the seeds not strophiolate. ;

Holotype: Byron Bay, Bauerlen 8.1896 (NSW 31556), in National Herbarium,
Sydney.

Distribution: QUEENSLAND: Scortechini (NSW 31548); Noosa R., Staer 9.1911
(NSW 31547); Currumbin Creek, White 9.1912 (NSW 31546); Mt. Lindsay, White
10.1921 (NSW 31549). New SourH Wates: Byron Bay, Rupp 9.1917 (NSW 31554);
Three Mile Scrub, near Byron Bay, Forsyth 11.1898 (NSW 31550, NSW 31557); Lismore,
Rothwell 10.1906 (NSW 31560); ? Alstonville, Tomlins, about 1903 (NSW 31552);
Richmond R., Betche 9.1884 (NSW 31553); Hat Head, Constable 1.1953 (NSW 31559);
_Crescent Head, Davis 10.1941 (NSW 31555); Port Macquarie, Boorman 11.1915 (NSW
31558); Kendall, Bailey 9.1932 (NSW 31551); North Haven, Sless 10.1956 (NSW 39517).

Bentham, Fl. Austr., II (1864), 16, Moore & Betche, Handb. Fl. N.S.W. (1893), 129,
and Bailey, Queensland Flora (1900), 336, include this species in O. ellipticum (Labill.)
R.Br. and/or O. ellipticum (ULabill.) R.Br. var. angustifolium Benth.

PROCEEDINGS OF THE LINNEAN SocteTy of NEw SoutTH WALES, 1958, Vol. Ixxxiii, Part 2.

188 NOTES ON SOME EASTERN AUSTRALIAN PAPILIONACEAE,

MIRBELIA Sm.
MIRBELIA PLATYLOBIOIDES (DC.), comb. nov.
Synonymy: Chorizema platylobioides DC., Prod., II (1825), 103; Mirbelia grandiflora
Ait. f. ex Hook. in Bot. Mag. (1827), t. 2771.

MIRBELIA BAUERI (Benth.), comb. nov.

Synonymy: Chorizema baueri Benth. in Ann. Mus. Wien, II (1838), 71 (as
Chorozema); Mirbelia jeaniae Blakely in Aust. Nat., X (19388), 132.

I am indebted to Dr. R. Meiville, of the Royal Botanic Gardens, Kew, who
indicated to me that Chorizema baueri appeared to be synonymous with Mirbelia
jeaniae. Dr. Melville dissected a flower from Bentham’s Type specimen and forwarded
a copy of his accurate camera lucida drawing which is now housed in the National
Herbarium, Sydney. A photograph of this specimen, Negative No. Kew 803, in the
possession of the Division of Plant Industry, C.S.I.R.O., Canberra, has been kindly lent
to me by Miss N. T. Burbidge.

PULTENAEA Sm.
PULTENAEA BLAKELYI, nom. nov.
Synonymy: Pultenaea trinervia Blakely in Contrib. N.S.W. Nat. Herb., 1 (1941),
122, non P. trinervis J. M. Black (1923).

PULTENAEA PALUDOSA, Sp. Nov.

Frutex parvus, plerumque quam 40 cm. brevior, caulibus teretibus vel minoribus
subtriquetris, pubescens pilis longis tenuibus, laxe appressis. Folia usque ad 0°8 cm.
longa alternata erecta ovata vel ovata-lanceolata, utrinque glabra vel rare pilis paucis
longis tenuibus, supra plerumque plana, marginibus plerumque planis et subtuberculatis,
apice obtuso vel rare subacuto, nervo primo inconspicuo. Stipulae minutae triangulares
pallido-virides. Flores 0-6 cm. longi in capites parvas densas terminales disposti.
Bracteae 0:3 cm. longae, ovatae vel lanceolatae vel lineares, pubescentes pilis longis
tenuibus, saepe lobis 2 lateralibus parvis. Bracteolae 0:2-0:3 cm. longae lineares
pubescentes, pilis longis tenuibus, ad pedicellum sub base calycis dispositae. Calyx
0-3 cm. longus, pilis longis tenuibus sed minus versus basem vestitus, lobis inferioribus
acuminatis, quam tubo longioribus, superioribus latioribus, breviter connatis. Ovarium
dense pubescens usque ad basem, stylo brevi pubescenti, compresso. Legumen 0-3-0-4
em. longum, turgidum.

A small shrub usually less than 40 em. in height, the branches terete or the
smaller ones somewhat three-sided, pubescent with long, fine, loosely appressed hairs.
Leaves 0-8 cm. or less in length, alternate, erect, ovate to ovate-lanceolate, the upper
surface usually flat, the margins usually flat and slightly tuberculate, the apex obtuse
or rarely somewhat acute, both surfaces glabrous or rarely with a few long fine hairs,
the mid-vein inconspicuous. Stipules minute, triangular, light green. Flowers 0:6 cm.
long in small dense terminal heads. Bracts 0:3 cm. long, ovate to linear, often with
two small lateral lobes, pubescent with long fine hairs. Bracteoles 0:2-0:3 cm. long,
linear, pubescent with long fine hairs, attached to the pedicel below the base of the
calyx. Calyx 0:3 em. long, covered with long fine hairs but less so towards the base
of the tube, the lower lobes acuminate, longer than the tube, the upper broader than
the lower and united higher up. Ovary densely pubescent to the base, the style
short, pubescent, flattened. Pod 0-3—0-4 cm. long, turgid.

Holotype: Swamps between Coogee and Bondi, Betche 16.9.1886 (NSW 38185), in
National Herbarium, Sydney.

Distribution: New South Wares: Myall Lakes, Sydney University Expedition
8.1934 (NSW 38183); Tenilba, Port Stephens, Harp 7.1953 (NSW 38182), 8.1954
(NSW 38180); Iter Australiense, Brown 1802-5 (NSW 38181); Centennial Park,
Chippendale 8.1953 (NSW 38187); Centennial Park, Forsyth 10.1896 (NSW 38186);
Botany, Fletcher 8.1888 (NSW 38189); Botany Bay, Camfield 10.1898 (NSW 38188);
La Perouse, Camfield 10.1898 (NSW 38190); Bulli Pass-Appin road, 5 miles from Bulli
Pass, Rodway 9.1934 (NSW 38184).

This species has been included in P. incurvata A. Cunn. by Australian authors.

BY JOY THOMPSON. 189

PULTENAEA BRUNIOIDES (Meissn.), comb. nov.
Synonymy: Dillwynia brunioides Meissn. in Lehm., Pl. Preiss., I (1844), 62.

DILLWYNIA Sm.
DILLWYNIA RETORTA (Wendl.) Druce var. PHYLICOIDES (A. Cunn.), comb. nov.
Synonymy: D. phylicoides A. Cunn. in Field, N.S.W. (1825), 347; D. ericifolia Sm.
var. phylicoides (A. Cunn.) Benth., Fl. Austr., II (1864), 148.

190

SOMATIC HYBRIDIZATION IN PUCCINIA GRAMINIS VAR. TRITICI.

By I. A. WAtson, The University of Sydney, and N. H. Luie, N.S.W. Department
of Agriculture, Sydney.

[Read 30th July, 1958.]

Synopsis.

Further studies have been made on somatic hybrids in Puccinia graminis var. tritici
resulting from a mixture of spores of two parental strains on screening varieties of wheat.
It is concluded that nuclear exchange alone cannot explain the diversity of strains obtained.
A nuclear fusion process followed by recombination and segregation to form dicaryons is
suggested to explain this happening. :

Somatic hybridization is shown to be important in the field since a prevalent field strain
was isolated from a mixture of spores from two other field strains. Sexual hybridization on
barberry, mutation and somatic hybridization are now to be regarded as three important
causes of variability in P. graminis var. tritici.

It has now been established that strains of Puccinia graminis Pers. var. tritici
(Hriks. and EH. Henn.) may undergo some form of hybridization when grown in
association on wheat seedlings under laboratory conditions (Nelson e¢ al., 1955; Nelson,
1956; Watson, 1957). When selected strains are used and an efficient screening
procedure is available it can be shown that the process is of relatively frequent
occurrence and highly pathogenic strains may appear in the progeny of avirulent
parents (Watson, 1957). The mechanism allowing such hybridization to occur is
not Known. Nelson (1956) conciuded that hybrids were arising by an exchange of
nuclei in the parents and Watson (1957) suggested, as a result of the diversity of
strains recovered from a mixture of two parents, the possibility of a sexual process
similar to parasexualism. Recently in studies with the leaf rust organism Vakili
and Caldwell (1957) have concluded that nuclear exchange cannot explain the multi-
plicity of new strains arising from a mixture of two parental types. The present
work extends what was done previously and demonstrates that the process, which
for the time being is called somatic hybridization, can result in new strains arising
under natural conditions in the field.

Mixtures of Red 111 and Orange NR-2.

Further mixtures have been made from single spore cultures of the strains used
previously by the senior author (Watson, 1957). The progeny from fifteen mixtures
in all have been studied and the techniques have been similar to those adopted in
earlier work. Strains unlike the parents were recovered from fourteen of the fifteen
mixtures.

In Table 1 is presented a summary of the reaction types of the various strains
that have been isolated from the mixtures. They have been tested several times on
critical varieties and no evidence of instability has been found. The diversity of types
is very striking. All the varieties representative of Stakman’s series reacted
differentially except Marquis and Khapli. The type 2 on Little Club is an unusual
reaction to P. graminis var. tritici but is common if P. graminis var. secalis is placed
onto seedlings of this variety. The virulence of this same strain on C.1.12632 is also
unusual and shows clearly that genes for virulence have segregated in the progeny
to give strains with a wider host range than that of the parents.

Two varieties with outstanding sources of resistance to the parents and their
progeny were Marquillo and Khapstein. The virulence on Thatcher shown by strain
NR-7 is difficult to explain at present, especially when a related variety Marquillo
showed such a high resistance. Possibly all the effective genes of Marquillo are not
present in Thatcher.

PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WALES, 1958, Vol. Ixxxiii, Part 2.

BY I. A. WATSON AND N. H. LUIG. 191

TABLE 1.

Reaction Types shown by Fifteen Wheat Varieties when Inoculated with Red 111 and Orange NR-2 and Eleven Strains
Resulting from Crossing them on Wheat Seedlings.

Parents. Progeny.
Variety.

Red Orange E : = i

sate | asl, ee 2 3 4 5 6 7 8 9 LOD tet
Little Club .. 4 4 4 4 4 4 4 4 4 2 4 4 4
Marquis 0; 3+ 3+ 3+ 3+ 3+ 3+ 3+ 3+ 3+ 3+ 3+ 3+
Reliance : 0; 38+ (0) 3+ 0 3+ 0 3+ 0 3+ 3+ 0 3+
Kota as 0; 3 3 3 3 3 3 3 : 3 3 3 3
Spelmar 0; 0; 0; 0; 3+ 3+ Bar || § : 3+ 3-5 3+ 3+
Kubanka 0; x 3-5 3+ Bar 3+ 3+ 3 2 2 x= 2-5 3+
Acme tg 0; 0; 0; 0; 3+ 3+ Sap || 8 3 3 ex 3 3+
Hinkorn ye 0; Bsr 0; 0; 0; 0; 0; 3 0; 0; 0; 0; 0;
Vernal eee 0; ar 3+ 0; x+ 0; x+ 3 0; 0; 0; 0; 0;
Yalta oe 0; 38+ 31 31 31 oTL 38+ 3+ 31 31 3+ 31 34+
Bokveld ere 0; 0; all afl 31 oil 31 31 31 31 3e 31 3c
Thatcher .. 0; 0; 0; 0; 0; 0; 0; xsi 3 x+ | 31 : x+
Marquillo .. 0; 0; : ; 5 F 5 5 5 Bil 5 X-+- 2
Cie 2632) 2. 0; il 31 call 31 il iL xa 1+ 3 a xx x—
Khapstein .. 0; sll se Silae || gil Sil sr 31+ BIL se il pL Se 31+ gi 45 ji+ 31
Strain designa-

tion ns Race NR-2 | NR-4| Race | NR-3| Race | NR-5| NR-7| Race | NR-6} Race | NR-8 | NR-9
111 196 34 54 222

In Table 2 are given the frequencies with which certain of the strains were
recovered from various mixtures. Figures were not obtained from mixtures 1 and 2
and no red pustules were found on Marquis in mixture number 6. ‘Two strains,
race 34 and the NR-6, have tended to predominate in the progeny of the mixtures as a
whole and the frequency of recovery of any one strain is most probably related to

TABLE 2.
Frequency of Isolation of Eleven Strains as Progeny of the Cross Red 111 x Orange NR—2 when Different Screening Varieties
are Used.
Mixture
Number. Sereening Varieties. Strains Recovered and Frequency.
1'* Standard differentials. Race 34, NR-3, NR-4.
2* ss on NR-3, Race 196.
3 is 5 Races 34 (3), 196 (6).
4 =3 ig NR-5 (1).
5 Emmer. NR-6 (1), NR-7 (38).
6 Marquis. Nil.
7 % and Emmer. Race 34 (1), NR-6 (3), Race 222 (2), Race 54 (2), NR-8 (2).
8 a AD as Race 34 (1), NR-6 (13), Race 222 (2), Race 54 (2).
9 45 », Kota. Race 34 (1), NR-6 (9), Race 222 (1), Race 54 (2).
10 5 35 be Race 34 (1).
11 3 OR, Race 34 (7), NR-6 (2).
12 58 56) 55 Race 34 (3), NR-6 (1), NR-7 (2), Race 54 (4), NR-9 (1),
NR-8 (3).

13 Reliance, Marquis and Kota. NR-6 (2).
14 as 3 nA mii Race 34 (2), NR-G (4).
15 % Ad 50 3 Race 34 (2), NR-6 (6).

* The frequency of recovery of the strains from these mixtures was not recorded.

192 SOMATIC HYBRIDIZATION IN PUCCINIA GRAMINIS VAR. TRITICI,

the variety used in screening. The table shows the varieties that were used in this
latter process and, since these were so few in number, it is unlikely that a random
sample of progeny was recovered from any one mixture. In mixture number 5, for
example, where the isolations were made from Vernal Emmer, three of the four red
pustules found were virulent on Hmmer. To obtain data on the relative frequency
of occurrence of various strains in the progeny the technique has been varied in
experiments now being undertaken.

The data from Table 1 clearly confirm and expand the previous results (Watson,
1957) in which two cultures avirulent on Arnautka, Mindum, Spelmar and Acme gave
rise to progeny to which these same varieties were susceptible. Since ability to attack
these varieties has been found to behave as a dominant factor (Johnson, 1954) this
would not be expected. The possibility of genes for pathogenicity other than those
reported by Johnson, being asscciated with the reactions on these varieties, must not
be overlooked. Their presence in this material is suggested from the complexity of
strains resulting from the selfing of red 111 on barberry.

Progeny of Red 111.

Johnson (1954) usually found that when strains avirulent on Arnautka, Mindum
and Spelmar were taken through barberries the progeny were also unable to attack
these varieties. While there were certain exceptions, it appeared that avirulence was
controlled by a recessive gene. Teleutospores of red 111 were produced at St. Paul,

TABLE 3.

Reaction of Six Varieties of Wheat to Red Race 111 and Seven Selected Strains appearing among the Progeny obtained
from Selfing.

Red 111 Selected Progeny Strains.

Parent.
Reliance 0; 4 3 4 0 4 4 4
Arnautka 0: 3 3 x 0) 3 0 4
Mindum 0; 3 1 1 0 x 0 1
Spelmar 0; 1 1 3 0 3 3 0
Acme 0; ; 3 1 3 1 3 0
HKinkorn 0; ; ‘ 1 3 : 3 0

* These strains were separated and the reactions taken by Messrs. Wilcoxson and Paharia and their help is grate-

fully acknowledged.

Minnesota, U.S.A., and used by the senior author to infect barberries. Among the ~
resulting progeny were several strains virulent on one or other of the three above
varieties and Acme (Table 3). From this table it is apparent that the parental
strain red 111 is carrying genes for virulence on these varieties of Triticum durum.
The effects of such genes only become manifest with segregation and recombination
in selfed progeny.

It is suggested that some process, still unexplained, has resulted in the segregation
of these same genes following the somatic hybridization that has taken place in the
experiments reported herein. The segregation of these genes has resulted in the
isolation of strains to which the varieties of JT. durum are susceptible. The occurrence
of unexpected strains among the progeny of red 111 could be due to genes for
virulence associated with the varieties of 7. durum being unlike those previously
reported by Johnson. Since red 111 has been recovered as progeny of the intervarietal
cross P. graminis var. tritici x P. graminis var. secalis (Johnson, 1949) it is possible
that the genes for avirulence of the latter on the 7. durwm varieties have become
associated with the genes for virulence on these same varieties from P. graminis var.
tritici in this strain. While these latter genes normally behave as dominants, it may
be that they are hypostatic to the genes from P. graminis var. secalis and strains
having both are avirulent on varieties such as Mindum. When segregation occurs
the presence of genes for virulence on the 7. dwrwm varieties again becomes evident.

BY I. A. WATSON AND N. H. LUIG. 193

The data of Table 3 show further that virulence on Arnautka, Mindum, Spelmar
and Acme is not controlled by the same gene or group of genes, even though the
reactions of the first three are usually the same to any one strain. The data also
show that the gene for avirulence on Hinkorn and Reliance are each in a heterozygous
condition. Such heterozygosity would explain how segregation in the somatic hybrids
between red 111 and NR-2 could result in strains virulent and avirulent on both these
varieties. It is similarly postulated from the mixing studies that the genes for
virulence on Vernal Emmer are also in a heterozygous condition. However, on selfing
red 111 no cultures virulent on this variety were recovered. Since such virulence is
controlled by two genes (Johnson and Newton, 1940), failure to find Vernal Emmer
susceptible may have been due to the small number of isolations made. In general the
results obtained from selfing red 111 complement and confirm those obtained from
crossing red 111 and NR-2 on wheat seedlings. They reveal the genetic constitution of
strain red 111 which would theoretically make. possible certain strains recovered as
somatic hybrids when this race is crossed with NR-2.

The Importance of Somatic Hybrids in the Field.

The cultures red 111 and orange NR-2 have been selected for studies of somatic
hybridization since they are well marked genetically and since their progeny can be
so readily isolated. Both are essentially laboratory cultures of little practical
importance. The results with them have illustrated important principles but they
have not demonstrated the significance of the process under natural conditions in
the field. :

Strains of rust prevalent in the field usually do not have the appropriate
characteristics for studies of this kind, and this fact probably accounts for the early
failure of attempts to demonstrate hybridization between dicaryons. The heterozygosity
of genes for virulence, the colour of the spores and the varieties suitable for screening
are all important factors which, if possible, should be considered in the selection. of
material for this type of experiment.

A search has been made to find strains from New South Wales which would be
suitable for this purpose and experiments have been carried out using two of them.
One was an orange biotype of strain 21 Anz-1 and the other a normal red biotype
of strain 222 Anz-2. Red 21 Anzl is now the most prevalent strain in the south-
eastern part of Australia (Watson, 1957a). Strain 222 Anz-2 occurs in the northern
part of the eastern wheat belt and has been prevalent in northern New South Wales
and southern Queensland where varieties having stem rust resistance similar to that
of Gabo have been grown.

Inoculum from single spore cultures of orange 21 Anz-1* and of red 222 Anz-2 was
increased separately on Little Club. It was then mixed mechanically and used to
inoculate the variety Acme on which orange 21 Anz-1 is virulent and red 222 Anz-2
is avirulent. Red pustules occurring on Acme were then isolated and examined. The
whole experiment was carried out in strict isolation and there was no evidence of
contamination from outside.

In Table 4 are given the reactions on five wheat varieties of six strains that
were isolated and it is evident that certain of them combine the characters of both
parents although none of the progeny was orange. Strains A, C and E were virulent
on C.1.12632, an unusual feature to be found in local strains, and strain F combined
in the one strain virulence on Celebration and on Yalta. B and D were unlike either
parent and the whole progeny of six new strains suggests that somatic hybridization
has occurred between orange 21-1 and red 222-2. On Stakman’s differential series,
strain F conforms most closely to race 21, and since it resembles strain 21 Anz-1
except that it has the additional ability to attack Yalta, it has been designated
21 Anz-2. Its appearance among the segregates was of considerable significance.

As reported earlier, red strain 21 Anz-1 was first recorded in 1954. It spread
rapidly in the south, and although it was common in the north of New South Wales

* Our colleague Dr. E. P. Baker first isolated this strain in 1955 from Griffith in New
South Wales.

H

194 SOMATIC HYBRIDIZATION IN PUCCINIA GRAMINIS VAR. TRITICT,

the rate of increase was retarded by the extensive cultivation of Gabo and varieties
with a similar type of resistance to it. Crops of the susceptible variety Bencubbin
were in sufficient quantity in the area to enable it to build up, and this variety
from time to time during 1955 and 1956 was found simultaneously infected by 21 Anz-1
and one or other of the prevalent strains in the area 126 Anz-3 and 222 Anz-2.
Facilities were available on a field scale to enable hybridization between strains
21 Anz-1 and 222 Anz-2 to occur.

Late in 1956, strain 21 Anz-2 was first recorded from Woodburn in northern
New South Wales from material sent in by Mr. H. Farrelly on Federation wheat.
Since it attacks Yalta, the varieties Gabo, Koda and Charter which have a similar
resistance and which are widely cultivated in the area are susceptible. This fact as
well as the remarkable survival and competitive ability of strain 21 Anz-2 have
enabled it to become by far the most prevalent strain of P. graminis var. tritici in
northern New South Wales and southern Queensland.

TABLE 4.

Reactions of Five Varieties of Wheat to Two Parental Strains of Rust and Six of their Progeny Resulting from Mixing
them on Wheat Seedlings.

Parents. Progeny.
Orange Red

21-1 299-2 A. B. C. D. E. F.
Little Club eiailhe ord 4 4 4 4 4 4 4
Reliance .. ab 0 3+ 3+ 0) 0 3+ 3+ 0
Acme a2 bi 3+ x= x+ x+ x— 3¢ oe 3e
Yalta ii a 3 3+ 3+ 3 2 3+ 3 3+
C.1. 12632 tid fer ee x x 3c 1 3e 1
Celebration fe 38+ 31 x+ 3+ xi— 3+

While the authors hold the view (Watson and Luig, 1958) that 21 Anz-1 originated
on barberries in Tasmania, the isolation of 21 Anz-2 in the north suggests a separate
origin for this strain. The production of strain 21 Anz-2 in the laboratory by
hybridizing two strains of rust almost identical with those occurring in the field
shows that somatic hybridization in the field between red 21 Anz-1 and red 222 Anz-2
must be considered as a probable mode of origin of this strain.

Discussion and Conclusions.

Somatic hybridization must now be assumed to be an important mode of origin °
of new strains of wheat stem rust. Certain parental strains can be shown to undergo
the process readily so that there is no difficulty in isolating the progeny from repeated
mixtures. . Other strains which are not so well suited for the experiments do not
allow progeny to be isolated with such frequency. At this stage, however, the frequency
of hybridization between different strains cannot be given. It is possible that some
strains will hybridize frequently, others less readily and some seldom or not at all,
as Holloway (1954) has found for strains of Neurospora crassa with regard to asexual
reproduction.

The results from this experiment show that nuclear exchange cannot satisfactorily
explain the origin of new strains among the progeny of red 111 and orange NR-2.
The diversity of types exceeds that possible by this process alone. The authors
conclude from the difference in frequency of the strains isolated in the progeny and
their wide range in pathogenicity that a process of segregation and recombination is
involved. So far no evidence is available to show that nuclear fusion has occurred
between the parents but the hypothesis suggested is that such fusion does take place.
The fusion follows anastomosing between the hyphae of the different parental strains
(Nelson et al., 1955) and the nuclei of opposite sex take part. The fusion nucleus
is short-lived and segregates immediately giving rise to other nuclei with different

BY I. A. WATSON AND N. H. LUIG. 195

gene combinations. By an association of these nuclei, dicaryons are formed having
new combinations of genes for virulence which become evident as new pathogenic
strains. The exact sequence of events cannot be given as Pontecorvo (1956) has done
for the Ascomycetes but so far we have been unable to isolate an unstable strain the
equivalent of the diploid in Aspergillus.

While the exact mechanism making somatic hybridization possible must await
further work, the synthesis of new strains important in the field by mixing inoculum
of prevalent natural strains has great significance. Sexual hybridization and mutation
must now be supplemented by somatic hybridization to make three important causes
of variability in P. graminis var. tritici under natural conditions. A particular strain
of rust, however, need not necessarily arise in one way only. When only a single
step in pathogenicity is involved mutation can account for changes in virulence
(Watson, 19576). Strain 21 Anz-2 could have arisen from 21 Anz-1 by mutation for
pathogenicity on one variety but we have not attempted to produce it in this way.
In 1958, however, a strain conforming with 21 Anz-2 has been isolated from barberries
in Tasmania. As 21 Anz-l1 is the prevalent strain on the island and as a 21 Anz-2 had
not previously been isolated there it appears likely that 21 Anz-2 is a segregate
obtained from selfing 21 Anz-1 on the alternate host. Strains conforming in their
reaction type with a well-known strain such as 21 Anz-2 but having evolved by
different means will ultimately be shown to have some but not all of their genes
in common. Hence minor differences in reaction type will always be found among them.

Acknowledgements.
We wish to acknowledge the help given by our colleague Dr. E. P. Baker. The
help made available from the University Research Grant is also acknowledged.

Referenres.
Houutoway, B. W., 1954.—Genetic control of heterocaryosis in Newrospora crassa. Genetics,
40: 117-129.
JOHNSON, T., 1949.—Intervarietal crosses in Puccinia graminis. Can. J. Research, Sect. C,
27: 45-65. :

, 1954.—Selfing studies with physiologic races of wheat stem rust, Puccinia graminis
var. tritici. Can. J. Botany, 32: 506-522.
, and NewrTon, M., 1940.—Mendelian inheritance of certain pathogenic characters of
Puccinia graminis tritict. Can. J. Research, Sect. C, 18: 599-611.
NELSON, R. R., 1956.—Transmission of factors for urediospore color in Pwecinia graminis var.
tritici by means of nuclear exchange between vegetative hyphae. Phytopath., 46: 538-540.
, WILCOxSON, R. D., and CHRISTENSEN, J. J., 1955.—Heterocaryosis as a basis for
variation in Puccinia graminis var. tritici. Phytopath., 45: 639-643.

PoONTECORVO, G., 1956.—The parasexual cycle in fungi. Ann. Rev. of Microbiol., 10: 393-400.
VAKILI, N. G., and CALDWELL, R. M., 1957.—Recombination of spore color and pathogenicity
between uredial clones of Puccinia recondita f. sp. tritici. Phytopath., 47: 536. (Abs.)
Watson, I. A., 1956.—The occurrence of three new wheat stem rusts in Australia. Proc.

Linn. Soc. N.S.W., 80: 186-790.

, 1957a.—Rust Investigations. Conf. of Cereal Breeders and Geneticists, Longerononge
Agr. College, Victoria, August, 1956. C.S.I.R.O., Melbourne.

, 1957b.—Mutation for pathogenicity in Puccinia graminis var. tritici. Phytopath.,
47: 507-509.

, 1957¢.—Further studies on the production of new races from mixtures of races of
Puccinia graminis var. tritici on wheat Seedlings. Phytopath., 47: 510-512.

, and Luic, N. H., 1958.—Widespread natural infection of Barberry by Puccivia
graminis in Tasmania. Proc. LINN. Soc. N.S.W., 83: 181-186.

196

A NOTE ON THE STATUS OF APHODIUS TASMANIAE HOPE.
By B. B. Given, Entomology Division, D.S.I.R., Nelson, New Zealand.
(Communicated by Mr. C. H. Chadwick.)

[Read 30th July, 1958.]

In a previous paper (Given, 1950) the status of several Australian species of the
genus Aphodius was reviewed. An error in this paper was noted by Mr. C. EH. Chadwick,
of the New South Wales Department of Agriculture, who informed me of his finding
and suggested the necessary correction.

The present correct status of the species concerned is as follows:

APHODIUS TASMANIAE Hope, 1847, Trans. ent. Soc. London., 4 (5): 285.

howitti Hope, 1847, Trans. ent. Soc. Lond., 4 (5): 285. australasiae Blanchard,
1853, Voy. Pole Sud, 4: 101. longitarsis Redtenbacher, 1867, Reise Novara, 2(A): 58.
andersoni Blackburn, 1904, Proc. roy. Soc. Vic., 17 (1): 154.

In my paper of 1950 I was in error in giving priority of name to howittti
(tasmaniae has priority over howitti, being described above it on the page), and in
the reference to Hope’s paper as being Proc. ent. Soc. Lond., 1846: 147.

The above corrections do not alter the status of other species mentioned in my
previous paper. 7

Reference.
GIVEN. 1950.—PrRoc. LINN. Soc. N.S.W., 75 (3-4): 158-157.

a
PROCEEDINGS OF THE LINNEAN Society oF NEw SouTH WALES, 1958, Vol. Ilxxxiii, Part 2.

197

SIR WILLIAM MACLEHAY MEMORIAL LECTURE.
TIMING IN HUMAN EVOLUTION.
By A. A. Appiz, Department of Anatomy, University of Adelaide.
(Seven Figures.)
[Delivered 19th June, 1958.]

—<____——_—.

SIR WILLIAM MACLBEAY.

Mr. President and members of the Council: Your invitation to deliver a lecture im
commemoration of our greatest benefactor tempers any inclination to personal pride
with a diffidence compounded of anxiety to do justice to that wise and far-sighted
man, and humility at the poverty of what I have to offer. I can only say how deeply
I appreciate the honour you have done me.

A first memorial lecture should be a simple recital of the man’s life and
achievements, an appreciative analysis of the benefits that have ensued upon his
generosity and a just picture of him as a man. This apparently straightforward task
is fraught with some difficulties. We know something of his life, and his public
achievements are officially recorded. The fruits so far of his benefactions are apparent
in the flourishing state and reputation of the Society, in the quality of the papers
published in the ProcrEprnes, in the records of the Linnean Macleay Fellows, and
so on. But of Macleay personally we Know only that his great modesty forbade
practically any intrusion of himself into public affairs. What little we know of him
we gain from the opinions of others, and those I have read have invariably been
high ones.

What little is known, indeed, has been avidly seized upon by all previous com-
mentators in an attempt to present a worthy appraisement. Haswell (1891), Fletcher
(18938 and especially 1920 and 1929), Walkom (1925, 1942) and others have gobbled
up every scrap, leaving little for a successor. Lately Macmillan (1957) has expanded a
small morsel into his delightful book on the Chevert expedition but it seems unlikely
that many more such morsels will give themselves up. Consequently, I must tell the
well-known story in my own way, acknowledging that I have Grawn freely on the
authors I have cited. Also, largely because the story is so well known, I feel that
much of the detail can safely be left for reference to their works. My account of
Sir William Macleay is, therefore, relatively brief, and I shall supplement it with a
short discussion of some matters that have interested me during the last few years.

William John Macleay, like any other living organism, cannot properly be assessed
apart from his environment. It was the context of his surroundings, current events.
and personal contacts—especially family ones—that brought out those particular
features that cause us to honour him tonight. Macleay’s context, appropriately enough,
takes us back to Linné himself so perhaps I might start at that point.

Elsewhere (Abbie, 1954) I have discussed briefly the influence of Linné in the
early history of Australian biology. That depended largely upon his association with
Sir Joseph Banks through Solander, Dryander and Fabricius. When Linné died
Banks was offered the entire Linnean collection of books, specimens and cabinets for
one thousand guineas (Smith, 1911). Banks did not purchase but a young Dr. Smith
was persuaded to do so. The collection reached England in 1784 and Smith was moved
to found a Linnean Society, which was established in London in 1788. Banks took
_great personal interest in the new society and defrayed many of its early expenses —
an example followed even more generously a century later by the subject of tonight’s
talk.

PROCEEDINGS OF THE LINNEAN Socipty or NEw SoutH WALES, 1958, Vol. Ixxxiii, Part 2-

198 SIR WILLIAM MACLEAY MEMORIAL LECTURE,

Alexander Macleay.

The story now really begins for, six years after the Linnean Society of London
was founded, the first relevant Macleay entered upon our scene. In 1794 Alexander
Macleay was elected a Fellow of the Linnean Society and in 1798 he became its second
secretary, a post he held until 1825.

Alexander Macleay was born in 1767 in the County of Ross; his father was
Provost of Wick and Deputy-Lieutenant of the County of Caithness. Alexander held
various public offices from 1795 until 1818, when he retired on a pension at the age
of fifty-one. In 1825 the Earl of Bathurst persuaded him to come to Australia as
Colonial Secretary, which position he filled until 1836. Alexander was elected the
first Speaker of the (old) Legislative Council in 1843 and carried out his duties with
ability, judgement and impartiality until 1846, when he retired because of his age.
Two years later he died in his eighty-first year.

From his first association with the Linnean Society Alexander had collected
biological material — chiefly insects —in Britain and from other countries by exchange
or purchase. His collection, already famous before he left England, formed the basis
of the Macleay Museum. He does not seem to have published any scientific papers
but was elected a Fellow of the Royal Society in 1809 and a Councillor in 1824. In
Australia (although some material was sent to the Linnean and Zoological Societies
of London) collecting appears to have languished somewhat in favour of horticulture,
in which Alexander gained high regard; but he was an ardent supporter of the
Colonial —later Australian— Museum. He owned property at Brownlow Hill and
Glendarewel Farm in the Camden district, and on a grant of land “near” Sydney
(one and a haif miles from Sydney Town) built Elizabeth Bay House, which was
occupied almost continuously by Macleays from 1837 to 1908.

Wiliam Sharp Macleay.

The eldest son of Alexander was born in London in 1792 and graduated from
Cambridge in 1814. Until 1825 he was in Paris on government service and became
friendly with Cuvier, Lamarck and other notable French biologists. Then he went to
Havana, Cuba, on a mixed English and Spanish Commission for the abolition of
slavery. After occupying various posts there he returned to England in 1836 and
retired on a pension the next year at the age of forty-five. In 1838—in the company
of his cousins William and John —W. S. Macleay left England for ever to settle in
Australia.

W. S. Macleay’s first inspiration to biology probably came from his father and
was stimulated deeply by his contact with the French biologists. However, he does
not appear to have begun collecting seriously on his own account until he went to
Cuba. He published a number of papers, including the Horae Hntomologicae, which
contained philosophical speculations on the “Circular System” and “Quinarianism”
so misapplied by over-enthusiastic friends that they fell into disrepute. When Macleay
returned from Cuba he met Darwin, also newly returned from his famous voyage on
the Beagle, and was one of those who urged Darwin to publish his diary (F. Darwin,
1887). Macleay was elected to the Councils of the Linnean and Zoological Societies of
London and was President of Section D at the 1837 meeting of the British Association
for the Advancement of Science.

On the voyage to Australia with his cousins W. S. Macleay collected marine
biological material assiduously. In Sydney he warmly embraced the new biological
field, especially in ichthyology, and made a wide circle of like-minded friends, of whom,
biologically speaking, the most important were Dr. George Bennett and Thomas Henry
Huxley. Huxley mentions him as “William Macleay” in a letter from Sydney in 1848,
and in 1851 wrote to “W. Macleay” in Sydney on the possibility of a professorship in
natural history at Sydney University (L. Huxley, 1900). This publication refers only
to “Sir William Macleay” in the index but it is obvious from the context and cross
checks that William Sharp Macleay was the person concerned. Curiously, there appears
to be no reference to W. S. Macleay in the Rattlesnake diary (J. Huxley, 1935).

While W. S. Macleay collected widely and made valuable contributions to taxonomy,
his real interest lay with the pre-Darwinian philosophical and systematic side of

BY A. A. ABBIE. 199

biology. A great service to Australia was his strong support of the Australian
Museum. He was a member of the Committee from 1841 to 1853; then he was
elected a Trustee, a responsibility he sustained until 1862, when ill health forced
retirement. He died in 1865 at the age of seventy-three and left most of his possessions
to his brother George, then in England. From our point of view it was most fortunate
that he bequeathed the whole of his own and his father’s natural history collection to
his biologically-minded cousin William John Macleay.

George Macleay.

George plays a less direct part in this story. The third son of Alexander, he
was born in London in 1809 and came to Australia at about the same time as his
father. At first he was mainly occupied in managing his father’s Camden property
with the help of a younger brother James. George also acquired a property of his
own on the lower Murrumbidgee. He went with Sturt’s second expedition down the
Murrumbidgee and Murray Rivers in 1829-30 and distinguished himself on that arduous
venture. So early as 1836 he was a member of “A Committee of Superintendence of the
Australian Museum and Botanical Gardens” and later was elected a Museum Trustee.
George became Member for the Murrumbidgee in the (old) Legislative Council in 1855
(Legislative Assembly in 1856), but he resigned all his appointments when he returned
to England in 1859. There he was elected a Fellow (later Councillor) of the Linnean
Society and was created K.C.M.G. Some time between 1869 and 1874 he made a brief
visit to Australia to help wind up the estate of his brother W. S. Macleay. He married
twice but had no children and died at Mentone in 1891 in his eighty-second year.

George collected for his father on the Murrumbidgee but does not appear to have
been an ardent naturalist. Nevertheless, the interest was there, as his election to the
Linnean Society shows, and his long support of the early Australian Museum merits
much commendation. We would consider that his most valuable contribution lay in
the guidance and encouragement he gave his cousin during the early days of squatting
on the Murrumbidgee, when the future Sir William was founding the fortune that
makes the rest of this story possible.

William John Macleay.

The chief figure in this account was Alexander’s nephew, born at Wick in 1820.
At-about seventeen he began to study medicine at Edinburgh University but the death
of his widowed mother inclined him to heed his uncle’s advice to migrate to
Australia. William and his brother John embarked with W. S. Macleay towards the
end of 1838, reaching Sydney in March, 1839. There was a big family reunion at
Elizabeth Bay House. John was delicate and was advised to take a sea voyage —
quaint advice in view of the fact that he had just completed one of some months’
duration! So he set off back for England but died on the way. Meanwhile, William
took up a property — Kerarbury — on the lower Murrumbidgee, no doubt on the advice
of his uncle and of cousin George, who was already established there. For fifteen
years William experienced the typical life of a squatter, coming to Sydney only as
occasion demanded. We know very little indeed of the Murrumbidgee era apart from
an episode of great personal heroism in the face of bushrangers. Fletcher (1929)
has recounted some details of property transactions at that time but they are too
complex to be discussed here. There is no doubt, however, that William prospered.
In 1855 he was elected Member for Lachlan and Lower Darling in the (old)
Legislative Council. Responsible government was introduced in 1856 and William
represented the same district in the Legislative Assembly. When George Macleay
departed for England in 1859 William became the Member for the Murrumbidgee until
he resigned in 1874. In 1877 he was elected to the (new) Legislative Council and.
held that seat for the rest of his life. He was an independent but public-spirited
parliamentarian, advocating many necessary reforms and serving actively on a number
of committees and commissions. He became a Trustee of the Australian Museum —
thus preserving the family tradition — and a member of the Senate of the University
of Sydney. He was knighted in 1889, two years before he died.

200 SIR WILLIAM MACLEAY MEMORIAL LECTURE,

From 1855 on, William’s parliamentary duties demanded more and more of his
time in Sydney; and on his marriage to Miss Susan Emmeline Deas-Thomson in 1857
he took up permanent residence here —first at Denison House, Phillip Street, and
later at 153 Macquarie Street. Thereafter, his Murrumbidgee property was left to the
care of a manager. When W. S. Macleay died in 1865, William and his wife moved
finally to Elizabeth Bay House.

William’s first interest in biology may have been aroused by his preliminary
medical studies in Edinburgh. At all events, such interest must have been stirred
on the voyage with W. S. Macleay, who pursued biological work en route. While on
the Murrumbidgee William probably undertook some collecting for his uncle but it
seems clear that any biological urge had little opportunity to express itself until he
was permanently settled in Sydney, where, in 1856, he became a member of the
Philosophical Society of New South Wales. In Sydney he had access to the collections
of Alexander and W. 8S. Macleay (which he inherited on the death of the latter)
and began active collecting on his own account. To this end he undertook personal
collecting trips, employed collectors in various parts of Australia and arranged
exchanges and purchases from overseas. (In 1873 the Senate of the University of
Sydney accepted his offer to bequeath the whole combined Macleay collection to the
University.) William was largely instrumental in getting the Entomological Society
of New South Wales started in 1862. He was the first president (W. S. Macleay having
declined because of his health) and contributed a number of papers to the Trans-
actions — which journal attracted the attention of entomologists elsewhere. However,
the entomological ranks in Sydney dwindled and the Society lapsed after eleven
years, the last number of the Transactions appearing in 1873.

Workers in wider biological fields were becoming more numerous, and Macleay’s
own collection and interests had gone far beyond entomology. It seemed that a
society less narrowly confined would be a more suitable forum for naturalists generally.
In 1874 he recorded that “Dr. Alleyne and Captain Stackhouse are trying to get up
a Society of Natural History’. This marked an epoch in William’s life. He resigned
his parliamentary seat and thereafter devoted himself almost entirely to natural
history. Perhaps the current visit of H.M.S. Challenger helped to determine the issue.

There seems to have been a proposal to call the new society “The Banksian
Society’. In view of Banks’s services to biology in general and Australia in particular
that would have been appropriate enough. Nevertheless, the decision to name it “The
Linnean Society’ was undoubtedly correct. lLinné’s system of classification provided
the firm basis for general biological work; the title is more explicit; Banks himself
had been a great supporter of Linné and of the Linnean Society of London; and all
the relevant Macleays were at some time Fellows of that Society. On 28th October,
1874, a preliminary meeting in the board room of the Sydney Public Library decided
to form the Society, fixed the subscription and chose a distinguished list of officers
with William Macleay as president. On 4th November the proposed rules were adopted;
on 13th January, 1875, the office bearers and council were formally elected, and on
25th January the first scientific meeting was held in Lloyd’s Chambers, 362 George
Street.

While all this was in train William had been looking for a vessel to take a
collecting expedition to New Guinea and he purchased the barque Chevert early in
1875. She proved stout enough but was not suitable for conditions on the New Guinea
coast. The story of that expedition has been fully told by Fletcher (1893, 1929) and
by Macmillan (1957) se I shall not delay you with the details. To some extent —
particularly in the failure to penetrate inland—the expedition fell short of expectations.
But, speaking as one who has had to organize more modest expeditions of a different
kind, I can say that few, if any, fulfil all the hopes of the planners. That apart, the
adventure was a signal success. A large quantity of material was collected, biological
interest everywhere was excited and it was shown that Australia could manage an
undertaking of that sort quite independently of outside help. Naturally, this enterprise
proved a great stimulus to the new Society, to which we must return.

BY A. A. ABBIE. 201

The Linnean Society’s changes of fortune and residence have been amply described
by Fletcher and Walkom, and by others incidentally. All this I need not go over
again. There are, however, some points that stand out and should be emphasized.
One is the unwavering generosity of William Macleay in furnishing from his own
pocket accommodation, secretarial assistance, help towards publication costs and liberal
support in the provision of an adequate reference library. And few societies have
enjoyed such munificence as almost complete replacement of the library after the
disastrous fire at the Garden Palace, or such a gift as the Linnean Hall at Elizabeth
Bay, or such a final bequest as seems to ensure the stability of the Society far into
the foreseeable future. In all these respects I feel that William Macleay surpassed even
his exemplar Sir Joseph Banks in generosity.

Yet Macleay was no mere Mycaenas. He gave freely to support the science he loved
but he was also an active worker in and for that science, whether in the field and
laboratory or at the secretary’s desk. In all this I think that we can see more than
ordinary generosity: it was, rather, an unwavering determination—jin the face of
setbacks that would have frightened lesser men—that the Society should survive at
all costs. But for this, biology in Sydney would have fared badly, for the University
had no medical school and, consequently, no school of biology. Macleay, with his
far-sighted vision of the importance of biology to Australia’s economy, supported what
was virtually an extra-mural school until the University revaired its omission. However
that may be, there is no doubt that through all these vicissitudes he ensured that
workers could continue with their research and publish it without hindrance. In the
outcome, the Society grew steadily in reputation, as it has continued to grow since,
to its present status of one of the foremost of such societies. To be able to say that
is ample justification for this meeting in Macleay’s honour.

At that point one should stop but there is a little more to be said. Macleay became
a@ member of the Senate of the University and was appointed to a committee on the
setting up of a medical school. This was realized in 1883 and the consequential
Department of Biology proved complementary to the Linnean Society, the two com-
bining splendidly to foster still more vigorously their common interests. Macleay took
the necessary steps to further that cooperation by leaving his museum to the
University and, above all, in establishing the Linnean Macleay Fellowships. One more
example of his amazing prescience should be recorded. He was one of the few to
foresee the potential importance of the newly founded science of bacteriology and he
provided money to support study of that branch of biology too.

When Sir William Macleay died on 7th December, 1891, he left behind the reputation
of a great man and a generous benefactor. But he left considerably more —a living,
growing memorial in the flourishing Linnean Society of New South Wales, and in the
distinguished scientists who owed their first opportunity to the Linnean Macleay
Fellowships.

References.

ABBIF, A. A., 1954.—The history of biology in Australia. Aust. J. Sci., 17: 1-9.

DARWIN, F., 1887.—The Life and Letters of Charles Darwin. I. Murray, London.

FLeTcHEeR, J. J., 1893—The Hon. Sir William Macleay, Kt., F.L.S., M.L.C. The Macleay
Memorial Volume, VII-LI. Linnean Society of New South Wales, Sydney.

, 1920.—The Society’s heritage from the Macleays. Proc. Linn. Soc. N.S.W., 45:

567-635.

, 1929.—The Society’s heritage from the Macleays. Part ii. Proc. LINN. Soc N.S.W.,

54: 185-272 (posthumous, edited by A. B. Walkom).

HASWELL, W. A., 1891.—Presidential Address. Proc. LINN. Soc. N.S.W. (second series),
6: 706-723. (Note: the title page of this volume gives the year as “1891” but the
Address is dated “Wednesday, January 27th, 1892.’’)

Huxiey, J.. 1935—T. H. Husxley’s Diary of the Voyage of H.M.S. Rattlesnake. Chatto and
Windus, London.

Huxiey, L., 1900.—Life and Letters of Thomas Henry Hualey. 1. Macmillan, London.

MACMILLAN, D. S., 1957.—The Squatter went to Sea. Currawong Pub. Co., Sydney.

SmiTH, E., 1911.—The Life of Sir Joseph Banks. Lane, London.

WALKOM, A. B., 1925.—The Linnean Society of New South Wales ... . Historical Notes of
its First Fifty Years (Jubilee Publication). Awstralasian Med. Pub. Co., Glebe.

, 1942.—The background to Sir William Macleay’s endowment of Natural History.
Proc. LINN. Soc. N.S.W., 47: iv-xv.

202 SIR WILLIAM MACLEAY MEMORIAL LECTURE,

After that I hesitate to obtrude myself but I take courage from the fact that the
eminent contributors to the Macleay Memorial Volume of 1893 felt that the best tribute
they could offer was an account of their own work. I must apologize that the subject
of my further discussion is so commonplace —a single animal, forming a single genus
of the Primates and represented by only one species. However, man is part of natural
history and I take refuge in the avowed object of the Society: ‘the cultivation and
study of Natural History in all its branches.”

TIMING IN HUMAN EVOLUTION.
INTRODUCTION.

In the famous tenth edition of his Systema Naturae Linné (1758, p. 20)
courageously included man with his most similar contemporaries in the single order
Primates. This he did on a purely objective study of physical characters.

That alone was a tremendous advance for which Linné is always to be honoured.
But he stopped short at classification, accepting in principle the view that species are
sui generis and have been so for all time. However, increasing knowledge brought
to light forms not so easily pigeon-holed, intermediates that aroused suspicion of
some mutability in the living order. The century that followed the publication of
Linné’s classification saw a great deal of work and speculation on the possibility of
an evolutionary process and exactly within the century Darwin and Wallace supplied
the first convincing evidence in favour of organic evolution. Then busy minds turned
to seek for the underlying mechanism, and so on to the nature of life itself. Another
century has passed, some of the mechanisms are becoming apparent and it is the last
problem that now arouses the greatest interest (Abbie, 1955).

Here it is not my ambition to probe so deeply. Instead, I wish to draw attention,
in a very limited way, to one mechanism — to the element of time (or, rather, timing)
in physical development. This is only a part of the problem but it is interesting in
itself and can, as I hope to show, give an indication of the sort of thing that has
happened — might even afford some indication of what could happen.

In 1916 D’Arey Wentworth Thompson (see Thompson, 1942) extended the
application of mathematical techniques to problems in morphology. His work may be
considered the culmination of a purely physico-mathematical school of thought which
failed to make allowance for such variables as genetics, adaptation, convergence, ete.
Nevertheless, he showed that by manipulation of appropriate Cartesian coordinates
it is possible to derive sundry recognizable variants of form-—even quite bizarre-
looking ones — from well-known basic patterns. These may be considered, in a purely
relative sense, distortions in space. Huxley (1932) followed this up with his work
on relative growth which gains by taking genetics into account and imports time into
the inquiry on such distortions. Medawar (1945) also drew attention to the incidence
of time. The progress made along these lines is seen in such publications as Tempo
and Mode in Evolution (Gaylord Simpson, 1944), Essays on Growth and Form (Le Gros
Clark and Medawar, 1945), Growth (1948) and many others.

It will be appreciated that instances of differential growth are really manifestations
of acceleration or retardation in the development of some limited region in respect to
the remainder of the body. That is, localized distortions in space could equally be
considered localized distortions in time—timing in onset, speed, duration and
cessation of growth. Indeed, it is difficult to see how distortions in space could occur
apart from distortion in timing. However that may be, it is my intention to consider
the effects of timing in development and I shall restrict my remarks chiefly to the
animal with which I am most familiar — man.

Before I start I should make a brief comment on the basis of this paper. In
1926 Bolk pointed out that many features of the human skull represent a retention
into adult life of characters distinctive of foetal stages of development. This he called
“foetalization”. The conception was extended more widely into the animal kingdom
by Garstang (1928), de Beer (1940), Hardy (1954) and others, who introduced the
terms “paedomorphism” and “neoteny” to embrace this general application. The

BY A. A. ABBIE. 203

reverse process was designated ‘“‘gerontomorphism”’. I have used such ideas in
considering problems of the human skull (Abbie, 1947, 1952a) but in the latter paper
I pointed out that not all peculiarly human features can be attributed to paedo-
morphism: many must be considered the result of gerontomorphism. Schultz (1957b)
has recently repeated this warning.

Paedomorphism represents a slowing down in differentiation —a delay in timing;
gerontomorphism is a speeding up in differentiation—an acceleration in timing. It
is the balance between these two in different parts of the body that produces the
distinctively human form among primates and, to a large extent, the distinctions of
physically different ethnic groups among humans. Clearly it is not possible to
consider all the examples of primate paedomorphism and gerontomorphism, even if we
knew them, so I must confine myself to a few of the most outstanding.

TIMING IN THE HEAD.
The Skull.

It is well known that the skulls of all major primates are closely similar up to
the time of birth (Abbie, 1952a). At that stage the calvariae are little more than
osteo-fibrous membranes for the brain (Abbie, 1947) and a rudimentary facial skeleton
is suspended below. Thereafter differentiation proceeds actively, but at different rates
in different animals. In most non-human primates the jaws, in particular, rapidly
become large and protuberant and the calvariae may show such developments as big
brow ridges and nuchal and sagittal crests, although these features are not necessarily
correlated (Abbie, 19520). Human skulls never attain to great extremes. As Bolk
pointed out originally: in the skull, and particularly in the relatively large round
brain case, the human retains more of the foetal character. In different ethnic groups,
of course, suppression of such features varies considerably (Fig. 7), and the same
applies within any ethnic group—a European may well have quite big jaws and
large brow ridges, an Australian aborigine may have relatively small jaws and a high
round forehead (Abbie, 1951). Nor is the shape of the head predetermined and
immutable. In 1911 Boas (see Boas, 1940) showed that human headform can change
with change in environment and this has since been confirmed by other observers (see
Kaplan, 1954). I have shown (Abbie, 1947) that the changes detected by Boas are not
haphazard but, in fact, with improvement in environment headform tends to adhere
more to the foetal type: neither much longer nor much shorter than the mean foetal
cranial index at about the middle of the human scale (Fig. 7). It seems, then, that
paedomorphism is still an active factor in determining at least one human character.

Apart from such obvious features, human adherence to foetal standards is disclosed
in more subtle ways—egenerally thinner cranial bones, delayed closure of sutures,
tendency to retain a metopic suture, more forward siting of the foramen magnum, more
open spheno-maxillary fissure (Hone, 1952), trend towards failure of third molar teeth
to erupt, and so on. Aiso, crznial paedomorphism is accentuated where growth is
arrested short of full differentiation as in dwarfs, pygmies and adult females.

Gerontomorphism in the human skull is shown in a number of less obvious
features. The relatively high nose has departed more from the foetal standard than
has the flatter nose of apes. This may be “adaptive’. The mastoid process is usually
much better developed than in most apes and here timing is deeply involved: Schultz
(19576) points out that the human mastoid process appears soon after birth and is
practically maximal by adulthood; in the gorilla mastoid development does not begin
until after the permanent dentition is completed and is not finished until old age,
when the process is as large as in a young man. In this respect man has far
outstripped the ape in attainment of a high differentiation, and this may well be
“adaptive” —to provide secure attachment for the sternomastoid muscles which help
to hold the head upright.

Particular interest centres upon the jaws, which betray an interesting mixture
of paedomorphism and gerontomorphism. Overall, the human jaws are relatively small,
a paedomorphic feature possibly aggravating the tendency to suppression of the third
molar teeth — from lack of time and/or space to get them through. Perhaps associated

204 SIR WILLIAM MACLEAY MEMORIAL LECTURE,

with this is suppression of the premaxillae as separate bones. This may be due either
to prenatal closure and obliteration of the premaxillo-maxillary sutures or to over-
growth of the premaxillae by the maxillae (for discussion see Johnson, 1937; Wood
Jones, 1938). In either case the result must be considered paedomorphic. It marks a
very decided departure from the regular primate pattern and is not, I think, to be
dismissed so lightly as Schultz (19576) suggests. In the lower jaw Murphy (1957)
has shown that there are two distinct growing parts, an alveolar border and a basal
portion, which should be treated separately. Only occasionally do humans show
much true facial prognathism— with the jaws as a whole protuberant; but they
frequently exhibit some alveolar prognathism — when the alveolar borders and teeth
protrude in front of the line of the jaws proper (Abbie, 1952a). In this case alveolar
growth exceeds basal growth. In apes alveolar (as well as facial) prognathism is
marked and the chin is left behind—is decidedly receding, in fact. In man, to a
varying extent in different ethnic groups, there is usually a well-marked chin which
indicates that basal growth in the mandible has kept up with, or even exceeded,
alveolar growth. The absolute amount of basal growth may be less than in apes but
the human chin is relatively more advanced and differentiated, and this must be
considered an instance of gerontomorphism.

Cranial Capacity and the Brain.

Human cranial capacity far exceeds that of any other known primate. No gorilla, .
even one three times the size of a big man, has a capacity much more than some
600 c.c. while the human average is around 1400 c.c. The human range is very wide —
from about 800 ¢.c. (an exceptional minimum recorded by Le Gros Clark, 1937) to well
over 2000 c.c.—but within that range size is no index of mental ability, and this
includes all putative “missing links’, such as Pithecanthropus, Sinanthropus,
Neanderthal Man and so on, but not the Australopithecinae, which are down towards
the ape series.

The human brain is not, of course, absolutely the biggest known —it is exceeded
by that of the larger whales and adult elephants. But here the factor of total body
bulk intrudes and raises the question of relative brain size. This is determined by
the brain weight : body weight ratio. On this criterion the human wins handsomely
over all other large brains. In the biggest whale the ratio is about 1: 25,000 (Wood
Jones and Porteus, 1929), i.e. brain weight is only 0:004% of the total body weight,
whereas in an adult human male the ratio exceeds 1:50, i.e. the brain is more
than 2% of the total hody weight (Vierordt, from Donaldson, 1895). But even in
this respect man is not supreme, since the little marmoset (Hapale) has a higher
ratio, although the brain itself is, naturally, very much smaller.

The human brain, then, is distinguished by two factors — great absolute size and
great relative size. What has timing to do with this? That is best determined by
comparing growth of the human with that of other primates. For this we may use
some figures supplied by Schultz (195706) with the qualification that he relates cranial
capacity in cubic centimetres (not brain weight) with body weight in grams (his
table 3 and fig. 9). The cranial capacity of the newborn human is 14% of the body
weight, and the figure is very similar for the great apes. Apes attain maturity at
about 11 years as compared with twice as long for man, yet the apes end up with a
cranial capacity of less than 0:25% of body weight whereas the human finishes
with a cranial capacity of 2% of body weight. Man shows considerable retardation in
physical maturation as compared with apes, and his brain departs much less from
the foetal proportion: this may legitimately be cited as an outstanding exhibition of
paedomorphism. The point is well illustrated by Vierordt’s figures (see Donaldson,.
1895, p. 69) which compare the relative weights of all the important viscera at
successive stages of human development.

Bopy AS A WHOLE.
Man shares with other animals an axial growth gradient of cephalo-caudal
differentiation during ontogeny (Child, 1915). This is shown first by the large head,
tapering trunk and absence of limbs (Fig. 1). As development proceeds, the trunk

BY A. A. ABBIE. 205

becomes distinguishable, then the upper limbs and finally the lower limbs appear,
enlarge and ultimately catch up to their proper proportions. That is, the peak of
the gradient gradually shifts backwards, and in the limbs similarly moving gradients
control proximo-distal differentiation.

For example (Fig. 2), the crown-rump length drops from over 70% to just over
65% of the total stature (crown-heel length) during the last six months of gestation.
Vertical head height, which starts near 50%, is only 30% at three months and has
dropped to just under 25% at birth. On the other hand, the upper and lower
extremities, which appear later, show a second-month acceleration—the upper ahead
of the lower — which continues during the third and fourth months and then tapers
off —again the upper before the lower. Within the extremities the first spurt is in
the proximal segments, which slow down as the intermediate and distal segments

} J
(
g (|
A B
Zz
i
F H

Fig. 1.—Comparative foetal stages: A-D, macaque at a days, 31 days, 36 days and 53
days respectively (drawn from Heuser and Streeter, 1941); EH-H, human at 5-2 mm., 7:3 mm.
(drawn from Streeter, 1945), 12:2 mm. and 30-7 mm. respectively.

successively take over the impetus and in turn catch up and then slow down (data
from Scammon and Calkins, 1929). The process is similar in other primates. Many
more illustrations of this distal shift in the growth gradient peak could be given
but the point is sufficiently made for our purpose.

_ It would be convenient to be able to describe this shift in a single word. I had
considered, and discarded as ugly, the term ‘“distalization’” when I came upon the
botanical adjective ‘“acropetal’ to express the idea of “extremity-seeking” growth. With
your permission I shall import it into zoology.

To continue: By the time of birth the human foetus has a head which occupies
nearly one-quarter of its total length, a trunk twice as long but inferior extremities
only a little longer than the head (Fig. 3, A). With the passage of time these
proportions change. Growth of the head slows down -— at six years it is about one-sixth
of the total stature and in the adult only just over one-eighth. The changing ratio
of brain case to face is also quite notable. Growth of the trunk and upper extremities
proceeds at a moderate pace without striking changes. But the inferior extremities
pursue vigorously their process of catching up, occupying progressively more and
more of the total stature and, in effect, pushing the irunk farther and farther from
the ground. In adult European males, in the outcome, the top of the pubis is just
above the mid-point and the sitting height is only a little over half the total stature.

206 SIR WILLIAM MACLEAY MEMORIAL LECTURE,

Various factors may modify the final proportions. Anything that accelerates
maturation or slows down the growth rate—e.g. hypergonadism, malnutrition, achondro-
plasia, hypothyroidism, etc.— hampers extension of the lower limbs which end up
relatively shorter than the norm. This is evident in several kinds of dwarfs, and also
in healthy adult females who stop growing some years earlier than males. On the
other hand, anything that delays maturation or accelerates the growth rate —e.g.
hypogonadism (eunuchoidism), hyperpituitarism — favours extension of the inferior
extremities which, to our eyes, become disproportionately long.

However, the European standard of bodily proportions is only one possibility and
other ethnic groups may depart from it quite distinctly, while still within terms
of the cephalo-caudal gradient. Some, particularly mongoloids in China, Japan, Alaska
and South America, whose growing period does not seem to be curtailed, exhibit
such slowing down that their inferior extremities are relatively short and they

B €

L.E.

Toe) Ue, ata

Percentage of total (<.H) length

Percentage of total length
Percentage of folal length

12.16 20 24 28 32 356 40 1216 20 24 28 32 36 40 12 16 20 24 28 32 36 40
Weeks of infrauterine life Weeks of intrauterine life Weeks of infrauferine life

Fig. 2.—Human: Comparative growth of body as a percentage of. total (crown-heel)
length. A, Body and total extremities; B, Upper extremity; C, Lower extremity. C.R.L.,
crown-rump length; L.E., lower extremity (total); U.E., upper extremity (total); V.H.H.,
vertical head height; U.A., upper arm; F.A., forearm; H., hand; Th., thigh; L., leg; F., foot.
(Data from Scammon and Calkins, 1929.)

retain more foetal proportions in the trunk and head (Fig. 4). Keith (1919) has
attributed this to a sort of ethnic hypothyroidism, but that seems unlikely. On the
other hand, many peoples, notably in Africa, within the limits of the normal human
growing period achieve so much acceleration that their limbs appear exaggerated in
comparison with the trunk and head (Fig. 4). This, Keith has suggested, may be
due to hyperpituitarism, but again that is unlikely. The Hottentot woman presents an
interesting anomaly. Although she belongs to an almost “pygmoid”’ group, her
extremities, particularly the inferior, are relatively very long. In this case it seems
that some “genetic acceleration” overrides the usual picture of dwarfism.

Among long-legged peoples, the most interesting to us are the Australian aborigines.
We have now (Abbie and Adey, 1953a; Abbie, 1957 and unpublished data) a large
collection of measurements and X-rays relating to aboriginal growth which afford some
insight on what happens. It is evident from figure 5 that in both sexes the proportion
of stature contributed by the inferior extremities in aboriginal adults greatly exceeds
that in Europeans. At birth the proportions are about the same in the two peoples
(Fig. 3, A and B), and the aboriginal growing period may even be slightly less than
the European (Abbie and Adey, 1953). Yet in aborigines of both sexes at about the
sixth year the inferior extremities make a sudden spurt to produce proportions com-

BY A. A. ABBIE. 207

parable with those of a Huropean child of twelve. Thereafter extension of the inferior
extremities does not appear to proceed any faster than in Huropeans. It is the sixth-
year spurt, superimposed upon the regular growth pattern, that determines the
aboriginal: advantage in this respect. Here the matter of timing is clearly important.

42 Years Fully Grown Eully Crown

(16 Years) (20 “Years)

eet Y Yr eo att@ @aeas iaak@' fanntG (Manes
B Just After 6 Years 12 Fully Grown
Birth (20 Years)
Fig. 3.—Progressive growth pattern in, A, Europeans (redrawn from Abbie, 1950), B,
Australian aborigines (personal data).

The Huropean child does show a spurt, but delays it until about the age of twelve,
when there, is not time enough left to catch up with the aborigine. There do not
seem to be sufficient data on other long-legged peoples to determine whether or not
they follow the aboriginal pattern.

Acropetalism extends to segments of limbs and, generally speaking, where limb
growth as a whole is accelerated in comparison with Europeans, so is growth of the

208 SIR WILLIAM MACLEAY MEMORIAL LECTURE,

distal segments, which become relatively longer. The different forms in figure 4
illustrate this sufficiently, particularly in the inferior extremities. The trend is less
noticeable in the superior extremities, for in most peoples the tips of the extended
fingers fairly constantly reach to just below the middle of the thigh, no matter how
long this may be (Figs. 3 and 4). Nevertheless, the trend does exist and is expressed

CHIRIGUAN EUROPEAN NUBIAN

a Rear t
JAPANESE EUROPEAN MusoKO HOTTENTOT

Fig. 4.—Comparison of proportions of various ethnic types, all drawn as far as possible
to the same length from photographs in Martin (1928). The dots indicate, so well as the
poses permit, the approximate ends of the various limb segments.

as a relatively longer forearm and hand. Conversely, those whose limbs remain short
betray suppression of acropetalism in their relatively shorter distal segments.

The great apes present quite a different picture (Fig. 6). Their acceleration is
most obvious in the superior extremities, which also show marked acropetalism. In
comparison, the inferior extremities are relatively stunted (although acropetalism is
evident in the foot: see Schultz, 1949, 1957a, 1957b, for data). This could be due to

BY A. A. ABBIE. 209
the fact that the shorter growing period (only half the human) limits the time
available for extension of the inferior extremities. It is equally likely that the total
pattern is genetically determined in “adaptation” to brachiation and the quadrupedal
mode of progress imposed by the inadequate feet. At all events, it is clear that
embryological timing has emphasized the anterior end of the growth gradient rather
than the posterior end. The accent on inferior extremities in the human must be
considered an example of gerontomorphism vis-d-vis the apes.

TIMING IN OTHER DEVELOPMENTS.

A glance at Table 1 will disclose a number of further features which betray
equally well the total slowing up of human development as compared with other
primates. Gestation, development of pigmentation and hair, ossification, onset and
completion of dentition, growing period and onset of senescence are all progressively
slowed down, as we go from monkeys, through apes to man, in whom the trend is
outstanding. A number of developmental features indeed — pigmentation, hair growth,
dentition — despite the extended growing period, may never reach finality at all.

TABLE 1.
Carpal
Gesta- Completion Complete Ossifica- First Second |Growing Life
Primate. tion. of Covering tion Dentition. | Dentition. | Period. | Span.
(Weeks.)| Pigmentation. of Hair. Centres (Months.) | (Years.) | (Years.) | (Years.)
at Birth.
Macaque 24 Early in ges- During gesta- All 0:6— 5-9 | 1-6- 6:8 7 25
tation. tion. centres.
Gibbon 30 Onset during Onset during 2-3 1 2= 9? 2 = 3398) 9 33
Orangutan. . 39 gestation, gestation, 2-3 4-0-13-0 | 3-5— 9:8 11 30
Chimpanzee 34 completed completed 2 2-7-12-3 | 2°9-10-2 11 35
Gorilla 37 after birth. after birth. ? 3:0-13-0 | 3-0-10-5 11 35
(Coloured.)
Onset mainly
Man a 40 after birth. Never com- 0) 6-0-24-0 | 6-0-20-0 20 70+
(White.) pleted.
Never.

Date from Bolk, de Beer and Schultz (slightly modified).

Here I should add in parenthesis that while pigmentation does become complete in
some coloured peoples this must be attributed to “natural selection’, not to alleged
affinities with sub-human primates. In many aborigines completion of pigmentation is
delayed until adolescence or later (Abbie and Adey, 1953b).

One might add that the tail, so well developed in most monkeys, is suppressed
almost to extinction in both the apes and man. That is an example of paedomorphism,
but it is also an “adaptive” process, since the remnants of the tail and its muscles
are modified to form the pelvic floor which supports the viscera in the upright posture.
This is of considerable importance in humans but less so in the more quadrupedal
apes. However, such a contrivance does not seem to be always necessary since
other orthograde animals, e.g. the kangaroo, manage very well without it.

Although the longer inferior extremities in man represent a manifestation of
gerontomorphism, some qualification is necessary in regard to the foot. As is well
known, in all non-human primates the great toe is widely separated from the others —
like the thumb in the hand—and this feature is established early in prenatal life
(Fig. 1). In man the great toe not only lies close to the others, it is firmly attached
to them by the deep transverse ligament of the sole (deep transverse metatarsal
ligament of Wood Jones, 1944). This is a serious stumbling block to those who would
derive the human foot directly from the ape’s. It has been claimed that in the
mountain gorilla (G. beringei) the great toe lies closer to the foot than in the lowland
gorilla (G. gorilla) and may represent an intermediate stage (e.g. Morton, 1935;
Schultz, 1957b and others). I feel that this would be hard to sustain.

I

Separation of

210 SIR WILLIAM MACLEAY MEMORIAL LECTURE,

the toes depends upon the development of radial splits around the periphery of the
footplate (Fig. 1). In non-human primates the split for the big toe extends deeply,
releasing a highly mobile organ. In man the splitting is partly suppressed. It is
true that in some ape feet a fleshy web extends across the interval but the deep
transverse ligament does not include the great toe within its grasp in any primate
other than man (Wood Jones, 1944; Raven, 1950). In man, fixation of the great toe
is an important factor in preserving orthograde stability and in walking in the
upright position. Apes, despite the fact that they may go for limited periods on
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Fig. 5.—Comparative proportion of adult male and female Huropeans and aborigines all
drawn to the same dimensions (redrawn and modified from Abpie, 1957).

Fig. 6.—Comparative proportion of man and other primates taking trunk length as a
common basis. (Redrawn and modified from Martin, 1928.)

their hind limbs alone, are essentially quadrupeds, supporting themselves on their
excessively long upper extremities and ‘‘walking”’ upon only the outer borders of the
feet. So far as man is concerned, the peculiarity of the great toe is functionally
“adaptive”; in our present context limitation of splitting off is an example of
suppression in development — paedomorphism.. Suppression in development is also
exhibited by the other toes, which never become as long relatively as in apes (Schultz,
1957b) but leave the “big” toe in a dominant position. On the other hand, modification
of the astragalus and os calcis to form the human talus and caleaneum, and so the

BY A. A. ABBIE. 211

human heel, implies a great advance, even upon apes, in differentiation and
specialization. i

Here there is space only to mention the high grade of specialization in the human
vertebral column, sacrum, thorax and pelvis, all in the interests of the upright
posture (Schultz, 19570).

OTHER CONSIDERATIONS.

A few other points demand attention. Acceleration of growth may lead to high
differentiation, i.e. gerontomorphism. But prolongation of growth at a lesser speed
may approximate to the same end result and it cannot be denied that prolongation
of the growing period really represents retention of the foetal tendency to grow, i.e.
it is paedomorphism. Yet it is conceivable that these two diverse processes could
produce a similar outcome. How can confusion be avoided here?

I think that the answer lies in considering what is achieved in a given time.

EUROPEAN

NEWBORN INFANT

Fig. 7.—Adult human skulls of contrasting types derived from a common foetal pattern.

To take a crude example, a gorilla, in most physical features, including size,
has achieved far more differentiation in 11 years than a human has in twice the time.
In this respect the human is strongly paedomorphic in comparison with the gorilla
am toto. Many peoples have longer inferior extremities than Europeans, but so far as
can be detected they all achieve that distinction within approximately the same growing
period. They are more differentiated in the same time and must be considered in
this respect gerontomorphic; those who end up with shorter inferior extremities
after an equal period of growth are paedomorphic relative to Huropeans. The superior
extremities of apes are obviously gerontomorphie in this respect, but when it comes
to the inferior extremities as a whole difficulty arises. The apes have much shorter
inferior extremities, but they also have a much shorter time in which to develop
them. Here it would be rash to decide without qualification whether the apes are
paedomorphic or whether “adaptation” outweighs everything else.

Perhaps when comparing animals of different genera it would be wise not to
depend too much upon absolute chronology but to introduce some form of physiological
chronology. This is a matter that requires more consideration than I have been able
to give to it.

A point that has not been discussed in detail is the effect of the environment on
the expression of genetically determined characters. The term “adaptation”, of course,
implies that the environment is involved, but for each animal it is a special

int

212 SIR WILLIAM MACLEAY MEMORIAL LECTURE,

environment. In the case of man I am concerned mainly with the improvement in
environment — including particularly nutrition—that has followed the advance of
civilization. There is little doubt that our changing environment is producing some
physical changes. This is evident, for example, in the increase in stature and weight
of modern school children as compared with their parents at the same age, and
probably in more subtle ways, such as the change in headform detected by Boas (1940)
in the children of American immigrants. Elsewhere (Abbie, 1948) I have considered
some aspects of that problem. Here I need only say that we have no idea yet of what
is the optimum environment for man—vuntil we have we cannot do more than
speculate on what he could become with his present genetic make-up.
For the moment I must ignore any possible effects from increasing radiation.

: CONCLUSION.

In tracing the development of primates we are watching different expressions of
what is essentially the same process. All start off at about the same point but are
endowed with different potentialities which become manifest during ontogeny as the
imposition of differences in the timing of secondary developments. In one part change
starts early and differentiation is advanced; in another it is delayed and differentiation
is correspondingly retarded or even suppressed.

Development can be looked upon as a cinematographic film which comprises the
whole of differentiation and can be run fast or slow as desired, or at different speeds
at different times. When the whole film, or any section of it, is run through fast,
development is accelerated, parts rush te completion, many details are blurred or lost,
others become exaggerated; nevertheless, much more film can be run through in the
time available and in terms of differentiation much more can be achieved. If the
film is run through slowly the whole process is drawn out, features are unfolded in
great detail and some hitherto unsuspected disclose themselves; however, much less
film can be run through before time is up—if the running is slow enough even an
extended showing, as in man, is inadequate—and the total achievement in final
differentiation is correspondingly reduced. We have no idea of the proper speed for
the film, or even its length — although the result seems to be reasonably satisfactory
for any particular animal—but it would be interesting to speculate on the results
if the speeds were changed. Certainly there is wide scope for variation and so far
as man is concerned we get some hints from various developmental disorders and
ethnic differences. At all events, the speed is evidently not constant: most animals
show retardation in some features, acceleration in others. That applies equally to
man but on balance I think that in man, as compared with other primates, the slowing
down far outweighs the speeding up. In other words, paedomorphism is the major
factor in deciding human peculiarities.

We can say that while paedomorphism sets the basic human pattern gerontomorphic
intrusions impose many decisively human specializations. Instances of both may
clearly be designated “adaptive”. But “adaptation” is only a rather teleological way
of saying that some genetically determined characters become emphasized one way
or another because that fosters survival in man’s particular context. It is noteworthy
that this applies not to the characters themselves—which are common to all primates—
but to the timing they are accorded during development.

The explanation for man’s distinctions, then, is to be found in his genetic make-up,
and particularly in that part of it which regulates the rate at which different elements
of the pattern unfold. Therefore, when looking for some common ancestor for man
and other primates it is necessary to seek among embryos, not adults (Abbie, 1952a,
19520). This is simply a special case of von Baer’s modification of the Meckel-Serres
“law”. If I am correct—and I hope that I have persuaded you that I have some
reason on my side—man’s ancestry and affinities are not to be discovered by comparison
of adult primates, particularly when the possibility of convergence is taken into
account. Conceivably, a minor embryological twist in any primate stock could
introduce the changes in timing necessary to produce the human stem and “selection”
would do the rest. Consequently I am sceptical about “missing links” and “sub-human”
forms of hominids.

BY A. A: ABBIE. 213.

However that may be, it is evident that the features that distinguish man — and
men — are, at least largely, due to differences in timing during development. I do not,
of course, think that that is the only factor, and I may have over-emphasized its
importance. If I am wrong then I can take comfort from Goethe: “Es irrt der
Mensch so lang er strebt.”

References.

ABBIn, A. A., 1947.—Headform and human evolution. J. Anat. Lond., 81: 233-258.

, 1948.—The individual and the environment. Med. J. Aust., (1): 321-327.

, 1950.—Principles of Anatcmy. 3rd Ed. Angus and Robertson, Sydney.

, 1951.—The Australian aborigine. Oceania, 22: 91-100.

, 1952a.—A new approach to the problem of human evolution. Trans. Roy. Soe.
S. Aust., 75: 70-88.

, 1952b6.—The problem of human origins. Aust. J. Sci., 14: 200-201.

, 1956.—Will science explain the nature of life? Awst. J. Sci., 18: 137-141.

, 1957.—Metrical characters of a central Australian tribe. Oceania, 27: 220-243.

and Adey, W. R., 1953a.—Ossification in a central Australian tribe. Human Biology,
25: 265-278.

, 1953b.—Pigmentation in a central Australian tribe with special reference to fair-
headedness. Amer. J. Phys. Anthrop., (n.s.) 11: 339-359.

DE BEER, G. R., 1940.—Hmbryos and Ancestors. Clarendon. Press, Oxford.

Boas, F., 1940.—Race, Language and Culture. Macmillan, New York.

BoukK, L., 1926.—Das Problem der Menschwerdung. Fischer, Jena.

CuHILD, C. M., 1915.—Individuality in Organisms. University Press, Chicago.

