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Proceedings of the

Linnean Society
of New South Wales

Issued 5th November, 1965

Marine Biological Laboraioy
LIBRARY
DEC 2 7 1965
WOODS HOLE, MASS.

~ VOLUME 90
PART |
q No. 407

Registered at the General Post Office, Sydney, for transmission by post as a periodical

The Linnean Society of New South Wales

Founded 1874. Incorporated 1884

‘‘ For the cultivation and study of the science of Natural History in
all its branches ”

OFFICERS AND COUNCIL, 1965-66

President
D. T. Anderson, B.Se., Ph.D.

Vice-Presidents

Elizabeth C. Pope, M.Sc., C.M.Z.8.; G. P. Whitley, F.R.Z.S.; Professor J. M.
Vincent, D.Sc.Agr., Dip.Bact.; T. G. Vallance, B.Sec., Ph.D.

Hon. Treasurer
A. B. Walkom, D.Sc.

Hon. Secretaries
W. R. Browne, D.Sc., F.A.A.; A. B. Walkom, D.Sc.

Council

D. T. Anderson, B.Se., Ph.D. Professor 8. Smith-White, D.Sc.Agr.,
R. H. Anderson, B.Sc.Agr. F.A.A.
W. R. Browne, D.Se., F.A.A. tN. G. Stephenson, M.Sc., Ph.D.
R. C. Carolin, Ph.D., B.Sc., A.R.C.S. EK. LeG. ‘Troughton, O.M.ZS.,
S. J. Copland, M.Sc. F.R.Z.S.
*L. A. S. Johnson, B.Sc. T. G. Vallance, B.Sec., Ph.D.
Professor F. V. Mercer, B.Sc., Professor J. M. Vincent, D.Sc.Agr.,

Ph.D. Dip.Bact.
A. K. O’Gower, M.Se., Ph.D. A. B. Walkom, D.Sc.
Elizabeth C. Pope, M.Sc., C.M.Z.S. H.S. H. Wardlaw, D.Se., F. RB A.C.I.
7E. Shipp, Ph.D. G. P. Whitley, F.R.Z. S.

* Elected 26/5/1965 in place of Professor W. L. Waterhouse.
+ Elected 22/9/1965 in place of Professor I. A. Watson.
{ Elected 26/5/1965 in place of Professor B. J. Ralph.

Auditor
S. J. Rayment, F.C.A., Chartered Accountant

The Society’s headquarters are in Science House, 157 Gloucester Street, Sydney
N.S.W., Australia

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Proceedings of the
Linnean Society
of New South Wales

VOLUME 90
Nos. 407-409

CONTENTS OF PROCEEDINGS, VOLUME 90

PART 1 (No. 407).
(Issued 5th November, 1965)

(Presidential Address and Papers read March—April, 1965)

CONTENTS

Annual General Meeting
Review of year’s activities is A ahs AG ye 1
Elections 66 50 <3 45 3 ae an 36 ate 4
Obituary Notices 5
Balance sheets 7

ELIZABETH OC. POPE. Presidential Address. A review of Australian
and some Indomalayan Chthamalidae (Crustacea: Cirripedia).
(Plates i and ii) a a ails A ee Bs a st)

M. J. WHITTEN. Chromosome numbers in some Australian leafhoppers
(Homoptera Auchenorrhyncha). (Plate ili) .. = ae Sa (es

A. W. SWEENEY. The distribution of the Notonectidae bamuben: in
south-eastern Australia .. ste st oy : whe ste ee Ot

LYNNE BEDFORD. The histology and anatomy of the reproductive
system of the littoral gastropod, Bembicium nanum (Lamarck)
(Fam. Littorinidae) ae ae iy ae ag ae go OD

D. T. ANDERSON. The reproduction and early life-histories of the
gastropods, Notoacmaea vpetterdi (Ten.-Woods), Chiazacmaea
flammea (Quoy and Gaimard) and Patelloida alticostata (Angas)
(Fam. Acmaeidae) .. <4 se eis ide A bye .- 106

MARGARET SPENCER. Malaria in the D’Entrecasteaux Islands, Papua,
with particular reference to Anopheles farauti Laveran ES Boe all

MARGARET SPENCER. <A note on blood preferences of Anopheles
farauti.. Ey. a oe as on Bh <a os elas

PART 2 (No. 408).
(Issued 18th February, 1966)

(Papers read June—July, 1965)

CONTENTS

Y. M. UpADHYAYA and E. P. BAKER. Studies on the inheritance of
rust resistance in oats. III. Genetic diversity in the varieties
Landhafer, Santa Fe, Mutica Ukraine, Trispernia and Victoria
for crown rust resistance ..

R. BASDEN. The occurrence and composition of manna in Hucalyptus
and Angophora

B. P. Moorkr. Australian larval Carabidae of the subfamilies Harpalinae,
Licininae, Odacanthinae and Pentagonicinae (Coleoptera) ek

R. Domrow. Some laelapid mites of syndactylous marsupials ..

R. TUCKER. Comparative studies on the external acoustic meatus.
I. The morphology of the external ear of the echidna (Tachy-
glossus aculeatus). (Plates iv-v) .. : as ee Be

A. E. H. PEDDER. The Devonian tetracoral Haplothecia and new
Australian phacellophyllids. (Plate vi)

R. Domrow. Some mite parasites of Australian birds

N. E. Mitwarp. Development of the eggs and early larvae of the
Australian smelt, Retropinna semoni (Weber). (Plates vii-ix)

ANN M. Cameron. The first zoea of the soldier crab, Mictyris longicarpus
(Grapsoidea : Mictyridae) : ee ae

E. J. ANDERSON. Plant parasitic nematodes in fruit-tree nurseries of
New South Wales

J. M. Marruews. Diurnal variation in the release of pollen by Plantago
lanceolata L. ..

M. G. BRooKkEeR and G. CauGHLEY. The vertebrate fauna of ‘‘ Gilruth
Plains ”, south-west Queensland ..

Page

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190

218

222

225

231

238

PART 3 (No. 409).
(Issued 22nd July, 1966)

(Papers read September—November, 1965)

CONTENTS

D. T. ANDERSON. Further observations on the life histories of littoral
gastropods in New South Wales. (Plate x) a : ;

P. J. Dart and F. V. MERcER. Observations on the fine structure of
the meristem of root nodules from some annual legumes. (Plates
Xi-XXV) ; of ae Bi

A. J. T. WRIGHT. Cerioid Stringophyllidae (Tetracoralla) from Devonian
strata in the Mudgee district, New South Wales. (Plate xxvi)

GWENNETH J. HINDMARSH. An Stace ay of five species of
Bassia All. (Chenopodiaceae) : ;

Y. T. TcHAN and D. L. JAckson. Studies of nitrogen-fixing bacteria.
IX. Study of inoculation of wheat with Azotobacter in laboratory and
field experiments. (Plates xxvii-xxviil) : 33 ot

N. H. Lurie and I. A. WATSON. Studies on the genetic nature of resistance
to Puccinia graminis var. tritici in six varieties of common wheat

F. R. Hiaernson. The distribution of submerged aquatic angie
in the Tuggerah Lakes system was bs ae i

R. C. JANCEY. Numerical methods in taxonomy ..

R. C. JANcEY. An investigation of the ad ages (DC.) Benth.
(Leguminosae) (Plates xxix-xxx) . : sts “3

Abstract of Proceedings ..

List of Members ..

List of Plates

List of New Genera and New Species

Index

SYDNEY
PRINTED AND PUBLISHED FOR THE SOCIETY BY
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Seamer Street, ee Sydney

an
SOLD BY tHE SOCIETY

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ANNUAL GENERAL MEETING
31st MARCH, 1965

The Ninetieth Annual General Meeting was held in the Society’s Rooms,
Science House, 157 Gloucester Street, Sydney, on Wednesday, 31st March,
1965.

Miss Elizabeth C. Pope, President, occupied the chair.

The minutes of the Highty-ninth Annual General Meeting (25th March,
1964) were read and confirmed.

President’s Introductory Remarks

It is customary for a retiring president to review the Society’s activities
during his year of office but in this rather special year, the Society’s 90th one,
I would like, first of all, to take a brief look forward.

The Council has decided to present our publication, The Proceedings of
the Linnean Society of New South Wales, in a new and more modern format.
An increase in the size of type used and a number of editorial changes should
result in a pleasing layout and make the Proceedings easier to read.

Since 1950, when the annual subscription was raised from £1 1s. to £2 2s.,
costs of printing and distribution of the Proceedings have risen steeply. About
each second year the printer has notified us of the necessity for an increase
in printing charges. In 1950 the basic charge was £1 10s. per page and at
the beginning of 1965 this had risen to £4 10s. 9d. per page. It has been
suggested that the Council should consider a review of membership sub-
scriptions, and that the result of their deliberations be submitted to members
in time for any changes to come into effect with the introduction of dollar
currency in February, 1966.

The Council is also exploring the possibility of changing the Society’s
programme of activities to meet the needs and wishes of members. Attendances
at Ordinary Monthly Meetings have fallen off so greatly, that it is apparent
that this type of meeting is no longer what the members want. This change
in the needs of members is being experienced also by many societies abroad
—it is a world-wide change. Some groups are meeting the challenge by
organizing one or two symposia each year, others by activities of a different
nature, and you can rest assured that your in-coming Council will be forward-
looking and will aim to introduce new ideas to test the needs of members.
Your response to these innovations will determine what the Council finally
decides. It could be that our Society may become merely a vehicle for
publication of research, and provision of a research library service to membegcs.
It is for the members to support or criticize Council’s attempts to find what
changes should be made so as best to serve the needs of the majority of our
members.

Before reviewing the year’s activities I should like to express, on behalf
of all members, our thanks to that small and devoted group of hard-working
people who do so much to keep the Society’s affairs running smootbly, as it
were, on roller bearings: to Miss G. Allpress, our Assistant Secretary, who
continually watches over the welfare of the Society and its members and
implements so ably and faithfully the considerable volume of clerical work
and the working of the library and its loans and exchanges which the Council
and its executive pass on to her. Our debt of gratitude to our Joint Honorary
Secretaries, Dr. W. R. Browne and Dr. A. B. Walkom, to whom we owe the
smooth running of the Society’s affairs in the past year, continues to mount

A

2 PRESIDENTIAL ADDRESS

with each year that they serve in this honorary capacity. Dr. Walkom is
also Honorary Treasurer and Editor of The Proceedings, so that we owe him
thanks on this score also. We thank them both most sincerely for their
services to the Society in the past year. Finally I offer my thanks to my fellow
councillors for their help and support during the year and to our Auditor, Mr.
S. J. Rayment, F.C.A.

REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR

The Society’s Proceedings for 1964, Vol. 89, Part I was published in 1964
and Parts II and III in 1965. Vol. 89 consists of 398 pages, eight plates and
180 text-figures. An increase in the cost of blocks of about 73% took place
in 1964 and notification was received from the Society’s printers of an increase
of 10% in printing charges as from 1st January, 1965. An anonymous con-
tribution of £525 was received towards the cost of printing the Proceedings
for 1964. Information was received of the death of the Society’s agent in
London, Mr. David Nutt. Council decided not to appoint another agent. The
State Government increased its annual grant to the Society from £200 to £400,
on condition that 100 (instead of 60) copies of the Proceedings be made available
for governmental distribution. The total net return from the Society’s one-
third ownership of Science House for the year ended 31st August, 1964, was
£1,356 6s. 1d.

During the year twenty-five new members were added to the list, two
died, one resigned, and two were removed from the list of members. The
numerical strength of the Society at 1st March, 1965, was: Ordinary Members,
265; Life Members, 33 ; Corresponding Member, 1; total 299.

It is with regret that the deaths of Professor Charles Baehni and Miss Vera
Irwin Smith are recorded. (See pages 5-6 for obituary notices.)

A total of 27 papers was read at the Ordinary Monthly Meetings. Lec-
turettes were given at the following meetings: June, Reptile Collecting in New
Guinea, by Mr. H. G. Cogger; July, Some Aspects of Forestry in New South
Wales, by Mr. G. Baur; September, Chemistry and Insects, by Associate
Professor E. W. K. Cavill, and October, Biological Studies in East Africa, by
Mr. H. J. de S. Disney. Interesting discussions followed the lecturettes. <A
symposium on the Natural History of Kosciusko was held in April, the following
speakers taking part : Dr. W. R. Browne, Dr. R. C. Carolin, Mr. D. K. McAlpine
and Dr. D. F. McMichael. We are grateful to the lecturers and speakers for
their contributions to the interest of the meetings.

On 21st August, 1964, the fourth Sir William Macleay Memorial Lecture
was delivered in the Large Hall, Science House, Sydney, by Professor H. G.
Andrewartha, Department of Zoology, University of Adelaide, the title of the
Lecture being ‘‘ How Animals can live in Dry Places ”’.

On 15th February, 1965, the resignation from the Council of Professor
B. J. F. Ralph was accepted with great regret.

On 24th March, 1965, the resignation of Professor W. L. Waterhouse, as
a member of Council, tendered on account of continued ill health, was accepted
with great regret and the Council “ put on record its high appreciation of
Professor Waterhouse’s devoted service to the Society as a member of Council
since 1930 and as president in 1935”.

Library accessions from scientific institutions and societies on the exchange
list amounted to 2,222, compared with 2,174 and 1,997 in the years 1963 and
1962 respectively. The total number of borrowings of books and periodicals
by members and institutions was 263 for the year. Members and others con-
tinued to consult publications in the Society’s rooms, and books and periodicals
were made available for photographic copying. Council decided that, com-
mencing with Part II of the volume for 1964, all copies of the Proceedings as
issued should be despatched directly by post ; hitherto certain foreign exchanges

PRESIDENTIAL ADDRESS 3

have been sent through the International Exchange Bureau by courtesy of
the Public Library of New South Wales. The following requests for exchange
of publications were acceded to during the year: Staatliches Museum fur
Tierkunde, Dresden, East Germany; Leyden Museum of Natural History,
Leyden, Netherlands (Abstract of Proceedings) ; U.S.S.R. Academy of Sciences,
Leningrad (two additional copies of the Proceedings commencing 1965 in
exchange for its ‘‘ Oceanology’’); Societas Entomologica Helsingforsiensis,
Helsingfors, Finland (Entomological Reprints for ‘‘ Notulae Entomologicae ’’)
and National Institute of Science and Technology, Manila, Republic of the
Philippines. Council arranged for a set of the Proceedings from 1941 to be
sent to Manila by the Museum of Applied Arts and Sciences, Sydney, as a gift
to the National Institute of Science and Technology, to replace volumes destroyed
during World War II. The University, Louvain, Belgium, is now forwarding
‘“‘ La Cellule”’ in exchange for our Proceedings. The Beaudette Foundation
for Biological Research, Moss Landing, California, U.S.A., was removed from
the Society’s exchange list owing to the cessation of its publication ‘ Pacific
Naturalist’. Mr. K. A. Hindwood presented a reprint of his paper ‘‘ George
Raper: An Artist of the First Fleet ” (from J. and Proc. Roy. Aust. Hist. Soc.,
vol. 50, pt. 1, 1964) and Mr. K. Mair, Director of the Royal Botanic Gardens,
Sydney, forwarded a copy of a valuable report by Dr. Joyce W. Vickery on
her observations of the behaviour of exotic grasses used for stabilization in
disturbed areas of the Snowy Mountains, for inclusion in the Society’s library.

During the year the Society: (1) supported the Iluka Softwood Rain
Forest Protection Committee in its efforts to prevent destruction of the local
native fauna and flora through possible rutile mining ; (2) sponsored a Photo-
graphic Conservation Exhibition at the Australian Museum; and (3) was
represented by invitation at the inaugural meeting in Canberra, on 21st August,
of the Australian Conservation Foundation, whose Provisional Council includes
two members of the Society. Council also sent two delegates to the Annual
Conference of the Nature Conservation Council of New South Wales. Owing
to the number of questions relative to Nature Conservation brought before it
for consideration, Council set up a Committee to examine and advise on matters
coming under this head.

At the Annual General Meeting on 25th March, 1964, the President (Mr.
G. P. Whitley) unveiled a framed, enlarged photograph of Dr. A. B. Walkom
to mark his 75th birthday and his 45 years of continuous service to the Society.
By resolution of Council a copy of the photograph was printed in Part I of
the Proceedings for 1964.

A collection of framed photographs, watercolour drawings and maps, a
photograph of Alexander Walker Scott, and a watercolour drawing of Hvolvulus
alsinoides by Rev. Julian Tenison-Woods—all of which had long been in the
possession of the Society—were presented to the Mitchell Library, the Macleay
Museum, and the Royal Botanic Gardens, Sydney, respectively.

Council has been considering the advisability of modernizing the format
of the Society’s Proceedings, and an ad hoc Committee has already submitted
a report on this matter.

Tinnean Macleay Fellowships

In November, 1963, Mr. P. J. Dart, B.Sc.Agr., was reappointed to a
Fellowship in Plant Physiology for the year commencing 1st January, 1964.
He continued his studies on nodule fine structure within the different legume-
Rhizobium cross inoculation groups, and his examination of fine structure
changes in Mo-deficient nodules. He resigned his Fellowship in May, 1964,
to take up a C.8.I.R.O. Overseas Scholarship to study in Denmark and U.S.A.

In November, 1964, Mr. A. J. T. Wright, B.Sc., was appointed to a Linnean
Macleay Fellowship of the Society in Palaeontology, tenable for one year from

4 PRESIDENTIAL ADDRESS

1st January, 1965. Mr. Wright proposes to continue his studies of the Devonian
sediments and faunas of the Mudgee district and of the Capertee Valley area.
We offer him our best wishes for a successful year’s research work.

Linnean Macleay Lectureship in Microbiology

Dr. Y. T. Tchan, Reader in Agricultural Microbiology and Linnean Macleay
Lecturer in Microbiology, University of Sydney, reported on his work for the
year ending 31st December, 1964, as follows: Progress has been made on the
cytology of Azotobacter, particularly on the ultrafine structure of flagella and
internal membranous organelles. The interference of Ca in the availability
of trace elements in medium for culture of Bewerinckia has been investigated.
It was found that it is related to the pH of the medium. Progress has also
been made on the study of pesticides by algal method. The method proved
to be suitable for testing of herbicide in soil.

The Honorary Treasurer (Dr. A. B. Walkom) presented the balance sheets
for the year ending 28th February, 1965, duly signed by the Auditor, Mr. S.
J. Rayment, F.C.S., and his motion that they be received and adopted was
carried unanimously.

PRESIDENTIAL ADDRESS
A Review of Australian and some Indomalayan Chthamalidae
(Crustacea : Cirripedia)

The review mentioned genera and species of the Family Chthamalidae
recorded from Australia and certain collections taken during the Dutch
Snellius Expedition of 1929-30, but dealt particularly with species occurring
in the upper half of the intertidal zone of the shores round Australia. The
possible significance of certain morphological characters as adaptations for
feeding at different intertidal hsights was discussed from the point of view of
the systematics of the Chthamalidae in general. (For full text see pp. 10 ef seq.)

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

President: _D. T. Anderson, B:Se., Ph.D.

Members of Council: R. H. Anderson, B.Sc.Agr. ; Miss Elizabeth C. Pope,
M.Sc., C.M.Z.S.; E. Le G. Troughton, C.M.Z.S., F.R.Z.S.; T. G. Vallance,
B.Se., Ph.D. ; J. M. Vincent, D.Sc.Agr., Dip. Bact. ; and G. P. Whitley, F.R.Z.S.

Auditor: 8S. J. Rayment, F.C.A.
The President then installed Dr. D. T. Anderson as President.
A cordial vote of thanks to the retiring President was carried by acclamation.

OBITUARY NOTICES

CHARLES BAEHNI

Professor CHARLES BAEHNI who died suddenly at Geneva on 23rd January,
1964, was elected to membership of this Society in April, 1952. For many
years he played a leading part in the life of his native city, Geneva, where he
received his early education and where he was destined to occupy high office.
Swiss botany has sustained a grievous loss. At the University his first botanical
investigations were carried out under his eminent predecessor, Prof. Robert
Chodat, and in 1932 he was awarded the degree of Dr. es Sc. In the same
year he was appointed as assistant at the Conservatoire et Jardin Botaniques.
During 1934-35 he studied in the Botanical Department of the Field Museum
of Chicago. He was conservator (1941) and later director (1943-1964) of the
Botanic Garden at Geneva. While director of the Gardens he was also
professor of systematic botany in the University of Geneva, and for a time
in the University of Lausanne. He published more than a hundred scientific
papers, his chief interest being in the Sapotaceae, although his contributions
extended to other families, including the Ulmaceae, Lacistemaceae and
Violaceae. He was a member of the Editorial Committee of the International
Code of Botanical Nomenclature from 1950.

A more detailed obituary notice appeared in Nature, Vol. 202, No. 4928,
April 11, 1964, p. 132.

VERA ADELAIDE IRWIN-SMITH

Miss VERA ADELAIDE I[RWIN-SMITH, B.Sc., F.L.S., who died on 2nd April,
1964, was born in Albury, New South Wales, in November, 1885. She was
the daughter of Irwin Smith, a surveyor in the Riverina district, and Adelaide
Smith, formerly Adelaide Riches. She was educated
at Glenair Ladies’ College in Albury. When about
18 years of age she went for a world tour witb her paternal
grandparents, and this trip evidently instilled in her a
love of travel which never left her. About 1910 Mr.
Smith moved to Sydney to live, probably for the benefit
of the children’s education; they lived at first at
Northwood, then at Valentia Street, Woolwich. Miss
Irwin-Smith had been told that her health would not
permit her to undertake a University course so she
took a course in architecture at the Sydney Technical
College, Ultimo. Having found that she could manage
that, she decided to do a science course at the University
of Sydney. She commenced the science course in 1912
and, on medical advice, took two years to complete the
first year. She graduated B.Sc. in 1916 with First
Class Honours in Botany and Second Class Honours in
Zoology. She was awarded a University Science
Research Scholarship which she held for the years
1916, 1917 and 1918. Miss Irwin-Smith was elected
to membership of the Society in 1916. From 1919 to 1923 she held a Linnean
Macleay Fellowship of the Society in Zoology, being the first woman to be
appointed to a Fellowship. She contributed thirteen papers to the Proceedings.

Her first published paper dealt with some new Chaetosomatidae. Recent
ecological studies on the minute animals inhabiting the interspaces between
sand grains on the littoral of oceans and lakes has led to a marked revival of

Miss Irwin-Smith

6 OBITUARY NOTICES

interest in many of the small and more obscure groups of animals. Among
these are the free-living worms of the group Chaetosomatidae, Australian
specimens of which were described in some detail in 1918 by Miss Irwin-Smith
in the Society’s Proceedings. Lately many requests have been received from
abroad for reprints of her paper, as workers are only now beginning to appreciate
the functional significance of many of the minute structures she described in
local Chaetosomes. In all she described four new species in two genera, one
of which was new.

In 1920 she was Senate representative on the Board of Directors of the
Sydney University Women’s Union.

During her tenure of a Linnean Macleay Fellowship she continued her
study of Neimatodes and allied worms, commenced under her Research Scholar-
ship. She had also undertaken the preparation of a report on the collection
of parasitic Nematodes brought back by the Australasian Antarctic Expedition.
She studied further collections of Nematodes parasitic on Australian hosts, and
undertook a series of studies of life-histories of Australian Diptera Brachycera.
Of the latter, she investigated Stratiomyiidae, dealing first with Metoponia
rubriceps, followed by a study of the Asilidae. Her studies of Nematodes
included the Nematode parasites of the domestic Pigeon, and also the genus
Physaloptera with special reference to those parasitic in reptiles. In
addition, she made large collections of dipterous larvae, with the object of
working out further life-histories, and much interesting information was
accumulated. In 1923 she went to England to continue her research work
and worked for a time at the Molteno Institute in Cambridge, but family
circumstances compelled her to return to Australia. After her return she
continued to collect and, in association with the late Mr. Luke Gallard of Epping,
Fruit Inspector in the Department of Agriculture, larvae of at least seven
families of Diptera were found. Many of these were reared to maturity and
records were kept of their development. Her father died shortly after their
return from England, and for some years she cared for her mother who became
an invalid. Later, her own poor health prevented her from completing any
further research work. She spent the latter part of her life in a convalescent
home. Miss Irwin-Smith bequeathed in her will the sum of approximately
forty thousand pounds to the Australian Academy of Science, Canberra, A.C.T.

bk

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PRESIDENTIAL ADDRESS

A REVIEW OF AUSTRALIAN AND SOME INDOMALAYAN
CHTHAMALIDAE (CRUSTACEHA : CIRRIPEDIA)

ELIZABETH C. POPE
Curator of Worms, Echinoderms, ete., The Australian Museum*

(Plates i and ii)
[Delivered 31st March, 1965]

Synopsis
All species of intertidal barnacles of the Family Chthamalidae known from Australian shores
are reviewed and discussed with reference to their geographical distributions and ecological
occurrence. Hight species in four genera are included but none is new to science. Considerable
extensions of the ranges of Chthamalus intertextus to New Guinea and of Chthamalus malayensis,
C. caudatus and C. withersi along the northern coast of Australia are recorded.

There are five tropical species in Australia in two genera, of which four are included in
genus Chthamalus and one in Octomeris. Only three species of the family occur m the temperate
southern half of the contiment, but each belongs to a different genus. Thus Catophragmus,
Chthamalus and Chamaesipho are represented. A second species of Chamaesipho occurs in New
Zealand, but these two species are the sole representatives of the Family Chthamalidae in that
region.

Certain species of Chthamalus (C. witherst and C. caudatus) and Octomeris brunnea possess
structures on their mouth parts and cirri which are believed to be adaptations for capturing
particulate food over a relatively short period and in seas showing only slight to moderate
degrees of water movement other than tidal flow except, of course, during storms. The
remaining five species from three genera have armouries of elaborate hooks or grapples well
adapted to the holding of food particles, in the ebb and flow of surf or other considerable water
movements. When these hooks, grapples and other food-holding adaptations are catalogued
for each species, it will be found that the species constantly subjected to roughest water move-
ments possess a greater variety of such adaptations than species from calmer situations.
However, many more Chthamalid species will have to be examined before any conclusions
about the phylogenetic significance of these structures, if any, can be drawn.

INTRODUCTION

The following genera have hitherto been included by general consent in
the Family Chthamalidae: Catophragmus, Chionelasmus, Octomeris, Chtham-
alus, Chamaesipho, Pachylasma and its closely related genus Hexelasma, and
the fossil genus Tessarelasma. Of these, four, namely Catophragmus, Chtham-
alus, Chamaesipho and Octomeris, are familiar intertidal dwellers on Australian
Shores, and Pachylasma and Hexelasma are represented in the deeper water
fauna off our coasts.

Of this deeper water group records are scattered, few specimens are known
of each species, and collecting has never been systematically carried out. Just
a few specimens have been taken by deep sea expeditions from time to time,
or by chance trawling operations, here and there. In these circumstances,
little is to be gained by reviewing them, until additional and adequate material
is collected. It is proposed here, merely to list the species occurring in
Australian waters and to pass on to a more detailed review of the extremely
common intertidal species.

The list of deep-water Chthamalidae occurring near Australia is as
follows: Pachylasma scutistriata Broch, 1922, eastern slopes of Bass Strait,
Tasman Sea, 70-160 fathoms ; P. integrirostrum Broch, 1931, Kei Islands and

* By permission of The Trustees.

PROCEEDINGS OF THE LINNEAN SoctetTy or New SoutH Watss, Vol. 90, Part 1

ELIZABETH C. POPE 11

Amboina, 140 metres; P. aurantiacum Darwin, 1854, ‘‘ New South Wales”
ie., east coast of Australia, ‘“‘ apparently from deep water’; Hexelasma
velutinum Hoek, 1913, NW. of Timor and Kei Islands, 245-390 metres; H.
arafurae Hoek, 1913, Arafura Sea, 560 metres.

Of the remaining genera listed in the Family Chthamalidae, Chionelasmus
(a deeper water form) has not been taken from seas off Australia and the
fossil genus Tessarelasma has been recorded no nearer to Australia than the
Indian subcontinent where it occurs embedded in rocks of the Surma Series
(Lower Pliocene) at a depth of approx. 2,000 feet, along the Arakan Coast
region of Eastern Bengal, Pakistan.

Terminology used in this review is defined in Pilsbry (1916).

Broadly speaking Australian Chthamalidae may be divided into two
groups differing markedly in their ecology, their abundance, and also in several
aspects of the basic arrangement of the plates forming their shells. The one
group, comprising the genera Chthamalus, Octomeris, Catophragmus and Chamae-
sipho, inhabit rocks or other hard substrates in the upper half of the intertidal
zone. Most of them thus spend more time surrounded by air than they do
under water. In fact, some individuals of Chthamalus antennatus in the
temperate zone or of Chthamalus withersi in the tropics in Australia spend
approximately 95-98% of their time exposed to air while Chthamalus
malayensis is exposed from 30-90% of the time*.

This is really an extraordinary state of affairs, when one considers that
they are marine animals, living cemented to the substrate and therefore
entirely dependent on the sea to bring them their food (for they are filter
feeders), and to carry away and distribute their planktonic young after they
have been brooded for a time. Presumably also fertilization of eggs can only
take place during periods when the barnacles are covered by seawater. Their
hermaphroditism must also have played a tremendous part in their successful
colonization of the high shore rocks. However, it is not intended to enlarge
on the mechanisms of reproduction in the Chthamalidae in this review. Also
no mention will be made of their respiratory needs since it is obvious that
plenty atmospheric oxygen is available to them. Rather their main metabolic
problems would appear to be concerned with obtaining sufficient food and the
need to conserve moisture during the long periods of subjection to the blazing
sunlight and constant high temperatures encountered in their environment.

As mentioned above, the other ecological group within the family inhabits
the deeper waters of the ocean which, from the point of view of filter-feeding
organisms, presents fewer problems. While it is not intended to review this
latter group, certain fascinating questions about their mode of life arise in
one’s mind. Why, for instance, has the genus Hexrelasma, in the course of its
evolution from an 8-valved ancestral type, reduced its shell plates to 6, by
the elimination, through amalgamation of the rostro-lateral pair of compart-
ments with the rostrum, whereas in the 6-valved intertidal genus Chthamalus,
the reduction from 8 to 6 has been achieved by loss of the carino-lateral pair
of plates—at the opposite end of the animal? Another enigma associated
with the Chthamalidae is the apparent failure of this otherwise evolutionarily
successful family to invade and colonize the lower half of the intertidal zone
of the shallower seas of the continental shelf, where the other sessile barnacle
family, the Balanidae, has become so successfully established. Whatever the
reason for their exclusion from the lower shore, the fact remains that some
genera of the Chthamalidae have, in the course of their evolution, become
highly adapted to living as marine animals out of water for the greater part
of the time. Can we by studying their field occurrence, the geographical and
vertical range of the various species, and their morphology learn anything

* From unpublished M.Sc. thesis by Miss Judy Bryan of Townsville University College,
Queensland.

12 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

about the adaptations that make for success in the high shore environment ?
The present study is an attempt to make a slight contribution in this field and
to arouse interest in these animals, so suitable for experimental studies.

Firstly, however, it is necessary to review the nature and composition of
the Australian Chthamalid fauna, for no comprehensive account of the family
has appeared since Charles Darwin’s Ray Society Monograph of the Sub-Class
Cirripedia was published in 1854. It is also apparent that the collections made
for, and sent to, Charles Darwin must have completely neglected mainland
Queensland in particular, and the tropical half of the continent in general,
for his work fails to record Octomeris brunnea and three species of Chthamalus
(which at that period were undescribed) although Barrier Reef species in
Family Balanidae were well represented.

The intertidal surveys made by Endean e¢t al. (1956, a and 6b) revealed for
the first time how extensively Indomalayan species of Chthamalidae range
along the eastern coast of Queensland and the prominent role they play in the
pattern of intertidal zonation. As a direct consequence of their work and
because of the zoogeographical importance of this group, collecting of cirripedes
in tropical Australia was extended by the author to the northern and western
coasts of the continent and the now extensive collections which resulted are
housed in the Australian Museum, in Sydney. It is chiefly on these collections
that the present study is based. The author has also made many field trips
over the years, studying the field occurrence of barnacles along 7,000 miles
of coastline.

The practice followed in this study has been to base descriptions on as
large a population of each species as possible, using many dissections and also
notes made of their field occurrence in varying environments, for slight changes
in environmental conditions have been found to cause marked differences in
the general appearance of Chthamalid shell plates.

Samples have been examined, wherever possible, in batches collected at
approximately hundred mile intervals throughout the Australian ranges of
each species. In most cases large Australian individuals in any one species
are aS much as 1$ to 2 times bigger than any specimen previously recorded
and it soon became apparent that the original descriptions of several species
had been based on fairly juvenile material—the original authors having been.
misled into the belief that their specimens were full grown by the fact that
they were frequently brooding developing larvae. Pilsbry’s description of
Chthamalus witherst (1916) and Darwin’s of Octomeris brunnea (1854) are two
such cases. If the descriptions of Australian specimens given below differ
markedly from the original accounts this is often the reason, for the larger a
barnacle grows the more rugged its shell becomes and the more characters
appear in the shell and in the internal soft parts.

The following systematic account includes all known intertidal species
of the Family Chthamalidae from Australia and New Guinea. They comprise
eight species divided between four genera of which only one genus (Chthamalus)
ranges into both the tropical and temperate areas of Australia and no one
species is common to both.

CHTHAMALIDAB OF AUSTRALIAN INTERTIDAL ZONE

Chthamalidae characteristic of the intertidal zone of temperate Australian
shores are Chthamalus antennatus, Chamaestpho columna and Catophragmus
polymerus and they are listed in the order in which they occur, from the high
shore down to mid-tide level. All three are species forming prominent bands
in the pattern of intertidal zonation in SE. Australia and all are adapted to
living on rocks which are usually subject to regular pounding by surf. To
a certain extent their vertical ranges overlap one another, so that C. antennatus
and Chamaesipho columna may be found together, likewise Chamaesipho and

ELIZABETH C. POPE 13

Catophragmus may occur in the one area, but only in areas subjected to especially
rough seas could C. antennatus and Catophragmus ever occur together in a
circumscribed area on the shore, for the latter could not survive for long on
the high level of the rocks frequented by C. antennatus.

Tropical species of the family represented on the Australian mainland are
Chthamalus withersi and Chthamalus malayensis on the upper surface of rocks,
with Chthamalus caudatus and Octomeris brunnea occurring either under boulders
or in shaded areas among boulders or under overhangs of rock. An area of
boulders offering the type of habitat frequented by C. caudatus and O. brunnea
is shown in Plate u, figure 4, and unless such areas are rather deliberately
searched, their occurrence in a locality can easily be overlooked. This probably
accounts for the few and scattered references to them in literature and it is
likely that they will be found to range more widely through the Indonesian
and Philippine Islands when systematic collecting is carried out in such places.

On the rocky shores of Queensland, Endean, Kenny and Stephenson (1956)
found a certain degree of interspecific competition between C. withersi and C.
malayensis, the former being favoured in areas subjected to wider fluctuations
in salinity and in turbid waters, and malayensis tending to be favoured by
increased wave action (i.e. less turbid areas) and more stable salinities. Thus
C. withersi invades the mangrove swamps and settles on the trees near the
mouth of coastal streams or colonizes wharf piles in river mouths and, indeed,
can occur in these habitats in great numbers, as shown in Plate ui, figures 2
and 5. On rocky reefs, fronting the more open waters, where turbidity is not
too high, C. malayensis becomes dominant intertidally, as shown in Plate ii,
figures 3 and 6.

Where these two species occur together, withersi occupies the higher levels,
ranging from the high water mark of spring tides down to high water neap
tide level, whereas C. malayensis ranges from approximately mean high water
mark down to high water mark of neap tides. Their vertical ranges thus
overlap to a certain extent and their distributions on any particular reef will
depend partly on the substrate and partly on the particular microclimates
offering within its area. Their occurrence in Queensland has been discussed
by Endean ef al. (1956, both papers). However, along the northern coast of
Western Australia, the behaviour of C. malayensis apperently differs, judging
by collections sent to the author. West of Darwin, Northern Territory, no
C. withersi have been taken but C. malayensis and C. caudatus colonize the
rocks between the same relative tidal levels as in Queensland but, owing to
the enormously greater tidal range (33 feet), the belt of intertidal rocks on
which they settle is, of course, much wider. In the absence of withersi in the
west, CO. malayensis and C. caudatus may colonize the roots and trunks of
mangroves. Records of Octomeris brunnea also occur on a similar substrate
in the New Hebrides and the Santa Cruz Island Group and would seem to
indicate either the needs of barnacles in the high tropics to find some degree
of shelter from direct sunlight and/or a lack of rocky substrates on which to
settle, so that they are driven to colonize tree trunks. The mangroves in these
areas occur, of course, on the open coastline in the high intertidal zone and
are not confined to the banks of inlets and rivers, where reduced salinities
obtain throughout the year.

Chthamalus malayensis and C. intertextus are the two species in tropical
Australia occurring where the water is consistently rougher. In general, how-
ever, at least inside the shelter of the rampart of the Great Barrier Reef of
Queensland, much of the mainland coast of Queensland must be judged as
relatively little exposed to wave action (except during cyclones).

Octomeris brunnea and C. caudatus may occur together in the one locality
since they occupy overlapping vertical ranges and tend to settle in shaded
areas (in Queensland, at least). The former species ranges from the level of

14 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

mean high water to the level of high water neap tides, while C. caudatus ranges
from the same upper level but extends lower down the shore to approximately
mean low water mark.

The remaining tropical species included in this account, C. intertextus,
has not yet been recorded farther south than the coast of Papua, not far from
Port Moresby, and although it occurs in the New Hebrides and Fiji it has not
been taken in New Caledonia. Some ecological data of its occurrence and
association with C. malayensis and O. brunnea in the Riu Kiu Islands are
recorded by Tokioka (1953) and Utinomi (1954) but no further data have been
supplied with the New Guinea material, other than the statement that it occurs
on rocks in the mid-tidal zone and C. malayensis occurs in the same locality.
It apparently is attached both on the open faces of rocks and on shaded areas
and under overhangs.

Systematic Account

In general, Australian members of this family have rather drably coloured
shells, generally coloured in varying tones of dirty grey, or greenish-grey where
erosion discloses underlying layers of horny lamina (in the case of Chthamalus
caudatus of a bright yellow tone) or where dark corium shows as a Series of
spots on the upper surface of greatly eroded shells. The only brightly coloured
member of the family is C. intertextus and then only when erosion reveals the
deep violet layers of shell laid down internally, during secondary calcification.

The practice has been followed of giving a table of measurements of a
series of barnacles, representative of the population of each species, since
maximum and minimum sizes mean little, when one is trying to make an
identification. They are given only where Australian material differed con-
siderably in size from previously described specimens.

No new species have been discovered in northern Australia. Those not
described by Darwin were described by Henry Pilsbry (1916) in another classic
monograph of Cirriped literature. Most of them range widely throughout the
Indomalayan Peninsula and Indonesia or the Western Pacific islands and it
was from there they were first described.

The following section includes descriptions of all intertidal species of the
Family Chthamalidae known to occur on Australian shores :—

Order THORACICA
Sub-order BALANOMORPHA
Family CHTHAMALIDAE

Genus CATOPHRAGMUS G. B. Sowerby, 1826

Sowerby, G. B. (1826) Catophragmus imbricatus, type of genus.—Darwin,
C., 1854; Pilsbry, H. A., 1916; Broch, H., 1922; Withers, T. H., 1935.

The genus Catophragmus was originally erected by Sowerby for C. imbri-
catus from Antigua in the West Indies.

It now comprises only three species—C. imbricatus Sowerby (1826) from
Bermuda and the West Indies, C. polymerus Darwin (1854) from Australia,
with a very doubtful record from ‘‘ Table Bay” (? South Africa) by Gruvel
in his 1905 Monograph, and C. pilsbryi Broch (1922) from Tobago, Panama.

Pilsbry (1907) described a fourth species, C. darwini, from several frag-
mentary specimens. However, he was unable to determine the number of plates
in the inner whorl of his species and guessed that it would be eight. In a later
work (1911) he expressed the opinion that, when whole specimens of C. darwini
were found, they might indicate the need for a new genus and he felt strongly
enough in the matter to suggest a name for the new genus Chionelasmus which,
meanwhile, he used as a subgenus of Catophragmus. When complete specimens

=

ELIZABETH C. POPE 15

of darwint came into the hands of Nilsson-Cantell (1928) and were found to
have only six plates in the inner shell whorl, he removed it from Catophragmus
and used Pilsbry’s proposed name, Chionelasmus, as the new generic name for
it. Chionelasmus differs from Catophragmus in the number of shell plates in
the inner whorl—six and eight respectively—and in the number of whorls of
supplementary plates—one in Chionelasmus and several in Catophragmus. The
genus Catophragmus is an intertidal one, whereas Chionelasmus is a deeper-
water species.

Pilsbry (1916) divided Catophragmus into three sub-genera Catophragmus,
Catomerus and Chionelasmus. This last sub-genus has now been elevated to
generic rank and the other two are distinguished from one another by the
presence in sub-genus Catophragmus of a caudal appendage and its absence
in sub-genus Catomerus. The Australian species thus falls into the sub-genus
Catomerus, while C. imbricatus and C. pilsbryi fall into sub-genus Catophragmus,
the former having a minute caudal appendage, fide Darwin, and the latter a
well-developed one, almost as long as the protopodite of cirrus VI, fide Broch.

In 1935 Withers erected another sub-genus within Catophragmus, sub-
genus Pachydiadema to accommodate his newly-described fossil species C.
(Pachydiadema) cretaceum from Ifo in Sweden. In the fossil species which was
described from a series of isolated plates, considerable differences in the structure
of the opercular valves and the absence of teeth on the basal edge of the shell
plates separated it from other species and sub-genera. However, it most
resembles the sub-genus Catomerus except in the opercular plates which are
close to those of pedunculate cirripedes, whereas in C. polymerus they are
clearly of the balanomorph type. The interest in C. (P.) cretaceum lies in the
fact that it is, to quote Withers, ‘‘ The earliest form, and the only Mesozoic
form, which can be regarded as belonging to the Balanomorpha, and shows
clearly its origin from the pedunculate stock. It is possibly the ancestor of
Catophragmus (Catomerus), which in turn is the most primitive member of the
Balanomorpha ”’.

Catophragmus (Catomerus) polymerus is the only species of the genus in
Australia and is easily distinguishable from other local intertidal barnacles by
the supplementary whorls of shell plates.

This species has a southern distribution in Australia and, in view of this,
it is perplexing to find that Broch (1927) regards C. polymerus as a representative
of the northerly intertidal zone of Australia. Guiler (1952), quoting P. H.
Fischer (1940), also lists this species as representing, in Tasmania, the “ relic
of a tropical fauna ’’, whereas it is endemic in temperate Australia.

The phylogenetic interest of Catophragmus has been discussed by Darwin
(1854), Withers (1928 and 1935) and nothing need be added to their statements.
The resemblance between a C. polymerus growing in its tall, narrow form
attached to the mussel Branchidontes rostratus and a short stalked species of
the Lepadormorph genus Mitella is most marked.

The world distribution of living species of this genus shows two isolated
groups with C. polymerus in Australia and the other two species in the Caribbean
Sea and Bermuda in the Atlantic. The fossil species C. (Pachydiadema)
cretaceum came from Ifo, Sweden, in Cretaceous rocks. The finding of this last
species does not alter Pilsbry’s and later Withers’s belief that the recent species
of Catophragmus represent survivors of an ancient genus once widespread in
its distribution.

No key to species of the genus Catophragmus is considered necessary as
Broch (1922, p. 301) makes the differences between the species quite clear.
These differences depend on obvious characters, such as the presence or absence
of caudal appendages, the sculpturing, and number of the supplementary plates
round the main whorl.

16 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

CATOPHRAGMUS POLYMERUS Darwin, 1854
(Plate i, figure 2; Text-figures 1,a,b; 2,a)

Catophragmus polymerus Darwin, 1854 ; Gruvel, 1903, 1905 ; Pilsbry, 1916 ;
Broch, 1922, 1927; Nilsson-Cantell, 1926; Pope, 1945; Dakin et al., 1948,
1952 ; Bennett and Pope, 1953, 1960; Endean, Kenny and Stephenson, 1956 ;
Womersley and Edmonds, 1958; Wisely and Blick, 1964.

Catophragmus polymerus, known as the “ surf barnacle”’, occurs in a well
developed horizontal band in the barnacle zone on intertidal rocks along the
south-east quarter of the Australian coastline.

In the original description Darwin lists Yetraclhita purpurascens among
other barnacles associated with Catophragmus and T. purpurascens does indeed
often occur in the same locality but is cryptic in habit, and not in an exactly
similar niche as the other species listed, e.g. Balanus nigrescens, Chthamalus
antennatus and Chamaesipho columna. These all attach to the upper surface
of the rocks in the barnacle zone. On the other hand, Tetraclita rosea occurs
in large numbers among the Catophragmus along the New South Wales coast
and it is felt that it may be the species Darwin intended to list as an associate
of OC. polymerus or, at least, it should be added to the list he gave.

Tetraclita rosea is favoured by warmer conditions and is able to survive
at a slightly higher shore level in New South Wales than C. polymerus. The
latter, on the other hand, is a southern species that is favoured by cooler con-
ditions. As a result, towards the northern, warmer end of their joint range,
notably from Ballina northwards, C. polymerus is greatly outnumbered by T.
rosea and, just north of the New South Wales border, in Queensland, 7. rosea
is the sole barnacle in this shore zone. Again, towards the south of New South
Wales, where the environment becomes progressively cooler, the numbers of
T. rosea fall off and C. polymerus becomes dominant. This tendency was noted
between Ulladulla and Batemar’s Bay. Farther south, at Mallacoota Inlet in
Victoria, 7. rosea is scarce and by Wilson’s Promontory it has disappeared
from the rocks, so that from there southwards C. polymerus competes for
settlement space with the mussel, Brachidontes rostratus, rather than with
another species of cirripede.

Local C. polymerus, from Sydney, have been shown to breed throughout
the year but peak breeding periods occur in winter and early spring (Wisely
and Blick, 1964).

Structure and Appearance of Shell

Darwin’s very detailed account of the shell of C. polymerus makes it
unnecessary to add more than a few comments on the variations seen.

Scutum and Tergum

There is some variation in the shapes of the opercular valves according
to the age of the barnacle and the degree of erosion it has experienced, but
their structures are basically similar to those shown in Darwin (1854, Plate
XX, fig. 4, e). The placement of the prominent articular ridge of the scutum
may vary from a central position on the tergal margin to a position above this
point—the corresponding furrow on the tergum varying in position in con-
sequence. Both tergum and scutum may be more nearly triangular in outline
and the tergum is generally somewhat bent about an axis running from the
apex to the basi-scutal tip of the valve, so that its two sides are inclined to one
another almost at right angles. The number of crests for attachment of the
tergal depressor muscles may be of the order of 13-15 in larger individuals.

Soft Body
The body of CU. polymerus occupies the space immediately below the
opercular plates only. The rest of the area of the base between the body and

ELIZABETH C. POPE IZ

the outer wall of the shell is occupied by the bases of the successive whorls of
supplementary plates and the enclosed spaces filled with corium. This
arrangement produces a structure well adapted to withstand severe battering
by surf. The basis of the shell is membranous and fairly tough and covers
the whole area below the shell. The milky white areas of the basis were tested
for the presence of calcium carbonate but none was present. The depressor
muscles of the tergum are very large and occupy approximately one-third of
the space inside the shell.