CLARK, W. H. Le Gros, 1937.—The status of Pithecanthropus. Man, 73.

and MEDAWAR, P. B. (Eds.), 1945.—Hssays on Growth and Form. Clarendon Press,
Oxford.

DONALDSON, H. H., 1895.—The Growth of the Brain. Walter Scott, London.

GARSTANG, W., 1928.—The morphology of the Tunicata, and its bearing on the phylogeny
of the Chordata. Quart. J. Micros. Sci., 72: 51-187.

GrRowTH, 1948.—Symposia of the Society for Haperimental Biology. No. Il. University
Press, Cambridge.

Harpy, A. C., 1954.—Escape from specialization. In Hvolution as a Process. Ed. J. S. Huxley,
A. C. Hardy, J. B. Ford. Allen and Unwin, London.

HEUSER, C. H., and STREETER, G. L., 1941.—Development of the macaque embryo. Carnegie
Inst. Wash. Pub. 525, Contrib. to Embryol., 29: 15-55.

Hone, M. R., 1952.—The postorbital wall. A comparative and ethnological study. Trans. Roy.
Soc. S. Aust., 75: 115-130.

Huxuey, J. S., 1932—Problems of Relative Growth. Dial Press, New York.

JOHNSON, H. H., 1937.—The narial margins in man. J. Anat. Lond., 71: 356-361.

JONES, F. Woop, 1938.—The fate of the human premaxila. J. Anat. Lond., 72: 462.

, 1944.—The Foot. Bailliére, Tindall and Cox, London.
and PorTeus, S. D., 1929.—The Matrix of the Mind. Edward Arnold, London.

KAPLAN, B. A., 1954.—Hnvironment and human plasticity. Amer. Anthrop., 56: 780-800.

KEITH, A., 1919.—The differentiation of mankind into racial types. Lancet, (ii) : 553-556.

LINNAEI, C., 1758.—Systema Naturae. Editio Decima, Tomus 1, Laurentiae Salvii, Holmiae.
Photographic facsimile, ed. G. de Beer, British Museum (Natural History), London.

MARTIN, R., 1928—Lehrbuch der Anthropologie, Bd. I. Fischer, Jena.

MeEpAwaAr, P. B., 1945——The shape of the human being as a function of time. Proc. Roy.
Soc., B, 132: 133-141.

Morton, D. J., 1935.—The Human Foot. Columbia University Press, New York.

MurpHyY, T., 1957.—Changes in mandibular form during postnatal growth. Aust. Dent. J.,
2: 267-276.

RAVEN, H. C., 1950.—The Anatomy of the Gorilla. Henry Cushier Raven Memorial Volume,
ed. W. K. Gregory, Columbia University Press, New York.

ScAMMON, R. E., and CALKINS, L. A., 1929.—The Development and Growth of the External
Dimensions of the Human Body in the Fetal Period. University of Minnesota Press,
Minneapolis.

ScHutTz, A. H., 1949.—Ontogenetic specializations of man. Arch. d. Julius Klaus-Stiftung,
24: 197-216.

, 1957a.—Die Bedeutung der Primatenkunde flir das Verstandnis der Anthropogenese.

Ber. wiber die 5. Tagung d. Deut. Ges. f. Anthropol., 13-28, Musterschmidt, Gottingen.

, 19576.—Past and present views of man’s specializations. JIrish J. Med. Sci., 341-356.

SIMPSON, G. GAYLORD, 1944.—Tempo and Mode in Evolution. Columbia University Press,
New York.

STREETER, G. L., 1945.—Developmental horizons in human embryos. Carnegie Inst. Wash.,
Pub. 557, Contrib. to Embryol., 21: 29-63.

THOMPSON, D’A. W., 1942.—On Growth and Form. 2nd Ed. University Press, Cambridge.

214

SOME MORE BARK- AND TIMBER-BEETLES FROM AUSTRALIA.
158. CONTRIBUTION TO THE MorRPHOLOGY AND TAXONOMY OF THE SCOLYTOIDEA.
By Kari H. ScHEDL, Lienz, Osttirol, Austria.
(Communicated by Dr. A. J. Nicholson.)

[Read 30th July, 1958.]

Synopsis.

This paper lists a number of new records of Scolytoidea in Australia, from both native
and imported timber intercepted at Australian ports. One new genus and one new species
are described.

Two more consignments of Bark- and Timber-Beetles from Australia, one submitted
by the courtesy of the Commonwealth Institute of Hntomology in London, the other
one originating from the collection of Mr. J. W. T. Armstrong, Callubri, Nyngan,
New South Wales, give the opportunity to enlarge our knowledge about the Australian
fauna and also to show which species have been introduced with imported timber in
various ports. The results of the identifications, as usual, are divided into new
records and the description of new species.

NEw REcorRDS.
Collection Mr. J. W. T. Armstrong.
Leperisinus tricolor Schedl, N.S.W., Acacia Plateau, J. W. T. Armstrong.
Leperisinus bimaculatus Schedl, N.S.W., Acacia Plat., J. W. T. Armstrong.
Xylechinus acaciae Lea, N.S.W., Waratah, 13:111.1940.
Acacicis abundans Lea, N.S.W., Acacia Plat., J. W. T. Armstrong; N.S.W., Waratah,
13.111.1940.
Acacicis minor Schedl, N.S.W., Acacia Plat., J. W. T. Armstrong.
‘-Hylesinus varians Lea, N.S.W., Acacia Plat., J. W. T. Armstrong.
Aricerus ficit Lea, N.S.W., Acacia Plat., J. W. T. Armstrong; Wide Bay, Queensland,
8 Palace, S., N.G.R., 25.11.1905.
Scolytogenes cryptolepis Schedl, N. Queensland, Cairns, Brooks.
Xyleborus testaceus Walk., N. Queensland, Cairns, Brooks.

Commonwealth Institute of Entomology.

Leperisinus bimaculatus Schedl, N. Queensland, Cairns, BH. W. Ferguson collection.

Diamerus interstitialis Lea, N. Queensland, Gordonvale, 1919, ex coll. S.A. Mus.; N.S.W.,
EH. W. Ferguson.

Zygophloeus australis, n. g., n. sp., N.S.W., Lisarow, 29.xi.1953, reared ex Acacia
decurrens, K. M. Moore.

Cryphalus hagedorni Hgg., C.8.1.R.O. Exp. Farm, Katherine, N.T., August, 1947, A. Wynn.

Stephanoderes melasomus Lea, N.S.W., Lisarow, 21.viii.1954, K. M. Moore, ex Muellerina
eucalyptifolia.

Xyleborus novaguineanus Schedl, N. Q’ld., Wongabel, 11.11.1930, R. H. Doggrell, boring
in Hndiandra palmerstonii, per A. L. Tonnoir; N. Q’ld., Cairns, BH. W. Ferguson
collection.

Platypus wmcompertus Schedl, N.S.W., Dorrigo, 238.111.1954, J. Cartwright, ex Hucalyptus
laevopinea.

Platypus pernanulus Schedl, Wongabel, 10.x.1930, ex Bolly Gum (Litsea reticulata).

Platypus queenslandi Schedl, N.S.W., Lisarow, 11.v.1954, P. Hadlington and K. M.
Moore, ex Hucalyptus saligna.

Platypus subgranosus Schedl, N.S.W., Bulga, 22.926, W. W. Froggatt, dead tree on
ground.

PROCEEDINGS OF THE LINNEAN SocIETy oF NEw SouTH WALES, 1958, Vol. Ixxxiii, Part 2.

BY KARL E. SCHEDL. 215

Imported with Lumber from the Pacific Islands or the Hast Indies.

Xyleborus cognatus Blandf., N.S.W., Sydney, 31.x.1924, W. W. Froggatt, in Red Lauan
(Shorea sp.); N.S.W., Vanikoro, Sydney, Vera Cruz,* Kauri (Agathis sp.), log,
1925, W. W. Froggatt; South Australia, Fiji origin, xi.1952, J. Wright.

Xyleborus laeviusculus Blandf., N.S.W., Sydney, Vanikoro, Vera Cruz,* Kauri (Agathis
sp.), log, 12.ix.1924, W. W. Froggatt; N.S.W., Sydney, 31.x.1924, W. W. Froggatt,
in Red Lauan (Shorea sp.).

Xyleborus fleutiauxi Blandf., N.S.W., Sydney, 31.x.1924, W. W. Froggatt, in Red Lauan
(Shorea sp.).

Xyleborus testaceus Walk., N.S.W., Sydney, 31.x.1924, W. W. Froggatt, in Red Lauan
(Shorea sp.); South Australia, xi.1952, J. Wright, Fiji origin; N.S.W., Lismore,
17.x.1952, K. Bootle; Solomons, Santa Cruz Isl., 1926, W. W. Froggatt, Kauri log.

Diapus pusillimus Chap., N.S.W., Sydney, 31.x.1924, W. W. Froggatt, in Red Lauan
(Shorea sp.).

Arizyleborus medius Keg., N.S.W., Balmain, Borneo Cedar (Shorea sp. (?)), 20.111.1928,
W. W. Froggatt.

Platypus jansoni Chap., Fumigator, A. Brooks, 1926.

Platypus pseudocupulatus Schedl, N.S.W., Balmain timber yards, 5.v.1953, M. B. Cappa,
ex Ramin (Gonystylus sp.) from Rejang River, Sarawak.

Platypus pseudopacus Schedl, N.S.W., Sydney, Wallis’ yards, W. W. Froggatt, 23.111.1923,
Pacific maple (Dipterocarpus sp.).

Platypus subgranosus Schedl, N.S.W., Borneo in Pacific maple, 3/22, Wallis & Co.

Crossotarus mniszechi Chap., N. Guinea, Kikori, G. N. ..., W. W. Froggatt.

Diapus 5-spinatus Chap., W. Australia, 1924, J. Clark, Borneo timber; Solomons, Santa
Cruz Isl., 1926, Kauri log, W. W. Froggatt collection; N.S.W., F.C., Pyrmont,
16.11.1948, P. Hadlington, ex Shorea sp.

Diapus pusillimus Chap., N.S.W., F.C., Sydney timber yard, ex N. Guinea Walnut
(Dracontomelum magniferum), 8.iv.1941, K. L. Taylor.

Xyleborus ferrugineus Fab., N.S.W., Lismore, 17.x.1952, K. Bootle.

ZYGOPHLOEUS, Nn. gen.

General appearance similar to that of Hypoborus Er. Head concealed under the
pronotum, the latter ascending from the apex to the base as in many Hylesinae,
trapezoid in outline, base bisinuate, scutellum not visible. Antennae with the scape
club-shaped, the funiculus 6-segmented, the club consisting of three joints separated
by distinct septa. Base of the elytra crenulate, the asperities becoming larger and
higher towards the suture, outline and declivity similar as in Hypoborus Er.

_ This new genus has to be placed in the Hypoborini between Glochicopterus Schedl
and Acacicis Lea, on one side, and those genera with a 5-segmented funiculus on
the other.

ZYGOPHLOEUS AUSTRALIS, Nn. Sp.

Dark reddish-brown when mature, 1:3 mm. long, 1:9 times as long as wide,
pubescence yellowish brown.

Front broadly convex, feebly aplanate in the centre, feebly shining, finely rugosely
sculptured, covered with very short scale-like setae.

Pronotum wider than long (18:12), widest at the base, postero-lateral angles not
rounded but closely attached to the elytra, sides very feebly and obliquely narrowed
in the basal half, thence with a rather strongly developed constriction, apex very
broadly rounded; ascending from the apex to the base, feebly convex, surface silky
shining, minutely punctulate, very finely punctured, the punctures bearing short
inclined scale-like setae.

* There is little doubt that Froggatt labelled these specimens Vera Cruz in error.
Vanikoro is in the Santa Cruz group, near the Solemon Islands. It should also be noted
that Froggatt frequently used his stock labels printed “Sydney, N.S.W.’’ to indicate that
specimens were collected in Sydney, even though they were bred from timber imported from
other places.

216 BARK- AND TIMBER-BEETLES FROM AUSTRALIA.

Elytra somewhat wider (20:18) and twice as long as the pronotum, sides parallel
on the basal half, apex broadly rounded, declivity commencing somewhat behind the
middle, evenly convex; disc striate-punctate, the striae more distinct towards and on
the declivity, the strial punctures moderately large, the interstices rather narrow,
finely and densely rugose, each one of them bearing a median row of remotely placed,
ascending spatulate scales, the smaller, inclined scales on each side of the median row
somewhat irregular in arrangement; declivity with pubescence inconspicuous, the suture
with more numerous inclined scales.

Types: Holotype in Division of Hntomology Collection, C.S.1.R.O., Canberra,
Australia; four paratypes in British Museum (Natural History) and four in collection
Schedl.

Locality: New South Wales, Lisarow, 29.xi.1953, reared ex Acacia decurrens,
K. M. Moore.

XYLECHINUS ACACIAE Lea.
(= Phloeophthorus acaciae Lea.)
The examination of the antennae of Phloeophthorus acaciae Lea has proved that
this species must be referred to the genus Xylechinus. Some specimens recently
collected at Waratah, Australia, 13.111.1940, belong to this hitherto doubtful species.

217

TWO NEW SPECIES OF HEMICYCLIOPHORA (NEMATODA; TYLENCHIDA).
By M. R. Saurr, Commonwealth Research Station, Merbein.
(Communicated by Mr. A. J. Bearup.)

(Two Text-figures.)

[Read 30th July, 1958.]

Synopsis.
Two new species of Hemicycliophora (H. tesselata and H. brevicauda) are described
which differ from known species of the genus in that the females have less than 200 body
annules.

INTRODUCTION.

The number of body annules in the female has long been accepted as an important
character in the separation of certain genera of the Criconematinae. Taylor (1936)
separated Criconemoides and Procriconema in part on this character: Criconemoides
with 160 or less annules, Procriconema with 200 or more annules. More recently, Loos
(1948) synonymized Procriconema and Hemicycliophora. The concept that females of
Hemicycliophora have 200 or more body annules has been sustained by Tarjan (1952)
in a review of the genus, and by Thorne (1955) in a paper describing 15 new species,
and including a valuable key to the known species. Lately, Colbran (1956) has
published the description of an Australian species, H. truncata, which appears to have
about 200 annules.

Chitwood and Birchfield (1957), in erecting a new genus, Hemicriconemoides,
within the subfamily Criconematinae, expressed opposition (in a footnote) to the idea
of setting apart Hemicycliophora as having 200 or more annules. The nematodes now
described appear to support the stand taken by Chitwood and Birchfield, in that these
nematodes appear clearly to belong to the genus Hemicycliophora, yet both have
between 140 and 160 body annules. In these two species the females have a sheath with
simply marked cuticle; the males have no sheath and no stylet, the spicules are
hooked, and caudal alae are prominent.

Acceptance of forms with about 150 annules as belonging to Hemicycliophora
increases the difficulty of distinguishing between females of Hemicycliophora and
Hemicriconemoides; however, Chitwood and Birchfield suggest that the presence of
a dorsal intestinal extension anterior to the base of the oesophagus may be a good
generic character for Hemicriconemoides. 'The two species described in this paper do
not show such an extension.

HEMICYCLIOPHORA TESSELATA, n. sp. (Fig. 1).

Measurements (on sheath): 5 99.—L = 0-850 to 0-950 mm.; a = 18-22; b = 5:3-5-9;
ce = 13-18; V = 90-92%. 3 g¢—L = 0:°690 to 0-735 mm.; a@ = 27-5-29; b = 5:9-6:8;
ce = 9-1-9-5.

Female: Body stout, cylindrical, arcuate when relaxed by heating. Larval cuticle
attached fairly closely to body except at tail. Annules about 150, very coarse, the
annules of the sheath rather flattened, those of the body cuticle more or less rounded.
Tail bluntly rounded, terminus hemispheroid. At the terminus the body cuticle carries
a hemispherical appendage whose structure could not be ascertained. This appendage
is readily stained in cotton blue lactophenol and therefore does not appear to be of
euticular origin. The head in most specimens, but not ali, is retracted in the sheath
which forms a protruding collar around it.

The cuticular annulations are marked by 20 rather deep and wide longitudinal
grooves, so that the surface appears as rows of rectangular blocks. A diagrammatic

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutH WALES, 1958, Vol. Ixxxiii, Part 2.

218 TWO NEW SPECIES OF HEMICYCLIOPHORA,

representation is given in Text-figure I, F, but the ful! number of grooves can be
determined only in transverse section, and only a few rows of cuticle blocks are visible
under high power of the microscope. No evidence of a lateral field was seen.

=

JOM mor
OvU0CGbG0a0
OoObe0enBnoH
JOO0000G00
QOooemr

Fig. 1.—Hemicycliophora tesselata, n. sp.
A, female; B, male; C, face view, female; D, anterior end, female; H, region of vulva;
F, surface structure, female; G, G,, head end male, ventral and lateral views; H, caudal alae,
ventral view; J, male tail; K, K,, lateral fields, male. ;

The head, surmounted by a large rectangular labial disc, appears to consist of
two annules, not set off from the body. It has a stout framework. In face view the
head framework is hexaradiate, and the amphids (?) appear near the edge of the
labial disc as barely visible slits. Spear rather stout, 95 to 105 microns, comprising a

BY M. R. SAUER. 219

long shaft and thick basal portion about 15 microns long. Spear knobs large, without
forward pointing processes. Dorsal gland opening about 12 microns behind the base
of the spear. Median oesophageal bulb about half body width, with large valve;
posterior bulb about one-third body width. The excretory pore is situated about
20 microns behind the junction of oesophagus and intestine. Hemizonid large.

Ovary single, outstretched, varying from 42% to 52% of total body length in
specimens observed. Vagina rather long, strongly cuticularized. Ventral contraction
of body at vulva. Oviduct long with prominent ovoid spermatheca containing
spermatozoa. Posterior end of intestine and rectum not seen. Anus visible only in
ventral view, opening some four body annules posterior to the vulva.

Male: No sheath. Body slender, arcuate when relaxed by heating, finely annulated.
In the head region and on the caudal alae the cuticle shows a paired annule formation,
every second annule being set off by a deeper constriction. Lateral fields marked by
four incisures, the inner lines more or less straight, the outer lines crenate at
intervals equal to two body annules.

Framework of head much reduced, no spear. Oesophagus reduced, no bulbs or
valve. The excretory pore opens opposite the anterior end of the oesphagus and a
prominent hemizonid is visible a few body annules anterior to it. Intestine vacuolate.
Testis quite short, 13% to 17% of total body length in specimens studied. Spicules
paired, long and strongly curved. Marked protrusion of the body cuticle at the cloaca.
Gubernaculum short, simple. Caudal alae thick, prominent. Tail long, conical to a
fairly blunt terminus.

Types—Holotype: Female, slide Hemicycliophora 1, Commonwealth Research
Station, Merbein. Collected 14th August, 1957. Allotype: Male, slide Hemicycliophora 1,
data as above. Paratypes: Five females, two males.

Type locality: Soil beneath Hucalyptus incrassata Labill., beside the Calder High-
way, one mile south of Hattah, Victoria.

HEMICYCLIOPHORA BREVICAUDA, nl. sp. (Fig. 2).

Measurements (on sheath): 12 99——L = 0-710 to 0-785 mm.; a = 18-20; b = 4:6-5:6;
c€=—; V = 96-97%. 7 6¢.—L = 0:500 to 0-590 mm.; a = 25-30; 0 = 5-8-7:2; e = 12-14.

Female: Body stout, cylindrical, marked by about 150 simple, coarse annules.
Annules of sheath somewhat flattened. Sheath fitting body fairly closely, sometimes
loose at terminus. Tail hemispherical. Lateral field indicated by a more or less
distinct, faint longitudinal groove which tends to be most prominent in the mid-region
of the body. All specimens show some irregularity of the annules — usually one annule
in the mid-region divides to two along the lateral groove. Some specimens show two
or three such irregularities, always widely separated. Other lateral lines are often
indicated by faint markings or irregularities in the annules; possibly there may be
four lines altogether but no more than three have been seen in specimens examined.
The specimen illustrated (Text-fig. 2, H) has one very prominent incisure and
indications of a second line. The divided annule is typical.

The head consists of two annules, not set off, surmounted by a large, nearly square
labial disc. Face view similar to H. tesselata but the head framework inclined to be
more slender. Two slits may represent the amphids. The spear is long, 85 to 92:5
microns, with the robust basal portion about 15 microns long. Dorsal oesophageal
gland opening about 10 microns behind the spear base. Excretory pore about 20
microns posterior to the oesophago-intestinal junction. Hemizonid large.

Ovary single, outstretched, 42% to 55% of total body length in these specimens.
Vagina strongly cuticularized. Ventral contraction of body at vulva. Long oviduct
including a distinct, nearly round spermatheca well filled with spermatozoa. Intestine
ending near vagina, rectum reasonably short but seldom visible. Anus seldom visible
in lateral view. Anus opens about two body annules behind the vulva.

Male: No sheath. Body slender, finely annulated. Lateral fields marked by four
nearly smooth lines, about one-third body width.

Head framework reduced. No spear, oesophageal bulbs, or valve present. Hemi-
zonid prominent a few annules anterior to the excretory pore, which opens just behind

220 TWO NEW SPECIES OF HEMICYCLIOPHORA,

the anterior and end of the intestine. Intestine vacuolate. Testis short, 16% to 21%
of total body length in specimens measured. Spicules long, strongly curved. Marked
protrusion of cuticle around the spicules. Gubernaculum short, simple. Definite
constriction of the body at the posterior end of the prominent caudal alae. ‘Tail rather
short, with acute terminus.

Vor
oven

>
ma

eg
ee end i
i= a TE

lees)
()

ii) come ae) AN, [p) ofl

Fig. 2.—Hemicycliophora brevicauda, n. sp.

A, face view, female; B, female; C, male; D, anterior end, female; E, male tail;
F, female tail; G, G,, male head lateral and ventral aspects; H, lateral field, female.

Types.—Holotype: Female, slide Hemicycliophora 2, Commonwealth Research
Station, Merbein. Collected 14th August, 1957. Allotype: Male, slide Hemicycliophora 2,
data as above. Paratypes: 45 females, 6 males.

Type locality: Soil beneath Codonocarpus cotinifolius F. vy. M., on the track to Lake
Mournpoul, Hattah, Victoria.

DIAGNOSIS.
Hemicycliophora tesselata and H. brevicauda are readily distinguished from species
of Hemicycliophora previously described by the low number of body annules in the
female, about 150 for each species. In other respects they resemble most closely

BY M. R. SAUER. 221

H. obtusa Thorne, 1955, in showing a hemispheroid terminus and a ventral contraction
at the vulva. Males are not Known in most species of the genus (including H. obtusa).

Hemicycliophora tesselata is distinguished as follows: Bisexual species. Female
with sheath, about 140-160 annules, hemispheroid tail, ventral contraction at the vulva,
body annules divided by 20 longitudinal grooves, sheath usually protruding around
head. Male without sheath, spicules much curved, caudal alae prominent, moderately
long conical tail, blunt terminus.

Hemicycliophora brevicauda is distinguished as follows: Bisexual species. Female
with sheath, about 140-160 plain annules, very short hemispherical tail, vulva near
terminus, ventral contraction at vulva. Male without sheath, spicules much curved,
short pointed tail constricted at the end of the prominent caudal alae.

Acknowledgements.
Mr. L. Smith, of Commonwealth Research Station, Merbein, washed the soil samples
for the author. These samples were processed in a modified Seinhorst extraction
apparatus (Seinhorst, 1956).

References.

CHITWOOD, B. G., and BIRCHFIELD, W., 1957.—A new genus, Hemicriconemoides (Criconematidae:
Tylenchina). Proc. helm. Soc. Wash., 24: 80-86.

CoLBRAN, R. C., 1956.—Studies of plant and soil nematodes. I. Two new species from
Queensland. Qd. Jour. agric. Sci., 13: 123-126.

Loos, C. A., 1948.—Notes on freeliving and plant parasitic nematodes of Ceylon. 3. Ceylon
Jour. Sci. (B), 23: 119-124.

SHINHORST, J. W., 1956.—The quantitative extraction of nematodes from soil. Nematologica,
1: 249-267.
TARJAN, A. C., 1952.—The nematode genus Hemicycliophora de Man, 1921 (Criconematidae)
with a description of a new plant parasitic species. Proc. helm. Soc. Wash., 19: 65-77.
Taytor, A. L., 1936.—Genera and species of the Criconematonae, a subfamily of the
Anguillulinidae (Nematoda). Trans. Amer. micr. Soc., 50: 391-421.

THORNE, G., 1955.—Fifteen new species of the genus Hemicycliophora with an amended
description of H. typica de Man (Tylenchida: Criconematidae). Proc. helm. Soc. Wash.,
22: 1-16.

222

A NEW SPECIES OF FROG OF THE GENUS CRINIA TSCHUDI FROM
SOUTH-EASTERN AUSTRALIA.
By Murray J. LittLeyoHN,* Zoolegy Department, University of Western Australia.
(Communicated by Mr. 8. J. Copland.)

(One Text-figure.)
[Read 30th July, 1958.]

Synopsis.

Differences in male call, as measured by objective sound analysis, initialiy indicated the
presence of a new species of Crinia in south-eastern Australia. This characteristic together
with other biological (breeding behaviour and in vitro crossing) and morphological data has
been used to confirm the status of the new form with respect to two closely related sympatric
species, C. signifera Girard and C. parinsignifera Main. The new species is grouped with the
C. insignifera superspecies. A description is given at the conclusion of the paper.

INTRODUCTION.

Differences in male call are generally considered to act as important reproductive
isolating mechanisms between related species of Anura (Blair, 1955, 1956). Thus the
revealing of distinct and discontinuous variability of call structure within one presumed
biological species leads to the suspicion that more than one species may be present.

While travelling through the Riverina district of Victoria and New South Wales
during August, 1957, with the aim to obtaining additional amphibian material and
tape recordings of male calls of Crinia parinsignifera Main and C. signifera Girard,
samples of a previously unrecorded Crinine call were obtained. The new call was
quite distinct from those of the other two species to which collected specimens
appeared to show close morphological affinity. All three call types were heard and
recorded calling together in several localities without any evidence of intergradation
in call structure. Subsequent examination of collected material showed that males
of the new call type could be readily distinguished on morphological grounds, but
that greater difficulty was experienced with femaies. The new form showed no close
relationships to any species of Crinia from south-western Australia.

RELATIONSHIPS.

On the basis of its morphology, biology and call structure the new call type
should be included within the C. insignifera superspecies of Main (1957).

This latter author has discussed at some length the highly variable texture
and patterns on the dorsal surface of these species and has separated distinct variants
or polymorphs into four categories, namely:

1. Striped: animals in which two continuous longitudinal ridges and several
variously coloured yellow, brown or red stripes traverse the back.

2. Lyrate: two raised lyre-shaped ridges are found on the shoulders. No
longitudinal ridges or stripes are present.

3. Warty: the back lacks either of the above conditions, but is covered in
numerous fine warts or spicules.

4. Smooth: no raised dermal areas are present and the pigmentation is
uniform.

These polymorphs may be present in varying proportions in all the previously
described members of the C. signifera-insignifera complex. There is also marked

* Present address: Department of Zoology, University of Texas, Austin, Texas, U.S.A.

PROCEEDINGS OF THE LINNEAN SocIETY oF NEw SoutrH WALES, 1958, Vol. Ixxxiii, Part 2.

BY MURRAY J. LITTLEJOHN. 223

variation in back colour and distribution of pigments. In the new species, however,
only one back pattern type has been found, namely the lyrate type, the light mustard-
brown back colour and occasional ochre-capped warts being consistent with the lyre
markings.

Main (1957) also considered the degree of pigmentation of the male throat and
the colour and pigmentation of both male and female belly to be of some assistance
in species diagnosis in the complex. These characteristics are here considered with
regard to the two sympatric species C. parinsignifera and C. signifera and the new
species. The throat of a breeding male of the new species has a pale grey-green
pigmented band around the mandible border with a fainter grey pigmentation in life
extending back to the forearms. In spirit the colour assumes a uniform light grey
appearance, while the belly is granular, dirty-white and sparsely flecked in black.
The female throat is white, immaculate, while the skin of the granular belly may
carry small pigmented black flecks. In males of C. signifera the very dark pigmenta-
tion of the throat extends posteriorly to or slightly beyond the forearms, where two
white pectoral spots may be present. The male belly is often darkly pigmented in
parts either in blotches or spots. The female has a white throat, but the belly is
usually heavily pigmented in an extremely variable pattern in which extensive blotches
may run together to give a dark appearance.

C. parinsignifera males show slightly reduced pigmentation of the throat compared
with that of OC. signifera, and a white belly, while the females have a white belly
and throat.

DISTRIBUTION AND HABITAT.

The new species has been collected and heard calling in the Murray River Valley
from Mulwala through to Echuca. It was not found at Mildura, Victoria, nor north of
Deniliquin and Finley, New South Wales. However, males were heard calling on the
plains country close to the foothills of the Great Dividing Range between Wangaratta
and Whitfield, Victoria. In the latter localities the habitat consisted of shallow
temporary ponds in clay soil, but in the other localities the species was generally
restricted to temporary ponds in the river valleys and up to five miles on either side
of the larger rivers.

Throughout the entire known range the new species is broadly sympatric with
either C. parinsignifera or C. signifera, often with all three species occurring together.

C. signifera is found in permanently wet situations throughout the Great Dividing
Range and south and east thereof with an extension into the higher rainfall areas
of the western foothills and along the greater river valleys farther west.

C. parinsignifera occupies the drier areas inland to the west of the Great Dividing
Range and generally inhabits temporary summer-dry ponds of this region.

BREEDING BIOLOGY.
Male Call Characteristics.

Samples of male calls were obtained for all three species under natural conditions
using portable tape recording apparatus. Calls of the new species and of C.
parinsignifera were analysed by cathode ray oscilloscope and the following acoustical
characteristics determined: call duration, number of pulses, and pulse repetition
frequency and approximate carrier frequency. These characteristics for C. signifera
were determined by the Ferrogram method of Frings and Frings (1956) (excepting
earrier frequency which was determined by oscilloscope). In all three species call
repetition rate was calculated by measuring with a stop-watch the time taken for an
individual to make ten successive calls.

The results of the analyses are presented in Table 1. The data used were
corrected to an effective recording temperature of 10°C. in order to minimize any
variation in call structure due to temperature effects, the correction factor being
determined by regressing each call characteristic against effective temperature using
the method of least squares. The oscillograms are presented in Text-figure 1, and
verbal call descriptions are given in diagnosis of the species.

224 A NEW SPECIES OF FROG FROM SOUTH-EASTERN AUSTRALIA,

Breeding Season.

The area was visited during August, 1957, when all three species were heard
calling strongly. Three mating pairs of the new form were seen during this time.
Main (1957) collected gravid females of C. signifera and C. parinsignifera during the
same month in 1955, so that it may be assumed that no marked seasonal isolation
is present.

Fig. 1.—Oscillograms of calls of C. signifera (upper left), C. sloanei (upper right), and
C. parinsignifera (lower). Each spot in the line across the top of the traces is equal to
0-01 second.

Calling and Breeding Temperatures.

All three species have been heard in chorus over the temperature ranges: air,
5:0-10-5°C.; water, 7:0-10-75°C.

Mating pairs of the new species were collected at air temperatures of 9-0°C., and
water 9-5°C., while Main (per. com.) collected females of ©. signifera and C.
parinsignifera at air 7:-5°C., and water 9-5°C. It seems reasonable to assume that here
no effective temperature isolation is operating.

TABLE 1.

Data from Call Analyses of Recordings from Victoria and New South Wales, Corrected to a Common Temperature of 10° C.
Mean value-tstandard error of the mean and ranges are given for each species.

Species.
Call Characteristic.
sloanet. parinsignifera. signifera.
Sample size (number of individuals) 17 35 14
Duration (secs.) 0-06+0-001 0:48-0:01 0-15 +0-006
(0-06-0- 08) (0:32-0:59) (0:12-0:19)
Number of pulses 13:0+0-4 78:0+1°8 5:0+0°3
(11-0-16-0) (56-0-100-0) (4-0-9-0)
Pulse repetition frequency (C.P.S.) 214-0+5-3 167:0+2:7 31-:0+1°6
(166 - 0—250-0) (141 - 0-219 -0) (25 - 0-49 - 0)
Call repetition rate (secs.) 9:7+0:28 61:5+2-7 6:0+0-2
(7- 7-11: 4) (39-0-115-4) *(4-7-8-2)
Approximate carrier frequency (C.P.S.) 2600 2800 2500
*N=24.

Sample compositions and effective temperatures.

ae

. parinsignifera
5, 16:5°C.;

io

sloanet bn .. Mulwala, N.S.W., 9, 10:-75°C.;
Mulwala 7, 10-75° C., 7, 6:75° C.; Echuca, Vic., 13, 9:0° C.; Kingston on Murray, S.A.,
Horsham, Vic., 3, 7:7° C.

‘. signifera .. .. Tocumwal, N.S.W., 7, 10:0° C.; Wangaratta, Vic., 7, 10:5° C.

Tocumwal, N.S.W., 8, 10°C.

BY MURRAY J. LITTLEJOHN. 225

Calling Position.

Males of the new species usually call while floating in open water of temporary
ponds. C. parinsignifera males call while out of water, 1—4 inches above the surface,
supported by tussocks of dried grass, either at the centre or periphery of the temporary
ponds. C. signifera males call while partly immersed in the water, sitting on submerged
vegetation at the borders of ponds, and generally under cover of overhanging grasses.

Some variation in calling position does occur, particularly when the males are
moving into the breeding sites. From the overall distinctness, however, it is suggested
that such behaviour could assist in maintaining efficient reproductive isolation.

In Vitro CROSSES.

Some crosses were made under field conditions, following as closely as possible
the methods of Rugh (1948), and, while giving poor larval survival in both the
experimentals and controls. do indicate that the cross 2? new species x ¢ C. signifera
can yield apparently normal offspring to metamorphosis (Table 2). In the cross 2 new
species x ¢ C. parinsignifera no embryos develeped beyond gastrula but it cannot
surely be said whether the failure was due to genetical breakdown or to poor
experimental conditions.

TABLE 2.
Results of In-vitro Crosses between the Three Species of Crinia.
Female. Male. Initial Hatched. | Metamorphosed.
Egg Number.
sloanet X parinsignifera oie ae 22 1 0
sloanet X signifera sa ee we 26 8 2
sloanet x sloanei (control) .. a 21 7 4 and
1 delayed

DISCUSSION AND DESCRIPTION.

From the above analysis it appears that efficient pre-mating reproductive isolation
is maintained principally by differences in male call and is probably assisted by
preference for distinct calling positions. There are not sufficient experimental data
available as yet to predict whether significant gene flow could occur if the pre-mating
mechanisms broke down and there is no evidence of interbreeding in the sympatric
field populations.

The distinctness of male call, sympatric occurrence with C. parinsignifera and
C. signifera without evidence of integradation and the consistent morphological
distinctness when considered together indicate that this new population warrants
species status for which the following description is given.

CRINIA SLOANEI, sp. nov.*

Types.—Holotype: 466/57. A sexually mature male in the Zoology Department
Collection at the University of Western Australia, taken in a temporary pond, on the
south bank of the Murray River by the Main Traffic and Railway Bridge adjacent
to Tocumwal on Murray, New South Wales. Allotype: 473/57. Zoology Dept. Univ.
W.A. Coll. Paratypes: 465/57, 467/57, 468/57, 470/57. Zool. Dept. Univ. W.A. Coll.

All specimens were collected by M. J. and P. G. Littlejohn on 6th August, 1957.

To be lodged with the Australian Museum, Sydney, New South Wales, together
with a tape recording of typical calls and photographs of oscilloscope traces.

Description.

Vomerine teeth absent. Snout short and rounded, 1:3 times as long as the eye;
canthus rostralis rounded; loreal region oblique; tympanum indistinct; fingers long,
free, with some subarticular tubercules; toes free, long, often with dermal fringes;
two metatarsal tubercules; tibio-tarsal articulation reaching eye or tympanic region.

*The name is designated in appreciation of the assistance of Mr. Ian F. Sloane, of
“Savernake Station’, Savernake, New South Wales, without whose co-operation collection in
this area would not have been possible.

226 A NEW SPECIES OF FROG FROM SOUTH-EASTERN AUSTRALIA,

Dorsum smooth except for two consistent (in life prominent) lyre-shaped ochre-
coloured ridges over shoulders; posterior regions sometimes bearing a few small
ochre-capped warts; dorsum a mustard-yellow colour in life, but pale grey-brown in
spirits.

Ventral surface prominently granular; throat of females white, while males have a
(greyish-green in life) grey border to mandible and a pale grey throat; belly of both
males and females white and sparsely flecked with small black spots.

Body Lengths.
Samples of breeding adults from Tocumwal showed the following mean snout-vent
measurements: Males: 15-6 mm. (S.D. 0-4 mm.); N=16. Females: 17:6 mm.; N = 2.

Range and Habitat.

Found associated with temporary ponds along Murray River from Wangaratta to
Echuca and in the south-eastern part of the plains adjacent to the western face of
the Great Dividing Range near Wangaratta and Whitfield, Victoria, and north of the
Murray River to Deniliquin and Finley, New South Wales.

Diagnosis.

a. Morphology: The consistent mustard-yellow-coloured back and the lyre-shaped
markings are characteristic of the new species, all the others of the complex being
polymorphic in back pattern and variable in colour. The pale pigmented area of male
throat may be of some assistance when comparing with the more extensive pigmentation
of throats of males of C. parinsignifera and C. signifera.

b. Call: The call of the new species may be described as a short metallic “chick”
regularly and rapidly repeated; that of C. parinsignifera is a long low “squelch” or
“buzz” repeated very slowly. C. signifera males make a short 4—7-pulsed call which is
rapidly repeated, described by Harrison (1922) as “Crick-ick-ick-ick-ick”’. The physical
characteristics are given in Table 1.

c. Inviability: There are indications that crosses C. parinsignifera g x new
species Q are inviable, while crosses C. signifera g x new species 9 show low Viability
but that some survivors reach metamorphosis (Table 2).

Acknowledgements.
The work was carried out while the author was in receipt of a research grant
from the University of Western Australia.
The author acknowledges the assistance of Patricia G. Littlejohn in field collecting.
He is also indebted to Dr. W. F. Blair for reading the manuscript.

References.

Buair, W. F., 1955.—Mating Call and Stage of Speciation in the WMWicrohyla olivacea-
M. carolinensis Complex. Evolution, 9: 469-480.

- —, 1956.—Call Difference as an Isolation Mechanism in Southwestern Toads (Genus
Bufo). Tex. Jour. Sci., 8: 87-106.

FRINGS, H., and FRINGS, M., 1956.—A Simple Method of Producing Visible Patterns of Tape
Recorded Sounds. Nature, 178: 328-329.

HARRISON, L., 1922.—On the Breeding Habits of Some Australian Frogs. Awst. Zool., 3: 17-34.

MAIN, A. R., 1957.—Studies in Australian Amphibia. 1. The Genus Crinia Tschudi in South-
western Australia and some Species from South-eastern Australia. Aust. Jour. Zool,
5: 30-55.

RuGH, R., 1948.—Haperimental Hmbryology (Rev. Hd.). Burgess Publ. Co., Minneapolis.
Pp. 1-480.

Proc. LINN. Soc. N.S.W., 1958. PLATE It.

Yad

EHupomatia vennettii EF. Muell.

227

ACARINA FROM AUSTRALIAN BATS.
By Rogpert DomrRow, Queensland Institute of Medical Research, Brisbane.

(Twenty-six Text-figures.)
[Read 24th September, 1958.]

Synopsis.

Twenty-one species of mites, belonging to seven families, are now known from Australian
bats.

In the Laelaptidae, the genus Ichoronyssus is recorded from Australia for the first time,
I. leucippe, n. sp., and J. aristippe, n. sp., being described from Miniopterus schreibersii
blepotis. Trichonyssus, n. g., is erected, with Chiroptonyssus australicus Wom. as genotype,
and also including 7. womersleyi, n. sp., based on males from M. s. blepotis and Nyctophilus
geoffroyi, which were originally ascribed by Womersley to Plesiolaelaps miniopterus. Plesio-
laelaps Wom. is considered to be a synonym of Spinolaelaps Radford, and the males of
S. miniopterus (Wom.) and Bewsiella fledermaus Domrow are described from M. s. blepotis
and Hipposideros bicolor albanensis.

In the Trombiculidae, the genus Trombigastia is recorded from Australia for the first
time, represented by TJ. alcithoe, n. sp., from H. b. albanensis. Trombicula dasyphloea, n. sp.,
is described from H. semoni.

In the Listrophoridae, Alabidocarpus recurvus (Wom.), originally described from a single
female, is redescribed in both sexes from Rhinolophus megaphyllus.

The present paper is intended to provide a summary of the known mite parasites
of Australian bats. Sixteen species have been described or recorded since 1931, while
five new species are described below. These 21 species are distributed among seven
families. All these species are discussed below, but there is some doubt whether
several of them are true parasites of bats.

Family SPINTURNICIDAE.

Two species of Spinturnix, a large, widespread genus found exclusively on bats,
have been recorded from Australia. They are S. antipodianus and S. novaehollandiae,
both described from unidentified hosts by Hirst (1931). The paper was posthumous,
and based on brief manuscript notes. Its publication was delayed for some time in
the hope of finding Hirst’s illustrations, but none appear to have been made.

Family LAHLAPTIDAHE.

In addition to the genera and species discussed below, I have recorded (1958)
Neolaelaps spinosus (Berlese) from Pteropus conspicillatus Gould in North Queensland,
and described Bewsiella fledermaus from Hipposideros semoni Matschie from Cape York
Peninsula.

Genus IcHoRONYSSUS Kolenati.
ICHORONYSSUS LEUCGIPPE, n. Sp. (Text-figs. 1-2.)

Types: Holotype female in Queensland Museum, Brisbane; from rump of the bat
Miniopterus schreibersti blepotis (Temminck), Yandina, S.E. Queensland, 10.iv.1958.

Description of female—A medium-sized, well-sclerotized species with slender legs;
length of idiosoma 530u, breadth 3244. Dorsuwm: Dorsal shield well developed, but
tapering somewhat in posterior quarter, and truncate posteriorly; with punctae forming
coarse irregular scale-like pattern over entire surface; with seventeen pairs of lateral
setae (the anterior ones much longer than the posterior), nine pairs of median setae
equal in size to posterolateral setae, and at least thirteen pairs of pores as shown.
Marginal cuticle with about 22 pairs of setae. Stigmata placed ventrolaterally between
coxae III and IV, but with peritremes running forward onto dorsum, and extending

PROCEEDINGS OF THE LINNEAN SOCIETY of NEw SoutTE WALES, 1958, Vol. lxxxiii, Part 3.

A

228 ACARINA FROM AUSTRALIAN BATS,

almost to vertex. Venter: Sternal shield preceded by striate zone behind tritosternal
base; with anterior margin ill defined, and slightly concave between sternal setae I;
posterior margin deeply concave; with usual six setae and four pores in addition to
two irregular porose areas (each composed of four smaller zones) outside anterior
pores. Metasternal setae and pores free in cuticle. Genital shield tapering to just
behind level of coxae IV; with one pair of setae, and irregular scale-like markings
anteriorly, which merge into longitudinal striae posteriorly (the two striae running
inwards and forwards from behind the genital setae are much stronger than the
remainder). Anal shield slightly flattened anteriorly, but rounded laterally and
tapering posteriorly to point, which is covered by minute spinules; anus set in anterior
half, with adanal setae near level of posterior margin of anus, and smaller than
postanal seta. Ventral cuticle probably with three pairs of pores and about 46 setae,

Text-figs. 1-2.—Jchoronyssus leucippe, n. sp. 1, Dorsum of female; 2, Venter of female.

the posterior two setae being slightly stronger than the rest. Legs slender, with II
and III slightly thicker than I and IV; I 398, II 345u, III 356u, IV 463 long.
Coxae II to IV with crescentic sclerotized are in posterior half, and coxae II with strong
spine-like process on anterodorsal margin as illustrated by Hirst (1921, p. 792) for
I. flavus. Femora I and II with two slightly stronger setae on dorsodistal edge as
figured by Hirst (1921, p. 782) for H. sternalis. Gnathosoma: Basal movable segment
of palp ventrointernally with anteriorly directed process as figured by Hirst (1921,
p. 793) for I. flavus. Chelicerae with ventral movable finger strong and unarmed, but
dorsal fixed digit weak, and with minute retrorse spines as figured by Furman (1950,
p. 481) for J. longisetosus.

Distribution.—Known only from the type host and locality in S.E. Queensland, but
see final sentence of remarks on Alabidocarpus below. This is the first Australian
record of Jchoronyssus.

ICHORONYSSUS ARISTIPPE, nN. Sp. (Text-figs. 3-6.)
Types: Holotype female and morphotype nymph in Queensland Museum, Brisbane;
both from rump of the bat Miniopterus schreibersti blepotis (Temminck), Teviotbrook,
S.E. Queensland, 10.x.1957.

BY ROBERT DOMROW. 229

Description of female——Similar to I. leucippe unless otherwise stated. Length of
idiosoma 6894. Dorsum: Dorsal shield with fine, regular striae forming scale-like
pattern over entire surface; with seventeen pairs of elongate lateral setae, eight pairs
of minute median setae, and seventeen pairs of pores as shown. Marginal cuticle with

Text-figs. 3-4.—Ichoronyssus aristippe, n. sp. 38, Dorsum of female; 4, Venter of female.
Text-figs. 5-6.—Ichoronyssus aristippe, n. sp. 5, Dorsum of nymph; 6, Venter of nymph.

about 380 pairs of setae. Venter: Sternal shield with anterior margin straight but
ill defined; with two undivided circular porose areas outside anterior pores. Genital
plate with irregular longitudinal striae of uniform strength. Anal plate evenly rounded
anteriorly. Ventral cuticle with about 50 setae. Legs: I 582u, II 507, III 535u, IV 671u

230 ACARINA FROM AUSTRALIAN BATS,

long. Anterior process of coxae II with minute backwardly directed barbule on
external edge. Gnathosoma: Basal movable segment of palp with ventrointernal surface
obscured during mounting, but apparently modified.

Description of early nymph.—Weakly sclerotized, idiosomal length 389u. Dorsum
with two shields. Anterodorsal shield pointed in front, but widening broadly, and
almost rectilinear posteriorly; with seven pairs of elongate marginal setae, four pairs
of minute median setae, and numerous irregular punctae. Postdorsal shield smaller,
and widely separated from anterodorsal shield; lateral and posterior margins straight,
but anterior margin produced medially into triangular process; with five pairs of
minute lateral setae, two elongate posterior setae, and several punctae. Dorsal cuticle
with five pairs of setae around peritremes, six pairs between two shields and two
terminal setae. Peritremes abbreviated and situated above coxae III and IV. Venter:
Intercoxal shield heptagonal, with five sides slightly concave, and tapering evenly to
a point between coxae III and IV; with three pairs of setae and at least one pair of
pores. Anal shield as in adult. Ventral cuticle with two smaller setae between
coxae IV, and six larger setae in front of anai shield. Legs somewhat stouter than in
adult; I 250yu, Il 242u, III 2144, IV 271u long. Otherwise as in adult.

Distribution.—Known only from the type host and locality in S.E. Queensland.

Remarks.—I have followed Baker and Wharton (1952) in accepting Lepronyssus as
a synonym of Ichoronyssus, although I prefer to keep Chiroptonyssus and Spinolaelaps
apart for the present. da Fonseca (1948) puts Ichoronyssus and Lepronyssus in the
same couplet of his key, separating them on the presence or absence of scale-like
markings on the genital plate. Hirst (1921) gives an excellent figure of these markings
in I. flavus (placed in Lepronyssus by da Fonseca). Furman’s figure (1950) of
I. longisetosus is semidiagrammatic, but shows that markings are present on the genital
shield, particularly a pair of stronger ones running forwards and inwards from behind
the genital setae. Radford’s (1941) figure of J. britannicus shows these two markings
clearly, but I feel his material is really conspecific with J. flavus.

da Fonseca (1948) makes much of the presence of two porose areas on the sternal
shield in delimiting his two monotypic genera Hirstesia and Lepronyssoides, but there
is no doubt that these porose areas are also regularly present in [choronyssus as here
understood. I. flavus (possibly, and certainly if britannicus is a synonym), I. granulosus
and I. longisetosus all possess these porose areas, as do the two species described above.
Figures of species which otherwise fit in I[choronyssus as here undertood, but do not
show these two characters on the genital and sternal shields, should not be accepted
without reserve.

Two other characters are also regularly present in [choronyssus —a ventrointernal
process on the basal movable segment of the palpi, and an anterodorsal spine on
coxae II. Both these characters are present in the genotype of Hirstesia, which, apart
from the additional setae on the genitoventral plate, is a typical Ichoronyssus.
Lepronyssoides is also very close to Ichoronyssus as here understood.

Several species of [choronyssus are recorded from Hurope and America, but they
are not at all well known. I have therefore preferred to describe the above two
species in detail as new. They may be separated by the number of median dorsal
setae present, the size of these setae relative to the lateral setae of the dorsal shield,
and the shape of the porose areas on the sternal shield. Minor differences are also
to be seen in the shape of the dorsal and anal shields, and the pattern of striae on
the genital shield.

TRICHONYSSUS, N. g. (OprE, a hair; vvoocw, to prick).

Diagnosis. —Laelaptid parasites of bats, with the following characters. Female:
Dorsal shield entire; sternal shield with two pairs of setae; sternal setae III free
in cuticle; genital shield with one pair of setae; coxae without heavy spines.
Male: Dorsal shield as in female; holoventral shield entire, slightly expanded behind
coxae IV; legs not abnormally enlarged, their coxae without heavy spines; femora IV
unarmed behind; ambulacral apparatus of tarsi II not modified; body cuticle posteriorly

with numerous extraordinarily long setae. Genotype: Chiroptonyssus australicus
Womersley, 1956.

BY ROBERT DOMROW. 231

In da Fonseca’s keys (1948) Trichonyssus runs close to Chiroptonyssus, but may be
separated in the female by having the metasternal setae free and not set on platelets,
and in the male by the compiete holoventral shield, the lack of a strong process on
femur IV, and the presence of extremely long opisthosomal setae. Two species are
included in the new genus, the genotype (which Womersley only tentatively assigned to
Chiroptonyssus Augustson) and a new species described below.

TRICHONYSSUS AUSTRALICUS (Womersley, 1956), n. comb.
There is nothing to add to the original description of this species, which was
described in both sexes from an unidentified South Australian bat.

TRICHONYSSUS WOMERSLEYI, 0D. sp.
Types: I designate as holotype male the specimen which Womersley (1957)
designated as allotype male of kis -Plesiolaelaps miniopterus from Miniopterus
schreibersiit blepotis (Temminck), Joanna, South Australia, 10.xii.1932, J. Hood coll.

Description of male—Length of idiosoma 422u, width 280u. Leg II stoutest; femur
IV without any process. Dorsal shield entire, with short setae. All ventral shields
fused to form holoventral shieid, with ventral area slightly expanded behind coxae IV.
With about 20 pairs of short setae on ventral cuticle, and about 16 pairs on ventral
portion of holoventral shield. Other ventral setae normal. Posterior margin with two
distinct groups of seven very long setae (100u). Digits of chelicerae unarmed, and
shorter than spermatephore carrier.

Remarks.—This species was deseribed as the male of Plesiolaelaps miniopterus,
but Mr. Womersley is in agreement with me that the sexes were wrongly correlated.
The true male of Plesiolaelaps is described below, the genus being reduced to a Synonym
of Spinolaelaps Radford. T. womersleyi may ke separated from 7. australicus (geno-
type and only other known species) by having elongate setae in two groups of seven
posteriorly instead of in a broad circlet of about 45 around the entire opisthosomal
margin. According to Womersley (in litt., June, 1958) the nymph described as
P. miniopterus also belongs to JT. womersleyi.

Genus SPINOLAELAPS Radford.
SPINOLAELAPS MINIOPTERUS (Womersley, 1957), n. comb. (Text-figs. 7-10.)

Types: Womersley’s holotype female is in the South Australian Museum, Adelaide.
Two plesiotype males are here designated, one in the South Australian Museum and
one in the Queensland Institute of Medical Research, Brisbane, both from Miniopterus
schreibersiit blepotis (Temminck), Teviotbrook, S.EH. Queensland, 10.x.1957. The
plesiotype male in the South Australian Museum could be regarded as the allotype.
The specimen designated as allotype of this species by Womersley is not congeneric
with the holotype, and has been designated above as the holotype of Trichonyssus
womersleyi, N. Sp.

Description of female—An oval, well-sclerotized, medium-sized species, idiosomal
length 410—-428u, breadth 261-278u. Dorsal shield entire, oval, but truncate and very
slightly concave posteriorly; with fine striae forming scale-like pattern over entire
surface, apart from heavily sclerotized vertex; with seventeen pairs of lateral setae,
twelve pairs of median setae (of which the inner posterior pair are very minute),
and fifteen pairs of minute pores. Marginal cuticle with 19-22 pairs of setae. All
dorsal setae subequal. Stigmata distinct, with rather narrow peritremes extending
forward to level of anterior margin of coxae II. Venter: Sternal shield with anterior
Margin evenly convex, and preceded by striate cuticle. Posterior margin concave.
With usual three pairs of sternal setae and two pairs of pores in addition to meta-
sternal pores, which are borne on posterolateral extensions of sternal shield. Surface
of shield with regular transverse striae. Metasternal setae borne on minute platelets.
Genitoventral shield flask-shaped, but somewhat narrower than genital operculum,
which has irregular markings. Shield proper with three transverse striae, and two
lateral and one posterior seta in addition to usual pair of genital setae. Anal shield
with central anus flanked by adanal setae, which are subequal to postanal seta.

232 ACARINA FROM AUSTRALIAN BATS,

Ventral cuticle with about 28 pairs of setae, some of which are borne on minute
platelets. Legs: Leg I enlarged and also with larger claw than II-IV. Tarsus I with
small sclerotized pit on dorsodistal face, presumably to accept tarsal claws when drawn

Text-figs. 7-10.—Spinolaelaps miniopterus (Womersley). 7, Dorsum of female; 8, Venter
of female; 9, Venter of nymph; 10, Venter of male.

Text-figs. 11-16.—Bewsiella fledermaus Domrow. 11, Dorsum of nymph; 12, Venter of
nymph; 13, Venter of male; 14, Postdorsal shield of female, amended; 15, chelicerae of
female on left, and of male on right (at twice indicated scale) ; 16, Venter of female, amended.

back. Femur I with two very strong setae dorsally, and femur II with one. Both
setae on coxae I and posterior seta on coxae II and III slightly thickened. Gnathosoma
with two stronger setae posteroventrally. Chelicerae slender, with weak unarmed
digits.

BY ROBERT DOMROW. 233

Description of male—The male described by Womersley belongs to the new genus
(Trichonyssus) defined above. The true male is as follows. Slightly smaller than
female, idiosomal length 346-357, breadth 225-232u. Dorsally (including peritremes)
as in female. Venter: Intercoxal shield with anterior margin as in female except for
median genital aperture; tapering evenly to terminate roundly between coxae IV;
with usual five pairs of setae and three pairs of pores; with regular transverse striae
anteriorly. Ventral shield tapering posteriorly and then expanding to fuse with anal
shield; with about 22 setae in addition to usual three anal setae. Ventral cuticle
with about five pairs of setae. Legs as in female. Gnathosoma: Chelicerae with fixed
finger reduced and somewhat shorter than stout spermatophore carrying finger.

Description of nymph.—Idiosomal length 328u, breadth 214y. Dorsal shields
indistinct, but setal pattern similar to adult. With reduced peritremes curved above
coxae III and IV. Intercoxal shield broadly rounded anteriorly and laterally; posterior
margin rectilinear between coxae III and IV; with three pairs of setae and transverse
striae. With two smaller setae between coxae IV and six larger ventral setae in
front of anal plate. Legs as in adult.

Material examined.—Six females, two males and one nymph from Miniopterus
schreibersit blepotis (Temminck), Teviotbrook, S.E. Queensland, 10.x.1957; also one
male from Hipposideros bicolor albanensis Gray, Bramston Beach, North Queensland,
13.x11.1957.

Remarks.—Spinolaelaps is in many respects similar to Bewsiella. Both are bat
parasites of similar facies, particularly in the characteristic sternai, genitoventral and
anal areas, the strong lateral peritremes, the enlarged front legs with their strong
claws, and the dorsal setation of femora I and II. Bewsiella, however, has a double
dorsal shield, while Spinolaelaps has one. Womersley (1957) described Plesiolaelaps
miniopterus aS a new genus and species, and compared it with Radfordilaelaps Zumpt
from a South African springhare, but Plesiolaelaps is here considered to be a synonym
of Spinolaelaps Radford, of which the genotype (S. jacksoni Rad.) is a parasite of an
African bat. It is of interest that several other genera discussed in this paper are
also found both on African and Australian bats, e.g., Calcarmyobia, Trombigastia and
Alabidocarpus.

In the female S. miniopterus may be separated from S. jacksoni by having three
additional setae on the genitoventra: plate instead of nine, and by its rather thicker
anterior legs. In the male the shape of the ventrianal shield differs in the two species.
Although the outline of the dorsal shield is probably not shown in either Radford’s
or Womersley’s figure, the general arrangement of dorsal setae in their two genera
is similar.

Note added 6th October, 1958.—I have since been able to see material from both
Womersley’s and Radford’s type series. The shape and setal and pore patterns of
the dorsal shield in both genera are as figured below. In addition Radford’s genus
shows the following characters in common with Womersley’s pit on dorsodistal face
of tarsus I: basal gnathosomal seta enlarged; stigmata sometimes clearly open to
exterior, with peritremes extending forward to anterior margin of coxa II; anal shield
similarly shaped, with tendency to encroach onto dorsal suriace; genital operculum
with irregular markings; some ventral setae on platelets. The variation in number of
ventral setae taken in by the genital shield is clearly only of specific value, and I now
have no hesitation in placing Plesiolaelaps as a synonym of Spinolaelaps.

Genus BEWSIELLA Domrow.
BEWSIELLA FLEDERMAUS Domrow, 1958 (genotype). (Text-figs. 11-16.)

Types: Allotype male in Queensland Museum, Brisbane, as is the holotype female,
and with collection data as given below.

Description of female.—While the original description is adequate for recognition,
the specimen figured is atypical in some respects. All six additional specimens show
the following features. Postdorsal shield slightly concave between two larger posterior
setae; with two very minute setae between and slightly in front of these larger setae.
Posterolateral corners of postdorsal shield rather more angulate, and provided with

234 ACARINA FROM AUSTRALIAN BATS,

small pore near edge of shield; additional paired pores as figured. The pattern of
pores on the anterodorsal shield is as in the original figure. Posterolateral corners
of sternal shield without indentation near sternal seta III, and accordingly evenly
convex, not concave. The posterior margin itself, however, is always quite concave.
With six setae arranged 2.2.1.1 on genitoventral shield in addition to usual pair of
genital setae, just behind which are a pair of small pores. Some ventral setae set
on minute platelets. Chelicerae with fixed digit well sclerotized basally, but apically
with delicate, leaf-like flap bearing two or three minute thorns. Movable digit rather
broader, and also with retrorse thorns.

Description of mate.—Slightly smaller than female, idiosomal length 314y, breadth
202u. Dorsum, peritremes and legs as in female. Venter with three shields. Intercoxal
shield convex anteriorly, but concave around genital aperture; lateral margins with
two concavities, one accepting coxae II, the other coxae III and IV. Surface of shield
with few transverse striae, three pairs of pores, three pairs of sternal setae, and one
pair of posterior setae (representing genital setae of female). Metasternal setae
generally free in cuticle, but on one side of two specimens borne on slender, posteriorly
directed extension of intercoxal shield. Ventral shield subcircular, with irregular
margin, and from 28 to 30 setae. Anal plate as in female. Ventral cuticle with about
sixteen pairs of setae. Gnathosoma: Chelicerae with fixed digit much reduced. Movable
digit fused with slender, but well-sclerotized spermatophore carrier.

Description of late nymph.—Two specimens of this stage were examined, both of
which were enclosed in the pelts of the early nymph described below. They were very
pale and inadequate for illustration. Intercoxal shield and its setation as in male, but
without genital aperture. Ventral area without ventral plate, but with numerous setae.
With two contiguous dorsal shields, the posterior one being somewhat smaller than in
adult. Peritremes as in early nymph. Otherwise similar to adult.

Description of early nymph.—Weakly sclerotized and bluntly oval; idiosomal length
260u, breadth 192u. Hysterosoma not greatly developed behind coxae IV. Dorsuwm with
two shields. Anterior shield much the larger, with one pair of minute vertical setae.
Posterior shield weakly defined, but apparently transverse oval; with four pairs of
setae in addition to a large and a very minute posterior pair. Dorsal cuticle with
twelve pairs of setae arranged as shown. Peritremes abbreviated and lateral, situated
above coxae III and IV. Venter with single intercoxal shield, which expands to level
of coxae III and then tapers rapidly to a point between anterior edges of coxae IV;
with three pairs of setae and two pairs of pores. Ventral cuticle with ten setae
arranged as shown, the posterior pair flanking the anal plate. Otherwise similar to
adult.

Material examined.—Six females, two males (including allotype) and four nymphs
from one of four bats, Hipposideros bicolor albanensis Gray, Bramston Beach, North
Queensland, 13.xii.1957.

Remarks.—This species was originally described from two females from Hippo-
sideros cervinus (Gould) from Cape York Peninsula; the host identification should,
however, be amended to H. semoni Matschie.

Family CHE YLETIDAE.
The only species of this family to be associated with bats in Australia is
Cheletonella vespertilionis Womersley, 1941, from an unidentified South Australian bat.

Possibly this predatory species plays a similar réle on bats to that of Onheyletiella
parasitivorax (Mégnin) on rabbits.

Family MYOBIIDAE.

Four species of this family are recorded from Australian bats. They were all
described by Womersley (1941) in the broad genus Myobia, but have since been
reassigned to different genera.

M. clara, a slender species described without detailed collection data, is now
placed in Neomyobia Radford, 1948.

M. chalinolobus, a very stout species from Chalinolobus gouldii (Gray) from South
Australia is the genotype of Pteracarus Jameson and Chow, 1952.

BY ROBERT DOMROW. 235

The third is a characteristic slender species with foliate dorsal setae, M. miniopterus,
from Miniopterus schreibersti blepotis (Temminck) from South Australia (Womersley
misspells the host genus and specific name of the mite). This species is accepted by
Radford (1954) and Jameson (1955) as a synonym of Myobia rhinolophia Radford,
1940, from Africa, the genotype of Calcarmyobia Radford, 1948.

The fourth, M. minima from C. gouldii, is considered by Radford (1954) to be a
deutonymph incerti generis.

Family TROMBICULIDAE.
Genus MyorroMBICULA Womersley and Heaslip.

This genus contains only a single species, WM. vespertilionis Wom. & Heas., 1943,
based on a single specimen lacking sensillae, and therefore of uncertain affinities.
Nor is it certain that it is a true parasite of bats, the specimen having been found in
the débris at the bottom of a jar of bats of uncertain origin.

Genus TROMBICULA Berlese.
TROMBICULA QUADRIENSIS Womersley and Heaslip, 1943.

This species was originally recorded (1943) from Rattus assimilis and Hydromys
chrysogaster in Queensland, but its synonym, 7. chiroptera Wom. & Heas., was described
with collection data similar to those of Myotrombicula above. Womersley later (1952)
recorded specimens of TJ. quadriensis (subsequently lost) from a bandicoot and a
possum from North Queensland, while I have identified it from Trichosurus vulpecula
(Kerr), D’Aguilar, S.E. Queensland, 4.iv.1957. Here again the evidence that this
species is a bat parasite is inconclusive.

TROMBICULA THOMSONI Womersley, 1954.
This species is close to 7. quadriensis and was taken from Chalinolobus gouldii
(Gray) occupying abandoned birds’ nests on Ayers Rock in Central Australia. <A
third closely related species is described below.

TROMBICULA DASYPHLOEA, Nl. Sp. (dacvddAo1os, With rough rind). (Text-figs. 17-19.)

Types: Holotype and one paratype larva in Queensland Museum, Brisbane; one
damaged paratype larva in Queensland Institute of Medical Research, Brisbane. All
specimens from perineum of a bat, Hipposideros semoni Matschie, Lockhart River
Mission, Cape York Peninsula, 13.vi.1956, M. J. Mackerras coll.

Description of larva—A small and delicate species of ovate shape, without heavy
sclerotization. Body setation: Dorsal setae simple, arranged 2.7.6.7 plus about 18
posteredorsal and caudal setae; HS 46a long, DS 40u long. With one pair of setae
between both coxae II and III, and numerous ventral setae behind them, which are
shorter (30:5u long), and with longer ciliations than DS. Gnathosoma: Chelicerae
stout, blade 23:2u long, without teeth apart from usual tricuspid cap. Galeal setae
well developed and ciliated. Maxillary setae strong and plumose. In addition to usual
ventroexternal tarsala, the palpal formula is B.B.bb(N)B.6B-N. The six setae on the
palpal tarsus comprise one dorsal and five ventral setae. The former is by far the
strongest, having a thick shaft and heavy distal ciliations. The five ventral setae,
apart from the two on the inner ventral edge, are very weakly ciliated, and may
occasionally be nude. Subterminaia absent. Palpal claw three-pronged, the strong
central prong flanked by small dorsal and ventral prongs. Scutwm rather broader than
long; subpentagonal. Anterior margin concave, with small convexity around AM seta,
which is set slightly behind AL setae. Both AL and PL setae set on prominent
“corners”. PL longer than AL and AM, which are subequal. Lateral margins quite
concave, and posterior margin bluntly pointed medially. Surface of scutum roughened
by heavy punctae, except for two subcircular areas behind sensillary bases and median
zone behind AM seta; punctae weaker near “corners” and along extreme anterior
margin. Sensillae set closer to level of PL than to that of AL setae, filamentous, and
with about ten approximately paired ciliations on distal half of shaft. The standard
data are given below. Eyes double, with anterior pair convex, about 15-34 in diameter;
posterior pair much weaker, about 7:34 in diameter. The eyes are well separated

236 ACARINA FROM AUSTRALIAN BATS,

from the scutum, but ali three specimens are fully engorged and somewhat distorted.
Legs all seven-segmented, with unisetose coxae. All tarsi with single basal bar; all
tibiae with two semi-bars; genua I and II with bar and III with semi-bar. The special
sensory setation is as follows: Tarsus I with pretarsala, subterminala and parasub-
terminala, tarsala and microtarsala. Tarsala exceedingly long (42:74) and slightly
tapering. Tibia I with one stout distal tibiala (18-3), microtibiala, and a finer, more
proximal tibiala. Genuw I with two fine genualae and one microgenuala. Tarsus II

Text-figs. 17-19.—Trombicula dasyphloea, n. sp. 17, Scutum and eyes; 18, Gnathosoma
shown dorsally on left and ventrally on right, with inset of humeral, dorsal and ventral
body setae; 19, Specialized setation of legs I-III.

with pretarsala, tarsala (19-54) and microtarsala. Tibia II with one stout tibiala (22)
and one fine tibiala. Genu II with one fine genuala. Tarsus III without specialized
setation. Tibia and genu III with one mastiseta each, 36u to 42u long.

Standard data in micra of larval scutwm of T. dasyphioea, n. sp.
AW PW SB ASB PSB SD AP AM AL PL SENS

Holotype .. 41-1 61-7 14-5 26-6 16-9 43-5 32:7 36:6 36-0 50-8 60-5
Paratype .. 43:6 60-5 13°3 26-6 15:7 42-3 32°7 33°9 33:5 46-0 —
Paratype sees Elio! — 13-3 27:8 15:7 43-5 32-7 — — 43-6 —
Means ao Zale) 61-1 13-7 27-0 16:1 43-1 32°77 35:2 34:7 46:8 60:5

Distribution—Knunown only from the type host and locality on Cape York Peninsula.

Remarks.—T. dasyphloea belongs to a characteristic group of parasites found
chiefly on bats. They have roughened, heavily punctate scuta, and follow after couplet 68
of Womersley’s key (1952, p. 45). It is possible that these species may be congeneric
with the lost genotype, 7. minor Berlese. Of the species known from Australia and
New Guinea, 7. thomsoni Womersley, 1954, from Chalinolobus appears to be most
closely related to the new species. The two may be separated by the scutal standard
data and the number of dorsal setae.

BY ROBERT DOMROW. 237

Genus TROMBIGASTIA Vercammen-Grandjean and Brennan.
TROMBIGASTIA ALCITHOE, Nn. sp. (Text-figs. 20-24.)
Type: Holotype larva in Queensland Museum, Brisbane; from wing membrane of
a bat, Hipposideros bicolor albanensis Gray, Bramston Beach, North Queensland,
13.xii.1957.

Text-figs. 20-24.—Trombigastia alcithoe, n. sp. 20, Scutum and eyes, with inset of dorsal
and ventral body setae; 21, Ventral view of palp; 22, Dorsal view of palpal tarsus;
23, Galeal seta; 24, Specialized setation of legs I-III.

Text-figs. 25-26.—Alabidocarpus recurvus (Womersley). 25, Lateral view of female;
26, Lateral view of male terminalia.

Description of larva.—A very small species even when engorged, but idiosomal
measurements unavailable due to rupture during mounting procedure. Dorsal setae
slender, to 37u long; with barbules stronger on convex side. Total number of humeral,

238 ACARINA FROM AUSTRALIAN BATS,

dorsal and caudal setae about 70. Ventral setae shorter (to 27-7u long), but rather
more curved and with longer ciliations; about 32 in number. Gnathosoma: Chelicerae
missing. Galeal setae nude and slender. Maxillary setae with about four slender
ciliations. In addition to the usual ventroexternal tarsala, the palpal formula is
B.b.bbb.7B—b. One of the two dorsal setae on the palpal tarsus is thick and heavily
ciliated, while the other dorsal seta and the five ventral setae are weakly barbed or
almost nude. Subterminala absent. Palpal claw 3-pronged, 20u long. Scutum trapezi-
form. Anterior margin concave, with small convexity around base of AM seta, which
is set somewhat behind level of the AL setae. Both AL and PL setae set on slight
“corners”. PL longer than AM, which is longer than Al. Lateral margins slightly
concave; posterior margin almost rectilinear between PL setae. Surface of scutum
with numerous small punctae, except behind AM seta and sensillary bases. Sensillae
set much nearer level of PL setae than to that of AL setae; shaft somewhat thickened,
but tapering distally; with barbules on basal third and ciliations on remainder of
shaft. The standard data of the holotype are given below. Eyes convex and double,
with anterior (17-14 in diameter) larger than posterior (13-34 in diameter). Legs all
7-segmented, with unisetose coxae. Intercoxal setae arranged in two pairs. All tibiae
with basal and distal semibar. Tarsi II and III with basal semibar, but tarsus I with
both basal and distal semibars. The special sensory setation is as follows. Tarsus I
with pretarsala, subterminala and parasubterminala, tarsala and microtarsala. The
tarsala is of moderate length (23-94), but does not reach the insertion of the
subterminala. Tibia I with a stout and a slender tibiala and one microtibiala. Genw I
with three slender genualae and microgenuala arranged as shown. Tarsus II with
pretarsala, tarsala and microtarsala. Tibia II with a stout and a slender tibiala.
Genu II with fine genuala. Tarsus III with mastitarsala bearing one or two weak basal
barbules (the other setae of comparable length have numerous barbs along their
entire length). Tibia and genu III with slender tibiala and genuala respectively.
Standard data in micra of holotype of Trombigastia alcithoe, n. sp.
AW PW SB ASB PSB SD AP AM AL PL SENS
53-9 76-2 21-6 26-9 20-8 47-7 44-7 35-4 30-8 43-1 67-8

Distribution—Known only from type host and locality in North Queensland.

Remarks.—There are six species in this compact genus according to Vercammen-
Grandjean and Brennan (1957), three from N.H. Africa and Arabia, and three from
Malaya. JT. alcithoe is closest to JT. harrisoni (Womersley, 1952) among the Malayan
species, but may be separated by its larger scutal standard data and short tarsala I.
The chelicerae of JT. alcithoe are unknown, but all three Malayan species have the
cheliceral blades short and broad-based.

This species and the two I[choronyssus species above are named after the three
daughters of Minyas, who were transformed into bats by Dionysus after declining
to join his revels.

Family ANOHTIDAHE.

In 1942 Womersley described Chiropteranoetus chaiinolobus from a _ single
deutonymph from the débris at the bottom of a jar of unidentified bats. This species
is not considered a true bat parasite, anoetid deutonympns (hypopi) being normally
found effecting their dispersal on insects.

Family LISTROPHORIDAE.
Genus ALABIDOCARPUS Ewing.
ALABIDOCARPUS RECURVUS (Womersley, 19438). (Text-figs. 25-26.)

Description of female.—Length 623-637u. Capitulum free, with two posterodorsal
angles heavily sclerotized. Postcapitular shield with midlateral margins extended
downwards between legs I and II, forming part of sockets wherein these legs are
articulated. With one or two pairs of minute setae immediately behind posterior
margin of shield. Legs I and II much expanded apically, without caruncles, but with
two setae each in posterodistal angles of expanded distal segment. With zone of
striated cuticle behind each of these legs. Leg III with four movable segments, the

BY ROBERT DOMROW. 239

penultimate segment with two setae on outer surface, and the distal segment with
an internal seta and three heavy apical claws, the outer one being twice as strong as the
inner two. Leg IV also with four movable segments, but with only one seta on pen-
ultimate segment (the one in the posterodistal angle), and one inner seta and two apical
claws on the distal segment, the outer being almost three times as long as the inner. All
three smaller claws with three or four minute denticulations on outer distal (convex)
edge. With two setae between coxae III and IV. Legs IV articulated on a curved ventral
apodeme with median longitudinal strut. Above coxae III are a weak and a very strong
seta (with their bases fused or discrete). Above these is a further minute seta, above
which again is sometimes another minute seta. Body cuticle with 52 to 58 annulations
dorsally and 28 to 35 ventrally. Apex of abdomen with two pairs of setae as shown,
and possibly a third small pair just above these. Hysterosoma in all three specimens
containing pale hexapod larva directed posteriorly, with structure as shown, although
some minute setae are probably invisible. The unmodified posterior legs of the larva
resemble the third pair of the adult, having three apical spines.

Description of male.—Exactly as in female illustrated, except for terminalia and
number of body striations. Length 441u. With 36 to 38 annulations dorsally and none
ventrally behind basal movable segment of leg IV. With two pairs of minute setae
on posterolateral annulations. Apex of body almost immediately behind legs IV with
two valves, each with sclerotized zone externally, with a heavily sclerotized bar
running downwards and arching up forwardly. Inner surfaces of valves each with
subcircular sucker and a single minute seta. Margins of valves with three pairs of
heavy setae, the central pair the strongest (125u), and the other two pairs subequal
(538-64). On one side of the male from Palmerston, however, it is the dorsal one
of the three setae that is strongest. Apodeme of leg IV as in female, but with median
longitudinal strut longer, reaching to insertion of two paired setae between basal
movable segments of leg IV. Almost immediately behind these setae (although not
clear due to position assumed by mite) a short longitudinal sclerotized structure
possibly representing penis; it is about as long but not as thick as the inner spur of
tarsus IV. In Figure 26, legs IV are inserted almost immediately in front of the region
figured, the hysterosoma being very short posteroventrally.

Material examined.—The holotype female from an unidentified bat from Bathurst,
N.S.W., 13.iv.1934, S. L. Allman coll, and two pairs of adults from Rhinolophus
megaphyllus Gray, Yandina, 10.iv.1958, and Palmerston, 28.v.1958. All four new
specimens were collected at the base of a different nasal vibrissa. The specimens
illustrated are the pair from Yandina. The slight variations in cuticular setation are
noted in the text. Otherwise I cannot separate the five specimens specifically.

Remarks.—Womersley’s Figure 44 does not show the minute setae on the body
cuticle, nor are the 80 dorsal and 48 ventral annulations the correct number — this has
led Lawrence (1948) into error in his key. The correct setation and relative sizes
of the claws and apical segments of legs III and IV are shown in Figure 4a, yet the
detailed figures of these structures in Figures 4B and 4c are incorrect. Legs Ili and
IV have only four movable segments. A. recurvus is very close to A. nasicolus
(Lawrence, 1938) from an African species of Rhinolophus. The two may perhaps be
separated by the relative lengths of the terminal setae and the denticulations of the
smaller tarsal claws. Miniopterus schreibersti blepotis was also present in the cave
in which the host bats were taken; Alabidocarpus, however, appears typically to be
found on the nasal vibrissae of Rhinolophus (see Lawrence, 1952).

Acknowledgements.

I am grateful to Messrs. H. M. Hale and H. Womersley for the loan of type and
other material and much kind advice; to Drs. I. M. Mackerras and H. H. Derrick for
their helpful criticism of this manuscript and their continued interest in my work;
to Dr. M. J. Mackerras for providing the specimens of Trombicula dasyphloea, nN. sp.;
to Dr. R. L. Doherty and Misses M. L. Emanuel and R. H. Smith for catching many
of the bats; to Mr. BH. Le G. Troughton for identifying several bats; and to Miss HE. Wood
for her care in typing the manuscript for this paper.

240 ACARINA FROM AUSTRALIAN BATS.

References.

BAKER, HE. W., and WHARTON, G. W., 1952.—An Introduction to Acarology. New York.
Macmillan. Page 85.
Domrow, R., 1958.—New and little known Australasian Laelaptidae (Acarina). Proc. LINN.
Soc. N.S.W., 82: 352-366.
DA FonsEcA, F., 1948.—A monograph of the genera and species of Macronyssidae Oudemans,
1936 (synom.: Liponissidae Vitzthum, 1931) (Acari). Proc. zool. Soc. Lond., 118: 249-334.
FuRMAN, D. P., 1950.—New mites (Acarina: Liponyssinae) from North American bats.
J. Parasitol., 36: 479-484.
Hirst, S., 1921.—On some new parasitic mites. Proc. zool. Soc. Lond., 769-802.
, 1931.—On some new Australian Acari (Trombidiidae, Anystidae, Gamasidae). Proc.
zool. Soc. Lond., 561-564.
JAMESON, E#. W., 1955—A summary of the genera of Myobiidae (Acarina). J. Parasitol.,
41: 407-415.
JAMESON, E. W., and CHow, C. Y., 1952.—Pteracarus, a new genus of myobiid mites (Acarina:
Myobiidae) from bats (Mammalia: Chiroptera). J. Parasitol., 38: 218-221.
LAWRENCE, R. F., 1938.—A new acarine parasite of bats. Parasitology, 30: 309-313.
, 1948.—Studies on some parasitic mites from Canada and South Africa. J. Parasitol.,
34: 364-379.
————, 1952.—Two new mite parasites of Natal bats (Listrophoridae, Sarcoptiformes).
Parasitology, 42: 136-143.
RADFORD, C. D., 1940.—Notes on some new species of parasitic mites. Part 3. Parasitology,
32: 91-104.
, 1941.—Notes on some new species of parasitic mites. Part 4. Parasitology, 33:
306-315.
, 1948.—A revision of the fur mites Myobiidae (Acarina). Bull. Mus. Hist. nat.,
Paris, (2) 20: 458-464.
, 1954.—Observations on the fur-mites (Acarina, Myobiidae). Ann. Mus. Congo
Tervuren, in 4°, Zool., 1: 238-248.
VERCAMMEN-GRANDJEAN, P. H., and PRENNAN, J. M., 1957.—Wight new chiggers from Fast
Africa and a new genus, Tromtigastia (Acarina: Trombiculidae). Ann. ent. Soc. America,
50: 484-496.
WoMERSLEY, H., 1941.—Notes on the Cheyletidae (Acarina, Trombidoidea) of Australia and
New Zealand, with Cescriptions of new species. Rec. S. Aust. Mus., 7: 51-64.
, 1942.—Miscellaneous additions to the acarine fauna of Australia. Trans. roy. Soe.
S. Aust., 66: 85-92.
, 1943.—Australian species of Listrophoridije Canest. (Acarina) with notes on the
new genera. Trans. roy. Soc. S. Aust., 67: 10-19.
, 1952.—The scrub-typhus and scrub-itch mites (Trombiculidae, Acarina) of the
Asiatic-Pacifie region. Rec. S. Aust. Mus., 10: 1-673.
, 1954.—A new species of Trombicula (Acarina: Trombiculidae) from bats from
Northern Australia. Ann. Mag. nat. Hist., (12) 8: 827-828.
, 1956.—On some new Acarina Mesostigmata from Australia, New Zealand and New
Guinea. J. Linn. Soe. (Zool.), 42: 505-595.
, 1957.—New genera and species of Acarina from bats from New Guinea, Philippines
and Australia. Trans. roy. Soc. S. Aust., 80: 67-72.
WoOMERSLEY, H., and HEASLIP, W. G., 1943.—The Trombiculinae (Acarina) or itch-mites of the
Austro-Malayan and Oriental regions. Trans. roy. Soc. S. Aust., 67: 68-142.
ZuMPT, F., 1950.—A new blood-sucking mite from the South African springhare. J. ent. Soc.
S. Afr., 13: 83-86.

241

A TURBIDIMETRIC METHOD FOR ESTIMATING THE NUMBER OF NEMATODE
LARVAE IN A SUSPENSION.

By C. D. Buaxe, Plant Pathology Laboratory, Faculty of Agriculture,
University of Sydney.*

(Two Text-figures. )
[Read 24th September, 1958.]

Synopsis.

A turbidimetric method for estimating the number of nematode larvae in an aqueous
suspension stabilized with 0:5% carboxymethyleellulose is described. A simply constructed,
modified Peter’s counting chamber is used to count the nematode larvae in a number of 10 ml.
standardizing suspensions. The light absorption due to the turbidity of each of these
suspensions is measured with a Hilger “Spekker” absorptiometer. A standard curve relating
the logarithm of absorption and the number of nematodes in the suspension is constructed
and used subsequently to predict the number of nematodes in a suspension, the logarithm
of absorption of which has been measured.

The results are rapidly obtained, amenable to statistical analysis and for Anguwina agrostis
(Steinbuch) Goodey, the percentage error in the prediction from the standard curve was less
than 5%.

INTRODUCTION.

During the course of an investigation into the aetiology of the plant parasitic
nematode Anguina agrostis (Steinbuch) Goodey the need arose for a rapid yet accurate
method for determining the number of larvae in a suspension. Conventional counting
methods depend for their accuracy on the precision with which very small volumes
of the suspension can be measured, the number of larvae in this volume estimated,
and the minimizing of sampling errors. Korsten et al. (1953) have described a colori-
metric technique for determining quantitatively the concentration of the stem eelworm
Ditylenchus dipsacit (Kitihn) Filipjev. This method has greatly reduced the laborious
task of counting, but in the construction of the standard transformation curve used
in the estimation, dilution counting is employed and hence the considerable errors
associated with dilution counts tend to be perpetuated in colorimetric estimations
using this standard curve.

A counting method is outlined employing a modification of Peter’s counting slide
(Goodey, 1957), which it is suggested allows the construction of a transformation
curve which can be used to estimate rapidly and more accurately the number of
larvae in a suspension whose turbidity has been measured.

EXPERIMENTAL.

Ten 10 ml. suspensions of the larvae in a 0:5% solution of pure carboxymethy]l-
cellulose (¢c.m.c.) were prepared, the eelworm concentrations of which were representa-
tive of the range over which the curve was to be used for estimating larval numbers.
The logarithm of the light absorption due to the turbidity of the suspension was
measured for each suspension using a Hilger ‘“Spekker’” absorptiometer fitted with
H508 filters. Hach reading was standardized against a solution of the suspending fluid.

The number of larvae in each suspension whose light ahsorption has been measured
must be determined by a direct counting method. A simply constructed modification
of Peter’s counting slide (Goodey, 1951) was introduced, to facilitate direct counting

* Present address: Biological Branch, New South Wales Department of Agriculture,
Sydney.

PROCEEDINGS OF THE LINNEAN Society or New SoutH WALES, 1958, Vol. lxxxiii, Part 3.

242 ESTIMATING THE NUMBER OF NEMATODE LARVAE IN A SUSPENSION,

and to minimize the error associated with the count. The counting slide consists
essentially of three parts:
(1) A microscope slide onto which two fine lines have been etched 12-5 mm. apart.
(If concentrated suspensions are being counted, an additional centre line can
be drawn.)
(2) Two traverse strips 10 mm. wide cut from a slide uniformly 1 mm. thick
cemented across the first slide, one at each end.
(3) A third slide is cut longitudinally 14 mm. wide and cemented to the cross-
pieces made by the portions of the second slide in a position as indicated in
Figure 1.
The stabilized suspension containing the nematodes was introduced between the
two slides in sufficient quantity to fill the volume between the two slides completely.
The number of larvae enclosed by the etched lines in strips selected at random were

Fig. 1.—Counting slide. A, General view; B, Side view; C, Plan view. For description
see text.
counted using a tally counter as the slide was moved across the low power field of
the microscope by means of a mechanical stage. By proper adjustment of the viscosity
of the c.m.c. suspending medium ail the larvae can be brought into the same optical
plane, thereby improving the rapidity and ease of counting.

From each standardizing suspension five slides were prepared and five strips
counted at random on each slide. The mean number of larvae per strip was contained
in a volume of fluid which equalled (the distance between the etched lines) x (the
depth of the cell) x (the diameter of the microscope field). Knowing the original
volume of the suspension, the number of larvae in the suspension can be calculated.

Using the logarithm of absorption and the number of larvae in each standardizing
suspension a standard transformation curve was constructed as shown in Figure 2
and used subsequently to predict the number of larvae in a suspension whose logarithm
of absorption only had been measured.

An implicit assumption of the method outlined is that the larvae are uniformly
distributed throughout the suspension whose absorption is measured. To judge the
effectiveness of ¢c.m.c. in stabilizing the lervae suspension, the number of larvae in

BY C. D. BLAKE. 243

five strips were counted at random on each of five slides. One set of slides was
prepared from a suspension where the suspending fluid was 0°5% ec.m.c. and the other
set where the suspending fluid was distilled water. If the larvae are uniformly
distributed then the slide counts will have a Poisson distribution and the index of
dispersion will be distributed as a x? (Fisher, 1948). The indices of dispersion have

TABLE 1.
A Comparison between Direct Slide Counting and Turbidity Measurement Methods for
Estimating the Number of A. agrostis Larvae in a Suspension.

Mean Number of Larvae per 16-875 mm.?*

Logarithm of of Suspension Determined by Percentage
Absorption. Error.
Direct Slide | Turbidity
Counting.+ Measurement.
0-045 2-60 2-69 3:3
0-095 5-35 5-51 2-9
0-215 13°45 13-93 3085)

* Volume of fluid per strip counted=1 x 12-5 1-35=16-875 mm.°
j Mean number of larvae per strip in 25 strips counted.

been calculated from the larvae counts per slide and with water as the suspending
medium was 39-358, P(x%) >39-358 is less than 0-01 and with c.m.c. as the suspending
medium was 3-218, P(x.) >3:218 falls between 0-5 and 0-7.

These results indicate that water was an unsatisfactory suspending medium. Also
there was the suggestion that the ¢c.m.c. suspension gave counts conforming to a
Poisson distribution. To establish c.m.c. as a satisfactory medium it would be necessary

in thousands per 19 ml.

Nematode larvae

on oe Os Oa OS

Logarithm of light absorption.

Fig. 2.—Standard transformation curve, showing the relationship between the number of
nematode larvae per 10 ml. of suspension and the logarithm of the light absorption. Regression
equation: Y = —168-3 + 39104X%. Where Y = the estimated number of larvae per 10 ml. of
suspension and X = measured logarithm of absorption of the suspension. Correlation
coefficient (r) = 0:96 (significant 0:1% level).

to have a large number of observations of five sets of larvae counts, to compute Fisher’s
index of dispersion from these data and then to match the distribution of these values
with that of the x2, distribution.
DISCUSSION.
The evidence presented confirms the findings of Korsten et al. (1953) that
absorptiometric methods can be used to estimate reliably the number of nematode

B

244 ESTIMATING THE NUMBER OF NEMATODE LARVAE IN A SUSPENSION.

larvae in a suspension. Figure 2 shows the amount of light absorbed by a suspension
is linearly related to the number of larvae of A. agrostis in the suspension. Such a
relationship allows the construction of a standard transformation curve which can be
used to predict the number of larvae in a suspension whose turbidity has been
measured. The reliability of this technique is shown by the results in Table 1.

In the design of the technique an attempt has been made to minimize counting
errors. Percentage errors as small as those given in Table 1 are possible for a
number of reasons:

(1) The use of c.m.c. as the suspending fluid reduces variability in the distribution
of larvae in the suspension and hence the light absorption which is measured
in a part of the suspension is representative of the whole suspension.

(2) When the counting slide is used large samples each of approximately 2 ml.
are taken randomly from the suspension. The large sampie taken from a
stabilized larval suspension reduces sampling errors.

(3) The accuracy with which the direct count is made using the counting slide
provides a sound basis for estimating the accuracy of the indirect method as
has been done in Table 1.

Satisfactory techniques have been described for making total counts of mixed
nematcde populations (Goodey, 1957) and which in addition enable the frequency of
the component genera in a population to be determined. Where a reliable estimate of
the total number of nematodes present is required, indirect turbidimetric counting can
only usefully be employed where a large and uniform nematode population is involved.

Acknowledgements.

The work described was carried out under the supervision of Associate Professor
N. H. White, of the University of Sydney, whose guidance and co-operation are
acknowledged. The counting slide described was constructed by Mr. H. V. Whitlock,
of McMaster Animal Health Laboratory, C.S.I.R.O., Sydney, and the statistical work
was done by Mr. C. H. Gray, Biometrician, Department of Agriculture, Sydney. I wish
to thank Associate Professor J. M. Vincent and Dr. Lilian R. Fraser for reading the
manuscript.

References.

FisHpr, R. A., 1932.—Statistical Methods for Research Workers (4th Edition). Oliver and

Boyd, Edinburgh.
Goopry, J. B., 1957.—Laboratory Methods for Work with Plant and Soil Nematodes. Pp. vi

+ 47. H.M. Stationery Office, London.

Kxorsten, L. H. J., SIEBEN, J. W., and Voskuy.L, K., 1953.—A Colorimetric Determination of
the Number of Eelworms in a Suspension. Huphytica, 2 (2): 135-138.

245

THE OVIPOSITION BEHAVIOUR OF AEDES AUSTRALIS (ERICKSON)
(DIPTERA, CULICIDAE).

By A. K. O’Gower, School of Public Health and Tropical Medicine,
University of Sydney.

[Read 24th September, 1958. ]

Synopsis.

The selection of an oviposition site by A. australis was influenced by the salinity of
the water and by the physical properties of the surface of the water container, namely, its
texture, reflectance and the presence of either a free, water surface or a moist, porous
surface. Fresh water was preferred to a range of salinities, a rough surface to a smooth one,
lower reflectances to higher reflectances, and a moist, porous surface to a free, water surface.

From a study of the interaction of these factors it was concluded that the preference
for a moist, porous surface over a free, water surface so influenced the oviposition behaviour
of this species that texture, reflectance and salinity had little effect upon it.

INTRODUCTION.

The occurrence of the larvae of Aédes australis (Erickson) (= concolor Taylor)
in the Sydney area in sandstone rock pools in which the salinity of the water varies
(Mackerras, 1926; Woodhill, 1936) must be due to its oviposition behaviour being
influenced by some factor of the environment other than salinity, because Woodhiil
(1941) has shown that fresh water is preferred to filtered sea water for oviposition.

Of the environmental factors which could influence the oviposition behaviour of
A. australis, those investigated were the salinity of the water, the reflectance of the
Oviposition site, the texture of its surface, and whether the surface was moist and
porous or a free water surface.

The factors reflectance, texture and either a moist, porous surface or a free, water
surface have been found to influence the oviposition behaviour of Aédes scutellaris
scutellaris (Walker) (O’Gower, 1955), which breeds in tree holes and coconut husks
(Penn, 1947); Aédes aegypti (L.) (Beckel, 1955; O’Gower, 1957; Wallis, 19540) which
occurs in rain water storage tanks, ete. (O’Gower, 1956); Aédes scutellaris katherinensis
Woodhill (O’Gower, 1957) which presumably breeds in similar situations as A. scutellaris
scutellaris; and Aédes albopictus (Skuse) (O’Gower, 1957) which breeds in tree-holes
and coconut husks (MacDonald, 1956). Salinity, however, usually has an inhibiting
effect upon oviposition behaviour (Wallis, 1954b; Woodhill, 1941), but Mathis (1934)
found Aédes meriae (Kd. and Ht. Sergent) (= desbansi (Seguy)) which occurs in salt
water rock pools, preferred saline water to fresh water for oviposition.

Other environmental factors such as pH, organic pollution and vegetation have
been found to influence oviposition behaviour (Lund, 1942; Manefield, 1951; Russell
and Rao, 1942), but these were not studied.

EXPERIMENTAL TECHNIQUE.

Pupae of A. australis were collected from salt water rock pools in the Sydney
metropolitan area, and the emerged adults maintained in a controlled temperature
and humidity room operating at 27 + 1°C. and 75 + 4 per cent. relative humidity, and
with fluorescent light twelve hours (6 a.m. to 6 p.m.) in every twenty-four (Backhouse
and O’Gower, 1956).

~The mosquitoes were given selections between two oviposition sites, the surfaces
of which varied in reflectance (black or grey), texture (rough or smooth), and
either a moist, porous surface or a free, water surface. Hither saline (one-half per

PROCEEDINGS CF THE LINNEAN Sociery or NEw SouTH WALES, 1958, Vol. Ixxxiii, Part 3.

246 OVIPOSITION BEHAVIOUR OF AEDES AUSTRALIS,

cent. by weight sodium chloride) or tap water was used in these oviposition sites.
A further range of salinities of one per cent., two per cent., and four per cent. saline
was also used for some experiments. However, the combination of a saline solution
and either a moist, porous surface or a free, water surface gave a choice between
either a saline, moist, porous surface and a free, fresh, water surface; or between a
fresh, moist, porous surface and a free, saline, water surface. In the former case
there was an increase in salinity of the moist, porous surface due to evaporation, and
in the latter case, because of an almost absolute preference for a moist, porous surface
rather than a free, water surface (experiment 1), the influence of salinity could not
be assessed. Therefore, as all combinations of the environmental factors involving
both salinity and either a moist, porous surface or a free, water surface were either
experimentally uncontrollable or else added little information on behaviour, they have
been omitted from this paper except in experiment 17. However, because the texture
of the surface has been found to be so important, and because the salinity of the
water cannot be ignored in determining oviposition behaviour, the interaction of these
two factors has been studied in further detail, by giving the mosquitoes selections
between oviposition sites of different textures and salinities.

The oviposition sites in experiments 1, 2, 3, 8, 9, 15 and 17 were formed by placing
into 9 cm. diameter Petri dishes either a pad of absorbent cellulose cotton with a
filter paper of the required texture and reflectance on top of it, or a filter paper of
the required reflectance on the bottom of the dish. Hither tap water or saline was
was then added to both containers, until the surface of the former was wet and
glistening, and the water level in the latter was as high as the wet surface of the
former. In experiments 4, 5, 6, 7, 10, 11, 12, 13, 14 and 16, the oviposition sites were
formed by folding 11 cm. diameter filter papers into cones, placing them in 50 ml.
beakers and adding either saline or fresh water until the water levels in the cones
were half their vertical heights.

The filter papers* had surfaces which were (i) black and smooth, (ii) black and
rough, (iii) grey and smooth, and (iv) grey and rough. Black “Tintex” dye was
used to obtain papers of low reflectance. A “General Electric’ spectrophotometer was
used to measure the reflectances of these papers when wet. At a wave-length of
555 millimicrons the reflectancest of these papers when wet were: (i) black smooth,
3%; (ii) black rough, 3%; (iii) grey smooth, 7%; and (iv) grey rough 6%. 2

Seven replicates of each experiment were done, and the variances were calculated
from the mean percentages of each. The significance of various preferences was
estimated by means of the t-test.

RESULTS.

1. Comparison between free water and a moist surface—In experiment 1 the
mosquitoes were given a choice between two water containers of similar reflectances.
One dish had a free, water surface, the other a moist, porous surface. The moist,
porous surface was significantly preferred (p < 0-001) to the free, water surface
(Table 1).

2. The effect of refiectance—In experiment 2 the mosquitoes were given a choice
between two water containers of similar textures but different reflectances. The
preference for these containers was inversely related (p < 0-001) to their reflectances
(Table 1).

3. The effect of textwre-—In experiment 3 the mosquitoes were given a choice
between oviposition sites of similar reflectances but different textures. A rough surface
was significantly preferred (p < 0-001) to a smooth surface (Table 1).

4. The effect of salinity.——In each of experiments 4, 5, 6 and 7 the mosquitoes were
given a choice between oviposition sites containing either fresh water or salinities
of 3%, 1%, 2% or 4%. The fresh water was significantly preferred (p < 0-001) to
the full range of salinities (Table 1).

* (i) Whatman, No. 5; (iii) Allnutt and Sons, No. Bl; (ii) and (iv) Allnutt and Sons,
No. D3.

+ Measured by C.S.I.R.O., National Standards Laboratory, University of Sydney.

BY A. K. O’GOWER. 247

TABLE 1.
Summary of Results.

Mean
Experiment Re- Number | Percentage Value of Prob-
Number. Surface. flectance | of Eggs of Eggs | Variance. Ste ability.
% Deposited.| Deposited.

A. The Influence of Water, Reflectance, Texture and Salinity on Oviposition by A. australis.

i Free water 7 95 2 3 100 <0-001
| Moist porous iG 4,598 98
2 Black .. 3 4,422 68 Pho5) 50 <0-001
Grey 7 2,146 32)
3 Rough 3 6,124 80 69 13-6 <0-001
Smooth 3 1,621 20 |
4 4% Saline water i 4,291 33 24 13 <0-001
Fresh water .. 7 7,443 67
5 1% Saline water 7 2,558 20 3 65 <0-001
Fresh water .. 7 10,080 80
6 2% Saline water 7 877 12 5 63 <0-001
| Fresh water .. 7 6,686 88
7 4% Saline water 7 504 4 3 99 <0-001
Fresh water .. U 10,256 96

TABLE 2.
Summary of Results.

Mean
Experiment Re- Number | Percentage Value of Prob-
Number. Surface. fiectance of Eggs of Eggs | Variance. moi Uinate ability.
% Deposited.| Deposited.

B. The Combined Influence of Water and Reflectance on Oviposition by A. australis.

8 Black free water .. 5 3 53 2 1 183 <0-001

Grey smooth moist porous 5,578 98

“I

C. The Combined Influence of Reflectance and Texture on Oviposition by A. australis.

| |
9 Grey rough .. an inal 6 5,897 78 147 | 8-6 <0-001
Black smooth ais mre 3 2,257 22 | |
|

D. The Combined Influence of Reflectance and Salinity on Oviposition by A. australis.

66 31 10-6 | <0-001

10 | Black 4% saline 50 oo | 3 8,100
34 |

| Grey fresh water .. fie 7 3,906

|

E. The Combined Influence of Texture and Salinity on Oviposition by A. australis.

11 Rough £% saline 3 12,562 73 38 13-3 <0-001
Smooth fresh water 3 4,944 27
12 Rough 1% saline 3 8,664 75 18 22-2 <0-001
Smooth fresh water 3 2,985 25
13 Rough 2% saline 3 8,737 78 8 By (52) <0-001
Smooth fresh water 3 2,357 22
14 Rough 4% saline 3 2,215 26 3 52 <0-001
Smooth fresh water 3 6,290 74
|

248 OVIPOSITION BEHAVIOUR OF AEDES AUSTRALIS,

5. The combined effects of reflectance and free water or a moist surface—In
experiment 8 the mosquitoes were given a choice between two water containers, one
with a free, water surface and a black background, the other with a grey, smooth,
moist, porous surface. The grey, smooth, moist, porous surface was significantly
preferred (p. < 0-001) to the free, water surface (Table 2), but this preference was
not different from that for a moist surface over a water surface of equal reflectance
(experiment 1).

6. The combined effects of reflectance and texture—In experiment 9 the choice was
between a water container with a grey, rough, moist surface and a water container
with a black, smooth, moist surface. The grey, rough surface was significantly preferred
(p < 0:001) to the black, smooth surface (Table 2), but this preference was not
different from that for a rough surface over a smooth one, both of equal reflectance
(experiment 3).

TABLE 3.
Summary of Results.

! Mean
Experiment Re- Number | Percentage Value of Prob-
Number. Surface. flectance of Eggs of Eggs | Variance. ib ce ability.
% Deposited.| Deposited.

F. The Combined Influence of Texture, Reflectance and Water on Oviposition by A. australis.

i

15 Black fresh water .. 56 | 3 87 2 2 127 <0-001
Grey rough moist .. Ae 6 7,181 | 98

1

G. The Combined Influence of Texture, Reflectance and Salinity on Oviposition by A. australis.

16 Black smooth fresh water .. 3 1,496 22 43 16 <0-001
Grey rough saline water .. 6 7,155 7

H. The Combined Influence of Texture, Reflectance, Water and Salinity on Oviposition by A. australis.

17 Black fresh water .. ms 3 25 1 0 ioe) <0-001
| Grey rough moist saline .. | 6 7,404 99
| i

7. The combined effects of reflectance and salinity.—In experiment 10 the mosquitoes
were given a choice between a water container with a black background with 4%
saline in it and a container with a grey background with fresh water in it. ‘The
container with the black background holding 4% saline was significantly preferred
(p < 0-001) to the container holding fresh water and with a grey background (Table 2).
However, this preference was not different from that for a black surface over a grey
surface (experiment 2).

8. The combined effects of salinity and textuwre—In each of experiments 11, 12,
13 and 14 the mosquitoes were given a choice between two water containers, one with
a rough surface holding either 3%, 1%, 2% or 4% saline, the other with a smooth
surface holding fresh water. In experiments 11, 12 and 13 the container with the
rough texture holding either 4%, 1% or 2% saline was significantly preferred
(p < 0-001) to the smooth-surfaced container holding the fresh water. In experiment 14
the smooth-textured container holding fresh water was _ significantly preferred
(p < 0-001) to the rough-textured container holding 4% saline (Table 2).

9. The combined effects of texture, reflectance and free water or a moist surface.—
In experiment 15 the mosquitoes were given a choice between two water containers,
one with a free, water surface with a black background, the other with a grey, rough,
moist, porous surface. The grey, rough, moist, porous surface was significantly
preferred (p < 0-001) to the free, water surface with the black background (Table 3).

BY A. K. O’GOWER. 249

However, this preference was not different from the preference for a moist surface
over a water surface irrespective of texture or reflectance (experiments 1 and 8).

10. The combined effects of texture, reflectance and salinity.—In experiment 16 the
mosquitoes were given a choice between a water container with a black, smooth surface
holding fresh water and a container with a grey, rough surface holding 4% saline.
The grey, rough-surfaced container holding saline water was significantly preferred
(p < 0:001) to the black, smooth-surfaced container holding fresh water (Table 3).
However, this preference was not different from the preferences for rough surfaces
over smooth ones irrespective of reflectances (experiments 3 and 9).

11. The combined effects of texture, reflectance, salinity and water or a moist
surface—Because only 4% saline in a container with a rough-textured surface was
less attractive than fresh water in a smooth-iextured container (experiments 11, 12,
13 and 14), the mosquitoes were given a choice in experiment 17 between a fresh
water surface with a black background and a 4% saline, moist, grey, rough surface.
The saline, moist, grey, rough surface was significantly preferred (p < 0-001) to the
fresh water surface with the black background (Table 3), but this preference was
not different from that for a moist surface over a water surface (experiment 1),
irrespective of reflectance (experiment 8), texture (experiment 15) or salinity (experi-
ments 4, 5, 6 and 7).

DISCUSSION.

Woodhill (1941) found A. australis preferred fresh water to a range of dilutions
of filtered sea water, and the present investigation, using a range of dilutions of
sodium chloride solutions, was in close agreement with this. Thus the preference
for fresh water over saline water was due to a chemotactile response by the gravid
females to sodium chloride in solution (see Wallis, 19540, for a detailed investigation
on the influence of salinity on oviposition).

However, because such behaviour could not explain the distribution of the larvae
of this species in salt water rock pools, other factors of the environment were also
studied. Thus, when the influences of the environmental factors of texture, reflectance,
salinity and either a free, water surface or a moist, porous surface were compared,
the preference for a moist surface over a water surface was found to be greater
than the preferences for rough surfaces over smooth surfaces, for low reflectances
over higher reflectances, and for fresh water over saline water.

Comparing the preferences of A. australis with those of A. aegypti (O’Gower, 1957)
and A. scutellaris scutellaris (O’Gower, 1955) one finds:

A. aegypti. A. australis. A. scutellaris.
Preference for rough over smooth .. ah 59% to 41% 80% to 20% 60% to 40%
Preference for black over grey as er 73% to 27% 68% to 32% 70% to 30%
Preference for water over moist at hs 75% to 25% 2% to 98% 54% to 46%
Preference for fresh over saline wy ae 74% to 26%* 67% to 338% Not available

* Mean of percentages from O’Gower (unpublished), Wallis (1954b) and Woodhill (1941).

It can be seen from these figures that while the preference for a water surface
over a moist surface was only slight for A. scuwtellavis, with A. aegypti the preference
was most decided, but with A. australis the preference was reversed and almost
absolute. This difference in behaviour of the two former species appears to be correlated
with their different larval distributions (O’Gower, 1957). However, the similarity in
behaviour of A. aegypti and A. australis with regard to salinity cannot be correlated
with their very different larval habitats.

A. meriae breeds in salt water rock pools and, because it lays its eggs on the
surface of the water (Mathis, 1929), the preference for salt water over fresh water
(Mathis, 1934) is understandable. <A. australis breeds in the Sydney area only in
sandstone, salt water, rock pools (Mackerras, 1926) and its eggs have been found
on the rock surface by the author either at or above the water line. Thus the almost

250 OVIPOSITION BEHAVIOUR OF AEDES AUSTRALIS.

absolute preference for a moist surface over a water surface, and the decided preference
for a rough surface over a smooth one, help explain the unimportance of salinity in
the oviposition behaviour of A. australis.

This explanation becomes more logical when the interaction of these environmental
factors is studied, for the attractiveness of a rough surface masks both the unattractive-
ness of saline water (except when the salinity equals that of sea water) and the
attractiveness of lower reflectances (experiments 11, 12, 13, 14 and 9), whilst the
preference for a moist surface over a water surface so influences oviposition behaviour
that texture, reflectance and salinity do not affect it (experiment 17).

Such behaviour can explain the occurrence of the larvae of A. australis in salt
water rock pools, but not their absence from fresh water rock pools, and although it
is possible that other environmental stimuli may influence the selection of an oviposition
site by this species, this investigation was limited to a study of the influence of
salinity and of the physical properties of the surface of a water container upon the
Oviposition behaviour of A. australis.

Acknowledgements.

The author wishes to thank Mr. D. J. Lee and Dr. P. Robertson, of the School of
Public Health and Tropical Medicine, University of Sydney, and Dr. A. R. Woodhill,
of the Department of Zoology, University of Sydney, for helpful criticisms in the
preparation of this paper, which is published with the permission of the Director-
General of Health, Canberra, A.C.T.

References.

BACKHOUSE, T. C., and O’Gowesr, A. K., 1956.—An inexpensive, easily constructed, controlled
temperature and humidity room for maintaining insect colonies. Bull. ent. Res., 47: 67-71.

BECKEL, W. H., 1955.—Oviposition site preference of Aedes mosquitoes (Culicidae) in the
laboratory. Mosquito News, 15: 224-228.

LuNnp, H. O., 1942.—Studies on the choice of a medium fer oviposition by Anopheles quadri-
maculatus Say. J. nat. Malar. Soec., 1: 100-111.

MacDoNnALD, W. W., 1956.—A mosquito survey of Kuala Lumpur airport with special reference
to Aédes aegypti. Med. J. Malaya, 10: 232-245.

MANEFIELD, T., 1951.—Investigations of the preferences shown by Aédes (Stegomyia) aegypti L.
and Culex (Culex) fatigans Wied. for specific types of breeding water. Proc. LINN.
Soc. N.S.W., 76: 149-154.

MACKERRAS, I. M., 1926.—The mosquitoes of the Sydney district. Awst. Nat., 6: 33-42.

MarTuis, M., 1929.—Biologie d’un moustique cdtier du Var. Aedes desbansi Seguy 1923. Bull.
Soc. Path. exot., 22: 179-183.

, 1934.—Influence du NaCl sur le determinisme de la ponte chez un moustique cdtier
du Var. Aedes desbansi Seguy 1923. Bull. Soc. Path. exot., 27: 757-759.

O’Gowrr, A. K., 1955.—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). Proc. Linn. Soc. N.S.W., 79: 211-218.

, 1956.—Control measures for Aédes aegypti: Surveys in northern Australia. Health,

Canberra, 6: 40-42.

—, 1957.—The influence of the surface on oviposition by Aédes aegypti (L.) (Diptera,

Culicidae). Proc. Linn. Soc. N.S.W., 82: 240-244.

—, 1957.—The influence of the surface on oviposition by Aédes albopictus (Skuse) and
Aédes scutellaris katherinensis Woodhill (Diptera, Culicidae). Proc. LINN. Soc. N.S.W.,
82: 285-288.

PENN, G. H., 1947.—The larval development and ecology of Aédes (Stegomyia) scutellaris
(Walker, 1859) in New Guinea. J. Parasit., 33: 43-49.

RUSSELL, P. F., and RAo, R. T., 1942.—On relation of mechanical obstruction and shade to
ovipositing of Anopheles culicifacies. J. exp. Zool., 91: 303-329.

WALLIS, R. C., 1954a.—Observations on oviposition of two Aedes mosquitoes. Ann. ent. Soe.
Amer., 47: 393-396.

———, 1954b.—A study of oviposition activity of mosquitoes. Amer. J. Hyg., 60: 13-16.

WoopHILL, A. R., 1941.—The oviposition responses of three species of mosquitoes (Aédes
(Stegomyia) aegypti Linnaeus, Culex (Culex) fatigans Wiedemann, Aédes (Pseudoskusea)
concolor Taylor) in relation to the salinity of the water. Proc. LINN. Soc. N.S.W.,
66: 287-292.

—————, 1936.—Observations and experiments on Aédes concolor Taylor. Bull. ent. Res.,
27: 633-648.

PALAEOZOIC GEOLOGY OF THE COOLEMAN CAVES DISTRICT,
NEW SOUTH WALKS.

By N. C. Srevens, Department of Geology, University of Queensland.
(Plates iii-iv; one Text-figure.)
[Read 24th September, 1958.]

Synopsis.
Stratigraphy and structure of Silurian strata, including fossiliferous limestones of
Cooleman Caves, are described, as well as Ordovician sedimentary rocks and Devonian lavas.
Brief notes on granitic rocks and minor intrusives are given.

INTRODUCTION.

The limestone of Cooleman (formerly Coolalamine) Plain was first reported on by
the Rey. W. B. Clarke (1860), and fossils collected by Clarke from this area were
described by de Koninck (1876-77). Further geological notes were published in a
report on the caves by Leigh and Etheridge (1894).

But it was not until recent years that any attempt was made to map the district
geologically. Ivanac and Glover (1949) and Walpole (1952) carried out reconnaissances
and produced maps, and in 1956 Snowy Mountains Hydro-Electric Authority geologists
began detailed work near Pocket Saddle in connection with the proposed Tantangara
Reservoir (Newbery, 1956). The writer extended mapping to the Cooleman and
Currango Plains and adjacent ranges during the summers of 1956-7 and 1957-8 (see
map, Pl. iii).

PHYSIOGRAPHY.

The western part of the area consists of broad, open plains with a general elevation
of 4,200 feet, bounded by wooded ridges which rise gradually to a maximum of 5,200
feet (Pl. iv, fig. 1). Cooleman Plain, remarkably level where undissected, is drained
by Cave Creek, a major tributary of the north-flowing Goodradigbee River. Close to
the southern margin of the area is the Murrumbidgee-Goodradigbee divide. It is a
wooded, east-west ridge, broken by two low saddles. The western saddle, Blue Waterhole
Saddle, lies between Cooleman and Currango Plains, the eastern one, Pocket Saddle,
which is almost flat, links Currango Plain with the Goodradigbee Valley.

The Goodradigbee River (known as Murray Creek in its upper reaches) rises in
the Bimberi Range and flows through granite in a general north-north-west direction.
On entering resistant Ordovician quartzites, it flows west through a gorge and then
pursues a northerly course through less resistant Silurian slates and limestones.

In this way it passes close to Pocket Saddle and has cut a valley with a base over
200 feet below the saddle.

The Goodradigbee Valley and the country east of the Blue Waterhole, with V-shaped
valleys and steep-sided gorges, contrasts with the “plain and marginal ridge” topography
of the rest of the area (PI. iv, figs. 3, 4).

Evidence of the 4,200 feet level is still visible on either side of the Goodradighbee
Valley downstream from Pocket Saddle. Much of this level has been preserved by a
covering of lavas, which still remains on the Rolling Grounds Spur.

Seattered outliers of Devonian lavas indicate that the topography of Cooleman
Plain is more or less the same as in Devonian times, the level limestone plain having
been formed before the outpouring of the lavas which gave rise to the surrounding
ridges.

Remnants of a higher level between 5,000 feet and 5,200 feet are widespread,
including Black Mountain and the Gurrangorambla Range north-west of Blue Waterhole
Saddle.

PROCEEDINGS OF THE LINNEAN Socrery or NEw SouTH WALES, 1958, Vol. Ixxxiii, Part 3.

252 PALAEOZOIC GEOLOGY OF THE COOLEMAN CAVES DISTRICT,

STRATIGRAPHY.
Ordovician.

Rocks classed provisionally as Ordovician are found on the eastern and western
margins of the area. The more important exposures are on the east, where quartzites
and slates extend south from the Rolling Grounds Spur and are known to be
continuous with the Ordovician Nungar Beds (Newbery, 1956; Stevens, 1958) 10 miles
to the south.

The beds consist mainly of quartzite and slate with sandy slates and siltstones.
The slates are usually buff or grey and the quartzites white, but black cherty types
are not uncommon. The rocks are strongly cleaved and show much contortion within
the beds. On the east they are bounded by granite, and to the west there is an
indistinct junction with Silurian slates of the Pocket Beds.

Another north-south belt of sediments of probable Ordovician age occurs in the
north-west of the district, apparently underlying Silurian limestone. These sediments
are mostly fine-grained, poorly bedded sandstones, similar to rocks regarded as upper-
most Ordovician in the Nungar area.

Silurian.

The majority of the sedimentary rocks are of Silurian age, including large limestone
belts, with formations of cherts, fine-grained sandstone, siltstone and shale.

Rapid facies changes from west to east complicate stratigraphical nomenclature
and sequence determination. The Cooleman Limestone, which is a thick and important
formation in the west, is represented by the Pocket Beds (mainly slate and limestone)
in the east, while the overlying Blue Waterhole Beds change from siltstone, shale
and sandstone in the west through cherts and sandstone to cherts (and perhaps
limestone) as one proceeds east.

TABLE 1.
Silurian sequence.

Area West Central East

Wilkinson Limestone

BS

Blue Waterhole Beds

bo co

(Siltstone, shale, (chert, sandstone) Blue Waterhole Beds
sandstone) (chert, limestone)
i, Cooleman Limestone 1. Pocket Beds

(a) Cooleman Limestone.

This formation, named by Walpole (1952), is exposed on Cooleman Plain out-
cropping over an irregular, arcuate area, which has been strongly dissected by Cave
Creek in the north-central parts to form Cooleman (Clarke’s) Gorge.

The limestone has been folded into broad, plunging folds and appears to be the
oldest Silurian formation. Near Blue Waterhole Saddle a few feet of chert and
micaceous tuff underlie the limestone, but it is not clear whether more iimestone
underlies these sediments and is covered or intruded by adjacent granophyre. In
most other places on the S.W. margin the limestone is bordered by younger dacite
lavas or granophyre.

The limestone is generally massive with occasional chert beds and thin shaly
layers; it is, like most lower Palaeozoic limestones in New South Wales, quite pure,
except for dolomitized zones which tend to follow joint planes. Near intrusive rocks
and lava flows it has been silicified, and parts of the original plain surface are capped
by silicified rock and also by ferruginous sediments and ironstone derived from
weathering of the limestone. Sink holes and caves are numerous and little water
flows in creeks upstream from the Blue Waterhole.

Fossils are abundant in many places between Spencer’s (White’s) Hut and Jones’
Hut. Brachiopods, stromatoporoids and corals are most prominent, but some gastropods
are found towards the base of the formation, also bryozoa and nautiloids. Large
bivalves occur in narrow beds north of Blue Waterhole Saddle and north-east of
Coolamine.

BY N. C. STEVENS. 253

Conchidium strictum is widespread and abundant, and, with Favosites, Striatopora
and (probably) Pycnostylus, gives a Silurian age to the formation.

In the Blue Waterhole area the limestone is very granular and lacks well-preserved
fossils.

(0) Blue Waterhole Beds.

These beds overlie the Cooleman Limestone and occur in a large, faulted basin
around Coolamine and in two smaller outcrops north and south of the Cooleman
Gorge.

At Harris’ Dam the lowest beds of the basin structure appear to be interbedded
with the limestone. They are micaceous siltstones or fine-grained sandstones, containing
flat lamellibranchs and traces of nautiloids. Dark, banded cherts with shales and
siltstones overlie the basal beds on Cave Creek, south of Coolamine. Slump structures
are common in this locality, and the shallow-water origin of some of the beds is shown
by mud-cracks at several horizons.

Above these rocks are more siltstones with narrow beds of black chert containing
irregular cavities, representing moulds of corals such as Heliolites. A compact sand-
stone outcrops in the centre of the basin, south of Coolamine. The formation’s
maximum thickness of about 2,000 feet is in this area.

Isolated outerops of shales and cherts which overlie the Cooleman Limestone north
of the Blue Waterhole have been correlated with the same formation. From siltstone
in one of these outliers small nautiloids have been collected.

On either side of Cooleman Gorge a formation ef bedded cherts with overlying
sandstone dips gently off the Cooleman Limestone.

The cherts are rhythmically bedded and show differential weathering (PI. iv,
fig. 6). They are highly fossiliferous with corals, both rugose and tabulate, pre-
dominating and show varying degrees of silicification. Similar rocks lie between the
Pocket Beds and a massive limestone along Blue Waterhole Creek west of its junction
with the Goodradigbee River.

From these cherts Favosites, Striatopora, Hercophyllum, ? Thamnopora and
(probably) Mycophyllum have been collected.

(c) Pocket Beds.

The Pocket Beds, named by Newbery (1956), consist of shales, slates, limestones,
cherts and tuffs, which outcrop in the Goodradigbee Valley north of Pocket Saddle.

They are found in two areas, separated by younger dacite. In the more southerly
area, slates and phyllites are the main rock types, with some bedded tuffs and a thin
discontinuous bed of limestone. They are bounded on the east by Nungar Beds and
on the west by younger dacite.

The slates are buff-coloured, highly cleaved and in places strongly jointed, resulting
in splintery fragments. Bedding planes are more obvious than in the Nungar Beds.

The limestone of the southern area is also highly cleaved, but contains corals,

including Propora sp., crinoid stems and small ? Pentamerids. The only fossils
recovered from the slates were a single trilobite pygidium and some poorly-preserved
brachiopods.

Because of doubt as to the base and top of the formation in this area a sequence
has not been established.

Further downstream the stratigraphic relations are clearer as cleavage diminishes
and the proportion of limestone and chert increases. Here the Pocket Beds have been
folded into a plunging anticline.

Slates, showing clearly the relation between bedding and cleavage, occur in the
core of the fold. On the north-west limb thin limestones are interbedded with slate
at a higher horizon and predominantly slaty strata are followed by cherty limestones
with minor amounts of slate. Both rock types contain abundant fossils — corals in the
limestones and brachiopods (including Conchidium and Stropheodontids) in the slates.
Trilobite pygidia have been found in both. Striatopora sp., Entelophyllum yassense,

254 PALAEOZOIC GEOLOGY OF THE COOLEMAN CAVES DISTRICT,

Plasmopora heliolitoides, Hercophyllum aff. shearsbyi, Favosites and Heliolites have
been determined from an horizon near the top of the Pocket Beds north-east of Black
Mountain. In addition, there are gastropods, Stropheodontids and straight conical
nautiloids.

(ad) Wilkinson Limestone.

At the top of the Pocket Beds are shales, overlain by cherts of the Blue Waterhole
Beds or by a massive limestone where the cherts thin out completely. The limestone
formation, which might be termed Wilkinson Limestone, is exposed in a steep-sided
gorge (Wilkinson’s Cliffs, Pl. iv, fig. 4) and is separated from the Cooleman Limestone
by a felsitic intrusion.

The two limestones have previously been mapped as a continuous bed (Walpole,
1952), but the dowstream outcrop is now considered a higher horizon, because it
overlies Blue Waterhole Beds (PI. iv, fig. 3), whereas the Cooleman Limestone underlies
these beds.

The only recognizable fossil collected from this limestone is a small, solitary
rugese coral of an acanthophyllid type, which may be Upper Silurian or Lower
Devonian.

? Devonian.

Lavas of acid and intermediate composition form some of the wooded ranges
bordering the Cooleman Plain and remnants rest unconformably on Silurian sediments
on the plain and on the edges of the Goodradigbee Valley. Angles of dip are gentle
and their sheet-like form is indicated by their distribution, occasional columnar
jointing (Pl. iv, fig. 5) and flow structures. As these lavas are post-Silurian and have
been gently folded, their age may lie between Devonian and Permian. Lithologically
similar dacites (Snowy River Porphyries) of probable Lower Devonian age (David,
1950) are known from eastern Victoria.

(a) Rhyolite.

A white rhyolite, porphyritic in quartz and orthoclase, underlies dacite on the
north-east side of Gurrangorambla Range near Harris’ Hut and forms a small outcrop
south of Spencer’s Hut. It has not been found beneath dacite on Cooleman Plain;
being more viscous, it probably did not flow so far.

(bo) Kellys Plain Dacite.

Dacite is the most widely distributed lava and has been named by Newbery (1956),
who studied it on Kelly’s Plain, south of the Murrumbidgee River. It extends
northwards, covering most of Currango Plain, and forms the lower parts of the
Gurrangorambla Range near Harris’ Hut. Outliers are found overlying Silurian lime-
stone on Cooleman Plain and a similar rock occurs north of Pocket Saddle.

Bedded rocks which are probably acid to intermediate tuffs are associated with
the dacite in the Gurrangorambla Range and interbedding of tuffs and several dacite
flows has given rise to terraces on the spurs.

The dacite behaves like a flow rather than an intrusive, except for the occurrence
north of Pocket Saddle where similar dacitic rocks are found in narrow tongues
associated with the Pocket Beds. These are strongly cleaved and extend down into
the Goodradigbee Valley. Quite possibly they are feeders to the flows or are epi-Silurian
intrusives which have not reached the surface.

(c) Rolling Grounds Andesite.

An augite-andesite which, because of its persistent occurrence about the 4,200-foot
level, is thought to be the youngest of the lavas, has been termed the Rolling
Grounds Andesite.

It occurs on the Rolling Grounds Spur and on the Cooleman Plain south and east
of Coolamine. No relations between it and other Devonian lavas have been noted.

The rock is a typical grey porphyritic andesite, and shows flow structure and
columnar jointing. Occasionally a little quartz is present, indicating its dacitic affinity.

BY N. C. STEVENS. 255

INTRUSIVE ROCKS.
Gingera Granite.

Only the western boundary of this granite was mapped for several miles along
the eastern side of the area. It is probably continuous with the granite which makes
the high peaks of Gingera, Bimberi and Morgan along the western boundary of the
Australian Capital Territory.

Lithologically, the types examined resemble rocks of the Murrumbidgee bathylith,
which is generally thought to be of Late Silurian age. The main type is a biotite-
granite which exhibits slight foliation with trend approximately parallel to that of
the cleavage of the country rocks (Nungar Beds). Seme muscovite aplite occurs along
the margin north of Oldfield’s Hut.

Black Mountain Granite and Related Granodiorites.

Black Mountain is composed almost entirely of pink granite, coarse and felspathic
along the lower western slopes, grading upwards into porphyritic pink granite, with
a fine-grained pink acid granite at the summit. No contacts with surrounding rocks
are exposed; granophyre adjoins it on the west and it is probably faulted against
Pocket Beds on the east. The variation from base to summit suggests that the
present summit is not far below the original roof of the stock.

Porphyritic pink granite is also found on the northern side of Blue Waterhole
Creek, associated with marginal granite porphyry.

In the north-west of the area examined, granite, granodiorite and granodiorite
porphyry outcrop near the U-bend in Cave Creek and in the Gurrangorambla Range.
The former intrusion has converted calcareous sediments of Blue Waterhole Beds to
cale-silicate hornfelses. Near its contact with dacite it becomes darker and more
fine-grained, and appears to grade into the dacite.

The granodiorite intrusion to the west is small, and grades southwards into a
larger mass of diorite. Here also the granodiorite is separated from the dacite by
rocks with texture and mineralogy intermediate between the two. A larger mass of
granodiorite forms most of the wooded ridge west of Coolamine.

All these intrusions, with the related diorite described below, appear to be later
than the Silurian sediments and are either associated with the extrusion of dacite or
closely following it. They may be assigned to the late Middle Devonian Tabberabberan
epoch.

Diorite.

The main dioritic intrusion lies to the east and south-east of Blue Waterhole
Saddle. It adjoins dacite to the south (with a transitional hybrid rock at one locality)
and granophyre elsewhere, and is connected by a narrow tongue to a granodiorite-
granodiorite-porphyry mass on the northern side of Seventeen Flat.

It intrudes the south-eastern tip of the Cooleman Limestone, producing some
interesting contact rocks, including vesuvianite-bearing types. The diorite itself is
epidotized near the contact.

Much variation in texture and some variation in composition are shown by this
intrusion. Felspathic veins and irregular areas containing elongated amphibole needles
also occur.

Another dioritic intrusion appears to intrude dacite and rhyolite in the range west
of Harris’ Hut. In each case field evidence (such as variation in the diorite towards
the contact) suggests that the diorites intrude the dacites, despite the lack of evidence
of any former substantial cover.

Gurrangorambla “Granophyre’’.

This name was originally given to the pink, felsitic rock with sparsely distributed
felspar and ferromagnesian phenocrysts which makes up much of the east-west portion
of Gurrangorambla Range.

The western part of this mass slopes east-south-east from Tom O’Rourke’s Peak,
falling 1,000 feet to plain level in a little over a mile. Elsewhere it behaves like a
subhorizontal sheet.

256 PALAEOZOIC GEOLOGY OF THE COOLEMAN CAVES DISTRICT,

A similar rock, with signs of fiow-structure, borders the granite on the western
slopes of Black Mountain and occurs to the south-east, where it is traceable into dykes
which cross the Goodradigbee River.

The texture of the rocks so far examined is not truly granophyric, with irregular
quartz-felspar intergrowths forming small spherical masses. There is no conclusive
evidence to show whether the “granophyre” is extrusive or intrusive.

Minor Intrusions.

Fine-grained, acid dykes intrude the limestone on Cooleman Plain, but have had
very little effect on it. Some show marked flow structure parallel to their walls, and
most contain pyrite. Similar recks intrude the limestone south of Cooleman Gorge.

Acid dykes up to 100 yards wide and a mile or more in length are found in the
Goodradigbee Valley east of Black Mountain. They, and a third, shorter dyke, radiate
from a hill of granophyre.

Andesitie intrusives, in the form of small dykes and sills, occur at a number of
places throughout the area, but basic dykes are relatively rare. Most of the latter
are dolerites.

STRUCTURAL GEOLOGY.

In this section the structure of each of the three groups — Ordovician, Silurian and
Devonian — will be discussed in turn, as each group has different structural charac-
teristics, which may be related to the Benambran, Bowning and Tabberabberan
orogenies respectively.

The overall picture is that of a major synclinal structure, with Silurian and
Devonian rocks preserved between long north-south trending belts of Ordovician
metasediments and granite.

Ordovician.

Little information is available on the structure of the Nungar Beds in this area,
as only their margin was mapped. The beds have been strongly folded, and the folds
may be isoclinal. Dips of bedding planes are steep, mostly to the east, and cleavage
planes dip in the same direction.

Silurian.
(a) Cooleman Limestone and Associated Beds.

The Cooleman Limestone occupies an irregular area due to its varying thickness
and folding. At its western extremity the thickness is only about 400 feet, but it
appears to thicken considerably to a maximum of 2,000 feet in the south-east
(Text-fig. 1).

per E 2

Text-fig. 1.—Sections along the lines ABC and DEF on the map (Plate iii).

West of a north-south line through Blue Waterhole Saddle, dips in the Cooleman
Limestone and the overlying beds suggest a basin structure, but this is complicated
by an anticline in the limestone west of Spencer’s Hut (Pl. iv, fig. 2) and abrupt
changes in strike and dip near the Blue Waterhole. Evidence of strong faulting is
seen at the boundary between Cooleman Limestone and Blue Waterhole Beds north-west
of Spencer’s Hut.

The eastern prolongation of the Cooleman Limestone down the gorge as far as the
felsitic intrusion is also due to a broad anticline which strikes east-south-east and

BY N. C. STEVENS. 257

plunges in that direction. Dips in the limestone and overlying cherts are gentle
(about 10°) in the gorge except ciose to a fault on the south-east margin of the
limestone, where a thin bed of overlying chert dips steeply to the south. This structure
does not continue downstream, probably because of igneous intrusions with associated
faulting.

Several minor folds and faults have been noted in the Blue Waterhole Beds, and
numerous small faults intersect the limestone along Cave Creek and in Cooleman
Gorge. Over most of its outcrop the limestone exhibits strong jointing, which locally
passes into a cleavage.

(ob) The Pocket Beds and Overlying Strata.

The Pocket Beds on the eastern side of Black Mountain have been folded into an
anticline with axial strike varying from north-south to north-east — south-west near the
Goodradigbee River, and plunging to the north-east. North of this axis most dips are
northerly, and the overlying cherts and Wilkinson Limestone dip in this direction.
South of the anticlinal axis the beds assume a north-south strike and easterly dip.
A syneclinal fold appears further east.

The structure is outlined, particularly in the northern limb, by cherty limestones
which are interbedded with the slates. Bedding and cleavage are easily distinguishable
in the northern outcrop, but in the southerly outcrop of the Pocket Beds cleavage
becomes more prominent and bedding less evident until, close to Pocket Saddle, the
degree of dynamic metamorphism approaches that of the adjacent Nungar Beds.

The strike of the cleavage in the Pocket Beds varies as the strike of the bedding
(i.e. from east-west in the north to north-south in the south), and may be up to
10-15 degrees different, but the dip of the cleavage is always steeper than the dip of
the bedding. The direction of dip of the cleavage changes gradually on passing from
the northern limit of the fold to the southern; the dip varies from southerly through
south-easterly to easterly as the river is followed upstream.

In the Wilkinson Limestone extremely complex close folds are outlined by thin
cherty layers in the cliff faces, but the bed, as a whole, does not seem to have been
strongly compressed. It is likely that the folding was caused by movement prior to
consolidation. Intraformational deformation caused by pene-contemporaneous slumping
has been described in laminated siliceous limestones from Texas and New Mexico
(Newell ef al., 1953).

? Devonian.

The structure of the ? Devonian lavas is in marked contrast to that of the older
beds, in that they are mostly gently dipping or sub-horizontal. In this area, folding
referable to the Tabberabberan orogeny seems to have been very slight. Probably at
this time the dacite suffered shearing of variable intensity. This is best shown in
the Pocket Saddle area, and all the occurrences of sheared dacite are close to slate
boundaries, which may also be fault boundaries. Beds of jasper and silicified rock on
the ridge between Pocket Creek and the Goodradigbee River suggest faulting, as does
a wide zone of ironstone and silicified dacite between Howell’s Peak and The Pockets
Hut.

Acknowledgements.

For financial support, transport, equipment and other facilities the writer is
indebted to the Snowy Mountains Hydro-Electric Authority, through Mr. D. G. Moye,
head of the Engineering Geology Branch.

In the field, able assistance was given at various times by M. B. Mc Mniery,
R. E. Cotton, I. P. Brown and G. Raymond. Thanks are due to Mr. and Mrs. T. Taylor,
of Currango, and Mr. Harris, of Coolamine, for hospitality and assistance.

Palaeontological determinations have been made by Dr. D. Hill, University of
Queensland.

Professor W. H. Bryan and Mr. D. G. Moye have kindly read through the manu-
script and the writer has also profited by petrological discussions with Dr. D. Lafeber,
of the Snowy Mountains Authority.

258 PALAEOZOIC GEOLOGY OF THE COOLEMAN CAVES DISTRICT.

References.

CLARKE, W. B., 1860.—Southern Goldfields of New South Wales (quoted by Leigh and
Etheridge, 1894).

DE KoNINCK, L. G., 1876-77.—Recherches sur les Fossiles paléozoiques de la Nouvelle-Galles
du Sud. Bruxelles.

IvAanac, J. F., and Guover, J. E., 1949.—Geological reconnaissance of the Proposed Hydro-
Hileetrie Works in the Tumut-Upper Murrumbidgee Area. (Unpubl. Rept.) Commonw.
of Aust. Bur. Min. Res., Geol. and Geophys., Rept. no. 12.

LEIGH, W. S., and ETHERIDGE, R., JuUNR., 1894.—Report on the Caves in Cooleman Creek,
Cooleman Plains, at the head-waters of the Goodradigbee River, with notes on the
surrounding district. Dept. Mines N.S.W., Ann. Rept. for 1893, 134.

Davip, T. W. H., 1950.—The Geology of the Commonwealth of Australia. London, Arnold,
1950, 2 vols.

NEwWBERY, J., 1956.—Murrumbidgee-Hucumbene Diversion. (Unpubl. Report.) Snowy Mountains
Authority Eng. Geol. Rept. no. S.G.69.

NEWELL, N. D., et al., 1953.—The Permian Reef Complex of the Guadalupe Mountains Region,
Texas and New Mexico. W. H. Freeman and Co., San Francisco.

STEVENS, N. C., 1958.—Geology of Part of the Murrumbidgee-Hucumbene Tunnel Line. (Unpubl.
Rept.) Snowy Mountains Authority Hng. Geol. Report. no. S.G.78.

WALPOLE, B. P., 1952.—Wee Jasper-Cooleman Caves Area. Commonw. of Aust. Bur. Min.
Res., Geol. and Geophys. Rec. 63. (Unpubl. Rept.)

EXPLANATION OF PLATES III-IV.
Plate iii.
Geological map of the Cooleman Caves district.

Plate iv.

1. Cooleman Plain, looking easterly from Jones’ Hut. Open grassland is Cooleman
Limestone, wooded ridge in mid-distance is granite and granodiorite. Bimberi Range (granite)
on horizon.

2. Faulted anticline in Cooleman Limestone, Cave Creek, west of Spencer’s Hut.

3. Cooleman Falls, Blue Waterhole Creek. Bedded rock in foreground is Blue Waterhole
chert, underlying Wilkinson Limestone (upper right).

4. Wilkinson Limestone, Wilkinson’s Cliffs. Granite outcrops on wooded hillside in
background.

5. Columnar dacite, Currango Plain, south of Blue Waterhole Saddle.

6. Bedded cherts of the Blue Waterhole Beds near Spencer’s Hut.

MELAMPSORA LINI (PHRS.) LEV. UREDOSPORE LONGEVITY AND
GERMINATION.

By H. B. Kerr.
(Three Text-figures. )
[Read 29th October, 1958.]

Synopsis.

During genetical investigations of rust resistance at Sydney University between 1948 and
1953 it was found that a boiled aqueous extract of host tissue greatly stimulated the
germination of stored uredospores of Melampsora lini (FPers.) Lévy. (Kerr, 1952). Exploratory
studies were carried out to determine the relative efficiency of water, gelatin solution and
aqueous host extract as germinating media for these spores. These showed that germination
of stored spores on water was highly capricious, that better but still somewhat capricious
germination obtained on gelatin, while maximum germination was obtained on the host extract.
Preliminary experiments were carried out to determine the nature of the substance or
substances responsible for the germination stimulating potential of the extract. These showed
that the substance was part of or comprised the ether-soluble oily fraction adsorbed by
activated charcoal.

Other experiments were set up to determine the longevity of uredospores of different
races of the pathogen under different storage conditions. These showed a _ significant
difference between races when temperature and specially humidity were carefully controlled.
Low temperature and intermediate relative humidity were most favourable to the maintenance
of uredospore viability during storage.

INTRODUCTION.

In any experiment depending on infecticn of host material or in which the results
are assessed in terms of the germination of fungal spores it is essential to obtain
maximum germination of the spores. Since water is the most common natural
germinating medium it is one of the most widely used in the laboratory (Hwang, 1942;
Prasada, 1948). Many workers have found that water was a poor germinating medium
even for fresh spores with no dormant period (Nobile, 1924; Wilcoxon and McCallan,
1943). Germination was often improved on a decoction of host tissue (Chiu and
Walker, 1949). Increases in germination often as good as that with host tissue were
sometimes induced by extracts of non-host planis. Germination was also improved,
in some cases, by the addition of chemicals to the germinating medium (Thiele and
Weiss, 1920; Noble, 1924). The favourable effect of the host tissue was not always
confined to an improvement in germination. Chupp (1917) found that spores which
failed to germinate at room temperature did so at this temperature when young
seedlings were added. Chiu and Walker found that the optimal temperature range
for germination of cucurbit black rot spores on water was 26°C. to 24°C., but was
24°C. to 28°C. on plant extract. Whitehead found that spores of Urocystis cepulae
germinated slowly on water, but much faster on onion juice.

Sharvelle (1936) found that the percentage germination, the length of the germ
tubes, and type of germ tubes of Melampsora lini uredospores sown on cold extracts
of different varieties of flax and linseed were correlated with the rust reaction of the
variety to the race used. The effect was destroyed by brief heating of the extract.
Cold extracts of immune and resistant varieties retarded the percentage germination
and length of the tubes, but extracts of susceptible varieties neither retarded nor
stimulated germination.

It is generally agreed that iow temperatures and low humidities are most
favourable for the maintenance of uredcspore viability. Bailey (1923) reported an
optimal humidity range of 20% to 40% RH for uredospores of Puccinia helianthi.
Raeder and Bever (1931) studied the effects of five humidities (100%, 76%, 49%, 25%
and 1:5% RH) on the longevity of P. glumarum uredospores. They survived best at
49% RH at most temperatures but in the range —2°C. to 5°C. they survived longest at
75% RH. Chester (1946), summarizing results of many workers, showed that uredo-

PROCEEDINGS OF THE LINNEAN SOCIETY oF NEW SoutH WALES, 1958, Vol. Ixxxiii, Part 3.

Cc

260 MELAMPSORA LINI (PERS.) LEV. UREDOSPORE LONGEVITY AND GERMINATION,

spores of P. triticina remained viable for over 1,000 days at 5°C. and for at least 100
days at temperatures of 0°C. to 20°C.

Longevity seemed to be affected by such factors as the time of year when the
spores were produced (Beauverie, 1924), the rust reaction of the variety from which
the spores were collected, and the degree of maturity of the spores (Chester, pp. 116-117).

Lyophilization is probably the most successful method of storing spores. It is
widely used for bacteria and fungi (Raper and Alexander, 1945) and has been
successfully applied to the storage of uredospores by Sharp and Smith (1952). After
500 and 300 days respectively uredospores of P. coronata and P. gr. avenae kept
in a vacuum at low temperature were scarcely less viable and no less infective than
they were soon after storage.

There has been no detailed study of longevity of uredospores of Melampsora lini.
Hart (1926) found that they remained viable for seven weeks at 7°C. and 60% RH.
Flor (1935) reported that they remained viable for two to four months at 4°C. and
for more than a year if kept in tightly stoppered tubes at —5°C. (1945). Waterhouse
and Watson (1943) observed that race A spores seemed to remain viable longer in cold
storage than spores of other races. Prasada (1948) concluded that spores stored best
at 5°C. to 7°C. but became inviable after 22 weeks, and appeared to remain viable
slightly longer at 10°C. to 15°C. than at 0°C.

HXXPERIMENTAL METHODS.

Uredospores used in these studies were taken from samples stored in a refrigerator
as reserves for the genetical investigations. They were collected from fully susceptible
varieties grown in glasshouses, with the exception of a few samples collected from
plants growing in a fluorescent light room. Spores were collected by dusting them
from pot-grown plants onto glazed paper, and stored in small glass phials 13” to 2”
long and about 4” diameter. They rarely occupied more than 25% of the total tube
volume, usually 10% or less. The tubes were stopped with labelled corks, and stored
within an hour or two of collection. The tubes were kept in a brass tray which
minimized slight daily fluctuations in temperature caused by periodic opening of the
door. The samples were removed as a group to room temperature about four times-
a year during defrosting. The temperature otherwise remained fairly constant at
0°C. to 2°C.

Samples selected for study were removed to room temperature and allowed to
adjust to this temperature for at least half an hour. Spores were shaken from the
tubes onto the surface of the germinating media, and dispersed evenly by vigorous
whisking with a looped piece of flexible wire. Whisking was continued until checks
under the microscope showed that all clumps of spores had been broken up. The
spores were germinated in closed petri dishes kept in the dark at a constant temperature
of 15°C. Percentage germination was measured (usually after 40 hours) by counts
of surface spores under the high power of the microscope from successive fields across
the diameter of the petri dish.

The linseed extract was prepared by bringing distilled water to the boil, adding
fresh seedling or young plant tips at the rate of one gramme green weight to 25 e.c.
of water, and boiling for twenty minutes. The solution was decanted off after removing
any surface scum and used without filtering or further treatment.

In the early studies, due to the pressure of the genetical and race survey work, it
was not always possible to make actual counts. But, whenever possible and unless
otherwise stated, germination on extract and gelatin was determined from counts of
600 to 700 spores. Spores sown on water were always too mobile to count. When it
became necessary to make estimates care was taken to overestimate rather than under-
estimate percentage germination on water, to guard against any bias in assessing the
germination stimulating potential of the linseed extract.

EXPERIMENTAL RESULTS.
Germination on Water, Gelatin and Host Tissue Extract.
Spores sown on linseed extract usually germinated much better than they did on
gelatin or water. It was assumed that maximum germination was obtained on the

BY H. B. KERR. 261

extract. Failure of spores to realize maximum germination on the other media was
attributed to storage hardening. This term is used to denote that conditioning of the
spore wall or spore contents by which it becomes increasingly difficult for spores to
germinate to the full extent of their viability. Samples tested during this project
sometimes became severely storage-hardened in less than a month. This was exceptional.
Most accessions were not noticeably storage-hardened until they had been stored for
more than a month.

TABLE 1.
Germination of Uredospores Stored for One Month.
Storage. Percentage Germination.
Race. ParanerAtine Humidity.
et e RH Water. Extract.
C.
Percentage.
2 0 25 40 97
1 0 50 25 97
2 0 75 20 91
2 BED) 25 5 98
1 3°5 75 0 52
6 10 75 5 15
Average 15 75

Samples of spores of three races, randomly selected from a range of different
storage conditions and tested on water and extract, showed that spores become storage-
hardened under most if not all conditions of storage. Spores stored for a month were
300% more germinable on extract than they were on water. Storage-hardening was
still more pronounced after four months’ storage.

TABLE 2.
Germination of Uredospores Stored for Four Months.
Storage. Percentage Germination.
Race. ‘ Tevineratiire | Humidity.
ot zZ : RH Water. Extract.
C.
Percentage.
1 0 25 3 71
6 0 25 5 83
2 0 25 0:5 74
1 0 50 15 82
6 0 50 10 95
2 0 50 3 93
6 3°5 50 5 94
6 10 25 0-2 41
1 10 50 0-1 38
Average 5 75

Spores stored for longer periods rarely germinated well on water, though some
samples gave up to 60% germination after more than four months’ storage. There
was no consistent correlation between the degree of storage hardening and the period
of storage.

Gelatin was used to counter the mobility of the spores on water, to obtain an
accurate measure of spore germination on an inert medium. Hxperiments were carried
out to determine the most satisfactory concentration for germination. One-eighth
per cent gelatin solution gave the most satisfactory results over the range 0°C. to 21°C.,
but there was little difference in the efficiency of concentrations from one-quarter per
cent to one-half per cent at the optimal temperature of 15°C. Spores gave a high

262 MELAMPSORA LINI (PERS.) LEV. UREDOSPORE LONGEVITY AND GERMINATION,

percentage germination on one-half per cent gelatin, but the tubes were stunted. At
higher concentrations germination was inhibited.

Cold plant extracts were prepared by macerating host tissue in solution in a
Waring blendor. This stimulated germination of some stored samples, but most samples
failed to germinate any better than they did on water. The same extract boiled
stimulated most of the stored samples. Four per cent linseed extracts gave the highest
germinations. Concentrations as low as 1% induced quite good germination. Below
this the solution lacked the surface properties of the standard extract, the spores were

TABLE 3.
Effect of Concentration of Gelatin on Germination.
Percentage Germination of Freshly Collected Race 1 Uredospores.
Concentration of Gelatin.

Temperature. ts % $% 4% £% 1% 2%
5° C 90 98 98 80* 15* 1*
15°C 89 93 91 91* 9* 3*
PAS (0; 65 86 74 80* 4* 1*

* Stunted germ tubes.

too mobile to permit accurate counts, and the percentage germination fell away.
Extracts prepared from immune and susceptible varieties, from typical flax and typical
linseed varieties were equally effective in stimulating germination.

Comparative studies were made on the three media at optimal temperatures.
Results in Table 4 were typical of those given by most if not all races stored for more
than five months.

Germination was always poorest on water. The spores were very mobile on the
surface of the water. Germination was negatively hydrotropic. Most of the tubes were
long, smooth, hyaline and almost unbranched. After several days they formed an
anastomosing network of apparenily fused tubes. Most samples germinated better
on gelatin than they did on water. The increase ranged from a very slight increase

TABLE 4.
Comparison of Percentage Germination on Gelatin, Linseed Extract and Water.
8.U. Percentage Germination on Media Listed.
Race. Accession Date Months
Number. Collected. Stored. Linseed Gelatin
Extract. 1%, Wee
2 492 2: 6:52 11 18 0:2 0
5 519 22:11:52 5t 75 19 2
2 548 22:11:52 5s 95 62 10
8 555 16:11:51 18 26 15 5
3 556 16:11:51 18 25 13 5
il 557 25:10:52 | 6 91 3 3
8 564 18:11:52 54 94 55 10
New Zealand
Race 3 22:11:52 54s 78 55 15
Average ae ata aS 63 28 6

to an increase almost as great as but never equal to that induced by the extract.
Germination was relatively negatively hydrotropic, but many tubes grew along and
some below the surface. The tubes like those on water were long, hyaline and almost
unbranched (Text-fig. 1).

The spores germinated faster on the extract than they did on the other two media.
Some tubes grew aerially and were typically unbranched. Most tubes grew along or
just below the surface. After reaching a length of two to three spore diameters they
branched profusely distally to produce a network best described as stag-horn (Text-
fig. 2). This type of branching was most pronounced among the Punjab-attacking (PA)

BY H. B. KERR. 263

races, and somewhat less pronounced with some non-Punjab-attacking (NPA) races
(Text-fig. 3).

Text-figures 1-3.

Maximum germination was obtained on each medium at 13°C. and 15°C. Experi-
ments were set up to determine the effects of higher temperatures on germination on
the different media.

264 MELAMPSORA LINI (PERS.) LEV. UREDOSPORE LONGEVITY AND GERMINATION,

In an early experiment samples of race 2 were tested on extract and water at
13°C. and 23°C. (Table 5). There was an appreciable reduction of germination on
both media at the higher temperature. The adverse effects of the higher temperature
were most pronounced on water.

The greater the storage-hardening of the spores and/or the more adverse the

conditions for germination, the greater the discrepancy between the observed germina-

TABLE 5.
Estimated Percentage Germination of Race 2 (Acc. 492) Uredospores on Water and Eatract at 13°C. and 23°C.
Period Stored at 0° C. in Months.
Medium. Temperature. SS | Average.
ACh 14 | 33] 4 4 4 4 4 4. 5 5 5 | 54 | 64
Extract 13 90 | 95 | 90 | 90 | 90 | 90 | 90 | 90 | 95 | 95 | 95 | 85 | 85 90
Water 13 10 | 20 | 15 | 10 | 30 | 10 | 60 | 80 | 60 |} 10 | 50 | 10 |} 50 | 32
Extract 23* 84 | 84 | 50 | 41 | 40 |} 90 | 73 | 88 | 80 | 71 | 84 | 66 | 73 71
Water 23 4 4! 0 0 0) 1 $ il il 4 0 1 0 if
|

* Count of 500 spores.

tion and the known viability assessed from the germination at 15°C. on extract
(optimal germination conditions). The experiment summarized in Table 6 showed
clearly that germination of the uredospores was conditioned by the temperature
tolerance of the race, the temperature of germination, the nature of the germinating
medium and the storage-hardening of the spores.

Hach sample gave maximum germination on the extract at 15°C. The percentage

germination under these conditions was taken to be the viability of the spores. Freshly
TABLE 6.
Germination of a High-Temperature-Tolerant Race and Four Other Races on Three Germinating Media at 15°, 20° and 25° C.
Punjab Non-Punjab-Attacking Races.

Race .. sits Se ae 1 5 6 5 13 11

$.U. Accession number 507 496 528 519 582 568
Period stored 2 months 2 months 2 months 1 week 1 week 2 weeks

Medium. Temperature.

Extract 15° C. 85 92 94 99 97 89
Gelatin 4% 24 hours. 66 63 62 47 58 i?
Water 65 50 50 20 10 5
Extract 15° C. 86 92 93 — — _
Gelatin 48 hours. 66 65 70 43 58 0)
Extract 20° C. 78 88 86 98 97 79
Gelatin 4% 40 hours. 60 47 34 40 58 10
Water 40 30 15 2 — 5
Extract 25° C. 70 10* 13* 0 30* 3*
Gelatin 4% 40 hours. 2 0 0 0 0 0
Water 1 | 0 0 0 0 0

* Germ tubes abnormal.

collected spores of any race germinated to the full extent of their viability at 15°C.
and 20°C. on extract. The slightly reduced germination at 20°C. on extract of races 1,
5 and 6 stored for two months, and of the short-stored race 11, indicated a slight
storage-hardening of these samples. The other two short-stored races 5 and 13 (stored
for only one week) germinated equally well at both temperatures on extract and were
presumably not storage-hardened. (The extreme storage-hardening of this sample of
race 11 after only two weeks’ storage is unusual but not without parallel. Its storage-
hardening was fully confirmed by its very poor germination on gelatin.)

BY H. B. KERR. 265

Very poor germination on gelatin was a reliable indication of marked storage-
hardening when the spores were highly germinable on extract. But percentage
germination on gelatin was no accurate indication of the measure of storage-hardening.
This was clearly indicated by the fact that non-storage-hardened races germinated
more poorly on gelatin at 15°C. in this experiment than spores of the races stored
for two months. Despite this, results on gelatin gave quite good indication of the
temperature effects on germination. Since the short-stored races 5 and 13 were not
storage-hardened and were virtually freshly collected a marked reduction in percentage
germination at the higher temperature was not expected. Germination of these races
on gelatin at 15°C. and 20°C. confirmed this. The two long-stored races 5 and 6
germinated much more poorly at the higher temperature. This agreed with the
general observation that 20°C. has an adverse effect on the germination of storage-
hardened spores. Race 1, however, owing to its high-temperature tolerance, germinated
almost as well at 20°C. on gelatin as it had at 15°C.

TABLE 7.
Effect of Temperature on Germination of Stored Spores Sown on Extract.
|
IRE ag 1 1 iL 1 1 17 13 18 ii 15 17
Temp. | Accession 613 613 507 621 624 604 582 607 550 599 604
Months

stored .. 9 14 3 4 14 4 1 4 4 4 11
UB Cs he ES 90 97 100 100 98 96 93 89 83 95 74

20° C. ae ae 90 97 100 99 99 95 91 61 80 96
25° C. 87 92 98 99 96 87 93 5 70 97 4

|

The adverse effect of high temperature was most pronounced at 25°C. None of the
NPA races germinated normally on any of the media. Some gave a slight percentage
germination on extract, but germ tube development was quite abnormal. Germination
Was entirely suppressed on gelatin and water. Race 1, however, germinated well on
the extract. But almost complete suppression of germination on water and gelatin
suggested that the temperature was beyond the optimal.