In freshly preserved C. polymerus the edges of the tergo-scutal flaps are
black. The body itself shows a marked contrast in the colouring of the cirri.
The first two pairs are a dark greyish-purple from pedicel to the tips of the
rami and bear thick bunches of comparatively long, light-coloured bristles on
each segment. Cirri III—-VI, on the other hand, have dark pedicels and as a
rule the rami are much lighter, varying from cream, with a few regular dark
patches, to light grey or fawn. Each segment has concentrations of dark
pigment cells round the bases of the spines. An occasional specimen has been
found with cirri III-VI having the outer third of the rami darker in colour
like the pedicels.

Trophi

The shapes of the various mouth parts vary slightly from those depicted
by Nilsson-Cantell (1926, text-fig. 2) but not significantly enough to warrant
new illustrations.

The main difference noted is, however, considered important since it
involves the setation of the trophi. In general, there are many more hairs
than are shown in his drawing, for C. polymerus is an extremely hirsute species.
The labrum is bullate and has a wide, deep groove anteriorly carrying along its
margins both hairs and a few small teeth (seen only under high magnifications
and from certain angles). The setae along the upper border of the palp are
so thickly bunched they resemble a brush. The mandible has three main teeth
and a coarsely pectinated lower angle in which the longest spines are at the
tip of the jaw. No secondary pectination was seen on teeth 1-3 as shown by
Nilsson-Cantell. Maailla I has two distinct notches and a double prominence
forming its lower angle. These divide the spines on the anterior border into
a number of distinct groups. The two upper pairs of spines are the largest.
Just below them, and above the first notch, are 3-4 pairs of considerably
smaller spines. Below this notch are 6—7 pairs of medium-sized spines which
arise from the anterior cutting edge between the upper and lower notches.
The spines on the double prominences of the lower corner of the jaw are much
smaller and comparable in size to those just above the first notch. The spines
on the upper prominence tend to point downwards, whereas those on the lower
one are directed horizontally. Mazilla II is somewhat pear-shaped, with a
notch in the upper half of the free edge. The notch is free of bristles but
numerous longish spines occur along the rest of the border. Those of the
ventral surface are about twice as long as those above the notch.

Cirri

Cirri I and If are much shorter than the remaining pairs and bear many
bristles on each segment of their rami. Both have unequal rami with the
anterior one exceeding the posterior one in length. The longer ramus is notably
broader than its fellow. The numbers of segments in the rami of cirrus I vary
but they are of the order of eleven in the anterior ramus and 7 or 8 in the
posterior one. The terminal segments, as well as several other segments, are
furnished with pinnate spines in which the central shaft bears alternating fine
side hairs (Text-fig. 1, b). These may be the barbed spines to which Darwin
(1854, p. 490) and Nilsson-Cantell (1926) refer but should not be confused with
the much stouter serrated spines of cirrus II (Text-fig. 1a). Cirrus IT has

)YD

18 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

its anterior ramus longer by about 3 of its segments than the posterior one,
the actual segmental numbers being of the order of 9-10 (anterior) and 8
(posterior). The 3 or 4 terminal segments of both rami of this cirrus carry
a number of very stout and coarsely pectinated spines in addition to the normal
and pinnate longer spines. These last tend to be more concentrated in segments
below the tips of the rami where the barbed stout spines are absent or few in
number. Cirri IJJ-VI are similar to one another in shape and the rami are
subequal. The numbers of pairs of major spines on each of their segments
is not always 5, as described by Darwin, but varies from segment to segment,

Fig. 1. Shapes of some stout spies and pinnate setae in intertidal Chthamalidae. (a-b), Cato-
phragmus polymerus. 'Tip of anterior ramus of cirrus II showimg stout serrated spines among
pimnate setae (a), while in (b) a pinnate seta is more highly magnified to show side hairs ;
(c), Octomeris brunnea, a stouter pinnate seta from cirrus II; (d-e), pmnate setae from terminal
segments of cirrus II in (d) Chthamalus withers and (e) Chthamalus caudatus ; (f-g), stout grapple-
spines from distal segments of cirrus II of Chthamalus intertextus (f) and of Chamaesipho columna (q) =
(h-2), stout, toothed spines from terminal segments of cirrus II m Chthamalus antennatus (h) and
in Chthamalus malayensis (1). (Spimes m view (f) seen from front view, while that in view (q) is.
seen in side view. Their structures are somewhat similar.)
F. J. Beeman del.

ramus to ramus, and cirrus to cirrus, even in the one individual. In a specimen
from Sydney cirrus III had four pairs on most segments of its rami; cirrus TV
had four pairs on its outer ramus and five on the inner one ; cirrus V was similar
to IV, while the spine pairs varied from four to five per segment on the outer
ramus while the inner ones had four pairs of major spines. In another barnacle
there were five pairs of spines on the segments of the inner ramus of cirri
IV-VI, while the outer ones had four. There is, however, one constant spine
character in all cirri from III to VI—namely, the presence of a thick brush-
like tuft of small spines of variable length, situated centrally on each segment
between the double row of major spines (Text-fig. 2, a). This character is
shared with the pedunculate species of the genus Mitella but is not present
in other Australian chthamalids.

ELIZABETH C. POPE 19

The penis is comparatively long, and towards its base is marked by a series
of narrow, dark rings and has a small tuft of hairs near its tip. There is no
caudal appendage.

Habitat

The preference shown by C. polymerus for areas of rock exposed to maximal
wave action has been indicated above. It grows on rocks just above the
distinctive Galeolaria Worm (Serpulid) Zone, in the upper half of the intertidal
zone, above mid-tide level, along with another so-called Surf Barnacle, Tetra-
clita rosea (Family Balanidae) but in Victoria and Tasmania and along the
mainland coast westwards from Bass Strait, C. polymerus occurs either above
or intermingled with bands of the mussel, Brachidontes rostratus (Dunker), to
which it is often attached. The absolute width of its vertical range varies in
Victoria proportionately with the variations in tidal range but it still occupies
relatively the same vertical proportion of the tidal range (Bennett and Pope,
1953).

Fig. 2. Central segments of cirrus VI of (a) Catophragmus polymerus showing small central tuft of

spies between bases of major spine pairs, and (6b) Octomeris brunnea.
F. J. Beeman del.

Distribution

Australian: Collections in the Australian Museum show that Catophragmus
polymerus ranges from Currumbin, near the border between New South Wales
and Queensland, southwards through Victoria and round into the eastern half
of the Great Australian Bight (Womersley and Edmonds, 1958). It also occurs
round Tasmanian shores. However, in Victoria and Tasmania the population
numbers may be considerably reduced where conditions favour the growth of
the mussel, Branchidontes rostratus, which occurs in beds on the shore rocks.
A covering of mussels prevents the Catophragmus larvae from settling and only
a few survive to settle on the shells of the mussels themselves.

Catophragmus polymerus is virtually absent also from approximately 200
miles of the Victorian coast immediately west of Cape Otway and only scattered
specimens are found in south-west Tasmania. These areas of coast are subjected
to prevailing cold winds from the south in summer, and probably air temperatures
are too low for survival of Catophragmus. Several other species of intertidal
animals are also missing in the same areas. However, insufficient physical
data are available to draw any firm conclusions in the matter.

In Darwin’s original description the locality ‘‘ Swan River?” is given for
material in the Cuming collection. However, no examples of C. polymerus
are to be found today, anywhere in the vicinity of Perth or Fremantle, and it
is absent from the southern coast of Western Australia. This anomalous record
of Darwin’s has always been a puzzle, for it is highly unlikely that the species
ever occurred there and it is not one that attaches itself to ships, so it is
unlikely that it could have reached the Swan River in this manner. Recent
Surveys on the intertidal zone in Tasmania, however, suggest how this record
may have occurred. Miss Isobel Bennett collected C. polymerus on Swan
Island in Bass Strait. Since Darwin’s original record has a query after the

20 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

word ‘‘ River’ he may have been unable to decipher the original label, with
the Cuming specimens, and they may actually have come from Swan Island.
If this were so, it would fit into the present geographical range satisfactorily
and remove the anomalous record for Western Australia. Another unfortunate
record occurs in the caption of Figure 468 in Broch’s contribution to Kukenthal
and Krumbach (1927) where it is stated that ‘*‘ C. polymerus is found in the
northern intertidal zone of Australia’, whereas it is limited to the southern
half of the continent.

As the growth of C. polymerus is favoured by exposure to turbulent seas,
it is not surprising to find it also missing along the sheltered part of the northern
coast of Tasmania where the shores are protected from the full force of oceanic
waves and the waters are sometimes turbid (Bennett and Pope, 1960).

World Occurrence: It is considered that Gruvel’s (1905) record of C.
polymerus from ‘‘ Table Bay ” (? South Africa) is probably due to an error of
labelling, for there are no subsequent reports of it from any locality outside
SE. Australia. Old records show that Mt. Wellington, Hobart, Tasmania, was
sometimes called ‘*‘ Table Mountain ” and this may account for Gruvel’s record.

Genus OCTOMERIS Sowerby, 1825

Nilsson-Cantell, 1921, q.v. for earlier bibliography.

Mention of this genus in connection with the Australian fauna was first
made in 1932 by T. H. Withers who recorded the fossil species Octomeris crassa
from beach-rock from Magnetic Island, North Queensland. However, in 1952,
a collection of living Octomeris, taken on a beach-rock boulder, from Magnetic
Island was brought to the Australian Museum and proved to be mature
specimens of O. brunnea Darwin.

Subsequent surveys along the Queensland coast by R. Endean, R. Kenny
and W. Stephenson (1956) recorded O. brunnea from Coral Point (Broad Sound)
northwards to the Cooktown area. Field work by the author has extended
the range southwards approximately to latitude 23° S.

In all, seven species of this genus have been described since 1825, but later
workers reduced the number by synonymy to four: O. angulosa Sowerby from
southern Africa; O. brunnea Darwin from Japan, Formosa, the Philippines,
Indo-Malayan Archipelago, Australia, Santa Cruz and the New Hebrides ;
O. suleata Nilsson-Cantell from Japan and Formosa; and the fossil species O.
crassa Withers recorded from beach-rock of Magnetic Island, off Townsville
in Queensland.

In this present paper O. crassa is synonymized with O. brunnea, reducing
the number of species in the genus to three.

Key to the species of Octomeris

1. Mandible with four main teeth and a pectinated lower border on which the lowest spines
are the longest. Bristles on the anterior border of Maxilla I divided into only two groups
by a single notch. Sutures between shell plates finely toothed. Basis membranous
Soasis Silent Bese SSS. Sieg 2S ERS LPR ROOTS Le aes cars ee eae O. angulosa Sowerby, 1825.

la. (1). Mandible with three teeth and a pectinated lower corner. Bristles of the anterior
margin of Maxilla I in three groups, either because two notches separate them as in O.
brunnea or, as in O. sulcata, because they are arranged in that way, although there is only

COANE ALO REL MUS Ea ue ec cE AIS, cus Bunt 620-5. oo Duo d ORG oid OF G6. oio.4 CIQIGINIT bo socimia FIDO EDS 950 cad (2)
2. No caudal appendage, sutures between shell plates coarsely toothed. Basis membranous
but with a central hole or thin area ........................ O. brunnea Darwin, 1854.
2a. (2). With cauda, appendage. Sutures between plates without teeth. Basis calcareous
Ehavole ql avleliG pry Ma ARMM hay by SME) Baal dee PAREN Bice Seis ID eu@end lero na O. sulcata Nilsson-Cantell, 1932.

OCTOMERIS BRUNNEA Darwin, 1854
(Plate i, figures 3 and 6; Text-figures 1,¢; 2,b)
Octomeris brunnea Darwin, 1854; Weltner, 1897; Gruvel, 1903, 1905;
Nilsson-Cantell, 1921, 1930 ; Hiro, 1932, 1939 ; Utinomi, 1954; Endean, Kenny
and Stephenson, 1956; Endean, Stephenson and Kenny, 1956.

ELIZABETH C. POPE 21

O. intermedia Nilsson-Cantell, 1921, 1932, 1938; Hiro, 1939.

O. crassa Withers, 1932. .

Coronula sp. Longman, 1930.

The comparatively late discovery of living O. brunnea on the Queensland
coast, by Dr. A. Keast in 1952, is explained by its cryptic habit and the lack
of systematic collecting of barnacles in Queensland before that time. The
fact that the intertidal surveys of Endean et al. (1956) discovered the presence
of O. brunnea along the east coast of Queensland from Coral Point, N. of Broad
Sound northwards to the Cooktown area in any locality offering suitable
habitats for its settlement and survival, leads one to believe that careful
collecting will reveal its presence throughout the Indo-Malayan Peninsula,
the Philippine Islands and south-east Asia from latitude 34° 5’ N., southwards
round the Asian coast line to the Mergui Archipelago in the Indian Ocean where
it has been recorded as O. intermedia by Nilsson-Cantell (1938).

The reasons for synonymizing O. intermedia with O. brunnea have been
set out by Hiro (1939) and it is only necessary to state that the extensive
Australian collections of Octomeris fully support Hiro’s action. There are
several series of barnacles from Queensland that were taken in the one locality,
sometimes from the same rock, that show individuals with the characters
described by Darwin (1854) for O. brunnea and others with the appearance
of O. intermedia of Nilsson-Cantell (1921). There are also intermediates between
these two kinds of shell structure.

Examination of the type specimens of both these species in European
museums shows them to agree closely with Queensland specimens in the
following way—the type of O. brunnea being referable to uneroded and generally
fairly juvenile barnacles (see Plate i, fig. 6) and that of O. intermedia, to older
eroded specimens (Plate i, fig. 3).

The soft parts of O. brunnea do not grow proportionately with the increase
in size of the shell. Their rate is slower than that of the shell plates, the inner
basal parts of which thicken greatly, so that the animal occupies only the small
central space immediately below the opercular plates. There is a close
correspondence in the morphology of mouth parts and cirri in all Australian
Octomeris, so that there is no doubt that only one species is present in the area.

The fossil species, O. crassa Withers (1932), is also included as a synonym
of O. brunnea for the following reasons :—Examination of the type material
of O. crassa in the Queensland Museum revealed complete similarity of shell
structures between the fossil barnacles from Magnetic Island, Queensland, and
specimens of well-grown and somewhat eroded O. brunnea growing today on
the beach-rock at the type locality of crassa (Magnetic Island, Queensland).
The ‘‘ fossil ” barnacles show no shell character not present in living O. brunnea.

It may be that Withers’s estimate that the rock forming the matrix is
‘‘ possibly of Pleistocene age’ is in error for, in the opinion of Dr. Graham
Maxwell (in a private communication to the author), the process of beach-rock
formation in Queensland has been continuous in geological time right up to
the present day and the beach-rock containing the O. crassa may, in fact, have
been younger than Withers thought. Associated fossil barnacles listed by
Withers as “ Tetraclita sp. and Chthamalus sp.” occur in the same rock with
his O. crassa. These proved upon examination to be Tetraclita squamosa Brug.
and Chthamalus caudatus Pilsbry and to differ in no way from present-day
specimens of these two species which are associated with living O. brunnea.

A fairly detailed description of the collections of O. brunnea in the Australian
Museum, Sydney, follows. It is based on a large series of specimens from a
wide geographical range.

Appearance and Shell Stuctures

The shell comprises eight sub-triangular plates with crenated sutures
between them. In uneroded and juvenile specimens the colour is brown and

22 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

the shell plates are finely ribbed (Plate i, fig. 6). There is a persistent brown
epidermis which projects as a comb-like row of tiny spines along the horizontal
growth-lines on the ribs. Eroded specimens tend to show a general grey colour
alternating with wavy dark lines where the dark laminae are exposed (Plate i,
fig. 3). In juveniles the orifice is often toothed but erosion leads to the wearing
down of the projecting apices and the orifice becomes wider. In eroded specimens
traces of the brown rib-folds are often still evident near the base of the shell.

As a rule, the outline of the base is almost circular but a series of measure-
ments shows that the carino-rostral diameter is generally slightly greater than
the width of the shell.

The shell has mostly a depressed conical shape except where crowding
modifies growth. Crowding tends to cause the shells to grow taller as
Table 1 shows. In Table 1 measurements are given of several Octomeris brunnea
exhibiting different growth habits, as being representative of numerous other
specimens that were examined in the present collection.

TABLE 1
Measurements of O. brunnea and Shell Appearance

Locality Carimo-rostral Breadth of Height of Deseription of
diam. mm mm. shell in mm. shell nm mm. shell
Chinaman’s Beach, Somewhat eroded normal
N. of Cairns, Q. 24 23 5-0 shape with parietes thickening
at base.
a 12 Bt 4-5 Uneroded. In a crowded
group.
North Keppel Is., Greatly eroded and parietes
E. of Yeppoon, Q. 25 23 5:5 much thickened, normal low
shape.
35 18 14 6-0 Uneroded. Shape distorted
and tall due to crowding.
Slade Pomt, near Shell eroded; parietes greatly
Mackay, Q. 26 22 7-0 thickened. Distorted by
crowding.
e 20 19 3-1 Shell eroded ; parietes thick-
ened, normal shape.
South of Nasowa, Fairly uneroded shell moder-
W. Central Maevo, 13 13 3-0 ately thickened at base.
New Hebrides Shape normal.

The eight shell plates are rarely equal in size, the carino-laterals and the
rostro-laterals are, more often than not, smaller than the others. Where shell
erosion is severe, it may be necessary to inspect the inner side of the shell to
see the crenated sutures. Radii and alae are but little developed and in older
specimens consist only of the projecting teeth which interlock with those of
adjacent valves. The basis is membranous but does not cover the entire area.
Centrally there is a thin patch or lacuna, as shown in Plate i, fig. 3, left
specimen.

Hiro (1939, pp. 252-4, fig. 3 and 4) gives an excellent account of the changes
occurring in the external appearance of the opercular valves and shell plates
with growth and with erosion and there is nothing to add to his description
in this respect.

The sheath extends to about half the depth of the shell and is lined by
2a brown membrane but below the sheath the interior of the parietes is a
translucent grey.

Scutum and Tergum

Basically the structures of the scutum and tergum in Australian O. brunnea
are similar to those depicted in Hiro’s paper (1939, fig. 3). However, slight

ELIZABETH C. POPE 23

variations occur in the shapes, articular ridges and furrows of these two valves,
for they may sometimes be situated nearer to the apices of the valves or, in
older specimens, nearer to the centre of the tergal margin of the scutum or the
scutal margin of the tergum. The depth of the scar of attachment for the
adductor muscle of the scutum is also variable, as are the number of crests
for the attachment of the depressor muscles of the tergum.

Soft Body

The flaps of the tergo-scutal membrane are bordered with white, round
the orifice, in newly preserved material and the cirri are light coloured, except
in the areas round the bases of the anterior spine pairs and the posterior spine
tufts. Here there are concentrations of dark pigment giving a distinctive barred
colour pattern to the cirri.

Trophi

The labrum has a well-defined wide, shallow groove which in all specimens
examined is deeper than that depicted by Nilsson-Cantell (1921, text-fig. 59a).
It is bordered by a single row of triangular teeth along its entire length. In
describing O. brunnea, Darwin (1854, p. 485) likened the 'abrum to that of O.
angulosa and as therefore having the “ Labrum bullate, with the crest hairy
and furnished with a few most minute teeth’. It must be remembered that
Darwin was describing immature specimens of this barnacle and this may
account for the smaller number and size of teeth seen. Palps.—The palps
of Australian O. brunnea bear more bristles along their straight anterior margins
and near the tip, than the number shown in Nilsson-Cantell (1921, text-fig.
59-61). Otherwise they are similar to those he described. Mandible-—The
mandible has three well developed teeth and a pectinated lower angle. There
is Some degree of variability in the structure of the second tooth for it may
carry a few small denticles on its upper margin or be quite smooth. These
denticles tend to disappear as the barnacle matures. The third tooth is generally
pectinated on its upper side. Mazilla I.—Two notches on the anterior border
of the first maxilla divide the bristles into three groups—two smaller bundles
on either side of a large central one. A pair of large spines and two pairs of
smaller spines lie above the first notch. The central group of bristles are second
in size only to the large dorsal pair above the first notch. The lower group
gives a pectinated appearance to the lower angle of the maxilla. Mazilla II
is aS shown by Nilsson-Cantell (1921, text-fig. 61) but is more hirsute.

Cirri

The first two cirri are considerably shorter than the remaining ones and
very setose. In each pair the rami are unequal, the anterior one being longer
by approximately two or so segments and about half as broad again as its
fellow. In spite of this, the two rami generally have an equal number of
segments (from 6 to 7) or occasionally they may differ by one. Many of the
longer and broader setae are pinnate as shown in Text-figure 1, c. Others
are pinnate but finer. In cirrus I there are a few of these broader setae on
the terminal segment but only a few occur in the segments below it. The
pedicel is also hirsute with numerous bristles along its anterior surface. Cvrrus
II resembles I in shape and size but the anterior ramus is not so broad, by
comparison with I. Although the segmental numbers of the rami tend to be
equal, the anterior one is longer by the length of more than two of its segments.
Many of its long setae are pinnate and are so numerous in the terminal segments
of the anterior ramus as to form a felted mass and an efficient sieving organ
for catching minute food-particles. Similar setae are found in the posterior
ramus but they are fewer in number. Cirri III to VI differ in shape and are
much longer than I and II. The two rami of each cirrus are comparable in
breadth and in shape and are almost equal in length except for cirrus III in

24 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

which the posterior one may sometimes be slightly the longer. Several of the
more basal segments of the anterior ramus of cirrus III carry tufts of pinnate
spines. Hach segment of the rami has four pairs of stout larger spines
anteriorly, of which the distal ones are longest and the proximal pair very
much the shortest, while posteriorly there is a small tuft of spines, placed at
the junction of adjacent segments (Text-fig. 2,b). One pair of spines in this
tuft is generally longer than the other ones. Concentrations of dark pigment
occur near the bases of all these spines, giving the characteristic darker colour
patches posteriorly, near the junction of adjacent segments.

The numbers of segments in the rami of the cirri vary slightly between
right and left sides of the same barnacle. Several specimens have been dissected
which had damaged cirri—shorn off at varying distances from the pedicel.
Damage and regeneration may account for the fact that the numbers of segments
making up the rami of right and left cirral pairs in one individual do not always
agree.

The penis is of moderate length, tapering to a blunt tip. Near the
extremity there are a few hairs scattered generally over the surface, with a small
tuft of hairs near the tip of the organ. There is no caudal appendage.

Habitat

Octomeris brunnea occurs either far back in crevices or on the undersides
of small boulders or overhanging rocks where it is sheltered from sunlight.
A series of specimens occurs on mangrove roots taken at Carlisle Bay, Santa
Cruz Island. Another batch was collected from the walls of an intertidal lime-
stone cave 14 miles south of Nasowa, W. Central Maevo, in the New Hebrides.
It occurs on the higher levels of the shore from approximately the level of mean
high water down the shore to the level of high water of neap tides. It is
rarely found in large numbers even in favourable localities.

Distribution

Australia and the Pacific: Collecting by the author has extended the
known range of O. brunnea in Queensland southwards for a further 200 miles
to the vicinity of the Keppel Islands in latitude approximately 23° S. There
are also specimens from the New Hebrides and Santa Cruz Islands in the
Australian Museum. It is not recorded, as yet, along the northern coast of
Australia.

World Occurrence: Southern Japan ; Formosa ; Philippine Islands ; Malay
Archipelago generally ; E. coast of Queensland, south to Tropic of Capricorn ;
New Hebrides and Santa Cruz.

Genus CHTHAMALUS Ranzani, 1817

Pilsbry, 1916, q.v. for earlier synonymy and references ; Nilsson-Cantell,
1921; Hiro, 1932; Kolosvary, 1941; Moore, 1944; Newman, 1961; Davadie,
1963 ; Zullo, 1963.

From the point of view of taxonomy the genus Chthamalus is regarded
as a difficult one. Certainly, so far as Australian species were concerned this
proved true. While four of the five species, namely C. antennatus, C. withersi,
C. caudatus and C. intertextus, posed few problems in identification and the
patterns of their geographical distributions were clear cut and logical, the
fifth, C. malayensis, proved most difficult to name. At one stage Australian
material of this species was thought to represent at least three distinct popula-
tions with confused and overlapping distributions, especially if stress were
placed on obvious shell differences, and one of these populations (typified by
the shell illustrated in Text-fig. 5,d) was not recorded in literature. These
three groups ultimately proved to be ecological or age variants of the one widely
ranging, polymorphic species, the true nature of which had not been appreciated

ELIZABETH C. POPE 25

by workers on the following important collections :—(1) the Dutch Siboga.
Expedition ; (2) additional collections from the Dutch East Indies (Indonesia)
housed in Dutch Museums; (3) Dr. Mortensen’s Pacific Expeditions (1914-16),
and (4) large collections from the Indomalayan area worked by C. A. Nilsson-
Cantell which were housed in various European museums. It was not recog-
nized by Darwin (1854) as being distinct from C. stellatus but he commented
on the pecularities of specimens from the Philippine Islands, although he did
not separate them from the European species. In all these earlier collections
samples had been taken from widely scattered geographical areas and little
notice was taken of any ecological background. Considerable confusion exists
in the literature documenting these collections and, even after examining all
of the specimens concerned, the author was still left in doubt as to the correct
name to apply to the species universally distributed round the tropical
Australian coast. However, many doubtful points were clarified when a
sufficiently large sample of this wide-ranging population was examined, and
the examination of the type of Pilsbry’s C. malayensis in the United States
National Museum in Washington, D.C. in 1963 provided the necessary clues
to clear up any doubts about the nomenclature of the northern Australian
species. Although the name C. malayensis had been applied by the author
to material collected during the intertidal surveys in Queensland made by
Endean, Kenny and Stephenson, the use of the name was still somewhat
tentative at that time. Fortunately it has proved correct so that records of
it in their surveys need no emendation. Under the specific description of C.
malayensis some slight indication is given of the confusion that has occurred
in its naming in previous literature and, while the synonymy quoted has been
made as complete as possible, carrying on from Utinomi’s excellent review
(1954), no attempt has been made to take each reference and state the nature
of the error it contains. To do so would require more space than is merited
in the present review. However, in the synonymy above, with the exception
of Stubbings (1963) and Daniel (1956), the actual specimens concerned have
been examined by the present author, before the step of synonymizing them
was taken.

When large populations of Chthamalus from all over the world are
examined without regard to names previously bestowed on them, it becomes
apparent that confusion of the species is in the literature, rather than inherent
in the barnacles themselves, provided that a certain combination of characters
of both shell and soft parts is used to determine the species. The chief
difficulty in determining the doubtful Australian Chthamalus hinged on being
able to distinguish with certainty the following closely related species :—C.
stellatus (Poli) s.s., C. antennatus Darwin, C. challengeri Hoek and C. malayensis
Pilsbry. During the present study the following characters were found useful
in distinguishing them: the general morphology of the mandible in the region
of the comb-like series of spinules ; the type of notching, below the uppermost
large spines on the first maxilla, and the crests for attachment of the depressor
muscles of the tergum, while the most reliable criteria, in fully mature specimens,
proved to be the minute structure of the stout, lanceolate spines associated
with the terminal segments of both rami (chiefly the anterior one) of cirrus
II and the sculpturing of the pit or depression on the interior of the scutum,
where the lateral depressor muscles are attached. These distinguishing
characters are tabulated in Table 2 which should assist the renaming of Indo-
malayan specimens in the collections of various European museums where
C. challengeri, C. stellatus and C. malayensis have often been very much confused.

Recent ecological-systematic surveys of cirripedes in the vicinity of Japan
and south-east Asia by Utinomi (formerly known as Hiro) have led to the same
general conclusions about the distributions of species within genus Chthamalus
as have emerged from the present study. After due allowance has been made
for the taxonomic confusion of the past, it is seen that there is a distinct tropical

AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

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ELIZABETH C. POPE 27

fauna common throughout the Indomalayan region which has its northern
boundary in the southern regions of Japan and SEH. Asia in approximately the
latitude of Taiwan.

This same fauna has now been found to range south of the equator into
the northern region of Australia and, in fact, colonizes the coasts of the northern
half of the continent. It contains no species of Chthamalus endemic to the
region but is characterized by C. malayensis, C. caudatus, and to the east by
C. withersi also. To the north in New Guinea C. intertextus also occurs.

In the more temperate seas of Japan a different group of species in genus
Chithamalus is represented on the upper shore, namely C. pilsbryi Hiro, C. dalli
Pilsbry and C. challengeri Hoek. In temperate Australia the genus is represented
by a single species, C. antennatus, and the rest of the upper shore Chthamalids
belong to different genera, namely Chamaesipho and Catophragmus. There is
thus a marked falling off from four species to one in the genus Chthamalus on
the southern half of the Australian coastline and corresponding levels of the
upper shore, while still occupied by representatives of the family, are relatively
poorly supplied with species of this genus. The sole representative, C. anten-
natus, is, however, endemic to southern* Australia and does not range across
the Tasman Sea to New Zealand where there is no representative of the genus
in the intertidal fauna. THarlier records of genus Chthamalus in New Zealand
by Cranwell and Moore (1938) have been subsequently shown by one of them
(Moore, 1944) to be juveniles of a second species of Chamaesipho—(C. brunnea).

The geographical distribution of Indomalayan and Pacific species of
Chthamalus can now be clearly defined, and Kolosvary’s (1941) arrangement
of them into formenkreise is found to be inaccurate as there is little tendency
for overlapping in the distributions of tropical and temperate species from one
major zoogeographic area to another. This author’s records of C. stellatus in
New Guinea, C. malayensis from Victoria, and C. moro from New Zealand in
his 1943 work should also be disregarded in the light of the present survey.

Among Australian species, C. withersi, C. intertextus and C. caudatus show
affinities in some of their structures, habitat preferences and feeding adapta-
tions with Octomeris, whereas C. antennatus and malayensis have corresponding
structures and adaptations more aligned with those of the most primitive of
all living genera, Catophragmus. Chthamalus intertextus also has features
relating it with both the three- and four-toothed groups of species within the
genus Chihamalus and also with Chamaesipho, since the structures of its short
stout spines on cirri IT and III are almost identical in shape with those described
by Moore (1944) for Chamaesipho brunnea. Zullo (1963) indicated in a paper
read before the X VI International Congress of Zoology that it was his intention
to divide the former genus Chthamalus into two, resurrecting Conrad’s name
Huraphia for the group of species with tridentate mandibles (Nilsson-Cantell’s
hembeli-group) and restricting the genus Chthamalus s.s. to those species with
quadridentate mandibles, of which C. stellatus is a typical example. Since
Zullo’s paper is not yet published the matter must be considered under review.
The present study has, however, shown that the line dividing species hitherto
characterized as having tri- and quadri-dentate mandibles is by no means so
clearly defined as was thought by Pilsbry (1916) and Nilsson-Cantell (1921).
The finding of large individuals of certain Australian species in which normally
4-toothed species have developed only 3 teeth, or conversely, of 3-toothed
species with 4 teeth, is going to make the drawing of distinctions between
Zullo’s proposed generic groups somewhat difficult. Moreover, if Chthamalus
intertectus is included with the hembeli-group of species (to which it shows

* The unfortunate use of geographical terms such as “South and West ” in the names of
certain Australian States can lead to confusion in discussions of distributions. The present
author’s practice is to use term “ southern’’ for the whole coast of the continent south of
approximately the latitude 30° S., and not just restricted to the State of South Australia.

28 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

undoubted affinities) the relating of the genus Chamaesipho more directly to
the stellatus-group by Zullo will require some explanation, since the structure
of its short stout spines on cirri IT and III and tendency of the sutures between
its shell plates to become obliterated and overgrown by later calcification.
would seem to indicate certain affinity with species like Chthamalus interteatus.

No published key to species of Chthamalus satisfactorily differentiates
C. malayensis from closely related forms and the key in Nilsson-Cantell’s 1921
monograph omits it. The following full synopsis and key to Australian and
New Guinea species has therefore been prepared and since it will largely be
used by non-systematists, unfamiliar with the vagaries and difficulties associ-
ated with the characters used to differentiate species in the genus Chthamalus,
@ rather full series of warnings and notes are appended in the form of footnotes.
With the aid of these, identification of Australian species of Chthamalus should
present little difficulty.

Key to Australian and New Guinea species of genus Chthamalus

1. (la). Chthamalus with much flattened shells without distinct ribs and having a mandible
generally with only three major teeth* above a coarsely and irregularly pectinate section
forming the lower tip of the jaw, with the longest spine generally at the tip of the jaw.
Teeth 2 and 3 often with secondary side teeth. Labrum generally not bullate but
flattened above the palps or developed as a muscular, semicircular funnel-like organ (see
Text-fig. 3.d). Rami of cirrus II without stout serrated, lanceolate spines on their terminal
segments except for C. intertextust. Pinnate setae very numerous ................ (2)

la. (1). Chthamalus with shells generally taller and of truncated conical shape which, under
conditions of crowding, may become tubular. Rarely flattened but in such individuals
ribbing generally more distinct than m species In group 2. Mandible with four major
teeth (occasionally five) of which the fourth is generally doubled*. Lower tip of jaw
generally armed with several (from two to four) spies of even size and distinct from the
other spines above them. Between these and tooth four, straight-edged, groups of closely
packed spinelets generally form comb-like sections as illustrated by Utinomi (1959, fig. 5,a)t.
Labrum typically rounded or bullate above the palps (Text-fig. 5,c). Terminal segments:
of both rami of cirrus I] armed with a bunch of stoutly built, lanceolate and serrated
spies” among. ythe pinnate Sevaen.0. 4a sc se ck ee eee oes ee Ort at hee Sl cree eee teint cae pe (4);

2. (2a). Sutures between shell plates with interlocking teeth or mterfolded lamimae. (If shells:
very eroded these structures may sometimes be seen only imternally.) Basis either
membranous or with an outer calcareous rim round an imner membranous section.... (3)

2a. (2). Sutures between shell plates simple and not interlocking and only loosely articulated
together. The lines marking the sutures between scuta and terga are straight, even im
worn fully grown individuals and not shaped like the Greek letter psi as in most species
of Chthamalus. Found in the highest intertidal zone. Shell much flattened and, if
uneroded, often cmnamon-brown with four white ribs projecting mto the orifice as teeth,
from the tips of the side valves. Rostro-laterals and lateral plates often with imternal
pits or calluses. Basis always membranous ................ C. withersi Pilsbry, 1916.

3. (3a). Sutures between shell plates with zigzag interlocking joint owing to teeth. Paired,
long slender, jomted caudal appendages present. Scutum and tergum separate and not
ankylosed. Basis wholly membranous .................. C. caudatus Pilsbry, 1916.

* Warning: exceptionally, large C. withersi may occasionally develop a fourth tooth on
the mandible (see p. 43) but this fourth tooth is rarely doubled and there is generally no trace
of comb-like arrangements of spmes below it.

7 C. mtertextus stands in a somewhat anomalous position since it clearly has only three teeth
on its mandible and has a peculiar semicircular labrum (like that of C. caudatus) which is not
bullate. However, the terminal segments of cirrus II possess a few stout spmes hidden among
the pimnate ones (much less obvious than those of species with four-toothed mandibles withim
the genus). These spines (Text-fig. 1,f) are not of the regularly serrate pattern, resembling
more the type of stout spme found in the genus Chamaesipho.

{ Very juvenile specimens or individuals with damaged and regenerating mandibles of the
stellatus-group may fail temporarily to show distinct comb-like sections in the lower part of
their jaws and may exhibit a somewhat random arrangement of the spinelets (see C. antennatus,
Text-fig. 4,9) reminiscent of the adult jaw im species with tridentate mandibles. Evidence from
the examination of a number of specimens would make it appear that a developing or regenerating
jaw of a typically “ stellatus-pattern ’”’ passes through a kind of “ hembeli-stage ’ of jaw develop-
ment before reaching the comb-like arrangement of spinelets normally seen in adults below
the fourth tooth. Generally im such cases, the doubled fourth tooth and lack of secondary
toothing on teeth two and three are clues which help to key a specimen correctly.

ELIZABETH C. POPE 29

3a. (3). Sutures formed by oblique interfolded laminae of radii and alae (when the shell is
not eroded) and marked by prominent vertical lines of growth, giving a chevron-like
appearance to the sutural areas of the shell (Text-fig. 3,2). When eroded, shell becomes
a deep violet hue due to secondary calcification internally and sutures are marked by
characteristically rounded wavy lines, as shown in Text-figure 3,6 and c. Scutum becomes
firmly ankylosed to its neighbouring tergum and basis often saucer-shaped with outer rim
calcified and inner section membranous (seen in side view in Text-figure 3,c). Cirrus II
and basal segments of cirrus III armed with stout, grapple-spmes of unusual shape
(exalt RMU Mee teerdevsctote Cuan Pact kl. soe auhiba Akpaem ot C. intertextus Darwin, 1854.

4. (1). With four teeth on mandible and sometimes even a small fifth tooth may be present.
Between the group of spies forming the lower tip of the jaw and the fourth tooth is
generally a fairly short comb-like section where the spmelets are of even size and length
but are not fine and hair-like*. Termmal segments of cirrus Il armed with very stout
SeLtaibcom@s pines) wiallan COALS aSIC en beetles aa nvr Sebo sey weirs pete ole wee easel a (5)

5. (5a). Inside of opercular valves not ruggedly sculptured and rarely pitted. Tergum
broad near its basi-scutal corner and generally with two (or, at the most, two plus a partly
developed third crest in larger individuals) for attachment of its depressor muscles.
Scutum with a distmet, but shallow, depression for the attachment of the lateral depressor
muscles, in which two low, incipient crests may be distinguished in full-grown individuals.
Serrated stout spines of cirrus II with a double row of blunt, short, peg-like spines in
which the two lowest are not separated from the rest by a distinct space (see Text-fig. 1,h)
Re es es ese bs ct ald any Jaud' whey 3 Gyaije) jsuchee te ons C. antennatus Darwin, 1854.

5a. (5). Inside surface of opercular valves with more rugged sculpturimg than in C. antennatus
and mostly pitted deeply. Tergum narrowing towards its basi-scutal corner and folded
along a line of deep pits along the axis drawn from the apex to the basi-scutal angle.
Generally at least three crests for attachment of the tergal depressor muscle with larger
and older individuals developing four crests. Scutum with a deep narrow pit for attach-
ment of lateral depressor muscles and no sign of ridges or crests within this pit. Lanceolate
spines of cirrus II stout and armed with a double row of stout pointed side spines, the bottom
pair of which may be worn down to mere bumps, but which are separated from those above
Dyeomsiohtecapm (Mextat or ey) er. rs see. 2 eee S119) 2, 52 ener cles C. malayensis Pilsbry, 1916.

CHTHAMALUS INTERTEXTUS Darwin, 1854
(Plate i, figure 1; Text-figures 1,f; 3,a-d)

Chthamalus wntertextus Darwin, 1854, illustrated; Gruvel, 1912; Hoek,
1913; Pilsbry, 1916, 1927; Hiro, 1939, illustrated; Utinomi, 1949, 1954;
Tokioka, 1953; Newman, 1961, illustrated.

The possession of a unique collection of obvious shell peculiarities such
as (1) a basis which is partly calcareous and partly membranous; (2) the
beautiful violet colour of the inside of the shell and the calcareous ring of the
basis; (3) the interlocking sutures between the parietes; and (4) scuta and
terga which tend to ankylose, so that only a trace of the original suture between
each scutum and tergum remains (generally only towards the basal parts of the
two valves) have contributed to the making of Chthamalus intertextus easy to
determine. It has therefore not been confused in past accounts.

However, recent work by Newman (1961) has shown that Darwin’s original
description was incorrect in one important respect. Darwin stated that every
full-crown. specimen. he investigated had the basal edge of the parietes “‘ inflected
rectangularly inwards, forming a smooth-edge ledge all round the basal mem-
brane, which, in proportion to the width of this ledge, was by so much reduced
in diameter’’. Newman has found that the calcareous basal shelf inside the
shell wall of C. intertextus was a product of secondary calcification of the basis,
rather than, as Darwin believed, an inflected extension of the parietal walls.
Newman also examined C. hembeli (Conrad) and Chthamalus calcareobasis Dora
Henry (1957), which are most closely related to C. intertextus in both having
calcareous bases, aS well as other shell structures in common. In both these

* Had it been necessary to include C. stellatus and C. challengeri mn this key, they would
have been separated from other species at this point by possession of the followimg characters :—
With four-toothed mandible, with a comb-like section between fourth tooth and spines of lower
tip of the jaw broader than the corresponding structure in C. malayensis and C. antennatus and
with much finer toothing—more hair-like than spime-like. Stout spmes on terminal segment
of cirrus II armed with double row of very fine spines (almost hair-like in structure)—never
peg-like or spine-like, as nm C. malayensis and C. antennatus.

30 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

species Newman pointed out that the limy base was also a product of secondary
calcification which, in the case of C. hembeli, was also marked by the deposition
internally of successive layers of shell material on top of the basis and the inner
parietal wall, which obscured the interlocking junction of the parietes and basis
and thus misled Darwin, and Henry who accepted Darwin’s statement.

It is not intended here to go into detail of the anatomy, or of the slight
variations which, no doubt, could be found in the present sample of C. interteatus
from New Guinea, for the simple reason that the material available is con-
sidered to be too meagre to be representative of the world population of this
species. Until collections of it are at least equivalent in their cover of the whole
geographical area—as they have been in the other species discussed in this
paper—it is sufficient merely to record additional facts which may help other
workers on the group.

As has been the case in all other species described in this paper, the size
of the specimens, in most cases, exceeds those seen by previous authors. Some
of the facts about this sample are summarized in Table 3.

TABLE 3
Measurements of 10 Chthamalus intertextus from Papua, New Guinea, taken in January, 1964

Carimo-rostral Width of shell Height, from

diameter, at right angles lowest point Breeding- Remarks
in mm. to the carmo- of basis to conditions
rostral diam- highest on
eter, m mm. shell, in mm.
eyed-nauplii
Uo eae 9-4 3-0 present
Colour: ash-grey to white
externally with, in places,
1-5 6-3* 3°2 no larvae persistent yellow epidermis.
Uneroded ; sutures interlock-.
ing
Almost rectangular in shape
10-8 9-4* 3-2 eyed-nauplii Central membrane of basis
present oval and dark—black or
navy-blue
eyed-nauplii Membranous part of basis
11-6* 11-2 4-0 present black
eyed-nauplii Shape: almost rectangular ;
11-8 oO 3-0 present membranous part of basis
oval
12-5 11-5* 4-0 no larvae
eyed-nauplii Caleareous basis -+ central
12-5 11-8 4-0 present membranous part, almost
basin-shaped
Shell sculpturing : raised ribs
eyed-nauplii or rows of white knobs—shell
12-5 12-0 4-0 present mauve when broken across,
powdery—white on surface.
Very worn
eyed-nauplii
13-0* ibe oy 3-0 present
Basis an outer rim of bright
eyed-nauplii mauve limy material and
14-0 910+ a7 2 present a central very dark mem-

branous pit

* Growth restricted im some way, e.g.

by erowding or by edaphic factors.

ELIZABETH C. POPE oul

Again it becomes evident that the presence of developing, nauplii inside
the shell is by no means a proof that the barnacle concerned has attained its
full size, as is so often assumed in earlier accounts. From Table 3 it may be
seen that while the smallest barnacle was brooding nauplii, it was approxi-
mately only half as large as the largest one in the group measured.

For a full description of the shell and of the soft body the accounts of
Darwin (1854, pp. 467-8, Plate 19, figs la and 1b) and of Hiro (1939, pp. 251-2,
fig. 2) should be consulted, and for ecological and zoological information papers
by Hiro (1939), Utinomi (1949 and 1954), and Tokioka (1953) will be found to
contain valuable additional information.

Appearance of Shell

Most specimens examined were greatly eroded and showed scant evidence
externally of the rather distinctive sculpturing on the shell, with which they
began their lives. The shell plates of an uneroded individual were irregularly
ribbed longitudinally and the whole shell was marked by numerous raised,
horizontal lines-of-growth, as shown in Text-fig. 3a. The deep violet of the
inner layer of the shell was visible only between these ribs where the yellowish
epidermis had worn away and allowed erosion to begin. The interlocking of
the sutures between adjacent plates begins as a most complex structure and,
before erosion occurs, the radii are seen to consist of a series of laminae,
paralleling the inner wall of the valve. These are “ interleaved ” rather than
‘interlocked ”’, with a series of similar laminae projecting in the opposite
direction, outside and parallel to the inner wall forming the alae of the adjacent
valve. All these projections carry numerous, obliquely-placed, fine lines of
growth. When viewed in situ externally, they look like a series of chevrons.
Usually only traces of the lines of growth and the ribs persist as longitudinal
rows of white raised bosses on the worn, bright violet background of the inner
layer of the parietes. After a certain amount of erosion, the sutures are simple
and straight from the orifice outwards for a certain distance (all trace of the
“‘ chevrons ”’ having disappeared), then each suture becomes wavy, with round
interlocking projections and continues thus to the periphery of the shell
(Text-fig. 3,5 and c). These projections of the interlocking joints are too
rounded to be called teeth. Internally the shell wall is coloured a deep violet
and is much pitted.

Basis

The most outstanding feature of the shell of C. intertextus is the basis
which is partly calcareous and partly membranous. Taken together, both
these sections form a saucer-shaped structure, with the violet-coloured,
calcareous section as an outer rim or flange and the dark membranous part
occupying and filling over the central hole (see side view of whole shell in
Text-fig. 3c). The width of the calcareous flange and, consequently, the sizes
of the central hole vary. However, no specimen has been seen in which the
central area was ever completely bridged over by limy structures.

Often the barnacle is found to be fastened over a small depression in the
substratum and this applies, both in the softer rocks like calcarenite (so-called
beach-rock) and in the much harder, dark volcanic rock of Hawaii (in a batch
collected by Miss Isobel Bennett at Nanakula Beach, Honolulu, Oahu), or in
material from the New Hebrides. The limitations placed on diametric growth
in the three species C. hembeli, C. calcareobasis and CO. intertextus, by secondary
and internal calcification have been discussed by Newman (1961) and pose
Some very interesting questions as to the possible function of a “ too-large ”
Shell, of which the internal volume would appear to be progressively reduced,
as the barnacle grows older. This is, in general, quite the reverse of what
happens in the Family Balanidae. The suggestion is here made that its function
may be concerned with reproduction for, when individuals are brooding nauplii,

2 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

Fig. 3. Species of Chihamalus with flattened labrum and with mandibles generally with three
major teeth. (a-d), Ch. intertextus, (a-b) half shells seen from above at different stages of shell
erosion, (a) being moderately eroded and juvenile with distinctly interlocking and overlapping
sutures with “‘ chevron ’’ appearance, (b) a moderately eroded older specimen with “* chevrons ”’
worn away; secondary calcification has produced a deeply violet coloured shell and sutures
with rounded interlocking processes ; (c) side-view with calcareous basal rmg of basis depending
below shell; (d) labrum (front view) with right palp removed to show pocket, funnel-like, semi-
circular protrusion, channelled towards the toothed groove leading to the mouth; (e-h) Ch.
withersi, (e) shell from above, (f) side-view of shell in silhouette, (g) rostro-lateral and (h) lateral
shell plates seen from inside, showing pocket-like outpushings of shell (occasionally they may be
secondarily calcified and be represented by a callus) ; (2-7) Ch. caudatus, (2) shell from above with
toothed sutures and (7) shell im side-view silhouette.
F. J. Beeman del.

ELIZABETH C. POPE 33

the space round the soft body is completely filled up by developing young.
The secondary sealing-off of the shell may in some way reduce evaporation
or help to retain moisture for brooding young.