An experiment summarized in Table 7 confirmed the conclusions drawn from the
preceding experiment. Five different accessions of Punjab-attacking race 1 and six
NPA accessions, including five different races, were tested on extract at 15°C., 20°C.
and 25°C. The spores held at 25°C. were pre-exposed to optimal germinating tempera-
tures for twenty minutes immediately after sowing before being placed at 25°C.
Percentage germination was checked after forty hours and the type of germ tube
development noted.

Hach race 1 accession gave excellent percentage germinations at each temperature
with only a slight falling off at the higher temperatures. Two of the six NPA
accessions were appreciably less germinable at 20°C. than they were at 15°C., and gave
extremely poor germination at 25°C. Both were extremely storage-hardened, though
one had been stored for only two weeks. Tests on gelatin fully confirmed their
storage-hardening. The other four NPA accessions germinated well at the higher
temperatures and were apparently not appreciably storage-hardened. The high germina-
tien of these accessions at 25°C. compared with the very peor germination of the NPA
races at the same temperature in the previous temperature must be attributed to
their pre-exposure for twenty minutes to optimal germinating temperatures. The
extract appeared to stimulate germination within the first twenty minutes, and once
stimulated adverse temperatures did not check the process.

The type of germ tube development summarized in Table 8 gave an excellent
indication of the temperature tolerance of the different accessions. Hach developed
typical stag-horns at 15°C., those of the race 1 accessions as a group being somewhat
more pronounced than the NPA accessions. At 20°C. each race 1 accession again
developed pronounced stag-horns. Stag-horn development of the NPA accessions was
completely suppressed, but germ tube development was otherwise quite vigorous,

266 MELAMPSORA LINI (PERS.) LEV. UREDOSPORE LONGEVITY AND GERMINATION,

slightly branched and apparently normal. At 25°C. germ tube development of the NPA
accessions was quite abnormal, very vesicular, and branching was completely inhibited.
Germ-tube development of the race 1 accessions was quite vigorous and normal at this
temperature, but stag-horns were suppressed.

Analysis of the Linseed Extract.

Detailed analysis of the extract was beyond the scope of the present investigations,
but several experiments were carried out to determine within very broad limits the
nature of the stimulating principle or principles. This involved ether extraction,
charcoal adsorption and charring of the aqueous extract, and tests of the residue to
determine whether the active properties of the extract had been eliminated or destroyed.

Since the residue after each treatment lacked the spore dispersing and immobilizing
properties of the original extract small amounts of gelatin were added to the
reconstituted solutions. A quantity slightly in excess of one-eighth per cent by weight
was added to offset the slight deterioration of the spreading qualities of the gelatin
induced by the salts, etc., in the solution. The ether extractions were made before
gelatin had been adopted as an inert spreading agent.

Special precautions were taken to ensure that the reagents were free from toxic
impurities.

Ether Hatraction.

The 4% aqueous extract was shaken up with ether and the aqueous residue
separated off. The process was repeated several times, fresh ether being used to
ensure removal of all ether-soluble fractions from the aqueous residual solution. The
residual solution was heated gently to eliminate all traces of ether, and finally made
up with distilled water to the original volume. The ether-soluble fraction was added
to water in an evaporating dish and the ether evaporated off slowly. This fraction
was tested in its concentrated form and the excess made up with distilled water to the
original concentration.

The percentage germination on the aqueous residue after extraction was lower
than that obtained on distilled water. A concentrated solution of the ether-soluble :
fraction induced a very high percentage germination. It was most remarkable for the
unusually large number of germ tubes which emerged from each spore. The tubes,
however, failed to develop after emergence. At the normal concentration this fraction
induced a rapid and high percentage germination, but the tubes again failed to develop.
When the ether-soluble and ether-insoluble fractions were added together and made up
to normal concentration the spores germinated rapidly and normally, almost as well
as they did on the original extract.

It seemed obvious from this experiment that the germination stimulating properties .
of the linseed extract derived principally from the ether-soluble fraction. This fraction
seemed to be responsible for the high percentage germination and the relatively rapid
rate of germination characteristic of the linseed extract. In concentrated form it
stimulated the germination process still more by inducing an unusually high number
of tubes from each spore. The gerin tubes failed to develop normally, however, on the
ether-soluble fraction. This might have been due to impurities in the ether, though
this does not seem likely in view of the precautions taken to eliminate impurities.
Results tend rather to suggest that some substance or substances present in the
ether-insoluble fraction were required in conjunction with the ether-soluble components
to induce normal development following germination.

Charcoal Adsorption.

B.D.H. activated charcoal and charcoal obtained by sulphuric acid charring of
sugar were used for this extraction. The B.D.H. charcoal had a slightly toxic effect
on germination not completely eliminated by frequent leaching with boiling water.
The charcoal prepared from sugar was thoroughly leached to remove all trace of acid
and then heated to over 600°C. to enhance its adsorbing properties. The 4% plant
extract was boiled with the charcoal for 20 minutes and the charcoal filtered off. The
B.D.H. charcoal was the most efficient adsorbing agent, and yielded a clear residual

BY

H. B. KERR.

TABLE 8.

267

Effect of the Temperature on the Type of Germ Tube Development on 4 per cent. Linseed Extract.

Temp.

15° C.

20° C.

25° C.

Race.

a

SEB eee

17

17

18

15

S.U. aos
Accession | Months Per-
Number. | Stored. centage
Germina-
tion.
613 9 90
613 14 97
507 2 100
621 4 100
624 1¢ 98
550 4 83
604 4 96
604 11 74
582 1 93
607 4 89
599 4 —
613 9 90
613 14 97
507 2 100
621 $ 99
624 1¢ 99
550 4 80
604 $ 95
604 11 65
582 1 91
607 $ 61
599 3 96
613 9 87
613 1¢ 92
507 £ 98
621 oy 99
624 14 96
550 4 70
604 4 87
604 11 4
582 1 93
607 4 5
599 4 97

Type of Germ Tube Development.

Activity.

Very active.

29 oH)

Normal.

Abnormal
hibited.
Abnormal
hibited.
Abnormal
hibited.
Abnormal
hibited.
Abnormal
hibited.
Abnormal
hibited.

in-

in-

in-

in-

Distribution.

Mostly surface.

Mostly surface, slightly
subsurface.

Surface mostly, mod-
erate amount just
subsurface.

Mostly surface.

Moderate amount just
subsurface.

A lot subsurface.

Slight amount
surface.

Mostly surface.

Mostly surface.

sub-

Mostly surface.

” 2

A lot subsurface.
Mostly just subsurface.
Mostly surface.

A lot subsurface, quite

a few long aerial
hyphae.
Mostly subsurface,

some aerial tubes.
Mostly just subsurface.

Mostly surface.

Plenty of surface
growth.

A few long aerial
hyphae, mostly
surface.

Quite a few long aerial
hyphae, mostly
surface.

Quite a few long aerial
hyphae, mostly
surface.

Subsurface.

Mostly subsurface.

Branching.

Very staggy.

Staggy.

Very staggy.
Staggy.

Slightly staggy.
Moderately
stagey.

Very staggy.
2 2
Staggy.
>

”

Long, slightly
branched.
Slightly
branched.
Long, slightly
branched.
Slightly
branched.

Long, slightly
branched.

Long, slightly
branched tubes.

staggy.
ot
agsy

; | slight
branch-
N o t ing.
staggy. |
|
No t|
staggy. |
Very vesicular,
no branching.
Very vesicular,
no branching.

Very vesicular,
no branching.
Very vesicular,
no branching.
Very vesicular,
no branching.
Very vesicular,

no branching.

268 MELAMPSORA LINI (PERS.) LEV. UREDOSPORE LONGEVITY AND GERMINATION,

solution lacking the surface properties of the original solution. The residual solution
after the sugar charcoal extraction was less clear and possessed some of the surface
properties of the original.

The spores of the four rust accessions tested germinated much more poorly on
the residual solutions from the two charcoal extractions than they did on the original
extract. Addition of gelatin to the residual solutions increased the percentage

TABLE 9.
Ether Extraction of Extract.
Race 2 Accession 492.

Germinating Medium. Percentage Type of Germination.
Germination.
1* Distilled water. 30 Very short germ tubes.
2; Normal strength linseed extract (boiled 95 Dense, very vigorous, staggy germ tube develop-
2 minutes). ment.
3* Residue of boiled extract after ether 5 Long and unbranched germ tube.
extraction.
4; Fraction extracted by ether and brought 90 Rapid germination but germ tubes failed to
back into aqueous solution. elongate.
57 3 and 4 combined. 90 Very rapid germ tube development, long, not
so even as 3 or 4, and slightly branched.
6; Concentrated solution of 4. H 85 Remarkable number of germ tubes from each
spore, but tubes failed to develop normally.

* Maximum estimate. Spores too mobile to count.
+ Minimum estimate.

germinations, but not appreciably beyond the figures obtained on pure one-eighth per
cent gelatin. The substance or substances responsible for the germination stimulating
potential of the plant extract had been adsorbed by the charcoal.

Charring of the Extract.
The plant extract was evaporated to dryness in a low temperature oven and then
heated to about 600°C. for an hour to eliminate all traces of organic matter. The

TABLE 10.
Charcoal Adsorption of Extract.

Percentage Germination after
48 Hours at 15°C.

$.U. Accession we ae 613 528 604 557

Race .. ae se ae 1 6 17 il
Months stored nie ae 9 3 11 94
Germinating Medium. Average.
il Normal linseed extract oF ¥s Me 92 75 80 52 75
2 Gelatin 4% .. ae He bf is 69 65 112, 40 47
3* Linseed extract residue after treatment with
activated B.D.H. charcoal ae rie 30 15 2 15 16

+ 3 with gelatin added. . x2 at aes 75 51 6 35 42

5 Linseed extract residue after treatment with
sugar charcoal ie me t 3 We 55 19 5 25 26
5 with gelatin added.. aie ba a 74 66 14 41 49

Ss

Germination based on count of 700 spores approximately.
* Spores too mobile on surface. Maximum estimate for 3.

residue was brought back into solution and made up to the original volume. It lacked
all the surface properties of the original solution.

Hight rust accessions were used to check the germination potential of the charred
extract. All of them gave very much lower germinations on the charred extract than

BY H. B. KERR. 269

they did on the original. Addition of gelatin to the charred extract increased
germination of six of the accessions appreciably, three of them to a level almost equal
to germination on the original extract. This need not be attributed to the fact that
the residue after charring together with the gelatin possessed the stimulating properties
of the plant extract. It could be attributed rather to the fact that the spores had not
become severely storage-hardened although they had been stored in some cases for
periods of 94 months. Particular importance attached to the results obtained with

TABLE 11.
Effect of Charring on Stimulating Properties of the Extract.

S.U. Accession .. ae .. | 507 557 613 496 528 550 568 604

Race oe ae oe ae 1 1 1 5 6 ih iil 17

Months stored .. ss an 3h 9k 9h 3h 34 34 3h 12

Germinating Medium. Percentage Germination after 48 Hours at 15°C. Average.
1 Normal linseed extract a a 95 70 84 90 84 64 69 64 | 78
2* Charred linseed extract Se ae 80 10 60 50 | 50 4 20 w 35
3 Charred extract plus gelatin ae 91 26 79 82 63 39 55 1 55

* Maximum estimate. Other percentage germinations based on count of 700 spores approximately.

race 17. Tests of the same sample of rust from which these spores had been drawn
showed consistently that they were severely storage-hardened. Tine charred extract with
and without gelatin failed to increase germination of the race 17 spores beyond 2%,
although the spores were still highly viable judging by their germination on the
original extract. It could be assumed that charring of the extract had destroyed the
substance or substances responsible for the germination stimulating potential of the
original plant extract.

TABLE 12.
The Effect of the Germinating Medium on Type of Germ Tube Development.
Race. SSE Linseed Extract. Charred Solution. Charred Solution and

Accession. Gelatin.

1 507 Moderately stagey. Super- | Long unbranched. Aerial | Long unbranched tubes. Aerial
ficial growth. growth. and superficial.

1 557 Very staggy. Superficial | Long unbranched. Aerial | Long unbranched tubes. Aerial
growth. ~~ growth. and superficial.

i 613 Very staggy. Superficial | Long unbranched. Aerial | Long unbranched tubes. Aerial
growth. erowth. and superficial.

5 496 Very staggy. Superficial | Long unbranched. Aerial | Long unbranched tubes. Aerial
growth. eTowth. and superficial.

6 528 Rather stagey. Superficial | Long unbranched. Aerial | Long unbranched tubes. Aerial
growth. growth. and superficial.

7 550 Rather staggy. Moderate | Long unbranched. Aerial | Long unbranched tubes. Aerial
amount subsurface. growth. and superficial.

ilil 568 Rather staggy. Superficial | Long unbranched. Aerial | Long unbranched tubes. Aerial
growth. growth. and superficial.

17 604 Slightly stagey. Superficial | Long unbranched. Aerial | Long unbranched tubes. Aerial
growth. growth. and superficial.

Preliminary Studies of Uredospore Longevity at 0°C. to 2°C.

Initial studies of uredospore longevity were based on results obtained with spores
stored in a refrigerator and used for the genetical investigations of disease resistance.
These results are listed in Table 13.

The results did not indicate any consistent correlation between the period of cold
storage and survival of the uredospores, although there was an obvious serious falling
off in viability by the end of twelve months. Some other factor or factors with time
played the dominant role in determining the survival of the spore samples. Various
possibilities were considered, e.g., race, spore bulk during storage, production at

270 MELAMPSORA LINI (PERS.) LEV. UREDOSPORE LONGEVITY AND GERMINATION,

different stages in the sporulation cycle, production at different times of the year,
pre-storage temperatures, freezing and thawing effects in the refrigerator, etc. 'These
possibilities are dealt with in the following section.

TABLE 13.
| |
Date i Time
Collected. Stored. Race, Accession and Percentage Germination.
| (Months.) |
21:11:50* | 6 1. 557 60% 586 50% 2. 544 0% 548 15% 552 75%
3. 511 0% 556 0% 4. 558 50% 578 50%
: 5. 514 85% 515 70% 554 60% 7. 550 15% 8. 555 1/10%
9. 559 80% 583 0% 10. 579 85% 11. 568 40%
{ 12. 580 10% 13. 582 50% 14. 584 38%
13:11:50* 6 2. 544 80% 3. 511 70% 556 80% 8. 555 70%
9. 583 50% 14. 584 60%
23:10:51 8 2. 552 54% 560 56% 4. 518 42%
5. 514 52% 515 53% 11. 568 57%
20:10:51 | 8 2. 552 59% 560 70% 548 72% 4. 518 58%
5. 514 67% 515 60% 11. 568 75%
13:11:50 10 1. 507 50% 586 60% 2. 496 81% 544 56% 548 76% 552 88%
4. 558 84% 578 81% 5, 5ilb 9496 5545 (82195 6. 551 59%
7. 550 74% 9. 559 1% 10. 572 83%
11. 568 20% 12. 580 57% 13. 582 79%
24:11:50 94 1. 586 87% 12. 580 838%
28: 7:50* 11 1. 557 30% 2. 548 4% 560 0% 3. 556 30%
6. 551 50%
31: 7:50* 11 1. 557 30% 8. 556 4%
3: 8:50* 11 1. 557 50% 2. 548 25% 560 50% 3. 556 40%
6. 551 70%
8: 8:50* 11 il, Hv = BI09% 2. 548 30% 560 40% 3. 556 20%
6. 551 25%
1: 5:50* 14 1. 507 15% 2. 492 30% 544 10%
5. 496 5% 515 10%
2: 6:50* 13 1. 557 251552 3. 556 4, 558 5. 554
8. 555 9. 559 All inviable.
17: 7:51 12 1. 557 34% 586 38% 2. 548 48% 552 56% 560 27%
3. 556 12% 4. 558 44% 578 14%
5. 514 69% 515 39% 554 18% 6. 551 4% 9. 559 18%
10. 579 13% 11. 568 11% 13. 582 30%
18: 8:50 143 1. 557 0% 2. 548 1% £560 4% 3. 556 20%
15: 8:50 144 il, By = BOGS 2. 548 9% 560 4% 8. 556 12%
11: 8:50 144 i, BBY? IBY 2. 548 4% 560 1% 3. 556 8%
31: 7:50 15 il, Be =O 2. 548 — 560 0% 3. 556 0%
12: 4:50 144 i, OY 9S SR, Ib oS OG ESR II) AG
1. 507 5. 514 515 6. 528 Inviable.
26: 4:50 14 , BIZ HY 5. 496 20% 519 25% 4. 518 1%
2. 492 6. 528 Inviable.
17:11:50 17 { 2. 560 12% 3, fli i% 8. 555 18%
1. 507 2. 548 3. 556 4. 558 578 5. 554 Inviable
6. 551 7. 581 550 9. 559 583 10. 579 Inviable
11. 568 12. 580 13. 582 Inviable.
13:11:50 17 1. 586 5% Wr, byl BOG
i, EXO" 577 2. 548 560 4, 558 5. 496 Inviable.
6. 551 7. 550 10. 579 11. 568 Inviable.
2:11:50 1743 5. 496 Lot A 18% Lot B 16%

* Estimated percentage germination.

Race Differences.

Race 2 (accession 492) appeared to survive better than other accessions. But
apart from this there were no consistent differences in the longevity of uredospores
of different accessions stored in the refrigerator. Race 9 (accession 559) was one of
the most viable of those tested after six months. It was one of the least viable
accessions after ten months’ storage. Several accessions were identified as the same
race, but different accessions of the same race often differed as much in viability
after prolonged storage as accessions of different races. It was obvious that a factor

BY H. B. KERR. 271

or factors other than race played the dominant role in determining the viability of
uredospores in cold storage.

Spore Bulk During Storage.
Small quantities of spores tended to survive better than spores stored in bulk. This
is shown in results obtained with race 2 accession 492, confirmed on many occasions

TABLE 14.
Quantity of Spores Percentage
Collected. Stored. Germination
on Extract.
10: 9:51 2% of tube volume. 91
10: 9:51 20% of tube volume. 10
2:10:51 25% of tube volume. 56

Count of 500 spores approximately.

by general observation. Uredospores of accession 492 were collected on 10th September,
1951. The material was divided into two lots and stored immediately. One lot of
spores occupied no more than 2% of the tube volume after corking. This lot was
-still 91% viable after eight months’ storage. The other lot, stored in a tube of equal

TABLE 15.
Viability of Spores of Race 2 (Accession 492) Collected During a Sporulation Cycle.
Period Date Date of Collections of Uredospores.
Stored. Tested. SSS
6:10:50. 10:10:50. 13:10:50. 16:10:50. 20:10:50.
9 months ae 11: 7:51* 80% 80% 80% 80% 80%
12 months is 17:10:51} 67% 17% 81% 83% 53%

* Minimum estimate of percentage germination.
+ Count of 250 spores approximately.

size, occupied about 20% of the total tube volume. It was no more than 10% viable
after the same period of storage.

But bulk storage was not always detrimental to uredospore longevity. The same
accession collected three weeks later was still 56% viable after 74 months’ storage,
although the spores occupied at least 25% of the total tube volume.

TABLE 16.
Viability of Spores of Race 6 (Accession 529) Collected During a Sporulation Cycle.
Period Date Date of Collections of Uredospores.
Stored. Tested. _
10:10:50. 13:10:50. 15:10:50. 19:10:50.
9 months .. 11; 7:51* 50% 30% 60% | 70%
12 months .. 18:10:51+ 25% 2% H% | 12%

* Minimum estimate of percentage germination.
+ Count of 250 spores approximately.

Production at Different Stages in the Sporulation Cycle.

It was thought that the longevity of uredospores might be somewhat predetermined
by the stage in the sporulation cycle during which they were produced, but this was
not supported by results obtained with races 2, 6 and 7. Uredospores of these races
had been collected at two- to four-day intervals during a single vigorous sporulation
eyecle, and were tested for viability after nine and twelve months’ cold storage. There
was no consistent correlation between the time in the sporulation cycle when the
spores were produced, and their viability after nine and twelve months’ cold storage.
Sometimes spores produced early survived better than spores produced later. Sometimes

272 MELAMPSORA LINI (PERS.) LEV. UREDOSPORE LONGEVITY AND GERMINATION,
the reverse was found. It could only be concluded that spores could remain viable
for considerable periods whether produced early, late or part way through the

sporulation cycle.

Production at Different Seasons.

Uredospores collected during winter often became inviable relatively quickly. It
seemed possible that spores produced during winter lacked the capacity for survival
of spores produced in spring and summer. But examination of results collected over
several years showed that spores produced during winter could remain viable as long

TABLE 17.
Viability of Spores of Race 7 (Accession 550) Callected During a Sporulation Cycle.
Period Date of Collection of Uredospores.
Stored. Date ;
Tested. 28:8:50. 30:8:50. 1:9:50. 4:9:50. 8:9:50.
9 months Eo == 60% 20% 40% 30%
12 months 18:10:517 18% 47% 4% 22% PALO,
|

* Minimum estimate percentage germination.
7 Count of 250 spores approximately.

as spores produced at other times of the year. Nine of the ten races collected in
mid-July were still moderately to highly viable after twelve months (Table 18a).
Spores of race 2 (accession 492) collected in early July were as viable after 134 months’
storage as spores collected in warmer months and stored for shorter periods (Table 188).

Prestorage for Several Days at Temperatures up to 20°C.

Uredospores of the different accessions were collected in general at two- to four-
day intervals. Temperatures in the glasshouse varied considerably from day to day,
and during each day. It seemed possible that these random variations in temperature
might affect the longevity potential of spores collected on different days, depending on
the period the spores were allowed to remain in situ prior to collection. The possibility

TABLE 18A.
Percentage Survival of 10 Races after 12 Months’ Cold Storage.
Percentage Germination of Accessions on 4% Extract
Collected. after 48 Hours. Period
(Count of 400 spores approximately.) Stored.
17:7:51 Race 1 <Ac5b57 34% Ac586 38%
Race 2 Ac5b48 43% Ac552 56% Ac560 27%
Race 3 Ac556 12%
Race 4 <Ac558 44% Acd78 14%
Race 5 <Ac5hb14 69% Ac5d15 39% Acb54 18%
Race 6 Ac551 4% 12 months.
Race 9 Ac5b59 18%
Race 10 Ac579 13%
Race J1 Ac5b68 11%
Race 1 Ac582 30%

was checked by exposing freshly collected spores of race 2 (accession 492) to tempera-
tures of 8°C. to 20°C. for varying periods before transferring them to the refrigerator.
Check samples were placed in the refrigerator without prior treatment.

It appeared from the results that exposure of spores to temperatures up to about
20°C. for periods of about ten days did not seriously affect the capacity of spores to
retain their viability in cold storage. Since the mean daily temperature rarely
exceeded 20°C. during those months when rusts were cultured in the glasshouse, and
since spores were seldom collected at greater than four-day intervals, it seemd highly
improbable that temperature conditions prevalent during the period of spore formation
in any way affected the potential longevity of spores in cold storage.

BY H. B. KERR. 273
Alternate Freezing and Thawing.

Freshly collected uredospores, and uredospores kept in cold storage for three
months were subjected to six cycles of freezing and thawing to determine the immediate
effect, if any, on viability. A parallel series of tests was carried out for each of three
races, races 1, 2 and 6. The spores were chilled in the tray beneath the cold box

TABLE 18B.
Viability of Spores of Race 2 (Accession 492) Collected at Different Times of the Year 1950.
|

Date Collected. 6:12. | 4:10. 23:9:1950. 25:8. | 30:8. | 29:8. | 15:8. | 8:7.
Months stored .. 14 34 44 5 5 5 5s 64
Percentage germ-

ination* 90 95 90 90 90 90 90 90 95 95 95 85 85

|

Months stored 84 104 11 1] 11 1] 11 J 12 12 12 124 134
Percentage germ-

ination; 77 85 85 68 87 90 63 — Si 85 79 5] 79

* Minimum estimate of percentage germination.
+ Count of 300 spores approximately.

of the refrigerator at a temperature of 0°C. to 2°C. for half an hour. They were then
removed and left at room temperature (approximately 15°C.) for another half-hour. A
sample was taken off for determination of percentage germination on extract before
returning the material for a further cycle. After six such cycles none of the races
freshly collected or stored for three months showed any detectable reduction of
viability when 600 spores were counted.

The process was continued for another six cycles, during which the half-hour
period was lengthened to more than an hour. At the conclusion of this series there
was still no detectable reduction of viability.

TABLE 19.
Effect of Pre-Storage at Higher Temperatures, Race 2.

Check Days Stored at the Temperatures Listed.
Temperature. Sample.
4 9 14 | 18 | 21
———————— |
13°-15° C. 90% = | 80% 80% | 70% 60%
18°-20° C. 95% 85% | 75% 35% > 1%
|

Estimated percentage germination after four months’ cold storage.

It was concluded that minor fluctuations of temperature in the refrigerator caused
by daily opening of the refrigerator door and major fluctuations which occurred three
to four times each year during defrosting would probably not have any serious effect
on the viability of the stored spores.

A Note on Uredospores Produced in the Fluorescent Light Room.

Uredospores collected during November, 1950, were produced by plants growing
indoors under fluorescent lighting. The excellent viability of these spores after
92 to 10 months’ storage showed that spores produced under artificial light had much
the same potential longevity as spores produced under natural light.

Summary.

The preliminary studies showed that most if not every race could remain viable
for at least a year, although unknown factors often caused them to become inviable
earlier. The percentage survival after a year ranged from 0% to almost 90%, but
could not be correlated with race differences, the time of the year, or stage in the
sporulation cycle when the spores were produced. There was no evidence to suggest

274 MELAMPSORA LINI (PERS.) LEV. UREDOSPORE LONGEVITY AND GERMINATION,

that periodic fluctuations in the refrigerator temperature or exposure of spores to
temperatures up to 18°C. for several days in any way significantly affected the capacity
of spores to remain viable in cold storage. Spores stored in bulk often became
inviable more quickly than spores stored in small amounts, but this was due to some
other factor than mere spore bulk. Some other major factor or factors were
responsible for the rather unpredictable survival of the spores in the refrigerator.
Spores stored in the refrigerator during the early stages of these investigations
probably varied considerably in their moisture content (at least for the first few
weeks) according to the humidity prevalent during their production in the glasshouse.
Spores collected during rainy weather were noticeably moister than spores collected
in fine weather. Since they were transferred into tubes and corked and stored almost
immediately after collection they had little opportunity to lose any excess moisture.

TABLE 20.
Effect of Pre-Storage at Higher Temperatures, Race 2.
Date | Period of Check Days Stored at 18° C. Days Stored at 8° C.
Collected. Cold Storage. Sample. rs
3 7 11 7 11
4:12:50 2 months* 90% 90% 90% 85% 90% 85%
| 9 months} 84% 85% 89% 81% 85% | 80%
Date Period of Check Days Stored at 18°C.
Collected. Cold Storage. Sample.
3 6 | 10
1:12:50 2 months* 85% 85% 80% 70%
9 months} 78% YEE 68% 68%

* Minimum estimate.
+ Count of 250 spores approximately.

Once this risk had been realized, spores stored in any bulk were allowed to dry out
in the tubes before corking or stoppered with cotton wool plugs. Careful observations
were subsequently made of spores collected in wet weather and stored immediately.
Drops of moisture frequently appeared inside the tube just below the cork. ‘The
moisture usually disappeared after a couple of weeks, but it is certain from the results
obtained later at 75% relative humidity in a carefully controlled experiment that
even at the low temperatures prevailing in the refrigerator there could have been a
serious loss of viability before the excess moisture had been transpired off by the
spores and escaped from the tubes. The volume of the tube and tightness of fit of
the cork would play an important role in determining the period during which the
humidity remained at a critically high level. These findings explain the common
observations that spores collected in winter (the rainy season in Sydney) often, but
not always, lost their viability sooner than spores collected in summer, and that
spores stored in bulk often became inviable sooner than spores stored in small
quantities.

Unless precautions are taken to ensure that the spores are reduced to the same
approximate moisture content at the start of longevity studies and stored under
conditions preventing local build-up of humidity around the spores in the storage
tubes, undetected variations in humidity are likely to obscure the effects of controlled
factors such as race and temperature. This was made very clear by the highly
capricious percentage survival of the races studied during the early investigations of
uredospore longevity at Sydney.

It is fairly certain that precautions taken during the final studies of longevity
detailed in the next subsection eliminated this risk. The use of linseed extract
should also have eliminated the equally grave risk arising from the capricious and
submaximum germination of M. lini uredospores on the commen germinating media.

BY H. B. KERR. 275

Uredospore Longevity of Three Races at Carefully Controlled Temperatures
and Humidities.
Experimental Methods and Materials.

The project was commenced in May, 1952, when it was decided to determine the
storage capacity of three races, 1, 2 and 6, at three humidities, 25%, 50% and 75%
relative humidity, and three temperatures, 0°C., 3-5°C. and 10°C. in all combinations.

The humidity was regulated in medium-sized desiccators with sulphuric acid of
known specific gravity (Maclean and Cook). The acid was made up to the requisite
specific gravity with distilled water by weight and checked with a hydrometer.
Temperature control facilities were very kindly made available by the C.S.1.R.0O.
Division of Food Preservation and Canning, Homebush. Under these conditions
temperatures did not fluctuate by more than +0-2°C. during the course of the experiment.

The uredospores of the three races were built up under identical conditions on
fully susceptible varieties in the glasshouse during mid-winter. Mild sunny weather
prevailed from the date of infection to completion of sporulation. The temperature
recorded on a thermograph showed a diurnal range, fluctuating from a maximum of
68°F. to 82°F. at 2 p.m. to a minimum of 46°F. to 56°F. about 8 a.m. Except for a few
brief cloudy periods the infected plants were exposed to uninterrupted direct sunlight
in the glasshouse from sunrise to sunset. There was no artificial lighting, but
temperatures were kept slightly above outdoor temperatures at night to ensure vigorous
sporulation.

The spores were collected at three-day intervals between 5.30 p.m. and 6 p.m.
The first collection was not retained for the studies, but the next four collections
were kept separately from the time of collection in small glass phials, stoppered
lightly with cotton wool, and placed in a desiccator at 50% relative humidity at 3°C.
Spores were collected only during the period of vigorous sporulation. After the last
collection had been stored at 50% RH for 24 hours the spores were prepared for
storage under the conditions mentioned above. A bulk sample of each race was
prepared by mixing together equal quantities of material from the four collections.
Thorough mixing was achieved when the bulk sample was sieved through a very fine
mesh wire screen to remove all traces of organic matter which might accidentally
have been collected with the rust.

The spores were set up in small glass tubes specially hand-made from narrow glass
tubing about one-twelfth inch internal diameter. Each tube, about one inch in length,
was identified for race, temperature and humidity with small colour patches painted
in a vertical series of three spots. They were left in a well-aired position for several
weeks after painting to eliminate the risk of fumes affecting viability during storage.

Equal amounts of rust were added to each tube to a depth of about one-tenth inch.
The tubes were then mounted unstoppered in wooden blocks bored with the requisite
number of holes to a depth of about half an inch. These blocks had been soaked in
liquid paraffin some time previously to eliminate risk of swelling or mildew development
at the higher humidity. Highteen tubes of each race were mounted in each of nine
such blocks, one block for each combination of temperature and humidity.

The blocks were mounted on glass trays inside the desiccators to avoid any risk
of sulphuric acid splashing onto the spores or onto the wooden blocks.

The desiccators were set up at the given temperatures at Homebush, and arrange-
ments were made for samples to be sent in for germination tests at Sydney University
at monthly intervals.

These samples were set up immediately on receipt on 4% host extract and kept
in the dark at 15°C. in closed small petri dishes for at least 40 hours before
germination counts were made. The spores were examined under the high power
of the microscope undisturbed on the extract. A minimum of 500 spores was counted
for each sample. In general about 600 spores were counted. The studies were continued
for 13 months.

D

276 MELAMPSORA LINI (PERS.) LEV. UREDOSPORE LONGEVITY AND GERMINATION,

Experimental Results.
The results have been graphed and summarized in three sections.
Graphs 7 to 15 give the viability curves of each race under the nine different
storage conditions, to highlight race differences. Graphs 1 to 6 were plotted from
the averaged percentage germination of the three races at each storage condition.

TABLE 21.
Percentage Survival of Uredospores of Races 1, 2 and 6 Stored at 0°C., 3:5° CO. and 10° C. and at 25%, 50% and 75%
RA (in all Combinations) after Storage for One to 13 Months.

0° C. 3:°5° C. 10° C.
Relative Months fala sien eliwetd | et mide Rabie Bendis —
Humidity. Stored. Race Race Race Race Race Race Race Race Race
1 6 2 1 6 2 1 6 2
1 95 97 97 95 99 98 91 99 99
2 94 96 97 96 97 95 92 96 96
3 84 92 90 82 94 90 80 92 87
4 71 83 74 58 84 69 35 41 23
5 47 73 45 44 60 37 16 29 1:6
25% 6 28 55* 26 5 34 12 0:8 0:9 0-1
7 23 49 18 7 27 6 0 0 0
8 8 27 8 2-1 6 1-2 0 0 0
10 4 12 0-8 0-1 0:3 0-1
11 1:3 3-4 1:5 0 0 0
12 1-4 2-9 0:9 ! 0 0 0 |
13 0 0-1 0-0
1 97 98 97 92 97 97 87 98 99
2 97 97 97 95 99 98 90 96 99
3 94 98 95 72 96 97 62 83 95
4 82 95 93 69 94 93 38 29 50
44 —_— — —_ —_— —_— — 22 5 34
50% 5 86 90 91 57 87 94 6 2°5 21
6 85 90 86 54 72 85 1:8 0-2 0:8
7 81 88 87 54 66 84 0 T T
8 55 79 80 13 17 42 0 0 0
10 50 61 76 4-4 2-4 12 0 0 0
11 72 72 76 5 1:3 6
12 58 80 80 20 0:3 3:2
13 44 42 40 1 TY T
1 90 93 91 52 64 65 1-4 15 20
2 58 65 65 io 24 28 0 0 0-1
3 19 26 26 T. 3°6 13 0 0 0
4 B397/ 13 11 0 0-1 0°3 0 0 0
75% 44 4-7 11 17
5 1-5 10 10 0 0 0
6 0 7 11 0 0 0
7 T 6 10
8 Lost
10 0 0 0

* Difficult to assess percentage germination of this lot owing to excessive clumping together of the spores.

Germination above 5% given to nearest 1%. Germination between 1% and 5% given to nearest 0:1%. Germina-
tion below 1% given to nearest 0-:1%.

“1."=Trace germination less than 0:1%.

Graphs 1 to 3 highlight the humidity effect on viability at each temperature. Graphs 4
to 6 were prepared to highlight the temperature effect on viability at each humidity.

Effect of Humidity on the Longevity of Uredospores.
0°C.

The effect of humidity on uredospore longevity was particularly marked at this
temperature. The spores retained a high level of viability for the first month at
75% RH, but deteriorated very rapidly at an almost linear rate during the next two
months to an average of 24% viability. By the end of the fourth, viability had entered
the final lag phase terminating between the seventh and eighth months of storage.

BY H. B. KERR. 277

The spores remained highly viable for the first three months at the lower
humidities, although survival was slightly better at 50% RH. After this time spores:
held at 25% RH began to deteriorate at an almost linear rate to 14% viability at the
end of eight months. There was a prolonged lag phase terminating after about thirteen
months of storage.

A high level of viability was maintained at 50% RH for the first twelve months.
Results at the end of thirteen months suggested that the spores had entered the
intermediate phase of rapid decline. But in view of the rather capricious germination
obtained earlier at eight and ten months’ storage it is possible that the spores were
more viable at the end of thirteen months than the germination figures indicated.

3°5°C.

There was marked deterioration within the first month at 75% RH. This deteriora-
tion appeared to proceed at a linear rate from the commencement of storage until at
least the end of the second month of storage, when viability had fallen to 18%.
Viability entered a brief lag phase terminating before the end of the fifth month
of storage.

Spores retained their viability equally well at 25% RH and 50% RH for the first
three months. After this period spores held at 25% RH began to lose viability at an
almost linear rate until the end of the sixth month, when the three races averaged
17% viability. ‘The curve then passed into a slight lag phase terminating before the
end of eleven months. Spores kept at 50% RH did not appear to enter the intermediate
phase until the end of the eighth month of storage and fell from 68% viability to
24% viability over the next month. There was a moderately long final lag phase and
the spores were virtually inviable after thirteen months’ storage.

10°C.

Spores held at 75% RH were only 12% viable at the end of the first month and
inviable by the end of two months. They retained a high level of viability for the first
three months at the lower humidities, but entered the intermediate phase of very
rapid decline shortly after this. ‘They were inviable at both humidities by the end
of seven months. Although there appeared to be only a slight difference between the
viability curves at 50% and 25% RH results from the individual races indicated
interesting differences.

Effect of Temperature on the Longevity of Uredospores.

25% RH.

Spores retained much the same level of viability at each temperature for the first
three months, then entered the intermediate phase of viabiltiy and deteriorated rapidly.
The rate of deterioration increased with increasing temperature. Spores held at 10°C.
were virtually inviable by the end of six months. There was no obvious final lag
phase. The spores were virtually inviable after ten months at 3-5°C. and after twelve
months at 0°C.

50% RH.

The spores survived equally well at each temperature for the first two months.
From the end of the third month differences between the viability curves at eack
temperature became increasingly pronounced. The spores were almost inviable at the
end of six months at 10°C. and after thirteen months at 3:5°C. They were still
approximately 40% viable after thirteen months at 0°C. As at 25% RH spores held
at 10°C. showed no obvious final lag phase, while spores held at 3:5°C. entered a final
lag phase of several months’ duration.

75% RH.

Spores held at 10°C. and 3:5°C. entered the intermediate phase within the first
month. By the end of the first month spores held at 10°C. had virtually entered the
final lag phase, but spores held at 3:5°C. were less than half-way through the
intermediate phase.

278 MELAMPSORA LINI (PERS.) LEV. UREDOSPORE LONGEVITY AND GERMINATION,

By the end of two months spores held at 10°C. were virtually inviable and spores
held at 35°C. had virtually entered the final lag phase extending until shortly after
the end of the fourth month.

Spores held at 0°C. retained a high level of viability for the first month, but soon
entered the intermediate phase which gave way to the final lag phase between the
third and fourth months. The final lag phase was considerably extended at this
temperature, terminating some time after the end of the seventh month.

g

s

PERCENTAGE CERMINAT TON

20

1 2 3 4 5 6 1 8 9 10 ll 12 13

1 2 3 4 5 6 7
3 NUMBER OF MONTHS STORED

10°C

PERCENTAGE GERMINATION

1 2 3 4 5 6 7 8 9 10 ll ous)
NUMBER OF MONTHS STORED

3.5°C 2

Race Differences.

Since the relative survival of the three races was remarkably constant at each of
the humidities irrespective of the temperature, the humidity-race interaction appeared
to be rather significant. Race differences have therefore been summarized according
to the humidity of the different storage conditions.

25% RH.

The relative survival of the three races remained remarkably constant at each
temperature at each stage of the viability curve. During the initial lag phase race 1
deteriorated faster than races 2 and 6. The last two races maintained much the same
level of viability for the first two months. By the end of the third month each race
was on the verge of the intermediate phase. During the following intermediate phase

BY H. B. KERR. 279
race 6 survived much better than the other two races. This was indicated not only
by its greater viability at each test, but aiso by its more normal spore colour. The
averaged viability of races 1 and 2 at the lower temperatures was for four months
20% lower than the viability of race 6, reaching a maximum difference of 29% after
four months and six months at 3:5°C. and 0°C. respectively. The superior survival
was maintained until the spores of each race had already entered the final lag phase.
Each race, however, became inviable at much the same time.

Deterioration of spore colour was characteristic of these storage conditions. Spore
colour did not deteriorate under the cther humidities or under any other set of
storage conditions during the current studies of longevity, etc. This deterioration was
obvious from the third month, and became progressively more pronounced until the
spores had become inviable and colourless. The spores were studied under the

TABLE 22.
Uredospore Colour after Storage at 25% RH at Three Temperatures.
Temper- Period Stored.
ature. | Race. |—
6 Months. 8 Months. 10 Months. 11 Months. 13 Months.
1 Capucine orange. | Capucine orange | Capucine buff. Pale yellow} Very pale yellow
to capucine buff. orange. orange.
0° C. 6 Orange. Capucine yellow. | Capucine orange. | Capucine orange | Capucine buff.
to orange buff.
2 Capucine orange. | Capucine buff. Capucine buff to | Pale yellow]! Pale orange
pale yellow orange. yellow.
orange.
il Capucine buff, | Capucine buff. Pale yellow | Pale yellow | Very pale yellow
capucine orange. orange. orange. orange.
3°5° C. 6 Mikado orange. Capucine yellow. | Orange buff. Capucine buff. Capucine buff to
pale yellow
orange.
2 Capucine orange. | Capucine buff. Pale yellow! Pale yellow | Pale orange
orange. orange. yellow.
1 Capucine orange | Pale orange | Pale orange | Pale orange | Very pale orange
to capucine buff. yellow. yellow. yellow. yellow.
10° C. 6 Orange buff. Orange buff to | Capucine buff. Pale orange
light orange yellow.
yellow.
2, _—— Pale orange | Pale orange | Pale orange | Very pale orange
yellow. : yellow. yellow. yellow.

Uredospores stored at 50% and 75% relative humidities at all three temperatures retained their original colour
Spores at 50% RH were little affected.

for at least 10 months.

Original colour: cadmium orange.
clature ’’, Plate III.)

microscope.

was impossible to determine accurately the number of colourless spores.

Spores kept at 75% RH then showed slight changes.
(Colour description based on Ridgway’s ‘*‘ Color Standards and Color Nomen-

Owing to the gradual gradation of colour between individual spores it

But the

figures obtained showed that race 6 had a much lower percentage of colourless spores

than the other races.

Careful checks under the microscope during the early stages

of germination failed to discover any colourless spores which had germinated.

50% RH.

Comparison of the relative survival of the three races at the different temperatures
at 50% RH was complicated by the fact that spores stored at 0°C. had only just
intermediate phase when the studies were concluded.

entered the

particularly of race 1 was often highly capricious.
Race 1 appeared to deteriorate faster than the other races during the initial lag

phase and the early part of the intermediate phase.

Germination

Race 6, which maintained much

the same level of viability as race 2 during the initial lag phase, was appreciably less
viable than race 2 during the intermediate phase, and slightly less viable than race 1
during the final lag phase. Race 2 was obviously better adapted than the other races

280 MELAMPSORA LINI (PERS.) LEVY. UREDOSPORE LONGEVITY AND GERMINATION,

for survival at this humidity and was considerably more viable than the other races
in the final intermediate phase from the end of the fourth month at 10°C. and from
the end of the fifth month at 3:5°C.

75% RH.

The rapid deterioration of all races at this humidity obscured race differences.
But the relative survival of the three races paralleled the results at 50% RH. Race 1
deteriorated most rapidly, race 2 the least rapidly of the three races. Races 2 and 6 on
the whole behaved remarkably alike at each temperature, and were appreciably more

TABLE 23.
Percentage of Colourless Spores after Storage under Conditions Listed for Varying
Periods.
Months Stored.
Temperature. Humidity. Race. ———
4* 5 6
1 25 31 45
0° C. 25% 6 10 12 22
2 15 28 51
1 0 0 0
50% 6 0 0 0
2 0 0 0
1 0 0 0
75% 6 0 0 0
2 0 0 0
1 25 32 61
3°5° C. 25% 6 10 15 23
2 20 30 50
1 0 0 0
50% 6 0 0 0
2 (0) 0 0
1 0 0 0
75% 6 0 0 0
2 0 0 0
1 30 42 65
10° C. 25% 6 10 17 25
2 20 30 68
1 0 0 0
50% 6 0 0 0
2 0 0 0
1 0 0 0
75% 6 0 0 0
2 0 0 0
|

* Estimate. The rest obtained from count of 400 spores approximately.

viable than race 1 at each test. The differences in percentage survival at any time of
race 1 on the one hand and races 2 and 6 on the other were greatest at the two
higher temperatures, but the difference in longevity of the two groups was most
pronounced at 0°C., the spores of race 1 being virtually inviable after five months
and the spores of the other two races still appreciably viable after seven months at 0°C.

Notes on Investigations of Uredospore Respiration.

A series of experiments was carried out using the Warburg apparatus to determine
the nature of the respiration curves associated with different phases of uredospore
germination. It was anticipated that these studies would show characteristic differences
between the respiration curves of spores sown on water, gelatin and extract and

PERCENTAGE GERMINATION

PERCENTAGE GERMINATION

PERCENTAGE GERMINATION

BY H.

NUMBER OF WONTES STORED
25% RELATIVE HUMIDITY

ge Stato 7 8 SmnTONNNT
NUMBER OF MONTHS STORED :

5Q% RELATIVE HUMIDITY

4 5 6 7 8 9 lo ll
NUMBER OF MONTHS STORED.

o°c 25% RELATIVE HUMIDITY.

B. KERR. 281

°,

o°c °
35°C @
eo 10°C ™

PERCENTAGE GERMINATION
mt

Oy)
o

ro
°

6 NUMBER OF MONTHS STORED

19% RELATIVE HUMIDITY

Raced ©
RACE@ @
RACEQ,

PERCENTAGE GERMINATION

TAS) 1 2 3 4.5 6 1 8 9 lo
8 NUMBER OF MONTHS STORED

325°C 25% RELATIVE HUMIDITY

rce2 o
RACEG @
RACER @

PERCENTAGE GERMINATION

12 13 1 2 3 4 5 6 7 8 9 10
NUMBER OF MONTHS STORED

9 10°C) =. 25% _RELATIVE HUMIDITY

282 MELAMPSORA LINI (PERS.) LEV. UREDOSPORE LONGEVITY AND GERMINATION,

possibly between Punjab-attacking and non-Punjab-attacking races. Despite many
modifications of technique, spores set up under otherwise optimal conditions failed
to germinate in the Warburg flasks. Spores gave a slight percentage germination
when sown on solution in very small quantities, but such quantities were inadequate
to give an appreciable respiration reading. Spores sown in bulk in a confined space
may produce an excessive amount of germination inhibiting substances or induce
local anaerobic conditions. Under normal conditions spores exposed to the air obtain
an ample oxygen supply and any deleterious substances are probably destroyed by
oxidation or rapidly diluted in the larger volume of solution.

Experiments were then carried out to determine whether dry and moistened spores
of Punjab-attacking race 1 accessions differed in their non-germination respiration
curves from non-Punjab-attacking races. It seemed probable that the difference in
temperature tolerance of Punjab and non-Punjab-attacking races would be reflected
in different curves in the temperature range 15°C. to 35°C., but repeated comparative
studies failed to determine any difference between the two race groups.

TABLE 24.
Uredospore Respiration Studies.
Showing the Rate of Uptake of Oxygen by 0-7 Gramme of Race 1 Uredospores in a Saturated Atmosphere and in
Aqueous Linseed Extract.

Period in minutes since
commencement of ex-
periment ie 6 0 20 40 60 70 80* 90 97°5 | 105 110 115 118

ul. of oxygen absorbed by
uredospores since 0 time | Set 47 97-4 | 149-7) 174 205 236 456 724 1195 | 1386 | 1657

at 0
Check extract .. ths 4-7 6:3 9-4 6-3 | 10-9 | 12-5
Period in minutes since i
commencement of ex-
periment pre oo || Jes 160 165 170 175 255 260 270 280 290 300 310

ul. of oxygen absorbed.. | Reset | 172 355 485 765 Reset | 33 52-3 | 64:5 66 71:5 | 76-5
at 0 at 0

* Extract tipped at 80 minutes.

Attempts were then made to study the respiration of the germ tubes of the two
race groups. Spores were germinated in petri dishes on extract. The weft of tubes
formed after 24 hours was floated off into a large container of water, washed gently
and transferred without distortion onto 7 cm. filter paper. The paper was lightly
rolled into a scroll. Six such papers were inserted into a Warburg flask to obtain a
measurable respiration. Respiration was studied over the range 25°C. to 35°C. The
results failed to reveal any significant difference between the Punjab-attacking and
non-Punjab-attacking races.

Results from one of the early experiments are illustrated in Graph 16. 0-70 gramme
of uredospores of race 1 were set up in a Warburg flask to which KOH had been
added in the central cup and 1 ml. of linseed extract in the side arm. The spores
had been stored for a month and were 80% to 90% viable. A check flask was set up
without spores to determine possible respiration of bacteria in the extract, and two
control flasks were also prepared to determine change in atmospheric pressure. The
temperature of the waterbath was maintained at 27°C. The usual procedures were
followed.

The respiration rate of the dry spores in a saturated atmosphere was measured for
80 minutes. The extract was then tipped onto the spores and the rotating arm set
in motion to disperse the spores with the solution. The results are given in Table 24
and Graph 16.

100

PERCENT AGE GERMINATION

PERCENTAGE GERMINATION

PERCENTAGE CERMINATION
~
°

a)
°o

3

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feo}

vw
fo}

DY
co}

we
°o

100}.

ES

s

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BY

H. B. KERR.

RACEX ©
e RACEG @
wy 0 RACER F
st e
o s &
°o
S
e Vv
e
e
fe}
°
3 4 5 6 7 9 N@ Ne NG)

o°c

4

3.5C

NUMBER OF WONTHS STORED

50% RELATIVE HUMIDITY

b)

6
NUMBER OF MONTHS STORED

7

7
NUMBER OF MONTHS STORED

50% RELATIVE RUMIDITY

19% RELATIVE HUWIDITY

RACEL
RACEG @
RACER WV

8

S

PERCENTAGE GERMINATION
w
°

12. NUMBER OF MONTHS STORED

10° 50% RELATIVE HUMIDITY

PERCENTAGE GERMINATION

1 2 3 4 5 6
| 4 NUMBER OF MONTHS STORED

3.5 C 75% RELATIVE HUMIDITY

g

3s

PERCENTAGE GERMINATION
~
fo}

1 2 3 4 5 6

15 NUMBER OF MONTHS STORED
10°C 79% RELATIVE HUMIDITY

283

284 MELAMPSORA LINI (PERS.) LEV. UREDOSPORE LONGEVITY AND GERMINATION,

Respiration was extremely constant in a saturated atmosphere and the spores were
absorbing oxygen at the rate of approximately 2-5 wl. per minute. Within 20 minutes
of tipping the extract the rate had increased to an average uptake of 66:2 wl. of oxygen
per minute. Measurements were interrupted after 118 minutes and resumed at 155
minutes. Further readings taken until 175 minutes showed that the spores were still
respiring rapidly at a slightly reduced rate. Readings were again interrupted. By
the time they were resumed at 260 minutes respiration had almost ceased.

SUMMARY AND DISCUSSION.
Uredospores survived longer at 0°C. than they did at higher temperatures in
humidities of 25% RH, 50% RH and 75% RH. They survived longer at 50% RH at
temperatures of 0°C. to 10°C. than they did at the lower or kigher humidities.

Uredospores

-T grammes

Absorbed by

Al__of Oxygen

16 20 40 Go 80 foo 8=—.: 120 155 RO 6025 CBO OSD 520

Time in Minutes from Commencement of Experiment

Uredospores stored at 0°C. and 50% RH remained highly viable for 12 months and
were still very viable after 13 months, though by this stage they appeared to have
entered the phase of rapid deterioration of viability. In view of the marked humidity
effects at 0°C. and the marked effect of temperature at 50% RH it seems highly
probable that still more favourable storage conditions could be determined by
experimenting with different combinations of temperature and humidity about a mean
of 0°C. and 50% RH respectively.

The very adverse effect of 75% RH on spore longevity stresses the need for
keeping humidity low, especially at higher temperatures. The very capricious survival
of the spore samples tested in the earlier experiments can almost certainly be
attributed to the effects of high humidity inside the spore-storage phials. Spores
collected during dull rainy weather, most prevalent in winter, have a higher than
usual moisture content. It was discovered during the course of the final tests that
spore samples collected under conditions of high humidity and stored in bulk in

BY H. B. KERR. 285

cork-stoppered phials often condensed small droplets of water inside the container.
This moisture disappeared within or shortly after a week. Serious deterioration would
undoubtedly occur under these conditions, the degree of deterioration probably being
directly proportional to the time of exposure to such humidities. Since this would be
determined by such highly variable factors as the original moisture content of the
spores, the spore volume and tube volume, and tightness of fit of the stopper variation
between samples would be considerable. This would certainly explain why spores
collected in winter and spores stored in considerable bulk usually deteriorated faster
than spores collected in spring or summer and stored in small quantities.

Low relative humidity had a decidedly deleterious effect on uredospore longevity
at 0°C. and 3:5°C. This effect, judging by the degree of deterioration of spore colour,
was just as pronounced at 10°C. But at this temperature spores of race 6 and to a
lesser degree spores of race 1 survived better at 25% RH than at 50% RH. Low
humidity caused a deterioration of spore colour, destroying during the process some
substance or substances responsible for normal germination. These substances may
have been part of the colour complex or may have been destroyed simultaneously
with these substances. Normal germination was accompanied by a gradual disappear-
ance of spore colour. MInviable spores retained their colour for long periods after
viable spores of the same sample germinated and became colourless. Abnormally
germinating spores commonly extruded the orange pigment into the germ tubes, but
no trace of orange pigment was ever found in the germ tubes of normally germinating
spores. Spores which had become colourless during storage at 25% RH appeared to
be totally inviable. The colour substance or substances tolerated a wide range of
conditions and were normally not destroyed during storage. They were therefore
rarely a limiting factor in the process of germination.

The rate of deterioration of the substances did not seem to be much affected by
temperature. At none of the temperatures was deterioration of colour or viability
marked prior to the end of the third month. But after this period there was rapid
deterioration, the rate increasing slightly with increasing temperatures.

The better survival of spores of races 1 and 6 at 25% RH as against 50% RH at
10°C. was particularly interesting. It hardly seemed likely in view of the pronounced
colour deterioration at 10°C. that this temperature checked the adverse effects of the
lowest humidity. It seemed rather that the lowest humidity countered at another level
the adverse effects of the highest temperature.

Under most storage conditions loss of viability probably resulted from a fall-off in
the energy-rich reserves following normal resting respiration. The rate of deterioration
would certainly have increased with rising temperature and rising humidity. Low
humidity probably countered this deterioration, permitting the spores to maintain a
higher level of energy-rich substances for a longer period than was possible at 50% RH.
'The existence of this reserve, however, became irrelevant for the maintenance of
viability once the viability-determining colour complex had dropped below the critical
level for maintenance of normal germination. In the case of race i, and more especially
race 6, deterioration of the colour substance was probably sufficiently delayed for
‘spores held at 25% RH to gain a slight survival advantage over spores stored at
50% RH. But this advantage was soon cancelled out and spores of both races
became inviable at much the same time at both humidities. Race 2 reacted differently.
At all stages of the intermediate phase of viability its spores were appreciably more
viable at 50% RH than at 25% RH. Since it is most unlikely that different processes
‘of deterioration occurred with the different races, the following seems to be the most
logical explanation. Race 2 possessed a higher level of respirable reserves than
races 1 and 6. This seems to be clearly indicated by a comparison of the viability
‘curves of this race and the other two races at the higher humidities. On the other
hand, race 2 deteriorated much more rapidly than race 6 and slightly more rapidly
‘than race 1 at 25% RH. These two factors would militate against 25% RH exerting
‘a positive influence on the maintenance of respirable reserves before deterioration of
the colour substance began to undermine viability of race 2 spores.

286 MELAMPSORA LINI (PERS.) LEV. UREDOSPORE LONGEVITY AND GERMINATION,

It is consistent with the data to assume that two different processes of deterioration
operated at 25% RH. The one process was conditioned by the low humidity, and
during the intermediate and final lag phase, but not the initial lag phase, was slightly
accentuated by rising temperatures. It affected a substance or substances which
comprised or were part of the complex responsible for spore colour. The other process
of deterioration was probably directly correlated with spore respiration and was
stimulated by rising temperature and humidity. At 50% RH and 75% RH loss of
viability was probably due to the latter process only.

It would be interesting in future experiments to determine the effect of humidities
slightly below and slightly above 25% RH on the longevity of spores stored at 10°C.
It is probable that lower humidity would increase the rate of deterioration of the
colour viability substance and cancel out the beneficial effects on spore respirable
reserves before any benefit to viability might accrue to the spores. On the other hand,
a slight rise in humidity above 25% RH might swing the balance in favour of longer
survival of all races at 10°C. at such a humidity as against a humidity of 50% RH.
Increase in temperature above 10°C. should favour greater longevity at 25% RH than
50% RH, since this sheuld cause deterioration of the respirable reserves to become a
limiting factor in germination before the colour complex had deteriorated below its
critical level for maintenance of normal germination, assuming that a rise in tempera-
ture would have a more pronounced effect on spore respiration than on deterioration
of the colour complex, since deterioration of spore colour is determined by humidity
rather than temperature.

Although races did not differ greatly in their longevity under a wide range of
storage conditions they did noi behave identically. Race differences seemed to be
correlated with humidity rather than temperature effects. The most significant dif-
ferences were observed at 25% RH. Race 6 exhibited a remarkable tolerance to the
adverse effects of this humidity on spore colour and viability. Race 1 deteriorated
faster than the other two races under most conditions of storage. It may have had a
lower respirable reserve than the other races. Race 2 deteriorated less rapidly and
remained viable slightly longer than the other races at the higher humidities. It may
have carried a greater reserve of respirable products. On the whole the races differed
only slightly in the over-all nature of their viability curves. They generally entered
the intermediate phase of rapid decline as a group and became inviable at much the
same time. There is certainly a need to use pure races in determining the effect of
different storage conditions on uredospore viability, especially during the intermediate
phase of viability. But the results given by any one race should be fairly typical
of the species as a whole.

The lower the temperature in the range 10°C. to 0°C., the better the survival at
each humidity, particularly at 50% RH. ‘This did not agree with results obtained by
Prasada (1948). He found that spores survived best at 5°C. to 7°C., and almost
equally well at 10°C. to 15°C. as at 0°C. His uredospores became inviable after
22 weeks at 5°C. to 7°C. The difference between Prasada’s results and these studies
is probably not due to differences in the races used. Races differed only slightly under
a wide range of storage conditions. The differences can probably be attributed to the
germinating media used. It is assumed that Prasada used water or some other inert
germinating media. The notoriously capricious germination of stored spores on such
media, especially water, emphasizes the need for using host extract to assess viability.

In the normal process of spore metabolism the resting spore respires and uses up
its respirable reserves. At the start these reserves appear to be greatly in excess of the
minimum required to sustain maximum germination. During storage the level of the
respirable reserves probably fails gradually to a certain critical level below which
further deterioration may seriously affect the capacity of the spores to germinate.
At this stage it is imperative that the spores germinate as rapidly as possible before
the impact of high humidity and intimate contact with the germinating solution has
so stimulated the respiration rate that the reserves fall below the level needed to
sustain vigorous germination. Anything tending to delay germination would cause
a serious deterioration in germination. Such a delay, among other things, could be
caused by storage-hardening of the spores. This might be due to physical or physio-

BY H. B. KERR. 287

logical deterioration of the spores tending to make them impermeable or dormant.
Storage-hardening generally developed after several months of storage. It increased
with age. But some samples became severely storage-hardened after only a few weeks
of storage. The very capricious germination (after prolonged storage) of spores of
race 1 kept at 50% RH at O°C. and 3:5°C. suggested that this humidity induced a
greater measure of storage-hardening than other humidities. But this may only have
been due to aging of the spores which remained viable longer under these conditions
than they did under other storage conditions.

Aqueous host extract countered the storage-hardening of the spores, stimulating
the spores to germinate before serious deterioration of the respirable reserves had
undermined the capacity of the spores to germinate. The physiologically active
substance or substances were present in or comprised the oily fraction of the extract.
The substances were readily extracted by boiling, but did not diffuse into solution
when cold extracts were prepared by macerating tissue in a Waring blendor. The
complex was not destroyed by prolonged boiling, but was destroyed by charring. It
was ether-soluble and adsorbed by charcoal. The stag-horn development of the germ
tubes at temperatures within the optimal range may have been a simple tactile response
to the aqueous extract. It may, however, have been conditioned by the same substance
or substances which stimulated germination. It should supply a simple means of
determining temperature tolerance of different races.

Poor germination on gelatin and water was probably due to the fact that the
preliminary period of conditioning of the spore contents leading up to actual emergence
of the germ tubes was unduly prolonged by the storage-hardening of the spores. This
permitted a considerable deterioration of respirable products. The better germination
on gelatin could not be attributed to its physiological activity, but suggested rather that
the physical spore-solution relationship played an important part in regulating the
germination of storage-hardened spores. The rate of uptake of water by the spores
may vary according to the germinating medium used.

References.

BaiLey, D. L., 1928.—Sunflower Rust. Tech. Bull. Minn. Agr. Exp. Sta., 16: 31 pp.

BEAUVERIE, J., 1924.—Sur la Germination des Uredospores des Rouilles du Blé. C. R. Acad.
Sci. Paris, 179: 993-996.

CHESTER, K. STARR, 1946.—The Cereal Rusts. 269 pp.

CHiu, W. F., and WALKgmR, J. C., 1949.—Physiology and Pathogenicity of the Cucurbit Black
Rot Fungus. Jour. Agr. Res., 73: 589-615.

CHupp, C., 1917.—Studies on Clubroot of Cruciferous Plants. N.Y. Cornell Agr. Eup. Sta.
Bull. 387 pp. —

Fior, H. H., 1935.—Physiologic Specialisation of Melampsora lini on Linum usitatissimum.
Jour. Agr. Res., 51: 819-837.

Hart, H., 1926.—Factors Affecting the Development of Flax Rust, Melampsora lini (Pers.)
Lév. Phytopath., 16: 185-205.

Hwane, L., 1942.—The Effect of Light and Temperature on the Viability of Uredospores of
Certain Cereal Rusts. Phytopath., 32: 699-711.

Keer, H. B., 1952.—Rust Investigations in Linseed. Nature, 169: 159.

NoBLE, R. J., 1924.—Studies on the Parasitism of Uvrocystis tritici Koern., the Organism
Causing Flag Smut of Wheat. Jour. Agr. Res., 27: 451-490.

PRASADA, R., 1948.—Studies in Linseed Rust, Melampsora lini (Pers.) Lév. in India. Jndian
Phytopath., 1: 1-18.

RAEDER, J. M., and Brver, W. M., 1931.—Spore Germination of Puccinia glumarum, with
Notes on Related Species. Phytopath., 21: 767-789.

RAPER, K. B., and ALEXANDER, D. F., 1945.—Preservation of Molds by the Uyophil Process.
Mycologia, 37: 499-525.

SHarp, EH. L., and SmirH, F. G., 1952.—Preservation of Puccinia Uredospores by Lyophilization.
Phytopath., 42: 263-264.

SHARVELLE, HE. G., 1936.—The Nature of Resistance of Flax to Melampsora lini. Jour. Agr.
Res., 53: 81-127.

THIELE, A. F., and WelIss, F., 1920.—The Effect of Citric Acid on the Germination of the
Teleutospores of Puccinia graminis tritici. Phytopatih., 10: 448-452.

WATERHOUSE, W. L., and Watson, I. A., 1943.—Further Determinations of Specialization in
Flax Rust Caused by Melampsora lini (Pers.) Lév. Proc. LINN. Soc. N.S.W., Ixxvil:
138-144.

WILcOxoN, F., and McCaAtLuan, S. E. A., 1934.—The Stimulation of Fungus Spore Germination
by Aqueous Plant HMxtracts. Phytopath., 24: 20.

288

ON SOME PERGINE SAWFLIES RHARED BY MR. M. F. LHASK
(HYMENOPTERA, PERGIDAB).

By Rosert B. BENSON, British Museum (Natural History), London.
(Communicated by Mr. K. EH. W. Salter.)
(One Text-figure.)
[Read 26th November, 1958.]

For many years now Mr. M. F. Leask, of Ballarat, Victoria, has been rearing and
studying Pergine sawflies and sending me the adults to name and samples of pickled
larvae for the British Museum collections. The opportunity thus afforded of seeing
long series of some of the species has shown me that I had underestimated the range
of variation of some of them in 1939 (Benson, 1939) and therefore below submit some
new synonymy.

I am indeed much indebted to Mr. Leask and it is with very great pleasure,
therefore, that I am now able to name after him a fine new Perga, one of his very
latest discoveries.

PERGA LEASKI, sp. nov.

Colour: Yellow becoming orange on head above and down-turned lateral portions
of tergites; black are tips of mandibles, ocellar area and middle part of postocellar
area, middle fore lobe of mesonotum (except for the impunctate patch behind}, lateral
lobes of mesonotum (except for the declivous sides), postscutellum, mesosternum and
lower margin of mesepisternum, middle and hind coxae, all abdominal sternites
together with sawsheath; and black with metallic violaceous lustre is the whole of the
upper side of the tergites except the extreme lateral margins. Wings flavescent with
yellowish-brown stigma and venation.

Length: 16:5 mm.; forewing 12-5 mm.; antenna 2 mm.

Pubescence on head, thorax, coxae and first tergite long and yellow (up to about
as long as the medial length of the clypeus).

Head (fig.): Shining between scattered punctures, denser on antennal crests.
Clypeus slightly emarginate medially. Antenna 6-segmented, almost as long as distance
between eyes in front; flagellum as long as width of clypeus; club as long as three
preceding segments together. Antennal crests large, rounded, subtriangular, not clearly
defined medially where they are scarcely separated from each other. Malar space very
short, only about half as long as second antennal segment. Hind ocelli about twice as
far from back of head as from each other. POL: OOL as 3: 2.

Thorax: Pronotum shining between large shallow punctures. Mesonotum covered
with small punctures about as far apart as their diameters, but becoming much denser
on the sides of the front lobe and the middle of the lateral lobes; hind portion of the
front lobe, however, has a raised impunctate area. Scutellum normal, convex without
medial furrow or depression, about 1:6 times wider than long; and together with
under-thorax between the scattered regular punctures with shining interspaces, medially
two or three times wider than the diameter of a puncture. Legs with very short tarsi;
hind tarsus little more than half as long as tibia (1:0: 1-8); basitarsus about as long
as three following tarsal segments together. Wings normel.

Abdomen: Shining but with transverse alutaceous sculpture above. Setae on sides
of sawsheath normal. Saw with about 80 marginal teeth, similar in pattern to that of
Perga dahlbomi (Benson, 1939, fig. 38) and other species of the same species group.

Clunes, Victoria 19 (holotype), bred from Jarva 20.11.1958 (larva no. 500), M. F.
Leask. (In British Museum.)

PROCEEDINGS OF THE LINNEAN SocieTy or NEw SoutH WALES, 1958, Vol. Ixxxiii, Part 3.

BY ROBERT B. BENSON. 289

In Benson (1939) this species would run to Perga brevipes Forsius, which is
undoubtedly the most closely related to it of known species, being the only other
known Perga with the head and thorax clothed in long pubescence. The new species
has longer antennae, almost as long as the distance between the eyes in front (in
P. brevipes an antenna is only about 0-8 this distance). It differs from P. brevipes also
in its flavescent wings, the dark violaceous tergites above and the denser punctation
of the thorax.

SOOO

Head of Perga leaski, sp. nov., from above.

PERGA KOHLI Konow and P. BRULLEI Westwood from Victoria.

Among other species reared by Mr. Leask in Victoria in 1958 were specimens of
Perga kohli Konow, previously only known from Queensland, and P. brullei Westwood,
previously known only from Western and South Australia.

SYNONYMY IN PERGAGRAPTA.

In Pergagrapta much material reared by Mr. Leask in Victoria gives an indication
of the possible range of variation in certain species and justifies the following
synonymy:

Pergagrapta glabra Kirby (= malaisei Benson, syn. nov.).

Pergagrapta turneri Benson (= hackeri Benson, syn. nov.).

Pergagrapta bella Newman (= nigra Benson, rossi Benson, and rohweri Benson,
syn. nov.).

Legends to Figures in Benson (1939).

In my previous paper (Benson, 1939) the legends to the figures were lost and did
not appear with the paper. They should have been as follows:

(Page 325.) Figs. 1 and 2: Labium and maxilla of Cerealces (fig. 1) and Perga
(fig. 2). Figs. 3-7: Antenna of Acanthoperga cameronii (fig. 3), Perga dorsalis (fig. 4),
P. brullei (fig. 5), Xyloperga halidaii (fig. 6), and Cerealces scutellata (fig. 7).

290 ON SOME PERGINE SAWELIES.

(Page 327.) Figs. 8-11: Portion of forewing to show stigma, radial and cubital
cells in Acanthoperga (fig. 8), Pergagrapta (fig. 9), Pseudoperga (fig. 10), and Perga
(fig. 11).

(Page 328.) Figs. 12-21: Mesoscutellum of Xyloperga halidaw (fig. 12), Paraperga
jucunda (fig. 13), Acanthoperga cameronii (fig. 14), Pseudoperga lewisii (fig. 15),
Pergagrapta bella (fig. 16), P. bicolor (fig. 17), P. spinoiae (fig. 18), Perga dorsalis
(fig. 19), P. kirbii (fig. 20), and Perga dahlbomiu (fig. 21). Figs. 22-27: Head from
above of Xyloperga perkinsi (fig. 22), X. halidaii (fig. 23), Pseudoperga lewisti (fig. 24),
Acanthoperga cameronii (fig. 25), Perga dorsalis (fig. 26), and P. brullei (fig. 27).

(Page 335.) Figs. 28-29: Apex of sawsheath from below in Perga affinis (fig. 28)
and P. dorsalis (fig. 29). Figs. 30-32: Incrassata bristles from sawsheath of Perga
affinis (fig. 30), P. dorsalis (fig. 31), and P. konowi (fig. 32).

(Page 336.) Figs. 33-39: Portion from middle of saw of Cerealces scutellata (fig.
33), Xyloperga perkinsi (fig. 34), X. amenaida (fig. 35), X. univittata (fig. 36),
X. halidaii (fig. 37), Perga dahlbomii (fig. 38), and P. affinis (fig. 39).

(Page 343.) Figs. 40-44: Portion from middle of saw of Antiperga antiopa (fig. 40),
Pseudoperga guerinii (fig. 41), Pergagrapta bicolor (fig. 42), P. bella (fig. 43), and
P. latreillei (fig. 44).

Reference.

BENSON, R. B., 1989.—A Revision cf the Australian Sawflies of the Genus Perga Leach sens.
lat. (Hymenoptera Symphyta). Awstralian Zoologist, 9 (iii): 324-57, 44 figs.

291

THE DIPTERA OF KATOOMBA. PART 2.
LEPTIDAE AND DOLICHOPODIDAE
By G. H. HArpy.
(Nine Text-figures.)
[Read 26th November, 1958.]

Synopsis.

Keys, discussions and descriptions concern chiefly, in Leptidae, Dasyomma flava Hardy and,
in Dolichopodidae, Heteropsilopus squamifer and Sciapus tumidus, n. spp. The females of
Arachnomyia longipes and Sympycnus separatus Parent are described, and some errors found
in literature are corrected.

INTRODUCTION.

The late O. Parent, publishing on Australian Dolichopodidae (1913-55), did not
realize how variable were the characters used in his descriptions, and the subsequent
attempts to identify from these have led to difficulties not foreseen by him. However,
he examined many of the types of early authors, confirming much synonymy previously
deduced from descriptions (Hardy, 1930), and he reduced to synonymy some of his own
names.

Most types of the Australian species are in the collection of the C.S.1.R.0.,
Canberra, and I am indebted for much information concerning these received, with
grateful thanks, through Dr. A. J. Nicholson, who also supplied the number of
specimens that were collected by the late A. L. Tonnoir.

Family LEPTIDAE.

Five genera were included under Leptidae by White (1914) but Metoponia (Hardy,
1919) was removed to Stratiomyiidae and two other genera were added. The inclusion
of Clesthenia White* was questioned (Hardy, 1921), leaving five genera only.

In a suggested new classification for the lower Brachycera (Steyskal, 1953) two
of these five genera, Atherimorpha and Austroleptis, were removed to Hrinnidae
(Xylophagidae) and this transfer was discussed (Hardy, 1955). It is anticipated that
these two small families, Leptidae and Hrinnidae, will become subfamilies under one
family unit when the world’s fauna becomes better understood. Three Australian
genera remain in the restricted unit, namely Spaniopsis, Dasyomma and Chrysopilus.

From the Blue Mountains only two species have been recorded, Dasyomma flava
Hardy (Blackheath) and Spaniopsis clelandi Ferguson (Wentworth Falls); three
more species are added below.

Key to genera and species of Leptidae.

1. Venation of wing complete: M, reaches the wing border .....................-.++---- 2.
Venation incomplete: M, reduced to a spur vein or absent. Anal cell closed. Abdomen
COTNGRU 6 ono Serer Sido tal ig. oun RRO TET OF OSGEE OEY noe oc EC REcn n CRE ERG eN RCE in nee CRO SCNORR CRC aM SPANIOPSIS. 3.

2. Anal cell open at wing border. Abdomen conical ........................ DASYOMMA. 5.

Anal cell closed. Abdomen iong and slender, tapering or about parallel-sided ...........
Ee ee Rrra ey csike ung atanan, Sreteh cals eu aes aahrel ohleg age eee onan ev cmsuen arcuate Chrysopilus aequalis Walk.

* Clesthenia aberrans White, found only on windows, was described as from both sexes,
but in the various collections in Australia only female specimens were to be seen. Some of
these look like possible males because of abdominal shrinkage, and as White only used a hand
lens when describing Diptera, this shrinkage may have deceived him, and he states that no
difference occurs between the sexes. White’s type male specimen needs investigation. A male
before me (13th January, 1955) has its hypopygium somewhat upturned (probably displaced),
conical with a pointed apex and is certainly Asiloidean in affinities. This specimen was
captured in the bush by sweeping in the Hobart suburb of Taroona and is the only male
specimen that I have seen.

PROCEEDINGS OF THE LINNEAN Society oF NEw SouTtH WALES, 1958, Vol. Ixxxiii, Part 3.

E

292 DIPTERA OF KATOOMBA. II,

8. Wings with the radial field area fuscous from the stigma to the wing tip. Antennae with
the style about one and a half times the length of the three segments ...............

REE OCI CSE ON ORO ache Coen GSU S cll Suemeranoseo Pike cho ERnler a ‘c oneturh yue.s Oreos olulalay S. marginipennis Ferg.
Wines) entirely, hyaline Wyjeeere ene chet yer ciency crevice totel eure en anchen seen acheteitereeler cu oley -Iteiiel ctiatctaish eee -eentens 4,

4, Antennae with the unsegmented style about twice the length of the three segments combined
BRE ACES CITE Ga Ce DIO OOO OFOrNE DG Dio Oia cio Gebinelasein conten biota do orcimoc ao S. longipennis Ferg.
Antennae with the style about the length of the three segments combined .. S. clelandi Ferg.

5. Legs dusky brown and the abdomen black on the male (the only sex known). (Coastal

EE Wat oY: 9) [Mn PERC RR rte erate a ogre bate eS 1 Ais CO ee Geol et ete eas oA BES LM D. abdominalis Hardy.
Legs bicoloured on both sexes. Abdomen black on male and normally yellow on female.
CMiounitaiin vara) iis sdehs ere SR Se eocmeigs a eects bese eek te ee Sprs tede Sh sleet es tetoeedehs D. flava Hardy.

CHRYSOPILUS AEQUALIS Walker.
Leptis aequalis Walker, 1848; Hardy, 1919 (Chrysopilus).
Hab.—Katoomba: occasional specimens occur around swamps from October to
December.

SPANIOPSIS MARGINIPENNIS Ferguson, 1915.

Hab.—Katoomba: 1 9, 21.4.1958.

Note——This specimen was carried onto the veranda and became trapped at a
window there. Sweeping shrubs in the area failed to yield more specimens, and the
presence of this species in the Blue Mountains was quite unexpected. Ferguson records
it from Gosford and Milson Island, occurring in May and June, and he gave the date
five weeks later than the present record.

SPANIOPSIS LONGIPENNIS Ferguson, 1915.
Hab.—Katoomba: 1 9, 30.1.1955.
Note.—This is the only specimen found, and occurred in company with S. cleland,
along the creek that runs to Minna-Ha-Ha Falls.

SPANIOPSIS CLELANDI Ferguson, 1915.

Hab.—Katoomba, numerous females are scattered from December to early April,
but no males have been found. These flies occur widely and regularly but not
concentrated in large numbers. They appear most frequently along mountainside
seepage areas.

Genus DAsyomma Macquart.

This genus seems to have species that are very variable in their colour characters,
but material so far collected is very scanty. The Tasmanian specimens have hairy
eyes, and mainland ones have them bare. The Tasmanian D. similis Hardy, 1919, was
found recently on Mt. Wellington (1 J, 12.1.1953, 1 9, 8.1.1953), adding to the scanty
material previously known.

From the mainland comes one species, a female from Blackheath, but the additional
material found at Katoomba indicates that the sexes are dimorphic and there is a
considerable variation in the colour pattern.

Malloch (1932) gives twelve specific names for South American specimens, nine
of them under new names (five species are based on unique specimens) without
attempting to ally the sexes. Under his grouping the Australian species come into
the typical Dasyomma, one of the three subgeneric groups proposed.

DASYOMMA FLAVA Hardy, 1933. (Text-fig. 1.)

do. Eyes bare, contiguous and with a well-defined line separating the small facets
below antennal level from the enlarged ones above. The face and triangular area of
frons, extending to the postocular region, are black-brown covered with a pulverulent
grey.

Thorax deep brown with black hairs and its pleura is black, varying to brown,
with long dark propleural hairs. Hairs on the mesopleura and metapleura are arranged
in a row of about seven each. The abdomen is entirely black with black hairs.

The coxae are black, this colour extending normally to the basal third of the
femora of the first two legs, and two-thirds of the hind femora, but varies to having
the colour restricted to the base of them all. The apex of the femur and the whole

BY G. H. HARDY. 293

of the tibiae are black. Each tarsal segment is black apically and the two last may
be entirely black. Elsewhere the legs are yellow. Light slender hairs occur on the
anterior coxae, dark ones on the intermediate, and there is an apical row of densely
packed bristly hairs on the hind coxae. One apical spine usually occurs on all tibiae
of both sexes, whilst a second very small spine, when present, is not very noticeable.

The wings are slightly grey with a faintly yellowish stigma, this latter colour
extending basally on the wing membrane.

Text-fig. 1—Dasyomma flava Hardy. Head, (a) front view, (b) side view of male,
(c) front view and (d) side view of female.

Text-fig. 2—Parentia tricolor Walker. Head of male.

Text-fig. 3—Heteropsilopus cingulipes Walker. Head of male.

Text-fic. 4—H. squamifer, n. sp. (a) Head of male, (6) head of female, (¢) hypopygium,
and (d) the two rows of hair fringes, hooked and spatulate, that extend full length of the
intermediate tarsi.

294 DIPTERA OF KATOOMBA. II,

©. Similar to the male, but the thorax and abdomen are yellow-brown. The head
is similarly coloured with the frons as wide as an eye-width, parallel-sided. A
conspicuous groove extends from the black-brown ocellar tubercle to the slightly
bulging triangular area just above the antennae.

Hab.—Katoomba: Allotype male (7.2.1957), paratype male (15.12.1957) and two
females (24.12.1956, 8.2.1957) form the basis of the above description, whilst other
specimens yield the variations in characters recorded. The latter bear on the label
the following dates: 1 9 (24.1.1956) which has the summit of the head slightly collapsed,
the abdomen blackish (presumably stained) and shrunken; 1 ¢ (612.1953), 1 9
(11.1.1952) which are normal in colour except that the femora have little black on
them, and the specimens are somewhat dilapidated.

These seven specimens were collected from windows, but a search in the neighbour-
hood on swamps, seepage areas, creeks, waterfalls and rocks has failed to yield more
specimens.

Family DOLICHOPODIDAE.

Six specific names have been applied to species from the Blue Mountains area,
namely, Diaphorus intactus and Sciopus plumifer (= ingenuus Erich.) from Springwood;
Condylostylus amoenus (= nigropilosus Macq.) from Mt. Victoria (all in Becker, 1922).
To these were added Sciopus ingenuus Erich. and Heteropsilopus cingulipes Walk.
(Hardy, 1930, 1951). Parent (1932) added Sympycnus callidus Par. from Mt. Wilson.

Subfamily CHRYSOSOMATINAE.

A satisfactory classification for Australian species of the Dolichopodidae has not
yet evolved, and the papers by the late O. Parent followed the general plan that
Becker used. This results in closely related species being placed in different genera
of the Chrysosomatinae, and some in Condylostylus, which does not occur in the
Australian region. The genus Chrysosoma is a complex of species well represented in
the tropical regions, and is unknown in the Katoomba area. Heteropsilopus extends
more widely and appears to form a homogeneous group that extends to Africa
(C. petersi and C. bacchi Dyte, 1957 (Hnt. mon. Mag., 93: 37) conform in wing character
and type of terminalia —see Hardy, 1953, ibid., 89: 7). Sciapus is another complex
genus covering some of the species originally placed under Condylostylus, the
remainder of which are now placed in Parentia.

Key to Genera.

1. Male with hook-shaped cilia along the costa of the wing and the first radial vein reaches
level with or beyond the apex of the median cell. Female normal in these respects.

Two pairs of apical scutellar bristles in both sexes ...................... PARENTIA.

Male with normal cilia along costa and the first radial vein is short, conforming with
the) femaile. in, these Characters. ~~ recipe sisec ieceia ecg cm Shenae re ate ose ose reece eel oie tem sway cust arene ame nee 2.

2. Median cross-vein of wings strongly sinuous, often angulated and with a spur vein. One
pair of apical scutellars but a second very weak pair may occur .... HETEROPSILOPUS.
Median cross-vein straight or slightly bowed. In a few species very Slightly sinuous. One

or two pairs of apical scutellar and occasionally discal bristles occur ....... SCIAPUS.

PARENTIA TRICOLOR Walk. (Text-fig. 2.)

Psilopus tricolor Walker, 1835; Hardy, 1935 (Parentia) (synonymy). Psilopus
nigropilosus Macquart, 1847; Hardy, 1935 (Parentia) (references). Condylostylus
amoenus Becker, 1922.

Synonymy.—Parent found that the type of gemmans Walker, 1849, conforms with
amoenus Becker, 1922, and that the type of tricolor Walker, 1835, was missing. The
description under the last name suggests that tricolor will be given precedence, but
at present it is not certain that specimens under the name nigropilosus also include
a second species to which one of the synonyms may apply.

Hab.—Katoomba: 1 4, 25.9.1955. This species is widely distributed from Tasmania
to New South Wales and, under the name gemmans Walker, from Western Australia
too, but apparently it is not often found in mountainous regions.

BY G. H. HARDY. 295

Genus HETEROPSILOPUS Bigot.

Bigot, 1859; Hardy, 1935, 1951.

The three species that occur in the Katoomba area are discussed below and are
readily identified on the male by characters contrasted in the key. The females conform
in wing characters with the males and can be readily allied, in so far as the ocellar
tubercle gives a good guide to affinities where there may be some doubt.

Key to species.

1. Male with a complete row of hook-shaped cilia, about as long as the tarsal width, on the
CACC Otters der ilGor ts eencammet sae ine ctsc eine eo sut hb aie seciere tne ace che sarciicr enti: eaten a eneyeene: e0 2.

The tarsi of the middle leg on male peculiarly formed; the two apical segments (often
appearing as one) flattened and bearing bristly hairs. The hypopygial tergite contains

a hook-shaped part on the short process. Both sexes have the wing marks reaching to,

or near to, the wing tip, at least as far as the apex of the middle radial branch,
though pale on newly emerged specimens. Ocellar tubercle normal in form ...........
ee ee Pere IAS Shes) Sab ee ndere SE ete Sea stte: sates: GP als GES aie ingenuus HErichson.

2. Male with a row of flattened, scale-like, hairs, adjacent to the cilia, along the full length
of the tarsi. Without an adjunct to the process of the hypopygial tergite. Wing marks

vary with age from clear wings (newly emerged) to banded, but not reaching the
wing tip. An oceasional exception in very old specimens shows a faint shadowing
reaching near to the tip. The ocellar tubercle is small, appearing like a small dome
TMOUMMCC COMMS VAG OMS) yo oe aycicchicrs seer ee ey fo fewest a oe stvener-n) cues laure’ siento: Sitasioh-ay ais) ey ie; tous aems yay is Squamifer, n. Sp.

Male without scale-like hairs adjacent to the cilia of the tarsi. Tergite of hypopygium
with a long prolongation of the process. On both sexes the wings are hyaline, and

the ocellar tubercle is very high and Slender .................... cingulipes Walker.

HETEROPSILOPUS CINGULIPES Walker. (Text-fig. 3.)

Psilopus cingulipes Walker, 1835; Hardy, 1935 (references and synonymy)
(Heteropsilopus ).

Parent records that Walker’s species is missing, and so the type is of uncertain
identity. Notwithstanding this, the form had been identified as identical with the
species which is normally found around Sydney and has the clear wings, as well as
agreeing with the description in other respects. The identity seems certain, but not
so all the synonymy so far applied.

It seems impossible te separate from descriptions those synonyms confirmed by
Parent, of which sydneyensis Macq. is the earliest published name, but it must be
noted that a difference occurs in the illustrations given by Becker and by Parent.

Synonymy.—Iin the notes below, the synonymy is clarified respecting plumifer
Becker, the description and illustrations being evidently based upon a complex that
proved difficult to resolve. In addition, doubt may arise concerning angulosa Bigot,
1890, and metallicum Parent, both of which are given as 6 mm. in length, whereas the
normal variation in size for the present form is 7 to 8 mm. The 6 mm, is not outside
possible variation for the species, and no other known form seems to be in agreement
with the descriptions, so as to permit removal of those two names.

Hab.—Katoomba: 1 2, 31.10.1955. Other specimens examined are from the Sydney
area. The species seems to be rare on mountains. The records give New South Wales,
the Federal Capital Territory and Victoria. Macquart gives Tasmania, but that was
an error, and probably should have been Sydney.

HETEROPSILOPUS SQUAMIFER, nh. Sp. (Text-fig. 4.)

Sciopus plumifer Becker, 1922, in part (wing figure 184); Hardy, 1935 (Hetero-
psilopus). Heteropsilopus cingulipes Hardy, 1951, in part.

Synonymy.—Becker erected the species plumifer on a complex found to correspond
with ingenuus. In 1952, another species agreeing with Becker’s species regarding the
wing figure was found to have a clear-winged form and cilia of the middle tarsi as on
cingulipes. A careful survey of very long series collected over several years showed
this new species squamifer was consistent in certain characters, showing a contrast
with both forms with which it had been confused, and these characters are given in
the key above.

Becker’s figure 183 is that of ingenwus, but missing (? hidden) the hook-shaped
process, and no mention is made of the middle tarsi in the description. The series of

296 DIPTERA OF KATOOMBA. II,

5 g and 2 Q is sufficient to contain the two species which are common and occur
together.

Description.—A blue-green metallic species, varying with some coppery colour and
with a reflecting, slight silvery, overlay, has abdominal cross-bands both basal and
apical on the male, and only apical on the female. At least the two basal segments
of the antennae, the mouth, most of the legs and the black-tipped lamellae are yellow.
The front coxae are yellow, the others are metallic. The tarsi are fuscous, which
colour extends onto the hind tibiae of the male, but if occurring there as a separate
band this does not exceed half the tibial length.

The wings vary from clear on newly emerged specimens to deeply marked,
including two bands crossing the area, as in Becker’s figure 184, and other markings
are also similar. The bands may be complete, fragmentary, very pale, but never joined
together. The marking may tend to extend along the costa towards the apex, but if
so it is only like a slight shadow in that area, and occurs on old specimens and may
deepen a little after death.

One pair of black bristles occurs, with a slightly shorter one on the summit near by
on the male; the latter bristle is very short on the female. Hairs behind the eyes
are white. Thoracic bristles are not consistent in arrangement, but normally 4
acrostichal (any additional ones are weak); 3 anterior dorsals; 1 each humeral and
posthumeral; 3 notopleural; 1 supraalar, which may be absent; 1 postalar and 1
scutellar pair. Submarginals of the abdomen are few and rather long, and all hairs
are white except on the legs, where they form black cilia in rows with an occasional
one outstanding and bristle-like. The femora are without bristles, but the tibia of
the middle leg has them, about 3 on male and 10 on female being discernible. On
the male the middle tarsi have a row of hook-shaped hairs that spread to, and grade
down in length on, the tibia, and a row of adjacent flattened scale-like hairs which
are pointed apically are limited to the tarsi.

Hab.—Katoomha: 36 ¢ and 36 2 form the type series. In addition about 40 more
of each sex have been examined. These occurred over several years, the 14th December
and ist February being the first and last dates of capture during the 1955-6 season.
Flood rains followed, ending in a short drought in 1956, when the 1956-7 season became
due. Few of these flies were seen during the last few days of December, then again
were missing from their haunts until the middle of January, lasting to the 1st March,
1957, but were not plentiful as in previous years.

Sydney: In the Australian Museum is a male specimen labelled ‘Northbridge,
19. Oct. 1924 (B. Bertram)’, bearing the typical wing marks and conforming in
terminalia.

Note on figures—The drawing of the hypopygium was made from a slide-mounted
specimen. Occasionally on drying the filaments become distorted and do not show the
true shape, and the outline of the short process may vary slightly in shape. The
filaments shown may be varied to slightly wider, but in form they resemble those of
H. cingulipes; those of H. ingenuus are much broader and a little shorter. The process
on the latter species (Text-fig. 7) is shown here with the hook-shaped adjunct which
is missing in the figure given by Becker, but which gives the filaments correctly formed.

HETEROPSILOPUS INGENUUS Erich. (Text-fig. 5.)

Psilopus ingenuus Hrichson, 1842; Hardy, 1930 (Sciapus) (references); Hardy,
1935 (references), 1951 (Heteropsilopus). Psilopus trifasciatus Macquart, 1849; White,
1916 (Sciapus); Parent, 1932c, 1933 (Sciopus); Becker, 1922 (Chrysosoma); Parent,
1932a (Chrysosoma); Hardy, 1935, 1951 (Heteropsilopus). Sciopus plumifer Becker,
1922, in part, Fig. 183 (terminalia). Sciopus gloriosus Parent, 1932a.

Synonymy.—Two species involved under plumifer Becker include the description
and figure of the terminalia which correspond with the present species, on which the
wing markings tend to vary. The normal markings are those given in Macquart’s
figure, but Parent redrew from Macquart’s specimens, giving a very expansive pattern,

BY G. H. HARDY. 297

not known on recent specimens so far seen. Possibly Parent’s figure was drawn from
another species and wrongly named. Parent’s other figures correspond with his
gloriosus, which is a synonym of trifasciatus acknowledged by him.

Hab.—Widely distributed over Tasmania to New South Wales and South Australia.
It is very abundant throughout the Blue Mountains, and the dates of capture at
Katoomba are from 4th November to the 8th March, but during the retarded season
ending in 1957 the last found was on 10th April.

b
d

oo

C

fo)
0°
v

d

Text-fig. 5.—Heteropsilopus ingenuus Erichson. (a) The normal form of the face and
frons, (b) the abnormally wide variation of the same, (c) the female head, (d) the hook-
shaped extension on the process of the hypopygium.

Text-fig. 6.—H. proximus Parent. Head of male.

Text-fig. 7.—Sciapus tumidus, n. sp. (a) Head cof male, (b) head of female, (¢) inter-
mediate tarsi of male, and (d) hypopygium.

Text-fig. 8—Arachnomyia lungipes Parent. Head of male.

Text-fig. 9—Sympycnus anomalipennis Becker. Part of the underside of the wing showing
the position of hairs each side of the vein R,;. m.c.: median cell.

Genus Scrapus Zeller.
Sciopus of authors.

Parent described as new, eleven species, four on males, six on females, and only
one on both sexes. He later reduced two of these names to synonymy, but in one
case he may have made an error. The key to species given here covers these eleven
species, and two of these, together with one new species, are found in the Katoomba
area. In addition, a species runs to mollis (2 9, December, 1955) but does not agree
with the description, and one runs to nigrociliatus (3 9, December, January, 1955, and
January, 1957); these latter names were based on two females each, and their identity
may be difficult to determine with certainty until the male is known. The new species,
tumidus, does not agree with any description but appears to be nearest to
quadrimaculatus.

298 DIPTERA OF KATOOMBA. II,

Key to Parent’s species of Sciapus.
1. Wings with markings. Coxa I yellow. Abdomen with basal bands on segments. Median

CROSS=ViEinRS Eradis te PLES coke whe sieht Toye ay OES eve eee Ste CRATES ANE UTTOS te esa eecreerin Sette feet pateLie pe Reet 2.
Wings without markings, at most slightly and uniformly darkened .................... 4.

A, Ai YE ClOrSOCemina)! lnisWles, I. m@mEeMel sscoscco sec 0dcge so ac soso nH ODDS EOaDaFODdS 3.
With four dorsocentral bristles. R, diverging at apex (= discretifasciatus according to
Parent, but this may be an error). Q@ueenslanidl, 2a cseners: syoksee ues e genevievei Co.

3. Wings with markings broadly extending along costa joining the two bands (when typical,
but may be reduced). New South Wales .......................+-c0-. proximus 3, Q.

Wing markings reduced to four spots, two reaching the costa. New South Wales ........

Bo Pe Sead a aS NIC RE OR GAMERA TICS Gl BEE PLR cllet tapraychs event cet etal Re er set Opam ais eletel Sih ceive mera ote Ore quadrimaculatus ®@.

4. Coxae I and II yellow, 4 dorsocentral bristles, 1 vair scutellar. Abdomen yellow at base.
Median cross-vein straight. New South Wales .......................... zonatus @.

Onay” GOR MLE VST OW. MI roc eteaaes oe res tse eae SCl A SPELT Oe ere Me trees Hea ae oY oc a ee red De aie c ti oe ice aM Shape earn 5.
All coxae dark. Five dorsocentrals, 2 scutellar pairs. Abdomen dark with broad basal
black bands. Median cross-vein straight (= nigrofasciatus). New South Wales ......

ry de ha iece b ce teh ogee ys sesh ch tm coeare isteactcreine eats Meee a Sine teams ate wore en Gta ogc pean site ee coy ei koeeiinge mney ee eye es regale co.

5. Median cross-vein slightly sinuous. Abdomen dark at base ..................2....--: S)-
Median cross-vein straight or slightly bowed. One pair of apical scutellars ......... 6.

6. Scutellum with discal bristles present. Five dorsocentrals. Abdomen dark at base. New
Souths Wiles pie ey veers ae ce ate Bem ane Upa ann eect Rey a ene ure chetiscutatum ®.
Scutellum without vdiseal bristles’ — sxeeyciveyeescsie mts ties tec aoe more ere) Guevara eu Su ARC ney eae Ue

7 Abdomen: syellOw “at ASE 9 sbis ha elsekid 2b hee s2 srepe als euereae een Ae esh pceertsl onebewel ©) shines cya eheheieseeeeanehee ne 8.
Abdomen entirely dark. Four dorsocentrals. Tasmania ............... nigrociliatus ¢.

8. Dorsocentrals 3, and the two radial lower veins diverging at apex. Queensland ..........
GALA ee NCHS cles ee ae A delvan erecs ede Saat eee TAS. ue ened sate we Mees ao Set cee te ere graciliventris &.

Dorsocentrals 4. Radial veins normal. New South Wales ......-............. mollis 9°.

9. One pair scutellar bristles. Dorsocentrals 4. Abdominal segments with apical bands. New
SOU WIALEST .Peyestte = Aoccystcuauacavenaho leis uedortat nace nett ak chan eae cae ieee onreel oeai ate Stirrcy ie erage sordidus 9°.

Two pairs of scutellar bristles. Dorsocentrals 6. Abdomen without bands. New South
AAV 2 TYR ie ehecick oes acece uC RAC CRE UES) CRORE ERTL ORME CCRC eae OR a Luce iericek Gasiaun tart yen difficilis @.

SCIAPUS DISCRETIFASCIATUS Macquart.

Psilopus discretifasciatus Macquart, 1849; White, 1916 (Sciepus); Hardy, 1930
(Sciapus); Parent, 1932a (discussion), 1932d (synonymy) (Sciopus). Sciopus depinctus
Becker, 1922. ?? Sciopus genevievei Parent, 1932a (probably a different species).

Synonymy.—The name depinctus Becker can be applied to this species with
certainty, but Parent saw Macquart’s specimens, and stated they were his genevievei,
giving no other comment. The wing pattern and description given by Macquart show
the wing tip is clear, whereas this is not so on Parent’s form.

The distinction Parent makes between the descriptions of Becker’s and Macquart’s
forms can have little value. This rests on size and abdominal bands. The largest
depinctus before me is 4 mm. (4:5 in Macquart) and the thin basal bands are present.
Becker gives 3 to 3-5 mm., which corresponds with specimens that have a shrunken
abdomen hiding the bands, this shrinkage being quite common on pinned specimens.

S. genevievei Parent is from Hidsvold (T. L. Bancroft), which district seems too
far north in Queensland for a species that appears to be confined to cooler regions,
and the name is likely to prove applicable to a distinct species.

Hab.—Katoomba: a series of 25 specimens, all females, were collected between
8th November and 22nd December, 1955-6, mostly found by sweeping low vegetation,
others occurring on windows and on leaves of trees.

Note.This species, which is very common in New South Wales, is liable to have
the wing pattern very pale, varying to practically clear winged. It seems curious
that, so far, no males have been found in the Katoomba area, though quite common
elsewhere.

ScIAPUS PROXIMUS Parent. (Text-fig. 6.)

Parent, 1928, 1933 (Sciopus); Hardy, 1930.

Hab.—Katoomba: 1 @, 21.12.1955, and 1 9, 18.12.1956.

Note—The female entirely misses the broad strip that, on the male, joins the
two transverse bands along the costa. It is evident that considerable variation occurs
in wing pattern because the pattern is recorded by Parent as being reduced in area
on the female.

BY G. H. HARDY. 299

SCIAPUS TUMIDUS, n. sp. (Text-fig. 7.)

Metallic green with blue and coppery reflections and with a restricted slight
whitish overlay. Mouth parts and halteres yellow. All coxae metallic, except on the
female the front ones are yellow. Femora yellow on female, but metallic on male with
the apex and sometimes trochanters yellow. Tibia and metatarsus entirely yellow, the
rest fuscous.

Dark incisions at the base of abdominal segments may occur on the male, which
has the hypopygium curiously formed and may appear to differ in accord with the
angle of view. The main tergite is long and the filaments are long and slightly curved.
Two apical appendages are present, the upper one is like a flattened scale and the
lower one is angulated, sinuous and bristle-like.

The bristles are slightly variable in the series, the rows normally having 3
acrostichals, 2 being presutural, 5 dorsocentral with the middle one very small, 1 or 2
humeral, 1 posthumeral on female, 2 notopleural and a line of 3 supraalar (possibly
the intraalar is included with this), 1 postalar and 1 or 2 pairs of apical scutellar
bristles. The apical bristles of the abdominal segments are not many but stand out
slender, long and black on the female.

The front coxae have only light-coloured hairs, and groups of long scattered hairs
are on the basal half of the femur and no bristles or long hairs occur on the tibia
and metatarsus. The middle femur is similar, and the tibia may have two bristles on
each dorsal, anterior, ventral and posterior surface. The male has, on the tarsi, two
outstanding long hairs on the first segment, and a row of small setae, three or four of
them outstanding. The second segment has a long bristle apically placed, and the
third is short and slightly curved. The fourth segment is relatively long with a
laterally flattened swelling on the basal half. The fifth segment is normal. On the
female these tarsi are normal in form with small apical bristles. The hind legs are
normal, with only apical bristles on segments, except one or two may occur on the
anterior surface of the tibiae and occasionally additional small bristles.

The wings are usually quite clear on the male, only one specimen having a faint
trace of marking. The female has shadow marking on cross-veins. The median cross-
vein is practically straight and is 13 to 2 times the length of the lower median branch
vein. One wing is aborted, showing a strongly bowed median cross-vein with detached
ends, and is displaced in position; also the lower median branch is occasionally
incomplete.

Hab.—Katoomba: 15 J, 14 9, 7th to 20th November, 1955. Nearly all these were
frequenting seven small red-currant-bushes in the mid-morning sunshine. Others
occurred on leaves of fruit trees, and the species has not been seen at other times.

Note.—The species shows considerable variation in chaetotaxy, which is a feature
liable to be greater than showing on the comparatively small series available for study.

Subfamily DOoLICHOPODINAE.

When the family is divided into the eight or nine subfamilies normally adopted,
no clarity is reached regarding generic relationships, so the partial rearrangement
previously employed (Hardy, 1939) is adopted here, bringing Neurogoninae and
Medeterinae into this subfamily. Into it come all genera, other than the Chryso-
somatinae, that have the male hypopygium free, not being embedded in a prior (sixth)
abdominal segment. This arrangement simplifies the taxonomy and may lead towards
a better understanding of genera.

ARACHNOMYIA LONGIPES Parent. (Text-fig. 8.)

Pleuropygius longipes Parent, 1933; Hardy, 1939 (Arachnomyia).

©. Agrees with the description of the male, except the narrow frons is parallel-
sided; the third antennal segment is shorter, being as long as wide at base; the
abdomen tapers to a slender apex and coloured as on male, except one specimen which
misses the yellow at the base.

Hab.—Katoomba: 1 ¢ (31.12.1956) and 4 9, two dated the same, the others being
14.1.1956 and 15.1.1957. ;

300 DIPTERA OF KATOOMBA. II,

The holotype was recorded from Victoria without other location, and these
additional specimens form the allotype and paratype females. Two were found on the
wall of the laboratory and a search on tree-trunks produced one other. One was on a
window in the laboratory.

Subfamily SyMPYCNINAE.

This is Campsicnominae of Becker, and is here extended to include all genera
which have the male hypopygium appearing to be embedded in a prior segment of the
abdomen. Lundbeck (1912) described it as somewhat embedded. The sixth sternite
forms a slight hollow to receive it, and this arrangement leaves exposed the entrance
to the genital cavity. The whole terminalia are quite syuimetrically arranged.

It is not clear yet which genera of the various proposed subfamilies belong to this
grouping, but probably Raphium Meigen, 1803, Hydrophorus Fallen, 1823, Diaphorus
Meigen, 1824, and Sympycnus Loew, 1857, together with most of the genera associated
with them, will come under one or two subfamily groups acceptable to most authors.

Genus SymMpycnus Loew.

Synonymy.—Genus Liparomyia White, 1916, is possibly Sympycnus, as the descrip-
tion of L. sedata White is near that of S. scitulus Parent, also from Tasmania. White
states that the hypopygium is curved forward under the venter for only a short
distance and this agrees with the genus. Also it seems to run to Sympycnus in the key
to Dolichopodidae given by van Duzee (1930), in “Diptera of Patagonia and South
Chile’. However, the description does not conform with any of the species described
by Parent.

Becker named one and Parent sixteen species, the latter being based on a limited
series in the Tonnoir collection. Fourteen are based on males only, and some defectively
described, making subsequent recognition difficult. The prescutellar depression is given
as absent from both Australian and New Zealand species, but all forms seen have that
depression. Some defects in Parent’s paper are rectified below, but others may yet
be found.

Key to species recorded from New South Wales.

These nine forms have the normal six dorsocentral bristles, and the approximate proportion
of the median cross-vein, given in parentheses, is based on ten units for the lower branch of
the median field, such as (6:10), but slight variations may occur.

1. Male with abnormal wing venation (3:10). Two hairs, or patches of hairs, occur on the
membrane, about level with the median cross-vein, one each side of the radial sector
on the under side of the wing. Female with normal wing venation (10:10). Both sexes
have a greenish-blue median thoracic stripe which occasionally may be a little obscure.
Without acrostichal bristles. Abdomen partly yellow. Third antennal segment little
longer than its maximum width. Front tarsi and tibia equal in length ..............
PARE OR OORT SCH Oe BOOT ech CoC BREE Ree Oa Coe Cae ec ORCA EAU ce Che OM LONE Oo anomalipennis Becker.

Wing venation normal on both sexes, and without the blue median thoracic stripe .... 2.

2. Species largely yellow and without acrostichal bristles. Front tarsi little longer than
til] 0} Fz ee eee AR reece ere ca CRI ccna eae Io MORE RCRA Seok ceca CCNA Seve APO CeO Ora SS 6 od 3.
Species metallic: ox dill tcoloumede | Sessck ioe ecsveas enone ome tease erica euevones a etemeneie ie ee eachoy ones eeeuenette 4,

3. Wings black at apex on male, clear on female (5:10). Abdomen dark with yellow side-
SPOUESHS fais oi PAR eh Ree AS Rta) cha eerc ee lone Solr Peaclleb one tyapie vena veut hem due nm meneneue sa areReetes marginatus Par.

Wings clear on both sexes (8:10). Abdomen yellow with broad black cross-bands .....
aL AT RICH oUt Rear atc oetatiist Caen a Ncee RiCy Sener aE MEA tea: qe Aircel lence icrteuy cinco okorg U-cyatarars G-o, caw ben callidus Par. oC.

4; Waithoutvaerostichaly bristles: \G6xLiONi wm Gise se hycatn eheWamobsker ate Metelkce Sie Fone edt cuclsLE ACEC UES uee iemeareueres 5.
With acrostichalsbristless Metallickspecicsi | see een neo eee eee 6.

5. Thorax metallic blue. Third antennal segment short. Front tarsi 1% times longer than
UID VE? | Faeete saves, “rrawe tenee adsl iotoon atray Secs tee kee ORR weTie) MORE c Cotnepicice CoE a POORER Meee ycaret tole setifemoratus Par. Cc.

Body entirely brown, varying to biack in parts. On male the third antennal segment is
twice maximum width, short on female. Front tarsi little longer than tibia. Eyes
below antennae contiguous on male, narrowly separated on female ..... separatus Par.

Brown species with blue frons and some yellow on the second abdominal segment or more.
Eyes separated on face. Third antennal segment short. Base of middle femur on male
with a group of outstanding hairs, sometimes tuft-like. Front tarsi slightly longer
than: bias?) sce ste eed Ri ein ae ce So gee eto a wiicoueesnentelonion allectorius Par.

6. Eront tarsi 24: times’ ‘the: Tene th’ of tility isc tvs cee cert ris er ee tereeeticr cen bree erence 7.

Front tarsi almost twice the length of tibia (6.10) ..................... infimus Par. ¢.

BY G. H. HARDY. 301

7. Front metatarsus distinctly more than half the total tarsal length, and with a small apical
ROTIGCHIOM (CSS). -obbesoor hones saeco conGo ooo od. ooOp om oaloo pple oo claudicans Par. ¢.
Front metatarsus only half the tarsal length, and without the apical projection (12:10)
Reed Sanaa nrai cess cniey cies one sicsy ss) ar tare SVeREWe) Suche rap omb oro Neprepeeyatestys ley. aiidith yoerlar a /ehehls eles capilliger Par. co.

Three species in the key occur at Katoomba, and amid other forms not identified
is a new species near S. praecipuus Becker (New Guinea).

The two figures of anomalipennis given by Becker and by Parent would suggest
that two species are standing under the name. Marked differences occur in these
drawings regarding the venation. The veins may alter slightly, but none have been
found with the venation as ziven by Becker, where the longitudinal veins appear to
be misplaced. Possibly the hairs on the wing membrane, a new character, may
help to isolate the males if two forms have heen incorporated.

SYMPYCNUS ANOMALIPENNIS Becker. (Text-fig. 9.)
Becker, 1922; Hardy, 1930; Parent, 1932a.
Heb. Katoomba, 2nd September to 22nd March, 1955-6.
This is quite a common species widely distributed over New South Wales, and
occurs even in winter. It is plentiful at Kateomba during November.

SYMPYCNUS ALLECTORIUS Parent, 1932a.

Correction.—A letter from Dr. A. J. Nicholson confirms the view that the holotype
has aborted front tarsi and that Parent’s figure of the wing does not belong, but
should have been attributed to S. tasmanicus Par. Seven of the nine males in the
Tonnoir collection have simple front tarsi and those collected at Katoomba agree,
except one has tarsi as illustrated by Parent and two are abnormal to a lesser degree.

When pinned and placed in store this species deteriorates more than normally.
Disintegration takes place more readily than with the other species under conditions
of the Katoomba climate. A cure was found in letting the specimens dry in the open
for three or four days before storing, a treatment that greatly reduces the trend for
antennae, legs and even head becoming detached. Artificial drying results in brittleness,
making subsequent handling hazardous. Description of this species awaits a longer
series in better condition than those now available.

SYMPYCNUS SEPARATUS Parent, 1932a.

®. Body colour, including head and coxae, entirely dark brown, deepening to
blackish in parts. Legs yellow-brown. Face narrow, normally parallel-sided and with
a transverse depression. Third antennal segment very short with the arista dorsally
placed.

One pair of long ocellar bristles, and another pair, shorter, on summit. About
10 postoculars and a few white hairs behind eyes. Thoracic bristles are 1 each humeral
and posthumeral, 6 dorsocentrals, 2 each notopleural and intraalar, 1 each supraalar,
postalar and scutellar pairs. Abdomen with long hair-like marginal bristles.

The legs have few bristles, only the apical row on front and middle, and a strong
outstanding bristle on rear coxae. In addition to apicals of femora and tibiae there are
only four outstanding dorsal bristles occurring on the hind tibiae.

The tarsi are simple, those of the first leg about one and a quarter, the middle
about equal to, and the hind leg three-quarters of their respective tibial length. ‘The
metatarsi are respectively half, half, and one-third of their total tarsal length.

The wings have the upper median branch about parallel with the lowest radial
branch, and median cross-vein proportional to the lower median branch vein is
slightly over 6: 10.

Hab—Katoomba: The earliest date of collecting was 25th September, 1956, and
the last in any season is dated 14th May, 1955. October and November are months of
greatest abundance, and after the middle of January they become very scarce.

About 200 specimens have been examined, of which 25 per cent. are males. The
females retained include the allotype and numerous paratypes. The Tonnoir collection
has six males.

302 DIPTERA OF KATOOMBA. II.

References.

BreckKgErR, TH., 1922.—Capita Zool., 1: 238 pp.
Bicot, J. M. F., 1859.—Ann. Soc. ent. Fr., 3: 215, 224.

, 1890.—Idem, 10: 285.
WRICHSON, W. F., 1842.—Arch. Naturgesch., 8: 273.
FERGUSON, E. W., 1915.*—Proc. roy. Soc. N.S.W., 49: 233-243.
Harpy, G. H., 1919.*—Proc. roy. Soc. Tasm., 119-129.
1921.*—Proc. LInn. Soc. N.S.W., 46: 285-6, 300.
1930.— Aust. Zool., 4: 124-134.
1933.*—Proc. LInn. Soc. N.S.W., 58: 408-9.
1935.—Idem, 60: 248-256.
—, 1939.—Idem, 64: 350-1.
1951.—Idem, 76: 223-5.
1955.*—Ent. mon. Mag., 91: 193-6.
Macquart, J., 1838-55.—Diptéres Exotiques, 2 vols., 5 suppl.
PARENT, O., 1926.—Ann. Soc. sci. Bruxelles (B), 46: 16, 18.
1928.—WMitt. zool. Stats. zool. Mus. Hamburg, 43: 191-8.
, 1929.—Ann. Soc. sci. Bruxelles (B), 49: 199, 201.
1932a.—Idem, 52: 105-176.
1932b.—Stettin ent. Ztg., 93: 233.
1932ce.—Ann. Soc. sci. Bruxelles (B), 52: 216-231.
1932d.—Bull. Mus. Hist. nat. Paris, 4: 875, 879.
1933.—Ann. Soc. sci. Bruxelles (B), 53: 170-187.
1934a.—Ann. Mag. nat. Hist., (10) 138: 1-38.
—, 1934b.—Encycl. ent. Dipt., 7: 114.
STEYSKAL, G. C., 1953.*—Ann. ent. Soc. Amer., 46: 237-242.
WALKER, F., 1835.—EHnt. Mag., 2: 471.

, 1848.*—List. Dipt. Brit. Mus., 1: 216.

, 1849.— Idem, 3: 644.
————., 1852.—Insecta Saundersiana Dipt., pp. 209, 211.
WHITE, A., 1914.*—Proc. roy. Soc. Tasm., pp. 38-47.

, 1916.—Idem, pp. 247-258.

* The references concerned with Leptidae have an asterisk, the remainder are concerned

with Dolichopodidae.

’

303

BAT-INFESTING ORNITHODOROS (1IXODOIDEA-ARGASIDAE) OF THE
ORIENTAL-AUSTRALIAN REGION.

By L. J. DumMBLETON, Entomology Division, Department of Scientific and
Industrial Research, New Zealand.
(Communicated by Dr. J. W. Hvans.)
(Highteen Text-figures.)
[Read 26th November, 1958.]

Synopsis.

Confirmatory evidence based on the reticulated structure of Haller’s organ in the larvae
supports Hoogstraal’s recognition, on adult characters, of a group of bat-infesting tick species
within the genus Ornithodoros. The genus Reticulinasus Schulze is reduced to rank as a
subgenus of Ornithodoros. Another member of the subgenus, O. solomoivis, is described from
the Solomon Islands.

Hoogstraal (1953) has drawn attention to a group of Oriental bat-infesting
Ornithodoros which are characterized in the adult stage by small size, pyriform shape,
small inconspicuous discs, mammillated integument and absence of eyes, cheeks, dorsal
hood and dorso-central tarsal humps. The species included are O. salahi Hoogstraal,
1953, O. batwensis Hirst, 1929, and possibly O. piriformis Warburton, 1918, which,
however, has distinct discs. Kohls (in litt.) considers that the Philippine species
mentioned by Hoogstraal is identical with batuensis.

The only other Oriental-Australian bat-infesting species known to me are Argas
steini Schulze, 1935, which for reasons given below is considered to be an Ornithodoros,
and another species from the Solomon Islands which is described below. Both of these
are known only in the larval stage.

In studying the Solomons larva it was found that Haller’s organ possessed the
characteristic reticulated structure of the capsule which Schulze (1941) described in
A. steini, for which he erected the genus Reticulinasus. The larvae of O. salahi and
the presumed larva of O. batwensis have similarly reticulated capsules. The larvae of
O. piriformis are not known. :

Hoogstraal’s grouping indicated on adult characters is therefore confirmed by the
characters of Haller’s organ in the larvae. Schulze’s genus Reticulinasus is reduced
to rank as a subgenus of Ornithodoros.

CHARTOTAXY OF TArsuS I IN ArRCASID LARVAE. (Text-figs. 1-5).

In the species of Argas and Ornithodoros studied the following setae are constant
in occurrence:

All ventral setae, viz., apical ventral (AV) 1 pair; basal ventral (BV) 1 pair;
mid-ventral (MV) 1 pair. In Argas spp. (Text-figs. 4, 5) these latter are basad of the
capsule whereas in Ornithodoros they are level with or basad of the trough.

Three pairs and 1 single dorsal seta, viz., apical (A) 1 pair; basal (B) 1 pair;
paramedian capsular (PC) 1 pair — level with or basad of the capsule; posterior median
(PM) —a single seta basad of and forming a triangle with PC.

There is more variation in the occurrence and position of the lateral setae and
the remaining dorsal setae. The lateral setae comprise anterior (AL) 1 pair—the
only lateral setae present in salahi and batuensis (Text-fig. 2) and the two species of
Argas (Text-figs. 4, 5) but absent in O. capensis (Text-fig. 3); posterior (PL) 1 pair —
basad of the capsule or posterior hair tuft, the only lateral setae in capensis and in
solomonis occurring in addition to AL.

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304 BAT-INFESTING ORNITHODOROS OF THE ORIENTAL-AUSTRALIAN REGION,

The remaining dorsal setae are: distal median (DM) —a single seta distad of
trough on the subapical tarsal prominence in four Ornithodoros spp. but absent in two
Argas spp.; posterior hair tuft (HT) —shorter setae with the bases membranous or
in a Membranous area and close together, situated between PC — absent in A. persicus,
2 in A. vespertilionis and O. solomonis, 3 in O. batuensis, salahi and capensis; pre-basal
(PB) 1 pair — present in A. persicus only.

HALLER’S ORGAN IN ARGASID LARVAE. (Text-figs. 1-5, 10, 11, 14, 18.)

The anterior trough or pit and the capsuie may be considered as constituting
Haller’s organ in a narrow sense but the following setae are in close and constant
relation to it: DM, PC, PM, HT. The anterior trough or pit is a membranous circular
or oval area posterad of the subapical prominence bearing up to eight conical or

Text-figs. 1-5.—Larva, tarsus I, chaetotaxy, dorsal of 1, O. solomonis; 2, O. batuensis ;
3, O. capensis; 4, A. persicus; 5, A. vespertilionis.

finger-like setae or sense cones. In the species studied it shows no obvious diagnostic
characters. The capsule is posterad of the trough. It consists of a subspherical pocket
or cavity, with heavily chitinized walls and floor, invaginated into the tarsus and roofed
with a thinly chitinized or membranous cover with a transverse oval orifice anteriorly.
Three or four modified finger-like setae arise from the wall of the capsule basad.
They are often difficult to distinguish and do not appear to offer diagnostic characters.
The capsule varies in shape from circular to transverse or asymmetrical, from shallow
to deep and narrow-necked and in position from a simple vertical-walled invagination
to an obliquely basad invagination, and these variations offer good specific characters.
In the subgenus Reticulinasus of Ornithodoros the walls and floor of the capsule bear
regular and strong reticulations, the roof or cover also has the reticulations well
developed marginally but they become indistinct as they approach the orifice of the
capsule.

Bat-infesting species are more numerous in North and South America, but none
of their larvae are so far reported as having a reticulated capsule. Schulze (1941)

BY L. J. DUMBLETON. 305

states that it was not present in O. hasei Schulze (? O. dunni Matheson). Reticulinasus
may prove to be a subgenus restricted to the Oriental-Australian regions. The
reticulated capsule is not reported from the genus Argas and is not present in larvae of
A. persicus and A. vespertilionis which I have examined.

Bopy CHAETOTAXY IN ORNITHODOROS LARVAE. (Text-figs. 6, 7.)
The following number and arrangement of setae is constant in the subgenus
Reticulinasus. Dorsal setae consist of 14 pairs: 4 pairs anterior —2 submedian and
2 sublateral; 2 pairs forming a transverse row before mid-length; 3 pairs sublateral;

Text-figs. 6-9.

a

6, 7.—Larva, body chaetotaxy of O. solomonis. 6, dorsal; 7, ventral.
8, 9.—Adult, Haller’s organ, dorsal of 8, O. batwensis; 9, O. salahi.

2 pairs forming a transverse row behind mid-length; 3 pairs posterior sublateral or
marginal. The ventral setae consist of 13 pairs; 1 pair on each coxa; 3 pairs paramedian
between coxae II and coxae III; 1 pair paramedian preanal close together; 1 pair
paraanal; 2 pairs paramedian postanal.

Other species of Ornithodoros, for example capensis and the species figured by
Cooley and Kohls (1944), show variation in the number of pairs of dorsal setae between
7 (turicata) and 18 (coriaceus) and capensis has more numerous ventral setae,
especially in the postanal field.

306 BAT-INFESTING ORNITHODOROS OF THE ORIENTAL-AUSTRALIAN REGION,

TARSAL CHARACATERS IN ORNITHODOROS ADULTS. (Text-figures 8, 9.)

The adults of only two species, salahi and batuensis, of the subgenus Reticulinasus
are correlated with the larvae. The adults of both differ from talaje and capensis in the
more numerous lateral setae on the sides of tarsus I. The depression of the roof of
the capsule below the level of both the trough and the posterior hair tuft, which is
common to both salahi and batwensis, may be a group character. MHaller’s organ is
similar in batwensis (Text-fig. 8) and salahi (Text-fig. 9) —four setae in the posterior
hair tuft, trough with eight sense cones, distal median setae two unequal. The capsule
of these two species does not differ markedly, except in shape, from other Ornithodoros
species.

The differences between the two species are that in salahi (Text-fig. 9) the tarsal
setae are more noticeably plumose and the capsule is asymmetrically invaginated basad
while in batuensis (Text-fig. 8) the capsule is invaginated vertically.

The occurrence may be mentioned here of modified ventral setae on tarsus I of
adults of O. gurneyi Warburton.

DESCRIPTIONS AND NOTES ON ORNITHODORUS LARVAE OF THE SUBGENUS
RETICULINASUS (Schulze, 1941).
ORNITHODOROS SOLOMONIS, n. sp. (Text-figs. 1, 6, 7, 10, 11-13.)

Larva (engorged).—Length, including capitulum 1-:9-2-2 mm., width 1:1-1:2 mm.
Colour (in life) slaty-grey. Shape elongate with slight constriction at level of
coxae II and stronger constriction immediately caudad of coxae III. Integument finely
striate except for smooth oval median area at mid-length on dorsum. Body chaetotaxy
(Text-figs. 6, 7) identical with that of other members of the group. The posterior
dorsal setae are submarginal though the median pair may reach the margin. These
setae, although the larva is larger, are shorter than those of salahi and batuensis.
Leg length 1:16 mm. Tarsus I total length including claw 0:48 mm., to base of
claw 0:34 mm. Tarsal setae (Text-fig. 1) 18 pairs plus 2 setae in posterior hair tuft.
Distal median seta (DM) apparently inserted on trough. Two pairs of lateral setae
AL and PL. Haller’s organ (Text-fig. 10) with capsule (Text-fig. 11) elongate, ovoid,
with floor walls and periphery of roof reticulated. Basis capituli with 2 paramedian
post-hypostomal setae and 2 longer post-palpal setae. Hypostome (Text-fig. 12) length
0:18 mm., dentition 2/2 with 8 teeth in outer file and 8 rounded teeth in inner file.
Palpi with joints 2 and 3 longer than wide, apical joint twice as long as wide, sides
subparallel, apical setae not half as long as joint.

Type.—Larva on slide mount in author’s collection. Paratypes: 15 larvae on slide
mounts in author’s collection. Other material: 10 larvae in Rocky Mountain Laboratory
collection.

Type Locality—Gill’s Plantation, Joroveto, Vella Lavella Island, Solomon Islands.
Coll. L.J.D.

Host.—Fruit bat ex cave. Larval ticks standing on head in caudal region of
dorsum, one inch from tail. Larvae of the Trombiculid mite Whartonia vellae (Dumb.)
also present.

O. solomonis may be separated from related species as follows:

(1) Palpal joint 4 twice as long as wide. Strong posterior marginal setae absent. Posterior

Inewie wi Wikia 4 SEAS coooanodoob po sooogoDD Gono oD DOGOF OOD ODOOS O. solomonis, n. sp.

Palpal joint 4 little longer than wide. Strong posterior marginal setae present. Posterior
hair stb! aywaitla3! GSCCACS. s. eve csua eves eae oo erencue owen ds soci ec oche paleo smouerewememoraue tte cicitcaet sy ame keine Montene (2).

(2) Capsule of Haller’s organ elongate, pyriform ....................... O. batwensis Hirst. -
Capsule of Halleris organ Subcircular .:..............-.-..-.-----= O. salahi Hoogstraal.

(O. steini Schulze is inadequately described but would apparently fall in couplet 2.)

ORNITHODOROS BATUENSIS Hirst. (Text-figs. 2, 14-17.)

Larva (presumed) (engorged).—Length 1:0 mm., width 0:‘7 mm. Body chaetotaxy
conforms to that of the group. Tarsus I (Text-fig. 2) length to base of claw 0:2 mm.,
chaetotaxy identical with that of salahi. Strong posterior marginal setae as in salahi.
Hypostome (Text-fig. 16) length 0:08 mm., dentition 2/2 with 6-7 teeth in outer file
and 6 in inner file. Haller’s organ (Text-fig. 14) with capsule (Text-fig. 15) elongate,

BY L. J. DUMBLETON. 307

pyriform, narrower basally. Palpi (Text-fig. 17) with joints 2 and 3 subquadrate, as
wide or wider than long; apical joint tapering, little longer than wide, apical setae
nearly as long as joint.

The specimens examined were collected 28/7/55 by J. L. Harrison from Batu Caves
(type locality), Malaya, from the bat Honycteris spelaea.

ORNITHODOROS SALAHI Hoogstraal. (Text-fig. 18.)
Larva elongate. Hypostome length 0:12 mm., dentition 2/2 with 8 teeth in outer
file and 4—5 in inner file. Tarsus I 0:23 mm. long to base of claw. Haller’s organ has

CAL

(

(
CA

ea CAG
od a4 ed

ie)

Text-figs. 10-18.
10-13.—O. solomonis larva. 10, 11, Haller’s organ, dorsal; 12, hypostome; 13, palp.
14-17.—O. batuensis larva. 14, Haller’s organ, dorsal; 15, capsule, dorsal; 16, hypostome;
17, palp.
18.—O. salahi, larva capsule, dorsal.

a Slightly transverse subcircular reticulated capsule. Posterior marginal body setae
are present and strong, and the dorsal and ventral body chaetotaxy corresponds to
that of solomonis and batuensis. There are 16 setae on tarsus I as in batwensis. There
are 3 setae in the posterior hair tuft. The proportions of the palpal joints are similar
to those of batuensis.

ORNITHODOROS STEINI (Schulze).
Only the larva, described from fruit bats from Timor, is known. The shape was
given as elongate and anteriorly truncate. The dentition is 2/2 with 7 teeth in the
outer file. Schulze’s (1941, fig. 36a) figure shows strong posterior marginal setae and

F

308 BAT-INFESTING ORNITHODOROS OF THE ORIENTAL-AUSTRALIAN REGION.

relatively short palpal joints. The capsule of Haller’s organ is described as subcircular
and reticulate. It is stated that there are five setae in the posterior hair tuft. If
this includes what I have called the paramedian capsulars (PC) then it is similar
to salahi and batuensis and differs from solomonis. If there are five setae present in
the tuft the species is distinct from the other three.

Schulze considered that larvae from Nycteris javanica in West Java were identical
with steini in that they had a similarly reticulated capsule but they could well be
another species.

The species is transferred to Ornithodores because the reticulated capsule of.
Haller’s organ is Known in several species of that genus but is not knewn in Argas.

The type is reputed to be in the Berlin Museum but I have been unable to see it or
to have comparisons made.

Acknowledgements.
I am much indebted to the following: Dr. H. Hoogstraal, of Cairo, for larvae of
O. salahi and A. vespertilionis and adults of O. batuensis; Dr. J. L. Harrison, of Kuala
Lumpur, for larvae of O. batuensis; Dr. F. H. S. Roberts, of Brisbane, for larvae of
A. persicus and adults of O. gurneyi; and Dr. G. M. Kohls, of the Rocky Mountain
Laboratory, for photostats of literature. |

References.

CooLny, R. A., and Kouus, G. M., 1944.—The Argasidae of North America, Central America
and Cuba. Amer. Midland Naturalist, Monograph No. 1, Univ. Press, Notre Dame,
Indiana, 152 pp.

HooGstrRaAu, H., 1953.—Ornithodoros salahi, sp. nov. (Ixodoidea, Argasidae) from Cairo
Citadel, with notes on O. piriformis WET SETS sre 1918 and O. batuensis Hirst 1929. Jour.
Parasitology, 39, 3: 1-8.

ScHuuzE, P., 1935.—Zur Vergleichenden Anatomie der Zecken. Zeits. Morph. Okol. Tiere, 30,
1: 1-40.

, 1941.—Das Geruchsorgan der Zecken. l.c., 87: 491-564.

309

NOTES ON AUSTRALIAN THYNNINAE.

II. THe GeneRA DIMORPHOTHYNNUS, RHAGIGASTER AND ETIRONE.
By B. B. Given, Entomology Division, D.S.I.R., Nelson, New Zealand.
(Communicated by Dr. A. J. Nicholson.)

(One hundred and twenty-six Text-figures.)

[Read 26th November, 1958.]

Synopsis.
Reference is made to the instability of the usually accepted tribal categories (subfamilies
of Turner, 1907, 1910, etce.). Selected species, including five new ones (Rhagigaster burnsi,
R. stradbrokensis, R. montanus, R. kiandrensis, and Hirone mulesi) are described and figured.

INTRODUCTION.

Since paper no. 1 of this series was published (Given, 1953), the types of Turner,
Smith and Westwood have been studied at the British Museum and Oxford University
Museum. While in some instances this has produced greater perplexity, in others
the situation has been clarified.

This series of papers cannot be considered as a revision of the group, but it may
encourage some other worker to undertake such a comprehensive task. Study has been
confined to a limited number of species, to define accurately those which can be
determined with reasonable certainty.

Acknowledgement is made to the trustees and staff of the British Museum (Natural
History) and to the staff of the Hone Department of Entomology, Oxford University,
for permission to study material in their collections and to publish results of these
studies. The writer is also indebted to the Director and the Senior Entomologist of
the National Museum of Victoria, Melbourne, for the loan of material of an undescribed

‘species for description. He is particularly indebted to Dr. P. B. Carne for his careful
eriticism of the draft of this paper.

REMARKS ON TRIBAL CATEGORIES.

It was hoped to begin this paper with a key to the tribes of the subfamily.
However, attempts to construct such a Key have failed. Although this could indicate
that the present tribal classification is unnatural, attempts to erect a new and more
satisfactory series of groupings have so far been unsuccessful. For females, separation
into three tribes (on Australian material) can be made as follows:

Females — Key to Tribes.

*1. QOcelli present, eyes more than half length of head capsule ..................... Diamini.
Ocelli absent, eyes much less than half length of head capsule ..................... Qe
2. Mesopleurae showing a distinct dorsal surface ...................--2e:- Rhagigasterini.
Mesopleurae not showing a distinct dorsal surface .....................200005 Thynnini.

* Ocelli are also present in the South American species, Anodontyra tricolor Westwood.

For males, no such clear-cut distinction can be made, and even the characters
listed by Turner for separation of the Diamini from the Thynnini (including the
Rhagigasterini) are not always very clear. The following tabulation of differences
is considered to be the best available:

Males — Key to Tribes.
First transverse cubital nervure broken, with no indication of a spur vein dividing first cubital
Colle Man diblesmtridemtaten srvcu- recta icicle) stem ees sweu sh eps tetten stere dec sneterete au sie er ebajieds: Genes Diamini.
First transverse cubital nervure entire, with a spur vein indicated or dividing first cubital
cells “Mandibles: not tridentate) Sak. se clenetew nis oo cals ere ee ay etsceeere mil ant da ere abanedatel Thynnini.
(including Rhagigasterini.)
It is hoped that after the completion of this series of papers the position will be

clarified, but until then the writer must consider two tribes only to be tenable, namely

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FE

310 NOTES ON AUSTRALIAN THYNNINAE. II,

the Diamini and the Thynnini. This compromise is a reversion from the opinions of
Turner in 1910 (p. 8) to that in 1907 (p. 210). However. even in his key of 1907,
Turner was incorrect in stating that in the Diamini the males are smaller than the
females. Such is usually, but not always, the case.

It must also be pointed out that Turner (1910) made a number of errors in his
subfamily key (corresponding to tribal designations in the present paper).

Turner stated that in the Rhagigasterini the females always have at least four
joints in the maxillary palpi. In the genus Hirone, several species have only three
joints.

In keying out the males, Turner used a number of characters which are considered
below in the light of the writer’s observations.

Diamini (comprising the single species, Diamma bicolor Westwood).

“Both recurrent nervures received by second cubital cell close together.’’—This
character is not sufficiently decisive. These nervures are actually not received very
close together, and both are received by the second cubital cell in species other than
Diamma bicolor. “Antennae short and stout.’—Many rhagigasterine males have
antennae equally short and stout. “First abdomina} segment slightly strangulated at
the apex.’”—This is not characteristic solely of Diamma. “Hypopygium not produced,
rounded.’’—This also applies to several species of the genus WHirone. “Mandibles
tridentate.’”—This character appears to be quite unique.

Rhagigasterini (as separated from Thynnini).

“Second and third cubital cells always each receiving a recurrent nervure.’”—This
is not confined to the tribe. ‘“Manaibles always bidentate.”—This also applies to most
Thynnini. “Hypopygium either not at all or very slightly produced and rounded at
the apex, unarmed; or else ending in a long, acute, strongly recurved apical spine.’—
The first type of hypopygium (not at all or very slightly produced and rounded at
apex) associates this grouping with Diamma. The second type (ending in a long, acute,
strongly recurved apical spine) is also found to a less extent in such genera of the
Thynnini as Thynnoides, Epactiothynnus and Zaspilothynnus. “Claspers with an apical
tuft of long hairs turned inwards.’—This character, like the last, appears to be merely
more prominent in the Rhagigasterini than in the Thynnini, and cannot he considered
as a key character.

Most characters which appear to separate natural tribal groups fail when all
species are considered. For example, the presence of ocelli in the female of Diamma
bicolor has always been regarded as unique, but in the South American species
Anodontyra tricolor Westwood ocelli are also present, and in other species their
positions are clearly represented by rudimentary structures. The reduction of female
mouthparts cuts across the tribal grouping of Turner (1910). It is fairly frequent
in the genus Hirone, rare in the South American Thynnini, but very common in the
Australian Thynnini. The ventral modification of the head of males of the Australian
Thynnini (Given, 1954) is completely absent in the South American fauna, and in
Thynnus and Megalothynnus in Australia.

Wing venation appears to be rather unreliable in the group, many long series of
males showing a surprising degree of variation in details.

Male characters in the clypeal and terminal abdominal areas have been tried, but
none can be satisfactorily employed as tribal characters. In females, characters of
the second and apical abdominal segments are similarly unsatisfactory. The answer to
the problem may lie in the study of genitalia, although inspection of these structures
from some fifteen selected genera does not show any great promise. There is doubtless
an answer to this problem, but it is not yet obvious.

Genus DIMORPHOTHYNNUS Turner, 1910.
Gen. Ins., 105, p. 5.—EHnteles Westwood, 1844, Arcan. Ent., 2, p. 143.—Rhagigaster
(part) Guérin, 1838, Voy. Coquille, Zool. 2, ii, p. 2138.
Genotype: Dimorphothynnus haemorrhoidalis (Guérin).
Characters of the genus.—In the male there appears to be little of reliable nature
to separate it from the genus Rhagigaster. The shape of the epipygium as used by

BY B. B. GIVEN. 311

Turner (1910, p. 5) in his key would be better stated “broadly truncate- posteriorly”
for Dimorphothynnus. Other characters appear to be even less useful. In the female
the transverse striation of the second abdominal segment appears to be general and
at the present state of our knowledge must be considered as the only reliable generic
character (Text-fig. A, 18-20) for separation from Rhagigaster and Hirone, but it does
not separate the genus from the Thynnini.

DIMORPHOTHYNNUS HAEMORRHOIDALIS (Guérin), 1842, and allied species.

The type of this species, which is in the Genoa Museum, is damaged (Guiglia,
1948). Turner evidently did not see it, and his determination of the species was
probably incorrect since he considered apicalis Smith to be a synonym (Turner, 1907).
He later altered his opinions somewhat, but did not improve the situation. His
changes of opinion in synonymy were as follows:

1907: EHnteles haemorrhoidalis Guérin, 1842.
apicalis Smith, 1859; bicolor Westwood, 1844; jfimbriatus Smith, 1859; ottonis
Dalla Torre, 1897; zingerlei Dalla ‘Torre, 1897; lecheri Dalla Torre, 1897.
1916: Dimorphothynnus bicolor Westwood, 1844.
? haemorrhoidalis Guérin, 1842; zingerlei ‘Dalla Torre, 1897; ? lecheri Dalla
Torre, 1897; haemorrhoidalis Turner, 1907.
Dimorphothynnus fimbriatus Smith, 1859.
apicalis Smith, 1859; sttonis Dalla Torre, 1897.

The 1916 synonymy involved the reduction of a genotype to synonymic status
beneath a later-described species, when the type of at least one of the two had not
been examined. Until the type of haemorrhoidalis Guérin is carefully compared with
that of apicalis Smith, which are both males, and the types of bicolor Westwood and
jimbriatus Smith (females) are compared and opposite sexes of the four types reliably
associated and checked, the confusion will persist. It is of interest to note that with
the type male of apicalis in the British Museum collection is a female which car
probably be considered as an allotype. The position then is that apicalis can be
compared with all other types of the complex.

From personal examination of types the following appears to be the situation
(see Text-fig. A):

1. D. haemorrhoidalis (Guérin), 1842. Type ¢ in Genoa Museum.

(Probably) D. bicolor (Westwood), 1844. Type 9 in Oxford University Museum.
2. D. fimbriatus (Smith), 1859. Type 2 in British Museum.
3. D. apicalis (Smith), 1859. Type @¢@ in British Museum.

In the general collections in the British Museum a series of Western Australian
specimens labelled by Turner as D. fimbriatus was examined. This series may be
divided into groups as follows: .

Males—
(a) Mandibles expanded. prosternum concave. (Text-fig. A, 4, 2.)
(ob) Mandibles normal, procoxae concave. (Text-fig. A, 5, 3.)
Females—
(a) Clypeus normal. (Text-fig. A, 15.)
(bo) Clypeus carinate (fimbriatus). (Text-fig. A, 16.)

An analysis of mounted pairs (taken in copuia) is as follows:
No. of pairs: 6.

Male Female
Type Type
(a) (a)
(a) (b) On account of frequency of occurrence, probable
(a) (0) correct pairing is
(a) (b) Male (a) with female (0)
(0) (a) Male (0) with female (qa)

(0) (a)

312 NOTES ON AUSTRALIAN THYNNINAE. II,

An analysis of specimens not indicated as having been in copula but considered in
accordance with locality is as follows:

Males Females Locality
3 (a) 3 (a) Wanneroo
4 (bd) 4 (b)

1 (a) 10 (0b) Yallingup
1 (0)

Text-fig. A. Dimorphothynnus.

1: D. apicalis (Smith) male, prosternum and procoxae. 2: D. fimbriatus (Smith) male,
prosternum and procoxae. 3: D. haemorrhoidalis (Guérin)? male, prosternum and procoxae.
4: D. fimbriatus (Smith) male, mandibles. 5: D. haemorrhoidalis (Guérin) ? male, mandibles.
*6: D. apicalis (Smith) male, head. 7: D. fimbriatus (Smith) male, clypeus. 8: D. haemor-
rhoidalis (Guérin)? male, eclypeus. *9: D. apicalis (Smith) male, epipygium. 10: D. fimbriatus
(Smith) male, epipygium. 11: D. haemorrhoidalis (Guérin)? male, epipygium. *12, 12a: D.
apicalis (Smith) male, hypopygium, lateral, ventral. 13, 13a: D. fimbriatus (Smith) male,
hypopygium, lateral, ventral. 14, 14@: D. haemorrhoidalis (Guérin)? male, hypopygium, lateral,
ventral. *15: D. apicalis (Smith) female, head. *16: D. fimbriatus (Smith) female, head.
17: D. haemorrhoidalis (Guérin)? female, head. *18: D. apicalis (Smith) female, abdominal
tergites 1 and 2. *19: D. fimbriatus (Smith) female, abdominal tergites 1 and 2. 20: D.
haemorrhoidalis (Guérin)? female, abdominal tergites 1 and 2. *21: D. apicalis (Smith)
female, pygidium. #22: D. fimbriatus (Smith) female, pygidium. 23: D. haemorrhoidalis
(Guérin) ? female, pygidium.

* Figures drawn from type specimens.

BY B. B. GIVEN. 313

The above analysis illustrates the danger of drawing conclusions from unpaired
material, as a casual examination of the second group (unpaired material) would
suggest that at Wanneroo pairing is in the reverse order to that of pairs taken in
copula.

It is both unfortunate and extraordinary that Turner did not note the mandibular,
coxal and prosternal male differences and the propodeal and elypeal differences in
females of Western Australian material, as these are excellent characters for separation.

Text-figure A, figures 2, 4, 7, 10 and 13 therefore probably represent D. fimbriatus 2,
while figures 3, 5, 8, 11 and 14 may represent D. haemorrhoidalis 8. (Of some interest
is the fact that mandibular variation very similar to that illustrated in figures 4 and 5
occurs in males of Rhagigaster mandibularis Westwood (Text-fig. C, 12) and R. wnicolor
Guérin. Turner failed to note the character in either genus.)

While apicalis (Smith) is a South Australian species, it is probable that haemor-
rhoidalis (Guérin) and fimbriatus (Smith) are confined to Western Australia. The
concave coxal and prosternal areas and expanded mandibles on the males, and the
curious clypeal carina of fimbriatus females, appear to occur only on Western Australian
species.

Other species of the genus are not considered in this paper as they have not
entered into the work of the writer. However, a note must be included concerning the
types of D. dimidiatus Smith. Both Oxford University and the British Museum have
specimens marked as types, and both are identical as regards structure. The British
Museum male is as much longer than Smith’s record for the type as the Oxford Museum
specimen is shorter.

Genus RHAGIGASTER Guérin, 1838.
Voy. Coquille, Zool. 2, ii: 213—Diamma (Part) Guérin, 1838, Voy. Coquille, Zool. 2,
ii: 235—Rhytidogaster Turner, 1907, Proc. Linn. Soc. N.S.W., 32: 229.
Genotype: Rhagigaster wnicolor Guérin.

Characters of the genus.

Male: General form elongate, the abdomen somewhat constricted between segments.
Head not ventrally modified, mouth parts normal, mandibles bidentate. Antennae much
shorter than head and thorax together. Hypopygium produced into a long recurved
apical spine.

Female: Head relatively large, broader than thorax. Mandibles usually bidentate,
sometimes simple. Mouth parts not usually greatly reduced. Thorax showing a
distinct dorsal mesopleural surface. Pygidium never markedly elaborated, not truncate
apically. Second dorsal and fifth ventral abdominal segments without characteristic
sculpture. Mesosternal intercoxal processes present.

RHAGIGASTER BURNSI, nN. sp. (Text-fig. B.)

Male: Colour black, the legs reddish. Head distinguished by the presence of an
irregularly arcuate, transverse, rugose carina above the frons (Text-fig. B, 3). Pro-
notum more strongly produced at the anterior angles than in any other species of the
genus (Text-fig. B, 1; cf. Fig. 4). First abdominal segment ventrally tuberculate,
approaching the subtuberculate state dorsally (Text-fig. B, 1 and 6; cf. Text-fig. B,
5 and 7). Terminal abdominal segment as illustrated in Text-fig. B, 1 and 8; cf. Fig. 9).
Length (excluding antennae) 19-22:5 mm.

Female: Colour black, the legs and antennae deep reddish, head with a brownish-
red transverse area between eyes behind antennae. Sculpture highly distinctive (Text-
fig. B, 2). Longitudinal post-orbital grooves absent from head. Length (excluding
antennae) 14-17-5 mm.

Relationships—Most closely allied to R. corrugatus Turner, but differs in the male
in the presence of an arcuate suprafrontal carina (Text-fig. B, 3), smoother clypeus,
strongly produced anterior pronotal angles (Text-fig. B, 1 and 4), more acute dorsal
prominence on first abdominal segment (Text-fig. B,; 1 and 5), ventral abdominal
tubercle less slender, intersegmental incision between abdominal segments one and two
shallower and more obtuse (Text-fig. B, 6 and 7) and differences in form and punctation

314 NOTES ON AUSTRALIAN THYNNINAE. II,

of terminal abdominal segment (Text-fig. B, 8, 9). The reddish colour of the legs is
also distinctive. Both burnsi and corrugatus may be separated from all other species
of the genus by the broadly rounded apex te the epipygium. In this they approach
the genus Dimorphothynnus in which this structure is broadly truncate apically.

A) Yas 3:

ne

91 eaUNEN

43 Py . ws i
0949)

Text-fig. B. Rhagigaster.

1-3: R. burnsi, n. sp. *1, male; *2, female; *3, male head, anterior. 4, 5: R. corrugatus
Turner. 4, male, pronotum; 5, male, abdominal segment 1, dorsal. *6: R. burnsi, n. sp., male,
abdominal segments 1 and 2, lateral. 7: R. corrugatus Turner, male, abdominal segments 1
and 2, lateral. *8: R. burnsi, n. sp., male, abdominal apex, lateral. 9: R. corrugatus Turner,
male, abdominal apex, lateral.

* Figures drawn from type specimens.

The female of R. burnsi can be confused with no species other than R&. corrugatus,
from which it differs in having even coarser head and thoracic puncturing (Text-
fig. B, 2), with more sparse puncturing on the abdomen, and in the colour of the head.
Whereas in burnsi the only light head coloration is a band between the eyes, in
corrugatus the entire dorsal area behind the antennae, the antennae themselves, and
the mandibles are light coloured.

BY B. B. GIVEN. 315

Locality, etc—All material of this species examined was collected by Messrs. Burns
and Pescott at Tubrabucca, New Seuth Wales, January 10-23, 1948.

Types.—Holotype 4, allotype ?, 19 paratype ¢ and 7 paratype 9 in the collection of
the National Museum of Victoria, Melbourne. Four paratype g, 3 paratype ? in the
collection of Mr. A. N. Burns, Melbourne (after whom the species is named). A
paratype pair in the collection of the Division of Entomology, C.S.I.R.O., Canberra,
A.C.T.

Acknowledgement.—For permission to examine and describe the above material
acknowledgement is made to Mr. A. N. Burns, Curator of Insects, and Mr. R. T. M.
Pescott, Director, The National Museum of Victoria.

RHAGIGASTER STRADBROKENSIS, D. Sp. (Text-fig. C.)

Male: Colour black, wings fusco-violaceous. Head (Text-fig. C, 1) with transverse
frontal carina above a sparsely rugose-punctate area. Pronotum (Text-fig. C, 2) not
strongly produced at anterior angles. First abdominal segment (Text-fig. C, 5) obtusely
tuberculate ventrally. Abdominal terminalia (Text-fig. C, 3, 4) as illustrated (note
widely spaced prominent lateral spines on hypopygium). Length (excluding antennae)
19 mm.

Female: Colour black with lateral testaceous areas between antennae and eyes.
Tips of mandibles reddish. Head (Text-fig. C, 13) shining, finely sparsely punctate
except on frontal area. Mandibles (Text-fig. C, 14) simple. Thorax (Text-fig. C, 13)
shining, slender, sparsely punctate. Abdomen smooth and shining, sparsely punctate.
First segment ventrally toothed (Text-fig. C, 16). Pygidium (Text-fig. C, 15) relatively
simple, shining. Length (excluding antennae) 11 mm.

Relationships—Prokably most closely ailied to R. unicolor Guérin.

Locality, etc—The only pair examined was collected by H. Hacker at Stradbroke
Island, 17th September, 1915.

Types.—Holotype 4, allotype @ in the collection of the Cawthron Institute, Nelson,
New Zealand.

RHAGIGASTER UNICOLOR LYELLI Turner, 1910. (Text-fig. C.)

Proc. zool. Soc. Lond., p. 260.

Male: Colour black, wings clear. Head (Text-fig. C, 6) with frontal area below
transverse carina closely, coarsely punctate. Pronotum (Text-fig. C, 7) with posterior
margin strongly curved, anterior angles not strongly produced. First abdominal
segment variably tuberculate ventrally. In Text-fig. C, 10, this condition is represented
in its maximum development for the species. Abdominal terminalia as in Text-fig. C,
8 and 9 (note depth of aciculus in profile (Fig. C, 9) and presence of lateral spines).
Length (excluding antennae) 14-19 mm.

Female: Black or piceous, meso- and metathorax and propodeum red, lateral areas
between antennae and eyes ferruginous. Head (Text-fig. C, 17) shining, sparsely
punctate except at bases of antennal prominences. Mandibles as in Text-fig. C, 18.
Thorax smooth and shining, as Text-fig. C, 17. Terminal area of abdomen smooth,
sparsely punctate, not highly distinctive. Length (excluding antennae) 10-13 mm.

Locality, etc.—Material examined (23 males, 21 females) was collected at Kiandra,
Jindabyne, Kosciusko, Cooma, Goulburn in N.S.W., and Dartmoor, Melton, Porpunkah,
Daylesford in Victoria. All were collected by the writer except a pair from Porpunkah
collected by F. E. Wilson and a pair from Goulburn collected by EH. F. Riek.

Types—The male originally described by Turner is in the British Museum. The
female has not previously been described, and the specimen from which the illustrations
in this paper were made is in the collection of the Division of Entomology, C.S.I.R.O.,
Canberra.

Relationships—Turner (1910a, 191060) divided R. wnicolor into three subspecies
and in 19100 he lists distinctions which appear to be sufficiently sound. In 1910a@ he
mentions proportions of the second and third cubital cells. This character, however,
is somewhat unreliable on account of its variability.

316 NOTES ON AUSTRALIAN THYNNINAE. II,

Turner was very much in error in confusing R. wnicolor with R. mandibularis
Westw., the material which he in 1907 ascribed to this latter species being obviously
misidentified.

Although the subspecies of R. wnicolor are related to stradbrokensis and
mandibularis the differences are considerable, as illustrated in Text-figure C.

Note.—Females of this species stridulate when not in copula.

Text-fig. C. Rhagigaster.

*1-5: R. stradbrokensis, n. sp. *1, male, head; *2, male, pronotum; *3, male, epipygium
and hypopygium, dorsal; *4, male, epipygium and hypopygium, lateral; *5, male, abdominal
segment 1. 6-10: R. wnicolor lyelli Turner. 6, male, head; 7, male, pronotum; 8, male,
epipygium and hypopygium, dorsal; 9, male, epipygium and hypopygium, lateral; 10, male,
abdominal segment 1. 11, 12: R. mandibularis Westwood. 11, male epipygium, dorsal;
12, male, left mandible, lateral. 13-16: R. stradbrokensis, n. sp. *13, female, head and
thorax; *14, female, mandibles and clypeus; *15, female, pygidium; *16, female, abdominal
segment 1. 17, 18: R. unicolor lyelli Turner. 17, female, head and thorax; 18, female, right
mandible. 19, 20: R. mandibularis Westwood. 19, female, head; 20, female, right mandible.

* Figures drawn from type specimens.

RHAGIGASTER MANDIBULARIS Westwood, 1844. (Text-fig. C.)

Arcan. Ent., 2: 105.

Male: Colour black, wings clear. Head similar to that of R. wnicolor lyelli (Text-
fig. C, 6) but with a distinctive process on the dorsal edge of the mandibles (Text-fig.
C, 12). Thorax and abdomen similar to that of R. unicolor lyelli except for terminalia.
Epipygium (Text-fig. C, 11) laterally produced, hypopygium without lateral spines.
Length (excluding antennae) 15-19 mm.

Female: Colour black or piceous, meso- and metathorax and propodeum and legs
red. No lighter markings between antennal bases and eyes. Head (Text-fig. C, 19)

BY B. B. GIVEN. 317

rectangular with posterior angles somewhat excavated. Mandibles as in Text-fig. C, 20.
Thorax and abdomen as in R. uwnicolor lyelli. Length (excluding antennae) 9-13 mm.

Relationships——Most closely allied to R. wnicolor, but readily separated in the
male by hypopygial and mandibular characters, and in the female by the differences in
the rear angles of the head, colour of legs, absence of colour patches between antennal
bases and eyes and shape of the mandibles.

Wing venation of males (variation).—In Text-figure G, 7-13, aberrations in wing
venation are illustrated. It is of interest to note that out of 23 males examined six
showed aberrations; Text-fig. G, 14 and 15 illustrate the range of variation shown in 23
males in the relative coincidence of junction of the second transverse cubital and
second recurrent nervures.

Locality, etc.—The type pair was collected at Port Phillip. Material in the writer’s
collection was collected at the following localities: Cavendish, Victoria Valley, Croydon,
Melton, Nigretta, Dunkeld and Hamilton, in Victoria.

Types.—The type pair is in the Oxford University Museum.

RHAGIGASTER ACULEATUS Saussure, 1868. (Text-fig. D.)

Reise Novara, Zool. 2, Hym.: 113 (¢)—Turner, 1910, Proc. zool. Soc. Lond., 1910:
264 (9).

Male: Colour black on head, thorax, legs and first abdominal segment. Remainder
of abdomen ferruginous, except for a dorsal infuscate mark on segments 2 to 5 inclusive.
Head closely punctate, not highly distinctive. No well-defined triangular area on
clypeal apex. Antennae shorter than thorax. Thorax closely punctate, pronotum
somewhat obtusely produced at anterior angles. Abdomen rather coarsely punctate,
second segment as in Text-fig. D, 3. Epipygium (Text-fig. D, 4) coarsely punctured.
Length (excluding antennae) 9-11 mm.

Female: Colour uniform light yellowish-brown, sometinies with the head slightly
darker infuscate. Head (Text-fig. D, 1) rather elongate, rectangular, puncturing
moderately dense anterodorsally except for a median longitudinal space. Thorax
(Text-fig. D, 1) rather sparsely and finely punctate, propodeum much broader
posteriorly than anteriorly. Abdomen finely punctate, smooth, anterior angles of
segment 1 acute (Text-fig¢. D, 1). Pygidium as in Text-fig. D, 2. Length (excluding
antennae) 5-8-5 mm.

Locality, etec——The four pairs examined were collected by the writer at Cavendish,
Victoria. The species also occurs in New South Wales.