Scutum and Tergum

Erosion of these shell plates leads to the loss of all trace of lines of growth
on the external surfaces of the opercular valves. Their most distinctive
feature, however, is that secondary calcification leads to the fusion of each
scutum with its tergum and the loss of any clearly defined line of articulation
between them, in adult specimens. In younger individuals (carino-rostral
diameter of 9-0 mm.) there may be no trace of the former suture in the apical
parts of the valves, while a trace may remain nearer to their basal margins.
It is, however, generally impossible to separate the scutum from the tergum
without breaking the valves at some point, other than the old suture. Inter-
nally they show little definite structure, except that areas of shell where muscles
attach are generally paler in colour than the remaining areas of deep violet.
This is specially noticeable in the case of the four crests for attachment of the
tergal depressor muscles and for the scar for the adductor muscles of the scuta.
The inner side of the tergum is generally considerably darker than the scutum
owing to greater secondary calcification. Both valves show internal pitting.
While the scuta and terga are firmly interlocked in C. calcareobasis and C.
hembeli, they are never ankylosed, as in the present species.

Soft Body

If the calcareous flange of the basis is well-developed, the soft body may
be completely obscured from below until the basis is broken away. In younger
individuals, where secondary calcification is only just beginning, the soft body
lies exposed and is seen, in material newly preserved in alcohol, to have a pale
hyaline blue colour, with occasional darker blue areas, e.g. both the rami of
cirri I and II and especially the inner surfaces of the pedicels of the remaining
cirral pairs. The tergo-scutal flaps are a navy blue with lighter brown margins.
The membrane lining the shell is black or navy blue, as is the membranous
part of the basis. The outer ends of the trophi are a pale brown, in contrast
to the general pale blue of the rest of the body, and to the bluish semicircular
funnel-like process on the labrum, above the groove.

There is no caudal appendage and the penis, which is of moderate length,
is pale brown proximally, fading out to white towards its tip. It shows
a ringed structure. The eyed-nauplii are cream-coloured. It should be
remembered that these colours refer to recently preserved material, as the
author has not handled live material.

Trophi

Viewed from in front, the labrum carries a distinctive, semicircular
projection, curved round to form a funnel or scoop rather than a rounded and
bullate structure. The anterior surface of the labrum, above the palps, is
mottled by dark punctate markings giving it an appearance similar to that
seen in certain beach-worn specimens of brain corals. It may be blue or
bluish-brown and is well furnished with muscles (internally). The actual
sroove of the labrum carries a row of peg-like teeth with a fringe of hairs above
and is a wide, shallow ‘‘—”’-shaped structure. The palps are carried adpressed in
slight depressions below the ‘“ funnel” which is seen to be channelled towards
the groove of the labrum, leading into the mouth (Text-fig. 3d). The palps
are a8 described by Darwin, and the mandible is similar to Hiro’s illustration
(1939, fig. 2A), with three major teeth above, and a coarsely toothed lower
angle. Teeth 2 and 3 carry secondary teeth along their upper margins.
Mazilla I varies only slightly from Hiro’s (1939) fig. 2B. In the New Guinea

Cc

34 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

material, there is clearly a pair of smaller spines between the notch and the
two much larger ones above. Mazilla II is swollen, carries numerous long
bristles, but the distinct notch is free from bristles.

Corri

The first two pairs of cirri are much shorter and more bristly than the
remaining four pairs, and have both rami dark blue, although their pedicels
are generally light coloured. In several individuals the posterior ramus in each
of cirri I and II had been damaged and the regenerating rami were only
approximately 1/3 of the length of the normal anterior ones. However, this
was exceptional. Normally eri I and IJ have their rami subequal, with
cirrus II slightly longer than I. The anterior ramus in both these cirri was
slightly longer and broader than the posterior one, in spite of the fact that
they normally contained the same number of segments—varying between 5 and
7 (or even more, in very large individuals). Marked inequality in the numbers
of segments between the anterior and posterior rami of any one cirrus is
generally indicative of recent damage and regeneration. Both cirrus I and
II bear longish, pinnate setae anteriorly on their distal segments. There are
generally many more of these pinnate bristles on the anterior than on the
posterior ramus. In addition, the terminal segment and some of the more
distal segments of the anterior ramus of cirrus II carry bunches of short stout,
forwardly-directed grapple-like spines with paired lateral hooks, round the
bases of the longer, fine pinnate ones. They resemble the short stout spines
seen in Chamaesipho columna or, more especially, those of Chamaesipho brunnea
which were first illustrated by Moore (1944, Plate 46). They are very numerous
on the most distal four or five segments of the anterior ramus and their numbers
fall off towards the sixth or seventh segment, below the tip. Their structure
is shown in Text-fig. 1f, and from circumstantial evidence their function is
believed to be to hold and concentrate the food particles, after they have been
sieved out of the water by the bunches of pinnate spines. Cvirri IJI-VI are
of structure normal to the group except for the three or four basal segments
of cirrus III, which are furnished with grapple spines similar to those on
cirrus II. Their rami are sub-equal in length and numbers of segments and
the more central segments of most rami carry four pairs of long spines anteriorly
with an occasional exceptional one with five pairs. In this last case the fifth
pair is generally very short and visible only under high magnification.

The batch of C. intertextus from Papua, New Guinea, collected by Miss
Judy Bryan, had obviously been taken after a period of feeding and in many
specimens the cirri were clogged with food particles which were arranged in
such a manner as to suggest that the pinnate spines, acting together, function
much as gill-rakers do in fish, and the grapple-like spines, at their “ roots’,
hold and concentrate the food.

Habitat

Specimens examined in the British Museum (Natural History) were
labelled ‘‘ Philippines—ex Museum Cuming” and presumably this batch is
the Type material for Darwin’s species. Some of them were growing on a
piece of coral-boulder or calcarenite and were aggregated so closely together
that they were a little reminiscent of the honeycomb-like aggregations of
Chamaesipho columna, to be described below. However, the shells of these.
crowded individuals did not appear, from inspection, to have coalesced as
happens in similarly crowded C. columna.

The specimens from Papua, New Guinea, were growing on intertidal rocks,
near mean tide level (fide Miss J. Bryan, who collected the first batch of
specimens in this area).

Hiro (1939, p. 252) records C. intertextus as growing on “ the under sides
of rocks, together with Octomeris brunnea”’ in Formosa. Utinomi (formerly

ELIZABETH C. POPE 35

Hiro) in his 1954 account of the cirripedes of the Tokara Islands, and Tokioka
(1953) give additional ecological information about its occurrence in the mid-
tidal zone (along with the pulmonate, Siphonaria subatra Pilsbry) where it is
one of the zone-indicators, in the area of shore, below the littorinid zone and
above the Ostrea zone.

Distribution

Australasia: Examples of C. interteatus, taken in this zoogeographical
area, are from Idlers’ Bay near Port Moresby in Papua, New Guinea, and were
collected by Miss Judy Bryan. A later batch from Mr. W. Filewood confirmed
that they are plentiful among the boulders there. It has not yet been taken
along the northern shores of the Australian continent.

World Occurrence: Chthamalus intertextus is recorded from the Philippines
Archipelago—(the type locality of Darwin, 1854) ; in the Bay of Kankamaraan
on the south coast of Kangean Island, in Indonesia (N. of Java). This latter
was considered a doubtful record by Hoek (1913), for it was allegedly “ dredged
in 22 m., from a muddy bottom ’”’, by the Siboga Expedition. This is indeed
very unlikely, because intertextus is an intertidal species. It is considered
that there is little doubt that Hoek’s identification is correct and that the
general locality where it was taken is correct. However, the data about the
dredging is doubtful. Other records are as follows :—Diamond Head by
Pilsbry (1916, 1927), and several other localities on Oahu Island, Hawaii,
recorded in collections of various museums of the world; at Kaiko, Taiwan
(Formosa), by Hiro (1939); at Genka and Benoki, W. coast of Okinawa-zima,
in the Ryukyu Islands by Utinomi (1954), and at Kapingamarangi Atoll, S.
of the Caroline Islands, by Newman (1961). A batch, showing weathering
half-way between the uneroded form (with chevron-like sutures) and extremely
worn individuals in which much secondary calcification has occurred, was
taken by Dr. Lane at Lolowai, Oba Island, New Hebrides, on volcanic rock.
One of these is illustrated in Plate 1, fig. 1. The present author has also had
a single, dried shell from Fiji Islands sent to her, but the exact locality within
the Fiji Islands is unknown. Gruvel (1912) has also recorded this species from
several islands in the Tuamotu Group. It will thus be appreciated that C.
intertextus ranges eastwards across the Pacific from the Philippines and Indonesian
Islands to Hawaii, through the Melanesian and Micronesian Islands.

CHTHAMALUS CAUDATUS Pilsbry, 1916
(Plate ii, figure 4 (habitat); Text-figures 1l,e and 3,?-7)

Chthamalus caudatus Pilsbry, 1916, figured ; Nilsson-Cantell, 1921, figured,
1930, 1932; Hiro, 1937, figured; Endean, Kenny and Stephenson, 195€ ;
Stephenson, Endean and Bennett, 1958; Zevina and Tarasov, 1963, figured.

Chthamalus sp. Withers, 1932.

Ecological surveys of Queensland shores revealed the widespread occurrence
of Chthamalus caudatus along the eastern Queensland coast whereas before, it
was only recorded in Australian Museum collections at Brampton Island (off
Mackay) and Hayman Island. The results of these surveys, published in 1956
(Endean. et al.) were the first literature records of C. caudatus as part of the
Australian fauna. Later it was also found during the resurvey of Low Isles
by W. Stephenson, Endean and Bennett (1958). Its somewhat atypical
occurrence at Low Isles may be explained by the presence on that reef of a
stand of Rhizophora and Bruguiera mangroves, for it is only on the roots and
trunks of these trees that C. caudatus finds a suitable substrate, within the limits
of its vertical range, where there is adequate shade.

Although C. caudatus has been adequately described and figured by Pilsbry
(1916), Nilsson-Cantell (1921) and Hiro (1937), it seems desirable to append

36 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

a short description of Australian material, since they may grow to one and
one-half times the size of previously recorded specimens and slight differences
in structure appear with age.

Appearance and Shell Structure

In spite of a superficial resemblance in shape and colour to eroded specimens
of Chthamalus withersi or to Chthamalus malayensis, OC. caudatus is readily
distinguishable by the possession of (1) zigzagged sutures between the shell
plates (distal to the sheath area of the shell) and (2) a pair of caudal appendages.
Tf the toothed sutures are not easily seen externally, they can usually be seen
internally. The caudal appendages are disclosed by dissecting the soft parts
away from the shell and looking between the bases of the pedicels of the cirri
VI and it may be necessary to use fine needles to separate them. They are
quite long.

The colour of the shell is generally a light ash grey, with wavy ledges of
yellow lamina, or darker lines showing between the successive layers of the
shell plates in eroded individuals (see Text-fig. 32). The outer rim of the shell,
however, is often a dark horny-brown and similar in colour to the shells of
moderately eroded C. witherst. As the wavy toothed sutures may sometimes
be visible only internally, it is possible to make a mistake in identification of
Chthamalus from the high intertidal zone. The rhomboidal orifice is compara-
tively large; the shell is generally oval in outline or may sometimes tend
towards a rectangular shape, and this is reflected in the table of measurements
(Table 4), in which the carino-rostral diameter always slightly exceeds the
breadth of the shell. The flattened low conical shape is reflected in the height
measurements. These were taken at the carinal end since this valve is generally
a little higher than the rostrum (Text-fig. 37).

TABLE 4
Dimensions of Australian Chthamalus caudatus

Locality Carino-rostral Breadth of Height in
diameter m mm. shell in mm. mm.
Poimt Vernon, South 12-0 10:5 2-5
Queensland

10-5 8:5 2-4.

e 12-1 11-5 2-2

Curtis Island, Q’ld. 15-0 10-0 2-4

~ 13:3 13-0 2-5

Low Isles, Queensland 13-1 12-0 2-9

” 15-0 12-0* 3.3

Cockatoo Island, 6-0 5-5 2-0
Yampi Sound, W.A.

as 6:5 5-1 2-0

* Growth laterally distorted by crowding.

In larger-sized barnacles, the outer margins of the shell plates often form
a broad flange-like area round the central cavity which rarely extends beyond
the immediate area, roofed by the opercular plates. On the lower surface of
this flanged area, the edges of the alae may thicken and grow downwards
towards the basis forming wavy, butress-like structures running from the lower
edge of the sheath to the circumference of the shell. Similar structures in
CO. depressus (Poli) have been described by Utinomi (1959, p. 393).

Internally the shell colour is either a dark purplish-grey or light brown,
while the soft body is light coloured with the cirri greyish. There are patches
of dark pigment round the bases of the spines on each segment. Developing
larvae are a creamy yellow. The sheath passes insensibly into the lower part
of the parietes. The basis is dark, often black, and completely membranous.

ELIZABETH C. POPE 37

The shell plates are fairly smooth externally and show little trace of ribs
or folds even near their outer margins. There is also little trace of radii
externally but the alae of the carina and rostrum are especially well developed.
The carina was taller than the rostrum in all specimens measured.

Opercular Valves

There is considerably more sculpturing in the opercular valves of well-
erown Australian C. caudatus than is shown in Pilsbry (1916, Plate 73, figs
1-la). In an uneroded barnacle, lines of growth are prominent on both the
scuta and terga and the layers of yellow lamina alternate with the layers of
calcareous matter.

The scutum has a longitudinal fold from the apex to the basal margin
externally and the tergum may have pits shaped like inverted ‘‘ V”’s near
their apices, for the reception of the tip of the scuta. However, most specimens
are too greatly eroded to show these structures. Instead, the well-marked
articular ridges and furrows between the scuta and terga are clearly to be seen
and a central, wide groove appears between the two scuta formed by the
inflected occludent margins. The darker coloured, underlying layers of the scuta
and terga may show through where the ash-grey shell matter has been eroded.

Internally the scutum is often yellow centrally, with a dark purplish margin
bordering the valve and there may be some pitting. The incurved basi-scutal
margin forms a small pit for attachment of the lateral depressor muscles. The
pit for the adductor muscle is clearly defined but has little suggestion of. a
ridge below it.

The tergum is generally darker internally than the scutum and is wider
above the articular furrow. There are a variable number of rather irregular
crests (five or more) for the attachment of the tergal depressor muscles and
they project quite noticeably below the basal margin. Internally a deep, narrow
groove runs longitudinally down the centre of the tergum but does not extend
right to the apex, though in larger barnacles it may reach the basi-scutal angle.
In a C. caudatus with a carino-rostral diameter of 6-5 mm. a groove is beginning
to develop, whereas in one with diameter 12-1, it is a marked feature of the
shell. The articular furrow is well developed.

Soft Body
The most interesting feature of C. caudatus is the pair of long, slender
caudal appendages, a feature not recorded in any other species in the genus.

Trophi

The labrum, when viewed from in front, has a distinctive, flattened, semi-
circular process above the palps, somewhat similar to that in C. intertextus. The
anterior surface of the groove of the labrum carries numerous triangular denticles
along its whole length and there are numerous fine hairs, especially in a central
area above the denticles. The semi-circular funnel-like projection is well
provided with muscles. Palps——The long bristles of the upper anterior
corners of the two palps overlap slightly centrally. The palps are clavate
and are as illustrated by Nilsson-Cantell (1921, Text-fig. 57a) but the Australian
Specimens tend to have many more bristles than are shown in his illustration.
The mandible has three main teeth without secondary side-teeth on them and
a pectinated lower angle in which the second spine from the lower angle is
longest and the 8 or so spines above it are nearly equal and similar in size to
the lowest one. There is little or no variation from Pilsbry’s illustration (1916,
fig. 92c). Mazxilla I has two distinct notches anteriorly so that the spines are
divided into three groups. Its shape and the arrangement of the spines are
closely similar in Australian specimens to Pilsbry’s drawing (1916, fig. 92b).
The only difference seen in the Australian material is the presence of three pairs

38 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

of shorter spines situated immediately below the two large upper spines above
the upper notch ; and the presence of a series of fine hairs along its lower border.
Mazilla II has a wide notch, free of spines, and is very similar in shape to that
depicted by Nilsson-Cantell (1921, Text-fig. 57b) except that larger Australian
specimens tend to have more long bristles along the straightish lower border
(Nilsson-Cantell’s figure is reversed and shows the lower border, above).
Viewed from in front, before dissection, C. caudatus has a pronounced “ beard ”’
hanging from the two maxillae IT.

Cirrt

In cirrus I the rami were unequal, the anterior one being longer and stouter
than the posterior one (segmental numbers of the order of 8 and 6 segments
respectively were found or, in a small specimen of 6-5 mm. diameter, from
Yampi Sound, Western Australia, 7 and 5). The more distal segments had
pinnate spines in addition to ordinary setae. The shape of cirrus II apparently
changes with increase in size. A small Yampi Sound specimen had the
anterior ramus slightly longer and certainly stouter than the posterior one,
as in cirrus I, although the numbers of segments in the two rami were almost
equal, e.g. 7 or 8 anteriorly and 8 posteriorly. In a larger individual the
posterior ramus was longer and slightly narrower than the anterior one and this
was reflected in the segmental count of 8 (anterior) and 11 (posterior). A still
larger specimen showed a tendency for the posterior ramus to grow even longer.
Again, there are some finely pinnate long spines on some segments of the anterior
ramus (Text-fig. le) but no trace was found of the short stout pectinate spines
of the type found in some other species in the genus. Cirrus III had sub-equal
rami and frequently equal numbers of segments, ranging from 13 to 18, in the
material examined. Segments of both rami were found to carry 3 or 4 pairs
of longer spines—often 3 pairs on the anterior one and 4 on the posterior but
this was not invariable. Cirri IV, V and VI each had sub-equal rami and
the numbers of segments of the anterior and posterior rami were nearly always
equal, except where there was evidence of damage and regeneration. There
were generally 4 pairs of spines on each segment of the anterior ramus and
sometimes 5 pairs on the segments of the posterior ramus of the same cirrus.
The two smaller pairs of spines are only seen under high magnifications.

The penis was moderately long (longer than cirrus VI even when slightly
contracted). It had a ringed appearance which was more pronounced when
it was greatly contracted. Distally it was not ringed and carried scettered
hairs along its length, with two tufts of fine hairs at its tip.

The caudal appendage is generally long, thin, and at least half as long as
cirrus VI or slightly more. At the junction between segments a series of fine
and, in some cases, long hairs ring the segments. Each segment generally
has a central, dark cross-marking on it. In a small specimen of 6-5 mm.
carino-rostral diameter there were 17 or 18 segments in its caudal appendages,
whereas one with the diameter of 12-0 mm. had 22 or 23 segments. Pilsbry’s
type specimen (U.S. National Museum 48087) is intermediate both in size and
number of segments in its caudal appendage, so that it would appear that the
number of segments in the caudal appendage increases propottionately with
shell growth.

Habitat

The occurrence of C. caudatus, within its geographical range, depends on
the presence of adequately shaded areas in the upper intertidal zone. Its
vertical range in Queensland is from mean high water (occasionally it may
occur higher where conditions are particularly favourable for its settlement)
down to approximately mean low water level. Its range thus overlaps those
of Chthamalus witherst and Chthamalus malayensis but these species can grow
fully exposed on the rocks whereas C. caudatus tends to be cryptic in habit.

ELIZABETH C. POPE 39

Distribution

Australian: Along the eastern coast of Queensland where suitable habitat
conditions prevail, from Point Vernon (near Great Sandy Island) in the south,
northwards along the mainland coast and on the high islands inside the Great
Barrier Reef to Port Douglas in the north. Collecting ends at this point on
the Queensland coast and it is likely that C. caudatus occurs right up the rocky
coast and round into Torres Strait. It also is recorded in the north of Western
Australia at Yampi Sound, on Cockatoo Island. No organized barnacle
collecting has been done east of this point towards Cape York. The absence
of records for this species in this area at present is therefore of little significance.
Withers (1932) recorded a species of Chthamalus as a sub-fossil in beach rock
from Magnetic Island, Queensland. It was found on examination to be C.
caudatus.

Extra-Australian Records: A few sporadic records of C. caudatus occur
in most areas of the Indo-Malayan Archipelago as follows :—Catabalonga,
Samar Island in the Philippines (Pilsbry, 1916); West coast of Sumatra and
at Pisang Island (opposite Maccluer Gulf) in West Ivian (botn records by
Nilsson-Cartell 1921 and 1930 respectively) ; the South China Sea (Zevina and
Tarasov, 1963); and Arakabyu Island in the Palao Islands (Hiro, 1937).

CHTHAMALUS WITHERSI Pilsbry, 1916
(Plate u, figures 1, 2, 5; Text-figures 1,d and 3,e-h).
Chthamalus withersi Pilsbry, 1916, figured ; Nilsson-Cantell, 1921, figured,
1930, 1932, 1938; Broch, 1931, listed only ; Hiro, 1937, figured ; Kolosvary,
1941; Endean, Kenny and Stephenson, 1956; Endean, Stephenson and
Kenny, 1956; Zevina and Tarasov, 1963, figured ; Southward, 1964.

? Chthamalus rhizophorae de Oliveira, 1940, figured, 1941, figured.

Having ex amined the Type of specimens of Pilbry’s Chthamalus withersi
(U.S. National Museum Number 48088 which included a Holotype and a
series of paratypes), it is obvious that his original (1916) description was based
on juvenile specimens. Now that a large series of C. withersi from Australia,
and the islands to the north of it, have been examined, it is necessary to emend
his description to include characters not usually evident in the barnacles until
the full adult size is reached. Some hint of the changes seen in larger specimens
was given by Nilsson-Cantell (1921, p. 296) who examined a specimen with a
carino-rostral diameter of 13-0 mm., whereas Pilsbry’s largest had measured
9-5 mm. in diameter. The collections examined by the author have, in addition
to the many small specimens with size ranges similar to those handled by Pilsbry,
many barnacles of large size, ranging in carino-rostral diameter from 10-0 mm.
to 21-0 mm. The largest specimens were collected by the Dutch Snellius
Expedition of 1929-30 on mangroves, at various points in the Celebes and
other nearby islands.

The largest Australian C. withersi seen so far also occur on mangroves,
in the Cairns district, Queensland. Here specimens of up to 16-5 mm. carino-
rostral diameter were taken by the author and careful searching would no
doubt, have revealed even larger specimens in the area.

Many details of occurrence, population size, interspecific reaction toa
second Chthamalus species (C. malayensis), and its environmental preferences
in Queensland are described in the two papers recording the ecological surveys
of Endean et al. (1956). It is extremely common on the upper parts of whart
and bridge piling and on the roots and lower trunks of the red mangrove,
Rhizophora, in Queensland as may be seen in Plate ii, fig. 1, 2, 5.

When the structures of full-grown C. withersi, newly recorded below, are
taken into account, the Brazilian species Chthamalus rhizophorae de Oliveira
(1940 and 1941) is found to resemble it extremely closely. The South American

40 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

Species also occurs on mangroves and has, so far, only been recorded in the
vicinity of Rio de Janeiro, a port. The possibility of its having been transported
by shipping from the Indo-Malayan area cannot altogether be ruled out,
especially as C. withersi may be regarded, in Queensland at least, as a fouling
species on wharf piles and therefore capable of being transferred to, and
transported by, shipping.

The author has unfortunately not been able to examine de Oliveira’s type
material, lodged in the Instituto Oswaldo Cruz, but has examined specimens
from Cananeia (Brazil, near latitude 25° S.), identified as C. rhizophorae by
Dora P. Henry, in the collections of the U.S. National Museum, Washington
D.C. (Number 101193). A large example of these, with a carino-rostral
diameter of 20 mm., was so similar in shell structure to the largest C. withersi
in the present Australian series, that it is suggested that C. rhizophorae de
Oliveira may have to be synonymized with C. withersi Pilsbry. The soft body
parts were not, however, available for dissection. In the absence till now in
literature of a description of fully-grown C. withersi, de Oliveira’s erection of
a new species for his South American material was logical and it will remain
to be seen if the new African species of Stubbings (the description of which
is said by Southward to be in press) is indeed new or whether it is in fact C.
rhizophorae or C. witherst.

Pilsbry’s description, with additions by Nilsson-Cantell (1921), is adequate
for the general run of C. withersi material collected on coastal rocks and wharf
piles on the east coast of northern Queensland and at Darwin, Northern
Territory (no organized barnacle collecting has been done between these two
areas), but their accounts are inadequate for specimens which have reached
full maturity (carino-rostral diameter 15-0 mm. upwards). Transitional
Stages during growth show gradual development of the structures present in
fully adult barnacles, so that any lingering doubts as to the inclusion of the
largest specimens in C. withersi Pilsbry are dispelled.

Emended description of C. withersi Pilsbry, 1916

Table 5 sets out a series of shell measurements of C. withersi of varying
sizes, ranging from 10-0 mm. to 21-0 mm. carino-rostral diameter.

It is clear from this table that Pilsbry’s material was far from full grown.
It shouid also be mentioned, that evidence of breeding is by no means a
criterion of full maturity in this species.

Appearance and Shell Structure

Chthamalus withersi has a much flattened conical shape; the basis is
generally sub-circular and wavy in outline, as shown in Text-fig. 3,e,f. The
carino-rostral diameter is in general slightly greater than the width of the
Shell. The shell tends also to be slightly narrower towards its scutal end,
when growth has been unrestricted. The general colour may vary from
cinnamon brown (uneroded or juveniles) to ash grey in adults.

In (. withersi, growing on mangroves, the outer layer of bark may appear
to ‘“‘ ride up” over the shell rim, as though the growing shell were ploughing
into it. However, in full-grown specimens this effect is rarely seen and the
barnacle appears to be attached, in the normal way, on top of the bark.

The shell tends to be slightly higher towards the rostral end, but this is
not invariable. The smooth shell plates are articulated very loosely, even
when the shell is more than 12 mm. in diameter, and this applies to the opercular
valves also which, when eroded, never display the capital psi (¥)-shaped
articulation pattern seen in so many other species of Chthamalus. This is
rather unusual in a genus in which the scuta and terga are so often interlocked
strongly together. Worn shells show a comparatively straight line of articula-
tion between the scuta and terga.

ELIZABETH C. POPE 4]

Older C. withersi may show low, poorly developed regular folds or ribs,
while juveniles show broad wavy folds, often visible only towards the outer
dark rim where growth is active and the shell is not yet eroded. Radii on
the shell plates cannot be seen externally, but in larger shells the alae of the
carinal and lateral valves are markedly developed.

The orifice is moderately large in full-grown specimens and almost diamond-
Shaped. In medium-sized and smaller individuals it tends to be rhomboidal
in outline, with the scutal angle somewhat rounded. Often 4 or 2 small
projecting white teeth occur where the summits of the lateral and rostrolateral
valves project slightly into the orifice. Internally these tooth-areas are white,
as opposed to the usual purple-grey or brown colour of the rest of the interior
of the shell. Sometimes the whole inner shell is white but even in these the
tooth-section is still distinguishable from the rest. The teeth usually mark

TABLE 5
Shell measurements of C. withersi

Carmo-rostral Width of shell Height of shell

Locality diameter at right angles at the rostrum
in mm. to carmo-rostral im mm.
i0 mm.

Halfmoon B., near

Cairns, Queensland, 10:0 8-0 Bo jl
Australia,
a 15-1 14-0 4-5
ws 16:0 15:0 3-6
4s 16-5 16:3 5-4
Darwin, Northern
Territory, Australia 11-0 10-0 1-5
ee 11-0 10:0 3:0
Fe 14-0 12-0 3-9
Mamudja, Celebes,
Indonesia 15-4 13-6 3-5
Bs 16-0 14-9 4-9
5) 17-0 18-4 5-0
(This diameter
“ distorted ” by
crowding)
A 19-9 - 16-0 5-0
FS 21-0 19-0 5:0

the summits of light-coloured stripes on the lateral and rostrolateral plates.
These four light stripes show best in uneroded, cinnamon-coloured individuals
but, while rarely evident in large eroded, grey-coloured specimens, the four
white tooth-patches still generally show internally.

Pilsbry (1916, Plate 73, fig. 2e) describes and figures the internal surface
of a rostrolateral compartment showing a sunken oval pit or callus below the
white summit of the valve. Such are indeed common in C. withersi, occurring
even in juveniles. As growth proceeds, however, a second and even more
distinct pit may appear on each lateral valve also (see Text-fig. 3,g,h). Some-
times the largest individuals show pits only on the laterals, those of the
rostrolaterals tending to be filled up, at maturity. This seems most marked
when the sideways growth of the rostrolateral valves is restricted. Perhaps
these pits are associated with breeding in some way, by providing additional
room for brooding developing nauplii.

Scutum and Tergum

In Australian specimens up to 10 mm. in carino-rostral diameter, the
opercular valves have few outstanding characters. There are, however,
generally considerably more structures than shown in the illustrations of

42 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

Pilsbry (1916, Plate 73, figs. 2a—2d) and Hiro (1937, fig. 4h, 7). There is an
oval pit for attachment of the adductor muscles, shallow but quite discernible ;
and a small, outwardly-curved surface with an extremely shallow pit on it,
near the basitergal corner of the scutum, where part of the lateral depressor
muscle is attached. The scutum becomes quite thick and strong in mature
individuals. The occludent margin always seems to be thickened and rolled
inwards, as shown by Pilsbry. In no specimen, however mature, was the
articular ridge or furrow ever developed to form anything more than a shallow
wavy fold and trough where it articulates with the tergum. It does not
interlock strongly like the pieces of a jigsaw puzzle, as happens in most other
species of Chthamalus. The articular ridge and furrow of C. withersi is more
pronounced in juveniles than in mature specimens which is unusual. In some
uneroded specimens there may be a light stripe on the tergum, parallel with
the scutal margin. The tergum is distinctive among species of Chthamalus
found in Australia in that it is nearly triangular, with the scutal and basal
margins nearly equal and both much longer than the carinal one. The basal
margin is slightly sinuous in well-grown individuals and, although there is a
slightly convex curve towards the basiscutal corner, it cannot be regarded as
a spur. The interior of the valve in adults is white and grey or purplish-grey
and is generally somewhat fretted and pitted, parallel to the margins. Along
the scutal and carinal border the inturned edges tend to become thickened
and to grow almost at right angles to the main plane of the valve. This is
especially true for the scutal margin, with the result that the tergum has a
deep V-shaped furrow along its length. The number of crests for the attach-
ment of the depressor muscles is amazingly constant throughout the size range,
being four, unless the valve has been damaged. Externally, if uneroded,
simple lines of growth may be seen on both scutum and tergum and, in juveniles,
traces of a hairy outer lamina may persist.

Soft Body

As in many chthamalids the animal body is very much smaller than the
shell-size would suggest. The tergo-scutal flaps are brown, but most of the
soft body is a pale creamy brown except for the following darkened areas :—
the outer sides of cirri I and II, the pedicels of cirri III to VI which are
darkened on their inner sides. The proximal part of the penis and a ring
round the mouth opening are also darker, and patches of dark pigment occur
on the segments of the rami of cirri III to VI round the bases of the spines.
The mouth-ring is made up of a broad, darkened band along the arched groove
of the labrum and the outer ends of the trophi—especially the tips of the second
maxilla which form the lower arc of the circle. However, the colour fades
fairly rapidly in mounted specimens. One specimen from Mamudju, Celebes,
taken in August 1929 by the Dutch Snellius Expedition, had large developing
embryos which were bright yellow. The basis is wholly membranous.

Trophi

The labrum is in general somewhat flattened anteriorly and semi-circular
Shaped above the palps, not truly bullate but not as distinctly channelled
towards the shallow U-shaped groove above the mouth, as in C. intertextus. This
groove is bridged across by a straight, clear chitinous structure which carries
at its centre a row of 28 or so peg-like teeth. Neither Pilsbry’s nor Nilsson-
Cantell’s illustrations show the structure of the labrum as it appears in specimens
examined from various Australian localities or from the Celebes, Indonesia.
It would appear that Pilsbry’s drawing (1916, fig. 91D) was made from below,
so that the normal U-shaped groove above the teeth and the flattened front
of the upper part of the labrum were obscured, while Nilsson-Cantell’s (1921,
Text-fig. 51a) fails to show the straight surface from which the teeth arise.
Australian specimens also show several triangular denticles on either side of

ELIZABETH C. POPE 43

the peg-like teeth, in well-grown specimens, but their size and number seem
to depend on the size of the barnacle. In a medium-sized specimen from
Mamudju, Celebes, up to six denticles occurred beside, and at a higher level
than, the central peg-like ones. There is also a fringe of hairs above the teeth,
which are seen even in smaller specimens. The palps are basically the same
as Shown in Nilsson-Cantell’s drawing (1921, Text-fig. 51b) except that the lower
anterior corner is generally slightly more rounded and there are considerably
more bristles on it. Their arrangement is, however, correctly depicted by him,
with longest bristles on the lower anterior part getting progressively shorter
towards the upper border and along the upper margin. Mandible.—There is
variation in the number of major teeth on the mandible, from 3 to 4 being the
usual range, though the latter number is encountered only in very large
Specimens and then only occasionally. The most frequent count for the large
teeth is 3, as stated by Pilsbry. However, several specimens from the Celebes
and Northern Queensland had 4 teeth. In one individual from Mamudju in
the Celebes, with an estimated carino-rostral diameter of 20-21 mm. (shell
damaged slightly), the mandibles had 3 large teeth on one side and 4 on the
other. In the latter, the 4th large tooth had obviously been derived by the
enlargement of the uppermost denticle of the pectinated lower angle. A
smaller individual, from Darwin, Australia, was found to have mandibles with
2 and 3 large teeth on the right and left sides, respectively. It would appear
from this evidence, that the numbers of large teeth on the mandible is not
definitive in C. witherst until the animal is fully grown. However, the majority
of specimens of this species will be found to have 3 large teeth on the mandible.
Mazilla I has its spines most distinctly divided into 3 groups by two well-marked
notches. In the upper group two large, strong spines are followed below by
two pairs of small spines, above a U-shaped notch. The central group of
medium-sized spines comprises 6-8 pairs, arising from the edge of the jaw which
is usually fairly straight in this section of maxilla I—not curved as in the lobes
above and below it. Below them is generally a V-shaped notch. The lower,
protuberant corner of maxilla I carries from 5 to 6 pairs of small spines,
comparable in size with those situated just above the upper notch. This
structure is substantially the same as was described and figured by Pilsbry
(1916, fig. 91lce) and the only character noted in the present material, not
figured by him, is the distinct fringe of hairs near the outer tip of the lower
margin. Mazilla IT has not been figured and Nilsson-Cantell’s description
(1921, p. 296) does not mention the large number of bristles present on the
anterior or free border above and below the wide, shallow notch. There is
also a dense patch of rather long bristles arising low down on the outer side
of the organ.

Cirri

The first two cirri are short and darker in colour than the remainder.
Cirrus I has its rami strongly incurved towards the mid-line, so that its effective
Sweeping action is almost at right angles to that of Cirri IJI-VI. Several of
the more distal segments and the terminal segments of both Cirri I and II
carry numerous pinnate setae, the structure of which is shown in Text-fig. 1d.
They are especially numerous and clumped along the inner sides of the rami
where their felted mass forms an efficient straining organ for food catching.
Ip many of the preserved specimens, almost every fine side-hair on the pinnate
spines had a particle entangled with it, so as to give the spines a knobbed,
rather than a plumose, appearance. It was at first thought that C. withersi
possessed spines intermediate in structure between pinnate and the lanceolate
Spines seen in C. challengeri or C. antennatus. However, in individuals free
from entangled food particles, they were found to be merely normal pinnate
Spines, the combined effect of which was to serve a function similar to that
of gill-rakers in fish. In cirrus I the anterior ramus is longer by at least two

44 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

of its segments and considerably stouter than the posterior one. The number
of segments in younger individuals is generally less than in older ones, with
7 in the anterior and 6 in the posterior ramus while fully-grown specimens.
tend to have 8 or 9 segments in both rami. Often segmental numbers vary
from right to left side in the same animal, so that too much importance should
not be attached to them diagnostically. Crrrus IJ is similar in general structure
to I, with the anterior ramus slightly longer than the posterior one, but only
a little wider. The numbers of segments in the two rami tend towards
equality in full-grown specimens, while in younger ones the anterior ramus
may contain one segment more than the posterior one, e.g. 7 to 8 segments
in the anterior and 6 to 7 in the posterior one. A large specimen from Mamudju,
Celebes, had 9 segments on the left anterior ramus and 8 on the right, while
both posterior rami contained 8 segments. Several individuals had 8 segments
in both rami and this seems to be the full adult number, where no regeneration
has occurred. Cirri ILI-VI are similar in structure, long and curled but III
is somewhat shorter than the remaining three pairs. Their rami are subequal
both in length and in the number of component segments of any one pair.
Anteriorly each cirral segment has a dark, pigmented area round the bases
of the three pairs of large spines arising there. Careful search, over a series
of specimens of varying sizes, failed to find any individual with more or less
than three pairs of large spines on the more central segments of the rami.
However, Nilsson-Cantell’s record of occasional cases of 4 pairs of spines (1921,
p. 296) cannot be disregarded, in view of the variation from 3 to 4 teeth on
the mandibles seen in present specimens. The penis is moderately long, dark
proximally and ringed, with scattered fine hairs along its distal section. There
are small tufts of hairs above and below the opening, on its tip. There is no
caudal appendage.

The discovery, in well-grown C. withersi, of occasional specimens in which
there are 4 major teeth on the mandible raises the question whether this species.
should still be associated with the hembeli-complex of species of genus Chthamalus,
all of which have 3 major teeth on their mandibles. However, a review of
other features of OC. withersi, and other species associated with the hembela
group, shows that it has the characteristic pectinated lower angle to its mandible,
with the largest denticle at, or near, the tip ; there is no trace of the comb-like
groups of spines as seen in the species belonging to the stellatus-complex ; it
mostly bas only 3 major teeth on the upper part of the mandible (only
occasionally 4); the distal segments of cirrus II have no stout, toothed lance-
olate spines. Instead there are numerous fine, pinnate spines on both cirrus
I and II. The balance is thus heavily in favour of retaining withersi in the
hembeli group, though it shows certain characteristics that place it in a somewhat.
intermediate position between these major sub-groups of the genus.

Habitat

Chthamalus withersi grows attached to intertidal rocks, wharf piles and.
Red Mangroves in the highest intertidal area in tropical eastern Australia
(Plate ii, figs 1, 2, 5). On rocks exposed to full sunlight, it tends to survive
only in crevices or areas of slight shade ; however, in the shaded areas of wharf
piles and on mangrove trees it reaches not only its largest size but also its
greatest density of population (see Plate ii, figs 2, 5). In Australia its vertical
range is from a little below the high water mark of spring tides down to the
level of high water neap tides or even mean sea level (Kndean, Kenny and
Stephenson, 1956). It is tolerant to considerable fluctuations in the salinity
and conditions of considerable turbidity in the seawater.

Distribution

Australian: Along the east coast of Queensland from Hervey Bay (near
Great Sandy Island) in the south, northwards right up the Queensland mainland.

ELIZABETH C. POPE AD

coast (as opposed to the Great Barrier Reef itself) to Cooktown. It also occurs
at Mandorah, near Darwin, Northern Territory. Between these two last
localities no systematic barnacle collecting has yet been done but its presence
there is likely. On the other hand, the absence of records of CO. withersi from
Yampi Sound in Western Australia, westward to Carnarvon and thence to
Shark Bay, is probably significant, since collections have been made along
this coast, at the author’s request, by trained collectors who were specially
looking for it and who found its niche on the mangroves occupied by another
species of Chthamalus, namely C. malayensis.

World Occurrence: The locality of the Type is “from a reef opposite
Cebu Philippine Islands’, Pilsbry (1916). It has also been recorded at
Tjilatjap, Java; Billiton, two new records from the Celebes area came from
collections made by the Dutch Snellius Expedition, 1929, from Maratoea
(N.E. Borneo) and Mamudju (Celebes) in Indonesia. It also occurs at Pisang
Island (opposite Maccluer Gulf in W. New Guinea); Baie van Dampelas, west
coast of Celebes; Chandipur, E. Pakistan; Bombay (in a collection sent to
the author for identification by Y. M. Bhatt of Bombay University), Thana
(near Bombay), Balasore, Orissa and Port Canningall in India; Elphinstone
Island and Port Maria, Mergui Archipelago. Otber than the specimens from
Bombay, the other records above occur in the writings of Nilsson-Cantell
(1921, 1930, 1938). Additional records are Mandarai Pier, Kororu Island, in
the Palao Islands (Hiro, 1937); the South China Sea (Zevina and Tarasov,
1963) and the following records are taken from museum collections :—on bark
of mangroves, near Old Panama; taken during the Pinchot Expedition of
1929 and identified by H. A. Pilsbry. This identification was confirmed by
the author, in 1963, at the Academy of Natural Sciences, Philadelphia, Penn.,
U.S.A. (This record near the Panama Canal suggests a possible route of
introduction of C. rhizophorae de Oliveira ? = C. withersi to Brazil.) Further
records are from Kei Islands, Arafura See; and as Chthamalus rhizophorae de
Oliveira, by Dora P. Henry at Cananeia, Brazil (in U.S. National Museum,
No. 101193, U.S. National Museum, Wasbington, D.C.).

It is thus the most widely ranging of species in the genus Chthamalus,
occurring in Australia.

CHTHAMALUS ANTENNATUS Darwin, 1854
(Plate i, fig. 4, Plate ii, fig. 7 (ringed specimen) and Text-figs 1,4; 4,a-q)

Chthamalus antennatus Darwin, 1854, illustrated ; Weltner, 1897; Giuvel,
1905, illustrated ; Broch, 1922, illustrated; Pope, 1945, illustrated ; Dakin,
Bennett and Pope, 1948, 1952 ; Bennett and Pope, 1953, 1960 ; Endean, Kenny
and Stephenson, 1956; Womersley and Edmonds, 1958, illustrated ; Wisely
and Blick, 1964.

Chihamalus antennatus was originally described by Darwin (1954, with
the shell illustrated in Plate XVIII, fig. 2, and Cirrus III on Plate X XIX,
fig. 3—nec “‘ fig. 2’ as given on page 460 of his text); by Nilsson-Cantell (1921,
1926, both illustrated) ; Broch (1922, illustrated) and by Pope (1945, illustrated).
Heological and distributional data are available in the following accounts ;
Dakin, Bennett and Pope (1948, 1952), Bennett and Pope (1953, 1960), Endean,
Kenny and Stephenson (1956), and Womersley and Edmonds (1958) and
information on the breeding period bas recently been published by Wisely and
Blick (1964).

Careful investigations of collections housed in a number of European
Museums has shown that the following records of C. antennatus are based on
misidentifications :—in Chile (Gruvel, 1905); at Amanu Is. in the Tuamotu
Group (Gruvel, 1912); at Santander, Spain (Gruvel, 1920) and at Broome in
north Western Australia (Broch, 1916).

Obvious misidentifications of Indomalayan and Pacific species of Chthamalus
are fairly common in literature prior to Pilsbry’s 1916 Bulletin. Early records

46 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

of C. antennatus in Chile, Patagonia, Spain and the Tuamotus Group are due
to misidentifications. Some characters which, in the keys given by previous
authors, would appear to be unique to certain species often are not so, and may
occur in several species, e.g. the long outer ramus on cirrus III occurs, for
instance, in C. antennatus and C. cirratus (Chile and Peru). Again, juveniles or
young of C. antennatus and C. malayensis and other species with 4-toothed
mandibles often appear superficially very alike, so that records of C. stellatus,
C. malayensis and even C. challenger: have often been confused with the present
species. In view of this, and of current interest in the genus Chthamalus and
the forthcoming regrouping of species by Zullo, some further anatomical details
are given, since this endemic Australian species is little known abroad.

Description

Chthamalus antennatus is one of the more heavily built and larger species
in the genus, although much smaller than either C. hembeli and C. calcareobasis.
Whilst the basal area of C. antennatus may be no greater than that of several
other Australian species in the genus its walls rise fairly steeply, practically
from the circumference, and its shape is that of a truncated cone. There is
no tendency to develop a flattened flange round a centrally raised area, as seen
in large C. caudatus or (. withersi. This accounts for its more massive
appearance and the larger soft body within the shell.

Table 6 gives the measurements of 20 well-grown specimens, taken from
localities noted for exposure to heavy wave action. Crowding is seen to cause
exaggerated upward growtb so that a tubular shell is formed, with the summits
of the rostro-laterals tending to form small “‘ teeth ’’ projecting into the orifice.
If growth is unrestricted the shell tends to have its length and width sub-equal,
and erosion of the tops of the shell valves may result in the formation of highly
polished areas resembling ivory. Inside, the shell is generally smooth, flesh
pink in colour and rarely pitted. The shell plates are very strongly articulated
by overlapping radii and alae which are not toothed but which may show
distinct lines of growth.

Darwin’s original account is so detailed and accurate that little need be
added except to clarify the characters needed to determine species within the
stellatus-complex. Text-figures 1,2 and 4,a-g illustrate the more important
features distinguishing C. antennatus from its nearer relatives.

Maximum sized specimens in the present series exceed the largest, so far
described, by 3-2 mm. in the carino-rostral diameter. However, there seems
to be no further modification of features described by Darwin in these larger
animals. However, some comment is called for on the fact that, while Pilsbry
(1916, p. 296) placed C. antennatus in the group, later characterized as the
hembeli-group by Nilsson-Cantell (1921), the latter author and Broch (1922,
p. 306) both placed it correctly in the stellatus-complex of genus Chthamalus.
Examination of large series has shown that most individuals have the mandible
similar to the illustration in Broch (1922, fig. 51a)—a typically stellatus pattern
of mandible. However, several large individuals in tubular shaped shells or
ones where the lower tip of the mandible had been damaged and was in process
of regenerating had mandibles in which the lower angle was coarsely pectinated
and had no clearly defined, comb-like section between the lower tip of the jaw
and the lowest major teeth. They were thus somewhat intermediate in structure
between a typical hembeli and a typical stellatus mandible. They have been
commented on by Darwin in his original description. Other shell and body
characters, however, definitely align C. antennatus with the stellatus group of
species, even in those which have atypical lower tips to the mandible.

ae

Opercular Valves

Darwin’s dismissal of the opercular valves as being “ hardly distinguishable
from those of C. stellatus”’ has encouraged certain of the misidentifications,

ELIZABETH C. POPE 47

such as that of Broch (1916, p. 14). Broch’s specimens were examined at the
Stockholm Naturhistoriska Riksmuseet (Registered No. 278) and found to be
juveniles of C. malayensis.