Type.—The type male has not been seen, and identification was based on the
original description and on material determined by Turner, in the British Museum.

Relationships—This species, and those which follow, are in the group which
Turner placed (1907, p. 211) in the genus Rhylidogaster. Most of the smaller members
of this group are closely allied, but aculeatws may be separated from them on colour
in the male, and on the acute angles of the first abdominal segment in the female.

RHAGIGASTER COMPARATUS Smith, 1859. (Text-fig. D.)

Cat. Hym. B.M. 7: 69.

Male: Colour black, abdominal segment 6 and terminalia reddish, tibiae and tarsi
brown. Head (Text-fig. D, 9) closely, rather coarsely and rugosely punctate. A small
triangular clypeal area present. Thorax closely and coarsely punctate, apical angles
not markedly produced. Abdomen more shallowly punctate than thorax. Epipygium
and hypopygium as illustrated (Text-fig. D, 7, 8). Length (excluding antennae)
9-5-10 mm.

Female: Colour black, abdominal segment 5 and terminalia red, legs ferruginous.
Head (Text-fig. D, 5) approximately square, uniformly punctate, with minute inter-
mediate punctures and striations, particularly anteriorly. Posterior angles broadly
rounded. Thorax uniformly coarsely punctate, the punctures elongate. Abdomen with
angles of first segment not acute; coarsely, longitudinally, rugosely punctate. Pygidium
as in Text-fig. D, 6. Length (excluding antennae) 6-5—7-5 mm.

318 NOTES ON AUSTRALIAN THYNNINAE. Il,

Locality, etc—The 16 pairs examined were collected by the writer at Wannon,
Cavendish and Woori Yallock, Victoria. Turner states the locality as being Victoria

and South Australia.
Types.—The type pair appear to be lost, but should be in the British Museum.

Remarks.—This species is commonly found feeding at Hakea blossom. It is most
plentiful during September and October.

pee,
ib 5K

Ee a

\

Text-fig. D. Rhagigaster.

1-4: R. aculeatus Saussure. 1, female; 2, female, pygidium; 3, male, abdominal segment 2;
4, male, epipygium and hypopygium. 5-9: R. comparatus Smith. 5, female; 6, female,
pygidium; 7, male, epipygium and hypopygium, dorsal; 8, male, epipygium and hypopygium,
lateral; 9, male, frontal detail of head. 10-14: R. iracundus (Turner). 10, female; 11, female,
pygidium; 12, male, head; 13, male, epipygium and hypopygium, dorsal; 14, male, epipygium
and hypopygium, lateral. 15: R. tumidus (Turner), male, epipygium and hypopygium.
16-22: R. montanus, n. sp. *16, female; *17, female, pygidium; *18, male, head; *19, male,
abdominal segment 2; *20, male, pronotum; *21, male, epipygium and hypopygium, dorsal;
+22, male, epipygium and hypopygium, lateral.

* Figures drawn from type specimens.

RHAGIGASTER IRACUNDUS (Turner), 1907. (Text-fig. D.)
Proc. Linn. Soc. N.S.W., 32: 237 (Rhytidogaster).—1910. Gen. Ins., 105: 7
(Rhagigaster).
Male: Colour black, abdominal segment 6 and terminalia dark red. Wings clear.
Head (Text-fig. D, 12) very coarsely punctate, with a smooth, depressed, cordate

BY B. B. GIVEN. 319

elypeal area with strongly elevated margins. Thorax closely, rugosely punctured.
Anterior pronotal angles moderately produced. Abdomen moderately coarsely punctate.
Epipygium and hypopygium as illustrated in Text-fig. D, 15 and 14. In some specimens
the aciculus is more uniformly curved than as shown. Length (excluding antennae)
12-14 mm.

Female: Colour ferruginous to piceous. Head almost square (Text-fig. D, 10),
uniformly punctate. Thorax (Text-fig. D, 10) uniformly punctate. Pronotum almost
square, depressed internal to posterior angles. Propodeum angled posteriorly. First
abdominal segment strongly angled anteriorly (Text-fig. D, 10). Abdomen uniformly
punctate. Pygidium as in Text-fig. D, 11. Length (exclusive of antennae) 8-9 mm.

Locality, etc—The type is from Melbourne. The four pairs examined by the
writer are from Hartley, South Australia (H. F. Lower), Canberra, A.C.T. (P. B.
Carne), and Nelson, Victoria.

Types.—The type male is in the British Museum. The female has not previously
been described and the specimen on which the above description is primarily based
is in the collection of the Division of Entomology, C.S.I.R.O., Canberra.

Remarks.—The Nelson pair is atypical. The male has abdominal segment 5 red,
and the female is not as angular as the genotype on the head or thorax. The male
from Canberra is similar to the Nelson specimen, and these may represent a different
but very closely allied species.

RHAGIGASTER TUMIDUS (Turner), 1907. (Text-fig. D.)

Proc. Linn. Soc. N.S.W., 32: 2386 (Rhytidogaster) —1910. Gen. Ins., 105: 8
(Rhagigaster).

This species superficially resembles R. iracundus but is readily distinguished as
follows.

Male: Head without cordate smooth clypeal area. Epipygium and hypopygium as
in Text-fig. D, 15 (compare Fig. D, 14).

Female: Colour dominantly black, head longer than broad, less densely punctured,
not so angular.

Locality, etc—The specimen (¢) in the writer’s collection is from Kiandra, N.S.W.
Turner records the species from Melbourne, Victoria; Swan River, W.A.; and Tempe,
N.S.W.

Types—In the British Museum.

RHAGIGASTER MONTANUS, Nn. sp. (Text-fig. D.)

This species is allied to R. tumidus.

Male: Colour black, wings clear. Head (Text-fig. D, 18) closely punctured, without
smooth clypeal area. Thorax closely punctured, moderately produced at anterior
angles (Text-fig. D, 20). Abdomen coarsely and rather densely punctured (second
segment, Text-fig. D, 19). Hypopygium and epipygium as in Text-fig. D, 21 and 22.
Length (excluding antennae) 11-12 mm.

Female: Colour black on head and abdomen. Thorax, legs, antennae and pygidium
red. Head (Text-fig. D, 16) oblong, moderately deeply punctate, aciculate between
punctures. Thorax (Text-fig. D, 16) sparsely irregularly punctate. Abdomen with
regular, elongate punctures, truncate anteriorly but without acute anterior angles.
Pygidium as in Text-fig. D, 17. Length (excluding antennae) 5:5-6:5 mm.

Locality, etc—The holotype and paratype pairs were collected by the writer at
Kiandra, N.S.W., at 4,500 ft., 9th February, 1952.

Types.——Holotype and allotype (pair taken in copula) in the collection of the
Division of Entomology, C.S.I.R.O., Canberra, and a paratype pair in the collection of
the Entomology Division, D.S.I.R., Nelson, New Zealand.

Relationships—Allied to the tumidus, aculeatus, iracundus group. The male is
separated from iracundus on account of its lack of smooth clypeal area, and from
tumidus, etc., by colour and terminal characters. The female is highly characteristic
on account of colour and punctation, as well as the form of the pygidium.

320 NOTES ON AUSTRALIAN THYNNINAE. II,

RHAGIGASTER KIANDRENSIS, n. sp. (Text-fig. HE.)

Male: Colour black, wings slightly infuscate. Head (Text-fig. EH, 3) rugosely
punctate, the frontal and supra-antennal carinae well developed, the apex of the former
subtuberculate. Pronotum (Text-fig. H, 4) broad and strongly but not acutely produced
at anterior angles. Abdomen shallowly and irregularly punctate (Text-fig. H, 5,
abdominal segment 2). EHEpipygium and hypopygium as in Text-fig. HE, 6 and 7. Length
(exclusive of antennae) 12-16 mm.

a
A
vey

Text-fig. H. Rhagigaster and Hirone.

*1-7: R. kiandrensis, n. sp. *1, female; *2, female, pygidium; *3, male, head; *4, male,
pronotum; *5, male, abdominal segment 2; *6, male, epipygium and hypopygium, lateral;
*7, male, epipygium and hypopygium, dorsal. 8, 9: R. pugionatws Saussure. 8, male, epipygium
and hypopygium, lateral; 9, male, epipygium and hypopygium, dorsal. 10-16: Hirone rufopicta
(Smith). 10, male; 11, female; 12, male, abdominal terminalia; 13, female, abdominal
terminalia; 14, female, pygidium; 15, female, head; 16, female, left mandible.

* Wigures drawn from type specimens.

Female: Colour black, metathorax and legs dark ferruginous. Head (Text-fig. H, 1)
almost square, posterior angles rounded, coarsely longitudinally rugosely punctate.
Minute punctures and lines between main punctures. Thorax (Text-fig. E, 1) unevenly
punctured, punctures coarse on pronotum, which is laterally somewhat depressed.
Abdomen more sparsely and finely punctate, anteriorly truncate, but anterior angles
not produced. Pygidium as in Text-fig. EH, 2. Length (excluding antennae) 8-11 mm.

BY B. B. GIVEN. 321

Locality, etc—The 16 type pairs were collected by the writer at 4,500 ft. at
Kiandra, N.S.W., in February, 1952, feeding at Leptospermum blossom.

Types—The holotype and allotype (taken in copula) and four paratype pairs are
in the collection of the Division of Entomology, C©.S.1.R.O., Canberra, and 11 paratype
pairs are in the collection of the Entomology Division, D.S.I.R., Nelson, New Zealand.

Relationships.—This species is allied to the tumidus, iracundus group, but is
larger and in some ways intermediate between them and corrugatus and burnsi:

RHAGIGASTER PUGIONATUS Saussure, 1868.

Reise Novara, Zool. 2, Hym.: 1138.

The determination of the two males studied is based on Turner’s interpretation
of Saussure’s description, having been named by comparison with Turner’s material
at the British Museum.

Male: Colour black, mesopleurae red, wings slightly violaceous infuscate. Head
closely rugosely punctate, no clypeal area or tubercle. ‘Thorax as in other members
of the twmidus, iracundus group. Abdomen moderately closely punctate. Hypopygium
and epipygium as in Text-fig. H, 8 and 9. Length (excluding antennae) 11:5 mm.

Locality, etc.—The two males examined were collected by EH. F. Riek at Orford
and Cole’s Bay, Tasmania, on 9th and 13th January, 1948. Turner also records this
species from New South Wales (Sydney).

Genus HIiRONE Westwood, 1844.

Arcan. Ent., 2: 44.Aelurus Turner, 1907 (not Klug) (part), Proc. LINN. Soc.
N.S.W., 32: 247—Aelurus (Lepteirone) Turner, 1907, Proc. Linn. Soc. N.S.W., 32:
299 Aelurus (Hirone) Turner, 1907, Proc. Linn. Soc. N.S.W., 32: 258.

Genotype: Hirone dispar Westwood.

Characters of the Genus.—The male may be separated from other members of the
rhagigasterine genera by the unspecialized hypopygium, which is apically broadly
rounded and rarely projects beyond the epipygium. However, this does not separate it
from all other genera of the Thynnini, as certain species of Zeleboria, Neozeleboria
and Phymatothynnus are very similar in this respect. From these genera, Hirone
males may be readily distinguished by the unspecialized anteroventral surface of the
head.

In the female the contiguous nature of the mesocoxae appears to be a distinguishing
character not only from other rhagigasterine genera, but also from the genus Ariphron,
to some species of which Hirone females are in other respects similar. The lack of
any conspicuous adornment, such as carinae or rugae on the second abdominal segment,
is a character shared with some species of Ariphron and with all species of
Rhagigaster.

Note.—The above remarks refer only to Australian genera.

HXIRONE RUFOPICTA (Smith), 1879. (Text-fig. E.)

Descr. N. Sp. Hym., p. 159 (Thynnus).—Turner, 1907, Proc. Linn. Soc. N.S.W.,
32: 251, Aelurus (Lepteirone) —1910, Gen. Ins., 105: 9 (Hirone).

Male: Colour as indicated in Text-fig. EH, 10. The line-shaded areas black, stippled
areas ferruginous and unshaded areas yellow. The relative extent of the colour areas
varies somewhat between individuals. Vestiture is golden. Head with the longitudinal
Median ridge below the antennal bases apically truncate-tuberculate. Depressed areas
between eyes and lateral ocelli. Punctation fine and close. Thorax (Text-fig. E, 10)
very finely and rather sparsely punctate on pronotum, elsewhere closely punctate.
Median segment very finely transversely rugose-striate. Abdomen smooth, very finely
and sparsely punctate. Terminalia as in Text-fig. EH, 12, not highly characteristic.
Length (excluding antennae) 9-12-5 mm.

Female: Colour ferruginous, usually darker infuscate on abdomen and anterior
discal area of head. Antennae and legs usually testaceous. Head (Text-fig. H, 15)
almost square, broadly rounded at posterior angles with posterior margin medially
incurved. Punctation rather variable and irregular, integument very finely aciculate
anteriorly behind antennal bases. Mandibles (Text-fig. H, 16) strongly tridentate. Eyes

322 NOTES ON AUSTRALIAN THYNNINAE. II,

very small. Thorax (Text-fig. EH, 11) slender, particularly the propodeum. Punctation
rather coarse but not close, dorsal surfaces finely longitudinally aciculate. Abdomen
more uniformly punctate than thorax, finely longitudinally aciculate. Pygidium (Text-
fig. H, 13, 14) relatively simple and finely punctate. Length (excluding antennae)
5-5-8:5 mm.

Locality, ete—Turner gives Adelaide and Melbourne as localities. The 32 pairs
collected by the writer were ali taken at Wannon, Victoria, in September and October
from 1949 to 1951. This species, like most of the genus, appears to be a honey-dew
feeder, rarely if ever visiting blossom.

Types.—The type male is in the British Museum. The above is the first description
of the female and is principally based on a specimen in the collection of the Division
of Entomology, C.S.I.R.O., Canberra.

EIRONE MAJOR Turner, 1919. (Text-fig. F.)

Rec. S. Aust. Mus., 1: 171.

Male: The following supplements Turner’s description. Colour black, with clypeus,
antennal prominences dorsally, anterior pronotal margin, mandibles basally and legs
yellow. Antennae ferruginous, darker apically. Tegulae and wing veins ferruginous.
Mandibles apically dark ferruginous. Head almost impunctate between ocelli and
antennae; much produced posteriorly (Text-fig. F, 6). Epipygium as in Text-fig. F, 5.
Length (excluding antennae) 12 mm.

Female: Colour of head and abdomen piceous; mandibles, antennae, thorax and
legs ferruginous. Head (Text-fig. F, 4) slightly broadest postericrly, somewhat sparsely
and irregularly punctate, sparsely and finely aciculate towards antennal prominences.
Thorax (Text-fig. F, 4) with rather large elongate punctures on pronotum and
mesopleurae, fine sparse punctures elsewhere except on posterior face of propodeum,
which is densely, finely punctate. Abdomen (Text-fig. F, 4) with scattered elongate
punctures, finer punctures toward anterior margins of each tergite. (Note: These fine
punctures may be covered by telescoping of segments in some specimens.) Pygidium
as in Text-fig. F, 2. Length (excluding antennae) 9:5 mm.

Locality, etc—Turner gives the locality for the type (collected by Lea) as Forest
Reefs, between Bathurst and Orange, N.S.W. The pair and single male collected by
the writer were both taken at Croydon, Victoria, in February, 1950, while sweeping
Hucalyptus foliage at about 25 ft. above ground level using a net with an 18 ft. handle.
The trees were infested with Hriococcus coriaceous Maskell, and it seems possible
that the specimens were feeding on honey-dew.

Types.—A cotype male is in the South Australian Museum, Adelaide, and another
in the British Museum. The female has not previously been described and the specimen
on which the above description is based is in the collection of the Division of
Entomology, C.S.1I.R.O., Canberra.

Variety of HiroNE MAgorR Turner.

Two males and a pair taken in copula at Kiandra, N.S.W., on 9th, 19th and 23rd
February, 1952, respectively, differ from the typical form as follows (see Text-fig. F).

Male: Larger size (14-5 mm. compared with 12 mm.), black antennae and antennal
prominences and difference in head shape (Text-fig. F, 7 and 6).

Female: Larger size (11 mm. compared with 9:5 mm.), black propodeum, differences
in punctation pattern (Text-fig. F, 4 and 1) and different pygidial characters (Text-
fig. F, 3 and 2).

The pair on which the above remarks are based is in the collection of the Division
of Entomology, C.S.I.R.O., Canberra. Two males are in the Entomolegy Division,
D.S.1.R., Nelson, New Zealand.

EXIRONE ARENARIA (Turner), 1907. (Text-fig. F.)
Proc. Linn. Soc. N.S.W., 32: 253, Aelurus (Lepteirone)—1910, Gen. Ins., 105: 9
(Hirone).
Male: Turner gives a good general description of this species, but the following
remarks will assist identification. Turner states that the apex of the clypeus is

BY B. B. GIVEN. 323

produced into minute spines. These processes would be better described as tubercles.
Text-fig. F, 8, of the head profile, illustrates the large antennal prominences. Text-
fig. F, 9, illustrates the epipygial structure, in which the reflex margin should be
noted.

The female is unknown.

Text-fig. FE. WHirone.

1-2: H. major Turner. 1, female; 2, female, pygidium. 3-4: H. major Turner, var. 3,
female, pygidium; 4, female. 5-6: H. major Turner. 5, male, epipygium; 6, male, head,
profile. 7: H. major Turner, var., male, head, profile. 8, 9: H. arenaria (Turner). 8, male,
head, profile; 9, male, epipygium. 10, 11: EH. schizorhina Turner. 10, male, head, profile;
11, male, epipygium. 12, 13: EH. ferrugineicornis Turner. 12, male, head, profile; 13, male,
epipygium. 14, 15: H. lucida (Smith). 14, male, head, profile; 15, male, epipygium. 16-18:
HE. dispar Westwood. 16, male, head, profile; 17, male, epipygium; 18, female.

Locality, etc——-The type was collected by French in Victoria, and the two males
examined were collected by the writer at Kiandra, N.S.W., 4,500 ft., on 3rd and 23rd
February, 1952. They were taken feeding on jassid exudations on Hucalyptus foliage.
One male is deposited in the collection of the Division of Entomology, C.S.I.R.O.,
Canberra, the other in the collection of the Entomology Division, D.S.I.R., Nelson,
New Zealand.

Type—tin the British Museum.

324 NOTES ON AUSTRALIAN THYNNINAE. II,

EIRONE SCHIZORHINA Turner, 1910. (Text-fig. F.)

Proc. zool. Soc. Lond., 1910, p. 264.

The following notes will serve to supplement Turner’s description. Male: Clypeal
tubercles and marginal depression (Text-fig. F, 10) are distinctive, as also is the
epipygium (Text-fig. F, 11). The antennal prominences are joined as a transverse
ridge.

The female is unknown.

Locality, etc—The type locality is New South Wales. Material examined was
collected at Blundeli’s, A.C.T. (EH. F. Riek, 10th February, 1948), and Kiandra, N.S.W.
(the writer, 23rd February, 1952). The former specimen is in the collection of the
Division of Entomology, C.S.I.R.O., Canberra, the latter in the collection of the
Entomology Division, D.S.I.R., Nelson, New Zealand.

EXIRONE FERRUGINEICORNIS Turner, 1910. (Text-fig. F.)

Proc. zool. Soc. Lond., 1910, p. 265.

Turner’s description is good, but the illustrations (Text-fig. F, 12 and 13) and the
following notes will make determination more simple.

Male: The pronotum, not the entire prothorax, is ferruginous. Antennae inserted
in rather deeply depressed areas, antennal prominences being scarcely present (as
illustrated by profile, Text-fig. F, 12). The abdomen is unusually short for the genus,
in the specimen examined being shorter than the thorax. Epipygium (partly retracted)
as in Text-fig. F, 13.

The female is unknown.

Locality, etc—The type, which is in the British Museum, was collected at
Hermannsburg, Central Australia. The specimen examined was collected by HE. F. Riek
on 27th October, 1949, 20 miles south-east of Bourke, N.S.W., and is in the collection
of the Division of Entomology, C.S.I.R.O., Canberra, A.C.T.

EIRONE LUCIDA (Smith), 1859. (Text-fig. F.)

Cat. Hym. B.M., 7: 36. Thynnus (Agriomyia)—Turner, 1907, Proc. Linn. Soc.
N.S.W., 32: 266. Aelurus (Hirone).—1910, Gen. Ins., 105: 9 (Hirone).

Male: Black, legs testaceous or light ferruginous except for coxae and trochanters,
which are black, mandibles testaceous with ferruginous apices, small yellow spot on
each antennal prominence, tegulae light yellow. Head (Text-fig. F, 14) shallowly,
rather finely punctate, with a smooth impunctate longitudinal band between anterior
ocellus and antennal prominences, clypeal margin strongly depressed beneath the
strongly forked and tuberculate clypeofrontal longitudinal carina. Thorax with the
pronotum only slightly narrowed anteriorly. Prenotum very finely and sparsely
punctate, mesonotum closely and deeply punctate and transversely rugulose. Abdomen
short and rather broad (equal in length to thorax), very finely and sparsely punctate,
minutely transversely rugulose. Epipygium as in Text-fig. F, 15. Length (excluding
antennae) 8 mm.

Locality, etc—The type locality is Tasmania. The specimen examined was collected
by Key, Carne and Kerr at Oatlands, Tasmania, on 15th January, 1948, and is in the
collection of the Division of Entomology, C.S.1.R.O., Canberra.

HEIRONE DISPAR Westwood, 1844. (Text-fig. F.)

Arcan. Hnt., 2: 144.

Male: Colour black, mandible apices deep reddish. Head (Text-fig. F, 16) completely
punctate, clypeus without tubercles or carina. Face strongly depressed below antennal
prominences, which are not markedly outstanding. Thorax rather finely punctate,
not rugulose between punctures. Abdomen longer than thorax, dorsally flattened,
finely shallowly punctate, finely shagreened on posterior discal areas of segments.
Epipygium as in Text-fig. F, 17. Length (excluding antennae) 10 mm.

Female: Colour ferruginous, head and abdomen somewhat darker suffused, legs
testaceous. Head (Text-fig. F, 18) irregularly punctate and finely aciculate. Mandibles
bidentate. Thorax (Text-fig. F, 18) more finely and sparsely punctate than head, finely
aciculate except for the posterior face of the propodeum, which is very finely and

BY B. B. GIVEN. 325

densely punctate. Abdomeu sparsely punctate and longitudinally aciculate except for
segment 5, which is finely and rather closely punctate. Pygidium extremely thin and
transparent. Length (excluding antennae) 4 mm.

Locality, etec-——The type locality is Adelaide, and the pair examined were taken by
the writer at Cooma, N.S.W., on 3rd February, 1952. This pair is in the collection of
the Division of Entomology, C.S.1.R.0O., Canberra, and the type pair in the Oxford
Museum.

13

Text-fig. G@. Hirone and Rhagigaster.

1-6: Hirone mulesi, n. sp. *1, male, head, profile; *2, male, head, frontal; *3, male,
abdominal outline, dorsal; *4, male, epipygium; *5, female, head and thorax; *6, female,
pygidium. 7-15: Rhagigaster mandibularis Westwood. 7-9, forewing aberrations; 10-13, hind-
wing aberrations; 14-15, range in position of insertion of second recurrent nervure on third
cubital cell of forewing.

* Figures drawn from type specimens.

EIRONE MULESI, n. sp. (Text-fig. G.)

Male: Colour black, shining. Head (Text-fig. G, 1 and 2) sparsely, finely punctate.
Antennal prominences very acute and prominent. A well-developed bituberculate, almost
crescentic clypeal prominence dominant below the antennae. Thorax sparsely punctate,
pronotum medially depressed towards posterior margin. Abdomen with the first
segment strongly constricted dorsally (Text-fig. G, 3), widest towards posterior.
Sparsely, irregularly punctate. Epipygium as in Text-fig. G, 4. Length (excluding
antennae) 7-5 mm.

Female: Colour of head, antennae and abdomen black, thorax ferruginous, legs
somewhat darker. Head (Text-fig. G, 5) almost square, posterior angles rounded.
Punctation irregular. Mandibles bidentate. Thorax (Text-fig. G, 5) almost impunctate.

326 NOTES ON AUSTRALIAN THYNNINAE. II.

Abdomen very finely punctate towards anterior of segments 2 to 5, elsewhere
irregularly and very sparsely punctate. Pygidium (Text-fig. G, 6) very broad, evenly
rounded and smooth.

Types——The type pair was taken in copula at Traralgon, Victoria, on 23rd
December, 1951, by Mr. M. W. Mules, and is in the collection of the Division of
Entomology, C.S.1.R.O., Canberra, A.C.T.

Note: Apart from the species mentioned above, the writer has material representing
at least 18 further species in the genus Hirone which it has not been possible to
determine with any degree of certainty, yet which cannot be described as new because
of the taxonomic difficulties of the genus. Three forms tentatively labelled as varieties
of H. ichneumoniformis (Smith), one labelled as a variety of H. lucidula (Turner),
and one labelled “near montivaga Turner” have been the only species in this indeter-
minate residue which have received even tentative labels. It would seem probable
that the genus will prove to be unsuspectedly large once collectors pay more attention
to coccid and similar exudations. One species has been taken only on young Banksia
leaves, where no sucking insects were present and no blossom was available.

The genus is an extremely difficult one from the systematic standpoint and requires
eareful revision.

References.

GIvEN, B. B., 1953.—Proc. LINN. Soc. N.S.W., 78 (5-6): 258-261.
, 1954.—Trans. R. ent. Soc. Lond., 105.
GUIGLIA, DELFA, 1948.—Ann. Mus. Stor. nat. Genova, 63.
TURNER, R. E., 1907.—Proc. LINN. Soc. N.S.W., 32 (2): 206-290.
, 1910.—Genera Insect., fase. 105.
, 1910a.—Proc. zool. Soc. Lond., 1910: 259-306.
, 1910b6.—Trans. ent. Soc. Lond., 1910: 407-418.

327

NOTES ON AUSTRALIAN THYNNINAHE.
Ill. THe Genus THYNNOIDES.
By B. B. Given, Entomology Division, D.S.I.R., Nelson, New Zealand.
(Communicated by Dr. A. J. Nicholson.)
(Highty-eight Text-figures. )
[Read 26th November, 1958.]

Synopsis.
The definition of the genus is discussed. Keys to Known males and females are given
with the characters used figured. Two new species are described, and one male and two
females are described for species previously Known from the other sex only.

INTRODUCTION.

Although all known species of Thynnoides are dealt with in this paper the exact
status of the genus is still not clear and it seems certain that it must ultimately
encroach on other genera such as Lophocheilus and Lestricothynnus. Indeed, the entire
generic classification of the subfamily requires revision.

The basis of this paper lies in figures and use of only diagnostic characters in
descriptions. It is intended to guide more extensive work which is sure to follow.

Acknowledgement is made to the trustees and staff of the British Museum (Natural
History) and the staff of the Hope Department of Entomology, Oxford University, for
permission to examine and publish data on their collections and for assistance in many
ways. To Mr. M. W. Mules, Mr. F. E. Wilson and Mr. J. C. Le Souef, of Melbourne,
Dr. P. B. Carne and Mr. E. F. Riek, of C.S.1.R.0., Canberra, and to Mr. G. Stephens,
of Hamilton, Victoria, acknowledgement is given for collecting specimens on request.

In particular, I am indebted to Mr. EH. F. Riek and Dr. P. B. Carne for detailed
eriticism of the draft of this paper.

Genus THYNNOIDES Guérin, 1838.

Voy. Coquille, Zool. 2, (2): 214—Thynnidea Ashmead, 1903, Canad. Ent., 35: 98.—
Thynnus (Thynnus) (part) Turner, 1908, Proc. Linn. Soc. N.S.W., 33: 198.

Genotype: Thynnoides fulvipes Guérin.

Male: Hypopygium strongly produced from lateral spines or shoulders to an apical
spine, broadest at lateral spines or shoulders. Abdomen narrowly fusiform, segments
strongly constricted at base, no spines on sixth ventral segment. Epipygium terminating
in a thin, transparent margin or plate. Antennal segments not arcuate.

Female: Fifth abdominal segment ventrally rugose; pygidium narrow, longi-
tudinally carinate or rugose; second abdominal segment with five or six strongly-
raised transverse carinae. Head and thorax not coarsely punctured; anterior margin
of pronotum bearing long hairs.

Remarks on the genus—The genus is an ill-defined one, particularly as regards
males. Confusion with Lophocheilus Guérin, Lestricothynnus Turner, Zaspilothynnus
Ashmead (a few species only) and Hlidothynnus Turner cannot readily be avoided.
Zaspilothynnus may be separated in the male by the prominent, almost bulbous nature
of the clypeus (Text-fig. D, 14), the depressed subapical area on the epipygium (Text-
fig. D, 15), and the presence of ventro-lateral tubercles or spines on the sixth abdominal
segment. Zaspilothynnus females have never less than seven transverse carinae on the
second abdominal segment, a broad pygidium, usually striate fifth abdominal sternite
and broad mesotibiae in most species (Text-fig. D, 16). Hldothynnus is in many ways

PROCEEDINGS OF THE LINNEAN Society oF NEw SouTH WALES, 1958, Vol. Ixxxiii, Part 3.

G

328 NOTES ON AUSTRALIAN THYNNINAE. III,

intermediate between Zaspilothynnus and Thynnoides, the males having the bulbous
clypeus of the former but lacking tubercles or spines on the sixth sternite, and having
genitalia which would place them in Thynnoides. They may usually be separated by
the presence of yellow markings on thorax and abdomen, and lack of marked
constriction between abdominal segments. Females are structurally similar to
Thynnoides, but have radially or longitudinally arranged carinae on the fifth sternite
as in most species of Zaspilothynnus. Lestricothynnus males are extremely difficult
to separate from Thynnoides, particularly species which are black in colour. The males
are more slender and genitalia appear to present differences, but no other distinctions
have been noted. The females appear to show no better characters for separation from
Thynnoides than do the males. Had all types concerned been seen by the writer, these
two genera would probably have been combined in this paper, under Thynnoides. The
genus Lophocheilus in some respects, particularly in one or two species, is intermediate
between Thynnoides and Zaspilothynnus. The principal difference between Thynnoides
and Lophocheilus lies in the broad pygidium and greater number of transverse carinae
on the second abdominal segment of females of the iatter genus. Lophocheilus males
are generally larger and broader than are those of Thynnoides, but neither this nor
the presence of tapering antennae in the former (Turner, 1910, p. 16) is reliable. It
would seem probable that Lophocheilus villosus Guérin (Text-fig. D, 17), at least as
interpreted by Turner, should be combined with Thynnoides and the majority of the
remaining species combined with Hemithynnus.

Key to Thynnoides. (Text-figs. A, B.)

Males.
1S Wower Maron son chy peus sGOoundeds Ge AG) ane: een eee eet preissw Turn.
lower marein: of ‘clypeus: ctrMMEate: | shire oe. Suse vosas syns ore Sie oes cao he dee eee ee eu roneaae 2.
2. Abdominal segments 4 and 5 ventrally subtuberculate (Text-fig. A, 9) ............ 3.
Abdominal segments 4 and 5 not ventrally subtuberculate .....................--- fe

(ot)

Fore-coxae strongly produced posteriorly (Text-fig. A, 4) (S.A., Vie., N.S.W.) .........-.-

Aoi ch CCl MAAR Gad CLEMO OL ACE CLET CE CHA tcROLa Oo DICKS Get cid ¢ been riotal ot cho tied big oss id Did gracilis (Westw.)
Fore-coxae ‘slightly if at all produced posteriorly (Text-fig. A, 38, 5) ................. 4,

4. Mesosternal intercoxal processes small, not completely covering coxal bases (Text-fig.

yA) an dr ey NE tee Ta ly eee ae en Ena ERA MID Mee Nne TCs AURA Neoa bar GemtalG Glow o 2 5.
Mesosternal intercoxal processes completely covering coxal bases (Text-fig. A, 6) ..... 6.

5. Anterior pronotal margin not reflexed medially (Text-fig. A, 10), tegulae testaceous
GQ NESE Wed i Sapeters secdie teciolatuanccn o Bevel os avalos ay oes APE Gi Pannen eee mesopleuralis Turn.
Anterior pronotal margin reflexed medially (Text-fig. A, 11), tegulae dark (S.A., Vic.,
INE SSW, ASCP B ta RS tink Apa REA Lae ee eens alt, pom me Sl) ab ane rujfithorax Turn.

6. Prenotum sparsely finely punctate, fore-trochanters not spinose (N.S.W., Vic., S.A., A.C.T.)
PS Ia 2 OUD OTOL Gio O Oto Gide OIGICP OID oO eh ice a EEDlOlOT oO naeG ey Ging aa Oe. Oreo Git Ola a Olea Ureran pugionatus Guér.
Pronotum closely finely punctate, fore-trochanters spinose (Text-fig. A, 3), Australia ....

Esa alee Zoliehiel firs tm, felts lev wide lel terts te aprotsmple retceurastes tei arses EPSRC een TI Ean Se aE en SR bidens (Sauss.).
7. First abdominal segment ventrally subtuberculate or sharply angled medially (Text-

PS ay eA BW)! eg coerce ee dar a Spence acbeep® Ai iad seen ata “erence emer ope ce RUS CR NORE SCE REA atl cae ei sone rae 8.

First abdominal segment not ventrally subtuberculate or sharply angled (Text-fig. A, 14)

On Ce HOIS TUM ERCP ONCE ON OCCT en cae ER eee hen icra a Gio Oro EL Sto Ou Ries Medio ara oig’a-6 12.

8. auess black or idark, int COlOUt cove oy eee tee eee ee OE oe eee S).

hegsi light inticolour 2. Aa ie eae LOA Re OR a POA Aiea, | a rea eure iil.

O.cHore=coxaedstron'slyi! CONCAWV ETRE + Scitks Sie terele tabs a bites Saee pCR aT ated ss Micke Ue pote ede Met ei Sherer meas 10.

Fore-coxae not strongly concave (QU.) .........0 cee ee ee ee ee eee juscocostalis Turn.

HOS: CihAKSOS MarEMinNh ye] NEYeler TONTSEo lio oo oo coe Goaoao ooo ob smb oa hoods O06 waterhousei (Turn.)

Chrous) maim; yelllony (CaS, Wie INGSEW.) so d5caccgduaccensbs5oganue senilis (Tirichs.).

ills MUSES UESIEIGOOUS (WIG, INDSEW 5) soogsacccmces0ccdcconsdouesoce fumipennis (Westw.).

eculae* black’ “GN.SoWe VGN) oc REP Ee Ro Setar eee tena tee anon fulwipes Guér.

PZ OAbdomen) marke digwaltin tyellown GWRAS)IN me ereinicia ciel ciiehetel a breieieih el een ieienei ey eel ee lanio Turn.

Abdomen, tnot:.marked wwithy vyellow? sch aloes. maki eek doctowterepeicel sac oars a epee aoe oe oe

1339 More-COXAe CONVEX pCWeARY ait tiiercteucnensienexcuegenepouetthor wok doierad Brome sicko oees sicasusietere hee berthoudi Turn

‘MOTEC=COKAC\ ICONGAN G15 y i elie 2 De elects ien aley abr ona Posen crater alee ensure reatemelirs Reg ec Or ae arate Rees cine ate ad oie Ren RCE RE 14.
14. Clypeus black, procoxal posterior process refiexed downward (Text-fig. A, 15) (Vic.)

AERC ICRC CHOR IEEE ADE Ce OC RENCE CM ceOR cuca AC Certain Ous polo steo OpOsCw NO cy cic tects “ote tatoo lugubris, n. sp.

Clypeus yellow or yellow margined, procoxal process not reflexed .................... 15.

15. Clypeus entirely yellow, wings dark (W.A.) .........6....00.0+-0-- mephelopterus Turn.

Clypeus narrowly yellow margined, wings almost completely clear (N.S.W.) ........
aera ei) i Wee ho ee eet me aS Oe Mee hc led Tar td ois Some alma yl oloaad ci Gt cao 5 Geoneco OO o wilsoni, n.sp.

BY B. B. GIVEN. 329

Females.

1. A mid-lateral tubercle on each side of head (Text-fig. A, 16) (Vic., S.A., N.S.W., A.C.T.)
COG OS CLOG GOD CO BE OO CO CRT CATO CHORT Ee CUS BI OES gh Om ieee RECON Ce GrRNCTE Cor PRHaOR SS coos eae rufithorax Turn.

2. Propodeum acutely produced at anterior angles (Text-fig. A, 17) (W.A.) .............
© EG 2618, SUF ENOL an GNM ONSU a cI Ute RIE ura ar te MI acest OR se AN ele MS nephelopterus Turn.

Propodeum not produced at anterior anmgleS ........... ce eee eee ee ee eee ee eee Bi.
See ATteriors pronotallaneles’ producedl guia. scala sae as eerie sleleas gielers os 6 GAs a eodiaiel ele euana 4,
AMILeHOMmpPLONOtalsaneles NOt sOEOGUCEGY Wai. cic cect cue «oe ce cio cus) sre iscg ik ler sy aware le een eevee ace 6.

4. Anterior pronotal angles obtusely produced (Text-fig. A, 18) ....................0000--
0), SOO reeto OMA TEE Ga CREE CREE RCM cute fulvipes Guér., moestus (Smith), *fumipennis (Westw.).

Anterior pronotal angles acutely produced (Text-fig. A, 19) ....................... 5.

5. Pygidium conspicuousiy laterally toothed (Text-fig. A, 20) (W.A.) .......... lanio Turn.
Pygidium not conspicuously toothed (Text-fig. A, 21) (Vic.) ............ lugubris, n. sp.

6. Diseal area behind antennae with numerous coarse punctures (Text-fig. A, 22) (Qu.,
INS WAVE) we 6 9-018 Grove OeELae tenet sect ie Gl Ge cre LE IR A ee ie eran eT a mesopleuralis Turn.
Disecal area behind antennae almost impunctate ...............ce eee e eee te ee ee eee Ue

7. Head strongly elevated medially behind antennae (Text-fig. A, 23) (Tas., Vic., N.S.W.)
RRS ee otras SN eRe eee Mee cer ct meee digk ny ceeans, tame tepid tnteteat nati, aie anehey 2 senilis (Hrichs.).

ECAC MIMI ORIN ly) 6c OnViexs COTFSAl yi as she oc sic) oc bos ols ora alae sens aeto devanausiereie eieitelsrsl sways eer alane 8.

8. Carinae on second abdominal segment strongly curved (Text-fig. B, 24) (S.A., Vie.,
BNE MAVEN MG rena NC Rian cir ae yt) AME NE we fesice ose aN a aoe eh Sei gee kk Aula I gracilis (Westw.).
Carinae on second abdominal segment not strongly curved (Text-fig. B, 25) .......... 9:

9. Head subcircular or broadest level with eyes (Text-fig. B, 26, 27) ................. 10.
Eeadmbroadestsposterior itomey.es (Cihext=fis, IBS 28) Bees sewe ease = de cuieies sc cu oe sccs ss ial,

10. Posterior margin of head bearing long hairs (Text-fig. B, 26) (N.S.W., Vic., S.A., A.C.T.)
09 DOO GOO OHH of RHE nso B.b.5 Eitho ploic oO OloRstO Cotto ctOnCES ID EME ELS, EeGhole: € orern Onno ors pugionatus Guér.

Posterior margin of head without hairs (Text-fig. B, 27) (W.A.) ....... bidens (Sauss.).
11. Apex of pygidium as in Text-fig. B, 29 (N.S.W.) ................. waterhousei (Turn.).
Apex of pygidium as in Text-fig. B, 30 (Qu.) ..................... jfuscocostalis (Turn.).

* See remarks on fumipennis (Westw.).

THYNNOIDES PREISSIZ Turner, 1910.
Proc. zool. Soc. Lond., 1910, p. 284.
The male only is known.
The type in the Berlin Museum was taken in Western Australia.
Distinguished by the rounded clypeal margin, according to Turner.
Neither type nor determined material has been seen by the writer.

THYNNOIDES GRACILIS (Westwood), 1844. (Text-figs. A, 4, 9; B, 24, 31, 42; C, 1.)

Arcan. Ent., 2: 139. Thynnus (fhynnoides)—Turner, 1910, Gen. Insect., 105: 46.
Thynnoides.—dallatorrei Schulz, 1906, Spolia Hym., p. 106. Thynnus.

The type pair in the Oxford University Museum is from Adelaide. Turner (1908)
records the species also from Melbourne and from Mittagong, N.S.W.

Distinguished in the male by the prominent pronotal angles (as in pugionatus),
the ventral abdominal tubercles (as in mesopleuralis, rufithorax and pugionatus) and
the acute posterior angles of the fore-coxae.

Female distinguished by the strongly curved carinae on the second abdominal
segment.

THYNNOIDES MESOPLEURALIS Turner, 1912. (Text-figs. A, 7, 10, 22; B, 34; C, 5.)

Ann. Mag. nat. Hist. (8), 10: 539.

Cotypes are in the British Museum, South Australian Museum, Western Australian
Museum and the Queensland Museum. All were collected at Brisbane, Queensland, in
September.

The key characters (small mesosternal intercoxal processes and lack of reflection
of pronotal margin) adequately separate mesopleuralis males from other species. The
reddish mesopleurae are not peculiar to the species.

For the female, the characters given in the key appear to be adequate.

A variant of this species, collected by P. B. Carne at Binalong, N.S.W., in October,
1950, is illustrated in Text-figs. B, 40, 44, and C, 15.

GG

330 NOTES ON AUSTRALIAN THYNNINAE. III,

THYNNOIDES RUFITHORAX Turner, 1910. (Text-figs. A, 8, 11, 16; B, 36; C, 2.)

Proc. zool. Soc. Lond., 1910, p. 284.

The type female of this species is in the Berlin Museum and has not been seen
by the writer. However, the characteristics of the species are so striking that there can
be little doubt as to the identity of material examined. The type was taken at
Ararat, Victoria. Material examined by the writer was collected at Cavendish, Wannon
(near Hamilton) and Hamilton in Victoria, St. Ives and Burra in N.S.W., Canberra,

WE

AQ

zy

Text-fig. A.

1, la: T. fuscocostalis male, fore-coxa, ventral, lateral. 2: JT. waterhousei male, fore-
coxa, lateral. 3: JT. bidens male, fore-coxa, ventral. 4: TJ. gracilis male, fore-coxa, ventral.
5: T. pugionatus male, fore-coxa, ventral. 6: T. pugionatus male, mesosternal intercoxal
process. 7: T. mesopleuralis male, mesosternal intercoxal process. 8: T. rufithorax male,
metacoxa, ventral. 9: ZT. gracilis male, abdomen, ventrolateral. 10: JT. mesopleuralis male,
pronotum. 11: TJ. rufithorax male, pronotum. 12: VT. pugionatus male, pronotum. 13: JZ.
fuscocostalis male, first abdominal segment, lateral. 14: 7. berthoudi male, first abdominal
segment, lateral. 15: J. lwgubris male, fore-coxa, lateral. 16: J. rufithorax female, head,
dorsal. 17: J. nephelopterus female, propcdeum, dorsal. 18: J. fwmipennis female (type),
pronotum, dorsal. 19: T. lanio female, pronotum, dorsal. 20: T. lanio female, pygidium.
21: T. lugubris female, pygidium. 22: T. mesoplewralis female, head, dorsal. 23: T. senilis
female, head, dorsal.

and Mt. Gambier and Glencoe in South Australia. Although this is the dominant
species on Leptospermum in the Hamilton district, western Victoria, during October
and November, it is rarely met with elsewhere.

The female is adequately described by Turner and may at once be distinguished
from all other species by the lateral tubercles on the head.

BY B. B. GIVEN. 331

Male: Colour black except for yellow lateral clypeal margins, yellow mandible
bases and leg colour which varies from black to ferruginous. Fore-coxae are flat or
slightly concave; hind-coxae highly characteristic (Text-fig. A, 8). Third, fourth and
fifth abdominal segments ventrolaterally tuberculate. Length (excluding antennae)
10-18-5 mm.

The specimens on which the above description of the male is based are in the
collections of the Division of Entomology, C.S.I.R.O., Canberra, A.C.T., and the
Entomology Division, D.S.I.R., Nelson, New Zealand.

\

AG
FZ
Z
Z
DZD
FF

Text-fig. B.

24: T. gracilis female, abdominal segment 2, dorsal. 25: T. fuscocostalis female, abdominal
segment 2, dorsal. 26: T. pugionatus female, head, dorsal. 27: T. bidens female, head, dorsal.
28: T. waterhousei female, head, dorsal. 29: TT. waterhousei female, pygidium. 30: T.
fuscocostalis female, pygidium. ol: TT. gracilis female, pygidium. 32: T. bidens temale,
pygidium. 33: 7’. fumipennis female (type), pygidium. 34: T. mesopleuralis female, pygidium.
35: T. nephelopterus female, pygidium. 36: T. rufithorax female, pygidium. 37: T. fumipennis
female, pygidium. 38: T. pugionatus female, pygidium. 39: T. senilis female, pygidium
(Daylesford). 40: T. mesopleuralis female, pygidium (Binalong, N.S.W.). 41: T. fuscocostalis
female, head, dorsal. 42: TJ. gracilis female, head, dorsal. 43: V. lwgubris female. 44: T.
mesopleuralis female (Binalong, A.C.T.).

THYNNOIDES PUGIONATUS Guérin, 1838. (Text-figs. A, 5, 6, 12; B, 26, 38; C, 6.)

Voy. Coquille, Zool. 2 (2), 234.

Type male in the Genoa Museum.

Turner’s material is from Sydney, and other material examined by the writer was
collected at Drik Drik, Victoria, Clare, S.A. (H. F. Lower), and Canberra, A.C.T.
(P. B. Carne).

For determination of material, reliance had to be placed on Turner’s identification
of the British Museum material.

332 NOTES ON AUSTRALIAN THYNNINAE. III,

Turner states (1908, p. 249) that the male is almost identical with gracilis, having
similar prominent anterior pronotal angles. ;

The red colour of the mesopleurae appears to occur in some specimens only, others
being completely black. The fore-coxae (Text-fig. A, 4, 5) appear to give the best basis
for specific separation of males of these two species. It is worthy of note that the
second abdominal segment, as well as the third to fifth, is ventrally tuberculate in
pugionatus.

The female is characterized by the head having a uniformly rounded posterior
margin bearing prominent hairs (Text-fig. B, 26) and an almost impunctate disc. The
abdomen is closely but unevenly punctate.

Variations.

A. Males vary in mesopleural colour. Specimens from Drik Drik and Nelson in
Victoria and Bordertown in South Australia have the mesopleurae red, while in those
from Canberra, Clare in South Australia, and Fernbank in Victoria they are black.
Of two males from Cavendish in Victoria, one possessed black mesopleurae, the
other red.

B. Fore-coxal posterior processes in some specimens are acutely pointed (as in
Text-fig. A, 5); in others these are shorter and rounded. The acute form is the more
general, but all material from Cavendish in Victoria and Bordertown in South Australia
has the rounded form.

C. Female punctation. Specimens from Bordertown have coarser head punctation
and more sparse abdominal punctatieon than other material.

D. Female colour. Specimens from Drik Drik, Nelson and Fernbank have the
head and thorax red, those from Cavendish have only the thorax red, while other
specimens are entirely piceous to black.

Note: Specimens of the various forms are deposited in the collection of the
Division of Entomology, C.S.1.R.0., Canberra.

THYNNOIDES BIDENS (Saussure), 1868. (Text-figs. A, 3; B, 27, 32; C, 3.)

Reise Novara, Zool. 2, Hym., p. 118. (¢) Thynnus.—Turner, 1910, Gen. Insect.,
105: 46. Thynnoides.—viduus Saussure, 1868, Reise Novara, Zool. 2, Hym., p. 123 (9).

Turner (1908, p. 249) first placed this species with viduus under T. gracilis but
in 1910 (p. 46) he altered his opinion to that stated above. The types have not been
examined by the writer. ee

In the male, the acutely produced fore-trochanters (Text-fig. A, 3) are sufficiently
characteristic for diagnosis, the distinctive hypopygium (Text-fig. C, 3) giving good
supporting evidence.

The female is more difficult to dotermine with certainty, although the posteriorly
broadly rounded head devoid of long hairs (Text-fig. B, 27) is fairly characteristic.

The type locality is Australia, and material determined by Turner is from south-
eastern Australia. .

THYNNOIDES FUSCOCOSTALIS Turner, 1912. (Text-figs. A, 1, la, 13; B, 25, 30, 41; C, 7.)

Ann. Mag. nat. Hist. (8), p. 540.

Cotypes are in the British Museum, South Australian Museum, and the Queensland
Museum. The type locality is Brisbane, collected in September.

In the male, the ventrally sharply angled first abdominal segment (Text-fig. A, 13)
groups the species with waterhousei, senilis, fumipennis and fulvipes, although in none
of these is this character so strongly pronounced. The rounded margin of the fore-
coxae (Text-fig. A, 1, la) separates the species from its nearest relative, waterhouset
(Text-fig. A, 2), in which the coxae are strongly concave and ventrolaterally keeled.
In senilis this keeling is evident but more rounded and less elongate in profile.

The females of fuscocostalis and waterhousei are very similar. The differences in
the shape of the head (Text-fig. B, 28, 41) and pygidium (Text-fig. B, 29, 30) are the
most obvious characters for separation.

BY B. B. GIVEN. 333

THYNNOIWES WATERHOUSEI (Turner), 1908. (Text-figs. A, 2; B, 28, 29; C, 11.)

Proc. Linn. Soc. N.S.W., 33: 244. Thynnus—1910, Gen. Insect., 105: 46.
Thynnoides.

The type pair in the British Museum was collected at Woodford, Blue Mountains,
N.S. W.

This species is very similar to fuscocostalis, under which species characters for
separation are discussed.

THYNNOIDES SENILIS (Hrichson), 1842. (Text-figs. A, 23; B, 39; C, 12, 17-27; D, 1-13.)

Arch. Naturgesch. Berlin, 8: 263. Thynnus—Turner, 1910, Gen. Insect., 105: 46.
Thynnoides.

This species is discussed on the basis of Turner’s identification of British Museum
material, the types not having been seen. Being an extremely variable species, final
conclusions must be reserved pending careful examination of the type compared with
the various forms in collections. The type is presumed to be in the Berlin Museum.

Confusion with waterhousei, fumipennis and fulvipes is likely. In what is taken
to be the typical form, characters used in the key should be found satisfactory.
However, even in particular localities, such as Adaminaby, N.S.W., the variation within
what appears to be a single community is considerable. It is highly probable that
with longer series available, several distinct species would emerge from what at
present appears to be a heterogeneous assemblage of forms. These forms are considered
below under locality headings.

A. Western Victoria.—The closest to British Museum material identified by Turner
are pairs from Daylesford, Hamilton and Ararat. Even in these three pairs differences
in female head shape (Text-fig. D, 1, 2) and the male fore-coxal process (Text-fig. C,
22, 27) are noted. In colour, females vary from piceous (Daylesford) to ferruginous
(Ararat) on legs, head and pronotum. The pygidia of these females are as illustrated
in Text-fig. B, 39.

Specimens from Dartmoor and Bochara (near Hamilton) are very similar to those
already mentioned, but the pygidia of the females (Text-fig. D, 10) are less slender
and the shape of the hypopygia (Text-fig. C, 17, 18) and the fore-coxal process (Text-
fig. C, 22-24) of the males are different. However, all these western Victorian specimens
could be considered to be conspecific. :

B. Snowy Mountains and Monaro Plains area.—Pairs taken from Adaminaby,
Jindabyne and Kiandra show marked variation from one another as well as certain
common characteristics which would seem to separate them specifically from Victorian
forms. The elongate fore-coxal processes (Text-fig. C, 25, 26) in the males, the head
shape, lack of small internal teeth on the mandibles (Text-fig. D, 4-9) and the form of
the apex of the pygidium (Text-fig. D, 11-14) in the females separate this group
from group A above.

The Victorian, specimens were collected during December and January, while those
from N.S.W. were taken during February.

The female of senilis has not been described, and the specimens on which illus-
trations and remarks in this paper are based are in the collections of the Division
of Entomology, C.S.I.R.O., Canberra, and the Entomology Division, D.S.I.R., Nelson,
New Zealand. :

THYNNOIDES FUMIPENNIS (Westwood), 1844. (Text-figs. A, 18; B, 33, 37; C, 8.)

Arcan. Ent., 2: 108. Thynnus (Thynnoides)-—Turner, 1908, Proc. LINN. Soc.
N.S.W., 33, 248. Thynnus.—1910, Gen. Insect., 105: 46. Thynnoides.

The types of the species are in the Oxford University Museum. Localities of
material examined are Melbourne, Sydney, Croydon, Woori Yallock. Castlemaine,
Cavendish and Nigretta.

The wings of the males are conspicuously ferruginous-fumed and the veins are
ferruginous, not black as in senilis. Tegulae are light in colour. In the typical form,
males have reddish legs, but specimens from western Victoria have the legs black.

Leg colour of females is as in the males, those from eastern Victoria being red,
those from western Victoria black.

334 NOTES ON AUSTRALIAN THYNNINAE. III,

There is some confusion concerning the female of this and related species. The
type female of fumipennis, the type of moestus (= fulvipes) and the British Museum
series of fulvipes (16 females) appear to be identical. The series of fumipennis
females in the British Museum are identica} with those of the writer and not similar
to the type. It therefore appears probable that Westwood’s pair are not conspecific,
his female belonging to fulvipes, not fumipennis.

The eleven pairs collected by the writer were all taken during December, localities
being Woori Yallock, Croydon, Castlemaine, Cavendish and Nigretta.

ALA Ane

Text-fig. C.

1-16: Hypopygium, ventral and lateral, of 1, 7. gracilis male; 2, JT. rufithorax male;
8, T. bidens male; 4, T. berthoudi male; 5, T. mesopleuralis male; 6, T. pugionatus male;
7, T. juscocostalis male; 8, T. fumipennis male; 9, T. lanio male; 10, T. nephelopterus male;
11, T. waterhousei male; 12, T. senilis male; 13, T. fulvipes male; 14, T. lugubris male;
15, T. mesopleuralis male (Binalong, N.S.W.); 16, ZY. wilsoni male. 17-21: Hypopygium,
ventral, of JT. senilis male. 17, (Hamilton); 18, (Dartmoor, Bochara); 19, (Jindabyne) ;
20, (Adaminaby 1); 21, (Adaminaby 2). 22-27: Procoxal process of JZ’. senilis male.
22, (Hamilton); 28, (Dartmoor); 24, (Bochara); 25, (Jindabyne); 26, Adaminaby) ;
27, (Daylesford).

THYNNOIDES FULVIPES Guérin, 1838. (Text-fig. C, 13.)
Voy. Coquille, Zool. 2 (2), p. 2338 (¢).—? rubripes Guérin, 1838, ibid., p. 233 (¢).—
labiatus Klug, 1842, Abh. Konigl. Akad. Wiss. Berlin, 1840, p. 23 (¢).—moestus Smith,
1859, Cat. Hym. B.M., 7: 36 (2 not ¢@).

BY B. B. GIVEN. 335

The type male not having been seen, it is impossible to be certain of this species,
which is closely allied to the variable species fumipennis and senilis. Confusion over
females is mentioned under fumipennis.

Turner states occurrence to be in Victoria and New South Wales.

THYNNOIDES LANIO Turner, 1910. (Text-figs. A, 19, 20; C, 9.)
Proc. zool. Soc. Lond., 1919, p. 286.
The type pair in the British Museum is from South Perth and was collected in
February on Hucalyptus blossom.
Colour distinguishes the male, the species having the clypeus, inner margins of
orbits, mandibles and antennal prominences yellow; yellowish lines laterally behind

Cap
‘CS C5 5

eo

Text-fig. D.

1-9: Head of JT. senilis female. 1, (Daylesford); 2, (Hamilton, Ararat); 3, (Dartmoor,
Bochara) ; 4, (Jindabyne) ; 5, (Kiandra) ; 6, (Adaminaby) ; 7, (Adaminaby 2); 8, (Adaminaby
1); 9, (Adaminaby). 10-13: Pygidium of T. senilis female. 10, (Dartmoor) ; 11, (Jindabyne) ;
12, (Adaminaby 2, ete.) ; 18, (Adaminaby 1). 14-15: Zaspilothynnus hackeri male. 14, head;
15, epipygium. 16: Zaspilothynnus gilesi female, mesotibia and tarsus. 17: Lophocheilus
villosus female, pygidium.

the eyes, a transverse yellowish mark on the pronotum; the tegulae, a lateral patch on
each side of abdominal segments 2-5 and a longitudinal mark on each side of the
same segments, reddish-yellow.

The acute anterior pronotal angles (Text-fig. A, 19) distinguish the females of
this species from others in Western Australia.

THYNNOIDES BERTHOUDI Turner, 1912. (Text-figs. A, 14; C, 4.)
Ann. Mag. nat. Hist., (8), 10: 540.
The type male was taken at Waroona, Western Australia, in December.
The male of this species, like the last, has distinctive colouring. The clypeus,
mandibles, antennal prominences and a line around the head from the mandible bases

336 NOTES ON AUSTRALIAN THYNNINAE. III.

behind the eyes and ocelli yellow. Wings are moderately dark. The convex anterior
coxae are also distinctive, although this is a character shared with lanio in Western
Australia.

The yellow line behind the eyes and the convex coxae separate this species from
nephelopterus, to which it is otherwise similar.

Female unknown.

THYNNOIDES LUGUBRIS, n. sp. (Text-figs. A, 15, 21; B, 43; C, 14.)

Male entirely black. Anterior pronotal angles not angular or prominent. Fore-
coxae slightly concave, the posterior processes or angles very strongly reflexed downward
(Text-fig. A, 15). Abdomen very finely punctate dorsally, the first segment flat
ventrally, tubercles lacking on all segments. Hypopygium (Text-fig. C, 14) highly
distinctive. Length (excluding antennae) 12-14-5 mm.

Female piceous or black, the legs and sometimes the head reddish. Mandibles
very slender, each with a small inner tooth half-way between base and tip. Head
strongly convex, laterally uniformly curved, very finely punctate, shining. Pronotum
with anterior angles acute but not as prominent as in lanio (Text-fig. A, 19). Five
long, uniform transverse carinae on the second abdominal segment, the fifth carina
the highest. Abdominal puncturing fine and uniform.

Holotype male and allotype female in the collection of the Divisicn of Hntomology,
C.S.I.R.O., Canberra. Paratypes in the collection of the Entomology Division, D.S.I.R.,
Nelson, New Zealand.

The type series (eleven pairs taken in copula) were all collected at Wannon (near
Hamilton), western Victoria, on honey-sprayed foliage of Hucalyptus sp. during
September and October, 1951. The number in this series is no indication of prevalence,
since they were selected from many hundreds of thousands of pairs representing nearly
forty species feeding at the sprayed tree over a number of seasons of observation and
collection. It is probable that this species is relatively rare.

THYNNOIDES NEPHELOPTERUS Turner, 1910. (Text-figs. A, 17; B, 35; C, 10.)

Proc. zool. Soc. Lond., 1910, p. 284.

The types, which were collected at South Perth during December, are in the
British Museum. This species is found on Leptospermum and sometimes Hucalyptus
blossom, and is said to be plentiful.

The male may be distinguished by the solid yellow clypeus, which is very strongly
convex, and the strongly concave procoxae. Apart from the yellow clypeus and mandible
bases the male is black.

In the female, the acute propodeal angles (Text-fig. A, 17) are highly distinctive.

The colour is dark reddish-brown, with the discal area of the second abdominal
segment lighter.

THYNNOIDES WILSONI, n. Sp. (Text-fig. C, 16.)

Male: Anterior clypeal margin and mandibles from base to towards apex, creamy
yellow; margins of antennal prominences narrowly translucent cream-white; otherwise
black. Anterior pronotal angles not produced, rounded. Fore-coxae strongly concave.
Abdomen very finely punctate, not ventrally tuberculate. Transparent termination of
epipygium very pronounced, hypopygial spine (Text-fig. C, 16) very slender.

The holotype male, collected by F. EH. Wilson at Merimbula, N.S.W., in February,
1950, in the collection of the Division of Entomology, C.S.I.R.O., Canberra.

References.
TURNER, R. E., 1908.—A Revision of the Thynnidae of Australia. Proc. LINN. Soc. N.S.W.,
33 (1) and (2): 70-256.
, 1910.—Gen. Insect., 105

337

AUSTRALASIAN CERATOPOGONIDAE (DIPTERA, NEMATOCHRA).
Part vit: A New GENUS FROM WESTERN AUSTRALIA ATTACKING MAN.
By WiLLis W. WirtH' and Davin J. LEE.’

(Five Text-figures. )

[Read 26th November, 1958.]

Synopsis.

This paper describes a new genus and species of sandfly (Austroconops mcmillani, gen.
et sp. nov.) collected by B. McMillan near Perth in Western Australia. In its diurnal biting
habit it resembles Leptoconops and Lasiohelea, but morphologically it shows some affinity to
both Leptoconops and Culicoides.

Genus AUSTROCONOPS Wirth and Lee, gen. noy.
Generic Diagnosis.

Female: Hyes densely hairy, contiguous for a short distance above the antennae;
vertex with a sparse row of hairs arching over eyes. Antennae 15-segmented; first
segment triangular, in head capsule; second and third large and globular, second larger
than third; remaining flagellar segments 4-15 almost uniform in size; short basal
verticils present on segments 3-15. Clypeus small, with four long hairs; proboscis
short, 0:75 as long as height of eye, mouth parts well developed, mandible with six
large teeth distally. Palp four-segmented, the usual fourth and fifth segments reduced
to a single, long, curved segment; third segment swollen with a long, open, sensory
area along mesal surface extending around to ventral surface anteriorly.

Thorax robust; humeral pits well developed. Legs unmodified, unarmed, with
short hairs; hind tibial spur plumose, tibial comb of four small spines. Hind basitarsus
1-25 times length of second segment; fourth cylindrical; fifth unarmed, with small,
sharp, equal claws; empodium rudimentary.

Wing with venation as figured; costa long, to 0-83 of wing length; radial veins,
base of media and r-m strong; r-m very oblique, forming almost a straight line with
base of media and posterior side of radial cells; two broad radial cells present,
second 2-2 times as long as first; media very faint in middle section of wing, where
it forms a long petiole to the medial fork, the fork a little posterior to the level of
the medio-cubital fork; anal angle broad, alula fringed, fringe of posterior wing
margin a simple row of alternating long and short hairs; macrotrichia absent, micro-
trichia numerous, erect and imparting a milky appearance to wing.

Abdomen short, stout; apex blunt with short, inconspicuous cerci. Two large
subequal, subspherical spermathecae and a very small, oval, vestigial one present, all
without sclerotized necks.

Male: Unknown.

Genotype: Austroconops mcmillani, n. sp., the only known species.

AUSTROCONOPS MCMILLANI Wirth and Lee, n. sp. (Text-figures 1-5.)
Description.
Length of wing 0-83 mm. (from basal arculus).
Thorax and abdomen jet black; head and its appendages, legs and halteres dark
brownish black; wings dull milky white to grey with anterior veins blackish.
-Mesonotum with sparse vestiture cf erect, short, bristly, blackish hairs; scutellum

1Hntomology Research Branch, Agricultural Research Service, U.S. Department of
Agriculture, Washington 25, D.C.
2School of Public Health and Tropical Medicine, University of Sydney, N.S.W.

PROCEEDINGS OF THE LINNEAN SocreTY oF NEw SoutH WALES, 1958, Vol. Ixxxiii, Part 3.

338 AUSTRALASIAN CERATOPOGONIDAE,

bare except for a lateral pair and two submedian pairs of long black hairs. Legs
with short black hairs.

Antennal segments 4-15 in proportion of 8-9-10-10-10-10-10-10-10-10-12-16, antennal
ratio (combined lengths of segments, 11-15 divided by 4-10) 0-87; breadth of segments
decreasing from 9 units at base to 7 units at apex of flagellum, last segment without
terminal nipple. Segments of palp in proportion of 5-8-28-12, third segment 2-5 times
as long as its greatest breadth. Legs with segments from femur distad in proportion
of 45-50-25-12-8-5-8 on forelegs, 50-50-25-12-8-5-8 on midlegs and 60-55-25-20-8-5-8 on
hindlegs.

Types.—Holotype female, National Park, Perth, Western Australia, 21:xii:1954,
B. McMillan, in School of Public Health and Tropical Medicine, University of Sydney.
Paratypes: 11 females (on slides), 5 females (pinned), same data as holotype; 9 females

Text-figures 1-5.—1, wing; 2, interorbital area; 3, palp (ventral view); 4, antenna;
5, spermathecae. (Fig. 1, x 75 approx.; figs. 2, 4, 5, x 300 approx.; fig. 3, x 500 approx.)

(on slides), Yanchep Caves Park, Western Australia, 23:xii:1954, B. McMillan (biting
on eyelids, 1300 hrs.). Paratypes in United States National Museum, British Museum,
C.S.1.R.O., Canberra, B. P. Bishop Museum, Hawaii, and Western Australian Museum,
Perth.

Distribution.—This species is also known from the following localities in Western
Australia: 10 miles S.H. Darban, 29:i1:1953, J. H. Calaby (11 females pinned) ;
Armadale, 7:1:1934, K. R. Norris (two females on slide).

Discussion.—The basic affinity of the genus Azustroconops seems to be closest to
Culicoides in the Ceratopogoninae but several characters strongly suggest an affinity
with Leptoconons. Its affinities may be summarized as follows:

(a) With Leptoconopinae. Palp with incrassate third segment bearing the sensory
organ and fourth and fifth segments combined in one slender segment; third antennal
segment large and swollen, nearly as large as second segment; wing with macrotrichia

BY WILLIS W. WIRTH AND DAVID J. LEE. 339

entirely absent, microtrichia similar to those of Leptoconops, dense, giving the wing
a milky appearance; humeral pits present; two large spermathecae and a minute
third one; claws small and equal; empodium vestigial; colour black; blood-sucking
habit diurnal. However, all Lepteconopinae have antennae with less than 15 segments
(12-14) and the radial field of the wing is drastically reduced.

(b) With Ceratopogoninae. Basically the wing venation is similar to that of
Stilobezzia, with long costa and radial cells and media with long petiole; the absence
of macrotrichia on the wing, with development of microtrichia, relates it to Ceratopogon.
The blood-sucking habit, presence of humeral pits, antennal segments without
sculpturing, and the small, equal claws without empodium place Austroconops closest
to Culicoides in the tribe Culicoidini. However, no Ceratopogoninae have palps of
the form common to Austroconops and Leptoconops. Culicoides also differs in that the
wing is more or less hairy and usually with a pattern and the tarsal ratio
approximately 2.

Final disposition of the genus Austroconops would best await the discovery of the
male or, perhaps better still, the immature stages.

340

MODE OF INHERITANCE OF RESISTANCE TO POWDERY MILDEW IN BARLEY
AND EVIDENCE FOR AN ALLELIC SERIES CONDITIONING REACTION.

By N. H. Luic, K. S. McWurrtrer and HE. P. BAKER, Faculty of Agriculture,
University of Sydney.
[Read 26th November, 1958.]

Synopsis.
Inheritance studies with Australian races 3 and 18 of powdery mildew revealed that a

further five highly resistant varieties, viz., Triple Bearded Lemma, Triple Awned Lemma,
Cheroff, Monte Cristo and Unnamed B.174, carried the Algerian (Ml,) gene.

Analysis of the mode of inheritance of resistance in Triple Bearded Lemma, however,
showed that it was more complex than that based on a single dominant factor behaviour.
In the resistant variety No. 22 a different gene, designated Ml,,, apparently at the Algerian
locus, conditioned resistance. Two further varieties (Gopal and Purple Nudum) were found
to possess a gene at the Algerian locus (or closely linked to it); from the phenotypic
expression of the reaction type and differences in mode of inheritance it was designated a
further allele (MIl,,). Thus the present studies suggest that a series of at least five alleles
at the one locus may condition reaction to mildew. :

The findings in relation to the varieties Goldfoil, Chevron, Moore H.76 and Hanna suggested
that they each possess a single factor for mildew resistance, independent of the Algerian
factor; the latter, however, was linked with the single incompletely dominant factor in
Psaknon. Linkage was also indicated between the single incompletely dominant gene in
Portugese and gene Mi,, in variety No. 22. Inheritance of resistance in Duplex was explained
by the action of two independent factors, one dominant and the other recessive. These two
factors appeared to be independent of the Ml, gene in Triple Bearded Lemma. Studies did
not indicate linkage of either of the genes Ml, and Ml, with seven and one morphological factor
pairs respectively involving linkage groups I, II, III, IV and V; nor was Ml, found to be
linked with genes for resistance to P. hordei and P. gr. secalis.

INTRODUCTION.

Little of detailed studies with powdery mildew of barley has been published by
Australian investigators. Consequently there is scant previous data on physiologic
specialization, and distribution of the pathogen throughout Australia, although the
incitant organism Hrysiphe graminis hordei El. Marchal has been recognized since 1910
(N.S.W. Dept. Ag., 1935). The presence in New South Wales of race 3 was reported
by Watson and Butler (1948). Waterhouse (1948) listed several resistant varieties to
an isolate of mildew. Certain of these were susceptible to race 3 in the present study,
suggesting that his culture may have been different from that currently used.

The pathogen is fairly widespread in cereal areas in New South Wales, and
economically the disease has been described as serious annually on coastal areas (N.S.W.
Dept. Ag., 1951). The disease occurs in other Australian States and in 1956 was
particularly severe in Western Australia (J. Reeves, personal communication). That
variability exists in the pathogen is indicated by the presence of two races—3 and
18 —in the field in 1956. Again, a critical study of collections from Castle Hill
Research Station showed variations in reaction type on certain differential varieties
when compared with the standard culture of race 3 under uniform conditions. Further,
the Western Australian collection gave a different reaction on a certain extra-
differential variety to the standard race 3 culture.

In Australia the critical stage for survival of the pathogen is undoubtedly over
the summer. In Canada, Cherewick (1944) has shown that the mycelium could survive
the winter and produce conidia in the following summer. However, the disease has
been observed to persist and spread throughout the winter at Castle Hill Research

PROCEEDINGS OF THE LINNEAN Society oF NEw SouTH WALES, 1958, Vol. Ixxxiii, Part 3.

BY N. H. LUIG, K. S. MCWHIRTER AND FE. P. BAKER. 341

Station. Hence the mild winter conditions locally would not seem to restrict
multiplication in the conidial stage. Perithecia which are frequently observed on
mature plants may account for the organism’s oversummering ability.

A feature of previous extensive studies overseas by Mains and Dietz (1930),
Honecker (1931), Mains and Martini (1932), Sarasola et al. (1946), and Newton and
Cherewick (1947) has been that relatively few barley varieties were resistant when
several races of mildew were employed.

In the present study all varieties resistant in the seedling stage were also resistant
to mildew in the adult plant stage, suggesting a physiological basis for seedling
resistance. However, some seedling-susceptible varieties gave rise to highly resistant
adult plants, indicating the action of an adult plant type of resistance. Studies on
the biochemical nature of physiological resistance are being pursued at this University
(Millerd and Scott, 1955). In these investigations certain of the varieties used were
those whose mode of inheritance of resistance is presented in the present paper. The
need for a prior knowledge of the inheritance of resistance pattern in these varieties
in conjunction with biochemical studies is obvious. Detailed investigations on the
mode of inheritance of resistance of several different genes have been carried out
overseas for many years. At least twelve genes for resistance are known at different
loci. Of these, five appear to be linked and are placed in linkage group II.

REVIEW OF LITERATURE.

The review of literature is restricted to studies pertinent to the present
investigations.

As early as 1930 Dietz and Murphy had shown that the variety Goldfoil possessed
a single recessive factor for resistance to race 4. Linkage data indicated that probably
the same factor existed in the variety Pflug’s Intensiv investigated by Honecker (1934),
who reported that temperature influenced the F., type of ratio observed. Briggs and
Barry (1938) reported that Goldfoil differed in one major factor from susceptible
Atlas, and found that resistance was incompletely dominant, thereby confirming the
results of Tidd (1937).

In 1935 Briggs reported a single incompletely dominant factor (Ml) for resistance
to race 3 in the variety Hanna C.I.906.

Briggs and Stanford (1938) identified the same dominant factor (Ml.) in two
varieties, Algerian C.1.1179 and §S.P.1.45492, and Briggs (1938) reported two major
factors in the varieties Arlington Awnless, Chinerme and Nigrate. As crosses between
any two of these three latter varieties gave no susceptible segregates they were thought
to have at least one factor in common. In 1940 Stanford and Briggs found that one
of the two major factors in these varieties was identical with the factor Ml, in the
variety Psaknon. In the same paper Stanford and Briggs presented evidence demon-
strating the action of three factor pairs for resistance to race 3 in the variety Duplex,
one recessive, which was designated mla, and two dominant factors, one identical with
the Hanna factor (Ml) and the other with the Psaknon factor (Ml,)). Huntley (1951),
who worked with race 9, however, found only two dominant genes in Duplex, linked
with each other with a crossover percentage of 35:82.

Briggs (1945) reported linkage between the Psaknon gene (Ml,) and the recessive
gene in Duplex (mla) (isolated in the variety Selection 175) with a recombination
value of 16:38 per cent. In addition both genes were found to be linked with the
factor pair Atat (normal vs. albino seedling) which placed these mildew genes in
linkage group II. The cressover percentages in linkages with at were 12:08 and 36-65
for the Duplex and Psaknon genes respectively. Therefore a gene order — ai—mlq—M], —
was indicated.

Further investigations indicated the presence of the Algerian factor in the varieties
India and Multan (Schaller and Briggs) (cited by Favret, 1949) ). The same
investigators also found single factors for resistance in the varieties Mubyan C.1.6316
and Black Russian. The factor in the latter variety, designated Ml,, showed incomplete

342 INHERITANCE OF RESISTANCE TO POWDERY MILDEW IN BARLEY,

dominance in crosses with susceptible varieties, but was found to be allelic with the
dominant Algerian factor (Ml.) (Schaller and Briggs, 1955). Therefore the designation
Ml,, was introduced for the previous Ml». In the same publication Schaller and Briggs
reported the Algerian factor to be linked with three other genes for resistance, viz.
Mi. (Kwan), mla (Duplex) and Ml, (Psaknon). The following arrangement of the
five genes was given.

10-4 + 2-81 U7 se ilots (seat se ile 7
Mi, Mi, ml, M1.
+, —- ——____-__’
M1, (M1, )
4-5 a= 257

In Argentine Favret and Vallega reported a recessive gene for resistance to Arg.
race 1 in the variety Nigrate C.1.2444, and a single dominant gene for resistance to the
two Argentine races in the variety Monte Cristo (Vallega and Favret, 1947; Favret,
1949). These genes were designated or" and Oi™ respectively. The latter (Oi™*) was
Subsequently discovered also by Favret in the variety Engledow India. In the variety
Nigrate, besides or", there was at least one modifying gene for resistance to Arg. race 1
(Favret, Joc. cit.). In the same paper Favret reported studies in California using
Californian race 3. He redesignated the Oi™* gene as Mlm and confirmed his previous
results showing that the two varieties Monte Cristo and Engledow India each carried
this gene. The Monte Cristo gene (Ml,.) was found to be linked with one of the two
dominant factors Briggs had found in the varieties Nigrate, Arlington Awnless and
Chinerme. Favret stated that probably the resistance of the variety Gopal to race 3

was controlled by two dominant factors which were linked with a crossover value of
15-20 per cent.

The variety Chevron C.I.1111 was found to have two dominant additive factors for
resistance to race 6 which seemed to be independent of each other (Hehn, 1948).

MATERIALS AND METHODS.

Reaction types were recorded macroscopically on the scale described by Mains and
Dietz (1930).

Two races of H. gr. hordei were used for testing material under study. From their
behaviour on the differential varieties listed by Newton and Cherewick (1947) they
were designated as races 3 and 18 respectively. Although the infection types for race 3
on these six varieties were consistent with those reported by Newton and Cherewick,
the Australian race appeared to be a different pathogenic entity from the North
American race 3; in the present study the varieties Bolivia, Nigrinudum, Quinn, Titan,
Arlington Awnless, Mulyan and Selection dd proved seedling-susceptible, whereas North
American workers have cited these as resistant.

Although the second race used conformed closely to race 7 of the 12 races on the
key proposed by Newton and Cherewick it more closely approximated race 18 on
Moseman’s (1956) key for race identification with the same six differentials.