The first illustration of opercular valves was published by Nilsson-Cantell
(1926, Text-fig. 3) and shows few definite characters, apart from the two crests
on the tergum for the attachment of the depressor muscles. Text-fig. 4,d and
e, Shows the opercular valves of a well-grown individual. The scutum is smooth
internally and but little sculptured in comparison with C. malayensis, but it
shows two incipient crests within the depression for attachment of the lateral

TABLE 6
Measurements of 20 well-grown C. antennatus from very exposed habitats

Carino-rostral Width at Height at
Locality diameter right angles tallest part Remarks
m mm. to carimo- of shell
rostral diam. im mm.
in mm.
Harbord, Sydney, 9-5 9-5 3-5 |
New South Wales. 11-0 11-0 4-9 |“ Normal’ shape not
19.x1.1964 11-6 12-3 3°5 >restricted by crowding
12-8 14-0 3-5 |
13-5 2 3-6 J)
15-0 15-2 5-0 ahe
15-0 15-0 5-9 (enone: tall
16-0 17-4 5-3 )
16-0 18-0 a:5 > Normal shape
18-5 16-0 5-0 J
Eucla, Great Australian 8-4 7:4 5-3 On a limpet shell
Bight, Western ) Tall, tubular form
Australia 10-0 10-0 9-0 fowimg to crowding.
10-5 9E3 10-5 | Tendency to have
J) toothed orifice
12-0 11-5, 7-0 Distorted. Average
height for species
should be nearer
4-5-5-0 mm. for
shells of this size.
13-0 9-6 6-5
Cape Forestier, S.E. 11-0 10-5 4-5
Tasmania 12-0 12-0 6-0 > Normal shape
15)565) 15-0 6-7 |
16-0 16-1 HoT J
18-8 19-0 9-5 Grown in restricted
space. Tall form

depressor muscles. The broad tergum has generally two crests for attachment
of its depressor muscles. Although more than 300 specimens have been
examined from areas throughout its geographical range, none has been seen
with three complete crests—two is the normal complement or at the most (and
only occasionally) two crests plus a small rudimentary ‘“ half-crest’’ may be
found. Other species in the stellatus-complex generally have four or more
crests on the tergum or, on rare occasions, three crests plus a rudimentary
‘* half-crest ”’.

Soft Body
The soft body is larger than in other Australian species of Chthamalus.

48 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

Fig. 4. Chthamalus antennatus. (a) Whole shell of a large and moderately eroded individual ;
(6-c) side views in silhouette of (b) * tall ” form resulting from crowding during growth and (c) more
“normal ”’ form; (d) scutum, interior view, showing comparative featurelessness of valve except
tor two incipient crests within the well-marked depression for attachment of the lateral depressor
muscles; (e) tergum, interior view, with two prominent crests for depressor muscles; (f) a
normally shaped mandible, with a short comb-like section composed of coarse spinelets in its
lower section ; (g) regenerating mandible after damage to lower tip (the pair of (f) but drawn to a
slightly larger scale), showing the coarsely pectinate (hembeli-stage) through which the stellatus-type
of mandible passes after damage and repair.
F. J. Beeman del.

ELIZABETH C. POPE 49

Colour

The basis is dark and wholly membranous ; the tergo-scutal flaps are dark
black with a white rim round the opening. There is another light rim round
the membrane at its point of junction with the lower end of the sheath and the
basal margins of the opercular valves. The prosoma is light and greyish but
cirrus I and most of IT and parts of the pedicels of the remaining four cirral
pairs are dark bluish-grey (fading into grey, after long preservation). The
penis is, by contrast, white with a darkened proximal part, where narrow,
raised dark rings occur. There are the usual dark areas round the bases of
the spines on each segment of the rami of cirri ITI-VI. In material freshly
preserved in 70% alcohol, the front of ‘‘ the face’ is a dark blue colour, because
the pigment seems to be concentrated round the bases of the bristles which
arise from the free surface of the trophi. There is a most distinctive, arched,
blue loop above the apex of the notch of the labrum.

Trophi

The labrum is typically bullate with a wide inverted V-shaped groove and
five moderately large, triangular denticles on either side of a central haired
section at the apex of the groove. There are also hairs above the teeth. The
rows of teeth are not continuous in all individuals from side to side, as shown
in the dorsal view by Nilsson-Cantell (1921, Text-fig. 53a). The palps are
almost rectangular in shape except for the median lower border which is slightly
rounded. The upper and median borders and outer tip carry longish hairs—
shorter along the upper border and becoming progressively longer towards the
lower, median edge of the organ. The mandible is variable and appears in
two main shapes. The commoner one has three large main teeth above,
followed by a fourth smaller, generally double tooth (Text-fig. 4f). Below
this is a comb-like section in which, however, the teeth of the ‘‘ comb” are
somewhat coarser than those of C. stellatus as shown in fig. 84 of Pilsbry (1916).
Below this again are generally four long, spine-like teeth, forming the lower
angle of the jaw. This is essentially a stellatus-pattern of jaw. Some large
Specimens, taken in areas subjected to maximum exposure to wave action,
frequently show damaged mandibles in course of regeneration, in which there
is a different pattern in the lower tip of the jaw—this variation was described
by Darwin (1854, p. 460). They have three upper teeth, the fourth (often
incipiently doubled) is almost indistinguishable as shown in Text-fig. 4,g, but
it is definitely present. Below it is a coarsely pectinated section, with other
spines showing at random between and behind the coarser spines—there is no
‘‘ comb-like’”’ section. Sometimes mandibles of the right and left sides may
vary and while the left one may have a stellatus-pattern for its lower tip, the
right may have a “‘ hembeli”’ one. However, in the individuals with somewhat
hembeli-like jaws the small, fourth double tooth can generally be seen, thus
enabling the real affinities of C. antennatus within the genus Chthamalus to be
recognized. The first maxilla has its spines separated into three groups by a
well-marked upper notch and a less obvious lower notch, situated just above
the bunch of small, bristle-like spines forming the lower corner of the organ.
The upper notch is shaped like the letter ‘‘U” on its side, with a rounded
bottom and there are no spines arising from its side-walls or within the notch.
There are two or three very large spines at the upper corner of the jaw and
generally two smaller pairs just above the notch. The central section carries,
between notches, a varying number (7-9) of medium-sized spines while the
nine or so bristle-like spines below the second notch form a brush. The upper
and lower borders of maxilla I carry a few hairs. Masilla JI is stouter basally
than is shown in Nilsson-Cantell’s illustration (1921, Text-fig. 53), the anterior
notch is distinct, rounded and free from bristles, and the spines along the lower
margin are longest.

D

50 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

Cirrt

Cirri I and II are slightly unequal but both are considerably shorter than
the remaining pairs, Cirrus I generally having six segments in both rami in
adults and each carrying a few long, pinnate bristles on its terminal segment.
Cirrus II has generally five segments in the anterior ramus and six in the
posterior one. Its pedicel is produced anteriorly into an oval-shaped outgrowth,
fringed with fine bristles. Both terminal segments of the rami carry very
stout, lanceolate spines furnished with a double row of stout peg-like serrations.
There are four (or five) of these denticulate spines on the anterior ramus and
even more on the posterior one. Their structure is distinctive among the
closely related species in the stellatus-complex and therefore diagnostic (Text-
fig. 1,h). The illustration of them in Broch (1922, fig. 51c) is not sufficiently
detailed to disclose the finer structures which have now been found to be
important. Cirrus III was adequately and ably described by Darwin (1854,
pp. 460-1) who made it very clear that the character of a very long, often tightly
coiled, outer ramus was by no means universally present in all individuals.
However, most subsequent authors, ignoring his warning, have placed undue
emphasis upon it as a diagnostic feature for the species. An investigation
was undertaken to determine the frequency of occurrence of the long outer
ramus of cirrus III and to look for other characters which might be used to
diagnose O. antennatus.

A total of more than 150 individuals in samples of 20 (except for one
locality in Southern Tasmania where only 10 were available), representing
five localities spread throughout the range of C. antennatus on the eastern
Australian coast, was dissected. All degrees of exposure to wave action were
represented in the localities chosen, ranging from sheltered oceanic (as at
Balmoral, Sydney) to very exposed (as at Harbord, also near Sydney). The
only clear result of this investigation was that in no case did 100% of individuals
in any locality possess the long outer ramus on cirrus III whereas in one quarter
of the samples, 100° did not have it, both rami being short and subequal in
length. In several batches only 15°% of the barnacles possessed the long ramus.
No clear-cut correlation could be found between the presence or absence of
the long cirrus and the age-groups of the barnacles or of any specific seasonal
or environmental factors. There was some indication, however, that increased
predation by the molluscs Dicathais orbita and Morula marginalba was taking
place in the late summer when local C. antennatus contained developing nauplii.
In late January, 1957, up to 70% of the population at Pearl Beach in Broken
Bay, N.S.W., had long outer rami on cirrus III and 80% were brooding
developing larvae. Samples taken in the same area, three weeks later, had
none and from 20-35% still contained developing nauplii. The predators
were very active in the area. The evidence is inconclusive and proper experi-
ments would be required to find out whether the long outer rami were lost
naturally after a moult or as a result of misadventure, either during release
of larvae or by the action of predators. The barnacle population contained
a range of sizes of shell, presumably of different ages, and all had lost the long
outer ramus. Here is a nice field investigation waiting for some student’s
attention. Wisely and Blick (1964) have established that the main liberation
of nauplii in C. antennatus occurs in Sydney specimens in late spring and con-
tinues intermittently until May, so that an investigation of predation over
this period might be fruitful.

After injury, regeneration leads to the formation of subequal rami on
cirrus III but what happens at the next moult is unknown. Does the outer
ramus tend towards the antenniform stage once more, and lengthen during
subsequent moults? This is possible because many large-sized individuals
do have long outer cirri, and it is felt unlikely that so fragile an organ could
escape damage throughout all the moults and other vicissitudes in the life of
a species living in such exposed conditions.

ELIZABETH C. POPE 51

Cirri [LV—VI are normally shaped, for the genus, and have subequal rami.
In cirrus IV most of the segments towards the middle of the rami have four
pairs of anterior spines ; cirrus V has some segments with four spines but the
majority generally have three pairs of spines, on both rami; but cirrus VI
almest invariably carries three pairs of spines on each of its segments.

Habitat

Chthamalus antennatus flourishes under conditions of maximum exposure
to high seas and to periods of long exposure to air and sunlight. The level
of shore rocks, where it lives, may be exposed to air for as much as 98% of the
time though in the warmest part of its range (S. Queensland) the author has
noted a tendency to settle in somewhat shaded areas on the rocks. In the
central coastal area of New South Wales, in localities exposed constantly to
rougher seas, it may be attached to rocks up to the level of extreme high water
mark of spring tides or in favourable localities, even in the splash zone above
it, on the side walls of regular drainage channels or areas where spray is con-
stantly blown.

Coastal surveys showed that the vertical range of C. antennatus had its
upper limit considerably lowered in Victoria and Tasmania, where air and sea
temperatures are somewhat lower than they are in the rest of its geographical
range (Bennett and Pope, 1953). In fact, from Cape Otway westwards for
200 miles in Victoria and at Maatsuyker Island, off SW. Tasmania where
prevailing winds, especially in summer, cause exceptionally cool air temperatures
to prevail during the greater part of the year, there are breaks in its horizontal
geographical range where it is virtually, if not quite, absent (Bennett and Pope,
1953, 1960; Womersley and Edmonds, 1958).

Distribution

Australian: This species ranges from Bustard Head (S. of Gladstone) in
Queensland southwards on the fully exposed, oceanic coasts of southern Queens-
land, through New South Wales, Victoria and Tasmania and thence westward
along the coast of South Australia in the Great Australian Bight into Western
Australia, at least as far as Eucla and a little beyond. It was not found farther
to the west, at Cheyne Beach or in the Albany area of Western Australia.

Specimens from Broome, Western Australia, named by Broch (1916) were
examined in Sweden and proved to be juveniles of C. malayensis.

World Occurrence: It is believed that all records of C. antennatus from
areas Other than in southern Australia—e.g. those of Gruvel (various dates
and countries)—are wrongly identified and even the records of an unknown
species of Chthamalus at Whangerei and Poor Knights Islands in New Zealand,
on the east coast of the Northland Peninsula, by Cranwell and Moore (1938)
were later found by Moore (1944) to be juveniles of her new species, Chamae-
sipho brunnea, which goes through a six-valved stage during its development.
There is no reliable record of this species outside Australian localities.

CHTHAMALUS MALAYENSIS Pilsbry, 1916
(Plate ii, figs. 3, 6, 7; Text-figs 1,2; 5,a-g)
See Utinomi (1954) for earlier bibliography and add the following :—

Chthamalus malayensis : Kolosvary, 1941 (part only of the material under
this name is true C. malayensis) ; Utinomi, 1949 (important distribution table),
1954; Endean, Kenny and Stephenson, 1956 ; Endean, Stephenson and Kenny,
1956 ; Stephenson, Endean and Bennett, 1958 ; Stubbings, 1961 ; Karande and
Palekar, 1963, illustrated ; Zevina and Tarasov, 1963, illustrated ; Southward,
1964.

C. antennatus: Gruvel, 1912; Broch, 1916, illustrated.

52 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

C. moro Pilsbry: See synonymy in Utinomi, 1954, but add Broch, 1922,
illustrated ; Utinomi, 1949.

C. challengeri: Nilsson-Cantell, 1921 (part only of C. challengeri), 1938.

C. challengert f. krakatauensis: Broch, 1931, illustrated.

C. stellatus: Darwin, 1854 (part only, including at least specimens from
Philippine Islands); Hoek, 1913, illustrated; Kruger, 1914 (recorded as C.
stellatus var. communis Darw.); Pilsbry, 1916 (? material from Port Cuyo in
the Philippines which should be malayensis?); Nilsson-Cantell, 1921, 1934,
1938 ; Stubbings, 1936; Daniel, 1956.

In the comments made on the genus some of the difficulties experienced
in attaching the correct name to the Australian material belonging to this
species were given and a glance through the list of authors in the above synonymy
and in the synonymy in Utinomi’s paper (1954) will show clearly the confusion
that has arisen over the identity of Chthamalus malayensis in written accounts
of European systematists. Several workers have allotted slightly differing
batches of C. malayensis to 2, 3, or even, in the cases of Broch and Nilsson-
Cantell, to no less than 4 different species. If one plots on a map of the South
China Sea and the Indomalayan island chain, the distributions attributed in
literature to C. malayensis (with its synonym C. moro), and those of C. stellatus
and ©. challengeri, the result is chaotic. It is as though a giant hand had
Shaken out a mixture of species at random like pepper out of a pepper pot,
and they had landed anywhere throughout the area. There is no pattern in
the distributions, and common sense appears to have been forgotten in the
process. Colder water species have been recorded at the equator and tropic
Species in the cool temperate seas.

No doubt this confusion has been in part due to the comparatively late
recognition of C. malayensis (1916) as a separate species, and in part to its
almost protean ability to acquire differing shapes and weathering under slightly
differing ecological conditions, coupled with the fact that many of the earlier
workers looked no deeper than shell structure and a few obvious characters
of the soft parts to determine their species.

The present review has covered the examination of large collections taken
along some 7,000 miles of the Australian coastline—from Great Sandy Island
in southern Queensland, northwards to Cape York and thence to several
scattered areas along the northern coastline (e.g. Groote Hylandt, Darwin and
Yampi Sound) into Western Australia, but from Port Hedland frequent sampling
points were possible, westwards and southwards to Garden Island, off Fremantle
in southern Western Australia. In addition material from New Caledonia,
New Guinea, Indonesia and India has also been dissected. This probably
constitutes the largest sample of OC. malayensis yet worked and after this
experience, and having examined Pilsbry’s types and struggled through the
tangled jungle of literature dealing with the species, it is the author’s considered
advice to future systematists, attempting to revise the genus Chihamalus, not
to waste time in trying to straighten out the errors of earlier writers. Although
this has been done in the present instance, and the source (or sources) of error
in each case listed in the above synonymy has been traced, it is felt that this
review is not the correct place in which to document these data. Rather should
future reviewers obtain reasonably large samples of each species belonging to
the stellatus-complex, from relevant localities and reexamine them for himself
in order to list the characters belonging to each of the species that have been
confused with C. malayensis. Only in this way will a clear picture be obtained
of their true relationships. A special warning is also necessary about several
misleading errors in Pilsbry’s original description of the soft parts of C. malay-
ensis. The shell parts of this species are clearly designated as type material
in the United States National Museum, in Washington, D.C., United States of
America (Registered No. 48084) but are dissociated from any soft body material.
The author was informed that the late H. A. Pilsbry had retained his micro-

ELIZABETH C. POPE 53

slide material and that it might be found in the Department of Mollusks of
the Philadelphia Academy of Natural Sciences, where Pilsbry formerly worked.
With the help of Dr. Tucker Abbott, in Philadelphia, Pilsbry’s microslides
were traced in bis former Department and a set, obviously associated with
work recorded in Bulletin 93 of U.S. National Museum (1916), was located
and examined for possible evidence. These microslides had originally been
mounted in glycerine jelly but were unringed and had deteriorated badly. In
fact it is doubtful if any measure could be found to restore them to a state
in which they could be usefully examined. They were not designated as part
of the type material, however, and, in any case, no trace could be found of any
microslide labelled ‘ C. malayensis”. As C. malayensis can fortunately be
distinguished by its shell parts alone with certainty, if necessary, by an expert
it has been possible to learn that there are serious incongruities if not errors,
in Pilbry’s original description and illustration of the soft animal parts of this
species. These errors have proved to be particularly misleading and are, no
doubt, the fons et origo of all subsequent confusion over this species—a confusion
that has been quite unnecessary since C. malayensis is, as Utinomi (1954) and
other subsequent authors have clearly been able to recognize, a valid species
well defined and demarcated from its most closely related species, C. antennatus,
and should never have been confused with the more distantly related C.
stellatus and C. challengert. Use of Table 2 (above) should help future workers
using collections already named in northern European Museums to rename
the specimens that have been incorrectly determined. In actual practice,
except for fringe areas of zoogeographical overlap, it should be almost possible
to determine closely related species of the stellatus-complex on zoogeographical
areas alone.

The errors specially noted in Pilsbry’s original account occur both in the
written material and in his fig. 90A (1916, pp. 310-312, since C. moro = C.
malayensis). The illustrations both for C. malayensis and C. moro on his Plate
72 are, however, correct. The following characters of C. malayensis, as described
by Pilsbry, are not correct :—(a) Cirrus II which, contrary to his statement,
does possess stout spines which are large-toothed and have a characteristic
Shape, whereas any of the C. challengeri, dissected by the author, do not have
similar spines (Pusbry’s 1916 drawing of C. challengeri, fig. 87C, is therefore
incorrect and much more like C. malayensis) ; (b) In the mandible the lowest
tip of the adult jaw generally has only two slightly larger teeth, not three as
Stated ; (c) the maxilla is not as illustrated (Pilsbry, 1916, fig. 90A) but has
a Clearly defined, upper U-shaped notch devoid of spinelets, above which there
are two very stout, large spines (nec 3 as he states) with three slightly smaller
ones below them followed by the clear notch; (d) Cirrus VI may have four
pairs of spines on each segment in one ramus and/or three or four on the other.

Under these circumstances it would seem best to redescribe C. malayensis
im some detail and to refer to more recent and reliable descriptions and
illustrations. However, before embarking on this it should be stated that the
author has compared the types of Pilsbry’s two species C. malayensis and C.
moro (held in the collection of the U.S. National Museum, Washington, D.C.,
in the U.S.A.) and there is no doubt at all that Utinomi’s (1954) action in synonym-
izing these two species is correct. In the Australian collections of C. malayensis,
the smooth-shelled, generally quickly-grown, taller shell of the ‘‘ moro” kind
occurs side by side with those with ribbed shells and smooth walls and the
flatter shape which is associated with the name C. challengeri Hoek forma
krakatauensis Broech (first used by Broch (1931) for specimens from Krakatau
Island). Broch’s illustrations show that his specimens from Krakatau Island
were indeed C. malayensis, and this was confirmed when Broch’s “ Type sett
and lectotypes”’ were examined in the Zoological Museum of the University
In Copenhagen, Denmark; Utinomi’s decision (1954, p. 18) in synonymizing
malayensis and C. challengeri f. krakatawensis Hoek is thus also confirmed.

54 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

Appearance and Shell Structure

Many Australian specimens of C. malayensis on the rocky substrate (Plate
li, fig. 6) have the shape of the specimens described by Broch (1931, as a form
of C. challengeri) and observation has shown that they are comparatively young
or half-grown specimens, even though they may be breeding. Further growth
and erosion lead to a loss of the distinct ribs with projecting processes round
the circumference of the shell and only traces of the ribs remain, near the
circumference of the shell as seen in Plate i, fig. 7 (the specimens not ringed
in the photograph) which shows fully grown somewhat eroded specimens from
Western Australia. The largest barnacle in this illustration is a specimen of
C. antennatus, from eastern Australia, deliberately placed on the timber among
the C. malayensis to emphasize the differences in shell ribbing, size, and general
facies, of the two species at a similar stage of their growth—differences which
are obvious to the eye, but hard to record in written descriptions. As juveniles,
while their shells are still thin and fragile, C. malayensis and C. antennatus are
similar externally and this accounts for Gruvel’s (1912) and Broch’s (1916)
mis-identifications of material from, respectively, Amanu Island, Tuamotu
Group and from Broome in north Western Australia.

It is interesting to follow the progress of Hiro—Utinomi’s ability to recognize
C. malayensis in its many forms, as his ecological experiences of it increased
over the years and the size of the population sample he handled grew greater,
for my own experience has paralleled his, in almost every respect. One sees
how, by placing too much reliance on the accounts and illustrations of earlier
writers, one could believe that anything from three to five distinct species
were present, but later found that they belonged to one highly variable species,
able to grow attached to rocks, or to mangroves (in areas where CO. withersi
was not present, to compete for lebensraum), or to molluses and other barnacles
and how in each of these situations the shape of the shell differs greatly in
appearance. Three of the more common variants of shell sculpturing are shown
in Text-fig. 5.a—d, but there are still other frequently recurring patterns of shell
erosion not depicted. The final stage is reached when senescent barnacles,
with carino-rostral diameters of 14 mm. or more, show a somewhat rectangular
Shape with only faint traces of ribs, often marked merely by a linear series of
black pits with dark corium showing through and with raised shell bumps round
them, the last remnants of the original ribs. The opercular valves are so worn
as almost to be characterless externally except for the deeply sinuous sutures
marking the interlocking of the scuta and terga. This last shell type is par-
ticularly common on mangroves or on the rocky shore between Carnarvon and
Broome, in the northern part of Western Australia, whereas in Queensland
C. malayensis rarely attains such dimensions and the shell form that is most
favoured in this environment (chiefly on intertidal rocks) is the more stellate,
smoother form shown in Plate u, figure 6, or in Text-fig. 5,b, which shows a
half-shell in which the ribs appear only near the circumference.

However, in spite of these varying shell shapes, internal structures are
constant and a detailed description of these will be given. Special stress will
be laid on diagnostic characters, for there is no complete recent description,
Since Utinomi’s important paper (1954, pp. 18-21), which did much to clarify
the nomenclatural muddle but did not redescribe the species. Plate ui, figs
3, 6 and 7 and Text-fig. 5 have been chosen to illustrate the appearance of
Australian specimens. The only widely differing form, not shown in the present
work, is the smoother rather juvenile-shaped kind usually found in localities
with less water movement—the prototype of Pilsbry’s original C. moro and
now recognized as a synonym of C. malayensis. It is illustrated (as C.
moro) by Hiro (1937, fig. 4,a-c). As may be seen in his figure, it has the rather
featureless opercular valves, typical of juvenile Chthamalus. In particular
only three of the usual four crests for attachment of the tergal depressor muscles
have been developed on the tergum. Examination of a series of barnacles

ELIZABETH C. POPE 55

wo |

Fig. 5. Chthamalus malayensis. (a-d) varied shapes, and sculpturimg of shell plates. (a) half
shell of a young and uneroded mdividual growing on rock (typical of the ribbed form designated
by Broch as C. challengeri forma krakataueusis); (b) half-shell of a slightly older and more
weathered individual, but far from mature, showing no pitting of the shell externally (typical
of majority of specimens occurring on rock in Queensland intertidal zone); (c) side view (silhouette)
of a typical C. malayensis ; (d) fully mature, much eroded specimen grown on a mangrove from
the north of Western Australia, much pitting showing externally ; (e) labrum of typically swollen
or bullate pattern, with left palp removed to show teeth and hairs along the groove above the
mouth ; (f) scutum, interior view showing rugged sculpturimg, pitting (as opposed to smoother
interior of C. antennatus) of general surface and the deep clearly defined pit for the imsertion of
the lateral depressor muscle ; (g) tergum, interior view, showing four crests for depressor muscles
attachment, and the prominent central row of pits about which the valve is bent at an obtuse angle.

F. J. Beeman del.

56 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAH

of increasing size would, however, have shown that, as they grow, first three,
then three plus a sort of ‘‘ half crest ’? would be found, then four, and sometimes.
even five full crests might occasionally be seen. However, four crests is the
more usual number in mature specimens. It should be remembered that this
species can also breed long before it reaches full size, so that the presence of
developing larvae in the mantle cavity does not necessarily mean that one is
dealing with a full-grown C. malayensis.

However, it is interesting to note that the characteristically deep pit on
the scutum for the attachment of its lateral depressor muscles is already well
in evidence in the “moro” form and at an early growth stage, and the small
ledge, below the attachment of the adductor muscle of the scutum, is almost.
non-existent. Correspondingly-sized and -aged C. challengeri are generally
already showing a well-raised ridge, below the pit for the adductor muscle,
in any material that has been worked by the author. This included “ co-type ”’
material (juveniles) kindly made available by Dr. J. P. Harding of the British
Museum (Natural History) of London.

The effect of special factors in the tropical environment on the shells of
intertidal barnacles would seem to be profound but, in the absence of experimental
work, we cannot even guess whether it is just the extraordinarily hot temperatures
and consequent high rate of evaporation, or whether some unknown factor
like more intense ultra-violet radiation or the effects of heavy monsoonal rains,
and so on, are operating. Whatever the controlling factor or factors may be,
it seems to be connected, if not primarily at least secondarily, with the intensity
of the sunlight, for the appearance of two batches of C. malayensis settled.
simultaneously in the same area (the one in shade and the other in full sunlight)
are often so different as to look like two different species i.e. the ones grown.
in shade resemble the moro kind of shell. Another special feature of tropical
shores is the presence, at least along the northern coast of Western Australia,
of mangrove trees in the higher intertidal zone, which are not necessarily
associated all year round with the soft mud of coastal rivers and much turbidity
in the sea. It is along this part of the Australian coastline that C. malayensis
has, as it were, taken to the trees and developed the larger rectangular-shelled
individuals with carino-rostral diameters of 15-0 mm. or more. Only traces
of the numerous narrow original ribs remain near the outer margin, and their
shells are deeply pitted internally. Often the pits open externally on the sbell
plates, at the end of one of the bosses marking the last remnant of the ribs
(Text-fig. 5d). The sutures between the shell plates are simple and untoothed
but are more firmly articulated together than they are in OC. witherst.

Uneroded C. malayensis are generally ash grey and have 4—6 ribs per plate
—narrower than the ribs or folds in correspondingly aged C. antennatus or C.
challengeri. There may be a tendency for the ribs of C. malayensis to bifurcate
towards the circumference of the shell so that the width of the ribs remains
constant, but their number per shell plate increases gradually as they age. The
basis of the shell is wholly membranous. Internally the colour of the shell
is a dark grey and it is generally considerably pitted.

Measurements of 10 individuals collected at Onslow in Western Australia
are given in Table 7. This sample showed considerable size variation and
differing shell weathering as set out in the last column. They were collected.
by Mrs. L. Marsh, on 29/9/1959 and none contained developing larvae. <A batch
taken towards the end of December 1964, however, from Cockatoo Island,
Yampi Sound in Western Australia (collected by N. Hoffman) had their mantle
cavities filled with well-developed, eyed-nauplii, almost ready for release.

These measurements of a truly tropical sample of C. malayensis indicate
its shape and generally low-growing habit. Only rarely will intense crowding
cause it to assume a more tubulo-conical form. In localities favourable to
O. malayensis, i.e. where there is some degree of water movement and slightly

ELIZABETH C. POPE Bye

less turbidity than usually occurs in many tropical seas, it may colonize the
rocks just as thickly as Chamaesipho columna does in the temperate zone, and
May appear as a broad whitish band along the rocky shore, just above or mingled
with the band of the tropical oyster, Crassostrea amasa (Iredale). Such an area
is illustrated in Plate ii, fig. 3 and was on an offshore rock stack at Yorkey’s
Knob, near Cairns in Queensland.

Opercular Valves

Reference has been made above to the variability in form and structure
of the opercular valves in shell with differing external appearance, but no
reference has been made to their more constant characters which can be used
to diagnose C. malayensis. Variability of form in these plates is well shown
in Utinomi’s drawings (1954, fig. 2,a-h) but constant characters can be recognized
in them too. Again it is best to ignore previous descriptions except Utinomi’s.

TABLE 7
Measurements of 10 C. malayensis from Onslow, Western Australia, 29.1x.1959

Carimo-rostral Width in Height in
diameter mm. mm. (at Remarks
in mm. highest point)
7-2 8-2* 3° 6* Uneroded—pale grey, ribbed
Tergo-scutal flaps jet black
8-5 8-5 2-5 Small, uneroded, ribbed form
Tots 11-0 5: 0* Very eroded
12-5 9-6 4-5 Shaped like C. antennatus due to
crowding. Very eroded
13-9 Tilofg) 6-5* Very eroded
14-5 12-0 4-5 Very eroded
14-6 13-5 4-3 Rectangular im shape and very
eroded
15-0 13-4 4-3 Very eroded
16-2 Il 5-0 Rectangular in shape, growth
crowded. Very worn. Rows of
black bosses, along former rib-
line
17-0 7 6-4 Basis dark (black) wholly mem-

branous and undamaged

* Distorted due to crowding.

The scutum always has a deep pit near its basi-tergal corner, directed upwardly
towards the apex, and this forms the point of attachment for the lateral depressor
muscle. It is deeper and more pronounced than the pits in related species
like C. antennatus and C. challengert and there is no trace within it of small
crests for attachment of the muscles, as seen in C. challengert. In C. antennatus
the corresponding lateral depressor pit is only moderately developed and traces
of raised crests may be seen within the pit, making this species intermediate
in structure in this respect between challengert and malayensis.

The scutum of C. malayensis, while lacking the very prominent and well-
marked ridge, seen in C. challengeri, below the scar of attachment for the lateral
depressor muscle, nearly always has a small ledge, especially in well-grown
Shells, below and towards its tergal side. The differences in this structure,
and the opercular valves in general, may be appreciated by comparing two sets
of illustrations by Fujio Hiro of C. challengeri in his 1932 paper (Text-fig. 1,a—d)
and C. malayensis in his 1939 work (Fig. 1,b-c). To these need only be added
the statement that Australian specimens of malayensis behave similarly to
the populations described by Utinomi (1954, pp. 18-21), remembering while
reading Utinomi’s account, that Nilsson-Cantell was confused over the system-
atics of this species and that Pilsbry had failed to note the coarsely built,

58 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

pectinated spines associated with cirrus II in C. malayensis, so that Utinomi’s
commept and comparisons with the descriptions of these authors should be
disregarded.

The tergum of C. malayensis (Text-fig. 5,9) in older individuals is generally
broad towards its apex but narrower towards its basi-scutal corner than in
the corresponding valves of C. antennatus. It is folded about an axis running
from its apex to the basi-scutal tip and this fold is often emphasized internally
by a line of very distinct pits which sometimes merge to form a narrow furrow.
The inner surface is much pitted in older barnacles and there are generally
four more or less parallel crests for attachment of the tergal depressor muscles
(occasionally 3$ in juveniles or 44 to 5 crests in very old individuals). This
contrasts sharply with the most closely related species in Australia, C. antennatus,
in which tbere are only two crests for tergal depressors, or at the most two
plus a partially developed third crest, in very old individuals.

Soft Body

Colour :—In newly preserved material (in 70° ale.) the prosoma is a light
creamy colour and the cirri are correspondingly light but with more of a pale
grey tone. Cirri I and II have rami a dark, somewhat purplish-grey, and
pedicels a lighter shade of the same general tone. Other cirri have darkened
patches on the inner sides of their pedicels and concentrations of dark pigment
round the bases of the large anterior paired spines on each segment of the rami.
The proximal part of the penis is also dark, the rest of it being startlingly white,
by contrast. It is very long, and tapers gradually to a fine tip. Developing
eyed-naupli were pinkish-white. There is no dark ring of pigment round the
mouth, aS in some other chthamalids ; instead, only the blades of the palps
and the outer tips of the mandibles and of maxilla II were slightly darker than
the rest of the head region. There is no caudal appendage.

Trophi

The labrum is of the “* normal ”’ bullate shape, i.e. bulging smoothly, above
the area where the spatulate tips of the palps le recessed in grooves and
closely adpressed to the labrum. Their lower borders are almost parallel to
those of the notch of the labrum which is broadly ‘“‘ A ”-shaped. ‘There is a
sight thickening of the walls of the V and a flattening at the peak of the
groove, the side walls of which carry a row of broadly triangular teeth with
a row of close-growing hairs above them. Frequently food particles and debris
are entangled between the teeth and the hairs and they have to be brushed
away before the structure of the notch can be seen. The lower borders of the
palps carry a fringe of long bristles which depend over the mouth opening.
These are longest medially. The mandible is essentially of the four-tootned,
stellatus pattern, in which the 4th tooth is smaller and is frequently doubled.
Below the 4th teeth, however vestigial they may be, there is generally an
appreciable change in the style of the dentition or spinulation except in the
case of damaged end regenerating jaws which will be described below. From
this point downward to the two large spike-shaped spines arming the lowest
tip of the jaw, the spines are generally placed parallel to one another like the
teeth in a comb and are of smaller size ; however, they are not fine and hair-
like as in C. stellatus and C. challengeri. Great variation bas, however, been
seen in this lower part to the jaw and even, as between right and left sides of
the one individual and this can usually manifestly be traced to regeneration
on the part of one mandible. Often the nature of the injury can still be seen.
As juveniles, or during regeneration or in certain individuals, the lower tip
of the mandible is reminiscent of the hembeli jaw pattern, i.e. there is no comb-
like section below the 4th tooth or, alternatively, it is so drastically reduced
as to be virtually absent, being replaced by a few moderately-sized spines
arranged as in the coarsely pectinated part of C. hembeli. This condition is

ELIZABETH C. POPE 59

shown (for C. moro) by Broch (1922, fig. 52a). Only with maturity and in the
absence of damage and regeneration does the ‘‘ normal” jaw shape develop.
It is as though juveniles and regenerating C. malayensis (like C. antennatus)
have to pass through a hembeli-stage during the development of their much
toothed and highly complex mandibles. Mazilla I is also somewhat variable
as regards the finer details of shape and the exact arrangement of the spines.
In addition the angle from which it is viewed can make a difference to the
appearance of the largest spines above the notch (from one angle there appear
to be three very large spines, whereas from another angle only two of the largest
Size may be seen). As a rule there are three of slightly smaller size below and
partly between them. Hiro’s illustration (1939, p. 251, fig. 1E) gives an
excellent picture of the shape of the mandible in the majority of specimens
examined from Australian localities. An interesting feature is the presence
of a well-marked upper notch, shaped like the letter U, lying on its side. The
walls of this notch are devoid of spines and only the tips of a fine hair or two
(from the side wall of the mandible) may project behind or in front of it. Such
hairs have no basal connection with the notch itself. This contrasts with the
structure of the maxilla of C. stellatus and C. challenger in which the notch
is much more obscure, more V-shaped and has several spines with their bases
originating along the upper borders of the notch indentation. They may even
obscure any view of the notch itself. The differences between C. malayensis
and for example C. stellatus (s. str.) may be appreciated by comparing Hiro’s
figure of malayensis, referred to above, with his own later illustration of C.
stellatus (Utinomi, 1959, fig. 5b) made after an intensive review of Mediterranean
specimens in which he established the nature of the differences between C.
stellatus (s.s.) and the rediscovered species C. depressus (Poli).

It is virtually useless to refer to published figures and descriptions of the
mandible of C. malayensis in earlier works than that of Hiro (1939) unless one
knows the details of the nomenclatural confusion in each of the papers concerned.
It is believed, for instance, that Pilbry’s original description (fig. 90a) is an
illustration of the mandible of C. challengerc or C. stellatus. It is certainly
unlike that of a true malayensis and, aS has already been outlined above, his
account of the soft parts in this species cannot be checked against microslide
mounts since this species alone seems to be missing from the batch used in
the preparation of his great 1916 monograph. Mazilla I] is of the usual bi-lobed
pattern with a wide, rounded notch free from bristles. Above this notch is
a row of stiffish, larger bristles (more spine- than hair-like) arranged lke the
teeth of a comb and sticking out more or less at right angles from the wall.
Below the notch the bristles are thickly clumped and hair-like, being shorter
near the notch and incieasing gradually in length round the lower tip of
maxilla II. Long hairs occur not only on the lower border but up the outer
Sides of each of these organs towards the rear, giving a bearded and be-whiskered
effect like a man with both a “ goatee-beard and mutton-chop whiskers ”’.
Seen from in front, the ‘ face” is very hirsute. From above, each of the second
maxillae is triangular in outline, with the base broad proximally. The two
maxillae II seem to be more closely adpressed to one another than is usually
the case in the genus and together they form a most effective lower lip, broader
than the average. The mandibles and maxillae I, on tne otber hand, are
comparatively smaller than those seen in the 3-toothed species of the genus.

Cirrr

The first two pairs of cirri are much shorter than succeeding ones and
exceedingly setose. They are approximately equal in length and in each the
anterior ramus is slightly stouter and a little longer than its posterior fellow.
Typically cirrus I has the number of segments as follows :—anterior ramus
eight and the posterior one seven. The numbers of segments may, however,
vary slightly in the population, but is of that order. Both rami of cirrus I

60 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

are heavily setose and the bunches of bristles, on segments, towards the centre
of the rami, form a felted mass of pinnate-type setae. These apparently act.
together to strain food particles from the water. They had frequertly to be
brushed clear of detritus and small plankton in order to see their structure.
There are also present, jointed, larger pinnate setae on the terminal segments
of both rami of cirrus I. Towards the posterior, inner margin of the anterior
ramus, six or so of the basal segments carry numerous peg-like, short stout
spines, the apparent function of which is to project into the fringe of setae
round the opposing posterior ramus, thereby locking the two together and
forming a broader and more rigid scoop-like structure of the remi which act
together and increase its efficiency as a feeding organ. Even the segments
of the pedicels of C. malayensis are more heavily fringed than those of C.
antennatus and some of these setae are of the pinnate type. The posterior
ramus of cirrus I lacks the stout peg-like spines posteriorly. In Cirrus II the
segmental counts are respectively eight and six (or of that order) for anterior
and posterior rami. The stouter anterior ramus carries setae and spines of
four main types, namely (1) stout, lanceolate, toothed spines; (2) pinnate,
hair-like bristles ; (3) ordinary straight spines; and (4) posteriorly placed peg-
like spines on several basal segments. Of these, the lanceolate spines merit
special attention, since their structure can be used in diagnosing the species.
Each of these lanceolate spines is armed by a double row of coarse, pointed
spikes, the lowest of which are separated from those above by a diastema-like
gap. In some species there is a trace of a joint in the spine at this point, and
the lower teeth-like projections are associated with the top of the “ basal ”’
section aS opposed to the distal or ‘ blade” part of the lanceolate spine. In
large individuals with very eroded shells (of the type depicted in Text-fig. 5,d)
the lowest spines may be reduced (by wear or erosion) to mere bumps, whereas
in younger specimens they are generally spine-like. The double row of serrated
spines above the gap may also be worn but are always coarser in build than
corresponding structures in either C. stellatus or C. challengeri. They are
generally, however, more pointed and not quite so coarse and peg-like as those
of C. antennatus. This last species lacks the two lower spines below the diastema
gap. It is hard to reconcile Pilsbry’s statement about cirrus II in the original
description (1916, p. 311) with what occurs in the material examined. His
statement, ‘‘ The spines of the terminal segments are as described and figured
for C. stellatus”’, is unfortunate when there is so marked a difference in the
coarseness of the serrations in the two species. If he merely meant that serrated
spines were present on cirrus II, as opposed to the absence of such spines (as
in the species C. withersi and C. caudatus), his remarks are understandable
but it is felt that much of the confusion in identifications of Chthamalus barnacles
from the Indomalayan Peninsula stems from this statement and one can fully
understand Nilsson-Cantell’s feeling (1921, p. 276) that he could not really
distinguish Pilsbry’s C. malayensis from other closely related species. In fact:
Nilsson-Cantell (1921) omitted it from his key to the species of Chthamalus
and from this time onwards appears to have been unable to recognize the
species correctly, referring it sometimes to C. challengeri but mostly to C.
stellatus. Moreover, his redescription of C. challengert in his 1921 monograph
(pp. 279-281) obviously includes both C. malayensis and C. challengeri, and
still further adds to the confusion. He specifically mentions the differences
in the structures of the lanceolate spines of cirrus II in specimens from Java.
[= OC. malayensis] in which the serrated side teeth are ‘ kraftig ’’ whereas in
Japanese [= C. challengeri] specimens they are ‘‘ schwacher”’ (see Table 2).
The number of stout spines in each terminal segment of the rami of cirrus I
varies slightly from barnacle to barnacle, but they are of the order of from six
to nine. They also occur in segments below the terminal ones, in varying
numbers, among the bunches of pinnate spines, but one has to explore several
planes of focus, under high magnification, before all may be viewed and counted..

ELIZABETH C. POPE 61

When a large sample of C. malayensis in varying age groups and from
varying localities is dissected, stout spines may be found varying in coarseness
and structure from a coarse pinnate-type, through a number of stages, where
the ‘‘ blade’’ part distal to the joint of the spines appears to have shortened
and thickened while its side hairs have correspondingly thickened and become
blunter, till they become tooth-like. The basic structural relationship between
the stout, lanceolate spines and the longer more delicate spines thus becomes
obvious, especially so where damage to the tip of cirrus II has led to regenera-
tion of the organ. It apparently takes some time and perhaps several moults
before the stout spines return to their normal, pre-damage shape. This again
may lead to confusion in specific determination unless one is experienced in
the group and underlines the necessity for examination of larger series of
specimens when determining species.

Considerable magnification and controlled lighting may be necessary to
disclose the presence of pinnate setae, but they are generally long and fine,
and occur in bunches, anteriorly on the segments, tending to form matted
elumps. They occur thus, from near the basal segment of each ramus to the
segment just below the tip, and are more thickly clustered on the anterior
than on the posterior ramus. Short stout, curved spines are again found along
the posterior border of the anterior ramus, opposite to the nearest border of
the posterior ramus, where they may be seen to interlock into spaces between
the bases of its setae. However, they occur only in pairs—generally one pair
per segment (not scattered in a random clump as they did on cirrus I) on the
lowest three or four segments. The lowest segment may sometimes carry
two pairs of spines. While not quite as setose as cirrus I, the second cirrus
of CO. malayensis is nevertheless a very efficient food-sieving organ. Cirri
III-VI are similar in structure to one another and of the ordinary pattern for
posterior cirri, sub-equal in the length and in the numbers of segments of the
anterior and posterior rami. Each segment, apart from a few basal ones has
four pairs of anterior spines of which the longest is the upper one and the
Shortest lowest. The lowest pair have their bases placed close together and
sometimes are both broken off so that the segment may appear to have
three spine pairs. However, the site of the missing fourth pair can generally
be seen. The penis is longer than cirrus VI in most specimens and is closely
annulated. There is no caudal appendage.

Habitat and certain aspects of ecology

When not restricted, by competition for settlement space (with C. withersi
from above and with the oyster Crassostrea amasa, below), Chthamalus malay-
ensis occupies a wide band on the open rock faces in Queensland and may be
found throughout the upper half of the tidal range of spring tides and even
into the splash zone in areas noted for constant choppy or rough seas, though
this is rather exceptional in Queensland. Populations of C. malayensis are
generally most dense between mean high water and the low water level of neap
tides. The occurrence of this species on the mainland coast of Queensland
and the effects of environmental factors on its distribution have been discussed
by Endean, Kenny and Stephenson (1956). It has also been the subject of
a more detailed autecological study by post-graduate student, Miss Judy
Bryan (University College of Townsville, Queensland), and the author has
read drafts of her thesis. The results of her work remain, as yet, unpublished
and are therefore unavailable. However, with due acknowledgement to Miss
Bryan a few of the most relevant facts are quoted here where they supply
actual figures for tidal heights, ranges and percentages of exposure for the
approximations noted in the present author’s field observations: Within a
spring tidal range of 12 feet 10 inches, C. malayensis may occur throughout
the upper half of the range and even beyond it into the splash zone, if such
a zone exists. It is densest between the levels 8-11 feet above zero tide mark

62 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

and the largest individuals occur most frequently towards the upper part of
its range. Populations tend to be most dense on near vertical rock faces (in
Queensland). Plate ii, fig. 3 shows an area where the numbers of C. malayensis
per unit area rival the most dense barnacle populations seen on temperate
Australian shores. The width of rock in the photograph was approximately
35 em. Such dense populations can, however, only be found in tropical
Australia, where the right substrate occurs within the barnacle’s tidal range
and where conditions of turbidity and salinity are suitable. One feels that
the ‘‘ paucity ’ of populations of barnacles in the tropics, often mentioned by
other authors, is attributable, more often than not, to the lack of suitable
substrates lying within the tidal range of the species which form dense
aggregations such as CO. malayensis and CO. withersi, rather than to a lack of
barnacle larvae which seek to populate the shores. Miss Bryan noted that
when the vertical range of C. malayensis was related to the graph of theoretical
exposure to air, some individuals might be exposed for up to 90% of the time
and the minimum exposure was of the order of 30%. Air temperatures in
Queensland can range to 35° C and even more, for short periods. Breeding
was most actively carried out in the warmer part of the year, larval settlement
occurring mostly from November, through the eight succeeding months to
June, with the period of densest larval settlement in November through to
February. Although Miss Bryan’s work covers the effects of predators and
experimental investigations of physical factors in the environment, it would
be premature to disclose her findings here.

According to field notes and observations supplied to the author by Mrs.
Loisette Marsh with her collections from the northern part of Western Australia,
C. malayensis behaves somewhat differently there from what it does in Queens-
land, for it occurs not only in the usual manner on rocks (between the 14 and
20 feet levels where the tidal range is 33 feet) but also invades the mangroves
and settles on their trunks and prop roots and on wharf piles. This habitat
is occupied in Queensland, as a rule, by the species C. withers which can tolerate
the turbidity and lowered salinities normally associated with the mangroves
there. However, C. witherst has not been recorded westward of Darwin
(Northern Territory). Some of these northern C. malayensis, taken from the
mangroves, attain relatively enormous sizes (see Table 7) but it has not been
possible, from the notes supplied by collectors and in the complete absence
of chemical and physical data of the environment from this area, even to hazard
@ guess as to why withersi 1s absent and its place on mangroves has been taken
by malayensis. It is hoped to investigate the area personally later, meanwhile
only the facts, as known, can be set down. The shells of many of the Western
Australian C. malayensis differ in appearance from those common in Queens-
land, as may be appreciated by comparing Text-fig. 5,b,d (d being a well-grown
individual from Onslow, W.A.). Itis also a matter of interest that at the extreme
southern end of its range in Western Australia (Garden Island, off Fremantle),
C. malayensis reassumes the shell shape and appearance seen in individuals
taken in central Queensland, i.e. it is smoother shelled and regularly ribbed,
as Shown in Plate ii, fig. 7. In C. malayensis, more than in any other species,
one can appreciate the profound changes caused in the cirripede shell by
varying factors of the environment. In the relatively high humidities that
obtain, desiccation seems to pose less of a threat to life than perhaps some
other effects of the environment, such as sunlight or the opportunity to catch
food. The higher a barnacle grows on the shore in the tropics, the less it seems
to be troubled by predacious molluscs and its distribution seems to be controlled
mainly by physico-chemical factors.

Distribution
Australian: This species is the most widely distributed Chthamalus on
mainland Australia, for it ranges from Hervey Bay (approximately 25° 30’ S).