Race 3 was maintained on the variety Abyssinian Intermediate C.1.5808 belonging
to the species Hordeum irregulare A. & W. The variety Goldfoil which is highly
resistant (‘‘O” reaction type) to race 3, but susceptible to race 18, served for checking
the presence of contamination in race 3. Race 18 was maintained in a separate
glasshouse on seedlings of Goldfoil, and in this case the variety Abyssinian Intermediate
was used to detect the presence of race 3, as it gives a resistant reaction (“l™’) with
race 18.

Notes on the resistance to mildew in the seedling stage and in the field were
obtained for 480 varieties of barley, and the varieties C.1.3933 (*B.402), C.1.4343-1
(B.414), India 4355 (B.344), Engledow India D.1.V.464 (B.570), Buff (B.496), Unnamed
(B.174), Monte Cristo C.1.1017 (B.569), Algerian (B.573), Triple Bearded Lemma

* B numbers refer to the varietal number in the University of Sydney Barley Accession
Register.

BY N. H. LUIG, K. S. MCWHIRTER AND E. P. BAKER. 343

C.1.6766 (B.278), Triple Awned Lemma (B.473), Multan (B.355), and Cheroff (B.326).
were outstanding for their high resistance to both races of the pathogen in the seedling
stage (‘“0” and ‘“0;” reaction) and for field resistance where both races were present
with race 3 more predominant. Other resistant varieties were characterized by a
higher seedling reaction type (‘“;n” to “1*’). WHighteen varieties, seven from the highly
resistant and eleven from the resistant group, were used in inheritance studies. These,
together with their characteristic reaction types to both races, are listed in Table 1.

TABLE 1.
Resistant Parent Reaction Types to Two Races of Powdery Mildew.
Variety. | Race 3. | Race 18.
Triple Bearded Lemma B.278 C.1.6766 ata ais ESOS er ESO ears
Triple Awned Lemma B.473 Oy? | se Osea
Monte Cristo B.569 C.1.1017 Oe oo @g
Algerian B.573 Ose AO en
Unnamed B.174 Ose RNB
Multan B.355 aay SaOsne
Cheroff B.326 ae 555 (Die g Ose,
Purple Nudum B.28 ©.1.2250 Sah aie
Gopal B.267 ere ORD
No. 22 B.69 ynn” Corea
Black Russian B.574 pene ei
Hanna B.225 C.1.906 Peis sc) eae
Duplex B.172 ee yh 2 Sen
Psaknon B.81 rE ey eid 2
Chevron B.513 C.1.1111 eel Oe ee eilier a iy
Moore H.76 B.296 ooh as BG Oey LSS
Portugese B.155 ii Ne itl esate
Goldfoil B.167 ana Gos yar 2

The varieties Goldfoil, Psaknon, Gopal, Chevron, Hanna, Black Russian, Duplex,
Algerian, Monte Cristo and Multan have been studied by overseas workers, but the
remaining eight resistant varieties used in the present study are apparently not
mentioned in previous publications 1elating to inheritance studies. Therefore a short
description is given:

Triple Bearded Lemma: Obtained from C. A. Suneson, Davis, California, U.S.A.
6-rowed, triple awned, rough awn, hulled, black lemma and pericarp.

Triple Awned Lemma: Obtained from D. Robertson, Fort Collins, Colorado, U.S.A.
2-rowed, triple awned, rough awn, naked, white lemma and pericarp.

Cheroff: Obtained from College of Agriculture, Davis, California, U.S.A. 2-rowed,
single awned, rough awn, hulled, white lemma and pericarp.

Unnamed B.174: Obtained from C. K. Vears, Wagga, Australia. 6-rowed, single
awnhed, rough awn, hulled, white lemma and pericarp.

Moore H.76: Obtained from W. H. Johnston, Manitoba, Canada. 6-rowed, single
awned, smooth awn, hulled, white lemma and pericarp.

Ne. 22: Obtained from J. T. Pridham, Cowra, Australia. 6-rowed, single awned,
rough awn, hulled, white lemma and pericarp.

Purple Nudum: Obtained from H. V. Harlan, Idaho, U.S.A. 2-rowed, single awned,
rough awn, naked, purple lemma and pericarp.

Portugese: Obtained from H. Wenholz, Cowra, Australia. 6-rowed, single awned,
rough awn, hulled, white lemma and pericarp.

The genotype of the following resistant varieties was determined in crosses with
susceptible parents: Triple Bearded Lemma, Triple Awned Lemma, No. 22, Purple
Nudum, Psaknon, Duplex, Goldfoil and Portugese.

In the following crosses both parents were resistant: Triple Bearded Lemma x Triple
Awnhned Lemma, and reciprocal; Triple Bearded Lemma x Cheroff; Triple Bearded
Lemma x Multan; Triple Bearded Lemma x Gopal; Triple Bearded Lemma x Purple

344 INHERITANCE OF RESISTANCE TO POWDERY MILDEW IN BARLEY,

Nudum; Triple Bearded Lemma x Monte Cristo; Triple Bearded Lemma x Unnamed
B.174; Triple Bearded Lemma x Algerian, and reciprocal; Triple Bearded Lemma x
Black Russian; Triple Bearded Lemma x No. 22; Triple Bearded Lemma x Goldfoil;
Triple Bearded Lemma x Hanna; Triple Bearded Lemma x Moore H.76; Triple Bearded
Lemma x Chevron; Triple Bearded Lemma x Duplex; Triple Bearded Lemma x Psaknon;
Triple Awned Lemma x Purple Nudum; Purple Nudum x Unnamed B.174; Purple
Nudum x Algerian; No. 22 x Psaknon; No. 22 x Portugese.

HWXPERIMENTAL RESULTS.
(a) Crosses Between Resistant and Susceptible Varieties.
(i) Triple Bearded Lemma B.278 in crosses with susceptible varieties.

F, studies.

F, seedlings of crosses between Tripvle Bearded Lemma and susceptible varieties
gave a highly resistant reaction type (“0;”) to both races. These F, seedlings when
transplanted to the field were at all stages free from any mildew infection. Thus
resistance was completely dominant in the F, generation. Susceptible parents used in
crosses were: II 21.15 Smooth Awn x Manchuria (B.36), Skinless (B.48), Kinver (B.49),
Arlington Awnless C.I.2280 (B.19), Third Outer Glume (B.582), Long Awned Glume
(B.583) and various translocation stocks (B.289, B.291, B.292. B.527, B.528, B.529).

F, and F, studies.

F, and F, investigations were carried out in crosses between Triple Bearded Lemma
and the varieties Smooth Awn x Manchuria B.36, Kinver B.49, No. 49 B.62 and
Unnamed C.J.2220B B.28. These results are presented in Tables 2 and 3 for F, and F,
studies respectively. The deviations of the F. totals for certain crosses are highly
significant when tested for single factor pair segregation. When analysed statistically
the data were homogeneous so that the figures were pooled and a single explanation
was attempted to satisfy all results.

Initially intermediate reaction classes were grouped with susceptible reactions.
The material was tested with mildew after rust readings were obtained, and in these
instances, since the plants were at the three- and four-leaf stage, critical mildew
reactions on the first and second leaves were difficult to assess. Subsequentiy, however,
segregating material was sown solely for mildew studies and it was then possible to
record an intermediate reaction class. Hence Table 2 is presented in two sections.

Whilst classifying F, lines, counts were made of resistant and susceptible plants
in certain randomly selected segregating lines. A total of 1141 resistant to 349
susceptible plants was recorded. When tested for a 3:1 ratio a non-significant P-value
(0:20-0:10) was obtained, but an excess of resistant plants is obvious.

When intermediate and susceptible plants were grouped together in the second
section of Table 2 and then considered in conjunction with the remainder of the table,
an overall ratio of 3-48 resistant: 1 susceptible was obtained. When observed and
expected results were compared on the basis of this figure all P-values were non-
significant.

Two possible explanations are advanced to explain both the excess of highly
resistant plants expected for single factor segregation, and the presence of plants with
an intermediate reaction. In the first hypothesis a single major gene is proposed with
a differential transmission rate for the gametes containing the different alleles.
Preferentially the dominant is favoured in transmission. If such a transmission rate
of 0:528:0-472 is assumed, a ratio of 3-48:1 (resistant:susceptible) is obtained. Segrega-
tion of modifying genes for resistance could explain the presence of plants with
intermediate type reactions in cases where susceptibility would be expected due to
absence of the major gene.

In the second case two factor segregation could explain the results observed.
If two independent factors (one dominant, the other recessive) were involved, then
only one F, line in 16 would be homozygous susceptible. The F, data exclude this

BY N. H. LUIG, K. S. MCWHIRTER AND E. P. BAKER. 345

TABLE 2.
F, Miidew Segregation in Crosses Involving T'riple Bearded Lemma and Susceptible Varieties.
Number of Seedlings.
Re- Inter- Sus-
Susceptible Cross No. | Family. | sistant. mediate. ceptible. yee P-value. =, P-value.
Parent. AEs se yn_g-n | « 3+_4” (Bs 1) (3-48 : 1)
Reaction | Reaction | Reaction
Type. Type. Type.
Smooth Awnx II 52.18 1 307 -- 82
Manchuria 2 314 — 86
B.36. 3 168 _ 38
A 311 _ 89
Total 2B a3 re 1,100 — 295 11-045 |<0-001 1-019 0-50-
0-30
Smooth Awn TI 53.69 ail 51 — 21
Manchuria 22; 24 — 5
B.36. 3 12 —— 3
4 14 —_ 1 i
5 83 — 20 |
6 16 —
od 12 —- 3
8 141 — 46
Total a a Ss 353 — 105 1-051 0-50-— 0-114 0-80-
0-30 0-70
Smooth Awnx II 54.8 all 139 7 46
Manchuria 22 169 6 42
B.36 3 94 4 13
4 240 9 65
m3) 208 8 48
6 137 3 21
oll 263 15 53
Total Se ae 1,250 52 288
Total (Res. v|s. Int. and
SUS!) Fe Pe 1,250 340 11-090 |<0-001 0-739 0-50-
0-30
Smooth Awn TT 55.26 Al 48 3 9
Manchuria 2 62 ~ 3 12
B.36. 3 59 5 13
A 56 6 22
5 7 5 19
Total oe ee 299 22 75
Total (Res. v|js. Int. and
SUS!) yee Bs we 299 — 97 0-054 | 0-90—| 1-118 | 0-30-
0-80 0-20
Kinver B.49 .. II 55.104 1 18 _ 6
2 60 6 17
3 24 — 7
4 29 2 a
5 a = 5
6 36 1 6
a 1 — —
8 28 — 8
| 9 38 2) 6
i 10 2 — 1
| lil 249 16 57
Total a a: oo 492 27 120
Total (Res. vjs. Int. and
SWS) os Be ae 492 — 147 os 0-30- 0-054 0-90-
| 0-20 0-80

346 INHERITANCE OF RESISTANCE TO POWDERY MILDEW IN BARLEY,

TABLE 2.—Continued.
F, Mildew Segregation in Crosses Involving Triple Bearded Lemma and Susceptible Varieties.

| Number of Seedlings. |
Re- Inter- Sus- |
Susceptible Cross No. | Family. | sistant. mediate. ceptible. ee P-value. x2. P-value.
Parent. “S07 % ee n-270 | “ 3r—4 >? (835 il) (3°48: 1)
Reaction | Reaction | Reaction
Type. Type. Type.
No. ATR oo |) TRAIT! Mh a 106 a 41
2 99 = 29 | |
3 166 — 49 |
4 58 — 28 |
5 142 — 41
Total A es 571 = 188 0-022 | 0:90- | 2-730 | 0-10-
0-80 0-05
Unnamed II 52.19 all 103 — 26
C.1.2220B !
B.23 |
H | !
Total oS oe as 103 — 26 . 1-614 0:30- 0-338 0-70-
0-20 0-50
Test for Heterogeneity.
| | ‘
Susceptible Variety. Resistant. | Susceptible. Total. X22 | P-value.
(3:48 : 1.)
Smooth Awn x Manchuria B.36 Fite a4 3,002 837 3,839 0-603 0-50-0-30
IN@, {0) IBY 6 oa a nt ve 571 188 759 2-623 0-20-0-10
Kinver B.49 .. ae Ae ou oe | 492 147 639 0-172 0-70-0-50
C.1.2220B B.23 ss 5% a ais 103 26 129 0-348 0-70-0-50
Total ae LP me Be 4,168 1,198 5,366 0-000 1-00
i |

Heterogeneity : ,%*=3-746; d.f.=3; P-value—0-30-0:20.

possibility. However, if linkage between two dominant genes, similar in effect, with
a low recombination value, is proposed, an F, ratio somewhat greater than 3:1 will
be expected. For a ratio of 3:48:1, approximately 5 per cent. recombination would be
required with segregation of modifying genes for resistance aifecting the susceptible
class. If modifying genes are considered to affect the resistant group causing an
intermediate reaction in certain cases, then the recombination value would be of the
order of 14 per cent.

TABLE 3.

F; Breeding Behaviour to Mildew of the Cross between Triple Bearded
Lemma (Resistant) and Smooth Awnx Manchuria B.36 (Susceptible).

F; Behaviour.

!
|

| | Total.
Homozygous | Searecatine ee ee |
Resistant. | greg = Susceptible. |
118 220 85 =

A modification of the two gene hypothesis would be to suggest two linked genes
unequal in effect—one dominant and epistatic and the other in the heterozygous

i

BY N. H. LUIG, K. S. MCWHIRTER AND E. P. BAKER. 347

condition showing an intermediate reaction. No linkage value can be calculated to
satisfy these conditions statistically, as too few intermediate type plants are observed
when considered in conjunction with the susceptible class. For this hypothesis,
segregation of modifying genes, which eause plants heterozygous for the second gene
(normally resistant) to become intermediate, would need to act.

F, data do not permit either of the two hypotheses to be definitely favoured. It
will be noted that the numbers of F, lines resistant:segregating:susceptible were
118:220:85; with the differential transmission rate postulated, the expected number of
lines would be 118:212:93 (x? = 0-990, P = 0:70-0:50), whilst with two linked genes
with 14 per cent. recombination the expected ratio would be 137:208:78 (x? = 3-956,
P = 0:20-0:10). Given 5 per cent. recombination, the corresponding ratio would be
123:205:95 (x? = 2-328, P = 0:50-0:30).

It would not be necessary to invoke the action of modifying genes on the two gene
hypothesis if a differential gametic transmission rate is considered in conjunction with
linkage. In this case the intermediate reaction class would be due to the second
gene being in the heterozygous state. However, it becomes impossible to estimate the
differential transmission rate and linkage value since both can vary simultaneously
in different ways to give the same F. and F, behaviour.

(ii) Triple Awned Lemma B.473 in crosses with susceptible varieties.

The variety Triple Awned Lemma was also crossed with susceptible Smooth
Awn x Manchuria B.36, and the F. segregation was studied (Table 4).

TABLE 4.
F, Mildew Reactions of the Cross Triple Awned Lemma (Smooth Awn x Manchuria) B.36.

i

i}
F, Segregation. | x2 | x2
= Total. | (3:1). P-value. | (3-48: 1). P-value.
Resistant. Susceptible. | | |
wees
112 | | 0-80-0-70 0-188 0-70-0-50
i}

35 147 me) erat |

The ancestry of Triple Awned Lemma in relation to Triple Bearded Lemma is
unknown. They differ in many morphclogical features. The triple awned condition
presumably arose in segregation from certain crosses (Martini and Harlan, 1942), and
these varieties may he different segregates from such material. On the other hand
one may have been a progenitor of the other. In any case there is no reason to
suggest that the inheritance pattern in these two varieties would be ‘lifferent. Although
a good fit statistically was obtaind in the cross with Smooth Awn x Manchuria for the
segregation of a single dominant factor, the number of plants was appreciably less than
that obtained in the cross of Triple Bearded Lemma with the same susceptible variety.
A ratio of 3:48 to 1 (resistant:susceptible) indicative of the behaviour of Triple
Bearded Lemma, however, also fits the observed results satisfactorily. Hence Triple
Awned Lemma is considered to possess the same factor(s) for resistance as Triple
Bearded Lemma.

(iii) No. 22 B.69 in crosses with susceptible varieties.
F, studies.

The F, generation in crosses with susceptible varieties was studied for reaction in
both seedling and adult plant stages. An intermediate reaction was exhibited as
seedlings but F, plants were fully resistant in the fieid to races 3 and 18.

F, studies.

Segregation results in F.. for crosses of No. 22 with susceptible varieties are set out
in Table 5. These results indicated that a singie factor was responsible for resistance
in No. 22. The F, seedlings were grouped into susceptible and resistant, with the
latter including the intermediate types. Due to the characteristic reaction type of
No. 22 it was not possible to distinguish intermediate and resistant reactions. Before

348 INHERITANCE OF RESISTANCE TO POWDERY MILDEW IN BARLEY,

the necrosis associated with the mildew reaction of No. 22 develops the reaction type
based on amount of mycelial development would be rated higher than that finally
assessed as characteristic for the variety.

TABLE 5.
Seedling Reaction to Race 3 of Powdery Mildew of F, Populations from Crosses between No. 22 (B.69) and Susceptible
Varieties.
| F, Reaction. x2
Second Parent. — (3:1). P-value.
Resistant. Susceptible. Total.

Smooth Awn x Manchuria B.36 .. bes 335 114 449 0-036 0-90-0-80
Smooth Awn x Manchuria B.37 .. my 63 18 81 0-333 0-70-0-50
Arlington Awnless B.19 .. aie ee) 122 42 164 0-033 0-90-0-80
Sel. 163 Manchuria B.40 .. ae oh 80 29 109 0-150 0-70-0-50
Minn. 184 Manchuria B.43 as i 91 28 119 0-137 0-80-0-70
Manchurian B.44 .. ee ae ms 23 6 29 0-287 0-70-0-50
No. 305 B.85 a6 a ve ie 31 11 42 0-032 0-90-0-80
Colsess B.97.. ih an Me arb 71 21 92 0-232 0-70-0-50
Orge 14J. B.101_ .. yh as ae 56 22 78 0-427 0-70-0-50
Featherston B.129 .. oe AS ie 29 13 42 0-794 0-50-0-30
Oderbrucker B.131 Re ay BE 36 12 48 0-000 1-00
Hooded Spring B.135 br a are 42 19 61 1-230 0-30-0-20
Horsford B.140 a es ae Ba 46 13 59 0-277 0-70-0-50
Glabron B.195 at si BA a 69 18 87 0-862 0-50-0-30
Oderbrucker B.254 ah ne Ae 63 23 | 86 0-139 0-80-0-70

Total ae A 3 ot 1,157 389 1,546 0-022 0-90-0-80

F, studies.

Three crosses between No. 22 and susceptible varieties were studied in the F,
generation for their behaviour to race 3. The data are outlined in Table 6 and fit a
1:2:1 ratio statistically.

TABLE 6.
Segregation of F, Lines of Crosses between No. 22 (B.69) and Susceptible Varieties for Reaction to Race 3 of Barley Mildew.

|
F,; Segregation.
Second Parent. — , x? P-value.
Re- Segre- Sus- (il 348 1b),
sistant. gated. ceptible. Rowell
Smooth Awn x Manchuria B.36 as 6 16 12 34 2-235 0-50-0-30
Smooth Awn x Manchuria B.37 ais 8 21 6 35 1-628 0-50-0-30
Arlington Awnless B.19 pee ch 13 22 4 39 2-795 0-30-0-20
Total .. Ne a ne 27 59 22 108 1-398 0:50-0:30
|

(iv) Purple Nudum B.28 in crosses with susceptible varieties.
FF, studies.

F, seedlings of a cress between the resistant variety Purple Nudum B.28 and the
susceptible variety Orge 14th J. (B.101) were tested with race 3. The results indicated
segregation of a single incompletely dominant factor pair for resistance (Table 7).
These results were confirmed in later studies when F, seedlings of crosses involving
Purple Nudum with seven susceptible varieties were tested for their reaction to Puccinia
graminis secalis and to race 3 or barley mildew. At the time the mildew reactions
were taken most seedlings had already reached the four-leaf stage and because of
difficulties in distinguishing between the resistant and intermediate classes they were
grouped together. Out of a total of 481 plants 109 susceptible were counted and the
fit for a 3:1 ratio was satisfactory (x2 = 1-403; P-value = 0:50-0:20).

BY N. H. LUIG, K. S. MCWHIRTER AND E. P. BAKER. 349

TABLE 7.
Reaction to Race 3 of F, Seedlings of Certain Barley Crosses.
F, Segregation. | 5
Parents and Total. 28 : P-value.
a : (Ratio).
Reaction. ReTICtAnE Intermediate. Srecentinle
: i (React.) E
Purple Nudum, “*;2’’x 54 119 69 242 1-925 0-50-0-30
Orge 14th J., “4” (38 Tar 9) (i322 1)
Skinless, ‘‘ 4’ x Goldfoil, 59 90 53 202 2-753 0-30-0-20
Su) ee (C5 The) (al 3278 1)
Skinless, ““4”x .. nit 97 — 35 132 5-225 0-05-0-02
(138: 3)
Duplex, “;,12” — — — — 0-162 0-70-0-50
(3:1)
Skinless, ‘‘ 4’ x Psaknon, 81 — | 25 106 0-113 0-80-0-70
Sosa 2 (3:1)
No. 49, “‘4’ x Portugese, 461 — 149 610 0-107 0-80-0-70
“qn” (82 1)
F, studies.

Evidence for a single factor pair for resistance in Purple Nudum was also obtained
when 179 F, lines of a cross between Purple Nudum and C.J.2220B were tested for
their behaviour to race 3 (Table 8).

TABLE 8.
Reaction of F., Lines of a Cross between Purple Nudum (B.28) and Variety C.1.2220B (B.23).
Homozygous | Segregating. Homozygous Total. x P-value.
Resistant. | Susceptible. | (ile 28 i)
47 | 88 | 44 | 179 0-154 0:95-0:90

(v) Goldfoil, Duplex, Psaknon and Portugese in crosses with susceptible
varieties.

The results of these investigations are presented in Table 7. In the case of the
cross involving the variety Goldfoil the tests were carried out with race 3; Goldfoil is
fully susceptible to race 18 as previously indicated. The results of Briggs and Barry
(1938), who worked with Californian race 3, to which the Goldfoil gene was incompletely
dominant, were confirmed.

The variety Duplex was found by Stanford and Briggs (1940) to combine three
genes for resistance to Californian race 3, one recessive gene, Mla, and two dominant
genes, Ml, and Mln. The recessive Duplex gene, mla, is excluded from inheritance
studies with Australian races, as the variety Selection dd, which carries this gene

TABLE 9.
F, Breeding Behaviour to Race 3 of Powdery Mildew of a Cross between Skinless (Susceptible) and
Duplex (Resistant).

Behaviour of F, Line. Observed. Expected.* 507.
All resistant = abs or ee we 61 56 0-446
Segregating approximately 3: 1 (resistant : sus-
ceptible) one a ee ae ss 45 48 0-186
Segregating mostly susceptible, some resistant 14 16 0-250
All susceptible ve a Eu ae ue 8 8 0-000
Total a at ay 10 af 128 128 0-882

P (3 d.f.) =0-90-0-80.
* The theoretical expectancy was calculated on the basis of segregation of two independent

factors for resistance, one dominant and one recessive. The x? fora ratio of 1 homozygous resistant :
2 segregating : 1 homozygous susceptible was 41-719 with a P-value less than 0-001.

350 INHERITANCE OF RESISTANCE 10 POWDERY MILDEW IN BARLEY,

alone, is susceptible to both races. From the behaviour of F, lines of a cross between
Duplex and Skinless (susceptible) clear evidence was obtained that two genes were
involved in conditioning resistance of Duplex to Australian race 3. The results are
presented in Table 9. One gene was dominant, the other recessive. Although the F,
figures presented in Table 7 favour statistically a single factor difference, F, analysis
precluded this hypothesis.

The figures for the cross Skinless with Psaknon confirmed the results of previously
cited work, which showed monofactorial segregation in crosses between Psaknon and
susceptible varieties. Moreover, 99 F, lines of this cross were tested with race 3 and a
segregation ratio of 31 lines homozygous resistant : 45 segregating : 23 homozygous
susceptible was obtained (x? for a 1:2:1 ratio = 2:111; P-value = 0:50-0:30). Among a
random sample of segregating F, lines 433 resistant and 151 susceptible were counted
(5° for a 3:1 ratio = 0-228, P = 0:70-0:50).

F, and F, generations of a cross between Portugese and the susceptible variety
No. 49 were tested with race 3 to determine the mode of inheritance of resistance in
Portugese. The F, resuits, presented in Table 7, indicated the presence of a single
incompletely dominant gene. The F, lines, although only 29 in number, gave 6
resistant : 14 segregating : 9 susceptible, thus agreeing with this hypothesis (x? for a
1:2:1 ratio = 0-205; P-value = 0:95-0:90).

(0) Crosses Between Resistant Varieties.
F, and F, seedlings of several crosses involving Triple Bearded Lemma with other
resistant varieties were tested with one or cther race to determine allelism or otherwise

TABLE 10.
Segregation in F, Populations of Certain Crosses involving the Resistant Varieties Triple Bearded Lemma B.278 and Purple
Nudum B.28.

F, Segregation. | Upper Limit
; | xe. of Recom-
Parents and Reaction. Highly Bol Total. (3:1). P-value. | bination at
Re- Sus-
Re- : ‘ 0-05 Level.
wa sistant. ceptible.
sistant.
Triple Bearded Lemma, “‘ 0; ’’ x
Triple Awned Lemma, ‘‘ 0; ”’ 874 — —- | 874 — — 11-83%
Triple Bearded Lemma, “‘ 0; ’’ x
Cheroff, “ 0;” near ciit 176 = = ae GG == es 26-05%
Triple Bearded Lemma, *‘ 0; ’’ x
Multan, “0;”’ of af 114 = = | ee = 32-26%
Triple Bearded Lemma, “‘ 0; ”’ x
Unnamed B.174, “0; ” as 496 — oe 496 — — 15-48%
Triple Bearded Lemma, “‘ 0;’’ x
Monte Cristo, “ 0; ”’ j 223 — — 223 = — 23-24%
Triple Bearded Lemma, “ 0; ”’
Black Russian, “‘ 2 ”’ é 138 22 — 160 10-800 0-001 SEilalie/ons
Triple Bearded Lemma, “‘ 0; ” | [
Algerian, “* 0;” ae 382 — - 382 -—— — ste
Algerian, “‘ 0; ’ x Triple Bearded | Hoga
Lemma, “ 0; ”’ ash bb 97 — a 97 — —
Triple ee Lemma, “‘ 0; ”’ x |
Gopal, ‘* ;n”’ ae 360 119 — 479 0-005 0:98-0:95 1°25%*
Triple Beane Tema, 7 “<Q: eX
Purple Nudum, “ ;n”’ s 392 165 — | 557 6-349 0-02-0-01 1:10%*
Triple Awned Lemma, “ 0; ” |
Purple Nudum, ‘‘ ;n”’ ae 429 147 a= 576 0-083 0-80-0-70 I 0796*
Purple Nudum, ‘“;n”’ x Al-
gerian, “* 0;”’ ore ate 144 — — 144 — = 4-19%*
Purple Nudum, ‘“;n” x Un-
named B.174, “‘ 0;” He? 231 — — 231 a —- 2-64%*
Purple Nudum, “ ;n’’ x Monte |
Cristo, “0; > 169 — = 169 == = 3-58%*
Triple Bearded Lame,” sel ZENG
No. 22, “* jon ae Aa 348 108 — 456 0-421 0-50 1-28%*

* On basis of linkage, one gene dominant. one incompletely dominant.

BY N. H. LUIG, K. S. MCWHIRTER AND E. P. BAKER. 351

of the factors for resistance. F., and F, results are presented in Tables 10 and 11
respectively. From a cross Triple Bearded Lemma x Triple Awned Lemma $874 F,
seedlings were tested but no susceptible segregates were observed. The presence of
the same (or an allelic) factor in both varieties is thus suggested.

Likewise no susceptible segregates were obtained in a cross between Triple Bearded
Lemma and the resistant variety Cheroff (B.326); all 176 F. seedlings tested were as
resistant as the two parents. No susceptible segregates were obtained in a cross
between the highly resistant Unnamed B.174 with Triple Bearded Lemma. The
absence of susceptible segregates in the F, and F, generations of the cross between
Triple Bearded Lemma and Multan B.355 indicates further that the genes for
resistance in these two varieties are probably allelomorphic and also identical. Since,

TABLE 11.
F, Breeding Behaviour to Race 3 of Mildew in Certain Barley Crosses.

i aes
F, Behaviour. S25 = 2
Parents and Reaction. BE Boe
Homozygous ; Homozygous Hose
ee x pee 5 z = al. <>)
Resistant. SERRA NY Susceptible. Hoel a
Triple Bearded Lemma, “‘ 0;’’ x Multan, “‘0;”’ | 48 — — 48 (6-10%)
Triple Bearded Lemma, “ 0;* x No. 22,170” | 51 — — 51 (5-80%)

as mentioned in the literature review, Schaller and Briggs (cited by Favret, 1949)
have identified a single dominant gene in Multan identical with that in Algerian, it
follows that the varieties Triple Bearded Lemma, Triple Awned Lemma, Cheroff and
Unnamed B.174 probably all possess the Algerian factor (Mla). Confirmation of this
fact is shown in the cross of Triple Bearded Lemma x Algerian and its reciprocal
when only highly resistant F, plants were observed in a population of 479 individuals.

Schaller and Briggs (1955) have shown that the variety Black Russian carries a
different allele (Ml..), conferring a resistance with a higher reaction type at the
Algerian locus. As would be expected, therefore, all 160 F., plants in the cross Triple
Bearded Lemma x Black Russian were resistant or moderately so. A significant excess
of highly resistant plants was obtained from a 3:1 ratio; however, a similar behaviour
of Triple Bearded Lemma in crosses with susceptible varieties has already been
mentioned. Some of the 22 F, plants in the resistant group were only moderately
resistant with a higher reaction type than the Black Russian parent, which itself
gives a moderately resistant reaction type to Australian races. This was probably
due to the segregation of modifying factors affecting reaction type.

When F, generation material of certain crosses between the highly resistant
varieties Triple Bearded Lemma, Triple Awned Lemma and Unnamed B.174 (reaction
types “0;’’) and two other resistant parents Gopal and Purple Nudum (reaction types
“;1-™’) were tested, no segregation for susceptibility was apparent. Similar results
were obtained with both races. As will be seen from Table 10, segregation of high
resistance versus resistance occurred in the cross with Gopal in a ratio of approxi-
mately 3 to 1, indicating dominance of the gene for high resistance. However,
significant deviations from the 3:1 ratio were evident in one cross involving Purple
Nudum and the total for crosses involving Purple Nudum does not statistically satisfy
a 3:1 ratio. However, it was noted that the reaction type of Purple Nudum (‘“;n’’)
under high temperatures is little different from the reaction of Triple Bearded Lemma
CBD

This behaviour in a total F, population approaching 1700 plants, in crosses
involving Purple Nudum with Algerian or varieties presumably carrying the Algerian
factor (Monte Cristo, Unnamed B.174, Triple Bearded Lemma, Triple Awned Lemma),
suggests that an allele at the Algerian (Ml.) locus is operative in the variety Purple
Nudum. In an F, population of 479 plants in the cross Triple Bearded Lemma x Gopal,
no reactions higher than the ‘“;"’ characteristic of the male parent were observed,
suggesting that an allele at the Algerian locus is also operative in Gopal. Since the

352 INHERITANCE OF RESISTANCE TO POWDERY MILDEW IN BARLEY,

reaction types of Purple Nudum and Gopal are apparently identical and both allelic
with the same gene, one may conclude that the genes for mildew resistance are
identical in these two varieties. Unfortunately, no material of a cross between these
varieties was available for study. Both appear identical in morphological charac-
teristics. Besides the difference in reaction type with the Algerian gene, studies
reported earlier in this paper show that the gene in Purple Nudum was only
incompletely dominant in crosses with susceptible varieties, whilst the resistance of
Triple Bearded Lemma behaved in a completely dominant fashion in such crosses.

Further, in a cross of Triple Bearded Lemma with the resistant variety No. 22
(infection type “‘1™’) no susceptible segregates apveared in the 456 F, plants, nor in
51 F, lines, suggesting that these varietics also possess allelic factors for resistance.
The behaviour of the variety No. 22 to mildew is characteristic and of a “1” type
reaction with pronounced necrosis, the reaction type being best described as “1™”. The
necrosis is so severe that in heavy infections it leads to the death of the leaves of
the seedlings. It is a very distinctive type of behaviour in comparison with a typically
hypersensitive reaction.

As mentioned in the literature review, the same single dominant gene (MJ],,) has
been shown to operate in the varieties Monte Cristo and EHEngledow India (Favret,
1949). The variety Monte Cristo (reaction type “0;’) was crossed with both Triple
Bearded Lemma and Purple Nudum but no susceptible segregates occurred in either
cross in the F, generation, indicating that the Algerian factor is operative in Monte
Cristo.

Summarizing these results on the inheritance of resistance in varieties possessing
the Ml. gene or an allele and adding, with certain modifications, results of other
investigators (Schaller and Briggs, 1955; Briggs and Stanford, 1938; Favret, 1949),
the following allelic series for all varieties carrying a gene for resistance to mildew
at the Algerian locus is suggested:

Phenotypic Expression to
Variety. Gene. Nature of ie
Resistance. Australian Californian
Race 3. Race 3.
Algerian ne 2s ae Ml, D SOS £303"?
India .. bb a a Mla D oo ()p. 22 Oe
Multan .. ee on Be Mla D 0 | 02
S.P.1.45492 ae at ara Mla D — STOR
Monte Cristo .. ys me Mla D + O22 ead ee
Engledow India ai ne Mla D ° @g SQ see
Triple Bearded Lemma Ne Mla D of Oe —
Triple Awned Lemma ad Mla, D S078 —_
Cheroff .. ape an ea Mla D “0: —_—
Unnamed B.174 PAS oe Mla D Fo (ORE _
Black Russian . . a We Mla» d Oo aie [2
No. 22 an a as Mlas d OS qpaal Se
Purple Nudum ae cs Mla; d payed 2 —
Gopal .. i Fe ae Mla; d aes =
(Susceptible variety) .. ae Ml, R uke Peet ee

D=dominant ; d=incompletely dominant; R=recessive.

Gopal was suggested by Favret as possessing two domirant genes linked with
15-20 per cent. crossing over. Present studies herein reported indicate that one of
these is allelic with the Algerian gene. With Australian races the reaction types of
Purple Nudum and Gopal are identical and as these varieties appear similar morpho-
logically to one another, Gopal may, like Purple Nudum, possess a single gene to local
pathogenic entities.

These studies further suggest that the Mlm gene (designated by Favret) in Monte
Cristo is allelic with the Algerian factor and probably identical with it. This conclusion
is based on the assumption that the same gene is operative in both studies. Hngledow

BY N. H. LUIG, K. S. MCWHIRTER AND E. P. BAKER. 353

India was shown by Favret to carry the same gene as Monte Cristo and hence is
likewise listed as possessing the Algerian gene.

Investigations in inheritance were carried out in crosses with Triple Bearded
Lemma and varieties carrying other than the Algerian or factors allelic to it. KF, results
for such crosses are included in Table 12. Before testing the behaviour of Triple
Bearded Lemma in relation to Goldfoil, the mode of inheritance of the latter gene
was studied in crosses with a susceptible variety, as reported earlier in this study.

As the Goldfoil gene has been previously placed in linkage group IV, and Triple
Bearded Lemma in these studies carries the Algerian factor (Ml.) which Briggs and
Stanford (1938) placed in iinkage group II, independent segregation of the two genes
was expected. This was indicated although only a small amount of material was
available for studies from a cross between the two varieties.

TABLE 12.
Segregation for Mildew Reaction in F, Populations of Certain Barley Crosses involving Triple Bearded Lemma B.278.
F, Segregation. |
; x*
Parents and Reaction. ie Inter- ani Total. (Ratio). P-value.
sistant NSE ceptible
: (React.). P 7
Triple Bearded Lemma, “‘ 0; ’’ x Goldfoil, 33 4 3 40 0-308 0-90-0-80
“<Q” (CU) (13 : 2:1)
Triple Bearded Lemma, “ 0;*’ x Hanna, 62 12 2 76 2-251 0-50-0-30
‘6 0; 2 (Ge in me) (13 SDE 1)
Triple Bearded Lemma, “ 0:’’ x Moore 380 74 26 480 4-056 0-20-0-10
TBIS@Ab5 RT (7 Tere >) (1B § Ae IW)
Triple Bearded Lemma, “‘ 0; ’’ x Chevron, 264 38 20 322 0-149 0-95-0-90
“=n (© 1+n”’) (i133 3 93.8 11)
Triple Bearded Lemma, “‘ 0; *’ x Duplex, 259 76 18 353 0-516 0-80-0-70
Sos iCal) | (48 : 18 : 3)
Highly Re- Inter- Sus-
Re- sistant mediate ceptible Total. xe P-value.
| Sua (il ™). (“ {tt 22) (EA), (Ratio).
f OG
Triple Bearded Lemma, “* 0; ”’ x 715 135 68 28 946 19-441 <0-001
Psaknon, “17-2” “Pre (15:1)
(Resistant + Intermediate) :
| (Susceptible)
| | H

Triple Bearded Lemma was also crossed with the variety Hanna C.1.906 (B.225),
which is highly resistant to both Australian races. Hanna also exhibited a “0;” type
of reaction. Briggs and Stanford (1938) found no evidence of linkage between the
Hanna factor (Mln) and the Algerian factor (Ml.) in the two varieties S.P.1.45492 and
Algerian. The results of the cross Triple Bearded Lemma x Hanna currently studied
are in agreement with their findings. The 12 seedlings in the population of 76,
exhibiting a ‘1’ type reaction are probably due to the incompletely dominant Hanna
gene conferring this type of reaction in the heterozygous condition.

The relationship of the factors for resistance in Triple Bearded Lemma and
Moore H.76 (B.296) were also studied. The latter variety is very resistant to both
Australian races; with race 18 a highly resistant reaction ‘0;’’ was recorded, whilst
with race 3, Moore H.76 showed a slight range from a “;” to a “1”’ type of reaction,
presumably due to environmental effects. F, seedlings of three families of this cross
were tested with race 3, and seedlings from a fourth family were tested with race 18,
The results with both races suggested independent inheritance of a single incompletely
dominant gene in Moore H.76. Of the 74 semi-resistant plants, approximately one-third
only could be due to the segregation of a possible second factor in Triple Bearded

354 INHERITANCE OF RESISTANCE TO POWDERY MILDEW IN BARLEY,

Lemma. The remaining two-thirds (approximately 50 in number) fit well a 2:1 ratio
for segregation intermediate : susceptible due to the Moore H.76 gene.

In the cross Triple Bearded Lemma x Chevron, in a total of 322 F. seedlings
38 semi-resistant (“1*™’) and 20 fully susceptible plants were recorded when race 18
was used. These figures are in close agreement with a 13:2:1 ratio. Chevron itself
gave a “1-"’ reaction but it was not possible in the F. generation to distinguish those
plants which showed the Triple Bearded Lemma from the Chevron type of resistance.
Hehn (1948), however, reported that two independently inherited factor pairs seemed
to control reaction to mildew (predominantly race 6) in a cross involving susceptible
Manchuria and resistant Chevron. He thought the genes to be additive in increasing
resistance. With this inheritance pattern it is not possible to explain the results
obtained in the present studies as one-sixteenth of the F., population was fully
susceptible. Crosses between Chevron and susceptible varieties have been currently
made to confirm the hypothesis that a single incompletely dominant factor governs
resistance in the variety Chevron to Australian races.

Two additional crosses involving Triple Bearded Lemma with Psaknon and Duplex
were studied as both offered material for linkage studies, since the literature review
indicated that these latter varieties have been reported to possess genes for resistance
linked with the Algerian gene. The variety Duplex was shown, as previously reported
in these studies, to possess two apparently independent genes against race 3, one
dominant and the other recessive.

The results from F. tests of the cross Triple Bearded Lemma with Duplex agree
with a 48:13:3 ratio assuming segregation of three genes, two dominant and one
recessive. The two genes of Duplex appear, however, to be independent of the Algerian
gene (Ml.) of Triple Bearded Lemma, although the 353 F, population is clearly
insufficient to test independent segregation of three or more genes.

There was strong evidence of linkage of genes for resistance in the cross Triple
Bearded Lemma x Psaknon. The F. results of this cross are presented in Table 12.
Only 28 susceptible plants were found in a total of 946, indicating linkage of the
factors for resistance in these varieties. With independence 44 plants would have been
expected, on the basis of 1 fully susceptible plant in 5:43 expected for the segregation
of Triple Bearded Lemma alone.

No. 22 B.69 was crossed with certain resistant varieties and in Table 13 are
presented the results of such F, studies. The reactions of 242 F, seedlings of the cross
No. 22 x Portugese to race 3 indicated linkage between the genes for resistance in
these two varieties. Previously the behaviour in F, and F, generations of a cross
between Portugese and the susceptible variety No. 49, when tested with race 3,
indicated that a single incompletely dominant gene controlled resistance in Portugese.

TABLE 13.

Seedling Reaction to Race 3 of Mildew of F, Hybrids of Crosses between No. 22 (B.69), Reaction Type “‘ 122”’, and Two
Resistant Varieties.

F, Reaction. | x2 |
Second Parent and Reaction. Total. 1 GBS 8 3), P-value.
, | Resistant. | Susceptible.
| — —— —
Portugese B.155, “1-2” ae ye 237 5 242 7-230 0-01-0-001
Psaknon B.81, “12” .. 50 | 66 1 67 2-588 0:20-0:10

With independence of the genes for resistance in Portugese and No. 22, 15 susceptible
F, plants would have been expected in a cross between these varieties. As only five
were observed, linkage with approximately 30 per cent. crossing over between the
genes fits the observced results.

Although statistical tests do not preclude independence of the genes for resistance
in No. 22 and Psaknon, the number of F., plants was small and it will be observed
that only one plant in the F, population of 67 was susceptible. As previous results

BY N. Hi. LUIG, K. S. MCWHIRTER AND E. P. BAKER. 355
in these studies indicate that the factor in No. 22 is allelic with the Algerian factor
(M1.) in Triple Bearded Lemma, linkage of the genes in No. 22 and Psaknon would
be expected as the Algerian factor (Ml.) has been shown in this report to be linked
with the Psaknon (Ml)) gene.

(c) Linkage Studies of Genes for Mildew Resistance in Relation to Other
Characters.
Table 14 presents a summary from the study of F,; lines to investigate linkage of
the Ml. gene in the cross (Smooth Awn x Manchuria) x Triple Bearded Lemma. This
table shows that the gene for resistance to race 40 of leaf rust, Puccinia hordei Otth

TABLE 14.
Summary of Inheritance of Five Factor Pairs in the Cross Smooth Awn x Manchuria B.36 x Triple
Bearded Lemma B.278.
(1) Inheritance of Resistance v. Susceptibility to Rust (Pa,pa,) and Resistance v. Susceptibility to
Mildew (Mlamlg).

Reaction to Rust.
Reaction to Mildew. Total.
Resistant. Segregating. Susceptible.
Resistant 36 60 22 118
Segregating 53 112 55 220
Susceptible 23 41 21 85
Total 112 213 98 425

x? (from a contingency table) for independence of genes Pa, and Ml, =2-752; P (4 d.f.)=0-70-0-50.

(II) Inheritance of Black v. White Lemma (Bb) and Resistance v. Susceptibility to Mildew (Ml,mlgq).

Reaction to Mildew.
Lemma Colour. —— Total.
Resistant. Segregating. Susceptible.
Homozygous black 31 49 21 101
Segregating 58 115 42 215
Homozygous white 29 56 PP 107
Total 118 220 85 423

x? (from a contingency table) for independence of genes B and Ml, =0-780; P (4 d.f.)=0-95-0-90.

(III) Inheritance of Rough v. Smooth Awn (Rr) and Resistance v. Susceptibility to Mildew (Mlgml,).

Reaction to Mildew.
Awn Roughness. Total.
Resistant. Segregating. Susceptible.
Rough awn 107 203 77 387
Smooth awn 11 17 8 36
Total 118 220 85 423

x? (from a contingency table) for independence of genes R and Mlz=0-447; P (2 d.f.)=0-80.

(IV) Inheritance of Normal v. Triple Awn (Trtr) and Resistance v. Susceptibility to Mildew (Ml,mlg).

Reaction to Mildew.
Awn Character. Total.
Resistant. Segregating. Susceptible.
Single awn 94 183, 70 347
Triple awn 24 37 15 76
Total 118 220 85 423

x? (from a contingency table) for independence of genes Tr and Ml,z=0-655; P (2 d.f.)=0-80-0-70.

356 INHERITANCE OF RESISTANCE TO POWDERY MILDEW IN BARLEY,

(Pa,) and the Ml, gene were inherited independently of each other. Independence of
Ml, was indicated in relation to the factor pairs for lemma colour (Bb— group II),
normal versus triple awn (Trtr— group I), rovgh versus smooth awn (Rr — group V).
Independence would be expected, except for the Bb factor pair in group II in which
the Ml. gene has previously been placed. However, these results confirm those of
previous workers, who have not shown linkage between the two gene pairs.

The possibility of linkage was also studied between resistance to race 14 of leaf
rust in variety No. 49 and resistance to race 3 of mildew in the varieties Purple
Nudum and Triple Bearded Lemma. The data are set out in Table 15 for segregation
of these two diseases. FF, seedlings were tested first with rust and then inoculated
with mildew. The statistical analysis presented shows that mildew and rust inheritance
are not associated and the genes concerned must be considered independent.

A further study was to determine if linkage existed between resistance to mildew
and resistance to rye stem rust in the variety Purple Nudum. The genes for resistance
entered the cross in the coupling phase in crosses between Purple Nudum and seven

TABLE 15.
Inheritance of Resistance in F, to Race 14 of P. hordei in Relation to Inheritance of Resistance to Race 3 of H. gr. hordei
in Two Crosses involving the Rust-resistant Variety No. 49 (B.62) with the Mildew-resistant Varieties Purple Nudum (B.28)
and Triple Bearded Lemma (B.278).

F, Reaction to Reaction to Rust.
Cross. Plant Mildew. Total.
No. Resistant. |Intermediate.| Susceptible.

all Resistant 7 26 61 39 126

Susceptible .. 17 17 13 47

Total we 43 78 52 173

II 54.19 52, Resistant 2 42 84 36 162

(No. 49x Triple Bearded Susceptible .. 11 26 20 57
Lemma). |

Total fe 53 110 56 219

3 Resistant Bc 35 74 29 138

Susceptible’ .. 10 19 11 40

Total me 45 93 40 178

Grand total (for rust) os FS oie & 141 281 148 570
x* for a 1:2:1 ratio (rust): 0:284; P=0-90-0-80.

Grand total (for mildew): 426 resistant; 144 susceptible.

x%* for a 3:1 ratio (mildew): 0-021; P=0-90-0-80.

oll Resistant Ha 35 95 51 181

Susceptible .. 24 30 14 68

II 54.3 Total Bs 59 125 65 249

(Purple Nudum x No. 49).

2 Resistant oe 24 48 28 100

Susceptible .. 7 17 6 30

Total mis 31 65 34 130

Grand total (for rust) ys 2 abn te 90 190 99 379

x* for a 1:2:1 ratio (rust): 0°430; P=0-90-0:80.
Grand total (for mildew): 281 resistant; 98 susceptible.
x” for a 3:1 ratio (mildew): 0:149; P=0-70.
Cross II 54.19: x? for independence (mildew and rust : from contingency table) = 3-674; P-value (2 d.f.)=
0-20-0-10.
Cross II 54.8 : x? for independence (mildew and rust: from contingency table) = 5-774; P-value (2 d.f.)=
0:10-0:05.

BY N. H. LUIG, K. S. MCWHIRTER AND E. P. BAKER. 357

susceptible varieties. 481 F, seedlings were inoculated with race “Sec. I” of P. gr.
secalis and after notes had been taken the plants were dusted with conidia of race 3
of #. gr. hordei. The data, set out in Table 16, failed to indicate linkage between
resistance to rye stem rust and resistance to powdery mildew. The plants intermediate
in their reaction to mildew were grouped with the resistant class.

TABLE 16.
Associated Behaviour of F. Hybrids of Crosses between the Variety Purple Nudum and Seven Susceptible Varieties to
Culture “‘ Sec. 1”’ of PB. gr. secalis and to Race 3 of HK. gr. hordei.

|
Reaction to Race 3. va
Reaction to “Sec. 1”. Total. (3:1). P-value.
Resistant. Susceptible. :
No reaction .. ae ae 4 2 6 0-222 0-70-0-50
Resistant $3 oe ae 261 74 335 1:513 0-30-0-20
Intermediate .. ey Fas 37 10 } 47 0-347 0:70-0:50
Susceptible are =o a 70 23 93 0-005 0-95-0-90
Total ae M. 372 109 481 1-403 0-30-0-20

x* for independence of inheritance to mildew and rust by contingency method=0-731; P (3 d.f.)=0-90-0-80.

Linkage was also not found between the Ml., gene in Purple Nudum and the three
morphological factor pairs purple versus white lemma (Pp), hulled versus naked
(Nn) and hooded versus awned (Kk) involving linkage groups {f, III and IV, respec-
tively (Table 17). The Mla; gene in No. 22 was also independent of rough versus
smooth awn, and the Psaknon factor (Ml,) in the variety Psaknon was not associated
with the factor pair hulled versus naked in inheritance (Table 17).

TABLE 17.
Linkage between Factors for Mildew Resistance and Morphological Factor Pairs.
(1) Linkage between Ml,, in Purple Nudum and Three Morphological Characters in Known Linkage Groups.

Linkage Mildew Segregation of Factor Pair. P-value
Group. Factor Pair. Segregation. x. (2 d.f.).
ib p. Total.
I Purple v. white | Resistant .. 26 16 42
lemma. Segregating 63 24 87
Susceptible 34 12 46
123 52 175 1-892 0-50-0-30
Linkage Mildew Segregation of Factor Pair. P-value
Group. Factor Pair. Segregation. x? (2 d.f.).
N. n. Total.
III. Hulled v. naked .. | Resistant .. 31 11 42
Segregating 64 19 83
Susceptible 27 18 45
122 48 170 4-329 0-20-0-10
Linkage Mildew Segregation of Factor Pair. P-value
Group. Factor Pair. Segregation. Pawn x*. (2 d.f.).
K. k. Total.
IV Hooded v. awned Resistant .. 36 11 47
Segregating 59 29 88
Susceptible 31 13 44
126 53 179 1-341 0-70-0-50

358 INHERITANCE OF RESISTANCE TO POWDERY MILDEW IN BARLEY,

TABLE 17.—Continued.
Linkage between Factors for Mildew Resistance and Morphological Factor Pairs.

(II) Linkage between Ml,, and the Factor Pair for Awn Barbing.

Linkage Mildew Segregation of Factor Pair. P-value
Group. Factor Pair. Segregation. x* (2 d.f.).
R. if Total.
V Rough v. smooth.. | Resistant .. 20 11 31
Segregating 53 15 68
Susceptible 21 13 34
94 39 133 3°603 0:20-0:10

(III) Linkage between Mlp and the Factor Pair Hulled v. Naked.

Linkage Mildew Segregation of Factor Pair. P-value
Group. Factor Pair. Segregation. i x. (2 d.f.).
N. n. Total.
Til Hulled v. naked Resistant .. 25 10 35
Segregating 34 PAIL @ 55
Susceptible 17 8 25
76 39 115 0-935 0-70-0-50

DISCUSSION AND CONCLUSIONS.

The mildew inheritance studies presented were mainly concerned with the three
apparently previously unstudied varieties, Triple Bearded Lemma (B.278), No. 22 (B.69)
and Purple Nudum (B.28). All three varieties were found to carry a factor apparently
at the same, viz. the Algerian, locus.

In the case of crosses of Triple Bearded Lemma with susceptible varieties an
approximate 3:1 F, ratio, indicative of the action of a single dominant gene for
resistance, was obtained. However, che totals deviated significantly from this segrega-
tion in several cases. With a ratio of 3:49:1 (resistant.susceptible) no statistically
significant deviations occurred. The F. segregation was further complicated by the
presence of a small proportion of intermediate reaction type plants.

The intermediate group has been suggested as being due to the behaviour of
modifying genes interacting with either a single dominant gene or with a second gene
linked with the major and dominant epistatic factor.

If a single factor pair alone is involved, then an unfavourable differential
transmission rate of the gamete containing the recessive mildew allele, or unfavourable
survival value in the case of the zygote carrying the homozygous recessive genotype,
would explain the aberrant ratio. Several previous investigators have proposed similar
explanations to account for aberrant ratios. For example, Hor (1924) suggested
differential viability as explaining a highly significant deficiency of awned segregates
in a cross which he studied; Smith (1956), in a study of inheritance of seed-coat colour
in cowpeas, stated that competitive elimination of homozygous recessives did not
explain the results of genotype analysis of F., plants. Instead, a hypothesis of unequal
efficiency of gametes containing dominant and recessive alleles was proposed. In the
case of the H:h pair the suggested ratio was 0:5407:0-4593. In the present investiga-
tions a transmission ratio of 0:528:0-472 would account for a 3:-49:1 F, phenotypic ratio.
Smith (1951) stated that smooth-awned varieties frequently exhibited a higher degree
of sterility than rough-awned varieties, presumably due to the association between
awn-barbing and feathering on stigmas. Although one of the susceptible parents in the
present studies was smooth-awned and this segregation deviated most from a 3:1
ratio, a connection between awn-barbing and mildew behaviour is difficult to formulate.
It would imply linkage between the factor pairs for awn-barbing (chromosome V) and
mildew reaction. The Algerian gene for mildew resistance has previously been located
in the second linkage group.

BY WN. H. LUIG, K. S. MCWHIRTER AND E. P. BAKER. 359

If two gene pairs are involved in conditioning the reaction of Triple Bearded
Lemma, F, behaviour immediately precludes two independent genes with one being
recessive. With two linked genes, the explanations advanced considered two closely
linked genes either both dominant or one incompletely so. In the latter case, action
of modifiers would affect the class heterozygous for the incompletely dominant gene.
No statistically satisfactory linkage value could be calculated to satisfy the ratio unless
the action of modifiers on this heterozygous group was postulated.

The action of modifiers would not be necessary to explain the observed results if
the two linked gene hypothesis is considered in conjunction with a differential gametic
transmission ratio. The second gene would then account for the intermediate class.

F, data were of no value in deciding between the different alternative hypotheses.
With assigned values for the differential transmission rate and possible linkages from
F, and F, data, all calculated values were statistically non-significant.

The variety No. 22 (B.69) on the basis of reaction type and mode of inheritance
of resistance possessed a different aliele at the Algerian locus. The mode of inheritance
was simpler than in the case of Triple Bearded Lemma and was satisfactorily explained
on the basis of a single incompletely dominant gene. The mode of inheritance appeared
to be affected by environmental conditions, presumably temperature, as at higher
temperatures the heterozygous genotype showed an increase in susceptibility. The
variety No. 22 itself did not show the same environmental effects on reaction type.

Allelic but different genes for resistance at the Algerian locus were first suggested
by Schaller and Briggs (1955) in the case of the highly resistant variety Algerian and
the semi-resistant variety Black Russian. They tested large F, and F,; populations but
did not find any susceptible segregates in a cross between the two varieties. As the
gene in Black Russian was incompletely dominant in contrast to complete dominance
of the Algerian (Ml.) gene, and was furthermore of a different phenotypic expression
(“2” type reaction) in the homozygous state, it was designated “Ml... The present
studies extend the series of alieles at the Algerian locus by a further two factors—
Mil., and Mla.

Failure to obtain susceptible segregates in crosses of Purple Nudum, Gopal and
variety No. 22 with varieties possessing the Aigerian gene indicated allelism of the
factors involved. The genes in these varieties cannot be considered identical with
the Algerian (Ml.) or the Black Russian (Ml.,.) alleles, as their phenotypic expression
in the homozygous condition is different. The reaction type of variety No. 22 was
extremely characteristic and designated “1™’’, indicating severe necrosis, with limited
mycelial development. ‘The gene in variety No. 22 was therefore designated as Mas.

Purple Nudum and Gopal gave a “;™’—‘“1="”’ reaction type and resistance was
conditioned by a single incompletely dominant gene pair. The reaction type of these
varieties was thus slightly higher than the practically immune reaction type (‘‘0;’’)
conferred by the dominant Algerian gene; the genes in these two varieties, which
were similar morpholegically, appeared to be identicai, and both varieties were
considered to possess the Mla, allele.

The Algerian (Mla) allele (or a gene very closely linked to it) was found to be
operative in the varieties Triple Awned Lemma B.473, Cheroff, Monte Cristo and
Unnamed B.174. The varieties Monte Cristo and Engledow India were reported by
Favret (1949) to possess the same single dominant gene for resistance, designated
Min. The present studies indicate that this gene is the Algerian factor; and with the
four varieties Algerian, Multan, India and S.P.1.45492 (reported by Briggs and
co-workers) the number of varieties known to possess the Ml, gene is now ten. All
these varieties were characterized by the same highly resistant reaction type (“0;”)
to both Australian races of mildew. The higher resistance of the Ml, allele was
epistatic to the higher reaction types.

The size of F, and F, plant populations in practice make it extremely difficult to
discriminate between allelism versus even moderately loose linkage. This fact has
been statistically considered by Mayo (1956). Estimations are based on linkage in

360 INHERITANCE OF RESISTANCE TO POWDERY MILDEW IN BARLEY,

the repulsion phase, with two dominant genes; the probability of getting no susceptible

2

F, plants if n are grown is (1 — —)", where p is the recombination value. At the

2

0:05 probability level, 0:05 = (1 — —)" and p can be calculated for different values of n.

For example if populations of 300 and 2,000 are considered, then p = 0-1980 and 0-0748
respectively. Thus the values of 19-8 and 7-48 are the maximum distances respectively
that the two genes are separated on the chromosome in each case, with the probability
that a misleading result will not occur more often than once in twenty times. Hence
it becomes impossible to establish allelism for two genes except when very large
populations, possible in microbial genetical experiments, can be grown. For example,
to establish linkage of 1 per cent. or less between two dominant genes, an F. population
of approximately 130,000 would need to be analysed at the 0-05 probability level. Thus
in Table 10 the upper limit of recombination at the 0-05 level for the separate crosses
is expressed as a measure of the maximum distance apart of the relevant genes in lieu
of allelism.

No established gene fer mildew resistance other than Ml, has been shown to
condition a practically immune (‘“‘0;”) reaction type and show complete dominance in
inheritance. Hence this is further evidence that the previously listed ten varieties
all carry the Ml, allele. In instances where one gene is incempletely dominant and
this reaction type can be readily distinguished from either parental one, the probability
of detecting recombinants is considerably increased. Such heterozygotes would occur

with an expected frequency of (1—p)-. Thus with 10 per cent. recombination between
2

the two genes, 4:5 per cent. of heterozygotes would be expected. This is a much greater
probability than for full susceptibility where only one plant in 400 is expected. Hence
the evidence for allelism between the incompletely dominant genes, Mla, in variety
No. 22 and Ml,, in Purple Nudum and Gopal, in crosses with varieties carrying the
Algerian gene when ho intermediate reaction type plants were detected, is statistically
on a sounder basis. If the varieties Triple Bearded Lemma, Triple Awned Lemma,
Algerian, Unnamed and Monte Cristo are all assumed to have the Algerian (Mla) gene
then a total of 2156 F, plants from crosses between any of these five varieties with
Purple Nudum (or Gopal) with the Ml., locus were studied. The upper limit of
recombination between the Ml, and Ml,, genes from this combined data at the 0-05

D 1
level is 0-28 per cent. according to the formula 0:05 = (1— (~—~—)?)", where one
my

gene is completely, and the other incompletely, dominant. The combined estimate from
F, and F, data of the upper limit of recombination between the Ml, gene in Triple
Bearded Lemma and the Ml., gene in No. 22 (B.69) at the 0-05 level is 1:10 per cent.

Fewer plants are required by backerossing to measure linkage with the same
accuracy when compared with F. methods. However, with small grain cereals it is
difficult technically to produce large numbers of backcrossed seeds. Hence estimates
must be based on F, behaviour.

In crosses between Triple Bearded Lemma and the resistant varieties Goldfoil,
Hanna, Moore 11.76, Chevron and Psaknon, the ratios obtained agreed statistically
with a two major factor segregation. A single factor difference in Goldfoil and Psaknon
was confirmed in their crosses with susceptible varieties. The Psaknon factor (Ml)
was found to be linked with the Algerian gene in Triple Bearded Lemma. If Triple
Bearded Lemma is considered to possess only a single factor for resistance, the
estimated recombination value between Ml, and Ml, is 34:81 + 6-17 per cent. However,
as inheritance of resistance in Triple Bearded Lemma in crosses with susceptible
varieties could not be explained on conventional single factor segregation, this value

BY N. H. LUIG, K. S. MCWHIRTER AND E. P. BAKER. 361

could not serve as an exact estimate of recombination. Nevertheless, Schaller and
Briggs (1955) presented data for linkage of Ml. and Ml, genes which are in close
agreement with the present results.

From its behaviour in a cross with a susceptible variety, Duplex (B.172) was
postulated to possess two factors, one dominant and probably identical with the Hanna
factor, and the other recessive. As indicated previously, the third reported gene in
Duplex (mla) is ineffective against the two Australian races. In a cross with Triple
Bearded Lemma the two genes in Duplex appeared to be independent of the Ml. gene.
Stanford and Briggs (1940) found in Arlington Awnless, Psaknon and Duplex the same
gene, namely Ml. It is not possible to correlate the results of the present study
with this hypothesis. Arlington Awnless is susceptible to Australian races in the
seedling stage and has only adult plant resistance. The incompletely dominant gene
in Duplex confers also a much higher type of resistance than the Psaknon gene (Ml).
However, it must be remembered that, whilst Stanford and Briggs (1940) also used
race 3, similar race designations do not necessarily mean that such races possess
identical factors for pathogenicity.

In certain respects the present findings may be compared with those of Favret in
Argentine. In this connection it is suggested that to Australian races employed the
variety Gopal possesses a single gene for resistance, Ml.,, allelic with the Algerian
factor, and not two linked genes as reported by Favret (1949) to his pathogenic types.
Furthermore, the variety Monte Cristo, seed of which was received from Argentine,
did not produce susceptible segregates in crosses with varieties carrying a gene for
resistance at the Algerian locus. The phenotypic expression of resistance indicated
that it possessed the Ml, gene.

No linkage was found either between the genes Ml, in Psaknon and Ml., in
Purple Nudum in relation to five morphological factor pairs involving linkage groups
I, Ill, IV and V. No association was found between the Ml, locus in Triple Bearded
Lemma and three morphological factor pairs involving linkage groups I, II and V,
nor between the two genes for leaf rust resistance and the Algerian locus. No
correlation existed between resistance to mildew and resistance to rye stem rust in
the coupling phase in crosses of Purple Nudum with susceptible varieties.

As indicated in the literature review, four genes for mildew resistance (Mlx, Mla,
mla and Ml,) are claimed to be located in linkage group II. Linkage between Mla
and Ml, was confirmed in the present investigations, but more precise placing of the
loci for mildew resistance is needed both from conventional linkage studies with
genetic markers and cytogenetical studies involving translocations. The use of trans-
locations in relation to semi-sterility and mildew reaction is being conducted with
this objective. These studies herein reported suggest that at least five alleles at one
of the loci in linkage group II niay condition mildew reaction. This is somewhat
unique for disease resistance genes, but a parallel case appears to exist from Flor’s
investigations on factors conditioning reaction to flax rust.

Literature Cited.

Briees, F. N., 1935.—Inheritance of resistance to mildew, Hrysiphe graminis hordei, in a cross

between Hanna and Atlas barley. J. Agr. Res., 51: 245-250.

, 1938.—Inheritance of resistance to mildew. Am. Nat., 72: 34-41.

, 1945—lLinkage relations of factors for resistance to mildew in barley. Genetics,
30: 115-118.

, and Barry, G. D., 1938.—Inheritance of resistance to mildew, Erysiphe graminis
hordei, in a cross of Goldfoil by Atlas barley. Zeit. Zucht. A., 22: 75-80.

, and STANFORD, E. H., 1938.—Linkage of factors for resistance to mildew in barley.
Jour. Genet., 37: 107-117.

CHEREWICK, W. J., 1944.—Studies on the biology of Hrysiphe graminis D.C. Canadian J. Res.,
22: 52-86.

Dietz, S. M., and MurpHy, H. G., 1930.—Inheritance of resistance to Hrysiphe graminis hordei.
IV (Abst.). Phytopath., 20: 119-120.

362 INHERITANCE OF RESISTANCE TO POWDERY MILDEW IN BARLEY.

FAvrRET, EH. A., 1949.—Herencia de la resistancia a Hrysiphe graminis hordei en cebada. Rev.
de Inv. Agr., 3: 31-42 (Spanish).

HeHN, EH. R., 1948.—Inheritance of agronomic characters in barley. Thesis (Ph.D.), Iowa
State Coll., 63 pp. (typed). (Cited by Smith, 1951).

HOoNECKER, L., 1931.—Notes on the mildew problem in barley with particular reference to the
aspect of breeding. Pflanzenbau, Pflanzenschutz wu. Pflanzenzucht, 8: 78-84, 89-106.
(P.B.A., 2: 442.)

, 1934.—On the modification of the infection and the occurrence of various physiologic
forms of barley mildew (Hrysiphe graminis hordei Marchal). Zeit Zucht. A., 19: 577-602.
(P.B.A., 9: 369.)

Hor, K. S., 1924.—Interrelations of genetic factors in barley. Genetics, 9: 151-180.

HUNTLEY, D. N., 1951.—Hrysiphe graminis in barley. II. Mode of inheritance of resistance.
Iowa State Coll. J. Sci., 25: 252-253.

Mains, E. B., and Dietz, S. M., 1930.—Physiologic forms of barley mildew, Hrysiphe graminis
hordei Marchal. Phytopath., 20: 229-239.

Mains, E. B., and MARTINI, M. L., 1932.—Susceptibility of barley to leaf rust (P. anomala)
and to powdery mildew (EHrysiphe graminis hordei). U.S. Dept. Ag., Techn. Bull. No. 295,
33 Dp.

MARTINI, M. L., and HARLAN, H. V., 1942.—Barley freaks. J. Hered., $3: 338-343.

Mayo, G. M. E., 1956.—Linkage in Linum usitatissimum and in Melampsora lini between the
genes controlling host-pathogen reactions. Aust. J. Biol. Sci., 9: 18-36.

MILuERD, A., and Scott, K., 1955.—A phytopathogenic toxin formed in barley infected with
powdery mildew. Aust. J. Sci., 18: 63-64.

MosEeMAN, J. G., 1956.—Physiological races of Hrysiphe graminis f. sp. hordei in North America.
Phytopath., 46: 318-322.

N.S.W. DEPARTMENT OF AGRICULTURE, 1935.—Plant diseases recorded in New South Wales.
Science Bull., No. 46.

N.S.W. DEPARTMENT OF AGRICULTURB, Biological Branch, Division of Science Service, 1951.—
Plant disease survey. Twenty-first Annual Report.

NEWTON, MARGARET, and CHEREWICK, W. J., 1947.—Hrysiphe graminis in Canada. Canad. J-
Res., ©. 25: 73-93.

ScHALLER, C. W., and Brices, F. N., 1955.—Inheritance of resistance to mildew, E. gr. hordei,
in the barley variety Black Russian. Genetics, 40: 421-428.

SARASOLA, T. A., FAVRET, H. A., and VALLEGA, J., 1946.—Reaccion de algunas cebadas con
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(Spanish with English summary.)

SmirH, F. L., 1956.—Inheritance of three seed-coat colour genes in Vigna sinensis savi.
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STANFORD, H. H., and Brices, F. N., 1940.—Two additional factors for resistance to mildew in
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WATERHOUSE, W. L., 1948.—Studies in the inheritance of resistance to rust of barley, Part II.
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363

SPORES AND POLLENS FROM A PERMIAN-TRIASSIC TRANSITION, N.S.W.
By J. P. F. HENNELLY, Coal Research Section, Commonwealth Scientific and
Industrial Research Organization.

(Plates v—vi; two Text-figures.)
[Read 26th November, 1958.]

Synopsis.

An account is given of an investigation of the microspore and megaspore contents of
some sections of strata above the Bulli seam, encountered during the drilling of a bore at
Appin. One new genus and six new species were encountered and taxonomic dscriptions
of these are given, with preliminary notes on three other microspore and megaspore types.

INTRODUCTION.

The material examined in this study of spores and pollens comprised the roof
shales of the Bulli seam and 87 feet of the overlying Lower Narrabeen sediments. It
was taken from Appin Bore No. 4, put down by Australian Iron and Steel Limited. An
indication of the sampling depths and of the quality of spore preservation is given
in Text-figure 1.

RETICULATUS FIBULATUS. BULLIENSIS HORRIDUS
P. NIGRACRISTATUS NUSKOISPORITES | GRANULATISPORITES

PAL.SER.

DEPTH BELOW NUMBER OEPTH

SURFACE |

RADIATUS. sP."A"

1919

1920

1916

1915

ele ees ea Te te Shae lay aie eral eioic lola a aloha este iene

IN Ulli Seam
I GOOD = 100 SPECIMENS POOR = € 100 SPECIMENS 2 Oo 50 100%
VERTICAL SCALE APPROXIMATE ONLY AND VARIABLE HORIZONTAL SCALE —=

Text-fig. 1—Sampling depths and the quality of spore preservation.
Text-fig. 2.—Distribution of sporomorphs in Appin Bore No. 4.

PROCEEDINGS OF THE LINNEAN Socrery or New SoutH WALES, 1958, Vol. Ixxxiii, Part 3.

364 SPORES AND POLLENS FROM A PERMIAN-—TRIASSIC TRANSITION,

Examination of the megaspore and microspore content showed that a transition
zone extended from 2 in. to 15 in. above the Bulli seam. This zone was characterized
by a dominant Apiculatisporites species and an easily recognized cristate tetrad, in
association with other types which were undoubtedly of Triassic age. The presence
of numerous small megaspores further indicated a Triassic rather than a Permian age.
In the sediments more than 15 in. above the Bulli seam only the obviously Triassic
sporomorphs occurred (Text-fig. 2).

It has long been recognized that the macroscopic fossil record indicates the
presence of a transition zone overlying the Bulli seam (EHtheridge, 1892; Dun, 1901,
1910). Walkom (1925) considers this flora to be more closely related to the Glossopteris
than to the Thinnfeldia. Though not identical with Apiculatisporites species found in
the various Permian floras, Apiculatisporites bulliensis, new species, shows some super-
ficial resemblance to Apiculatisporites cornutus of the Greta Coal Measures. It would
be of great interest if fertile strobili of Schizoneura gondwanensis were available for
comparison with both the above-mentioned common transition bed sporomorphs.

The microspore assemblage in the flora of the roof shales of the Bulli seam is
much less varied than that of the Thinnfeldia flora of the Middle and Upper Narrabeen
Formations. Of the samples examined, few contained well-preserved spores, and in
even the best of these the specimens were not numerous. MHolotypes and paratypes of

the new species are preserved in the palynological herbarium of the Coal Research
Section.

TAXONOMIC DESCRIPTIONS.
(a) SPoRITES.

Division SPORITES H. Potonié, 1893.
Subdivision TRILETES Reinsch, 1883.

Genus APICULATISPORITES (Ibrahim. non Bennie and Kidston) Potonié and Kremp.
APICULATISPORITES BULLIENSIS, n. sp. (PI. v, figs. 3-5.)

Amb circular or slightly oval. Trilete, laesurae indistinct, extending almost full
radius. Exine 2—4u, opaque. Ornamented with blunt conical processes 1u or less in
diameter, 1-13u in length, and 2u apart. Dimensions (27 specimens): Diameter 20—40u
(mean 29).

Probably owing to its small size and comparatively thick exine, this sporomorph
is particularly resistant to corrosion and forms a useful indicator of the sediments of
the Bulli transition zone. It somewhat resembles A. cornutus (Balme and Hennelly,
1956), of the Greta Coal Measures, except for its slightly smaller size, smaller conical
processes, and wider spacing between spine bases. It is much more prolific in the
Bulli transition zone than A. cornutus has been found to be in any of the Permian
sediments examined by the author.

Type Locality: Appin Bore No. 4 from 1696 ft. 10 in. to 1697 ft. 5 in.

Genus QUADRISPORITES, Nn. gen.

The name Quadrisporites is proposed as a form genus for persistent tetragonal or
hexagonal tetrads of microspores of vascular plant origin but otherwise of unknown
affinities, and whose dehiscence mechanism is either alete or so obscured as to be
indeterminate.

Genotype, Quadrisporites horridus, 0. sp.

The morphology of the tetrad suggests the monolete condition, but the cohesion
of the tetrad even under adverse conditions of preservation indicates that it must be
at least functionally alete. This unusual characteristic results in isolated spores being
in a very small minority. Such spores are so poorly preserved that ruptures in the
exine may not be related to dehiscence. As the type normally encountered is the
tetrad it is preferable to classify it as such, rather than to attempt correlation
with individual form genera based on dehiscence and ornamentation (Potonié and
Kremp, 1954).

>?

BY J. P. F. HENNELLY. 365

The term Tetrasporites has been applied to spores of presumed Ulvacean affinity
(Fliche, 1886).

QUADRISPORITES HORRIDUS, nN. Sp. (PI. v, figs. 6-7.)

Tetragonal tetrads of spores approximately spherical and of similar appearance
and dimensions. Exine luz in thickness but opaque owing to heavy ornamentation.
Examination of detached individuals does not disclose the type of dehiscence apparatus.
Probably functionally alete. Ornamentation, a confused system of filiform cristae.
Processes lu in diameter and 2-34 in length superimposed on an interrupted micro-
reticulum. Dimensions (20 individual spores, one each from 20 tetrads): Diameter
23-374 (mean 28).

This sporomorph is very useful as a means of identifying the Bulli transition zone
in the Appin Bore. The tetrad formation is persistent even in corroded material, and

TAPLE 1.
Palynological summary: Lowest Triassic overlying the Bulli seam (Permian),
showing the transition zone and sediments to 75 feet above in green and grey
shales (Lower Narrabeen Formation).

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Type of sediment: 1917-1888, Clastic sediments, mainly green-grey shales; 1889,
Bulli seam S.S.A.; 1890, Bulli seam minus dirt; 1891, Bulli seam plus dirt.

366 SPORES AND POLLENS FROM A PERMIAN—TRIASSIC TRANSITION,

quite characteristic in appearance. In detached corroded specimens the remains of
the interrupted reticulum forming the cristate system resemble corroded specimens
of Apiculatisporites bulliensis and Granulatisporites sp. “A’’.

It is recommended that only coherent tetrads be recorded in palynological
examinations. In the palynological summary (Table 1) each tetrad has been recorded
as a Separate entity.

Type Locality—Appin Bore No. 4 at 1697 ft. 2 in.

Subdivision ZONALES Bennie and Kidston emend. Potonié and Kremp.
Genus CIRRATRIRADITES (Wilson and Coe) Potonié and Kremp.
CIRRATRIRADITES FIBULATUS, nN. sp. (PI. v, figs. 8—9.)

Spore complex, consisting of a central body with an equatorially attached trans-
parent membranous wing. Amb of central body circular or rounded triangular, wing
approximately concentric with body. Trilete sutures fairly straight, extending to
apices of wing. Lips lw. Ornament, bedy and wing granulate with grana about lu
and closely spaced, and microreticulate with muri lu and lumens of 4u over the body.
This reticulation extends also to the outer wing where the lumens are reduced to
about 24 and are scarcely discernible. Wing overlap 1u or more, and often difficult
to detect. Dimensions (50 specimens): 30-354 overall diameter (mean 40), body
diameter 22-474 (mean 29y).

Superimposition of muri on grana gives an appearance of grana larger than 1p.

A somewhat similar but psilate-winged and slightly smaller-dimensioned Cirratri-
radites sp. is recorded by de Jersey (1949) as type T32C, from the Triassic of Ipswich,
Queensland.

Type Locality—Appin Bore No. 4 from 1697 ft. 24 in. to 1697 ft. 9 in.

(60) Division POLLENITES.
Subdivision SACCITES Erdtman.
Genus NUSKOISPORITES Potonié and Klaus.
NUSKOISPORITES RADIATUS, nN. sp. (PI. v, figs. 10-12.)

Synonymy: Type 34C Taylor, 1953.