ELIZABETH C. POPE 63

in 8.E. Queensland, northwards to Torres Strait and thence westward along
the northern tropical coast right round to Shark Bay in Western Australia.
From there southwards, sporadic records occur down to the vicinity of Garden
Island (off Fremantle) but it is rare in this last locality. Chthamalus malayensis
also occurs in Papua, New Guinea (taken by Judy Bryan), but little systematic
collecting for barnacles has been carried out round this vast island, and absence
of records of it there from areas other than Papua, is not significant, for the
present.

World Occurrence: Owing to the tremendous confusion of O. malayensis,
by a number of previous authors, with C. stellatus, C. challengeri and C. anten-
natus no detailed listing of localties will be given from earlier literature. It
will be necessary to resurvey all previous collections to be certain of their
identifications. This applies to most specimens in Huropean and some American
Museums. However, in principle, it may be taken that C. malayensis ranges
throughout the Indomalayan region as follows: From the Persian Gulf
(Stubbings, 1961) along the coasts of India and Pakistan, Malaya, and the
South China Sea (Zevina and Tarasov, 1963) to Formosa (Hiro, 1939). Hiro
points out that its ecological equivalent in Japan’s more temperate seas is
C. challengert, whereas in temperate Australia it is replaced by C. antennatus
and Chamaesipho columna in the south-east. It also ranges widely in Indonesia,
the Philippines, Palao Islands, and several Islands in the Arafura Sea. To
these literature records the following new ones should be added: Kambang
Island, near Timor (taken by the Dutch Snellius Expedition of 1929 on 26—28th
November, on mangroves) ; New Caledonia in the following areas where it was
collected by the author: Reef north of Heinghéne (E. Coast) 13/7/1960, on
intertidal rocks and on rocks and mangroves at Baie des Citrons and at Ricaudy
in July 1960 (both in the vicinity of Noumea) ; at Carlisle Bay, Santa Cruz Island,
Santa Cruz Group (29/7/1926, collected E. Troughton and A. Livingstone),
and Suva, Fiji, on mangroves in Kumbuna Creek (Station 24, Te Vega Expedi-
tion, 26/8/1963, collected by Isobel Bennett). It thus ranges widely in tropical
waters.

Genus CHAMAESIPHO Darwin, 1854
Chamaesipho columna (Spengler) Darwin, 1854, Type species of genus ;
Gruvel, 1905; Moore, 1944. .
C. seutelliformis Zevina and Tarasov, 1963.
Lepas columna Spengler, 1790.

Type species for Darwin’s genus Chamaesipho was Lepas columna Spengler
(1790). Allegedly it came from a Tahitian locality (Otaheite). With it, in
genus Chamaesipho, Darwin associated a new species, C. scutelliformis, which
he believed was ‘ probably from the seas of China’. Zevina and Tarasov
(1963) recently confirmed the occurrence of scutelliformis in the South China
Sea and Gruvel (1905) extended its range to the Indian Ocean. P. H. Fischer
(1884) recorded it in the New Caledonian Archipelago where, however, recent
collecting by the author has failed to find it.

In his 1854 monograph Darwin assigned Australian and New Zealand
Specimens of Chamaesipho to Spengler’s species C. columna, but did so with
considerable reluctance, since the original description was very incomplete.
This matter will, however, be discussed fully when the species is described
below.

Only one species has been added since the two listed by Darwin, namely
C. brunnea from New Zealand, described by Lucy Moore in 1944, the type
locality being Lyall Bay, near Wellington. Previous New Zealand authors
had confused O. brunnea with, and included it in their accounts of, C. columna.
Miss Moore’s paper clears up the earlier confusion between these two species
and defines their geographical ranges in New Zealand. She has shown that
C. columna occurs throughout New Zealand, whereas C. brunnea is limited to

64 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAEK

warmer waters of the North Island and the north-eastern segment of the
South Island. The present survey shows that its range does not extend to
Australia.

Although there are only two species in Australia and New Zealand, the
genus Chamaesipho figures most prominently in the intertidal zone, because
of its dense populations (e.g. in the case of C. columna up to 3,000 per square
foot of rock) in New South Wales, frequently forming a light-coloured frieze
along the rocks, as seen in Plate 7, fig. 1 of Dakin, Bennett and Pope (1948).

The Australian species of Chamaesipho is adapted to withstand a fair
degree of desiccation since it lives above mean sea-level where it is often sub-
jected to long periods of hot sunshine and drying winds. The fusing of the
four shell plates into a tubular shell wall may possibly be a survival factor by
cutting down evaporation. Only the one species, C. columna, occurs in Australia.

Key to the species of Chamaesipho
1. (2). Four-valved (at least in adult stage), depressed Chthamalids of small size (five mm.
carino-rostral diameter) with the rostrum much smaller than the other three shell plates.
The upper part of the sutures adjacent to the rostrum persist when their lower sections
grow together and become obliterated. Compartments other than the rostrum pierced
by a series of four oval apertures—one im each lateral and two m the carma. These extend
as Shelly tubes to the base of the barnacle. Tergum with a wide, blunt, centrally placed
spur and without pits between the attachment crests for the depressor muscles. Orifice
relatively mgs ually: eee een eee teens sence ahem ae cee eee ee C. scutelliformis Darwin
2. (1). Four-valved Chthamalids when adult, though juveniles commence with six, with large
rostral valves and no orifices in any valves. All traces of sutures between the parietes
generally obliterated at an early stage of growth. Shell often tubular and tall, with the
orifice proportionately larger than in C. scutelliformis, and often only slightly smaller than
the basis. Tergum with little trace of a spur and with pits between the ridges for the
abuachimentiormoheldepressormauscles! seri eect eer ei ae ae ro eeeeeeer (3 or 4)
3. (4). Carmo-rostral diameter of adult shell generally no more than six mm., and im tall,
tubular specimens the height may be more than three times the diameter. Soft body
with navy blue coloration, even after preservation. Tergum with four to six deep pits
between the crests for the depressor muscles. There is a small but distimet adductor ridge
on the scutum. Mandible with four to five teeth of which third and fourth may be doubled.
Grapple-like spies on the anterior ramus of cirrus II with only one or two (and occasionally
three) pairs of side “teeth” or hooks (see Text-fig. 1g) ........ C. columna (Spengler)
4. (3). Shell of adult generally larger than in foregoing species. Carino-rostral diameter up
to 24 mm. and height up to 19 mm. The greater width of the shell proportionately to
the height, and the thicker parietes make it easy to distinguish from C. columna. Soft
body light and dark brown in colour. Scutum has almost no trace of an adductor ridge.
The tergum has no adductor ridge and no spur. Crests for the depressor muscles about
seven in number with pits between them. Cirrus II with grapple-like spines with three
OP IMAGINE) (OAHUES OL ln@olieeel HCW WEGWA aaooddacosossddavcpoodses5ooboobs 4 C. brunnea Moore.

CHAMAESIPHO COLUMNA (Spengler), 1790
(Plate i, figure 5; Text-figure 1,q)
Lepas columna Spengler, 1790.

Chamaesipho columna Darwin, 1854; Gruvel, 1905 ; Broch, 1922 ; Nilsson-
Cantell, 1926; Moore, 1944; Pope, 1945; Dakin, Bennett and Pope, 1948,
1952 ; Bennett and Pope, 1953, 1960; Womersley and Edmonds, 1958 ; Wisely
and Blick, 1964.

The species Lepas columna was first erected by Spengler in 1790 for
specimens which had probably been collected during one of Cook’s Expeditions.
Apparently, however, this material was incomplete, for Spengler did not refer
either to the characteristic structures of the opercular valves or to the features
of the soft body parts. The locality quoted was ‘“ Otaheite ”’.

In his monograph (1854), Darwin placed Australian and New Zealand
specimens of Chamaesipho in Spengler’s species, L. columna, but did so with
some reluctance because, although Spengler’s original description fitted his
material in all but one of the few characters described, namely size, Spengler’s
account was not sufficiently detailed for Darwin to be sure that Australasian
specimens belonged to the same species.

ELIZABETH C. POPE 65

The dimensions given by Spengler for his Lepas columna were: height
26-5 mm. and breadth 17-0 mm. The largest Australian specimen in the
present series has the following measurements: height 19 mm. and diameter
6-0 mm., which are considerably less than Spengler’s figures. It is by no
means certain that they represent the largest specimens in Australia but it
is unlikely that any would ever be found as large as those of Spengler. On
this count alone Darwin’s use of Spengler’s specific name for Australasian
material must come into question.

In discussing New Zealand Chamaesipho, Lucy B. Moore (1944, p. 317)
also expressed doubts about the advisability of using the name (. columna
(Spengler) for the commoner New Zealand species of the genus Chamaesipho.
She stated “It seems highly likely that our barnacle is not ZL. coluwmna in
Spengler’s sense’’. However, she continues to use the C. columna (sensu
Darwin) for New Zealand and Australian material, since this name is so widely
current in Australasia. It has been thought worth while in the present review
to re-examine the position, and to attempt to clarify the situation.

Identification of Australasian Chamaesipho with L. columna Spengler

In the first place, it is not likely that a species of Chamaesipho from a
tropical locality such as Tahiti (given as the type locality) would be identical
with one from the southern shores of Australia and temperate New Zealand,
especially when it is known that C. columna (sensu Darwin) is strictly limited
in its Australian distribution to the temperate region reaching the northern,
warmer end of its range some 400 miles south of the Tropic of Capricorn.

Although one or two isolated specimens of 0. columna have been collected
by the author at Lord Howe Island, on intertidal rocks, the species has failed
to establish itself there. The two specimens found were stunted and atypical
in shape, being more like C. scutelliformis in basal outline than regularly-shaped
Australian C. columna. The seas around Lord Howe Island (lat. 32° 8.) are
warm enough to allow limited development of patches of reef corals and, at
the time of collection, the author attributed C. coluwmna’s failure to become
established as being due to its inability to survive in the warmer sea tempera-
tures there.

Tahiti lies well to the north of the Tropic of Capricorn and has a tropical
molluscan shore fauna (fide Dr. H. Rehder in a personal communication to
the author). The Australian species is favoured by the cooler conditions
obtaining on the shores of southern New South Wales and Victoria judging
by its size and density. In southern Tasmania, however, conditions may in
farthest south be too cold for the development of large communities. On the
grounds of sea temperatures alone the present author would agree with Moore
that a New Zealand and Australian species of Chamaesipho is unlikely to be
identical with one from the tropics. There is, however, the possibility that
the locality quoted by Spengler for his L. columna may have been incorrectly
recorded by the collector or have been confused during the long voyage to
Europe. A similar case of a presumably transposed locality occurs in a molluse
described by Spengler (fide Dr. D. F. McMichael, Curator of Molluses, the
Australian Museum). <A species of Austrocochlea which was collected by Cook’s
party, almost certainly at Kurnell, New South Wales, had been attributed to
a wrong country. This mollusc material passed to Humphrey in England
and thence to Spengler. It was finally described by Chemnitz (1781, Conchylien
Cabinet (Martini), 5: 230, Plate 185, figs. 1850, 1851). It was stated there,
to have come from New Zealand where no such shell occurs.

In the absence of the Type material, which the author was unable to
locate in European Museums, it is only possible to re-examine available facts,
and to take into account any recent findings and see if further light can be
cast on the problem. The biggest objection to the acceptance of Spengler’s

E

66 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

specific name for Australasian material has been the disparity in the size of
the specimens described by Spengler and that of Australasian Chamaesipho.
It is, however, believed that there is a possible explanation for this which will
be set out below. .

A re-examination of Spengler’s description, as translated from the original
Danish by Dr. T. Gislén (quoted in Moore, 1944, p. 316-7), reveals a statement
(probably the one that convinced Darwin that he could use Spengler’s name
for Australian material): ‘‘ A beautiful Patella... . is exteriorly completely
covered by small cylinder-shaped Lepades, like those I have just described.
They stand as small columns close to each other, and resemble a honeycomb
because of their white angular openings. These also are from Otaheite”’.

As limpets are absent from certain tropical islands, the author consulted
Dr. Harald A. Rehder (Curator, Division of Mollusks, U.S. National Museum,
Washington, D.C.) who has recently carried out extensive field work on
Tahitian shores. He stated that limpets, of the genus Cellana (formerly known
as Patella) are found in Tahiti as well as on Pitcairn Island but, he said, he had
never seen any of them covered by barnacles of any kind. This statement is
considered significant since, in his letter, Dr. Rehder said ‘‘ Last year (1963)
I covered fairly carefully the entire coastline of Tahiti”? and as he was working
on molluses especially, it is felt he would have seen C. columna on limpets at
least, even if he had not noticed them on the rocky reefs.

Several other scientists, at the author’s request, have searched for C.
columna on the shores of Tahiti but have also failed to find it. One is thus
led to the supposition that it does not occur there today and probably never
did. Spengler’s material, judging from the date of publication, is probably
material taken during one of Captain Cook’s voyages, during which both Tahiti
and New Zealand were visited and the suggestion is made that an error has
occurred in giving the locality of the barnacles. Had it been ‘‘ New Zealand ”’
and not ‘“‘ Tahiti’ everything in the original description would be explicable.
A mistake, such as is suggested, is fairly likely, as both countries were inhabited
by Polynesians and any place names obtained from the indigenous peoples
would sound similar. Confusion between the two would be easy if labelling
was inadequate at the time of collecting and cases of wrong localities have
already been shown to have occurred in other specimens handled by Spengler
from Cook’s Expeditions.

Supporting New Zealand as the original locality for Spengler’s L. columna
are the following facts :—

(1) Spengler described limpets with their shells covered with closely
packed C. columna, as being also from Tahiti. Such an association of barnacles
and limpets is common in New Zealand (and Australia) but (fide Dr. Rehder)
is not found in Tahiti.

(2) Chamaesipho columna has not been recorded in Tahiti since Spengler’s
original record. It is, however, one of the commonest intertidal animals in
New Zealand, occurring in the honeycomb-like aggregation described.

(3) Only in recent years has Lucy Moore shown that not one, but two,
species of Chamaesipho occur on New Zealand shores. Till that time both
Species were included in C. columna but an ecologist (Oliver, 1923, p. 535) had
noted that barnacles from the upper part of the Chamaesipho range were of
considerably larger size. It was this larger type that proved to be C. brunnea
Moore.

It is suggested that herein may lie the explanation for the apparently
impossibly large size given by Spengler for his species L. columna, which no
subsequent worker has been able to equal in specimens from Australasia. If
Spengler’s original sample were New Zealand in origin, he may have been
examining a mixed batch of C. brunnea Moore and OC. columna (sensu Darwin).
The large measurements quoted, may have applied to a specimen of C. brunnea,

ELIZABETH C. POPE 67

which is much more likely to approximate to Spengler’s dimensions than C.
columna. Later workers in New Zealand failed to recognize the second species
in the Chamaesipho zone until 1944, so it is quite likely that Spengler could
also have made a similar mistake, especially if the material he had was eroded
and had the opercular valves and soft parts missing, aS apparently it had.
The acceptance of a New Zealand origin for Spengler’s L. columna would make
his original description much more credible but would still leave doubts as to
which of the two New Zealand species should carry the specific name columna.
Many of the shell characters given are equally applicable to both the species
from New Zealand but, of the few remaining characters, the following apply
better to the smaller more ubiquitous New Zealand species of Chamaesipho :
(1) the description of the orifice ; (2) the barnacles’ habit of growing in honey-
comb-like aggregations ; (3) the orifice is more nearly equal in size to the basal
opening of the shell and sometimes is larger. Spengler described the opening
of the shell thus, ‘“‘ It is wide and much larger than the lowest part of the
shell’ ; and finally (4) its habit of forming a honeycomb-like covering on the
shell of a limpet. Chamaesipho brunnea is of a size too large to crowd enough
specimens on to a limpet shell to produce a honeycomb appearance ; moreover
Moore (1944) does not record it on limpets. The only one of Spengler’s
characters that would better fit C. brunnea rather than the smaller species is
that of size.

The majority of Spengler’s characters therefore apply to the smaller
Australasian species and it is felt by the present author that the specific name
C. columna should rightly be restricted to it. This would fortunately fit in
with Darwin’s (1854) use of C. columna for the small Australian and New
Zealand barnacles, of which he gave an excellent augmented description, and
Miss Moore’s use of the names C. columna and C. brunnea for the two New
Zealand species would be valid. The use of C. columna (sensu Darwin and
Moore) is and has been in wide use in Australia and New Zealand for
approximately 100 years. It is therefore desirable to continue this usage if
it can possibly be done without infringing the International Code. It is hoped,
at a later date, to take the necessary steps and, after due enquiries, to establish
neotypes for the species and set the current use of name on a proper basis.

Description of Australian C. columna

Darwin supplemented the original description of C. columna by adding
considerably detailed accounts of the opercular valves and soft parts as well
as the shell plates, and Broch (1922) and Nilsson-Cantell (1926) have added
further details. A more detailed description of New Zealand material was
given by Moore in order to differentiate between C. columna and C. brunnea,
and there is little need to add to her account. Shell characters and ecological
data for Australia are described and illustrated in Pope (1945); Dakin et al.
(1948) ; Bennett and Pope (1953, 1960) ; and Womersley and Edmonds (1958).
A few comments on similarities or differences between Australian and New
Zealand specimens will be given.

Shell

In shape and appearance of the shell and orifice, Australian C. columna
are comparable with the New Zealand ones but the dimensions of the largest
Australian specimen from a collection made at Cape Bridgewater in Western
Victoria are slightly larger than measurements quoted by Moore, namely,
height 19 mm. and width6mm. The surface of the parietes is only occasionally
pitted in Australian specimens and there is little trace of regular ribbing on
_ the shell; however, the lower edge of the shell may have a crimped, wavy
outline in crowded specimens. As in New Zealand, juvenile specimens may
Show six shell valves but these are soon reduced to four and all trace of sutures
between them may subsequently be lost.

68 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

The plates of the shell may be thicker in the low-growing individuals
forming part of a ‘‘ honeycomb” or in solitary specimens, but in crowded,
tall specimens the walls are generally thin and brittle. The sheath is compara-
tively short and lined by a dark membrane and has a series of darker slightly-
raised rings, marking the successive positions of attachment of the opercular
membrane during growth. The term “ rifled’’ (used in Gislén’s translation
of the original description) implying a spiral marking on the sheath is not
strictly correct, at least in Australian material. The lower margin of the
Sheath does not depend freely into the shell cavity. The structure of the
opercular valves is distinctive among Australian Chthamalidae especially in
the case of the tergum, owing to the presence of the 4-6 deep pits which, with
their separating crests, form the attachment for the depressor muscles and
occupy about half the length of the basal margin. The illustrations of these
valves by Moore (1944, Plate 46, left column, a) and Darwin (1854, Plate 19,
fig. 3,b,c) should be consulted, rather than those of Nilsson-Cantell (1926,
Text-fig. 4,h,7) which show the tergum with crests without associated pits for
the attachment of the depressor muscles. In several Australian specimens
there was a slight demarcation of a spur from the margin of the tergum. In
most, however, the spur and the basi-scutal corner of the valve are one and
the same.

Soft Body

The structures of the soft body have been adequately described by Moore
and only shght variations from New Zealand material are found in the Australian
population. Colour: The colour is navy blue in general but the free margins
of the scuto-tergal folds may be a horny brown colour. The cirri and the
proximal part of the penis are also navy blue but the prosoma and thoracic
region are lighter in colour.

Trophi: The labrum is bullate with a wide, V-shaped groove, the anterior
border of which is fringed by numerous crowded hairs, especially towards
the peak of the V. Moore reports five denticles on the labrum, and in the
present examples there are also five but they are asymmetrically placed,
with one denticle placed on one arm of the V and four on the other. Nilsson-
Cantell (1926, Text-fig. 4a) shows a considerably larger number of denticles.
The mandible has four, or sometimes five, teeth of which teeth three and four
may be doubled. The fifth tooth, when present, is centrally placed in the
pectinated lower section of the jaw as shown in Nilsson-Cantell (1926, Text-
fig. 4c). This pectinated area of the jaw has regular, fine teeth except where
a fifth tooth is present and where the strong tooth-like spine or spines form
the lower angle of the mandible. There is no secondary side-toothing on the
main teeth and the structure of the whole organ is strongly reminiscent of the
jaw pattern found in the stellatus subgroup of genus Chthamalus. The first
Mazxilla is as described by Moore (1944). The two large upper spines are
followed below by 3-5 pairs of small spines in the “ notch”. A second small
notch delineates, below, a central, fairly straight cutting edge which may carry
up to seven pairs of larger spines, second in size only to the topmost ones.
The lower rounded point of the maxilla I has 6—7 pairs of finer spines. The
upper and lower margins are haired. Mazilla II in the present sample differs
in shape from Text-fig. 4e in Nilsson-Cantell (1926) in that its free tip is
generally more pointed and projects further than the rest of the cutting edge.
The notch is distinct and without spines, although there are numerous spines
above and below it. The spines on the anterior border are in general longer
than shown by Nilsson-Cantell.

Corre

Cirri I and IT are distinctly shorter than Cirri IV—VI, while cirrus IIT is
intermediate in size and structure between the other two groups. Cirrus I
has the anterior ramus longer by several segments than the posterior one and

ELIZABETH C. POPE 69

is stouter towards its base. Both rami carry numerous fine spines, bunched
anteriorly, many of which are pinnate. In addition, peculiar grapple-like,
stout short spines also occur on the central segments, as described by Moore
(1944). Cirrus II generally has its anterior ramus shorter by at least three
to four segments (or even more) and is stouter than the posterior one which
may be similar in general build or longer and more filiform, as noted by Darwin,
Broch (1922) and Moore. Many had cirrus II with both rami similar in build
{except that the anterior one is longer and carries considerably more setation)
but one or two had the posterior ramus longer and with the more distal seg-
ments similar in structure and setation to the rami of the more posterior cirri
but in these, the three or four basal segments carried bunches of pinnate setae
plus crowded groups of grapple-like spines of the type shown in Text-fig. 1,g.
The anterior ramus of cirrus II carries dense bunches of spines (both pinnate
and ordinary) anteriorly, together with numerous grapple spines on the more
distal segments. They are specially abundant round the bases of segments
4-6. These grapple-like spines are obviously similar in origin to the stout
lanceolate spines seen in certain species of the genus Chthamalus and in fact
are closely similar in build to those of Chthamalus intertextus. In all the grapple
spines the paired recurved side spines are less numerous than the serrations
in the lanceolate spines of other Chthamalids. In Australian material grapple
Spines vary both in the number of segments on which they occur and also in
the number found on each segment. They appear to serve as grapples to hold
food particles. Cirrus III generally has sub-equal rami and otherwise is as
described by Moore, possessing both the plumose spines and the hooked, three-
or five-pointed grapple spines described for cirrus II. The latter occur on the
basal segments amidst the thick tuft of spines. No specimen with antennae-
form posterior rami have been seen in the present batch. Each segment of
cirrus III towards the centre of the ramus carries five pairs of long normal
spines. Cirri IV to VI are normal in structure and nearly equal in size ; more-
over, the paired rami are sub-equal. Each segment is bullate and bears four
pairs, sometimes five pairs, of stout spines of which the distal pair is longest
and size decreases towards the proximal part of the segment.

The penis is stout, ringed and even in a somewhat contracted state is
longer than cirrus VI. It tapers rapidly towards its tip which has scattered
hairs along it. There are two tufts of moderately long hairs at the very tip.
There is no caudal appendage. In the Sydney area of New South Wales,
Wisely and Blick (1964) record C. columna breeding in winter and early spring.

Habitat

Chamaesipho columna shows a preference for attaching to a rocky substrate
on exposed coasts in Australia and its range does not extend far inshore, in
inlets or harbours. Moore records it “‘ very occasionally ” on wood, but none
have yet been recorded on this substrate in Australia. It grows in the upper
half of the intertidal zone approximately up to the mean or the lower high
water levels of spring tides. The upper level of its distribution depends tre-
mendously on the amount of wave action in the area. Its range extends
downshore to a varying degree, according to environmental conditions and
whether it meets competition for attachment from other species. In New
South Wales its vertical range is apparently limited below, by competition
with the two surf barnacles, Catophragmus polymerus and Tetraclita rosea for,
im areas of western Victoria where lower sea and air temperatures exclude the
surf barnacle, Catophragmus polymerus, C. columna ranges further down the
Shore to the short algal turf near low water of neap tides (Bennett and Pope,
1953, p. 117).

Geographical Distribution
Australian: Chamaesipho columna occurs on the temperate shores of
Australia from the region of Cape Byron in northern N.S.W., southwards to

70 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

Victoria and Tasmania, where it occurs on all exposed coasts and, finally, it
ranges westward along the southern coast of the continent to Point Sinclair
(in the eastern half of the Great Australian Bight) where it is recorded by
Womersley and Edmonds (1958, Plates 1, 2). It has also been taken on one
occasion at Lord Howe Island in the Tasman Sea but had failed to establish
itself here.

World Occurrence: Moore has recorded C. columna on all the shores of
both the North and South Islands of New Zealand and in the Kermadecs. The
validity of the record of this species by Spengler for Tahiti (Otaheite) has been
challenged above and, until it is confirmed, it is believed that it should be
disregarded.

DISCUSSION

The collections studied for the present review contained individuals in
each species of markedly greater size than any previously described. As the
specific distinctions in many species of Chthamalids become increasingly clear
in fully grown material, the descriptions have been modified, where necessary.
It is felt that the recording of details of anatomy for each species will be of
special interest to workers presently engaged in a phylogenetic review of the
family. The importance of basing specific identifications on aS many characters
as reasonably can be used, cannot be overstressed because of the undoubtedly
close relationships evident between species from widely separated zoogeograph-
ical regions as well as those from adjacent localities. The differences between
Species are realities but are often very difficult to define, and are best shown
in such details as the structure of the serrated spines of cirri I and IT or the
fine differences to be seen in the pinnate spines (see Text-fig. 1,b-e) which are
less likely to vary than the more evident shell characters generally used.

While the general patterns of the trophi of most chthamalids are basically
alike, the fine differences between species can often best be appreciated by
viewing the barnacle’s ‘‘ face’’ from the front, before beginning dissections
of the mouth parts. The shapes of the ‘‘ beard” and ‘‘ side whiskers ”’ are
often quite characteristic and some species are notably more bristly than others.
Catophragmus polymerus, for instance, is extremely well provided with setae
round its mouth parts. There is often also a characteristic colour pattern
round the mouth.

The degree to which the rami of cirri I and II are curved in over the mouth
is also characteristic in certain species and the type of teeth and general arrange-
ment of the teeth and hairs above the groove of the labrum can also be of
specific importance. However, the necessity of focusing the microscope
through several planes should be remembered, as there may be teeth on two
separate levels and hair on yet another. The very distinct triangular teeth
of Chthamalus withersi, lying on either side of the peg-like teeth, seem to have
been overlooked previously because of failure to explore the different levels
of the groove of the labrum.

In existing keys to species of the genus Chthamalus, stress has been laid
on the structure of the mandible as a means of subdividing this somewhat
unwieldy genus into sub-groups. The present study has shown, however,
that the dividing line between those species with the so-called tridentate or
hembeli jaw and those with the quadridentate or stellatus kind of mandible is
sometimes a rather hazy one, especially if one is dealing with juveniles or
individuals which are regenerating jaws after damage (by no means a rare
occurrence). It is considered that some other characters might be preferable
to use for the primary separation of Chthamalus into sub-groups. For example,
the presence or absence of calcareous layers in the basis, even if it is a product
of secondary calcification, might result in more natural subgroups since it
would bring together Chthamalus hembeli (Conrad), C. intertextus and C.
caleareobasis Henry which have been shown by Newman (1961) to have many

fond

ELIZABETH C. POPE AA

shell structures in common that differ rather markedly from remaining species
in the genus, and this might be borne out by comparisons of the morphology
of their soft bodies which were, unfortunately, not available in the present
instance for examination.

Another interesting character which may be obscured, unless special care
is taken during the making of microslides of the labrum, is seen in species like
Chthamalus intertextus and C. caudatus, as contrasted with species like C.
malayensis and OC. antennatus which have the normal rounded or bullate labrum,
characteristic of the family. It is the semicircular, rather funnel-like structure
on the front of the labrum above the palps, best seen when the ‘ face ”’ of the
barnacle is examined before dissection. The muscles which can regulate it
and cause it to assume the funnelled shape which leads towards the mouth,
can be seen in cleared preparations, and in C. intertextus the whole of this
peculiar structure, with its pitted surface (like a brain coral in appearance),
seems to function as a hopper to channel the food particles (swept towards
it by the incurved cirri I and II and by the palps) towards the mouth.

Other species of Chthamalus may also possess the peculiar type of labrum
described above, for one author remarks, without being specific, that the
semicircular extension on the labrum of C. caudatus is seen in several other
species of Chthamalus. If it is of more widespread occurrence, it might prove
to be a significant character for differentiating subgroups within the genus
since, so far as Australian material is concerned, it is associated with species
included by Nilsson-Cantell (1921) in his so-called hembeli group.

Living, as they do, in the upper half of the intertidal zone the actual times
available to the various species of shore chthamalids for feeding is limited to
their periods of immersion. This ranges from a mere 2% of the time at the
highest levels to a maximum of approximately 30° of the time for mid-tide
species. That the Chthamalidae have managed to flourish in such conditions
is a clear indication of the efficient food-capturing adaptations that have been
evolved within the group. It was therefore of interest to find that only two
broad patterns of food-capturing have apparently been evolved and that they
can be correlated with the degree and force of water movements experienced
by the barnacles during feeding. The two groups of species showed morpho-
logical difference in their mouth parts and cirri which are considered to be
of an adaptive nature. :

Some of the adaptations enabling chthamalid barnacles to catch sufficient
food appear from circumstantial evidence to be very effective. Attention
was first drawn to this matter during the preparation of microslide mounts.
Many cirri and mouthparts were entangled with food particles and debris and
much patient brushing and picking clean of serrated spines was necessary to
reveal the underlying structures. It was noticed that there were two fairly
well defined patterns of distribution of the particles and one could be associated
with the requirements of feeding on the high shore for very short periods under
relatively calm conditions, and the other could be associated with the need
to capture and hold food in turbulent waters—turbulence due either to surf
or to choppy seas, or to the currents caused by strong tidal ebb and flow.
These last are greatest in the mid-tidal zone.

In Octomeris brunnea, Chthamalus caudatus and C. withersi food particles
were found fairly evenly distributed and entangled in the setation of the cirri
and mouth parts and were not aggregated markedly into bolus-like clumps.
The exception was O. brunnea, where a slight tendency was noticed for particles
to be caught and amassed in the bunches of pinnate setae towards the base
of the anterior ramus of cirrus III. However, these aggregations of food were
not as large as those seen in species to be mentioned below.

Feeding in these high shore, tropic species is apparently carried out in a
hurried and indiscriminate way by the sweeping of all particles forwards

72 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

towards the labrum. In those species with the semicircular processes above
the palps the food can be quickly diverted towards the mouth and even the
possession of a flat-fronted labrum, rather than a bullate one, would have
adaptive advantages for feeding. There is no evidence of a tendency to “‘ pick
over’ the food particles. It is perhaps significant that among the Australian
species of Chthamalus those with the most highly developed semicircular funnels
on their labra (C. caudatus and C. intertextus) tend to be hypobiotic in habit.
Is this adaptation of the labrum connected in some way with their need to feed
in an ‘upside down” position? However, this would have to be verified
experimentally.

In the turbid and briefly slack water conditions obtaining on the high
shore during normal conditions of high tide in tropical seas, there is no lack
of suspended detritus or other particles, as may be verified by standing on
this part of the shore when the tide is at its highest or by looking at the deposits
of organic matter left at high tide mark on beaches. One can well imagine
the effectiveness of making a series of quick, sweeping dips with very setose
cirri in this concentration of food particles and the brushing of them quickly
towards the mouth, as outlined above. In the swifter flowing water movements
of the mid-tide zone or under the action of surf, the retention of fine detritus
would be almost impossible owing to the flushing action of the water. Here
other feeding mechanisms designed to entangle and hold larger food particles
as well as detritus have been developed.

In Catophragmus polymerus, Chamaesipho columna, Chthamalus antennatus,
C. intertextus and C. malayensis, in addition to the particles of food distributed
randomly among the pinnate setae, there were compact, rounded clumps of
food, entangled and held by the serrated spines and these aggregations were
most difficult to dislodge by brushing. The spines often had to be picked
clean by means of fine needles in order to show their structure. The fact that
these serrated or grapple spines are situated opposite the level of mouth and
within reach of the jaws is also considered significant. Small planktonic
organisms and detritus were detected in the food clumps on the spines. It
would appear that particles sieved and “ dip-netted ” from the water by the
tufts of pinnate setae are gradually passed downwards on the ramus and
accumulated and held by the stout serrated spines till they form small rounded
masses. Such an accumulation could later be picked or brushed off by the
toothed and setose mandibles or other mouth organs and passed into the
mouth. Indeed, several such balls of food matter were discovered between
the jaws of individuals which had been feeding just prior to preservation. In
such species where food is picked over and transferred to the mouth by the
trophi from the anterior cirri, a bullate labrum would be no disadvantage.

Serrated spines or grapple-spines occur in Catophragmus, Chamaesipho
and Chthamalus antennatus which live on surf beaten rocks and also in Chthamalus
malayensis and C. intertextus, the two species which live on those parts of the
tropical intertidal zone most exposed to water movements. While the wave
action they experience is not comparable with that consistently endured by
the three southern Australian species mentioned, it is nevertheless the maximum
experienced within the Australian tropics and the one most consistently
subjected to strong ebb and flow of tidal currents.

No ecological information was supplied with the New Guinea material of
Chthamalus intertextus, except that it occurred with C. malayensis in the mid
tide zone. Hiro (1939) records the species also with Octomeris brunnea ‘‘ on
the underside of boulders ” at Taiwan, so the peculiar combination of feeding
adaptations it possesses may be a reflection of its special needs for feeding both
in an inverted position and in waters showing a considerable degree of movement.

If serrated spines and extra setation are indeed effective adaptations for
feeding in turbulent seas, one would expect to find most adaptive structures
of this kind in the surf barnacle, Catophragmus polymerus, since of all Australian

ELIZABETH C. POPE 73

intertidal chthamalids it is subjected to roughest seas for the longest periods
and, conversely, one would expect to find fewest of them in Chthamalus
witherst since it occurs in areas subjected to least water turbulence.

This is indeed so, for Catophragmus (Text-fig. 1,a, b) not only has cirri and
mouth parts that are extremely setose, with many setae of the feathery pinnate
type, but it also carries numerous stout barbed spines on the distal segments
of the rami of cirrus II; the groove of the labrum, above the mouth carries
both hairs and teeth. In addition, each of the segments of the rami IJI-VI
carries a small thick tuft of shorter bristles situated centrally between the rows
of paired longer spines. These short bunches of spines serve to trap and hold
food particles (Text-fig. 2,a). Such bunches of short spines also occur in stalked
barnacles in the genus Mitella, but they are not found generally in any other
species of the family Chthamalidae in Australia. Darwin (1854) mentions
that in Catophragmus imbricatus the intermediate spines on each segment of
cirri III to VI are fewer in number than in Catophragmus polymerus. By
contrast O. brunnea (generally considered as a closely related genus) has tufts
of short intermediate spines on only a few of the basal segments of the anterior
ramus of cirrus III and not on every segment of cirri ITJ-VI. Their limited
occurrence in Octomeris brunnea can perhaps be linked with the fact that such
structures are unnecessary for food capture in calmer tropical seas. In fact
they might be a hindrance to the quick propulsion of fine food particles towards
the mouth.

In Chthamalus withers: the front of the labrum above the palps is somewhat
flattened, but in no specimen among those dissected did any individual show
signs that the labrum could assume the funnel-like shape seen in C. caudatus
and C. intertextus. The smaller central teeth and wide shallow groove of the
labrum could be adaptations allowing free movement of minute food particles
into the mouth. As organs to strain the particles from the water and propel
them towards the mouth, the unspecialized but very setose cirri I and IT are
very effective and the incurving of their rami would be an added advantage
for the quick and effective gathering and swallowing of food in a short time.
Short, serrated spines are completely lacking in C. withersi which would appear,
therefore, to be unable to hold aggregations of food or catch and imprison
active plankton of slightly larger size.

Intermediate between Catophragmus and Chthamalus withersi in the
adaptations developed for holding and concentrating food particles are
Chamaesipho columna, Chthamalus antennatus, C. malayensis and C. intertextus.
All have either short spines with serrations or grapple-like spines on their
second cirri, as well as the tufts of pinnate setae. In addition C. intertextus
and Chamaesipho columna have grapple-spines on a few basal segments of the
anterior ramus of cirrus III, directly opposite the mouth, though in neither
of these species are there tufts of spines on all the other cirral segments.
Chthamalus malayensis has extra stout spines posteriorly on certain of the basal
segments of the anterior ramus of cirri I and IJ, which are thought to act as
devices locking together the two rami much as hamuli may lock together the
wings of certain insects. In this way a wide and more rigid structure could
result for dipping food particles from the water. Such stout posterior spines
occur only on the anterior rami and are often found interlocked with the fringe
of setae of the posterior ramus but, as with the other structures mentioned
as possible feeding adaptations, much experimental work and detailed study
of corresponding organs in other barnacles is necessary before their effectiveness
or otherwise as adaptations for feeding in the rigorous environment of the upper
half of the intertidal zone can be truly evaluated.

The different combinations of adaptive feeding structures found, for
instance, in each of the four tropical species in the genus Chthamalus in Australia
point to the fact that each of the species may indeed be feeding in a slightly
different way or on a slightly differing diet, and hence be occupying a different

74 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

ecological niche, even when, as has occasionally happened, three of them have
been collected within a circumscribed area of a few square centimetres. An
intensive study of the group Chthamalidae could do much to clarify some of
our ideas about intertidal niches and related problems.

Acknowledgements

Many people and staffs of museums and university departments of
zoology have been of great assistance during the making of Australian
collections and the study of relevant research collections abroad, during the
preparation of this review. It is impossible to mention them all by name.
The author is very grateful for their assistance and this opportunity is taken
of thanking them. While it is invidious to select names of persons and
institutions for special mention the assistance given by the following has
merited special thanks: In Australia, my colleagues of the Australian Museum
who have made collections for me, wherever possible, Miss Isobel Bennett,
Mrs. Loisette Marsh, Mr. Bert Jackson, Dr. E. P. Hodgkin and the Directors
and curators in charge of Crustacea from the Western Australian Museum,
the Queensland Museum and the National Museum of Victoria.

While in Europe I was greatly indebted to numerous people for allowing
me to study important collections of barnacles and for their helpful assistance
in obtaining literature. To the following, special thanks are due: The Director
and Dr. J. P. Harding of the British Museum (Natural History), London for
permission to examine their most important collections of cirripedia named
by Darwin and Nilsson-Cantell; Dr. C. M. Yonge and staff of the Millport
Marine Biological Station; the Director and staff of the Marine Biological
Association’s Laboratory, Plymouth, U.K.; Dr. Denis Crisp of the Marine
Biological Station of North Wales and the British Council in London for a
small grant, in aid of research in Great Britain. In the Netherlands, assistance
was received from the following: The Director and Dr. Engels of the Zoologisk
Museum, Amsterdam, for permission to examine the large collections of
cirripedes taken by the Siboga Expedition and in Leyden the Director and Dr.
Lipke Holthuis of the Rijksmuseum van Natuurlijke Historie for permission
to examine their extensive collections from Indonesia (including those of the
Snellius Expedition) and special thanks is due to Dr. H. Boschma for help in
obtaining valuable literature. In Denmark, thanks are due to the Director
and to Dr. Torben Wolff of Universitetets Zoologisk Museum in Copenhagen
for permission to examine the extensive collections of cirripedes, resulting from
Dr. Mortensen’s Pacific Expedition of 1914-16 (worked by Dr. H. Broch).
In Sweden, help was received from the Director and curator of Crustacea of
the Naturhistorika Riksmuseet in Stockholm and permission given to examine
the very valuable collections forming the basis of Dr. C. A. Nilsson-Cantell’s
(1921) monograph, in addition to the collections from Dr. Mjéberg’s Swedish
Expedition to Australia, 1910-1913. In the United States of America, thanks
are due to the following institutions and people for allowing me to work on
their extensive and valuable collections: The Director and Dr. John Garth
of the Allan Hancock Foundation of the University of Southern California
(cirripedes from the eastern Pacific islands and western seabord of North,
Central and South America); the Director and Dr. Fenner Chase Jun. and
Dr. D. Squires of the Division of Marine Invertebrates for permission to examine
the collections forming the basis of the famous 1916 Monograph by the late
H. A. Pilsbry, which included most of the type material relevant to tropical
Australian species of Chthamalus ; the Director and Dr. Tucker Abbott of the
Department of Mollusks, Academy of Natural Sciences of Philadelphia, Penn.,
for access to further Pilsbry material ; and finally for sending relevant collec-
tions of barnacles to me, for examination, thanks are due to Dr. Huzio Utinomi
of Japan, Drs. Zevina and Tarasov of U.S.S.R. and Dr. Y. M. Bhatt of the
Institute of Science, Bombay, India.

ELIZABETH C. POPE rs)

Thanks are also due to the Director, and certain officers of the Australian
Museum for specialized help during the preparation of this review, to Mr. John
Beeman for preparation of the line drawings, and Mrs. E. Brown for lettering
on Text-figures, to Messrs. H. Hughes and C. Turner for photographs and help
in preparation of the plates and especially to my assistant Miss Janet Walsh
for help in cataloguing distributions and for help in preparation of the
manuscript and bibliography. Thanks are also due to the staff of the Library
for help with literature and the bibliography.

Finally, grateful acknowledgement is made to the numerous friends and
acquaintances who aided me during field trips and by making collections at
many inaccessible places round Australia or in neighbouring Pacific islands.
Acknowledgement for two photographs used in Plate i, figs 4 and 5 is also
made to Mr. Justice F. G. Myers.

It is hoped that no important acknowledgement has been overlooked and
if any such oversight has occurred it is unintentional.

Selected References

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Brocu, H., 1916.—Results of Dr. E. Mjoberg’s Swedish Scientific Expeditions to Australia
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, 1931.—Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-16. LVI. Indo-
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CRANWELL, L. M., and Moors, L. B., 1938.—Intertidal communities of the Poor Knights
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Daxin, W. J., BENNETT, ISOBEL, and Pops, EvizapetTH, 1948.—A study of certain aspects of
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Darwin, C., 1854.—‘‘ A monograph of the sub-class Cirripedia, the Balanidae, and the
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DavaDIE, CLAUDE, 1963.—‘Systematique et structure des Balanes fossiles d’Europe et
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ENDEAN, R., STEPHENSON, W., and Kenny, R., 1956.—The ecology and distribution of inter-
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FiscHer, P., 1884.—Cirripédes de L’archipel de la Nouvelle-Calédonie. Bull. Soc. zool. Fr.,
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GRuvEL, A., 1903.—Revision des Cirrhipédes Opercules. 1. Partie Systematique. Nouv. Arch.
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Guiter, BE. R., 1952.—The nature of intertidal zonation in Tasmania. Pap. roy. Soc. Tasm..,
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76 AUSTRALIAN AND SOME INDOMALAYAN CHTHAMALIDAE

Henry, Dora P., 1957.—Some littoral barnacles from the Tuamotu, Marshall, and Caroline
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Hiro, F., 1932.—Report of the Biological Survey of Mutsu Bay. 25. Cirripedia, Scz. Rep.
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KonosvAry, G., 1941.—Die Formenkreise der Chthamaliden. Zool. Anz., 133, 3-4: 67-81.

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HLIZABETH C. POPE V7

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li: 122-124.
Womerstey, H. B. 8., and Epmonps, 8. J., 1958.—A general account of the intertidal ecology
of South Australian coasts. Aust. J. Mar. Freshw. Res., 9, 2: 217-260, Pls. 1—12.
Zevina, G. B., and Tarasov, N. I., 1963.—Marine foulmg and Teredo. The cirripede barnacle
fauna (Cirripedia: Thoracica) from the continental littoral of S.E. Asia. Transactions
of the Institute of Oceanology, \xx, Moscow: pp. 76—100.
ZuLLo, V., 1963.—A classification and phylogeny of the Chthamalidae (Cirripedia : Thoracica).
Summary only of a paper read before the XVI International Congress of Zoology, in Wash-
ington, D.C. Proceedings Vol. 1, p. 190.

EXPLANATION OF PLATES [I AND II

Plate I. Five Species of Australian Chthamalidae. 1, Chthamalus intertextus growing on
basalt. Traces of imterlocking laminae (like lines of growth) appear adjacent to the wavy
suture, to the left of the lowest shell plate. 2, Catophragmus polymerus. 3, Well-grown and
heavily eroded Octomeris brunnea (originally described as O. intermedia but proved to be adult
of species in fig. 6). Left (from below). showing central hole in basis, and right from above.
4, Fully grown Chthamalus antennatus from Sydney. Note light enamel-like apices on certain
plates of shells (3 top left barnacles). Accompanying littormid is Melarapha unifasciata.
5, A group of Chamaesipho columna with accompanying littormid molluscs. 6, Juvenile
Octomeris brunnea (not to same scale as those in view 3) with narrow ribs and well-defined
interlocking sutures.

Plate II. Some Tropical Chthamalus spp. and their habitats. 1, Roots of the Red Man-
grove, Rhizophora, provide a suitable substrate for Chthamalus barnacles. A stream near Cairns,
Queensland. 2, Closer view of the root system of Rhizophora, showing Chthamalus withers
generally on the lower side of the roots. 3, Dense populations of Chthamalus malayensis im a
favourable habit, tropical Queensland (see text). 4, Typical habitat for O. brunnea and
Chthamalus caudatus m shaded sides or under the overhang of tumbled boulders, on the higher
areas of tropical shores. These two barnacles are not visible on the rocks normally, and are
generally overlooked. 5, Pier of road bridge over a coastal creek in central coastal Queens-
land, showing dense barnacle populations in upper intertidal zone. From the poimting finger
upwards, Chthamalus withersi covers the surface, while below this a species of Balanus replaces
it entirely. 6, The smooth, ribbed form, common in juvenile Chthamalus malayensis or m Queens-
land specimens generally. 7, Eroded adult Chthamalus malayensis from Western Australia
wharf timber. A single Chthamalus antennatus from Sydney has been placed alongside (in the
‘white circle) for comparison in size and sculpturing.

CHROMOSOME NUMBERS IN SOME AUSTRALIAN LEAFHOPPERS
(HOMOPTERA AUCHENORRHY NCHA)

M. J. WHITTEN
Department of Botany, University of Tasmania

With an Appendix by J. W. Evans, Australian Museum, describing a
new genus and species of Eurymelidae

(Plate iii)
[Read 31st March, 1965]

Synopsis
Chromosome numbers are recorded for twenty-seven species of Hurymelidae, thirty species:
of Cicadellidae and eight other species of Homoptera Auchenorrhynecha. A study of meiosis
reveals bivalent co-orientation at Anaphase I m the Auchenorrhyncha while a consideration of
chromosome numbers suggests several trends related to the phylogeny of the Cicadellidae.
Having diffuse centromeres (or polycentric chromosomes), chromosome number may be a more
variable character in leafhoppers than in other groups.

INTRODUCTION

The Australian flora has for some time been subject to cytological
investigation (for review see Smith-White, 1959), while the fauna, in particular,
insects, has suffered neglect in this respect. The incomplete taxonomy of most
insect groups and the unavailability of keys, continue to make such studies.
difficult.

Australian leafhoppers, however, constitute a welcome exception for,
although keys are not yet available, the group has been studied extensively
by Dr. J. W. Evans and the identification of specimens has become practicable.

In the present work an attempt has been made to examine representatives
of all distinctive groups but efforts were especially concentrated on collecting a
representative sample of the only family of leafhoppers endemic to Australia,
the Eurymelidae.

Interest has been added to the problem because leafhoppers belong to the
Hemiptera, the only known order of insects whose chromosomes lack localized.
centromeres (Hughes-Schrader & Ris, 1941; White, 1954). It was hoped the
study might reveal the phylogenetic significance of chromosome number and
its variation in groups where the restrictions imposed by localized centromeres.
are not present.

METHODS

Material for cytological examination (usually the whole insect) was fixed in
ethanol-acetic acid (3:1) for 24 hours and then washed and stored in absolute
alcohol at —10°C. Storage under these conditions did not result in any
detectable deterioration of the material. Some specimens were fixed and stored
in chloroform: alcohol: acetic acid solution (4:3:1). This was equally
satisfactory, except that some difficulty was encountered in spreading the cells
after several months of storage.

Spermatogonial squashes were made in aceto-orcein. Although both
meiotic and mitotic cells were examined where possible, the bulk of the
chromosome determinations were made on meiotic cells of 1st and 2nd division.

PROCEEDINGS OF THE LINNEAN SociETY oF New SoutH Watss, Vol. 90, Part 1

M. J. WHITTEN

TABLE |

List of Chromosome Numbers

Haploid
Family and Species Chromosome Locality*
No.
CICADELLOIDEA
1. EURYMELIDAE
EURYMELINAE
Hurymela
fenestrata Le Pelletier & Serville. . 10+X an 45, & Se WO. let id Gs ue,
20, 22, 53
distincta Signoret Ss 104+X 23, 24
erythrocnemos Burmeister . . 104+X 25
Eurymeloides
pulchra Signoret 104+ X “Le, AO, Bi BS
bicinctus Erichson 10+ X 29, 32
punctata Signoret 104+ X 40, 41
perpusilla (Walker) 104+X 47
Hurymelops bicolor (Burmeister) 10+xX 5, 19, 41, 42
Paureurymela parva Evans : 104+X 25
Pauroeurymela amplicinta (Walker) 104+X 39
Hurymelita terminalis (Walker) 10+X 43, 44, 45
Aloeurymela gearyi Evans} 94+xX 43, 46
Hurymelessa moruyana Evans 104+ X 47
Hurymelella tonnotri Evans 104+X 18
POGONOSCOPINAE
Pogonoscopus myrmex China 10+X 31
IPOINAE
Ipoella
fidelis Evans a 104+ X 4, 34, 43, 46, 5, 41
sp. (novo) (62.11.200) 104+X 5
sp. (novo) (62.4.33) 10+X 13
Ipoides
hackert Evans 104+ xX 43
honiala (Kirkaldy) 10+X 48
Anipo
pallescens Evans 10+xX 5), ls
brunneus Evans 104+ X% 2, 28
Katipo
rubrivenosa (Kirkaldy) 104+X i, @, 3)
Signoretti Kvans ae ie 104+X 47
Anacornutipo lignosa (Walker) 104+X 19, 20, 43, 46
Opio multistrigia (Walker) +X 5
2. CICADELLIDAE
ULOPINAE
Taslopa montana Evans 9+ xX I, oe
8+XY 38
Cephalelus
minutus Evans 10+X B, IN, ey
sp. (64.2.35.2) 10+X 38
J ASSINAE
Batrachomorphus sp. (64.1.34.4) 9-- xX 34
APHRODINAE
Euacanthella palustris Evans 9+X 15
TARTESSINAE
Tartessus
flavipes Spanberg 13--X 18
fulvus (Walker) 12+xX 1, 30, 33, 36, 38, 48, 49, 55
sp. (62.4.69) 13+X 44
AUSTROAGALLOIDINAE
Austroagalloides brunnea Evans J14+X 29

* See end of Table 1 for key to localities.

+ See Appendix.

80 CHROMOSOME NUMBERS IN AUSTRALIAN LEAFHOPPERS

TaBLE 1—Continued

List of Chromosome Numbers

Haploid
Family and Species Chromosome Locality*
No.
LEDRINAE
Thymbrini
Rhotidoides dongarrensis Evans .. 10+X 30
Rhotidoides sp. (64.9.30.2) es 104+-X% 30
Putoniessa sp. (64.4.33.3) oe 10+ xX 33
Stenocotini
sp. (64.9.30.5) SE as Ee 10+X 30
‘CICADELLINAE
Cicadella angustata (Evans) a 8+X 1
IDIOCERINAE
Idiocerus sp. (63.4.17.4) ae is 8+X 17
HECALINAE
Paradorydium brighami Kirkaldy 104+X% 34
‘TYPHLOCYBINAE
Erythroneura sp. (63.4.15.6) a +X 15
Typhlocyba sp. (63.3.4.2) .. Ns 94+ xX 4
sp. (64.3.38.4) .. ae 8+XY 38
DELTOCEPHALINAE
Deltocephalus
longuinquus (Kirkaldy) .. se 4+X 17
3+ xk 17
taedius (Kirkaldy) ate 2 5+ Xx 35, 15, 37
sp. (64.1.34.2) Py: ine be 4+X 34
sp. (64.3.18.4) Je Re ye 5+X 18
sp. (64.3.1.14) Bs Se zi 5+xX I 3}, Be}
sp. (64.3.15.5) ts Me ae 6+X 15
sp. (64.3.15.1) ee ae Se 5+X 15; 35
Phrynophyes kirkaldyi Evans oes 9--xX 3
Aconurominus flavidiventris (Stal) .. 7+xX 17
Balclutha sp. (64.1.34.5) oh ae 7+ 34
Nesoclutha obscura Evans .. sre 8+X iL7/

3. MEMBRACIDAE

Sextius virescens Fairmaire a a 10+xX 5, 48, 50, 51, 54
Hufairmairia fraternus Distant es 104+X 19, 49
Other AUCHENORRH Y NCHA
CERCOPOIDEA
Philagra parva Donovan .. os Tap2X 5, 48
CICADOIDEA
Cyclochila australasiae Donovan .. 9+X 5
FULGOROIDEA
FLATIDAE
Siphanta sp. .. of 5 36 13--xX 5
EURYBRACHIDAE
Dardus erebus Distant SC e 13+X 51, 52
FULGOROIDEA sp. (64.9.30.6) .. a5 13+XY 30
COLEORRHYNCHA
PELORIDITDAE

Hemiodoecellus fidelis (Evans) 2N=22, 23 m males 2

* See end of Table 1 for key to localities.

Key to Localities
1. Collin’s Bonnet, T.; 2. Shoobridge Bend, Mt. Wellington, T.; 3. Mt. Nelson, 7. ;
4. Sandy Bay, T.; 5. Rookwood, N.S.W.; 6. Spring Hill, T.; 7. Hollow Tree, T.;
8. Homebush, N.S.W.:; 9. Huon River, Franklin, T.; 10. Parramatta, N.S.W.; 11. Geeveston,
T.; 12. Arve River, Hartz, T.; 13. Roseville, N.S.W.; 14. Como. N.S.W.; 15. Springs,
Mt. Wellington, T.; 16. Blakehurst, N.S.W.; 17. Bruni Island, T.; 18. Lake St. Clair, T. ;
19. Wellington, N.S.W.; 20. Coonabarabran, N.S.W.; 22. Sale, Vic.; 23. Murrurundi, N.S.W. ;

M. J. WHITTEN 81

24. Lueas Heights, N.S.W.; 25. Barjarg, Viec.; 26. Bankstown, N.S.W.; 27. Windsor, N.S.W. ;
28. Dubbo, N.S.W.; 29. St. Ives, N.S.W.; 30. King’s Park, W.A.; 31. Red Hill, W.A.;
32. Mt. Colah, N.S.W.; 33. Adamsfield, T.; 34. Deniliquin, N.S.W.; 35. Cradle Mt., T.;
36. Huon Valley, T.; 37. Huonville, T.; 38. Plateau, Mt. Wellington, T.; 39. Warrah, N.S.W. ;
40. Regents Park, N.S.W.; 41. Chester Hill, N.S.W.; 42. Canberra, A.C.T.; 43. Bourke,
N.S.W.; 44. Nyngan, N.S.W.; 45. West Wyalong, N.S.W.: 46. Walgett, N.S.W.;
47. Grampians, Vic.; 48. Glenmorgan, Qld.; 49. Gunnedah, N.S.W.; 50. Trangie, N.S.W. ;
51. Westmar, Qld. ; 52. Toowoomba, Qld.; 53. Launceston, T.; 54. Bendigo, Vie. ; 55. Cobar,
N.S.W.

LOCALISED CENTROMERE
CO-ORIENTATION

DIFFUSE CENTROMERE

AUTO-ORIENTATION

DIFFUSE CENTROMERE
CO-ORIENTATION

Fig. 1. Diagrammatic representation of bivalent behaviour at first division of meiosis: (a), when
the centromeres are localized as occurs in most organisms; (b), in Aphids and Coccids and the
plant genus, Luzula ; (c), im Auchenorrhyncha and Psyllidae indicating the two ways of forming
the metaphase bivalent which result in co-orientation. (c) was interpreted from meiotic cells of
Hurymela fenestrata.

CHARACTERISTICS OF MEIOSIS

During meiosis and mitosis in Homoptera the spindle attaches along the
entire length of the chromosome, producing characteristic configurations at
metaphase and anaphase (Fig. 1, b, ce, and Plate iti). In aphids and coccids
(Fig. 1, b) the homologues of each bivalent orient independently on the metaphase
plate (auto-orientation) resulting in post-reduction (Ris, 1942, 1945; Brown,
1954; Hughes-Schrader, 1948). In Psyllids (Whitten, unpublished) and in
the Auchenorrhyncha the bivalents orient in the normal manner (co-orientation)
resulting in pre-reduction (White, 1954; Halkka, 1960a). Rhoades (1961)
questions the occurrence of co-orientation in the Auchenorrhyncha and suggests
careful observation may reveal that the diffuse centromere is always associated

F

82 CHROMOSOME NUMBERS IN AUSTRALIAN LEAFHOPPERS

with auto-orientation. Unequivocal evidence has been found to show this is
not the case.

In Deltocephalus longuinquus (Cicadellidae), males normally possess four
bivalents plus an X chromosome. However, one individual was found with three
bivalents plus an X while another had two bivalents, a trivalent and an X (see
Fig. 2 and Plate iii, h). Since all the other species of Deltocephalus so far examined
have four or more bivalents (see Table 1) it is suggested that a fusion has taken
place between two chromosomes in D. longuinquus and the population is
polymorphic for this fusion.

An analysis of anaphase I and metaphase II cells from the heterozygous
individual indicated that the fused chromosomes go to one pole while the two
single chromosomes are drawn to the opposite pole. This outcome is not possible
if auto-orientation were operating. <A careful analysis of chiasma terminalization
and bivalent orientation in Hurymela fenestrata confirms this finding (Fig. 1, ¢
and Plate iii, b, ¢, d).

The phylogenetic significance of co-orientation in the Auchenorrhyncha
and the Psyllidae will be discussed elsewhere.

°
sf
e8

(vi

as é@
ses g-
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Fig. 2. (i-tii), Different karyotypes in the one population of Deltocephalus lonquinquus. In (ii)
there is heterozygosity for a chromosomal fusion, while in (ii1) there is homozygosity for the fusion ;.
(iv), diakinesis in the heterozygous individual ; (v), (vi), early anaphase showing co-orientation.

DISCUSSION OF CHROMOSOME NUMBERS
1. EURYMELIDAE

The Eurymelidae comprise approximately one hundred species. Although
the twenty-seven species examined constitute only a small sample of the family, |
they are representative, including nine of eleven genera of the EHurymelinae,,
six of the fourteen genera of the Ipoinae, and one of the genera of the
Pogonoscopinae.

With two exceptions, chromosome numbers are constant in the family, and
it may be argued that the haploid number of 10 autosomes and an X chromosome:
is basic and primitive in the family and the two deviant species, viz. Aloewrymela
gearyt (9+X) and Opio multistrigia (6+ X) are derived forms.

It is interesting to note that the monotypic genus Opio is distinctive
morphologically and ecologically from the remaining genera and it is probable
that Casuarina has served as its food plant for a considerable period while most
representatives of the Eurymelidae are associated with Hucalyptus (Evans, 1959,
and personal communication).

The low chromosome number in Opio probably indicates a reduction in
recombination index, and, in consequence, an increased genetic stability. The.

M. J. WHITTEN 83

Stability of its environment (i.e. of its host, Casuarina) may well have offered the
conditions favouring such a reduction in recombination index. A comparison
of karyotypes (Fig. 3, a, b) suggests the reduction was accomplished by four
independent fusions involving two chromosomes to give two large and four
small bivalents at metaphase 1. A simple fusion would explain the karyotype
of A. gearyi although it is not obvious which chromosomes may have been
involved (Fig. 3, b, ce).

Fig. 3. Karyotype of (a) Opio multistrigia showmg six bivalents and an X chromosome ;
(6) Eurymela fenestrata, with ten bivalents and an X chromosome; (c) Aloewrymela gearyt, with
nine bivalents and an X chromosome.

2. CICADELLIDAE

The Cicadellidae comprise approximately ten thousand species and, of
these, some two hundred have been examined cytologically (Halkka, 1959,
1960b). Consequently it is both difficult and dangerous to draw conclusions
from the present data. Nevertheless several trends can be recognized.

Some primitive cicadellids (e.g. some Ulopinae) feed on moss and reeds
(Evans, 1947): those groups represented in Australia which are of later
evolutionary development are predominantly arboreal (e.g. Jassinae, Tartessinae,
Austroagalloidinae) while, according to Evans, ‘ those leafhoppers of most
recent development feed on grasses and herbaceous plants though not limited
to these plants”’. This advanced group (e.g. the Deltocephalinae) are adapted
to a broad range of environmental conditions and consequently are more widely
distributed than the more primitive cicadellids which tend to be relict in their
distribution.

Halkka (1959) had noted that the chromosome numbers of the Delto-
cephalinae are more varied than those of the ‘ primitive” groups. This
variability is understandable in the light of the morphological and zoogeographical
evidence. He has further noted that the chromosome numbers are lower in the
Deltocephalinae and has concluded that recent evolution has been associated
with a reduction in chromosome numbers. In fact, this finding forms the basis
for most of his phylogenetic considerations.

The present results (Table 1) support Halkka’s hypothesis. The Delto-
cephalinae have a lower mean chromosome number and their range (3-++X to
9+X) is larger than other groups. Opio multistrigia and A. gearyi in the
Eurymelidae, as previously mentioned, are further examples of derived forms
with reduced chromosome numbers.

The reduction in chromosome number is undoubtedly due to fusion of
chromosomes. Hither the fusion of the X chromosome to an autosome giving
a Neo-XY sex mechanism, or simply the fusion of two autosomes, is possible.
Halkka (1959) cites several examples of sex chromosome-autosome fusion while
Taslopa montana (Table 1) provides another. An example of autosome fusion is
found in D. longuinquus where, of eight individuals examined, six were 4-+X,
one was 3+ X and one a “hybrid” (Fig. 2).

Variation of chromosome number within the same population of a species
with localized centromeres should normally lead to the elimination of one or other
type or else lead to the establishment of two distinct races because of the increased

84 CHROMOSOME NUMBERS IN AUSTRALIAN LEAFHOPPERS

incidence of meiotic non-disjunction. Species with diffuse centromeres would
not encounter this difficulty when different chromosome numbers are present.
White (1957a, 1957b) has shown the occurrence of individuals heterozygous for
a broken/fused chromosome is extremely rare in the grasshopper Moraba seurra
even though the two races, which have different chromosome numbers, are
contiguous for many miles. Selection must be relatively strong against such
individuals although it would appear non-disjunction may not be responsible
for the reduced fitness in this case. White (1956) cites several other instances
(e.g. the mantid Ameles heldreichi, the grasshopper Trimerotropis sparsa, and the
mollusc Purpura lapillus) where broken/fused heterozygotes occur in natural
populations, but these examples must be rather exceptional.

Thus it may well be that, having diffuse centromeres, chromosome number
in leafhoppers is a more adaptable character than it is in other organisms and
hence is more subject to variation associated with environmental requirements.
Nevertheless it is still of some significance in phylogenetic considerations. An
examination of a large sample of the remaining 9,800 or more species will no doubt
shed light on the problem.

SUMMARY

1. Chromosome counts have been recorded for 64 species of Homoptera
Auchenorrhyncha.

2. An examination of meiosis in the group reveals that the bivalents are
co-orientated. In particular the behaviour of chromosomes heterozygous for a
fusion supports this conclusion.

3. 10+X is suggested as the base number for the Hurymelidae.

4, The evidence presented supports Halkka’s thesis that the more advanced
leafhoppers have lower chromosome numbers and that the numbers are more
varied in these groups.

5. The presence of diffuse centromeres renders chromosome number more
adaptable as a character for selection.

Acknowledgements

The bulk of this work was done during tenure of a Wheat Industry Research
Council Fellowship.

IT am indebted to Dr. W. D. Jackson and Professor S. Smith-White for reading
the draft manuscript and offering invaluable advice.

In particular I wish to thank Dr. J. W. Evans for identifying the material
and for his continued interest and helpful criticism in reading the draft manuscript.

References

Brown, S. W., 1954.—Univ. Calif. (Berkeley) Publ. Botany, 27: 231-278.
Evans, J. W., 1947.—A natural classification of leafhoppers, Part 3. Trans. Roy. Ent. Soc.
London, 96, Pt. 6: 105-271.
, 1959.—Evolution in the Homoptera. “The Evolution of Living Organisms.”
(Melbourne University Press.)
HatKKa, O., 1959.—Chromosome studies on the Hemiptera Homoptera Auchenorrhyncha. Ann.
Acad. Sci. Fennicae, A IV, 43: 1-72.
, 1960a.—The structure of bivalents in the Homoptera Auchenorrhyncha. Chromosoma,
II: 245-262.

, 19606.—Chromosome evolution in the Cicadellidae. Hereditas, 46: 581—591.
HuGuHES-SCHRADER, S., 1948.—Cytology of coecids. Advances in Genet., 7: 127-203.
Hucues-ScuraDER, 8., and Ris, H., 1941.—J. Haptl. Zool., 87: 429-456.

Roaves, M., 1961.—‘‘ The Cell’’, III. (Academic Press, N.Y. and London.)
Ris, H., 1942.—J. Haptl. Zool., 90: 267-322.

, 1945.—Biol. Bull., 89: 242-257.

Swarn-Walte, S., 1959.—Cytological evolution im the Australian flora. Cold Spr. Harb. Symp.
Quant. Biol., 24: 273-289.

Waitt, M. J. D., 1954.—“‘ Animal Cytology and Evolution.”” (Cambridge University Press,
Cambridge.)

M. J. WHITTEN 85

Wuitr, M. J. D., 1956——Adaptive chromosomal polymorphism in an Australian grasshopper.
Evolution, 10: 298-313.

, 1957a.— Cytogenetics of the grasshopper Moraba scurra. Aust. J.Zool., 5,3: 285-304.

-and Cuinnicr, L. J., 1957b.—-Cytogenetics of the grasshopper Moraba scurra. Aust. J.

Zool., 5, 3: 338-347.

Explanation of Plate w

(a) Bouquet stage of meiosis in Hurymela fenestrata (Eurymelidae) ; (6, c) Two consecutive
stages of diakinesis in H. fenestrata; (d) Metaphase I in same; (e) Diakinesis in Deltocephalus
taedius: (f) Metaphase II in same; (g) Diakinesis in Aconurominus flavidiventris (Stal) (Cicadel-
lidae); (hk) Diakinesis in Deltocephalus longuinquus showing trivalent.

APPENDIX

In response to a request from Mr. Max Whitten, a new genus and species of
Eurymelidae are described below.

A NEW GENUS AND SPECIES OF EURYMELIDAE
(Homoptera, Cicadelloidea)

J. W. EVANS
Australian Museum

EURYMELINAE
ALOEURYMELA, gen. nov.

On the face of the head the labium terminates between the middle coxae
and the anterior margin of the ante-clypeus is depressed below the rest of the
sclerite. The crown of the head is only slightly wider against the eyes than in
the centre. The tegmen has a well developed appendix. The hind tibiae have
one spur and a few additional small spines. The male genitalia have oval sub-
genital plates bearing terminal hook-like styles arising from the ventral margins.

Type species.—Aloeurymela gearyi, sp. nov.

In coloration and general appearance Aloeurymela resembles genera
comprised in the Ipoinae rather than those in the Eurymelinae. It is included
in the last-named subfamily because of the characters furnished by the male
genitalia, in particular the presence of a well developed ventral accessory clasping
process associated with the sub-genital plates.

ALOEURYMELA GEARYI, sp. nov.
(Fig. 4)

Length, g, 9 4-83 mm. General appearance long and narrow, sometimes
with a characteristic diamond-shaped marking on the folded tegmina. Face of
head pale apricot, or dark brown, mottled with yellow; lora and maxillary
plates pale brown. Crown and pronotum pale or dark brown, or black, mottled
with pale brown or greyish-white. Scutellum concolorous with the pronotum

86 CHROMOSOME NUMBERS IN AUSTRALIAN LEAFHOPPERS

but a darker shade. Tegmen basally concolorous with the head and thorax with
two irregular transverse whitish fasciae, which may be confluent in the costal
area. Male genitalia as in Fig. 4 (the aedeagus may have an additional spine

to the one shown in the figure).

Fig. 4. Aloewrymela gearyi. 1, subgenital plate and paramere; 2, aedeagus.

Holotype 3 and Allotype 9 from Cunnamulla, Queensland (coll. N. Geary,
11/41) in the Australian Museum.
_ Known distribution elsewhere.—Perth (Western Australia); Gilruth, Moo-
looka (Queensland) ; Walgett (New South Wales).

THE DISTRIBUTION OF THE NOTONECTIDAE (HEMIPTERA)
IN SOUTH-EASTERN AUSTRALIA

A. W. SWEENEY
Faculty of Agriculture, University of Sydney

[Read 28th April, 1965]

SYNOPSIS

The distribution and relative abundance of species of Notonectidae found in south-east
Australia are discussed. Fourteen species occur in this region, eleven in the genus Anisops,
two in the genus Hnithares and one m the genus Paranisops.

INTRODUCTION

The Notonectidae (‘‘ back swimmers ”’) are common in freshwater habitats
of Australia, such as rivers, creeks, pools and waterholes. Several authors
have discussed the taxonomy of the species found in Australia, including
Kirkaldy (1897, 1904) and Hungerford (1934, 1940). Hale published several
papers on the Notonectidae (1923, 1924, 1925). He described all the known
Australian species, several of which were new, and established the genus
Paranisops. Brooks (1951) revised the genus Anisops on a world basis and
described twelve new species from Australia. He solved many taxonomic
problems associated with this genus.

There are eight genera in the Notonectidae, five of which have been recorded
from Australia. Australian species of Nychia and Notonecta have been des-
cribed by Hale (1925) and Kirkaldy (1897), but they were not encountered
during the course of this study. Awnisops, Enithares, and Paranisops are the
only genera known to occur in south-eastern Australia.

The distribution of the Australian Notonectidae is not well known and
this paper describes the known distribution of species found in south-eastern
Australia.

MATERIALS AND METHODS

This study is based on specimens collected from more than 120 localities
by the author and colleagues, during 1962 and 1963. The collection has since
been deposited in the Macleay Museum, University of Sydney. The area
visited includes most of New South Wales and Victoria, with some localities
in the eastern part of South Australia, and Brisbane, Queensland. The
Notonectid collections of the Australian and South Australian Museums were
inspected but were not examined in detail.

Most specimens were preserved in 70% alcohol although representatives
of each species were pinned. The technique adopted for the examination of
the male foreleg of Anisops was that described by Brooks (1951). After
removal from the insect the organs were cleared in 5% caustic potash solution
and dehydrated in alcohol. Except where otherwise stated, specimens from
the various localities mentioned in the text were collected by the author.

PROCEEDINGS OF THE LINNEAN Society or New SoutH Watss, Vol. 90, Part 1

88 DISTRIBUTION OF NOTONECTIDAE IN SOUTH-EASTERN AUSTRALIA

Genus ANISOPS

Anisops is the dominant Notonectid genus in Australia and the following
twenty-two species have been recorded from this continent :

malkint Brooks *
nasuta Fieber *
nodulata Brooks *
occipitalis Breddin *
ocularis Hale *
paracrinita Brooks *
semita Brooks *

stala Kirkaldy +
tasmaniaensis Brooks t
thienemanni Lundblad +
windt Brooks *

A. barrenensis Brooks *
A. calcaratus Hale +
canaliculata Brooks + *
deanet Brooks +
doris Kirkaldy +
elstont Brooks F
endymion Kirkaldy
evanst Brooks t
gratus Hale +
hackert Brooks +
hyperion Kirkaldy +

i
De fee fa fe a he

* North Australian species
+ South-east Australian species
{i Tasmanian species

Many of these species have been collected only in the far north, in the
Cape York and Darwin areas. Ten of them were collected in the south-east
of the continent during the present study, as well as A. tahitiensis which has
not previously been recorded from Australia.

Only the males of the genus have reliable diagnostic characters. They
can be distinguished from the females by the single tarsal segment of the
foreleg and the lateral prongs on the third segment of the rostrum. A key to
the genus Anisops was presented in Brooks’ (1951) paper, but as this includes.
almost eighty species it was felt that a simplified key of the local species would.
be of some value.

Key to males of south-east Australian Anisops

1. Small species, less than 5 mm. in length .................:...-5------+e--eeeees 2
Larger species, greater than 5 mm. in length ............................----.. 3
2. (1) Facial tubercle * with a median groove, without median depression of pronotum
a ait cae eee el ROR Cea Ne eRe cio SRE INOS A eiereys OrMno 6 oar: one A. canaliculata
Facial tubercle without median groove, with median depression of pronotum A. elstoni

3. (1) Greatest width of head slightly greater than width of pronotum .......... A. doris
Greatest width of head less than width of pronotum .......................... 4
4. (3) Large species greater than 8 mm. in length ............................4.-.-. 5
Medium-sized species less than 8 mm. in length .............................. 6
5. (4) Vertex + extends beyond eyes into a cephalic horn, without spur on tibia of foreleg
RS Cais Ohne oat oie cote a SOT DIRE. Sle PRAT Ra o a eR OAc ao ed Gg 8 A. stalv
Vertex not extended beyond eyes, with spur on tibia of foreleg ...... A. calcaratus
6. (4) Dorsal surface of pronotum with a median depression ................-.. A. gratus
Dorsal surface of pronotum without median depression ........................ a
7. (6) Facial tubercle raised mto a median carina ........................- A. tahitiensis

Facial tubercle not raised into a median carina ......................+++-0---
8. (7) Front femur broad ; dorsal and ventral margins almost parallel for basal three-fourths

Co) Maat tsp Cera Ve rl cee erarsuaoencclsrca cic tus. ceenO Lic eeaane Go ty old, obs ojo NOD OS GU Erba tro A. thienemanni
Dorsal and ventral margins of front femur not parallel ........................
9. (8) Apex of third rostral segment wider than base of fourth segment ........ A. deanet
Apex of third rostral segment equal to base of fourth segment ................ 10
10. (9) Anterior tarsus with a median row of five setae .................... A. hyperion
Anterior tarsus with more than five setae in two rows ................ A. hacker

* Facial Tubercle: The modified region of the space between the eyes, immediately
above the labrum.
+ Vertex : The anterior dorsal margin of the space between the eyes.

ANISOPS THIENEMANNI Lundblad
(Fig. 1a)

This is the most common and widespread species in south-east Australia and is
most abundant on the western plains of New South Wales. It also occurs
on the slopes and tablelands of the Great Dividing Range but is rare on the
east coast, having been collected from only two localities in that area (Morwell
and Narooma).

A. W. SWEENEY 89

Localities :-—NEwW SouTH WALES: Hartley, 5/8/62; Narooma, 30/12/62 ;
Goulburn, 26/12/62; Lake Bathurst (W. Williams), Jan. 1962; Canberra,
A.C.T., 26/12/62 and 20/1/63; Gunning, 20/8/62; Taralga, 4/8/62 ; Cooma,
26/12/62 ; Thredbo, Crackenback River, 28/12/62 ; Glanmire, 11/8/62 ; Koora-
watha, 16/1/63 ; Wellington, 28/8/62 ; Forbes, 26/4/62 ; Young, 16/1/63 ;
Brungenbrong, 19/1/63 ; Gulargambone (J. Anderson), 26/12/62 ; Coonabarabran
(J. Bishop), 28/6/62 ; Warren (P. Bailey), 30/6/62 and (J. Bishop), 12/8/62 ;
Wagga Wagga, 16/1/63 ; Yerong Creek, 16/1/63; West Wyalong, 25/4/62 ;

Fig. 1. Distribution of a, A. thienemanni; b, A. deanei; c, A. hyperion; d, A. stali.

Leeton, 23/4/62; Yenda, 22/4/62; Lake Cargelligo (J. Bishop), June, 1962 ;
Cobar, 27/8/62 ; Nyngan, 1/7/62; Bourke (J. Bishop), 5/7/62; Wilcannia,
Caltigena Tank, 27/8/62 ; Menindee, 26/8/62; Euston, 22/8/62. VICTORIA :
Sale, 18/1/63 ; Morwell, 18/1/63 ; Nareil, 19/1/63 ; Benambra, 19/1/63; Mt.
Hotham (W. Williams), Jan. 1962 ; Violet Town, 16/1/63 ; Winton, 16/1/63 ;
Wunenhu, 21/8/62; Echuca, 21/8/62; Lake Cooper (W. Williams), Jan. 1962 ;
Nyah, 22/8/62; Hattah Lakes (W. Williams), 27/8/62 ; Lake Hindmarsh (Ww.
Williams), Jan. 1962 ; Casterton (W. Williams), 19/8/61 ; Hamilton (W. Williams),
Jan. 1962. SoutH AvsTraLiA: Mingary, 28/8/62; Oodla Wirra, 25/8/62 ;
Truo, 23/8/62 ; Sheoak Log, 23/8/62.

90 DISTRIBUTION OF NOTONECTIDAE IN SOUTH-EASTERN AUSTRALIA

ANISOPS DEANE! Brooks
(Fig. 1b)

A common species, particularly on the coast. It is often found on the
Slopes and tablelands but rarely occurs on the inland plains.

Localities :—NEw SoutH WALES: Bulahdelah, 22/8/62 ; Raymond Terrace,
22/8/62; Marsden Park, 24/6/62; Waterfall, 5/5/62; Narooma, 30/12/62 ;
Bemboka, 29/12/62 ; Milton, 30/12/62 ; Nimmitabel, 29/12/62 ; Kanangra Walls,
20/1/63 ; Goulburn, 26/12/62; Collector, 20/1/63 ; Gunning, 20/8/62 ; Cooma,
26/12/62 ; Berridale, 19/1/63 ; Jindabyne, Saw Pit Creek, 27/12/62 ; Glanmire,
11/8/62 ; Wellington, 28/8/62 ; Cootamundra, 16/1/63 ; Forbes, 26/4/62 ; Gular-
gambone (J. Anderson), 24/4/62; Gilgandra, Castlereagh River (J. Bishop),
29/6/62; Trangie, 27/8/62; Warren (P. Bailey), 30/6/62; Young, 16/1/63 ;
Yerong Creek, 16/1/63; Mullengandra, 20/8/62; Nyngan, 1/7/62; Hermidale,
27/8/62 ; Leeton, 23/4/62. Vicroria: Bairnsdale, 18/1/63; Nareil, 19/1/63 ;
Chiltern, 16/1/63 ; Morwell, 18/1/63 ; Heyfield (W. Williams), 25/11/62 ; Violet
Town, 16/1/63. SourH AUSTRALIA: Oodla Wirra, 25/8/62; Burra, 25/8/62 ;
Sheoak Log, 23/8/62. QUEENSLAND: Brisbane, 27/5/62; Petrie, 31/5/62.

ANISOPS HYPERION Kirkaldy
(Fig. 1e)

The distribution of A. hyperion is similar to that of A. thienemanni. It is
rare on the coast but common on the slopes and ranges and also occurs on the
western plains, though not as frequently as the latter species.

Localities :—NEw SoutH WALES: Mittagong, 3/8/62 and 20/8/62 ; Kanan-
gra Walls, 20/1/63 ; Goulburn, 26/12/62 ; Gunning, 20/8/62 ; Collector, 20/1/63 ;
Canberra, A.C.T., 20/1/63; Glanmire, 11/8/62; Bathurst, 28/8/62; Manildra
(J. Bishop), June 1962 ; Forbes, 26/4/62 ; Trangie, 27/8/62 ; Warren (J. Bishop),
12/8/62; West Wyalong, 25/4/62; Yerong Creek, 16/1/62; Mullengandra,
20/8/62 ; Bourke (J. Bishop), 5/7/62 ; Cobar, 27/8/62 ; Mount Hope (J. Bishop),
June, 1962; Wilcannia, 27/8/62. VicTorIA: Bairnsdale, 18/1/63 ; Benambra,
19/1/63 ; Chiltern, 16/1/63 ; Violet Town, 16/1/63 ; Winton, 16/1/63 ; Glenrowan,
16/1/63 ; Yarrawonga, 21/8/62; Wungnhu, 21/8/62; Echuca, 21/8/62. SouTH
AUSTRALIA: Oodla Wirra, 25/8/62; Sheoak Log, 23/8/62.

ANISOPS STALI Kirkaldy
(Fig. 1d)

A. stali is most common on the inland plains of New South Wales and similar
areas of the surrounding States. It has been collected near the coast (Wingham
and Bemboka) and on the western slopes of the Dividing Range. Altitude may
be a factor affecting the distribution of this species as it has not been found in
the higher areas of this region.

Localities :—NEw SoutH WALES: Wingham, Jan., 1962; Bemboka,
29/12/62 ; Cudgegong River (J. Bishop), 5/11/62 ; Cootamundra, 16/1/63 ; Junee,
16/1/63 ; Gulargambone (J. Anderson), 24/4/62; Wellington, 28/8/62 ; Coona-
barabran (J. Bishop), 28/6/62 ; Trangie, 27/8/62; Warren (P. Bailey), 30/6/62
and (J. Bishop), 12/8/62; Lake Cargelligo (J. Bishop), June, 1962 ; Hermidale,
27/8/62; Mount Hope (J. Bishop), June, 1962; Cobar, 27/8/62 ; Wilcannia,
26/8/62; Menindee, 26/8/62; Euston, 22/8/62. Victoria: Chiltern, 16/1/63.
SoutH AUSTRALIA: Oodla Wirra, 25/8/62.

ANISOPS CALCARATUS Hale
(Fig. 2a)
This species is uncommon and is found mostly in the arid inland areas.
It appears to have a distribution similar to A. stalv.
Localities :—NEw South WALES: Canberra, A.C.T., 20/1/63; Manildra
(J. Bishop), June, 1962; Wellington, 28/8/62; Coonabarabran (J. Bishop),

A. W. SWEENEY 91

28/6/62 ; Cootamundra, 16/1/63 ; Junee, 16/1/63 ; Trangie, 27/8/62 ; Hermidale,
27/8/62 ; Cobar, 27/8/62 ; Menindee, 26/8/62. VicToRIA: Bairnsdale, 18/1/63 ;
Chiltern, 16/1/63. SouTH AUSTRALIA: Oodla Wirra, 25/8/62.

ANISOPS ELSTONI Brooks
(Fig. 2b)

This is a comparatively rare species. It has been collected from several.
localities near Sydney but does not occur far inland. (The most westerly
collection was made at Narrandera, N.S.W.)

Localities :—NEW SoutH WALES: Badgery’s Creek, 7/8/62 ; Marsden Park,
1/7/62; Raymond Terrace, 22/8/62; Narooma, 30/12/62; Valley Heights,
5/8/62; Mittagong, 3/8/62; Narrandera, 25/4/62. VicToRIA: Benambra,
19/1/63 ; Winton, 16/1/63.

Fig. 2. Distribution of a, A. calcaratus; b, A. elstoni; c, A. gratus; d, A. hackerv.

ANISOPS GRATUS Hale
(Fig. 2c)
This uncommon species occurs throughout the inland plains. It has not
been collected on the coast or tablelands.
Localities :—Nrw SoutH WatEs: Yerong Creek, 16/1/63 ; Leeton, 23/4/62 ;
Nyngan, 1/7/62 ; Wileannia, 26/8/62. Vicrorta: Nareil, 19/1/63 ; Yarrawonga,
21/8/62. SoutH AUSTRALIA: Oodla Wirra, 25/8/62; Burra, 25/8/62.

92 DISTRIBUTION OF NOTONECTIDAE IN SOUTH-EASTERN AUSTRALIA

ANISOPS HACKERI Brooks
(Fig. 2d)

This species is uncommon in south-eastern Australia but it is the dominant
Notonectid of the Brisbane area. Its distribution seems confined mainly to
the north-east of the region.

Localities :—NEW SOUTH WALES: Bulahdelah, 22/8/62; Raymond Terrace,
22/8/62; Kanangra Walls, 20/1/63; Forbes, 26/4/62; Gulargambone (J.
Anderson), 24/4/62. QUEENSLAND: East Ithaca Creek, Brisbane, 24/5/62 ;
Upper Brookfield, Brisbane, 30/5/62; Toowong Creek, Brisbane, 26/5/62 ;
Ferny Grove, 27/5/62; Petrie, 31/5/62.

ANISOPS DORIS Kirkaldy
(Fig. 3a)

This is a rare species with distribution similar to A. hackerit. It is common
in Hawkesbury Sandstone creeks in the Sydney area where it is often found
in association with P. inconsians.

Localities :—NEw SoutTH WALES: Booral, 22/8/62; Waterfall, 5/5/62 ;
Appin, March, 1962; Picton Lakes, 7/10/62; Manilla (J. Bishop), 25/6/62 ;
Cudgegong, 5/11/62; Warren (P. Bailey), 30/6/62. QUEENSLAND: Upper
Brookfield, Brisbane, 30/5/62.

ANISOPS CANALICULATA Brooks
(Fig. 3a)
The type locality of this species is Barron River, North Queensland. It
has only been collected by the author at Petrie, Queensland, and may be a
‘“northern ” species whose distribution extends as far south as this locality.

Localities :—QUEENSLAND : Petrie, 31/5/62.

ANISOPS TAHITIENSIS Lundblad
(Fig. 3a)

This species has been found in New Guinea, the New Hebrides and the
Solomon Islands (Brooks, 1951; Lansbury, 1963) but has not previously been
recorded from Australia. It is common in the Brisbane area and probably
occurs in other localities in Queensland and northern New South Wales.

Localities :—QUEENSLAND : East Ithaca Creek, Brisbane, 24/5/62 ; Toowong
Creek, Brisbane, 26/5/62; Ferny Grove, 27/5/62 .

Genus ENITHARES

More than 40 species in this genus are distributed through Africa and
southern Asia (Hungerford, 1956). Two species have been recorded from this
country, both of which occur in south-eastern Australia.

Key to Australian Emthares

1. Vertex not extended markedly beyond the eyes in both sexes, narrow ridge of ventral
abdominal keel, males without spur on anterior trochanter ............ E.. bergrotht
2. (1) Vertex extended markedly beyond eyes in both sexes, broad ridge of ventral abdominal
keel, males with spur on anterior trochanter .......................... E.. hackert

ENITHARES BERGROTHI Montandon
(Fig. 3b)

This is a common species with a distribution similar to A. deanei. It is
found on the east coast as well as the tablelands and slopes but is rare on the
western plains.

Localities :—NEw SoutH WALES: Raymond Terrace, 22/8/62; Waterfall,
5/5/62 ; Valley Heights, 5/8/62 ; Manilla, 25/6/62 ; Richlands, 4/8/62 ; Taralga,
4/8/62; Canberra, A.C.T., 20/1/63; Glanmire, 11/8/62; Cudgegong, 5/11/62 ;

A. W. SWEENEY 93

Cowra, 16/1/63 ; Gulargambone (J. Anderson), 24/4/62 ; Mullengandra, 20/8/62 ;
Wagga Wagga, 16/1/63. Victoria: Tambo River (W. Williams), 23/2/62 ;
Morwell, 18/1/63 ; Chiltern, 16/1/63 ; Winton, 16/1/63 ; Wangaratta, 16/1/63 ;
Dalyston (W. Williams), 2/9/62 ; Tallarook, 17/1/63 ; Kul Kyne (W. Williams),
27/8/61. SoutTH AUSTRALIA : Sheoak Log, 23/8/62. QUEENSLAND : Hast Ithaca
Creek, Brisbane, 24/5/62 ; Toowong Creek, Brisbane, 26/5/62.

ENITHARES HACKERI Hungerford
(Fig. 3b)
This species is rare in south-eastern Australia and was only collected in
three localities during this study. The type locality is Brisbane, Queensland.
Localities :—NEwW SoutTH WALES: Manilla, 25/6/62; Leeton, 23/4/62.
VicTORIA : Wangaratta, 16/1/63.

Fig. 3. Distribution of a, @ A. doris, © A. tahitiensis, @ A. canaliculata ; b, @ H. bergrothi,
O HE. hackerit, © P. inconstans.

Genus PARANISOPS

This genus was established by Hale (1924) from specimens obtained at
Epping, N.S.W. There is only one known species, P. inconstans, which is of
special interest as it occurs in two distinct morphological forms. There is a
black winged variety as well as a pale form which lacks functional hindwings.
This species has only been found near Sydney (type locality) and Brisbane
(Hungerford, 1934). Only the pale form, which is quite common in Hawkes-
bury Sandstone creeks around Sydney, was collected during this study.

PARANISOPS INCONSTANS Hale
(Fig. 3b)
Localities :—NEw SoutH WALES: Waterfall, 5/5/62 ; Appin, March, 1962 ;
Heathcote, March, 1962.

DISCUSSION

Most species of Notonectidae found in south-eastern Australia are more
common within restricted areas of this region. Several species (A. thienemanni,
A. hyperion, A. stali, A. calcaratus and A. gratus) are mor2 prevalent on the
inland plains and the slopes of the Great Dividing Range. Others (A. deanei,
A. elstoni, A. hackeri, A. doris and E. bergrothi) have more easterly distributions
and are usually found near the coast. A. hackeri and A. doris seem confined
to the north-east.

Suitable aquatic habitats in inland areas usually consist of man-made
dams and waterholes which are often isolated and many miles apart. The
species found in these areas may have better dispersal powers than the ‘‘ coastal ”’

94 DISTRIBUTION OF NOTONECTIDAE IN SOUTH-EASTERN AUSTRALIA

species and thus be better adapted to invade an arid environment where habitats
are witlely separated. The other species may be limited to the coast and ranges
where the heavier rainfall ensures an adequate supply of freshwater habitats.

Several species are often found together in the same locality. This is
partictlarly so im western areas, where the scarcity of habitats may be
responsible for this gregariousness.

There are several species of Anisops with type localities in North Queens-
land. A. canaliculata is the only one of these which has been found outside
this area and is probably a ‘“‘ northern ” species whose distribution range extends
farther south than the others.

P. inconstans commonly occurs in Hawkesbury Sandstone creeks and seems
rigidly restricted to this habitat as it has not been found elsewhere in the region.
This limited distribution may be explained by the predominance of the pale
form which is incapable of flight.

Acknowledgements

I wish to acknowledge the generous assistance of Dr. A. R. Woodhill,
Reader in Entomology, University of Sydney, and Mr. J. Bishop, Zoology
Department, University of Sydney, who encouraged this work in many ways.
I am indebted to the following people who collected specimens for this study :
Dr. W. Williams, Zoology Department, Monash University, Mrs. J. Anderson,
Macleay Museum, and Mr. P. Bailey, C.8.I.R.O. Division of Wild Life Research.
I wish to thank Dr. I. Lansbury of Hope Department of Entomology, Oxford
University Museum, for the gift of a specimen of A. tahitiensis.

References

Brooks, G. T., 1951.—A revision of the genus Anisops (Notonectidae-Hempitera). Univ.
Kansas Sci. Bull., 34: 301-519.
Hats, H. M., 1923.—Studies m Australian aquatic Hemiptera. Rec. S. Aust. Mus., 2: 397-424.
, 1924.—Two new Hemiptera from New South Wales. Proc. Linn. Soc. N.S.W..,.
xlix: 461-467.
, 1925.—The aquatic and semi-aquatic Hempitera. Arch. Zool., 17: 1-19.
Huncerrorp, H. B., 1934.—Concerning some aquatic and semi-aquatic Hemiptera from
Australia. Bull. Brooklyn Ent. Soc., 29: 68.
, 1940.—A new HEnithares from Australia (Notonectidae-Hemiptera). J. Kansas Ent.
Soc., 138: 130-131.
, 1956.—Some interesting aspects of the World distribution and classification of
aquatic and semi-aquatic Hemiptera. Proc. Tenth Int. Con. Ent., 1: 337-348.
KirKatpy, G. W., 1897.—Revision of the Notonectidae, No. 1. Trans. Ent. Soc., 1897, 393-426.
, 1904.—Uber Notonectiden. Wien. Hnt. Zeit., 22: 93-135.
Lawnsspury, I., 1963.—Notes on the water-bugs of the Solomon Islands and New Hebrides.
Pacific Insects, 5(1): 5-10.

Author’s present address: Malaria Institute, P.H.D., Rabaul, T.P.N.G.

THE HISTOLOGY AND ANATOMY OF THE REPRODUCTIVE
SYSTEM OF THE LITTORAL GASTROPOD BEMBICIUM NANUM
(LAMARCK) (FAM. LITTORINIDAE)

LYNNE BEDFORD
University of Sydney

[Read 28th April, 1965.]

Synopsis
In Bembicium nanum the sexes are separate and easily distinguishable ; the female by the
yellow ovipositor, the male by the conspicuous penis.

The histology and anatomy of the male reproductive system of B. nanum, apart from the
shape of the penis, the absence of penial glands, the compact nature of the testis and the germinal
epithelium of the testis, are similar in general to other littormids. However, in the female
reproductive system, greater differences are found ; the renal oviduct is lined by a phagocytic
syncytium, the receptaculum seminis is modified for storage, nourishment and phagocytosis of
sperm, and the bursa copulatrix functions only as an organ for reception of spermatozoa.

Time of spawnmg appears to be mdependent of the time of year and different for each
individual. Egg masses are slightly more numerous in the sprmg and summer months. Large
seasonal differences in the size of the reproductive system and extensive resorption of gametes
do not occur. In B. nanum resorption of spermatozoa is restricted to the vesicula seminalis
of the male system and to the receptaculum seminis of the female system. Slight resorption
of ova appears to occur in the renal oviduct of the female reproductive system of B. nanum.
This limitation of phagocytosis of gametes in B. nanum, in comparison with North Sea littorinids,
may be related to the milder climatic variations in the Sydney coastal areas.

INTRODUCTION

Little is known of reproduction and development in Australian littorinids
(Anderson, 1960). Apart from a brief and rather inaccurate description by
Kesteven (1902) and a brief reference by Anderson (1958) in a taxonomic survey
of the genus Bembicium, no details of the reproductive system of B. nanum have
yet been described. In the following work, the anatomy and histology of the
reproductive system of B. nanum are described. Differences from North Sea
littorinids are noted and related to continuous breeding throughout the year
in B. nanum.

METHODS

Males and females of B. nanwm were collected at intervals during 1961 and
1962 on the rock platforms of the ocean coast near Sydney. Animals, removed
from their shells and relaxed in fresh water, were dissected under a binocular
microscope. For histological studies, Smith’s formol-bichromate, 5°% formol
saline and Baker’s formaldehyde calcium were found to be the most suitable
fixatives. To prevent hardening, material was taken to 95% alcohol, trans-
ferred to 1% celloidin in methyl benzoate, followed by benzene, then embedded
in paraffin (M.P. 56°C). Sections were cut at 8u and stained in Ehrlich’s
haematoxylin and eosin or Heidenhain’s azan stain.