Amb, of both central body and bladder, circular. Trilete sutures indistinct.
Dehiscence, sometimes by loss of the whole proximal face in the manner of an
operculum, otherwise a triangular rent formed by the proximal face rolling back
from the fully opened sutures, which extend the full radius of the body. Exine of the
central body translucent and about ilu in thickness; ornament very finely granulate
on the proximal face and faintly reticulate on the distal face. Grana less than lu.
Equatorial bladder is also finely granulate and fincly microreticulate with muri less
than 1 in width and lumens 1—23y, exine transparent and thinner than that of body.
Overlap 7-12u. Radial folds usually present in the bladder. Dimensions (20 specimens) :
Total diameter 70-110u (mean $1), central body 26-60u (mean 45z).

The general appearance is granulate except under high magnification, and super-
ficially resembles N. dulhuntyi Klaus (Klaus, 1953). Size range is smaller, and radial
folds are fairly characteristic. When ruptured with loss of the proximal face this
species can only be recognized by the finely granulate wing ornament.

Type Locality—Appin Bore No. 4 from 1697 ft. 0 in. to 1697 ft. 9 in.

Genus PIryOSPORITES Seward.
PITYOSPORITES NIGRACRISTATUS, nh. sp. (PI. v, figs. 13-15.)

Pollen bisaccate. Central body circular or subcircular, proximal cap unthickened
and very finely granulate, with grana less than lu. Exine about 1p in thickness.
Crests prominent, and ranging up to 7u in width. Bladders symmetrically disposed.
Furrow indistinct, dehiscence often by irregular rupture with varying body opening.
Bladder length is usually not greater than the longitudinal diameter of the central
body, and bladder width is not greater than this dimension. Biadders ornamented
with an irregular reticulum, often interrupted and sometimes appearing vermiculate.

BY J. P. F. HENNELLY. 367

The muri are less than lu and lumens vary from 2—h5u. Dimensions (20 specimens) :
Central body polar diameter 30—45u4 (mean 38). total span 45-724 (mean 61y).

P. nigracristatus differs from P. elliptica (Cookson) Balme (Cookson, 1947; Balme,
1957) in that it has more obvious wing reticulation, slightly thicker wing tissue, and
the wings do not project beyond the poles. Apart from its rather more robust
structure, P. nigracristatus differs from P. similis Balme in having larger and irregular
reticulation of wing and more prominent crests.

Type Locality.—Appin Bore No. 4 from 1696 ft. 9 in. to 1697 ft. 9 in.

PITYOSPORITES RETICULATUS, n. sp. (PI. v, fig. 16; Pl. vi, figs. 17-18.)

Pollen grain bisaccate. Central body circular or tending to oval. Ornament finely
granulate and microreticulate, with muri about lw in width and lumens 4-6y. Exine
within the lumen about iu thick and translucent to transparent. Furrow normal.
Bladders symmetrically disposed on either side of the furrow, with prominent crests
about 5u in thickness. Outline of the whole pollen is approximately oval. Bladder
width is variable but generally not as great as body diameter. Bladders externally
microreticulate with prominent muri lw in width. In proximal view the reticulum
obscures the central body. Dimensions (25 specimens): Body 52-884 (mean 63), total
span 63-1374 (mean 96z).

The wings are prone to separate from the body with the crests attached, giving
the appearance of a monolete sporomorph with thickened iips. In oxidized samples
the muri and erests are darkened and render this pollen easily identifiable even in
the fragmented state. It is the dominant of the sediments 34 ft. and 74 ft. above the
Bulli seam in the Appin Bore No. 4.

Type Locality—Appin Bore No. 4 16238 ft. to 1697 ft. 9 in.

(c) NorEs oN Two MEGASPORES AND ONE TRILETE MICROSPORE.
MEGASPORE TYPE “A’’. (PI. vi, figs. 21-22.)

Amb circular to rounded trianguiar. Trilete sutures about 35u in width, extending
almost full radius. Exine opaque, 20—45u in thickness; contact face and laesural ridge
visible in some specimens. Silhouette shows prominent muri. Ornamentation deeply
reticulate with muri 4-84 in width and 10—15y in height. Lumens 10-15u. Exine
within the lumen is rugose. In corroded specimens the muri are persistent, the exine
within the lumen becoming translucent. This megaspore is very common in the
Appin Bore Transition Zone.

Type Locality—Appin Bore No. 4 at 1697 ft. 4 in.

MEGASPORE TYPr “B”. (PI. vi, figs. 19-20.)

Amb rounded triangular, straight sutures 10u in width extending almost full
radius. Exine translucent to opaque, 10-20 in thickness. Contact area (poorly
developed) covers most of proximal surface. Ornamentation, reticulate with muri 2u
in width and lumens 5y or less. Within the lumen the exine is granulate, with
grana lp or less, 1--2u apart.

Both megaspores appear to be comparatively small, with diameters of 250—500y,
but as most of the specimens examined are incomplete it is inadvisable to attempt
to specify the size range more definitely at this stage.

Type Locality—Appin Bore No. 4 at 1696 ft. 11 in.

Genus GRANULATISPORITES (Ibrahim) Potonié and Kremp.
GRANULATISPORITES sp. “A”. (Table 2; Pl. vi, figs. 23-24.)

Amb circular, with a faintly irregular outline due to ornament. Trilete sutures
indistinct, extending almost full radius. Lips thin, lu or less. Contact area rarely
identifiable. Dehiscence often by rupture. EXxine 24 in thickness, translucent.
Ornamentation, granulate with grana 1u or less, about 2u apart. Dimensions (30
specimens): 20-744 diameter (mean 45u).

368 SPORES AND POLLENS FROM A PERMIAN-—TRIASSIC TRANSITION,

In the poorly preserved specimens obtained at Appin Bore No. 4 the corrosion of
the exine in patches presents a false reticulate appearance in high focus. The
silhouette, however, shows only grana of less than lu in height and no indication of
muri. It is possible that this sporomorph includes two varieties, the smaller 20—45y
(mean about 35u) and the larger 36-744 (mear about 54) in diameter; the difference,
apart from size, being a slightly thinner exine in the larger variety.

Type Locality—Appin Bore No. 4 1694 ft. 11 in. to 1697 ft. 9 in.

Acknowledgements.

Thanks are due to Australian Iron and Steel Limited, for making available the
sample material and all necessary bore log data; to Mr. H. R. Brown, Officer-in-Charge
of the Coal Research Section, Commonwealth Scientific and Industrial Research
Organization, for his interest and encouragement; and to Dr. G. H. Taylor, for advice
and for the loan of his specimens from the Leigh Creek deposits. Mr. C. H. Noble
was responsible for the photomicrographs.

TABLE 2.
Distribution of Appin Sporomorphs in Other Australian Triassic Sediments.
Thinnfeldia
Flora, Callide.+ Leigh Creek.t
N.S.W.*
Pityosporites reticulatus al bs — — —
P. mgracristatus ae bee Ais x — —
Cirratriradites fibulatus Me nts — — —
Nuskoisporites radiatus BH He ? — x
Apiculatisporites bulliensis oa: at — —- —
Granulatisporites sp. “A” .. bs x — —
Megaspore type ‘“* A ”’ hr ae — — | =
Megaspore type “B”’ x — —
Lueckisporites spp. x — —
Lycopodium spp. ie aye S — —
Quadrisporites horridus ye ls = aa =

* Two samples, a mudstone and an anthraxolite shale.
+ A single sample.
{ Discussions with G. H. Taylor, and a few samples of his Leigh Creek specimens.

Bibliography.

BALME, B. E., 1957.—Spores and Pollen Grains frem the Mesozoic of Western Australia.
C.8.I.R.O. Coal Research Section, Report Ref. T.C.25.

BaLMeE, B. E., and HENNELLY, J. P. F., 1956.—Trilete Sporomorphs from Australian Permian
Sediments. Aust. J. Bot., 4: 240-60.

Burees, N. A., 1935.—Additions to our Knowledge of the Flora of the Narrabeen Stage of the
Hawkesbury Series in N.S.W. Proc. LINN. Soc. N.S.W., 60: 257-64.

Cookson, I. C., 1947.—Plant Microfossils from the Lignites of Kerguelen Archipelago.
B.A.N.Z. Antarct. Res. Exped., 2, pt. 8: 129-42.

Dun, W. S., 1901.—A Specimen of Shale from the Sydney Harbour Colliery Shaft. Proc.
LINN. Soc. N.S.W., 26, Notes and Exhibits, pp. 738-9.
, 1910.—Notes on some Fossil Plants from the Roof of the Coal Seam in the Sydney
Harbour Colliery. J. Roy. Soc. N.S.W., 44: 615-19.

ETHERIDGE, R., JR., 1892.—Plant Remains from the Bulli Coal Measures. Ann. Rep. Dept. Min.
and Agric. N.S.W., 1891: 269.

FLicHr, P., 1886.—Note sur les Flores Tertiares des Environs de Mulhouse. Soc. Indus.
Mulhouse Bull., 56: 348-62.

DE JERSEY, N. J., 1949.—Principal Microspore Types in the Ipswich Coals. Univ. of Queensland,
Dept. of Geol., Papers 3, No. 9.

KuLAus, W., 1953.—Uber die Sporendiagnose des deutschen Zechsteinsalzes und des alpinen
Salzgebirges. Zeit. Deutsch. Geol. Gesell., 105: 776-88.

PoToNi£, R., and Kremp, G., 1954.—Die Gattungen der Paldozoischen Sporae Dispersae und
ihre Stratigraphie. Geol. Jahrb., 69: 111-94.

BY J. P. F. HENNELLY. 369

Taytor, G. H., 1953.—The Spore Content of the Leigh Creek Coal. S. Aust. Dept. of Mines,
Min. Rev. No. 99: 155-70.

WaLkom, A. B., 1925.—Fossil Plants from the Narrabeen Stage of the Hawkesbury Series.
Proc. LINN. Soc. N.S.W., 50: 214-24.

EXPLANATION OF PLATES V-VI.

Magnification is 500 diameters in every case except Figures 19 and 21, where it is
50 diameters.

Plate v.

3: Apiculatisporites bulliensis, showing ornament. 4: A. bulliensis, showing sutures.
5: A tetrad of A. bulliensis. 6: Quwadrisperites horridus, showing dark cruciform of contact
faces. 7: Q. horridus, ornament. 8: Cirratriradites fibulatus, amb. 9: C. fibulatus, wing and
body ornament. 10: Nuskoisporites radiatus, trilete dehiscence. 11: N. radiatus with dehiscence
by rupture, showing the proximal portion still hinged along the lower margin and opened in
the manner of an operculum. 12: N. radiatus, ornament. 13-15: Pityosporites nigracristatus.
16: P. reticulatus.

Plate vi.

17, 18: P. reticulatus. 19: Megaspore type “B” in reflected light showing contact face.
20: Megaspore type “B” fragment, showing ornament in transmitted light. 21: Megaspore
type “A” in transmitted light. 22: Megaspore type “A” fragment, showing ornament in
transmitted light. 23, 24: Granulatisporites sp. “A’’.

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371

ABSTRACT OF PROCEEDINGS

ORDINARY MONTHLY MEERTING.
26th Marcu, 1958.

Dr. S. Smith-White, President, in the chair.

The Chairman announced that library accessions amounting to 59 volumes,
463 parts or numbers, 33 bulletins, 10 reports and 13 pamphlets, total 578, had been
received since last meeting.

PAPERS READ (by title only).
1. Seed Coat Anatomy and Taxonomy in fHucalyptus. By E. Gauba and
L. D. Pryor.

2. A New Species of Aédes (Finlaya) from Northern Australia (Diptera, Culicidae).
By Elizabeth N. Marks and Ernest P. Hodgkin.

ORDINARY MONTHLY MEBRTING.
30th Aprin, 1958.

Professor F. V. Mercer, Vice-President, in the chair.

The following were elected Ordinary Members of the Society: Messrs. C. D. Blake,
B.Sc.Agr., Sydney; E. L. Jones, B.A., North Sydney; F. J. Moss, Roseville Chase,
N.S.W.; and Mrs. Margaret B. Reye, B.Sc., Brisbane, Queensland.

The Chairman announced that the Council had elected the following office-bearers
for the 1958-59 session: Vice-Presidents: Mr. S. J. Copland, Professor F. V. Mercer,
Professor J. M. Vincent and Dr. Lilian Fraser; Hon. Treasurer: Dr. A. B. Walkom;
Hon. Secretaries: Dr. W. R. Browne and Dr. A. B. Walkom.

The Chairman announced that library accessions amounting to 22 volumes, 141
parts or numbers, 15 bulletins and one report, total 179, had been received since the
last meeting.

The Chairman also announced that there will be no Ordinary Monthly Meetings
in June and August this year. The Sir William Macleay Memorial Lecture, entitled
“Timing in Human Evolution’, will be given by Professor A. A. Abbie on Thursday,
19th June, 1958, in the Main Hall, Science House, 157 Gloucester Street, Sydney, at
8 p.m. All interested are invited.

PAPERS READ.
1. Inheritance of Oil Characters in Hucalyptus. By L. D. Pryor and L. H. Bryant.
2. The Status of Nitrogen in the Hawkesbury Sandstone Soils and their Plant

Communities in the Sydney District. II. The Distribution and Circulation of Nitrogen.
By Nola J. Hannon, Linnean Macleay Fellow in Botany.

LECTURETTE.

An illustrated lecturette entitled “National Parks in New South Wales” was given
by Mr. Allen A. Strom, Honorary Secretary, National Parks Association of N.S.W.
(Central Region).

ORDINARY MONTHLY MEETING.
28th May, 1958.
Dr. §. Smith-White, President, in the chair.

Messrs. J. W. Green, B.Sc., Armidale, N.S.W., and R. K. Pengilley, Armidale, N.S.W.,
were elected Ordinary Members of the Society.

The Chairman offered congratulations to Miss Nola J. Hannon on obtaining the
degree of Ph.D. of the University of Sydney.

J

372 ABSTRACT OF PROCEEDINGS.

The Chairman announced that Library Accessions amounting to 16 volumes, 125
parts or numbers, 10 bulletins, 1 report and 61 pamphlets, total 213, had been received
since last meeting.

PAPERS READ.
1. Migration and Utilization of Reserve Substances during Flight in Aphis
craccivora Rock. By Max Casimir.
2. Widespread Natural Infection of Barberry by Puccinia graminis in Tasmania.
By I. A. Watson and N. H. Luig.
3. The Species of the Genus Hrodium L’Hér. endemic to Australia. By R. C. Carolin.
4. Pollen and Pollination in the Hupomatiaceae. By A. T. Hotchkiss.

NOTES AND EXHIBITS.

Mr. N. H. Luig exhibited a slide of a rust-infected seedling blade of wheat. In
the course of taking rust notes on wheat seedlings an F3 plant was found where one half
of the seedling leaf blade showed a resistant and the other half a susceptible reaction
to wheat stem rust, Puccinia graminis tritici, strain 21-Anz 2. In the present case the
F3 plant came from a cross between a susceptible variety and a highly resistant line
carrying 44 chromosomes; the two extra chromosomes being derived from the grass
Agropyron elongatum. The F3 line in which the unusual plant was found segregated
for resistance and susceptibility. Since it is known that resistance is associated
with the extra chromosome from Agropyron it is assumed that the plant observed is a
sectorial chimera, and during an early cell division loss of the extra chromosome
occurred. Thus one half of the leaf carries the extra chromosome and is resistant, the
other half lacks this chromosome and is susceptible.

Mr. G. H. Hardy exhibited some Platypezidae (Diptera), stating that Microsania,
the smoke flies, are world-wide in distribution but not often found in Australia. The
bush-fires of last year appear to be responsible for their appearance on his windows
at Katoomba, N.S.W., and more recently they occurred in the smoke of burning blackberry
near-by. The literature on these flies is very extensive but many mysterious features
in their habits still need resolving. One is the extraordinary numbers of mites carried
beneath the abdomen which seems specially designed for that purpose, the venter having
a flange-shape that accommodates the pearly pink mites on the specimens exhibited.
They are minute flies. A mite-infested specimen mounted on a micro-slide was exhibited
with mites in sitw and also some pinned specimens without mites.

Mr. A. J. Bearup demonstrated the life cycle of Acanthoparyphium spinulosum
S. J. Johnston. While examining the Gastropods of Narrabeen Lagoon for trematode
parasites, it was noted that about 10% of Salinator fragilis had cysts containing an
Hchinostome metacercaria with 23 collar spines. This number agreed with those of a
cercaria which occurred in 8% of Pyrazus australis, living in the same habitat. Under
experimental conditions nearly every Salinator became infected with several cysts. The
cercariae also penetrated into larval polychaetes and encysted in the body cavity. When
the matured cysts were fed to a young seagull a pure infection with A. spinulosum was
obtained.

Dr. Smith-White exhibited specimens of Astroloma conostephioides and of Astroloma
pinifolium from the Grampians, western Victoria. A. conostephioides has a range
extending from central Victoria to the west coast of South Australia. Throughout
this range, the great majority of plants are heterozygous for chromosome inversions,
and there must be heterotic value in the inversion heterozygosity. The association
of inversion hybridity with the maturation of pollen in tetrads is of interest. The
specimens of A. pinifolium from the Grampians were compared with specimens of the
same species from La Perouse, near Sydney. The specimens from the two regions show
rather small but quite distinctive morphological differences. In the mature flowers of
the Hast Coastal form, the stigma is entirely enclosed, and the species is probably
inbreeding. In the Grampians form, mature flowers have an excerted and exposed
stigma, and are possibly outbreeding. The morphological differences of the species in

ABSTRACT OF PROCEEDINGS. 373

the two regions, and the possible difference in breeding systems, are associated with
differences in meiotic behaviour which are being studied.

Dr. D. F. McMichael exhibited specimens of an unidentified freshwater animal
which he had collected in New Guinea. Its body resembled that of a freshwater
gastropod, but it lacked a shell. A muscular visceral mass is situated on the dorsal
surface of the body. Investigation of its anatomy is proceeding.

ORDINARY MONTHLY MEETING.
30th Jury, 1958.

Dr. S. Smith-White, President, in the chair.

Mr. A. G. Lyne, B.Sc., Ph.D., C.S.I.R.O. Sheep Research Laboratory, Parramatta,
N.S.W., was elected an Ordinary Member of the Society.

The Chairman offered congratulations to Dr. J. M. Thomson on obtaining the
degree of Doctor of Science of the University of Western Australia.

The Chairman announced that Library Accessions amounting to 22 volumes, 271

parts or numbers, 30 bulletins, 6 reports and 21 pamphlets, total 350, had been received
since last meeting.

PAPERS READ.

1. Catalogue of Australian Mammals and their Recorded Internal Parasites. Parts
1-4. By M. Josephine Mackerras. (Communicated by Dr. I. M. Mackerras.)

2. A Note on the Status of Aphodius tasmaniae Hope. By B. B. Given. (Communi-
cated by Mr. C. H. Chadwick.)

3. A New Bird-flea from Tasmania. By F. G. A. M. Smit. (Communicated by
Mr. D. J. Lee.)

4. Two New Species of Hemicycliophora (Nematoda:Tylenchida). By M. R. Sauer.
(Communicated by Mr. A. J. Bearup.)

5. A New Species of Frog of the Genus Crinia Tschudi from South-eastern Australia.
By Murray J. Littlejohn. (Communicated by Mr. 8S. J. Copland.)

6. Some More Bark- and Timber-Beetles from Australia. 158. Contribution to the
Morphology and Taxonomy of the Scolytoidea. By Karl HE. Schedl. (Communicated by
Dr. A. J. Nicholson.)

7. Studies of Nitrogen-fixing Bacteria. VII. Cytochromes of Azotobacteriaceae.
By F. J. Moss and Y. T. Tchan.

8. Somatic Hybridization in Puccinia graminis var. tritici. By I. A. Watson and
N. H. Luig.

9. Systematic Notes on some Eastern Australian Members of the Papilionaceae.
By Joy Thompson.

SYMPOSIUM.
A symposium on the “Origin and Distribution of Australian Fauna and Flora”

was held in which a leading part was taken by Professor R. L. Crocker, Dr. J. W. Evans
and Mr. Gordon Packham.

ORDINARY MONTHLY MEETING
24th SEPTEMBER, 1958.
Dr. S. Smith-White, President, in the chair.
Messrs. J. P. F. Hennelly, B.Sc., West Pennant Hills, N.S.W., and K. M. Moore,
Lisarow, N.S.W., were elected Ordinary Members of the Society.
The Chairman announced that the Council is prepared to receive applications for
Linnean Macleay Fellowships tenable for one year from ist January, 1959, from qualified

374 ABSTRACT OF PROCEEDINGS.

candidates. The range of actual (tax-free) salary is up to £800, according to qualifications.
Applications should be lodged with the Honorary Secretary not later than Wednesday,
5th November, 1958. Hach applicant must be a member of this Society and be a
graduate in Science or Agricultural Science of the University of Sydney.

The Chairman announced that Library Accessions amounting to 28 volumes,
144 parts or numbers, 4 bulletins, 6 reports and 3 pamphlets, total 185, had been received
since last meeting.

PAPERS READ.
1. Palaeozoic Geology of the Cooleman Caves District, N.S.W. By N. C. Stevens.

2. The Oviposition Behaviour of Aédes australis (Erickson) (Diptera, Culicidae).
By A. K. O’Gower.

3. Acarina from Australian Bats. By Robert Domrow.

4. A Turbidimetric Method for Hstimating the Number of Nematode Larvae in a
Suspension. By C. D. Blake.

LECTURETTE.
Miss Elizabeth C. Pope delivered a Kodachrome-illustrated lecturette on “The
Natural History of Australian HEchinoderms’”’.

ORDINARY MONTHLY MEETING.
25th October, 1958.

Professor F. V. Mercer, Vice-President, in the chair.

Mr. B. J. G. Marlow, B.Se., Australian Museum, Sydney, was elected an Ordinary
Member of the Society.

The Chairman announced that the Council is prepared to receive applications for
Linnean Macleay Fellowships tenable for one year from ist January, 1959, from
qualified candidates. The range of actual (tax-free) salary is up to £800, according to
qualifications. Applications should be lodged with the Honorary Secretary not later
than Wednesday, 5th November, 1958. Hach applicant must be a member of this
Society and be a graduate in Science or Agricultural Science of the University of
Sydney.

The Chairman referred to the death on 14th October, 1958, of Sir Douglas Mawson,
who at the time of his death was the second oldest member of the Society, having
been elected in 1905. In 1951, having had more than 40 years of membership, he was
made an honorary life member; also to the death on 28th September, 1958, of Dr. W. G.
Woolnough, who had been a member of the Society from 1899 to 1933 (with a short
hiatus) —a total of 34 years, when he resigned. Of recent years he had given material
help to the Society by gratuitously furnishing translations of communications in foreign
languages.

The Chairman announced an invitation to members to visit the Australian Atomic
Energy Commission Research Establishment, Lucas Heights, on Open Days (8th—-10th
December, 1958, inclusive), by ticket only.

The Chairman announced that Library accessions amounting to 10 volumes, 92
parts or numbers, 8 bulletins, 4 reports and 5 pamphlets, total 119, had been received
since last meeting.

PAPER READ.

Melampsora lini (Pers.) Lév. Uredospore Longevity and Germination. By Harland

B. Kerr.

Note—The paper by P. Hadlington set down for reading at this meeting was
withdrawn.

LECTURETTE.
A lecturette entitled ‘“Whales and Whaling”, illustrated by coloured transparencies,
was delivered by Mr, W. H. Dawbin.

ABSTRACT OF PROCEEDINGS. 375

ORDINARY MONTHLY MEETING.
26th NovempBrr, 1958.

Dr. S. Smith-White, President, in the chair.

The following were elected Ordinary Members of the Society: Messrs. P. W.
Hadlington, B.Se.Agr., Hurstville, N.S.W.; J. F. Rigby, B.Sc., Keiraville, N.S.W.; and
Frank Wilson, C.S.I.R.O., Canberra, A.C.T.

The Chairman offered congratulations to Miss Joyce W. Vickery on obtaining
the degree of D.Sc. of the University of Sydney.

The Chairman announced that the Council had appointed Miss Alison McCusker,
M.Sce., to a Linnean Macleay Fellowship in Botany for one year from ist January, 1959.

The Chairman announced that Library Accessions amounting to 8 volumes, 84 parts
or numbers, 1 bulletin, 3 reports and 7 pamphlets, total 103, had been received since
last meeting.

PAPERS READ.
1. The Diptera of Katoomba, Part 2. Leptidae and Dolichopodidae. By G. H. Hardy.

2. Notes on Australian Thynninae. 2. The Genera Dimorphothynnus, Rhagigaster
and Hirone. By B. B. Given. (Communicated by Dr. A. J. Nicholson.)

3. Notes on Australian Thynninae. 3. The Genus Thynnoides. By B. B. Given.
(Communicated by Dr. A. J. Nicholson.)

4. Mode of Inheritance of Resistance to Powdery Mildew in Barley and Evidence
for an Allelic Series Conditioning Reaction. By N. H. Luig, K. S. McWhirter and
E. P. Baker.

5. Australasian Ceratopogonidae (Diptera, Nematocera). Part VIII. A New Genus
from Western Australia attacking Man. By Willis W. Wirth and David J. Lee.

6. Spores and Pollens from a Permian-Triassic Transition, N.S.W. By J. P. F.
Hennelly.

7. Bat-infesting Ornithodoros (Ixodoides-Argasidae) of the Oriental-Australian
Region. By L. J. Dumbleton. (Communicated by Dr. J. W. Evans.)

8. On some Pergine Sawflies reared by Mr. M. F. Leask (Hymenoptera, Pergidae).
By Robert B. Benson. (Communicated by Mr. K. EH. W. Salter.)

NOTES AND EXHIBITS.

Mr. G. H. Hardy exhibited slides of small Diptera and of a male scale insect
that had been cleaned in turpentine-phenol and mounted directly from this on a
microscope slide, preserving the natural colours and markings. The flies can be left
in the medium indefinitely without obvious defects clearing the fatty matter, and it
leaves the cuticle coloration unharmed.

Dr. H. S. McKee exhibited a copy of the diary of Charles Moore of his journey
to the Pacific Islands in 1850.

Miss Isobel Bennett exhibited Kodachrome slides of Echinoderms taken at Heron
Island whilst the animals were still alive and submitted the following note:

Echinoderms from the Capricorn Group, Queensland, 23° 27’ S—Some New Locality
Records.

In August, 1958, members of the Sydney University Biological Society, whilst
working at the Marine Biological Station at Heron Island, were taken out in the prawn
trawler ‘“Challenge”’—a vessel carrying out penaeid exploratory work under charter
to the Commonwealth Government.

A 30-minute trawl was made in 25 fathoms off the eastern end of Wistari Reef,
just to the south of Heron Island. It was noteworthy in that the bulk of the catch
consisted of Echinoderms. Since these differed considerably from the common species
of the reef flats of Heron and Wistari, a collection was brought back and presented
to the Australian Museum.

376 ABSTRACT OF PROCEEDINGS.

Endean (Aust. J. Mar. Freshw. Res., 8:233, 1957) has given the known distribution
of the Queensland shallow-water Hchinoderm fauna, and since the list of species taken
are not recorded from the Capricorn Group, these were considered worth noting.

ECHINODERMATA—ASTEROIDEA: Iconaster longimanus (Mobius), Stellaster incei Gray,
Anthenea aspera Doderlein, Pentaceraster australis (Lutken), Metrodira subulata Gray,
Ophidiaster propinquus Livingstone and Acanthaster brevispinus Fisher. HCHINOIDEA:
Salmacis sphaeroides (L.).

As “reef species” none of the above is recorded south of Low Isles (16° 23’ S.) and
Acanthaster brevispinus has not previously been recorded from Australian waters.

Mr. J. P. Freeland exhibited a series of Kodachrome slides showing the habits and
behaviour of various types of ants.

Dr. HE. J. Reye exhibited an improvised power aspirator for collecting large numbers
of sandflies attracted to man. The stimulus for this gadget was a request for a
“tablespoon” of Culicoides ornatus Taylor made by Commonwealth Laboratories. They
required females and that the material be reasonably free of other insects. Conventional
collecting methods such as light-trap and mouth-operated aspirators proved too laborious.
Six feet of plastic garden hose was attached to a funnel stuck into the air intake of
an electric paint-spraying pump. At its other end the hose fitted to a similar funnel
and a 13 oz. beer can open at both ends. This formed the aspirator casing. A 2%”
rubber plug carrying a few inches of 4” copper pipe fitted the intake end and the
Pipe projecting through the plug carried the insect cage. The insect cage is made of a
cream tin (forming its lid) with right-angled slits in rubber sheet, and a small
tomato-juice tin largely cut away and lined with organdie. When insect activity was
high, catches of one-twelfth to one-sixth tablespoon could be made in three-quarters to
one hour. As the attackers were nearly 100% C. ornatus females, no further sorting
was required.

Dr. I. V. Newman exhibited a shoot from each of two plants of Acacia spectabilis
A. Cunn. ex Benth. These are from a group of twelve plants on the railway cutting
north-west of Pymble station and on the south-west side of the line. Hleven of the
plants are normally erect and upward branching, whether on the sloping side of the
cutting or on the level ground at the top. The twelfth and most distant plant is
completely prostrate, mostly trailing closely to the sloping surface of the cutting, but
also creeping up and beginning to extend through the grass on the level ground at
the top of the cutting. The prostrate form flowered profusely, possibly more so and
earlier than the erect forms. The exhibits represent the prostrate plant and one of the
erect plants. Seeds from the prostrate plant have been collected for sowing and future
genetical study.

Mr. W. J. Peacock exhibited a specimen of Brunonia australis R.Br., from Bell,
N.S.W. Seventy per cent. of a population at Bell are structural heterozygotes. The
structural hybrids exhibit association of 4 and 6 chromosomes at metaphase I of
meiosis as well as the normal bivalents.

Mr. R. Carolin exhibited a specimen of Lawrentia avzillaris Lindl. (Lobeliaceae)
to show a specialized development of the normal Lobeliaceous pollinating mechanism.

Dr. A. R. H. Martin exhibited a card-sorting index for fossil flora of Australia
and submitted a note on evidence of a marine transgression at Myall Lakes, N.S.W.

1940

1927
1940

1922
1927
1952
1912

1952

1952

1949
1950

1955
1956
1954

1935

1946
1940

1952
1948
1954
1958

1941
1929

1946
1955
1924
1949

1911
1952

1949
1931

1955
1927

1934

1949

1956
1957

377

LIST OF MEMBERS.
(15th December, 1958.)

ORDINARY MEMBERS.
(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, Hlizabeth 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., Ph.D., 92 Warrigal Road, Surrey Hills, E.10, Victoria.

Aurousseau, Marcel, B.Sc., 229 Woodland Street, Balgowlah, N.S.W.

Baas-Becking, L. G. M., Ph.D., D.Se., Bureau of Mineral Resources, Civic Centre,
Canberra, A.C.T.

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.

Barlow, Bryan Alwyn, B.Sc., Department of Botany, Sydney University.

Barnard, Robert Alexander Stephen, 25 Alt Street, Ashfield, N.S.W.

Baur, George Norton, B.Se., B.Se.For., Dip.For., Research Section, The Forestry Office,

Coff's Harbour Jetty, N.S.W.

*Beadle, Professor Noel Charles William, D.Se., University of New England, Armidale, 5N,
N.S.W.

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

Beattie, Joan Marion, D.Se. (née Crockford), c.o. Mr. G. A. Beattie, Golden Plateau
Mine, Cracow, Queensland.

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

Besly, Miss Mary Ann Catherine, B.A., Department of Zoology, Sydney University.

Black, Roger Foster, B.Se., C.S.I.R.O., Irrigation Research Station, Griffith, 5S, N.S.W.

Blake, Clifford Douglas, B.Se.Agr., Biological Branch, N.S.W. Department of Agriculture,
Farrer Place, Sydney. 5

Blake, Stanley Thatcher, D.Se. (Q’ld.), Botanic Gardens, Brisbane, Queensland.

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

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

Briggs, Miss Barbara Gillian, 13 Findlay Avenue, Roseville, N.S.W.

Browne, Ida Alison, D.Sc. (née Brown), 363 Edgecliff Road, Edgecliff, N.S.W.

Browne, Lindsay Blakeston Barton, Ph.D., C.S.I.R.O. Division of Entomology, P.O. Box
109, City, Canberra, A.C.T.

Browne, William Rowan, D.Sc., 363 Hdgecliff Road, Edgecliff, N.S.W.

Bunt, John Stuart, B.Se.Agr., Ph.D., School 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.

Cameron, Miss Beryl Marlene, B.Sc., Department of Zoology, Sydney University.

Campbell, Thomas Graham, Division of Hconomic Entomology, C.S.1I.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.

Carolin, Roger Charles, B.Sc., A.R.C.S., Department of Botany, Sydney University.

Casimir, Max, B.Sec.Agr., Flat 2, 36 Benelong Road, Cremorne, N.S.W.

LIST OF MEMBERS.

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

Chambers, Thomas Carrick, M.Se. (N.Z.), Department of Botany, Sydney University.

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

*Churchward, John Gordon, B.Sc.Agr., Ph.D., 55 Belmont Street, Mosman, N.S.W.

Clark, Laurance Ross, M.Sc., c.o. C.S.1.R.O., Division of Entomology, P.O. Box 109, City,
Canberra, A.C.T.

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

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

Clinton, Kenneth John, School of Public Health and Tropical Medicine, Sydney
University.

Cogger, Harold George, 4 Blane Street, Granville, N.S.W.

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

Colless, Donald Henry, Department of Parasitology, University of Malaya, Sepnoy Lines,
Singapore, Malaya.

Common, flan Francis Bell, M.A., M.Sc.Agr., C.S.I.R.O., Division of Entomology,
P.O. Box 109, City, Canberra, A.C.T.

Copland, Stephen John, M.Sce., 15 Chilton Parade, Warrawee, N.S.W.

Costin, Alex Baillie, B.Sc.Agr., C.S.I.R.O., Division of Plant Industry, P.O. Box 109,
City, Canberra, A.C.T. 4

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

Cramp, Miss Marion Falconer, B.Sc., Department of Zoology, Sydney University.

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

Crocker, Professor Robert Langdon, D.Sc., Department of Botany, Sydney University.

Crook, Keith Alan Waterhouse, M.Se., Department of Geology, University of New
England, Armidale, 5N, N.S.W.

Davies, Stephen John James Frank, B.A. (Cantab.), 61 Abbotsford Road, Homebush,
N.S. W.

Davis, Professor Gwenda Louise, Ph.D., B.Sec., Faculty of Science, University of New
England, Armidale, 5N, N.S.W.

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

de Bavay, Mrs. Jill Armson, B.Se. (née Whitehouse), Department of Botany, University
of New England, Armidale 5N, N.S.W.

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

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

Domrow, Robert, B.A., B.Sc., Queensland Institute of Medical Research, Herston Road,
Herston, N.9, Brisbane, Queensland.

Donnelly, Robert Bede, 32 Victoria Street, Waverley, N.S.W.

Douglas, Geoffrey William, B.Agr.Sci., 226 Clarendon Street, East Melbourne, C.2,
Victoria.

Drover, Donald P., Ph.D. (W.A.), ¢c.o. Institute of Agriculture, University of Western
Australia, Nedlands, Western Australia.

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

Dyce, Alan Lindsay, B.Se.Agr., C.S.I.R.O., Wildlife Survey Section, P.O. Box 109, City,
Canberra, A.C.T.

Ealey, Eric H. M., M.Sc., c.o. C.S.I.R.O., University Grounds, Nedlands, Western Australia.

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

Elliott, John Henry, 16 Malton Road, Beecroft, N.S.W.

Endean, Robert, M.Se., Department of Zoology, University of Queensland, Brisbane,
Queensland.

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

Evans, Miss Gretchen Pamela, c.o. Department of Botany, Sydney University.

Evans, John William, M.A., D.Se., Se.D., 47 Bundarra Road, Bellevue Hill, N.S.W.

*Wairey, Kenneth David, Box 1176, G.P.O., Sydney.

Filewood, Lionel Winston Charles, 62 Dickson Avenue, West Ryde, N.S.W.

Frame, William Robert, Goroka, 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.Sce., ‘““Hopetoun’”, 25 Bellamy Street, Pennant Hills, N.S.W.

Freeland, Jonn Percy, 19 Central Avenue, Como, N.S.W.

LIST OF MEMBERS. 379

*Garretty, Michael Duhan, D.Se., Box 763, Melbourne, Victoria.

Green, John William, B.Sc. (Adel.), Department of Botany, University of New England,
Armidale 5N, N.S.W.

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

*Grifiiths, Mrs. Mabel, B.Sc. (née Crust), 89 Stock Road, Bicton, Western Australia.

Griffiths, Mervyn Edward, M.Sc., C.S.I.R.O., Wildlife Survey Section, P.O. Box 109, City,
Canberra, A.C.T.

*Gunther, Carl Ernest Mitchelmore, M.B., B.S., D.T.M., D.T.M. & H. (England), 29
Flaumont Avenue, Lane Cove, N.S.W.

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

*Hansford, Clifford Gerald, M.A., Sc.D. (Cantab.), D.Sc. (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.

Hennelly, John Patten Forde, B.Sc., Highs Road, West Pennant Hills, N.S.W.

Hewitt, Bernard Robert, B.Sc, M.Sc. (N.S.W. Univ. Tech.), Department of Agriculture
and Stock, Toowoomba, Queensland.

Heydon, George Aloysius Makinson, M.B., Ch.M., 9 Sirius Avenue, Mosman, N.S.W.

Hill, Miss Dorothy, M.Se., Ph.D., Department of Geology, University of Queensland,
Brisbane, Queensland.

*Hindmarsh, Miss Mary Maclean, B.Sce., Ph.D., 79 Onslow Street, Rose Bay, N.S.W.

*Holder, Miss Lynette Anne, B.Sc., 48 Rutiedge Street, Eastwood, 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.

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

*Hotchkiss, Arland Tillotson, M.S., Ph.D. (Cornell), Department of Biology, University
of Louisville, Louisville 8, Kentucky, U.S.A.

Hotchkiss, Mrs. Doreen Elizabeth, Ph.D., B.A. (née Maxwell), 2440 Longest Avenue,
Louisville, Kentucky, U.S.A.

Humphrey, George Frederick, M.Se., Ph.D., C.S.I.R.O. Marine Biological Laboratory,
Box 21, Cronulla, N.S.W.

Jacobs, Maxwell Ralph, D.Ing., M.Se., Dip.For., Australian Forestry School, Canberra,
ARCADE

Jessup, Rupert William, M.Sc., 12 Berrigan Crescent, O’Connor, Canberra City, A.C.T.

Jobson, Arthur Edgar, 3 Wellington Road, East Lindfield, N.S.W.

Johnson, Bruce, B.Se.Agr., Ph.D., Waite Agricultural Research Institute, University of
Adelaide, Private Mail Bag, Adelaide, South Australia.

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

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

Jones, Edwin Llewelyn, B.A., 491 Alfred Street, North Sydney, N.S.W.

Jones, Mrs. Valerie Margaret Beresford, M.Se. (née May), 36 The Hsplanade, Narrabeen,
N.S. W.

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

Keast, James Allen, M.Se., M.A., Ph.D. (Harvard), Australian Museum, College Street,
Sydney.

Kerr, Harland Benson, B.Se.Agr., Ph.D., 41 Badminton Road, Croydon, N.S.W.

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

Kesteven, Hereward Leighton, D.Se., M.D., Ch.M., 326 Beaconsfield Terrace, Brighton,
Brisbane, Queensland.

Kindred, Miss Berenice May, B.Sc., 58 Caroline Street, Kingsgrove, N.S.W.

Langdon, Raymond Forbes Newton, M.Agr.Se., Ph.D., Department of Botany, University
of Queensland, George Street, Brisbane, Queensland.

Langford-Smith, Trevor, M.Sc., Department of Geography, Sydney University.

*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.Sc. (née Melvaine), Manor Road, Hornsby, N.S.W.

Lee, David Joseph, B.Se., School of Public Health and ‘Tropical Medicine, Sydney
University.

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

Luig, Norbert Harold, 184 Falcon Street, North Sydney, N.S.W.

Lyne, Arthur Gordon, B.Sc., Ph.D., C.S.1.R.O:, Sheep Biology Laboratory, P.O. Box 144,
Parramatta, N.S.W.

LIST OF MEMBERS.

Macdonald, Colin Lewis, 378 Wilson Street, Albury East, N.S.W.

Macintosh, Professor Neil William George, M.B., B.S., Department of Anatomy, Sydney
University. 5

Mackerras, Jan Murray, M.B., Ch.M., B.Sc., Queensland Institute of Medical Research,
Herston Road, Herston N9, Brisbane, Queensland.

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

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

Martin, Anthony Richard Henry, M.A., Ph.D., Department of Botany, Sydney University.

Martin, Mrs. Hilda Ruth Brownell, B.Se. (née Simons), Lot 1, enr. Villiers and Chamber-
lain Roads, Padstow, N.S.W.

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

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

MeCulloch, Robert Nicholson, M.B.E., D.Se.Agr., B.Sc., Roseworthy Agricultural College,
Roseworthy, South Australia.

McCusker, Miss Alison, M.Sec., Department of Botany, Sydney University.

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

McGarity, John William, B.Sce., Agr., c.o. Faculty of Agriculture, University of Adelaide,
Adelaide, South Australia.

McKee, Hugh Shaw, B.A., D.Phil. (Oxon.), c/- Department of Botany, Sydney University.

McKenna, Nigel Reece, 78 L’Estrange Terrace, Kelvin Grove, Brisbane, Queensland.

McMichael, Donald Fred, B.Se., Ph.D. (Harvard), Australian Museum, College Street,
Sydney.

MeMillan, Bruce, M.B., B.S., D.T.M.& H. (Eng.), D.A.P. & E., F.R.E.S., School of Public
Health and Tropical Medicine, Sydney University.

Menzies, Miss Barbara Patricia, M.Se. (N.Z.), Ph.D. (Cantab.), i6 Lucerne Road,
Remuera, Auckland, S.E.2, New Zealand.

Mercer, Professor Frank Verdun, B.Sc., Ph.D. (Camb.), Department of Botany, Sydney
University.

Mercer, Mrs. Greta, B.A., B.Sc. (née Baddams), 13 Hampton Road, Keswick, South
Australia.

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

*Meyer, George Rex, B.Sc., Dip.Ed., B.A., M.Ed., 91 Bowden Street, Ryde, N.S.W.

*Miller, Allen Horace, B.Sc., Dip.Ed., 6 College Avenue, Armidale 5N, N.S.W.

Miller, David, C.B.E., Ph.D., M.Se, F.R.S.N.Z., E.R.E.S., Cawthron Institute, Nelson,
New Zealand. 5

Millerd, Miss Alison Adéle, M.Se., Ph.D., Waite Agricultural Research Institute, Private
Mail Bag No. 1, Adelaide, South Australia.

Millett, Mervyn Richard Oke, B.A., 18 Albion Road, Box Hill, #11, Victoria.

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

Moore, Kenneth Milton, Cutrock Road, Lisarow, N.S.W.

Morgan, Mrs. Eva, M.Sc., 4 Haberfield Road, Haberfield, N.S.W.

Moss, Francis John, 70 Victoria Road, West Pennant Hills, N.S.W.

Moye, Daniel George, B.Se., Dip.Ed., 6 Kaling Place, Cooma North, N.S.W.

Muirhead, Warren Alexander, B.Sce.Agr., Soil Conservation Service, Box 49, Hay 6S, N.S.W.

Mulder, Rudolph Herman, Box 31, P.O., Kogarah, N.S.W.

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

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

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

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

Newman, Ivor Vickery, M.Sc, Ph.D., F.R.M.S., F.L.S., Department of Botany, Sydney
University.

Nicholson, Alexander John, D.Sc., F.R.E.S., C.S.I1.R.O., Box 109, Canberra, A.C.T.

*Noble, Norman Scott, D.Sce.Agr., M.Se., D.I.C., C.S.1.R.O., 3814 Albert Street, East
Melbourne, C.2, Victoria.

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

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

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

O’Gower, Alan Kenneth, M.Sc., 21 Gaerloch Avenue, South Bondi, N.S.W.

Oke, Charles George, 34 Bourke Street, Melbourne, C.1, Victoria.

Osborn, Professor Theodore George Bentley, D.Sc, F.L.S., St. Mark’s College,
Pennington Terrace, North Adelaide, South Australia.

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

1952
1954
1940
1957

1957
1922

1947
1957
1937

1935

1938

1949

1956

1929
1951

1952
1957
1953

1958

1957
1946

1958
1936
1932

1932
1919
1950
1948

1930
1947
1957
1953

1955
1955

1952
1953
1916
1943
1945
1937

1926

1932
1956
1956

1949
1958

1935
1952

LIST OF MEMBERS. 381

Packham, Gordon Howard, Department of Geology, Sydney University.

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

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

Paterson, Miss Betsy. Rivers, B.Sc., Department of Botany, University of New England,
Armidale, 5N, N.S.W.

Peacock, William James, Department of Botany, Sydney University.

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

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

Phillis, Miss Elizabeth Robin, 22 Balmoral Street, Waitara, N.S.W.

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

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

Pryor, Lindsay Dixon, M.Sc., Dip.For., Parks and Gardens Section, Department of the
Interior, Canberra, A.C.T.

Purchase, Miss Hilary Frances, Ph.D. (Lond.), B.Se.Agr., Faculty of Agriculture,
Sydney University.

Rae, Mrs. Clare Annetta, B.Sc., Department of Zoology, University of New England,
Armidale 5N, N.S.W.

Ragegatt, Harold George, C.B.E., D.Sec., 60 Arthur Circle, Forrest, Canberra, A.C.T.

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

Ramsay, Mrs. Helen Patricia, M.Sc. (née Lancaster), 34 Haig Avenue, Ryde, N.S.W.

Reed, Eric Michael, 7 Frederick Street, Campsie, N.S.W.

Reye, Eric James, M.B., B.S. (Univ. Qld.), School of Public Health and Tropical
Medicine, Sydney University.

Reye, Mrs Margaret Burnett, B.Sc., c.o. A. & N.Z. Bank, Cnr. Queen and Creek Streets,
Brisbane, Queensland.

Reynolds, Miss Judith Louise, 132 Homer Street, Earlwood, N.S.W.

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

Rigby, John Francis, B.Se., 5 Banool Street, Keiraville 5C, N.S.W.

Roberts, Noel Lee, 43 Hannah Street, Beecroft, N.S.W.

*Robertson, Rutherford Ness, B.Sc., Ph.D., F.A.A., Food Preservation Research Labora-
tory, C.S.I.R.O., Private Mail Bag, Homebush, N.S.W.

*Salter, Keith Eric Wellesley, B.Sc., Department of Zoology, Sydney University.

*Scammell, George Vance, B.Sc., 7 David Street, Clifton Gardens, N.S.W.

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

Shaw, Miss Dorothy Edith, M.Se.Agr., Ph.D., Department of Agriculture, Stock and
Fisheries, Port Moresby, Papua-New Guinea.

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

Shipp, Erik, 23 Princes Street, Turramurra, N.S.W.

Shortman, Kenneth Douglas, Cooriengah Heights Road, Engadine, N.S.W.

Simonett, David Stanley, M.Sc, Ph.D., Department of Geography, The University of
Kansas, Lawrence, Kansas, U.S.A.

Slack-Smith, Richard J., c.o. Department of Fisheries and Game, Melbourne, Victoria.

Slack-Smith, Mrs. Shirley Margaret (née Ryan), c.o. Department of Fisheries and Game,
Melbourne, Victoria.

Slade, Milton John, B.Sc., 10 Blizabeth 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, D.Sc.Agr., Department of Botany, Sydney University.

Southcott, Ronald Vernon, M.B., B.S., 13 Jasper Street, Hyde Park, South Australia.

Spencer, Mrs. Dora Margaret, M.Sc. (née Cumpston), Mapamoiwa, Fergusson Island,
via Samarai, Papua.

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

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

Steinberg, Miss Joan Emily, B.A., 850 38th Avenue, San Francisco, California, U.S.A.

Stephenson, Neville George, M.Se. (N.Z.), Ph.D. (Lond.), Department of Zoology,
Sydney University.

Stevens, Neville Cecil, B.Sc., Ph.D., Department of Geology, University of Queensland,
St. Lucia, Brisbane, Queensland.

Stewart, Alexander Drennen, B.Sc., c.o. Mrs. R. Hastick, 108 Brook Street, Coogee, N.S.W.

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

Sullivan, George Emmerson, M.Sc. (N.Z.), Department of Histology and Embryology,
Sydney University.

382

1911
1955

1940
1950

1950
1956
1949
1944
1943
1946
1921
1950

1952
1949
1917
1930
1940

1934

1952

1909
1946

1947
1930
1911
1936
1947
1927
1941

1949
1946

1926
1954
1954
1952
1950
1947
1934
1949
1932
1936
1949
1949

1957

LIST OF MEMBERS.

*Sulman, Miss Florence, 11 Ramsay Street, Collaroy, N.S.W.
Sutherland, James Alan, B.A., B.Sc.Agr., ‘Skye’, Galloway Street, Armidale 5N, 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., 6S Sciences (Paris), Microbiology Laboratory, Faculty of Agri-
culture, Sydney University.

Thompson, Mrs. Joy Gardiner, B.Sc.Agr. (née Garden), Botanic Gardens, Sydney.

Thomson, James Miln, D.Se. (W.A.), c.o. C.S.I.R.O., Box 21, P.O., Cronulla, N.S.W.

Thorp, Mrs. Dorothy Aubourne, B.Se. (Lond.), “Sylvan Close’, Mt. Wilson, N.S.W.

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

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

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

*Troughton, Ellis Le Geyt, C.M.Z.S., F.R.Z.S., No 1 Hall Street, Bondi Beach, N.S.W.

Tugby, Mrs. Hlise Evelyn, M.Se. (née Sellgren), Department of Geography, Australian
National University, Box 4, G.P.O., Canberra, A.C.T.

Valder, Peter George, B.Sc.Agr., Ph.D. (Camb.), Biological Branch, N.S.W. Department
of Agriculture, Box 36, G.P.O., Sydney.

Vallance, Thomas George, B.Sc., Ph.D., Department of Geology, Sydney University.

Veitch, Robert, B.Sc., F.R.E.S., 24 Sefton Avenue, Clayfield, Brisbane, Queensland.

Vickery, Miss Joyce Winifred, D.Se., Botanic Gardens, Sydney.

Vincent, Professor James Matthew, D.Se.Agr., Dip.Bact., Faculty of Agriculture, Sydney
University.

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

Walker, John, B.Se.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.Se., Institute of Agriculture, University of Western
Australia, Nedlands, Western Australia.

Ward, Mrs. Judith, B.Sc., c.o. H.E.C., Wayatinah, Tasmania.

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

Wardlaw, Henry Sloane Halcro, D.Se., F.R.A.C.I., 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.Sc., “Burragillo’, Merrywinebone 6N, N.S.W.

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

Watson, Professor 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 Laboratery,
Kuantan. Pahang, Federation of Malaya.

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

Williams, John Beaumont, B.Se., University of New Hngland, Armidale, 5N, N.S.W.

Williams, Mrs. Mary Beth, B.Se. (née Macdonald), Department of Botany, University
of New England, Armidale, 5N, N.S.W.

Williams, Owen Benson, M.Agr.Se. (Melbourne), c.o. C.S.ILR.O., Box 229, P.O., 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.

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

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

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

Zeck, Emil Herman, 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.Sc.Agr. (Copenhagen), State Laboratory of Plant Culture,
Department of Bacteriology, Lyngby, Denmark.

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

I.—Seed coat anatomy in Hucalyptus.

LIST OF PLATES.
PROCEEDINGS, 1958.

Il —Hupomatia bennettii F. Muell.
III.— Geological Map of the Cooleman Caves District.

IV.—Cooleman Caves District.

V.—ViI.—Spores and pollens from a Permian-Triassic transition.

LISTS OF NEW GENERA, SPECIES, SUBSPECIES, AND NAME.

Atellana (Listrophoridae)
Austroconops (Nematocera)
Quadrisporites or as

alcithoe (Trombigastia)
aristippe (Jchoronyssus)
aureum (Hrodium)

australis (Zygophloeus)

brevicauda (Hemicycliophora)

britteni (Aédes (Finlaya) )
bulliensis (Apiculatisporites)
burnsi (Rhagigaster)
crinitum (Hrodium)
dasyphloea (Trombicula)
fibulatus (Cirratriradites)
horridus (Quadrisporites)
kiandrensis (Rhagigaster)
leaski (Perga) :
leucippe (Ichoronyssus)
lugubris (Thynnoides)
memillani (Austroconops)
montanus (Rhagigaster)

dium)

VoL. 83.

Genera.

Page
43

Trichonyssus (Laelaptidae)

so BOT Zygophloeus (Scolytoidea)
.. 364

Species.
5a Bae mulesi (Hirone)
5 HORS nigracristatus (Pityosporites)
5 oF paludosa (Pultenaea) 5
50 CAs papilio (Atellana)
5 PAL) perkinsi (Austrochirus)
so) radiatus (Nuskoisporites)
. 364 reticulatus (Pityosporites)
. 313 robustum (Oxzylobium)
oo} 6B sloanei (Crinia)
mez solomonis (Ornithodoros)
.. 366 squamifer (Heteropsilopus)
. 365 stradbrokensis (Rhagigaster)
Renny) tesselata Cet LED):
. 288 tumidus (Sciapus) :
50 LA vandiemeni (Hoogstraalia)
.. 336 wilsoni (Thynnoides)
50 BO womersleyi (Trichonyssus)
. 319

Subspecies.

eygnorum ssp. glandulosum (Hro-
Rie oe RAMA oye Bat ECO
New Name.
(Pultenaea) . 188

blakelyi

383

Page

. 230

. 215

. 325

_. 366

. 188
43
41

_. 366

. 367

_. 187

. 225

. 306
.. 295
. 315
.. 217
. 299
.. 173
-- 336
. 231

384

INDEX.

1958.

Page
Abbie, A. A., Timing in Human
Evolution PRES LEO
Abstract of Pr aceaitings . 371-376
Acarina from Australian Bats .. 5 eal
Aédes australis (Hrickson), The Ovi-
position Behaviour of .. 245
Aédes (Finlaya) from Northern “Aus:
tralia, A New Species of 33
Annual General Meeting 1
Aphis craccivora Koch, Wieration
and Utilization of Reserve Sub-
stances during Flight in 165
Aphodus tasmaniae Hope, A Note on
the Status of . : 196
Atopomelinae, A Smarty of the 40
Australasian Ceratopogonidae. VIII.
A New Genus from Western Aus-
tralia attacking Man 337
Australian Bats, Acarina from 227
Australian Mammals, Catalogue of,
and their Recorded Internal
Parasites, I-IV 101
Australian Thynninae—
Notes on II, 309—III, 327
Bacteria, Studies of mas Rta ey
VII 161
Bacteriology aang, srepant re salle oe
Ramsgate property 4
Baker, EH. P., see Luig, N. HH.
McWhirter, K. S., and Baker,
E. P.
Balance Sheets for the Year ending
28th February, 1958 : 6-8
Barberry, Widespread Natural Tnviac.
tion of, by Puccinia graminis in
Tasmania ; Srcamreanem Loi
Bat-infesting Ornithodoros. of the
Oriental-Australian Region . 303
Bats, Australian, Acarina from . 227
Bearup, A. J., see Notes and Exhibits.
Beetles, Bark- and Timber- from
Australia, Some More .. he eae:
Bennett, Isobel, see Notes and
Exhibits.
Benson, R. B., On some Pergine Saw-
flies reared by Mr. M. F. Leask 288
Benson, W. N., obituary notice 4
Bird-flea from Tasmania, A New 173
Blake, C. D., elected 2 member, 371—
A Turbidimetric Method for
Estimating the Number of Nema-
tode Larvae in a Suspension .. 241
Browne, W. R., elected Honorary
Secretary Bee a hony tush ARAL eA RON AL
Bryant, L. H., see Pryor, L. D., and

Bryant, L. H.

Page

Carolin, R. C., The Species of the
Genus Hrodium L’Hér. endemic
to Australia. (With a Key to all
the Taxa known to occur in
Australia.), 92—see Notes and

Exhibits.

Casimir, M., Migration and Utiliza-
tion of Reserve Substances
during Flight in eae cracci-
vora Koch

Catalogue of ungimeiien Oviemmoale
and their Recorded Internal
Parasites, I-IV

Ceratopogonidae, Australasian, VIII
Citrus in Australia, Virus Diseases
of

Congratulations “0 mentivers

2, 371, 378,

Cooleman Caves District, New South

Wales, Palaeozoic Geology of the
Copland, S. J., elected a Vice-
President
Diptera of Katoomba, 2. Leptidae

and Dolichopodidae ak
Domrow, R., Acarina from Aus:

tralian Bats, 227—A Summary

of the Atopomelinae 5
Dumbleton, L. J., Bat-infesting Ome

thodoros of the Oriental-Aus-
tralian Region
Elections nN sags Pe 5,
Hrodium WL’Her. eailennie to Aus-

tralia, The Species of the Genus
Eucalyptus, Seed Coat Anatomy and
Taxonomy in, 20—Inheritance of
Oil Characters in ..
Eupomatiaceae, Pollen sail Pollina
tion in the :
Evolution, Human, mMming in
Exchange Relations ae
Exhibits—see Notes and EeRipitee

Fraser, Lilian R., elected a Vice-
President, 3871—Virus Diseases
of Citrus in Australia, 9—Sum-
mary of Address ee

Freeland, J. P.—see Nowes: andl x

hibits.

Frog of the Genus Crinia from
South-eastern Australia, A New
Species of

Gauba, E., and Pryor, L. D., Seed

Coat Anatomy and Taxonomy in
Hucalyptus, 1 ..

. 165

. 101

337
9
375

251

. d71

. 291

40

. 803

371

92

55

86

5 IY

2

. 222

20

INDEX.

Page

Given, B. B., A Note on the Status of
Aphodius tasmaniae Hope, 196—
Notes on Australian Thynninae,
II, 309—III

Green, J. W., elected a Cae

Hadlington, P. W., elected a member

Hannon, Nola J., Linnean Macleay
Fellow in Botany, summary of
work, 1957, 2—reappointed for
1958, 3—resignation, April, 1958,
3—The Status of Nitrogen in the
Hawkesbury Sandstone Soils
and their Plant Communities in
the Sydney District. II. The
Distribution and Circulation of
Nitrogen Leonie, otc dlr OCA abe

Hardy, G. H., The Diptera of
Katoomba. 2. Leptidae and Doli-
chopodidae, 291—see Notes and
Exhibits.

Hawkesbury Sandstone Soils and
their Plant Communities in the
Sydney District, the Status of
Nitrogen in the, II :

Hemicycliophora, Two New Species
of a

Hennelly, 7. P. F, ‘elected < a “member,
373—Spores and Pollens from a
Permian - Triassic Cee
N.S.W. 2

Hodgkin, HE. P., see ‘Warks, Ceiianeh
N., and Hodgkin, EK. P.

Hotchkiss, A. T., Pollen and Pollina-
tion in the Eupomatiaceae

Inheritance of Oil
Eucalyptus

Characters in

Jones, HE. L., elected a member

Katoomba, The Diptera of, 2

Kerr,
Lév. Uredospore Longevity and
Germination ?

Lecturettes Be To eR on, Mere
Lee, D. J., see Wirth, W. W.,
Lee, D. J.
Library Accessions ..
Linnean Macleay Fellowships:
Reappointments for 1958, 3—appli-
cations invited for 1959, 374—
appointment for 1959 .. F
Linnean Macleay Lectureship in
Microbiology, Summary of re-
ports of Dr. Y. T. Tchan from
Ist August, 1955, to 31st De-
cember, 1957
List of Members : ee
Lists of New Genera, Species, Sub-
species and Name ..
List of Plates Siirlo es
Littlejohn, M. J., A New Species of
Frog of the Genus Crinia
Tschudi from South-eastern Aus-
tralia

and

H. B., Melampsora lini (Pers.)~-

. 327
. 371

375

65

65

217

. 363

. 2, 371-375

_. 317

. 383
. 383

. 222

385

Page

Luig, N. H., see Watson, I. A., and
Luig, N. H.—see Notes and
Exhibits.

Luig, N. H., McWhirter, K. S., and
Baker, HK. P., Mode of Inheritance
of Resistance to Powdery Mildew
in Barley and Evidence for an

Allelic Series ET emn Re-
action
Lyne, A. G., slaciedl a camera

Mackerras, M. Jesephine, Catalogue

of Australian Mammals and
their Recorded Internal Para-
sites, I—IV
Mammais, Australian, Cahaldene Of
and their Recorded Internal
Parasites ... Ae one ane
Marks, Elizabeth N., aad Hodgkin,

E. P., A New Species of Aédes
(Finlaya) from Northern Aus-
tralia NO ETRE SPOR I 6
Marlow, B. J. G., elected a member
Martin, A. R. H.—see Notes and
Exhibits.
Mawson, Sir D., reference to death ..
McCusker, Alison, appointed Linnean

Macleay Fellow in Botany for
1959 cele! UR i. Aiea es

McKee, H. S.—see Notes and
Exhibits.

McMichael, D. F.—see Notes and
Exhibits.

McWhirter, K. S., see Luig, N. H.,
McWhirter, K. S., and Baker,
EH. P.

Melampsora lini (Pers.) Lév. Uredo-
spore Lengevity and Germina-
ION 66

Members, List of

Mercer, F. V.,
President

Migration and Utilization of Reserve
Substances during Flight in
Aphis craccivora Koch ;

Mode of Inheritance of Resistance in
Powdery Mildew in Barley and
Evidence for an Allelic Series
conditioning Reaction

Moore, K. M., elected a member

Moss, F. J., elected a member

Moss, F. J., and Tchan, Y. T., Qaadiies
of Nitrogen- fixing Bacteria, VII

elected a _ Vice-

Nematode Larvae in a Suspension,
A Turbidimetric Method for
estimating the Number of

New Bird-flea from Tasmania ..

Newman, JI. V.—see Notes and
Exhibits.
New Species of Aédes (Finlaya)

from Northern Australia ;
New Species of Frog of the Genus
Crinia Tschudi from South-
eastern Australia ..
Note on the Status of ‘Aphodius
tasmaniae Hope

.. 340
. 373

101

. 101

33
374

374

. 375

so ZY)
. 317

371

. 165

.. 340
. 373

371

161

. 241
Sis

33

222
196

386 INDEX.

Page
Notes and Exhibits:

Bearup, A. J.—Demonstration of
the life cycle of Acanthopary-
phium spinulosum S. J. Johnston 372

Bennett, Isobel — Kodachrome
slides of Echinoderms taken at
Heron Island and a Note on
Hchinoderms from the Capricorn
Group, Queensland, 23° 27’ S.—
Some New Locality Records .. 375

Carolin, R. — A _ specimen . of
Laurentia aaillaris Lindl. to
show a specialized development
of the normal Lobeliaceous pol-
linating mechanism .. . 376

Freeland, J. P—Kodachrome ‘slides
showing habits and behaviour of
various types of ants ‘ 376

Hardy, G. H.—Slides of small Dip-
tera and of a male scale insect
that had been cleaned in turpen-
tine-phenol and mounted directly
from this on a microscope slide,
preserving the natural colours
ana markings, 375—Some Platy-
pezidae; also a mite-infested
specimen mounted in a micro-
slide with mites in situ and
pinned specimens without mites 372

Luig, N. H.—Slide of a _ rust-
infected seedling blade of wheat 372

Martin, A. R. H.—A card-sorting
index for fossil flora of Australia
and a note on evidence of a
marine transgression at Myall
Lakes, N.S.W. ae 376

McKee, H. S.—Copy of the diary of
Charles Moore of his journey to
the Pacific Islands in 1850 .. 375

McMichael, D. F.—Specimens of an
unidentified freshwater animal
from New Guinea Me 373

Newman, I. V.—A shoot from “each
of two plants of Acacia spec-
tabilis A. Cunn. ex Benth. .. 376

Peacock, W. J.—A specimen of
Brunonia australis R.Br. from
Bell, N.S.W. .... . 316

Reye, EH. J.—An improvised power
aspirator for collecting large
numbers of sandflies attracted
COs anaes Oe EEE oe ie a er OG

Smith-White, S.—Specimens of
Astroloma conostephioides and
of A. pinifolium from the Gram-
pians, Western Victoria .. 372

Notes on Australian Thynninae. II.
The Genera Dimorphothynnus,
Rhagigaster and Hirone, 309—
Ill. The Genus Thynnoides .. 327

Obituaries: W. N. Benson; D. G.
Stead SRA MAL ttl sae al 4

O’Gower, A. K., The Oviposition
Behaviour of Aédes australis
(Erickson) a . 245

On some Pergine Sawflies ‘reared by
Mr iM. i. leask @en74 ee ee aes 288

Page
Oriental-Australian Region, Bat-
infesting Ornithodoros of the .. 303

Ornithodoros, Bat-infesting, of the

Oriental-Australian Region oo oR}
Oviposition Behaviour of Aédes
australis (Wrickson) .. .. .. 245

Palaeozoic Geology of the Cooleman
Caves District, New South Wales 251

Papilionaceae, Systematic Notes on
some Hastern Australian Mem-
bers: (Of thee, Ay er Te ee ea yi

Peacock, W. J.—see Notes and
Exhibits.

Pengilley, R. K., elected a member.. 371

Permian and Triassic Transition,
N.S.W., eaten and Pollens from

a ; Lit” Rola pea ERE OOO
Plates, List of ec LIS 383
Pollen and Pollination in ang ane

matiaceae ae 86

Powdery Mildew in Banley, Mode of
Inheritance of Resistance to,
and Evidence for an Allelic

Series Conditioning Reaction .. 340
Presidential Address .. .. ae 1
Pryor, L. D., see Gauba, E amd

Pryor, L. D.

Pryor, L. D., and Bryant, L. H.,
Inheritance of Oil Characters in

Hucalyptus Pm CS Rae eRe vo nt)
Puccinia graminis var. tritici,
Somatic Hybridization in .. .. 190

Reye, EH. J.—see Notes and Exhibits.
Reye, Margaret B., elected a member 371
Rigby, J. F., elected a member... 375

Sauer, M. R., Two New Species of
Hemicycliophora eee es een

Sawflies, On some Pergine, reared
by Mr. M. F. Leask .. 288

Schedl, K. E., Some More Bark- and
Timber- beetles from Australia 214

Science House us 5 2
Seed Coat Anatomy and Taxonomy
in Hucalyptus, I .. . 20

Sir William Macleay Memorial Lec-
ture established, 1—announce-

ment of, 371—lecture, 1958 .. 197
Smit. F. G. A. M., A New Bird-flea
from Tasmania Aa 173

Smith-White, S., elected President,
5—see Notes and Exhibits.

Somatic Hybridization in Puccinia
graminis var. tritici .. . . 190

Some More Bark- and Timber-beetles
from Australia. 156. Contribu-
tion to the Morphology and
Taxonomy of the Scolytoidea .. 214

Species of the Genus Hrodiwm L’Hér.
endemic to Australia. (With a
Key to all the Taxa known to

occur in Australia) .. .. .. 92
Spores and Pollens from a Permian-
Triassic Transition, N.S.W. .. 363

Page

Status of Nitrogen in the Hawkes-
bury Sandstone Soils and their
Plant Communities in the Syd-
ney District. II. The Distribu-
tion and Circulation of Nitrogen

Stead, D. G., obituary notice

Stevens, N. C., Palaeozoic Geology of

the Cooleman Caves District,
New South Wales ..

Studies of Nitrogen-fixing Bacteria:
VII. Cytochromes of Azotobac-
teriaceae 5

Summary of the Accom olince
Summary of year’s activities

Sydney District, Status of Nitrogen
in the Hawkesbury Sandstone
Soils and their Plant Communi-
ties in the, II

Systematic Notes on some Hastern
Australian Members of the
Papilionaceae

Tasmania, A New Bird-flea from, 173
—Widespread Natural Infection
of Barberry by Puccinia gra-
minis in

Tcehan, Y. T., Linnean hacia ite.
turer in “Microbiology, Summary
of reports, 3—see Moss, F. J., and
Tehan, Y. T.

Thompson, Joy, Systematic Notes on
some Hastern Australian Mem-
bers of the Papilionaceae

. 161

65

.. 187

. 181

. 187

INDEX. 387

Page

Thynninae, Australian, Notes on, II,
SOON 55° oo Boe kerr OvAtL
Timing in Human evolution ee ite OT

Turbidimetric Method for Hstimat-
ing the Number of Nematode
Larvae in a Suspension .. .. 241

Two New Species of Hemicycliophora 217

Vincent, J. M., elected a _ Vice-
President Sah ae 668 edd

Virus Diseases of Ghtars in petra 9

Walkom, A. B., elected Honorary
Treasurer and ae Secre-
LAT: Sete Uae > Sal

Watson, I. A., and Tones N. i, Wide.
spread Natural Infection of Bar-
berry by Puccinia graminis in
Tasmania, 181—Somatie Hybrid-
ization in Puccinia graminis var.
tritici * seal OL) bea, ve eee OD

Widespread NaCUE A Infection of
Barberry by Puccinia Ee
in Tasmania .. .. . 181

Williams, Mary B., Tinian Masieees

Fellow in Botany, summary of

work, 1957, Auge for

DIO EMER ms a is 3
Wilson, F., elected a ee emienareye 375

Wirth, W. W., and Lee, D. J., Aus-
tralasian Ceratopogonidae (Dip-
tera, Nematocera). VIII .. .. 337

Woolnough, W. G., reference to death 374

ii
rea) oe
’ * ¥ ’
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Proc. Linn. Soc. N.S.W., 1958. PLATE It.

GEOLOGICAL MAP
OF THE
COOLEMAN CAVES DISTRICT

%

ONE MILE

FAULTS BOUNDARIES "|
F——F isreraco oo
F Fo operimite = -—-—

11 cave)
ANN, 1 fe

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a

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. Oo,

SY
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o
e
°

LEGEND

ja

d FERRUGINOUS SEDIMENTS (Qf),
GRAVELS(Q9)

INTRUSIVE

KELLYS PLAIN DACITE

\ Na] SURRANGORAMBLA
RHYOLITE (Or) GRANOPHYRE, FELSITE

CH v
= rah

m
x
v A Ye
A WILKINSON LIMESTONE CA DIORITE <a ee e \ ZS yi /
\ ee
BLUE WATERHOLE BEDS BLACK MTN. GRANITE .
AND RELATED GRANODIORITES

POCKET BEDS WITH [#4] GINGERA GRANITE

LIMESTONE ( Spl)
COOLEMAN LIMESTONE MINOR INTRUSIONS a= ocid
DSS i—intermedicte —b—basic

NUNGAR BEDS (On) SILICIFIED AND.
SANDSTONE (Os) DOLOMITIZED ZONES

== 3 -:DEVONIAN,

Proc. Linn. Soc. N.S.W., 1958. PLATE Iv.

Cooleman Caves district.

Proc. Linn. Soc. N.S.W., 1958. PLATE V.

Spores and pollens from a Permian-'Triassie transition.

Proc. Linn. Soc. N.S.W., 1958. PLATE VI.

Spores and pollens from a Permian-Triassic transition.

Thay

4 5
f Sarg as 3
a un
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ax
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(Issued 24th July, 1958.)

; Parne easteAt nt
LIBRA Bi as A
1y59

Pan EXXXHE | DEC.
Part 1. | WOODS HOLE, MASS.

PROCEEDINGS

OF THE

LINNEAN SOCIETY

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FOR THE YEAR

1958

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BY NOLA J. HANNON. 85

Haren, A. B., 1955.—The Influence of Plant Litter on the Jarrah Forest Soils of the
Dwellingup Region, West Australia. Leafl. For. Bur. Canberra, 70, 18 pp.

Jacogs, M. R., 1936.—The Primary and Secondary Leaf Bearing Systems of the Eucalypis
and their Silvicultural Significance. Bull. For. Bur. Aust., 18: 1-78.

, 1955.—“‘Growth Habits of the Eucalypts.” Forestry and Timber Bureau. Common-
“wealth Govt. Printer. 262 pp.

JHNNY, H., 1941.—‘‘Factors of Soil Formation.” McGraw-Hill, New York. 281 pp.

JENNY, H., BineHam, F., and PAaDILLA-SARAVIA, B., 1948.—Nitrogen and Organic Matter
Contents of Equaterial Soils of Colombia, South America. Soil. Sci., 66: 173-186.

JENNY, H., GzsseL, S. P., and BineHam, F., 1949.—Comparative Study of Decomposition
Rates of Organic Matter in Temperate and Tropical Regions. Soil. Sci., 68: 419-432.

KITTREDGE, J., JR., 1948.—“Forest Influences.” McGraw-Hill, New York. 394 pp.

Lutz, H. J., and CHANDLER, R. F., yr., 1949.—“‘Forest Soils.” John Wiley and Sons, New York.
4th printing. 514 pp.

McNamara, P. J., 1955.—A Preliminary Investigation of the Fauna of Humus Layers in
the Jarrah Forest of Western Australia. Leaf. For. Bur., Canberra, 71. 16 pp.

Merz, L. J., 1952.—Weight and Nitrogen and Calcium Content of the Annual Litter Fall of
Forests in the South Carolina Piedmont. Proc. Soil Sci. Soc. Amer., 16: 38-41.

Rennis, P. J., 1955-—-The Uptake of Nutrients by Mature Forest Growth. Plant € Soil,
7, 1: 49-95.

RussELL, HE. J., 1950.—*Soil Conditions and Plant Growth.” Longmans, Green and Co.,
London, 8th ed. 635 pp.

Scott, D. R. M., 1955.—Amount and Chemical Composition of the Organic Matter contributed
by Overstory and Understory Vegetation to Forest Soil. Bull. Yale Univ. Forestry School,
PAD Ot (NG 99 8%

Simons, Hiupa R., 1951.—A Study of Fungi Active in the Decomposition of Litter of
Casuarina littoralis Salisb. (suberosa Ott. and Dietr.). Honours Thesis, Botany Dept.,
University of Sydney.

SPECHT, R. L., 1953—The Ecology of the Heath Vegetation in the Upper South Hast of
South Australia— A Preliminary Investigation. Ph.D. Thesis, University of Adelaide.
TURNER, J. C., 1954.—Some Aspects of the Nutrient Cycle in Rainforest in New South Wales.

Honours Thesis, Faculty of Agriculture, University of Sydney.
- Wisk, L. E., and JoHN, EH. C., Hd. 1952.—‘“Wood Chemistry” I. Am. Chem. Soc. Monograph
Series. Reinhold, N.Y. 1-688.

PROCEEDINGS, LXXXIII, PART 1, 1958

CONTENTS.
Pages
Presidential Address, delivered at the Highty-third Annual General Meeting,
26th March, 1958, by Dr. Lilian R.. Fraser:
SuMMary VOLU ear SwACHVITLTES Hy cei. eh liared tie van ehes tne nuM EEC cite, uterine ays 1-5
Virus) Diseases 01 .Citrus) in Australia Wy. uieten) jens icici ken eveieieats 9-19
NTO CELONIS Uehara cise e es Rod deste etait Cone h Lea Mneele Vide yi agatel P RALOTA , <ciys Satta nga aR, UOC ae Baas 5
Balance Sheets for the Year ending 28th February, 1958 .. .. .. .. .. .. 6-— 8
Seed Coat Anatomy and Taxonomy in Eucalyptus. I. By E. Gauba and L. D.
Prvyorcelate ieanineteen Lext-ficuress) mee weni a oie ee ei tees) aie eee Uae
A New Species of Aédes (Finlaya) from Northern Australia (Diptera, Culicidae).
By Elizabeth N. Marks and Ernest P. Hodgkin. (One Text-figure.) .. .. 33-39
A Summary of the Atopomelinae (Acarina, Listrophoridae). By R. Domrow.
(Nine Text-figures.) aie kere Pantapeitaiel cl Me ta Vues tay Col wc true HH CALAIS | ease aie BR 40-64
Inheritance of Oil Characters in Hucalyptus. By L. D. Pryor and L. H. Bryant.
(Five Text-figures.) PRET WHOL OMMU AER ctu AWAY: ASO HUAI Lees ARIE a go ag IO 55-64

The Status of Nitrogen in the Hawkesbury Sandstone Soils and their Plant
Communities in the Sydney District. II. The Distribution and Circulation
of Nitrogen. By Nola J. Hannon, Linnean Macleay Fellow of the Society
in “BotanysAn Cl won LextSuress) icc saci tae uoupete te beccan te par mies) ie le Maser dete 65-85

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BL/WHOI LIBRARY

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