RESULTS
Male Reproductive System

As in all male prosobranchs, the testis (Fig. 1) in B. nanum lies in the visceral
Spire over the digestive gland, its tubules being grouped around the visceral
arterial system (Anderson, 1958). The wall of each tubule is a flattened
germinal epithelium. Within this lies a dense layer of spermatocytes, then a
layer of spermatids, while mature spermatozoa (Fig. 2) and nurse cells (Fig. 3)
with finely vacuolated cytoplasm and attached spermatids and spermatozoa
occupy the lumen.

PROCEEDINGS OF THE LINNEAN SocrETy oF NEw SoutH Wates, Vol. 90, Part 1

96 REPRODUCTIVE SYSTEM OF BEMBICIUM NANUM

The small tubules unite and open into the coiled vesicula seminalis which
runs along the axial surface of the spire and opens anteriorly into the vas
deferens (Fig. 1).

The vesicula seminalis is about 200u in diameter and is lined by cuboidal,
vacuolated epithelial cells (Fig. 4). Spermatozoa are found in the vacuoles
together with nurse cells, penetrating the epithelium, and in an unorientated
mass in the lumen.

HEAD

BUCCAL MASS PENIS

CTENIDIUM FOOT

MANTLE

Se

KIDNEY

RECTUM

VESICULA SEMINALIS HYPOBRANCHIAL GLAND

PROSTATE

DIGESTIVE GLAND
AS DEFERENS

| A SaaS
TESTIS

Fig. 1. Male reproductive system of B. nanum. The mantle cavity has been opened dorsally and
the kidney has been folded to the right.

The vas deferens (Fig. 5) is short, about 100u in diameter, lined by a
ciliated cuboidal epithelium and surrounded by a thin layer of circular muscle,
followed by a layer of connective tissue. It passes into the connective tissue
under the kidney and continues as the prostate gland on the right side of the
mantle cavity.

The prostrate gland looks like a complete duct in dissected specimens
(Fig. 1), but is composed of an attached right and freely-hanging left lobe,
separated ventrally so that the lumen of the gland opens into the mantle
cavity. Hach lobe has a deep ciliated sperm groove on its inner edge (Fig. 8).

LYNNE BEDFORD 97%

Anteriorly, the prostate becomes a closed tube, the free edges of the two lobes
fusing in the ventral midline.

The columnar epithelium lining the lobes of the prostate gland has two
types of cells, gland cells and supporting cells (Fig. 6). Each gland cell has
a basal nucleus, a single nucleolus and granular cytoplasm containing numerous
eosinophil granules, especially in the narrower distal parts of the cell. The support-
ing cells, regularly placed between the gland cells, are expanded distally and
compressed to thin cytoplasmic strands proximally. The free surfaces of the
cells are densely ciliated, the cilia being longest on the edges of the sperm
groove and in the groove itself. Many mucous cells are found in the epithelium

\ <_ SPERMATOZOON

“

ch

SPERMATIDS

SPERMATOG ONIUM

ERMINAL EPITHELIAL
CELL |

Fig. 2. B. nanum. T.S8. through the wall of a testicular tubule.

of the edge of the left lobe (Fig. 7). No muscles appear under the prostate
epithelium, but circular and longitudinal muscles are found under the mantle
epithelium on the outer edge of the left lobe.

At the mouth of the mantle cavity, the prostate gland continues as the
anterior vas deferens (Fig. 1), a narrow duct 75py in diameter, lined by ciliated
cuboidal epithelium and surrounded by a muscle sheath composed of circular,
longitudinal and transverse muscle (Fig. 9). The anterior vas deferens passes
along the foot to the right of the buccal mass, in a ridge of dense connective
tissue and opens anteriorly into the penis.

The penis (Fig. 1), a conical projection slightly dorso-ventrally flattened,
is covered by a ciliated columnar epithelium. Underlying the latter is a thin
layer of circular muscle, then layers of dorso-ventral and oblique
muscles, while longitudinal muscles run through the connective tissue

@

98 REPRODUCTIVE SYSTEM OF BEMBICIUM NANUM

internally. The connective tissue has numerous blood spaces and _ the
penial nerve lies in a ventral position. The penial duct, which runs through
the penis dorsally, is 200u in diameter and is lined by ciliated columnar cells,
75u high (Fig. 10). A thin coat of circular, lengitudinal and transverse muscles
lies immediately external to this epithelium.

‘.
2

SSs

NURSE CELL

VACUOLE

SPERMATOZOON

CYTOPLASM

NUCLEUS

NUCLEOLUS

A 10
ee

1Op 3
Figs 3,4. B.nanum. 3, Nurse cell with attached spermatozoa ; 4, T.S. through the wall of the
vesicula seminalis.

SUPPORTING CELL
ren aT NUCLEUS
|

iil
{|
|
! | GLAND CELL
| ILIA

CIRCULAR MUSCLE

CONNECTIVE TISSUE

Figs 5,6. B.nanum. 5, T.S. through the wall of the vas deferens; 6, T.S. through the wall of
the prostate gland.

Female Reproductive System
Like the testis, the ovary (Fig. 11) lies over the digestive gland and its
tubules are grouped around the visceral arterial system, this arrangement
being seen in young specimens only (Anderson, 1958). The wall of each tubule
is a flattened germinal epithelium, with developing oocytes projecting into

LYNNE BEDFORD 99

the lumen of the tubule and mature oocytes lying in the lumen (Fig. 12). The
ovarian tubules open into a single duct, the renal oviduct (Fig. 11), which
runs through the connective tissue under the kidney.

The wall of the renal oviduct is composed of a syncytial epithelium with
scattered oval nuclei, each with a single nucleolus, and an external sheath of
circular muscle. Large vacuoles, 10u in diameter, filled with large, yolky,
eosinophil granules, also occur in the epithelium cytoplasm (Fig. 15). Anteriorly,
the renal oviduct dilates, then narrows and opens into the pallial oviduct or
‘uterus’? (Fig. 11). In its narrower region the renal oviduct has a typical
ciliated cuboidal epithelium.

Immediately behind the opening of the renal oviduct into the pallial
oviduct is the opening of the receptaculum seminis (Fig. 11). This is a short
blind duct lying in the mantle cavity between the renal oviduct and pallial
oviduct, in which it is embedded anteriorly. Posteriorly, the receptaculum

f PROSTATE GLAND CELL

: SUPPORTING CELL
Dy, [P é
? Wi ig, C\LiATED SPERM GROOVE

UY ff | (GQ LONG CILIA

UCOUS CELL

MANTLE EPITHELIUM

CIRCULAR MUSCLE LAYER

LONGITUDINAL AND TRANSVERSE
MUSCLE LAYER

CONNECTIVE TISSUE

ui lou
Fig. 7. B. nanum. T.S. through the left lobe of the prostate gland.

seminis ends in a bulb, 300 py in diameter, lined by unciliated, cuboidal epithelial
cells and filled with unorientated spermatozoa (Fig. 17). Anteriorly, it is
laterally compressed, with diameters of about 300u and 150u (Fig. 13). In
this anterior portion, its ventral epithelium, which lines a deep groove, is a
syneytium with large oval nuclei. Numerous spermatozoa are found with
their heads embedded in the syncytium (Fig. 16). Dorsally, the epithelium
is columnar and is composed of three cell types : mucous cells, cells containing
granules and spermatozoa, and ciliated supporting cells (Fig. 14). The mucous
cells each have a granular cytoplasm staining evenly with haematoxylin and
a basal nucleus containing a single nucleolus. The cells containing granules
and spermatozoa, presumably phagocytically digesting the latter, also have a
basal nucleus containing a single nucleolus, but their cytoplasmic granules
stain strongly with eosin. Neither of these cell types is ciliated. The supporting
cells, between them, are expanded distally and bear cilia about 10u high. A
circular muscle sheath, 10u thick, surrounds the receptaculum seminis.

The pallial oviduct has, running ventrally along its length, a channel with
densely ciliated cuboidal epithelium, 5y high (Fig. 18). Its lateral and dorsal

100 REPRODUCTIVE SYSTEM OF BEMBICIUM NANUM

walls, on the other hand, are enlarged to form a posterior albumen gland and an
anterior jelly gland. The two glands lie to the right of the rectum on the
dorso-lateral wall of the mantle cavity and can be seen through the thin mantle
tissue (Fig. 11). The albumen gland (Fig. 19) is a large mass of much folded
epithelium, the lumen between the epithelial folds connecting with the ventral

TE :
MANTLE ROOF

LEFT LOBE OF
PROSTATE

RIGHT LOBE OF PROSTATE

HYPOBRANCHIAL \

GLAND

SPERM GROOVES ———
CONNECTIVE TISSUE

CS

MANTLE EPITHELIUM

100).
>

Fig. 8. B. nanum. Diagrammatic T.S. through the roof of the mantle cavity showing the
relative positions of the prostate gland, rectum and hypobranchial gland.

rn re

TT KT

CYTOPLASM

NUCLEI

NUCLEOL!

CIRCULAR MUSCLE

LONGITUDINAL AND TRANSVERSE &
MUSCLE

9 Su CONNECTIVE TISSUE ie) lop

Figs 9,10. B.nanum. 9, T.S. through the wall of the anterior vas deferens; 10, T.S. through
the wall of the penial sperm duct.

channel of the oviduct. Both glandular and supporting cells occur in the
epithelium. The densely ciliated supporting cells are each expanded distally
and are connected to the basement membrane by thin cytoplasmic connections
and have a spindle-shaped, densely staining nucleus at the base of the distal
expansion. The albumen gland cells are filled with secretion droplets staining
heavily with haematoxylin, these droplets being more numerous in the distal
parts of the cells. The nucleus of each cell is oval, basal and has a single

LYNNE BEDFORD 101
nucleolus. The epithelium of the albumen gland is covered externally by a
coat of circular muscle, 5y thick.

The jelly gland (Fig. 11) is composed of the two thickened, folded, lateral
walls of the pallial oviduct, with the lumen between them opening into the
ventral channel of the pallial oviduct. The gland cells (Fig. 20) are filled with

OVIPOSITOR

JELLY GLAND

MANTLE RECTUM

CTENIDIUM HYPO BRANCHIAL GLAND

OPENING OF VAGINA

PALLIAL OVIDUCT——_

BUCCAL MASS
BURSA COPULATRIX

ALBUMEN GLAND

KIDNEY

RECEPTACULUM SEMINIS

RENAL OVIDUCT

DIGESTIVE GLAND
<—__rurr.
| | ‘Ov ARY

Fig. 11. Female reproductive system of B. nanum. The mantle cavity has been opened dorsally
and the kidney has been folded to the right.

secretion droplets which stain heavily with haematoxylin. The supporting
cells are similar to those of the albumen gland.

Opening anteriorly into the ventral channel of the pallial oviduct is the
bursa copulatrix, a large blind duct, 300u in diameter, which lies to the right
of the oviduct in the mantle cavity (Fig. 11). Its ciliated columnar epithelium
(Fig. 22) is thrown into folds of varying size (Fig. 21), which are supported by

102 REPRODUCTIVE SYSTEM OF BEMBICIUM NANUM

@i\<-SHED OOCYTE

DEVELOPING OOCYTE
OOGONIUM

l2 < 100M,

Fig. 12. T.S. through the wall of an ovarian tubule of B. nanum.

ciLiaA————1 MA \ uN |

MUCOUS CELL “qa een

CELL CONTAINING
GRANULES AND

SPERMATOZO ON es =

RCULAR MUSCLE
| \4

Figs 13-16. B.nanum. 13, T.S. through the receptaculum seminis ; 14, T.S. through the dorsal
epithelium of the receptaculum seminis ; 15, T.S. through the wall of the renal oviduct ; 16, T.S.
through the ventral syncytium of the receptaculum seminis.

LYNNE BEDFORD 103

extensions of muscle from a surrounding muscular sheath. Unorientated
spermatozoa are found in the lumen of the bursa copulatrix.

At the mouth of the mantle cavity the bursa and oviduct unite as a short
vagina (Fig. 11). From this, a ciliated groove runs between two lobes on the
foot, to the right of the buccal mass and head. These lobes comprise the

CILIA

SUPPORTING CELL

GLAND CELL

Figs 17-22. B. nanum. 17, T.S. through the wall of the posterior bulb of the receptaculum
seminis ; 18, T.S. through the wall of the ventral channel of the pallial oviduct ; 19, T.S. through
the wall of the albumen gland; 20, T.S. through the wall of the jelly gland; 21, diagrammatic
T.S. through the wall of the bursa copulatrix ; 22, T.S. through the wall of the bursa copulatrix.

Ovipositor. Hach lobe has a ciliated epithelium, 50u high, in which many
mucous cells are found. The epithelium at the edge of the groove is similar,
but about 20u high. Beneath the epithelium of the ovipositor is a ridge of
circular and transverse muscle.

Numerous large, granule-filled cells were recorded in the tissues of the
reproductive system of Littorina rudis by Linke (1933), who supposed them

104 REPRODUCTIVE SYSTEM OF BEMBICIUM NANUM

to be amoebocytes of excretory function. Similar cells were seen in the tissues
of B. nanum, especially in the epithelium of the albumen gland, jelly gland,
ventral channel of the pallial oviduct and bursa copulatrix.

DISCUSSION
Male Reproductive System

The histology and anatomy of the reproductive system of B. nanum are
similar in general to those of Littorina littorea, L. obtusa, L. rudis and Cremno-
conchus syhadrensis, described by Linke (1933, 1935). Anderson (1958), how-
ever, has already noted the following differences between the male reproductive
systems in Littorina and Bembicium: the shape of the penis, the absence of
penial glands and the compact nature of the testis, which lies over the digestive
gland in Bembiciwm. Anderson’s results are confirmed here and further
differences have also been noted. The germinal epithelium in the testis of
B. nanum is a flattened epithelium, not the syncytium described by Linke in
other littorinids. Linke also described seasonal phagocytosis of spermatozoa
in the testes of littorinids. No seasonal phagocytosis of spermatozoa was
observed in the testis of B. nanum.

As in ZL. littorea, L. obtusa, and LD. rudis (Linke, 1933) and in Ocinebra
erinacea, Nucella lapillus, Nassarius reticulatus and Buccinium undatum (Fretter,
1941), phagocytosis of spermatozoa and degenerating nurse cells occurs in the
vesicula seminalis of B. nanum. The muscular vas deferens apparently acts
as a sphincter, regulating flow of spermatozoa from the vesicula seminalis into
the prostate.

Kesteven (1902), who briefly described the male reproductive system of
B. nanum, made no reference to the prostate, referring to it as a closed vas
deferens. In fact, the prostate is open and glandular. Spermatozoa move
along the ciliated groove of the prostate gland and receive the prostate secretion
before moving into the closed penial sperm duct.

Female Reproductive System

Kesteven’s (1902) description of the anatomy of the female reproductive
system of B. nanum contained a number of errors, namely, his attribution of
reduction in overall size of the system to non-breeding periods, his description
of the ‘ uterus” and his omission of the receptaculum seminis and bursa
copulatrix.

As in L. obtusa and L. rudis (Pelseneer, 1911 ; Linke, 1933), sexual ripeness
is independent of the time of year and is different for each individual in B.
nanum. Absence of seasonal phagocytosis of eggs in the ovary of B. nanum
(cf. other littorinids, Linke, 1933) may possibly be due to the milder climatic
variations in the Sydney coastal area, as compared with those in the North Sea
coastal areas. Kesteven (1902) noted a marked overall reduction in the size
of the female reproductive system, especially in the ovary, but no such reduction
was noted in this examination of B. nanwm. His diagram of the female system
appears to be of an immature specimen and his observations on size reduction
were probably due to either immaturity or examinarion of parasitized specimens.

No phagocytosis of eggs occurs in the renal oviduct of other littorinids
(Linke, 1933), but Fretter (1941) described it in the ingesting gland of other
prosobranchs. While no ingesting gland is found in B. nanum, phagocytosis
of eggs appears to occur in the renal oviduct.

Using the methods outlined above, the difficulties encountered by Linke
in the histological examination of the ‘‘ uterine’ glands in the pallial oviduct
were overcome. Using these techniques, it was possible to identify the anterior
jelly gland and the posterior albumen gland in the pallial oviduct of B. nanum.
A similar arrangement of glands was found in the pallial oviduct of B. auratum
by Anderson (1958), but she gave no details of histological structure. The

LYNNE BEDFORD 105

enlarged ‘‘ uterus’ recorded by Kesteven in Bb. nanuwm is in fact these glands
of the pallial oviduct. The muscular supporting bands he described are not
muscular tissue, but folds of glandular material (Fig. 11). The glands are
surrounded by a very thin coat of muscular tissue. During secretion of the
albumen gland in Littorina, Linke described extrusion of the upper parts of
the gland cells into the lumen of the gland. Presumably, this was a fixation
artifact as no such extrusions are found in the albumen gland of B. nanum.
The structure of the jelly gland is similar to that described by Linke (1933)
in L. obtusa and L. littorina.

Kesteven (1902) made no reference to the receptaculum seminis and bursa
copulatrix in. B. nanum. The position of the receptaculum seminis and bursa
copulatrix is similar to that recorded by Anderson (1958) in B. auratum. In
Tittorina Linke (1933) described a receptaculum seminis lined by a syncytial
epithelium, in which sperms are embedded. The receptaculum of B. nanum
differs from that of Littorina in a number of ways: in B. nanum the recep-
taculum swells to form a bulb lined by cuboidal epithelium and filled with
unorientated spermatozoa ; the anterior part is lined ventrally by a syncytium
in which spermatozoa are embedded, and dorsally by an epithelium composed
of mucous and ciliated supporting cells, and cells containing granules and
spermatozoa. The latter cells are presumably phagocytic. No similar recep-
taculum seminis has been described in other prosobranchs.

Unlike the bursa copulatrix of Littorina, which is lined by a syncytium
and contains orientated spermatozoa (Linke, 1933), the bursa copulatrix of
B. nanum is lined by a ciliated columnar epithelium, contains unorientated
spermatozoa, and functions only as a receptor organ for spermatozoa.

Acknowledgements

This work was supported by a research grant from the University of Sydney
and was part of a thesis accepted by the University of Sydney for the M.Sc.
degree. The author wishes to thank Dr. D. T. Anderson for his advice and
criticism of the manuscript.

References

AnpeErson, D. T., 1960.—The life histories of marine prosobranch molluses. Journ. Malacol.
Soc. Aust., 4: 16-29.

ANDERSON, H., 1958.—The gastropod genus Bembictum Phillipi. Aust. J. Mar. Freshw. Res.,
9: 546-568.

FrettEer, V., 1941.—The genital ducts of some British stenoglossen prosobranchs. J. Mar.
Biol. Assoc. U.K., 25: 173-211.

Kesteven, H. L., 1902.—Notes on Prosobranchiata, No. 2 Littormacea. Proc. Linn. Soc.
N.S.W., 27: 620-636.

Linke, O., 1933.—Morphologie und Physiologie des Genitalapparates der Nordseelittorinin.
Wiss. "Meeresuntersuch., 19: 1-60.

Linke, O., 1935.—Zur Morphologie und Physiologie des Genitalapparatus der Susswasser-
littorinide Cremnoconchus syhadrensis Blanchford. Arch. Naturgesch., 4: 72-87.

PELSENEER, P., 1911.—Reserches sur l’embryologie des Gastropodes. Bruaelles Mem. Acad.
Roy., Ser. 2, 6: 1-167.

THE REPRODUCTION AND EARLY LIFE HISTORIES OF THE
GASTROPODS NOTOACMAHA PETTERDI (TEN._WOODS),
CHIAZACMAEHA FLAMMEA (QUOY AND GAIMARD) AND
PATELLOIDA ALTICOSTATA (ANGAS) (FAM. ACMAEKIDAR)

D. T. ANDERSON
Uniwersity of Sydney

[Read 28th April, 1965.]

Synopsis

Notoacmaea petterdi, with an externally fertilized egg 150u in diameter, develops into a
planktonic lecithotrophic trochophore in 16 hours, remains planktonic for 30 hours, during which
it transforms into a simple veliger, then settles, alternating crawling with intermittent swimming,
during a period of eight days. Feeding does not begin until settlement is permanent, by which
time metamorphosis is well advanced. Development in Chiazacmaea flammea, with a 130u egg,
and Patelloida alticostata, with a 150u egg, follows a similar course, but in C. flammea permanent
Swimming is maintained for 60 hours and permanent settlement attamed after four more days,
while in P. alticostata swimming for 60 hours is followed by settlement over three more days.
The form and dimensions of the eggs and larvae in these three species resemble those of Acmaea
testudinalis, but associated with a more prolonged subsistence on yolk, C. flammea, P. alticostata
and especially N. petterdi have a more extended swimming-distributive phase than A. testudinalis.

INTRODUCTION

Although several species of acmaeid limpet are commonly represented
along the New South Wales coast (Dakin, 1953), their reproduction and early
life histories have not been investigated (Anderson, 1960). Little is known
of larval development in the Acmaeidae, the only comprehensive description
being that of Kessel (1964) for Acmaea testudinalis. The present study of
Notoacmaea petterdi, Chiazacmaea flammea and Patelloida alticostata shows that
their larval development differs from that of A. testudinalis in a number of
interesting ways.

MATERIALS AND METHODS

For N. petterdi, which inhabits upper littoral vertical rock faces exposed
to the ocean surf, animals collected from the rock platform at Harbord, N.S.W.,
in July and August 1964 and in January 1965 were found to contain ripe
gametes at both periods, suggesting that breeding occurs throughout the year.
For Chiazacmaea flammea, which lives in association with oysters intertidally
in estuarine waters, animals collected from the shores of Middle Harbour,
N.S.W., in August 1964 and January 1965 also contained ripe gametes at both
times, similarly indicating a prolonged breeding season. For Patelloida alti-
costata, which lives at very low levels on intertidal coastal rock platforms, animals
containing ripe gametes were obtained from Long Reef, N.S.W., in January
1965, but have not yet been examined at other times of the year.

Larvae of each species were obtained by artificial fertilization, after
releasing eggs and sperm by dissection of the adults. Eggs were divided into
batches of about 100 and allowed to stand in 300 ml. of filtered sea-water for
30 minutes before adding a few drops of sperm suspension. Swimming
trochophores resulting from successful fertilization were transferred by means
of a pipette to Petri dishes of filtered sea-water, a similar transfer to fresh
sea-water being effected each day until permanent settlement had occurred.
All cultures were maintained at 20°C.

PROCEEDINGS OF THE LINNEAN Sooiety or NEw SourH Watss, Vol. 90, Part 1

D. T. ANDERSON 107

RESULTS
Notoacmaea petterdi

The mature oocytes of NV. petterdi (Fig. 1) are pink in colour, uniformly
yolky and opaque, covered by a thin egg membrane and, after immersion in
sea-water for 30 minutes, spherical, with a diameter of 150u. Sea-water
immersion also causes a thin layer of colourless jelly to swell up on the surface
of the egg, but there is no indication of adhesion between eggs, suggesting
that they float freely in the water following natural spawning. Sixteen hours
after fertilization, free-swimming yolky trochophores (Fig. 2) are found swimming
near the water surface by means of a prototroch of long coarse cilia. The
action of the prototroch is intermittent, the larva either swimming at random
in short darting bursts or drifting with the prototroch folded forwards. The
prototrochal ciliary beat has a clockwise metachronal rhythm but the
larva itself does not rotate while swimming. Behind the prototrochal ring is
a ring of short vibrating cilia, while anteriorly and posteriorly lie apical and
terminal tufts of stationary cilia, held extended while the larva is in motion.
The episphere with its paired antero-lateral protuberances is also covered by
short, slowly waving cilia. Due to the internal mass of yolky macromeres,
the trochophore retains the pink coloration of the egg, but its outer parts are
colourless.

At about 19 hours, the foot rudiment begins to grow out ventrally behind
the prototroch and during the next five hours the trochophore develops into
a simple early veliger (Fig. 3). The prototroch shows little change of form,
but its action becomes more vigorous and continuous, so that long periods of
steady swimming in a semi-upright position near the water surface are inter-
spersed with short resting periods during which the larva gradually sinks through
the water. No unidirectional response to light is observed at this or any other
stage of development. The episphere in the early veliger is already reduced
in size, and has lost its paired protuberances, although its apical tuft and general
ciliation are retained. The hyposphere, in contrast, is enlarged and elaborated,
with a conspicuous, bilobed, ventral foot rudiment and a globular, colourless,
dorsal shell. Due to dorsal enlargement of the hyposphere, the original terminal
tuft of cilia is pushed postero-ventrally and now sprouts:from a small pro-
tuberance behind the foot. The interior of the veliger is still occupied by a
pinkish mass of yolky macromeres.

During the second day of development, the veliger continues to swim in
an upright position near the water surface, with the velar cilia maintaining
their clockwise metachronal beat, rising through the water, then sinking again
at intervals when the velar cilia come to rest. At the same time, the veliger
(Fig. 4) increases greatly in size and shows numerous structural changes. In
the velum, the velar cilia grow longer and retain their vigorous activity, but
the velar cells become much smaller, indicating the onset of a gradual meta-
morphosis. The episphere, although retaining its ciliation, also becomes smaller,
being much flatter at the end of the second day than at the beginning. The
foot, in contrast, increases in size as a triangular wedge incorporating the
terminal tuft and its protuberance at the apex, and secretes an operculum on
its posterior face. The colourless larval shell is greatly enlarged, with the
visceral mass, attached posteriorly to the shell by paired columella muscles,
occupying only part of it, the remainder being occupied by the mantle cavity.
Torsion occurs during the second day, so that the mantle cavity becomes dorsal
in position, but neither withdrawal into the shell nor muscular movements
of the animal are observed during this time. The main mass of yolk is now
concentrated in the visceral mass, but the remainder of the tissues are also
Semi-opaque and not obviously differentiated.

Further progress in development and metamorphosis during the third
day (Fig. 5) is accompanied by a change in behaviour. From continuous
Swimming, the behaviour of the larva alters to long periods of sedentary attach-

108 REPRODUCTION AND EARLY LIFE HISTORIES OF SOME GASTROPODS

ment to the bottom interspersed with brief slow swimming excursions upwards
through the water. Even agitation of the dish in which the larvae are main-
tained fails to alter this pattern. The larval shell grows no larger, but the
velum becomes slightly smaller and its cilia begin to shrink. The episphere

|

5

Figs 1-5. Notoacmaea petterdi. 1, Mature oocyte; 2, trochophore, 16 hr., ventral view ;
3, veliger, 24 hr., ventral view ; 4, veliger, 41 hr., ventrolateral view ; 5, veliger, 65 hr., lateral view.

\

becomes flattened and shows the onset of differentiation of the eyes as a pair
of dark brown dorso-lateral pigment spots and tentacles as a pair of blunt,
short protuberances ventral and median to the eyespots. The head and visceral
mass retain the pinkish-brown opacity indicative of continued lecithotrophy,

D. T. ANDERSON 109

but the beginnings of differentiation of the gut can be discerned in the visceral
mass and the columella muscles become contractile, producing complete with-
drawal into the shell in response to stimulation. The foot also becomes highly
muscular and mobile, elongates slightly in a posterior direction, and develops
a layer of short, continuously-beating cilia over its ventral surface. Slow creeping
in an exploratory manner over the substratum on the ciliated sole of the foot
begins towards the end of the third day, but the larva is unstable in the creeping
position and frequently tips over to one side or the other. Spasmodic beating
of the velar cilia during creeping appears to assist in maintaining balance while
the foot is in this rudimentary condition.

During the fourth day, although brief swimming excursions continue,
metamorphosis becomes more evident and the capacity for strong attachment
and creeping in a straight line on the foot is enhanced (Fig. 6). The velum
continues to shrink, its cilia becoming finer and shorter, while the tentacles
elongate and become highly mobile. The ciliated foot is also greatly elongated
antero-posteriorly, while in the visceral mass, although some yolk remains,
the coil of the intestine leading to the anus becomes conspicuous. Crawling
on the foot occurs in the typical'snail manner, the shell being held upright and
the tentacles extended forwards, outwards and downwards, rhythmically tapping
the substratum in front of the animal. In contrast to their activity during
the earlier phase of more unstable crawling, the velar cilia now remain at rest,
partly covered by the shell, as the animal creeps along. While attached by
the sole of the foot, however, the animal cannot withdraw fully into the shell,
part of the foot remaining uncovered when columella muscle contraction occurs
and the shell is clamped down on the body. Full withdrawal is possible only
if the foot becomes detached from the substratum.

During the fifth day, the same type of crawling behaviour is pursued more
vigorously, with the tentacles and the anterior end of the foot pushing out from
side to side in what appears to be an exploratory manner, and with frequent
changes of direction. At the same time, internal differentiation proceeds
rapidly in the visceral mass, the coiled gut becoming more obvious, and the
tentacles grow longer and the eyespots larger. A general increase in muscular
activity is also evident during this time. The velum, however, does not appear
to undergo further reduction.

Progressively, development in the same direction continues during the
next four days, growth of the foot and tentacles being accompanied by further
differentiation of the visceral mass (Fig. 7). The velum reduces in size only
very slowly during this time, although its cilia become finer and shorter and
the swimming excursions made become more and more infrequent and more
and more feeble. On the last of these days, further addition to the margin
of the shell begins, giving it the circular marginal outline of an incipient adult
shell, and following this, on the tenth day, the capacity for swimming is lost.
The animal (Fig. 8) crawls actively on its foot, and if dislodged, immediately
reattaches. The gut is by now very well developed and although feeding has
not yet begun, it is obvious that it must soon do so. Development was not
followed beyond this point. :

Chiazacmaea flammea

The mature oocytes of C. flammea (Fig. 9) are brown in colour, uniformly
yolky and opaque, covered by a thin egg membrane and, after immersion in
sea-water for 30 minutes, spherical, with a diameter of 130u. A layer of
colourless jelly covering the egg swells to a thickness of about 10u, but there
is no evidence of adhesion between eggs. Like those of NV. petterdi, they probably
float singly and demersally after natural spawning. Fourteen hours after
fertilization, free swimming trochophores (Fig. 10) are found near the water
surface, swimming by a metachronal clockwise beating of the long thick cilia
of the prototroch, short curving bursts being interspersed with periods of rest.

110 REPRODUCTION AND EARLY LIFE HISTORIES OF SOME GASTROPODS

The brown coloration of the egg is retained in the yolky interior of the trocho-
phore, occupied by macromeres. On the conical episphere, in addition to a
general motionless ciliation and paired antero-lateral protuberances, is an apical
tuft of long cilia, normally held stiffly upright during swimming but also showing
slow bending and waving movements. The hyposphere is unciliated save for
a terminal tuft of long motionless cilia borne on a small postero-ventral pro-
tuberance. In all general respects the trochophore of C. flammea is similar
to that of N. petterdi.

\\
Wane’

\ fie
poe

Figs 6-8. Notoacmaea petterdc. 6, Karly metamorphosis, 90 hr., lateral view; 7, continuing
metamorphosis, 64 days, ventral view ; 8, first permanently settled stage, 10 days, lateral view.

Development proceeds rapidly. Within the next five hours (Fig. 11), the
hyposphere enlarges dorsally, the globular larval shell is secreted, and a small
simple foot rudiment grows out mid-ventrally behind the prototroch. The
latter undergoes little change, but swimming in a semi-upright position becomes
more or less continuous, with periods of slow swimming interspersed with short
faster curving bursts. The episphere shows no change other than loss of the -
apical tuft.

D. T. ANDERSON 111

During the remainder of the first and throughout the second day (Fig. 12),
steady swimming near the water surface continues as the veliger becomes
progressively elaborated. In spite of this, the prototroch undergoes reduction
in size during this time, although its cilia remain long and active. The episphere
becomes flattened and unciliated, while behind the prototroch, the foot rudiment

I3

Figs 9-13. Chiazacmaea flammea. 9, Mature oocyte; 10, trochophore, 14 hr., ventral view ;
11, veliger, 19 hr., dorsal view ; 12, veliger, 44 hr., lateral view ; 13, early metamorphosis, 4} days,
lateral view.

enlarges, growing posteriorly, secretes an operculum, and becomes ciliated
over its ventral surface. The larval shell is greatly enlarged and the visceral
mass undergoes torsion and begins to show differentiation of the gut. Some
muscular activity also becomes evident in the foot and visceral mass, but with-
drawal into the shell does not occur. In swimming, the velum is projected
antero-dorsally, with the visceral mass and shell suspended below it, and the

112 REPRODUCTION AND HARLY LIFE HISTORIES OF SOME GASTROPODS

beating of the velar cilia draws the animal along in a vertical position with
the foot trailing. The visceral mass is still opaque, due to the presence of
brownish yolk reserves. —

Steady swimming and progressive development continue during the third
day, the visceral mass becoming more differentiated and the foot longer and
more muscular. During the fourth day (Fig. 13), paired dark brown eyespots
are developed dorso-laterally on the episphere, while ventral and median to
them paired tentacles grow out. At the same time, the velum begins to shrink
and swimming becomes interspersed with periods during which the larva settles
and crawls very slowly and feebly on its foot, with the shell held upright.
Gradually over the next three days, with little further change in appearance,
the capacity for swimming is lost and crawling greatly improved. The mode
of crawling is similar to that of N. petterdi at the corresponding stage. By
this time, most of the yolk reserves have been utilized but there is no evidence
that feeding begins before settlement has become permanent. Development
was not followed beyond this point.

Patelloida alticostata

The mature oocytes of P. alticostata (Fig. 14) are yellow in colour, uniformly
yolky and opaque, covered by a thin egg membrane and, after immersion for
30 minutes in sea-water, spherical, with a diameter of 150u. Sea-water
immersion also causes a double layer of colourless jelly to swell up around the
egg, a uniform, dense, inner layer being covered by a less dense, irregular, outer
layer. Some tendency to adhesion is observed between eggs, and it is possible
that in natural spawning the eggs adhere temporarily as a gelatinous egg mass
until hatching and escape of the trochophores occurs.

Trochophores (Fig. 15) are found swimming actively near the surface of
the water 18 hours after fertilization. They are semi-opaque, filled internally
with a mass of yellow yolky macromeres, and differ from the trochophores of
N. petterdi and C. flammea in a number of ways. The prototroch is more pro-
tuberant, with a large number of much finer cilia and, although these beat in
clockwise metachronal rhythm in the usual way, the swimming of the trochophore
is a slow continuous straight line progression, without the darting, curving
movements characteristic of the other species. The conical episphere is finely
ciliated, with low paired antero-lateral protuberances bearing long cilia, and
with little development of an apical tuft. The hyposphere bears the usual
terminal tuft of fine cilia.

During the second day of development (Fig. 16), the prototroch enlarges
slightly as a velum, its cilia become more powerful and steady swimming near
the water surface continues. At the same time, the episphere flattens and
loses much of its ciliation, while the hyposphere enlarges and differentiates as
a dorsal visceral mass secreting a globular shell and a ventral foot rudiment
secreting a posterior operculum. The visceral mass is still very yolky. Torsion
occurs towards the end of the second day.

During the third day of development (Fig. 17), steady swimming in a semi-
upright position continues, but the velum begins to shrink. The episphere
also becomes more flattened and develops a pair of brown eyespots. Little
change occurs in the foot rudiment other than development of a fine ventral
ciliation, but the larval shell becomes further enlarged and the visceral mass
further differentiated, showing retention of yellow yolk reserves mainly ventrally.
Muscular movements in general are not conspicuous in the larva at this stage,
and, although complete withdrawal into the shell is possible, accompanied by
Sinking through the water, there is no attachment or creeping on the sub-
stratum. The early phases of metamorphosis, however, proceed rapidly during
the fourth day, with further reduction of the velum, outgrowth of paired
tentacles on the head, elongation of the foot, and further differentiation of
the organs of the visceral mass. Muscularity is greatly increased, and by the

D. T. ANDERSON alates

end of the fourth day crawling predominates over swimming. After two
further days, although the reduced velum is still retained, the crawling habit
has become permanent and the young animal is closely similar to the young

of NV. petterdi illustrated in Figure 8. Feeding does not begin until settlement
is complete.

WU

soby

|7

Figs 14-17. Patelloida alticostata. 14, Mature oocyte; 15, trochophore, 18 hr., anteroventral

view; 16, veliger, 42 hr., ventral view; 17, veliger just entering metamorphosis, 66 hr., lateral
view.

DISCUSSION

In N. petterdi and C. flammea, eggs are probably spawned singly into the
water, as in Aemaea virginea and Acmaea fragilis (Boutan, 1898 ; Willcox, 1898,
1900), since they show no tendency to adhere after artificial release. In P.
alticostata, in contrast, released eggs adhere temporarily by their outer jelly
coats, and it is possible that in natural spawning they aggregate as a transient
egg mass, aS in Acmaea testudinalis (Kessel, 1964).

H

114 REPRODUCTION AND EARLY LIFE HISTORIES OF SOME GASTROPODS

For Acmaea testudinalis, Kessel (1964) has shown that at 12°C, the 140u
egg hatches as a free-swimming trochophore in 10-13 hours. It remains
lecithotrophic for about 50 hours, attaining during this period a well developed
pretorsional veliger stage with a circular monotrochal velum. Planktotrophy
then begins, torsion occurs and the veliger remains planktotrophic, with further
development of the shell, foot and visceral mass, for about 25 hours. Towards
the end of this period, eyes and tentacle rudiments begin to differentiate in
the head and swimming begins to alternate with periods of crawling on the
now well developed foot. Permanent settlement, with crawling and feeding,
is established within 15 hours (i.e., by the time the larva is four days old) but
metamorphosis, with loss of the velum and operculum, further elaboration of
the head and foot and onset of secretion of the adult shell, does not become
obvious until 11 days after settlement.

Development in JN. petierdi, C. flammea and P. alticostata, while generally
similar in the three species, differs from that of Acmaea testudinalis in a number
of ways. Although egg dimensions, mode of spawning and fertilization, and
early hatching as a lecithotrophic free-swimming trochophore are shared in
common, and planktonic life is equally brief (about 30 hours in NV. petterdi and
60 hours in C. flammea and P. alticostata at 20°C, compared with about 75 hours.
in Acmaea testudinalis at 12°C), lecithotrophy is maintained throughout plank-
tonic life. Development of the eyes and tentacles and onset of velar shrinkage
are more precocious than in the planktotrophic larva of A. testudinalis. At
the same time, the transition to permanent settlement, preceding the onset of
feeding, occurs more slowly, taking three days in P. alticostata, four days in
C. flammea and eight days in JN. petierdi, and is accompanied by gradual meta-
morphosis and functional differentiation of the organs of the visceral mass.

Thus N. petterdi, C. flammea and P. alticostata are adapted to a more
economical utilization of yolk reserves than Acmaea testudinalis. The bio-
chemical basis of this difference is obscure, but if we regard the planktotrophic
development of A. testudinalis as primitive, it is a difference which appears
to offer certain advantages. In the absence of velar elaboration, planktotrophic
life in A. testudinalis is necessarily brief and rapid permanent settlement is
essential to the transition from planktonic to bottom feeding, even though the
onset of metamorphosis is delayed for several days. In C. flammea, P. alticostata
and especially N. petterdi, with similar larval dimensions and a similar simple
velum, permanent planktonic life is equally brief, but planktonic feeding is
obviated and the delayed onset of bottom feeding is associated with inter-
mittent swimming excursions during the several days before secretion of the
adult shell begins. Thus development is more direct but the distributive
planktonic phase is more prolonged. From such a mode of development it is
but a short step to the ovoviviparity and birth as a crawling juvenile described
for the Arctic Acmaea rubella by Thorson (1935).

Acknowledgements

It is a pleasure to acknowledge the assistance during this work of Miss
E. ©. Wood, and the advice of Miss I. Bennett on the collection of specimens.
The work was supported by a research grant from the University of Sydney.

References

AnpeErson, D. T., 1960.—The life histories of marine prosobranch molluscs. Journ. malacol.
Soc. Aust., 4: 16-29.

Boutan, L., 1898.—Sur le développement de l’Acmaea virginea. C. R. Acad. Sci. Paris, 126:
1877-1879.

Daxtn, W. J., 1953.—Australian Sea Shores. (Sydney, Angus and Robertson).

Kessen, M. M., 1964.—Reproduction and larval development of Acmaea testudinalis (Miller).
Biol. Bull. Wood’s Hole, 127: 294-303.

THorson, G., 1935.—Studies on the egg capsules and development of Arctic marine prosobranchs.
Medd. Grnland., 100: 1-71.

Wittcox, M. A., 1898.—Zur Anatomie von Acmaea fragilis Chemnitz. Jen. Zeitschr. f. Natur-
wiss., 32: 411-456.

. 1900.—Notes on the anatomy of Acmaea testudinalis Miller. Science, 11: 171.

MALARIA IN THE D’ENTRECASTEAUX ISLANDS, PAPUA, WITH
PARTICULAR REFERENCE TO ANOPHELES FARAUTI LAVERAN

MARGARET SPENCER

Formerly Entomologist, Department of Public Health,
Territory of Papua and New Guinea

[Read 28th April, 1965]

INTRODUCTION

The observations set out below were made during the years 1956-59, as
part of the investigations into the status of malaria in the indigenous popula-
tion of the D’Entrecasteaux Islands, carried out jointly with the Medical Officer,
at the direction of the Department of Public Health.

Detailed studies were made to obtain understanding of the pre-operational
norm, necessary for appraisal of any noticeable events which may occur after
vector control is begun. Previous observations of other workers on the
behaviour of A. farauti are related to observed events in the D’Entrecasteaux
Islands.

About 40,000 anophelines were taken resting or biting outdoors, indoors
and in window traps in more than 50 hamlets and several European stations.
Extensive larval collections were also made. The majority of the collections
were from coastal hamlets. Attention was also paid to garden houses and to
mountain hamlets. The inland anophelines taken on Fergusson Island occurred
at a height of about 400 feet above sea level. An anopheline species distribution
list for the D’Entrecasteaux Islands has been published (Spencer and Spencer,
1960).

GENERAL DESCRIPTION

The D’Entrecasteaux Islands lhe roughly 60 miles to the north and north-
east of Milne Bay, and have an extended coastline totalling nearly 100 miles
in length, a submerged mountain range with three main islands. The highest
mountain peak rises to over 8,000 feet. The coastal strip varies in width.
In many places it is characterized by swamps and lagoons of varying size.
Steep mountain drainage results in bars and deltas at the mouths of streams.

The population of 30,000 indigenes is scattered in small hamlets, often of
about ten houses only. Many of the hamlets are coastal; the mountain
dwellers frequently visit the coast and may have fishing shacks. The majority
of houses are on stilts. The roofs are usually of overlapping sago leaf, the
walls of parallel upright sago stems, and the floors of palm slats. Most
activities are carried on outside the house. The village people may retire
about 9 p.m., or may sit outside until late in the night, especially on moonlight
nights. They do not become accustomed to the bites of anophelines which
worry them a great deal. Mosquito nets are not commonly used.

Recruiting from these islands, as a work force for other areas, has been
heavy. The period spent away is about 18 months at a time. There are also
trading movements to and from the “‘ mainland ” (the island of New Guinea)
and the other islands. Important features of the social organization here are
the small size and physical separation of the hamlets, the restricted number
of people inhabiting them, and their placing relative to anopheline breeding
grounds.

PROCEEDINGS OF THE LINNEAN Society oF New SoutH Watss, Vol. 90, Part 1

116 ANOPHELINES AND MALARIA IN THE D’ENTRECASTEAUX ISLANDS

The coastal hamlets and fishing shacks are built near small mountain
streams with their attendant backwaters and lagoons. Separate garden
houses are a feature of some groups (often distant from the hamlet house),
and these may be built without walls. Hamlet houses in some places may
lack walls. In some places the yam harvest is stored in a separate room or
on an overhead platform within the house.

Annual rainfall has varied from 59” to 149". The average is probably
about 100”. The northern end of Goodenough Island has a well-marked wet
and dry season, with rain particularly concentrated in the months December
to March. Elsewhere in the D’Entrecasteaux Group rain falls throughout the
year, with no particular pattern, except a possibility of lower and higher rain-
fall cycles extending over periods of three or more years. The average
maximum and minimum monthly temperatures vary between about 70° and
90° F, the range being between 62° and 95° F. Relative humidity averages
vary from 74 per cent to 88 per cent, with 57 per cent as the lowest figure.

OBSERVATIONS IN UNSPRAYED AREAS

Anophelines were collected resting or biting, outdoors and irdoors, and
in window traps, in more than 50 hamlets and several European stations, of
the three main islands of the D’Entrecasteaux Group. Extensive larval
collections were also made. The majority of the collections were from coastal
hamlets. Attention was also paid to garden houses and to mountain hamlets.

Standard techniques were used before and after spraying. Mosquitoes
collected by sucking-tube were transferred to small mosquito-net cages tied
by tapes to rigid frames attached to a plywood base. These cages were
wrapped in moistened cottonwool and carried in a calico bag. Fed mosquitoes
survived very well in them. Where hourly time-groups were required, as for
man-biting catches throughout the night, 12 such cages were used.

We measured the incidence of association of the vector species with man
by means of man-biting catches, as described by Pampana (1963). We
standardized our technique on a 12-hour catch, 6 p.m. to 6 a.m., during the
course of these investigations. Two collectors alternated throughout the night.
They were dressed in shorts and shirts, and each caught from his own legs for
a three-hour period before resting. These men were skilled and reliable.
Initially they used mouth sucking tubes, later the battery-operated suction
devices which made the collecting both much easier and more efficient.
Wherever possible, the collectors sat among, or near, the village people at
their normal activities.

Outdoor biting

Over one series of pre-spraying catches covering a 14-month period, a
total number of 76 outdoor biting catches was recorded. These catches were
made at varied localities on the perimeter of the three large islands of the
D’Entrecasteaux Group. Within this series were two series of repeated catches
carried out at Uiaupolo hamlet (nine catches), and Bwalalea hamlet (20 catches),
these two hamlets being considered typical coastal hamlets reasonably repre-
sentative of many others.

Indoor biting

Over 11 months of the pre-spray period, a total series of 31 indoor biting
catches was made. A few comparative catches were made initially outdoors
and indoors on the same night.

Night house-resting

The pre-spraying resting pattern, or position within and on the house
relative to time, was investigated in (a) a number of different hamlets, and

MARGARET SPENCER WAG

(6) in the same hamlet, by all-night catches and _ limited-period calgles
(10 p.m.—2 a.m.) from all parts of the house.

1. For resting position, state of feeding, and a gross ime: tlistibution
(before and after midnight, etc.), the results from the different hamlets were
combined.

2. For the hourly resting time-pattern, collections were made from the
same three houses over three successive nights, making a total of nine house/
nights. A number of collectors maintained as nearly as possible a continuous
collection from all parts of the house on each night. Disturbance would
probably contribute to lowered numbers of resting mosquitoes.

Pre-spraying Tindbit -trap studies

Window traps were fixed to ‘‘ window” apertures on ordinary hamlet
houses. A continuous series was carried out in the hamlet of Bwalalea. The
results were averaged.

Dissections were carried out on time-group samples of the window-trapped
mosquitoes, to form the basis for an estimation of the movements of nulliparous
and parous individuals. The ovarioles were examined in saline to determine
if they were nulliparous or parous, to arrive at a value for “ p”’, and to attempt
age estimations for individual mosquitoes. vt

Pre-spraying sporozoite rate

The full series of dissections embraces three ‘‘ anopheline ecological units ”’,
the hamlets of Bwalalea and Uiaupolo, and the hamlet of Mapamoiwa together
with Mapamoiwa station. These units lie in a line along the north-western
coastal strip of Fergusson Island, separated by intervening bush, but the
interlinking human traffic means that from the point of view of infection risk
these three areas must be combined: together they may be regarded as
reasonably representative of the island group. The anophelines dissected
included those caught biting in hamlets and in garden houses, resting in
houses by day and by night, and resting outdoors by day. The window-trap
series was from Bwalalea hamlet only.

Human blood index

To establish the pre-spraying human blood index, blood meal smears
from outdoor resting mosquitoes were taken according to standard procedures
recommended by WHO, and precipitin testing was carried out by the Lister
Institute (England).

ANOPHELES SPECIES RECORDED

A. farauti—widespread, abundant, and the major vector.

A. punctulatus—also widespread, with distinct fluctuations of population :
of more limited and more localized importance than A. farauti, but contributes
to the transmission of malaria in these islands.

A. koliensis—adults found in very low numbers in a few places.

A. subpictus—widespread, and at times abundant: importance as a
possible vector not known. Showed a decided tendency to attack man and
to rest within houses.

A. annulipes—adults rarely taken, larvae sometimes numerous ; apparently
little affinity for human blood.

A. bancrofti—larvae taken in a few places: no adults taken: adults were
collected on Goodenough Island during the war.

A. longirostris—both adults and larvae taken in low numbers from a few

laces.
The A. punctulatus complex needs further taxonomic investigation. We
found a number of specimens, from several localities, of A. punctulatus with

118 ANOPHELINES AND MALARIA IN THE D’ENTRECASTEAUX ISLANDS

irregular black scaling on the apical pale part of the proboscis, sometimes
extensive. These specimens resemble Woodhill’s suggested hybrids between
A. punctulatus and <A. farauti. The geographical distribution of species
recorded during the pre-spray survey is shown in Figure 1, Spencer and Spencer,
1960.

_ In a total of 6591 anophelines taken biting out of doors throughout the
night, from 83 catches spread over two years, the species composition was as
follows: A. punctulatus (2-5 per cent), A. farauti (94-6 per cent), A. subpictus
(2.:7 per cent), A. koliensis (seven specimens only), A. longirosiris (two specimens
only).

In a smaller series of indoor biting catches throughout the night, less
widely spread, A. farauti formed 97-7 per cent of the total of 1926 anophelines
taken from 31 catches spread over two years, with a few A. punctulatus and
A. subpictus and one specimen of A. annulipes.

In night house-resting catches with a total of 1668 anophelines from 49
houses, A. farauti again dominated (92-8 per cent) with A. punctulatus (0-3
per cent) and A. subpictus (6-9 per cent). These catches included a period
when A. subpictus numbers rose due to a prolonged dry spell. Surprisingly,
two adult A. subpictus were taken resting at night in houses about 400 feet
above sea level and well inland.

A. farauti is thus by far the dominant anopheline of the D’Entrecasteaux
Islands, as it is also of all the outer islands to the north, east and south-east
—the perimeter of the Milne Bay District—and beyond.

With suitable rainfall, localized population explosions of A. punctulatus,
limited in time, have been observed in the D’Entrecasteaux Islands. Although
this species may appear to be absent from an area, colonization of transient
surface pools can be rapid. It is widely distributed throughout the D’Entre-
casteaux Islands, normally occurring in small numbers.

BEHAVIOUR OF ADULT MOSQUITOES

Interrelationship of human and anopheline ecology

It is important to have a clear idea of the normal dispersion and behaviour
of anopheline populations in relation to areas inhabited, or not inhabited, by
human beings, and of the normal manner and rate of build-up of anopheline
populations under favourable conditions.

In the D’Entrecasteaux Islands there are large gaps between clusters of
human population, of forest, kunai grass and secondary growth, with a low
density of indigenous mammalian fauna. The distance between individual
hamlets, or hamlet groups, may be greater than the expected range of flight
of anophelines, with consequent isolation of the populations. The coincidence
of highly suitable breeding places, and a concentration of suitable hosts, is
likely to result in a convergence of anopheline activity within the hamlet area
and around its edge. Outside the hamlet areas—and the tracks radiating
from them where interference with bush, and night movements of people, offer
a suitable coincidence of hosts and breeding opportunity—it is less likely that
large concentrations of anophelines can occur.

Under the circumstances outlined above, A. farauti populations can build
up closely dependent upon individual human communities and their domestic
animals. The anopheline populations can be expected to become less dense
as the distance from the hamlet becomes greater, unless the hamlets are situated
elose together with overlapping anopheline populations. The closer the spacing
of the hamlets, the looser the association of any individual A. farauti with a
particular hamlet.

Observations show high adult and larval densities in and around hamlet
areas (the ‘‘ domestic” populations) allied usually with low densities in garden
and bush areas (the ‘ wild’’ population). Residual spraying should cause

MARGARET SPENCER 119

reductions in anopheline density in hamlet areas, and the effects of spraying
should extend some distance into the perimeter areas.

Hamlets represent the more permanent human communities. Temporary
human communities are also formed, to engage in fishing, gardening, making
of sago, or other extra-hamlet activities. If such a temporary community
remains in one area long enough and conditions are suitable, it will cause the
formation about itself of a concentration of breeding from the base level
‘wild ’’ anopheline population. Such places are usually not protected by the
insecticide umbrella. A residual focus of infected mosquitoes here may survive
the comings and goings of different family groups, and infect the later comers.

Outdoor and indoor biting in hamlet areas, biting in garden houses, and day-time
biting

The data discussed in the next two sub-sections throw light on the normal
patterns of biting and the mosquitoes’ basic relationship to man. They also
suggest the degree of contact with the insecticide that the mosquitoes are
likely to have.

The peak period of attack for A. punctulatus was 12 midnight to 3 a.m.
For A. subpictus there was abrupt rise after 9 p.m., but no noticeable peak
period.

The results of outdoor and indoor human bait catches, relating to A.
farauti alone, are shown in Table 1. The calculated average value of the out-
door biting incidence is 62-0 bites per man per night, and of the indoor biting
incidence is 65-2 bites per man per night. The mean value is 63-6 bites per

TABLE |

A. farauti: Night-biting catches and indoor-resting catches

Results arranged to show hourly fractions of biting cycle
(Pre-spray catches in various localities)

Outdoor biting Indoor biting Night mdoor resting
June 1957—Aug. 758 Oct. 1957—Aug. 758 May 1959
Time (catches = 58) (catches = 25) (catches = 9)

Number Percentage Number Percentage Number Percentage

taken of total _ taken of total taken of total
6-7 Duley 135 3°8 77 4-7 71 S07
7-8 p.m. ‘ 213 5-9 86 5°38} 78 6-3
8-9 p.m. 289 8-0 105 6-4 134 10-8
9-10 _ 389 10:8 151 9-3 122 9-8
10-11 p.m. ~ 418 11:6 164 10-1 157 12-6
ll pm —l2 saanidnent 431 12-0 231 14-2 187 15-1
12 Hien Gem 395 11-0 187 ie 144 11-6
1-2 a.m. a 392 10-9 160 9-8 120 9-7
2-3 a 291 8-1 132 8-1 92 7:4
3-4 a.m. 269 7-5 133 8-2 70 5:6
4-5 a.m 236 6-6 112 6-9 43 3°5
5-6 a.m 133 3°7 90 D°5 24 1-9

120 ANOPHELINES AND MALARIA IN THE D’ENTRECASTEAUX ISLANDS

man per night. A reasonable average figure, relating to the whole island group,
and to all seasons, is 65 bites per man per night. This figure represents an
assessed over-all risk for a large area in which malaria is endemic, and in which
there is constant movement and interchange of the people.

The incidence of attack in night outdoor biting by A. farauti tended to
increase to a plateau lasting from about 9 p.m. to 2 a.m., followed by a fairly
steady decline (Table 1). There is no real peak of attack. Pre-midnight attack
is slightly greater than post-midnight attack, approximately 53 per cent : 47
per cent for a total of 4600 A. farauti.

It is suggested that sample outdoor leg biting catches of A. farauti can
be made for any convenient hours of the night, and the total density per man
per night calculated from the percentages shown in Table 1. Alternatively,
the catcher may sit outside while the village people do, then move inside when
they go into their dwelling for the night. The nature of the dwelling allows
free entry and exit to the mosquitoes. In effect, as shown below, mosquitoes
associated with inhabited areas follow the movements of the people. A. farauti
adults, after biting outdoors, will rest on objects near their hosts—stones, tree
buttresses, grass, etc., and have been seen fully engorged in large numbers.

From Table 1 it will be seen that the indoor attack builds up more slowly
than the outdoor attack, there is an intensification of biting after midnight,
and a fall in activity after about 1 a.m. Pre-midnight and post-midnight
attacks are the same, 50 per cent : 50 per cent.

A small series of night biting catches in garden houses was carried out to
determine whether there is a bush population which will attack under those
circumstances. The attack rate was low ; nevertheless in the relatively remote
and isolated garden house, attack does occur. Garden houses and other
temporary shacks are occupied for varying periods of time by varying numbers
of people. These structures often lack walls.

Day-time biting by A. farauti has been noted in a number of different
places—Solomon Islands, Australia, New Hebrides, Trobriand Islands (see
Black, 1955). We did not experience day-time attack in the D’Entrecasteaux
Islands.

Night resting in and on houses, entry and exit, age composition and longevity

House-resting catches were continuous throughout the night and were
from all parts of the house, including eaves, stumps, and under the floor. With
continuous collection, the resting rate largely reflects arrival and entry rate
(Table 1).

High house-resting densities tend to precede high indoor-biting incidence,
although the peak period of resting is at the time of heaviest indoor biting.
Pre-midnight resting is, however, at a higher density than post-midnight resting
(60 per cent : 40 per cent). This ratio was observed in all parts of the house
except the cross beams.

About 60 per cent of anophelines taken resting on houses at night were
taken from inside the houses (verandas are included). The favoured resting
position, 60 per cent of total resting indoors, is on the walls below the level
of three feet. Unfortunately this is the region from which most insecticide is
rubbed off by people sitting on the floor and leaning against the walls. Female
anophelines also rest on such objects as yams stored inside the house. Examina-
tion of distribution throughout the night shows no especially favoured time
for resting underneath the house, but there is a concentration on the eaves
and inside the roof between 10 p.m. and midnight, and on the inside walls
between 11 p.m. and 1 a.m. The fall in resting density between 9 p.m. and
10 p.m. (Table 1) is most marked in those resting on the walls, and is probably
due to the movements of people inside the house as they settle down for the
night.

MARGARET SPENCER AL

As would be expected, the numbers of resting and biting anophelines varied
greatly in individual houses, with the type of construction, relative distance
from the breeding places, number of people and domestic animals associated
with the house, and presence of a domestic fire. Apparently a fire affected
the anophelines by lowering the humidity in its vicinity rather than by the
smoke produced.

Under the conditions of maximum disturbance caused by continuous all-night
collecting from all parts of the house, only 26-5 per cent of those taken during
the night were either partially or fully fed. With intermittent collecting, the
proportion of fed females rose to an over-all 61-2 per cent of the total catches.
Also, the proportion of empty females decreased throughout the night, from
50-5 per cent of the total in the first quarter of the night to only 18-8 per cent

TABLE 2

A. farauti: Pre-spray catches in outlet window traps on houses, Bwalalea,
July-August 1959 ([wo series, maximum of four traps in each)

All-night trap catches All-night trap catches
(traps removed every hour) (traps removed every
Time three hours)
(12 catches) (38 catches)
Number Percentage Percentage Number Percentage
taken per hour per 3 hours taken per 3 hours
6-7 — 63 7-3 Q
7-8 — 26 3-0
8-9 7 38 4-4 14-7 456 16-4
9-10 — 33 3-8
10-11 pains 47 5:5
11-12 sentinel 29 3-4 12-7 491 17-7
12-1 ada 65 7-6 .
J-2 sam 51 5-9
2-3 nthe 90 10-5 24-0 581 20-9
3-4 _ 61 Tell
4-5 2am 116 13-5
5-6 aah, 239 27-9 48-5 1,246 44-9
Total ini 858 2,774 |

in the last quarter. These results indicate resting before and after feeding,
and also suggest that females entering early in the night may rest longer before
feeding than those entering later in the night. Metselaar (1957) records unfed
anophelines in West Irian resting up to 110 minutes. A noticeable number
of fed resting females had apparently not taken a full blood meal. (The
percentage of partially-fed females may be on the high side because of disturbance
due to the collectors.)

Selection of the host may be at random. As well as cats and roosting
fowls, pigs and dogs may be available. However, pigs are not numerous in
most villages. The results of precipitin tests on night-resting A. farauti in
one village without pigs showed a human blood index of 0:69 and a dog blood
index of 0-31. In another village the human blood index of night-resting
A. farauti was 0:97 (Spencer, 1962).

ju
i)
bo

ANOPHELINES AND MALARIA IN THE D’ENTRECASTEAUX ISLANDS

Window-trap studies of exit from houses (Table 2) show that 66 per cent
of the total exit takes place after midnight. The exit rate jumps after mid-
night, with a sharp climax between 5 a.m. and 6a.m. Although these window-
trap studies were on open hamlet houses, with a multiplicity of possible exits,
the results parallel closely those from the carefully controlled trap hut experi-
ments in Hollandia carried out by Van Thiel and Metselaar (1954).

Most females entering houses to feed leave again the same night, and usually
few adults can be found resting in houses by day, even where night populations
are heavy. Metselaar (1957) found in West Irian that 12-3 per cent of 1607
anophelines remained in the trap hut the day following entry. Few adults
appear to remain within the house for the whole period of the gonotrophic
cycle, as 94 per cent of A. farauti females found by day within houses were
recently fed. Day-time house-resting anophelines were usually found low on
the walls in the room most used by the occupants. Black (1955) has discussed
previous records of day-time house-resting A. farauti. We did not find males
resting in houses during the day, although an occasional male has been found
inside a house or a window trap at night.

TABLE 3
A. farauti: Age-grading of parous females entering outlet traps
before and after midnight

Number in each age-group

Number of

dilatations Entering trap Entering trap Total
6-12 p.m. 12-6 a.m.
1 14 27 4]
2 24 18 42
3 11 8 19
4 1 6 7
5) 1 0 1
>) 2 1 3

About 10 per cent of females entering houses leave again without feeding.
Dissections show that these unfed females may be nulliparous or parous. The
majority of females taken in window traps are fully fed, but partially-fed
females (both nulliparous and parous of different ages) occur (Table 3). The
dusk exodus (6-7 p.m.) is likely to include those females which have rested
in houses during the day. Females remaining indoors to mature their eggs
likewise include both nulliparous and parous specimens.

Dissections from window-trap catches show that only 34-7 per cent of
the nulliparous mosquitoes leave the house by midnight. Among parous ones
nearly 50 per cent of the total catch has already left the house by midnight,
and with a falling rate of arrival and accelerating rate of departure after mid-
night, the stay of the parous mosquitoes (and consequently their contact with
the insecticide) may be generally shorter than that of the nulliparous mosquitoes.
It is possible that bursts of females which have just laid eggs arrive to swell
the numbers in houses between 9 p.m. and midnight.

Daily variations in the proportion parous in window-trap catches were
also recorded for a period of two months (Table 4). The relative abundance
of nulliparous A. farauti tended to vary with the density of population, as
would be expected. During breeding flushes, nulliparous and young parous
females predominated ; as the over-all density fell, the proportion of older
females increased.

Supposing a two-day gonotrophic cycle for the majority of A. faraute
under observed conditions, ‘‘p’’ can be derived by taking the square root

MARGARET SPENCER - 123

of the proportion parous.' Three derivations were made from window-trapped
mosquitoes: (1) for one population cycle, covering one full rise and fall in
numbers ; (2) for a period of steady population ; (3) for the whole period of
observations (two months). The value of ‘‘p” was 0-85 in each case, and
this is therefore considered a reasonable pre-operational figure for A. farauti,
although it is derived from a selected sample (window-trapped females only).

Day-time resting out of doors

Both males and females can be found resting among the secondary growth
at the edges of the hamlet clearing, scattered in damp and sheltered situations.
It is likely that both males and females move out from the breeding grounds
after emergence. We did not find resting females in any numbers on the edges
of breeding places, in day-time searches, unless these were situated near a
collection of native dwellings.

TABLE 4

A. farauti: Pre-spray weekly variations in numbers trapped and proportion
parous in window-trap catches, in relation to rainfall

Period Number caught/ Proportion parous Rainfall

trap/night (and No. dissected) (inches)
14-17 July 36 0-70 (67) 0-85
19-20 July 101 0-41 (29) 0-55
21-31 July 17 0-81 (117) --
l— 5 August 9 0-76 (55) —
8-14 August 20 0-71 (173) —
16-19 August 17 0-86 (95) —
25-29 August 10 0-52 (89) 4-78
1— 3 September 24 0-85 (104) 11-92

Mean proportion parous 0-734 (729)

In a series of 848 A. farauti taken outdoors by day around the edges of
a native hamlet clearing (on Fergusson Island), 292 were males. In the females,
36 per cent were gravid, 56 per cent were fed but not fully gravid, and 8 per
cent unfed. The proportion gravid is much higher than in females resting
in houses during the day-time (one per cent) or taken in window traps (two
per cent).

Oviposition and gonotrophic cycle

Fully gravid females probably leave their day-time resting place about
dusk to lay their eggs. The flaccid and distended appearance of the ovaries
in A. farauti females dissected immediately after they were caught attempting
to bite suggests that females return immediately after oviposition to feed again.

The gonotrophic cycle for the majority of parous coastal A. farauti is almost
certainly two days. Partially-fed females may need a longer period to complete
the cycle. The possibility must be considered that the gonotrophic cycle may
not be regularly completed in two days when the opportunities available to
the female to lay eggs are restricted. It appears probable that the nulliparous
period extends over five days. Dissections of females made within 12-15 hours
of their entering window traps indicated that the time taken to reach stage III
of ovary development might be shorter for parous females than for nu'liparous.

SPOROZOITE RATES
The monthly sporozoite rate of A. farauti, in a series of 6049 dissections
extending over two and a half years, varied between zero and about 1-6 per
cent (Table 5). The over-all sporozoite rate from all catches was 0-63 per

1 According to Davidson (1954, Nature, 174: 792 and 1955, Ann. trop. Med. Parasit.,
49: 24), this interpretation is valid only if the sample dissected is limited to females caught
when the ovaries are in stage III of development.

124 ANOPHELINES AND MALARIA IN THE D’ENTRECASTEAUX ISLANDS

cent. This is probably too low, and a more representative estimate would
be one per cent in an average year. There was some indication that infected
anophelines are particularly likely to occur in June-July. In 1958, within
the monthly fluctuations there appeared high proportions of infected anophelines
in different localities at the same time, e.g. in February at the hamlet of Uiaupolo
and at Mapamoiwa station, in March at the hamlet of Bwalalea and at Mapamoiwa
station.

TABLE 5
Sporozoite rates in A. farauti and A. punctulatus in pre-spray periods
Quarterly A. farauts A. punctulatus
period -
No. dissected % positive No. dissected % positive
1956/1V2 18 0-0 145 0
1957/1 123 0-8 43 0
1957/11 360 0-28 4] 0
1957/I11 383 0-52 5 0
1957/1V 183 0-0 12 0
1958/1 564 1-4 2 0
1958/11 482 1-6 4 (1/4)
1958/11» 210 0-95 0 =
1958/I1V 0 — 0 =
1959/Le¢ 647 0:77 0 —
1959/11 3,079 0-36 ] 0
Total 6,049 0-63 253 0-4
a November and December only. ? July and August only. © February and

March only.

Among 18 sporozoite-positive A. farauti found in the period February to
July 1958, the rates of positivity were eight positive (in 154 dissected) caught
biting in hamlets, five positive (in 48 dissected) caught resting indoors, and five
positive (in 92 dissected) caught resting out of doors.

Between March and July 1959, in a total of 774 dissections of A. farauti
taken resting on houses at night, the sporozoite rate was 0-9 per cent. In the
same period, in a total of 2733 dissections from window traps in the same hamlet
area, the sporozoite rate was 0-29 per cent. The difference suggests that the
catches from the window traps contain a higher proportion of nulliparous and.
young parous females than the night-resting catches. In 253 salivary gland
dissections of A. punctulatus, one positive specimen occurred. This was in a
night-biting catch (Table 5).

POST-OPERATIONAL OBSERVATIONS
Reduction in anopheline densities on Goodenough Island

Spraying began on Goodenough Island with DDT in November, 1958.
The second spray round using dieldrin was on schedule six months later. The
third spray round, also using dieldrin, was apparently delayed until it was
four months overdue.

Since spraying began, a marked and persistent reduction in anopheline
density throughout the inhabited parts of the island has occurred.* Both
adults and larvae are relatively scarce. This can be taken as the effect of
spraying under the existing ecological conditions, superimposed upon natural
fluctuations due to climatic factors.

In April 1959, about four and a half months after the first spray round
of DDT, the total number of A. farauti taken resting on the walls of houses
between 10 p.m. and 2 a.m. had been reduced (in four villages) from an average
of 25 per house in 1957 to an average of 0-5 per house, and A. subpictus was

* Written at the end of 1960.

MARGARET SPENCER 125

not found at all. The night-resting searches were extended to a total coverage
of 21 villages around Goodenough Island. Of 187 houses searched, only 31
had anophelines resting on the walls between the hours of 10 p.m. and 2 a.m.
The total number of anophelines taken from the 21 villages was 135 A. farauti,
three A. punctulatus and two A. subpictus. This indicated a marked reduction
in anopheline density. In most of the positive houses, the total number resting
on the walls during the four hours of observation was less than five in each
house. Only a minority of positive houses were new, or had new walls. In
some of them the spraying was patchy.

The above observations were followed up in June 1959 by an all-night
biting catch in 11 villages where similar catches in 1958 had yielded averages
of 93 A. farauti and 16 A. subpictus per man per night (Table 6). Seven
observations now gave a negative result, and the over-all average biting

TABLE 6

Results of man-biting all-night catches in hamlets
of Goodenough Island, surveyed in two successive years

A. Average biting density in seven hamlets
surveyed twice before spraying

Mosquitoes per man per night

Year
A. farauti A. subpictus
1957 174 1
1958 82 12

B. Average biting density in eleven hamlets
surveyed once before and once after DDT-

spraying
Mosquitoes per man per night
Year
A. farauti A. subpictus
1958 93 16
1959@ 9 0

4 Seven of the 11 hamlets gave a negative result
—see text.

densities were nine A. farauti and no A. subpictus. In the four positive villages
above the average catches, per man per night, were 24 A. farauti and no A.
subpictus, compared with 1958 averages in these four villages of 87 A. farauti
and 22 A. subpictus.

One year later, in June-July 1960, immediately following the third spray
round (dieldrin), an experienced collector reported a marked scarcity both of
larval and of adult anophelines. Small numbers of mature A. farauti larvae
were found in five places. A total of five adult A. farauti for the whole island
was taken in a “ leaking’ mosquito net. Limited biting catches from 10 p.m.
onwards showed correspondingly small numbers of adult A. farawti, the only
Species taken.

Since the start of spraying, A. farauti occurred in such small numbers
and with such a very scattered distribution, as to discriminate against any
study of post-operational behaviour patterns.

Reduction of anopheline densities on Fergusson Island

Spraving began on Fergusson Island in March 1959 with dieldrin. A
“‘ check ” area was retained during the first spray round where pre-operational
entomological observations were continued until June 1959. The second spray

126 ANOPHELINES AND MALARIA IN THE D’ENTRECASTEAUX ISLANDS

round was carried out on schedule in September of 1959, and dieldrin was used
again. On this round the former ‘‘ check” area was included. The third
spray round on Fergusson Island had not begun at the time of the anopheline
survey.

This survey of Fergusson Island during July and August 1960 investigated
larval numbers and distribution, and a ‘ leaking ’’ mosquito net was used for
a rough estimation of adult numbers, as on Goodenough Island.

From this survey an over-all reduction in anopheline densities was evident
on Fergusson Island also. It seems that adult anophelines, while reduced in
number and distribution over the whole of Fergusson Island, remain most
apparent in the former ‘‘ check ”’ area, where catches of up to 39 anophelines
in one night were recorded from the ‘ leaking”’ mosquito net. The total
number of adult anophelines from 20 villages sampled in this way—including
some villages which previously had dense anopheline populations and heavy
biting rates—was 177 A. farauti, 13 A. punctulatus, and five A. subpictus. Of
the total of 177 A. farauti more than half came from within the former ‘‘ check ”
area.

Mature A. farauti larvae were taken from 14 places on the island’s peri-
meter, only three of these within the former ‘‘ check” area. Mature larvae
were more apparent, both in numbers and in distribution, than on Goodenough
Island. Two A. punctulatus larvae were found outside the former “ check ”
area. No larvae were recovered well inland, although A. farauti adults
occurred in noticeable numbers in one mountain village.

DISCUSSION

The age composition of the hamlet or ‘‘ domestic” populations of A.
farauti is variable, depending upon the rate of emergence of new individuals
and the rate of dying off of all age-groups. The nulliparity ratio tends to
reflect the numerical trend shown by trap catches and expected from rainfall
distribution: it is by itself a useful comparative measure of events. The
estimated mortality rate is 15 per cent per day. The oldest individuals seen
to date were estimated to be three weeks old.

The pre-operational data form a basis for deriving the following entomo-
logical indices. Macdonald’s (1957) symbols are used :

(1) ‘‘m” (anopheline density in relation to man) for coastal hamlets
AY

and all seasons, is equal to the man-biting rate (ma) divided by the

65 (bites per man per night) 173:

-375 (estimated man-biting habit) d

= blood preference x gonotrophic cycle

== (oti) < Ob == Uosgars

(3) “‘s” (sporozoite rate, or proportion with sporozoites in their glands)
== (oil 2

(4) ‘‘p” (probability of survival through one day, estimated as the square
root of proportion parous) = 0-85.

man-biting habit (a), viz.

0
(2) “a” (man-biting habit)

From the epidemiological viewpoint, attention should be focused on two
problems, (a) transmission within villages, and (b) transmission outside villages.
These twin problems may be overlapping, or quite separate, according to the
distances involved and the degrees of interference with natural ecology, between
the village areas and the areas of temporary habitation outside the village.
We may have to think in terms of separate ecological units, or overlapping
or continuous ones.

For example, if a sago marsh suitable for anopheline breeding, temporarily
but regularly inhabited by members of one village group, lies within flight
range of the village, transmission within that village overlaps with trans-
mission outside that village and in the sago marsh area.

MARGARET SPENCER 127

On the other hand, if the gardening area lies some miles away from any
village and is separated from all villages by virgin forest, transmission of
malaria by an anopheline population established in relation to the gardening
area will in all probability be quite separate from transmission within the
villages depending upon those gardens.

A third situation is that in which villages sited in a swampy coastal area,
where there are abundant breeding places to produce a dense anopheline
population, may be well separated from each other, but joined into a continuous
ecological unit by an abundant wild mammalian fauna (such as pigs and
wallabies) which sustains a dense anopheline population between the villages.

Each of the three situations presents difficulties in control, and one or
other of them will be present in all parts of this Territory. The best theoretical
chance of good control of transmission within the villages occurs when the
villages are isolated units set in clearings in virgin forest. Both A. farauti
and A. punctulatus are evidently very susceptible to residual insecticides in
settled village areas. The marked reduction in adult numbers which occurs
after spraying is evidence of this.

These factors must all be taken into account when evaluating the sequence
of events which follow upon spraying. They are also relevant in determining
where we should concentrate our attention. It may be that control of trans-
mission by residual spraying can never be adequate because we cannot fully
protect the people by this means alone during their extra-village activities.

Acknowledgements

This paper was issued as No. 454 in the World Health Organization’s
mimeographed ‘‘ WHO/Mal.” Series, 25 June 1964. The work was carried
out as part of the preliminary assessment for the Malaria Eradication Campaign
in the Territory of Papua and New Guinea. Indigenous staff under the leader-
ship of the Papuan Health Assistant, Mr. Jonathan Baloiloi, carried out the
arduous routine work of mosquito collection and examination. The writer
records her debt to her husband and colleague, Dr. T. Spencer, and also grate-
fully acknowledges the advice of Mr. C. Garrett-Jones of the World Health
Organization on the editing of this paper, which is published now with the
permission of the Director of Public Health for the Territory of Papua and
New Guinea.

References

Brack, R. H., 1955—Observations on the behaviour of Anopheles farauti Laveran, an important
malaria vector in the Territory of Papua, New Guinea. Med. J. Aust., 42: 949.

MeETSELAAR, D., 1957.—Pilot project of residual msecticide spraying in Netherlands New
Guinea. (Thesis, Leyden.)

Pampana, EH. J., 1963.—‘ A textbook of malaria eradication.” (London.)
Spencer, M., 1962.—Blood preferences of Anopheles farauti. (Unpublished document WHO/
Mal/377.)

SpencER, M., 1964.—Malaria eradication : Blood preferences of Anopheles farauti. Papua N.
Guinea med. J., 7: 19.

SPENCER, T., and SpENcER, M., 1960.—Malaria assessment methods. Papua N. Guinea med.
J., 4: 55.

Van Tuite, P. H., and Metseraar, D., 1954.—A pilot project of residual spraying as a means
of controlling malaria transmitted by anophelines of the punctulatus group in Netherlands
New Guinea. Docum. Med. geogr. trop. (Amst.), 7: 164.

Author’s present address: 1 George Street, Tenterfield, N.S.W.

A NOTE ON BLOOD PREFERENCES OF ANOPHELES FARAUTI

MARGARET SPENCER

Formerly Entomologist, Department of Public Health, Rabaul
New Guinea

[Read 28th April, 1965]

A total of 702 blood meals from A. farauti females collected on Fergusson
Island, New Britain, and Nissan Island, were sent to the Lister Institute, London,
under arrangement with the World Health Organization. The results of
precipitin tests on these are set out and discussed.

The aim was to establish a definite figure for the ‘‘ human blood index ”’!
in the villages of the D’Entrecasteaux Islands in the Territory of Papua and
New Guinea.

Copies of this paper were distributed by the World Health Organization as a paper in the
WHO/Mal series of documents (WHO/Mal/337, 1962), a series which does not constitute formal
publication.

Unfortunately, after the paper was accepted for publication in the PROCEEDINGS, it appeared
in the Papua-New Guinea Medical Journal, vol. 7, no. 1, dated December 1964. It is therefore
unnecessary to reprint it here. :

Kp.

1“ Human blood index ”’ is suggested by the World Health Organization (1959) as a better
term for the closeness of relationship of vector to man than “‘ anthropophilic index”’, and is defined
as the proportion of freshly fed Anopheles giving a positive precipitin reaction for human blood.

Ho —.
AUSTRALASIAN MEDICAL, PUBLISHING CO. LTD.
SEAMER AND ARUNDEL STS., GLEBE, SYDNEY

PLATE I

Proc. Linn. Soc. N.S.W., Vol. 90, Part 1

Five species of Australian Chthamalidae.

ak

%

Proc. Linn. Soc, N.S.W., Vol. 90, Part 1 PLATE

Some tropical Chthamalus spp. and their habitats.

Proc. LINN. Soc. N.S.W., Vol. 90, Part 1 PLATE II

Chromosome numbers in some Australian leafhoppers.

:
iy 5

STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS

Tf. GENETIC DIVERSITY IN THE VARIETIES LANDHAFER, SANTA FE, MUTICA
UKRAINE, TRISPERNIA AND VICTORIA FOR CROWN RUST RESISTANCE

Y. M. UPADHYAYA and H. P. BAKER
Faculty of Agriculture, The University of Sydney

[Read 30th June, 1965]

Synopsis

Segregation in the F, and F, generations for rust reaction was studied in certain crosses
between members of the group of crown rust resistant varieties comprising Landhafer, Santa
Fe, Trispernia, Mutica Ukraine (Ukraine) and Victoria, all resistant to the prevalent Australian
races, to assess their genetic diversity with regard to genotypes for resistance. Behaviour in
the seedling stage to several races as well as adult plant field reaction was studied. The two
factor pairs in Landhafer conditioning adult plant resistance, one of which conferred seedling
resistance in addition, were independent of the factors in the varieties Santa Fe, Trispernia and
Victoria. The seedling reaction type of Landhafer was epistatic over those of Trispernia and
Victoria. The factor for both seedling as well as adult plant resistance in Santa Fe was
independent of the factors in Victoria and epistatic to them. Certam modifying gene(s), how-
ever, resulted in the expression of a reaction type similar to that characteristic of Victoria by
suppressing the Santa Fe gene. The Santa Fe factor was considered allelic with the factors
for seedling resistance in the varieties Ukraine and Trispernia, no susceptible segregates occurring
within the limits of the population size studied. The factors were not considered identical,
however, since Trispernia exhibited a higher reaction type and the allele in Ukraine conditioned
resistance to fewer races than that in Santa Fe. The reaction type of Santa Fe was dominant
over that of Trispernia in tests with four races, but with race 203 the Santa Fe gene was inhibited
by the action of a pair of complementary factors, one contributed by each variety. The three
factor pairs in Ukraine, two acting in complementary fashion, involved im adult plant resistance
were independent of the Santa Fe gene. Indirect evidence indicated that the factors responsible
for seedling resistance in Santa Fe and Victoria were genetically independent. The independence
of the factors conditioning adult plant resistance in Landhafer and Ukraine and likewise the
independence of the Ukraine and Victoria adult plant factors could not, however, be established
in the absence of studies on the appropriate crosses.

INTRODUCTION

In a previous paper (Upadhyaya and Baker, 1962b) the mode of inheritance
in the resistant varieties Landhafer, Santa Fe, Mutica Ukraine, Trispernia and
Victoria was reported in the seedling and adult plant stages to certain of the
most prevalent field races of crown rust (Puccinia coronata avenae Erikss.) in
Australia. The relative merits of these varieties in their role in breeding for
resistance depend, in addition to the mode of inheritance they exhibit, largely
on their diversity with regard to their genotypes for resistarce. Information
on this latter aspect was obtained from intervarietal crosses between them
and is currently presented to show whether the genes which they possess are
identical, allelic, or distinct and non-allelic to Australian races.

It also has been pointed out previously that such knowledge is vital to an
understanding of the basis and significance of information revealed by physiologic
race Surveys since these varieties, together with the variety Bond, of which
the inheritance will be subsequently reported, form the nucleus of the varieties
in the current set used for such surveys.

LITERATURE REVIEW

Results of crosses of the varieties under study with susceptible varieties
were reported by Upadhyaya and Baker (1960, 19620). Seedling resistance
to various Australian crown rust races was conditioned by a single tactor pair

PROCEEDINGS OF THE LINNEAN SocieTY OF NEw Soutsa Watss, Vol. 90, Part 2

130 STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, II

in each of the varieties Landhafer, Santa Fe, Mutica Ukraine (Ukraine) and
Trispernia, and by four factors, Vea Vep (linked complementary) and IVc, Ve,
(linked) in the variety Victoria. For adult plant resistance the variety Land-
hafer possessed an additional recessive factor and Victoria two additional factors
Ve, and Ve;. IVe, Ve, were also operative in the adult stage but not Vea Vep.
Ve, was linked with Veg Vey. The factors for seedling resistance in Sant Fe
and Trispernia, but not Ukraine, also conditioned adult plant resistance. The
adult plant resistance of Ukraine was conditioned by three dominant factors,
two acting in complementary fashion.

Several investigators have presented results of studies on crosses between
certain or all of the resistant varieties currently being reported. Litzenberger
(1949) and Simons and Murphy (1954) found that the factors for resistance in
Landhafer and Santa Fe were independent. Finkner (1954) reported that the
resistance of Landhafer was genetically independent of those of Santa Fe,
Trispernia and Victoria: the factors involved in Ukraine and Victoria were
also considered independent. Simons and Murphy (1954) noted complicated
inheritance in certain crosses between these varieties. Landhafer x Trispernia
gave transgressive segregation with plants more resistant than either parent.
A cross between Santa Fe and Trispernia did not indicate allelism between the
genes in these varieties.

However, some of the factors found in certain varieties were considered by
various investigators to be allelic with, but different from, those in other of
the varieties. Finkner (1954) proposed genotypes thus: Ukraine MMUU,
Santa Fe M,M,U,U, (or M,M,) and Trispernia M,M, and/or two other factors.
Both factors in Ukraine were dominant over those with which they were allelic
in Santa Fe; similarly M, was dominant to M,. Finkner, Atkins and Murphy
(1955) reported that one (M,) of the two linked genes in Santa Fe was allelic
with the single gene M in Ukraine, and recessive to it.

With race 57 of the pathogen, and representing the single genes found to
condition resistance in Landhafer and Victoria as L and V respectively, Finkner
(1954) concluded that the allelic and non-allelic relationships with regard to
dominance or epistasis were in the following order: M or U>M, or U, or
L>V>M,. The immune reaction of Ukraine was dominant or epistatic to
that in the other varieties.

MATERIALS AND METHODS

F,, F, and F, generations were studied for crown rust reactions in crosses
between the five varieties under study. The fol!owing crosses were not
included because either the cross was not made or the F,s failed to set seed due
to adverse environmental conditions: Ukraine < Landhafer, Ukraine x Tris-
pernia, Ukraine x Victoria and Trispernia x Victoria.

Crown rust races employed for testing were 203, 226, 237, 237-4, 259 and
286. These are described by Baker and Upadhyaya (1955) and were built up
from field isolates.

The experimental procedures were set out by Upadhyaya and Baker (1960).

EXPERIMENTAL RESULTS
F, reaction types

The reaction types in the F, of the various crosses in the seedling and also
the adult plant stages, together with those of the parents to different specific
races as well as field inoculum, are presented in Tabie 1.

Genes conditioning resistance in certain of these varieties will be suggested
below to be allelic. In these cases the F, behaviour was an indication of the
dominance relationship. In cases of non-allelism the degree and type of epistasis
manifest was evident from the F, behaviour. From the data in Table 1 in

131

Y. M. UPADHYAYA AND E. P. BAKER

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STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, III

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Y. M. UPADHYAYA AND E. P. BAKER 133

certain cases the lower reaction type (higher resistance) was either completely
dominant or epistatic or the F, reaction type was intermediate between the
parental types. In tests against races 259 and 286, Victoria being susceptible
to the former and Landhafer and Santa Fe susceptible to the latter race, the
Fs of crosses of Victoria with Landhafer and Santa Fe showed intermediate
reaction types. In other cases F, reaction types slightly or distinctly less
resistant than either parent were observed. This occurred in the crosses Land-
hafer <x Santa Fe, Santa Fe « Ukraine and Santa Fe x Trispernia tested
with certain races.

F,, segregation
(i) Seedling tests

The results of studies at the seedling stage on F, populations of crosses
tested with one or several different races to which both parents were resistant
are presented in Tables 2 and 3. Table 2 pertains to crosses involving the
variety Landhafer.

In the cross Santa Fe <x Landhafer the presence of approximately one
susceptible plant in 16 in all tests suggested that the single factor pairs for
resistance in both these varieties revealed in their cross2s with susceptible
varieties were genetically independent. Thus 7/16 of the population were
expected to be resistant similar to the parents giving a “;” to ‘‘1=” reaction
type, and one-quarter semi-resistant resulting from the heterozygous effect
of each incompletely dominant gene singly. Doubly heterozygous plants on
this hypothesis would comprise one quarter of the F, population and their
reaction type might be expected to be resistant corresponding to that shown
by the F, seedlings which varied from ‘“ ;”’ to “ 1—in”’ according to particular
tests. However, when tested to this predicted 11 resistant : four semi-
resistant : one susceptible seedling ratio in the F, generation it was obvious that
in two out of the four cases the number of semi-resistant plants was
considerably in excess of that calculated, indicating that some of the doubly
heterozygous class had a higher reaction than that predicted, due probably either
to environmental effects or to the segregation and action of modifying
genetic factors.

In this cross also, whilst individual tests showed good agreement with the
predicted 15 (resistant + semi-resistant) : one susceptible F, seedling, there was
a small, though not statistically significant, excess number of susceptible plants
in all cases and, due to this, the total of all tests did not show a good fit to a
dihybrid 15:1 ratio, the P value being 0-:02—0-01. It is difficult to explain
this result since, if any linkage were envisaged between the genes in Landhafer
and Santa Fe, they would be expected to be present in the repulsion phase,
resulting in a deficiency rather than an excess of susceptible plants. In this
cross a few F, seedlings of an intermediate (‘‘ 2— ”’) reaction type were observed
and for statistical tests these were grouped with the semi-resistant class.

In F, segregates of the cross Landhafer x Trispernia, the factor pair in
Trispernia, in the absence of that in Landhafer, in the heterozygous condition
was expected to show an intermediate reaction type varying from ‘‘2” to
‘3c’ from the behaviour of Trispernia in crosses with susceptible varieties ;
only those seedlings homozygous for the Landhafer factor pair were expected
to give a resistant (‘‘;’’) reaction type. Thus the expected F, ratio was four
resistant : nine semi-resistant : two intermediate: one susceptible plant. The
deviations were not significant at the five per cent level. However, by grouping
the two middle classes and comparing with a four resistant : 11 intermediate : one
susceptible plant ratio, a better fit was obtained statistically, the P value
being 0-5—0-3 compared with 0-1—0-05.

Since in the cross Landhafer x Victoria the F, behaviour showed epistasis
or partial epistasis of the Landhafer reaction type, according to the particular
race employed, three-quarters of the F, seedlings in this cross were expected

134 STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, TI

to show the homozygous or heterozygous reaction type of the Landhafer gene
similar to that in its crosses with susceptible varieties. One-quarter would
therefore be expected to show a “;” to ‘“1—” reaction type and one-half a
““ 1” reaction type or one approximating this at normal temperatures (below
75° F). The remaining 25 per cent was expected to show segregation for the
Victoria type crown rust resistance in the ratio of 71-9 per cent resistant (‘ In ”’
to “2” reaction types): 28-1 per cent susceptible (Upadhyaya and Baker,
1960). Due to the presence of the large number of segregating factors involved
in the cross, the distinction between the ““;1—” and “;1” reaction types
was not clear cut and the two classes were combined for statistical calculations.
Hence the expected F, ratio in this cross was 75 per cent ‘‘;” to “1” reaction
types (due to Landhafer), 18 per cent ‘‘1n” to “2” (due to Victoria), and
seven per cent susceptible. Results with three races separately and the
combined total agreed well with this hypothesis. The tests involving race
237-4 were conducted at temperatures between 75° and 85° F. At these
temperatures it was observed previously that the Landhafer factor alone in
the heterozygous condition gave plants of an intermediate (“‘2” to ‘ 3—e”’’)
reaction type. In this instance the expected ratio was 25 per cent resistant
(“;” to ‘“*1” reaction types): 68 per cent intermediate (“2—” to “3c”
reaction types): seven per cent fully susceptible, and the observed results
agreed satisfactorily with this hypothesis.

In the cross Santa Fe x Ukraine 570 and 147 seedlings respectively were
tested to races 237-4 and 237 and no susceptible segregates were observed,
indicating that the single factor pairs in each case were allelomorphie or closely
linked. Although to races to which both were resistant the reaction type was
very similiar in both varieties, the factors were not identical since that in
Ukraine conditioned resistance to fewer races. It has been shown previously
that the resistance of Santa Fe to a large number of races to which it is resistant
is due to the same factor pair (Upadhyaya and Baker, 1962b).

Results involving F, seedling segregation in crosses of Santa Fe with Tris-
pernia and Victoria are presented in Table 3. In the total F, population of
840 seedlings involving tests with four races, no susceptible segregates were
observed in the cross Santa Fe x Trispernia. In tests involving race 259,
13 plants were noted with a slightly higher reaction type than Trispernia but
this may have been due to segregation of modifying genes or to high temperature
effects on reaction type. The absence of susceptible segregates in I’, indicated
that the factors in Santa Fe and Trispernia were allelic or closely linked. The
genes were considered to be distinct in view of the consistently higher reaction
type of Trispernia. Hence the single factor pairs in each of the three varieties
Santa Fe, Ukraine and Trispernia conditioning seedling resistance were con-
sidered to constitute an allelic series.

Since F, seedlings in the cross Santa Fe x Victoria were tested only with
races 259 and 286, to neither of which were both parents resistant, no evidence
was available on epistastis of reaction types. However, following the observa-
tions on a Victoria x Landhafer cross where the Victoria reaction type with
characteristic necrosis was hypostatic, the hypothesis of 25 per cent Santa Fe
reaction types, 50 per cent intermediate between Santa Fe and Victoria (resistant
reaction types with no necrosis), 18 per cent Victoria reaction types (resistant
with necrosis) and seven per cent susceptible segregates in F, was adopted.
On this hypothesis only in tests involving race 237-4 was a good statistical
fit obtained. In the other two tests the number of Santa Fe types was too
few and the Victoria reaction types in excess of that expected. A satisfactory
statistical fit was obtained in these cases only when the resistant classes of
reactions were grouped and the hypothesis of 93 resistant : seven susceptible
F, plants adopted. This suggested that the Santa Fe reaction type was not
completely epistatic to that in Victoria or that segregating modifying genes
from one or both parents were operative.

135

Y. M. UPADHYAYA AND E. P. BAKER

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136 STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, II

(uu) Adult plant tests

Results of F, segregation in the adult piant stage under field conditions
are presented in Table 4.

It was shown in a previous paper (Upadhyaya and Baker, 19626) from
data on the cross Burke x Landhafer that, under field conditions, Landhafer
possessed an additional factor (recessive in action) for crown rust resistance
operative in the adult plant stage only. With the operation of three independent
factor pairs conditioning adult plant resistance (one recessive in action) in crosses
involving Landhafer with Santa Fe or Trispernia, a ratio of 61 resistant : three
susceptible plants was expected for F, field segregation. In the cross Santa
Fe x Ukraine the expected ratio was 249 resistant : seven susceptible with four
factors involved (three from Ukraine, two acting in dominant complementary

TABLE 4

F, segregation for adult plant field reaction to Crown Rust in certain crosses involving the resistant
oat varieties Landhafer and Santa Fe with other resistant varieties

Adult plant field reactions Expected
Cross Total ratio P value
I R MR MS-S
Santa Fe x
Landhafer of 343 96 10 23 472 61:3 0:9-0:8
(22-1)
Landhafer x
Trispernia as 184 50 19 16 249 61:3 0-3-0-2
(11-7)
Victoria, x
Landhafer xe 201 3 — 2 206 98-88: 1-12 0:9-0-8
(2-3)
Santa Fe x
Victoria .. a 207 9 28 14 258 94-05 : 5:95 0-8-0-7
(15-4)
Santa Fe x
Ukraine .. ~°.. 56 6 5 2 69 249: 7 0-7-0°-5
(1-5)
Santa Fe x
Trispernia! O16 — 453 —- 0 453 all R —

1 No separate classification for the different types of resistance carried out
(Expected values in brackets.)
I= Immune, R= Resistant, MR = Moderately resistant, MS = Moderately susceptible,
S = Susceptible.

fashion and one from Santa Fe). No susceptible segregates were expected in
F, in the cross Santa Fe x Ukraine since the same factors conditioned both
seedling and adult plant resistances in each case and the seedling resistances
were previously indicated to be allelic (or closely linked). The segregations in
crosses involving the variety Victoria were based upon the presence of four
factor pairs in this variety, Ve,Vc, and IVc,I Vc, linked with ten per cent
recombination, and two independent factor pairs, Ve,Ve, and Vc3,Ve, giving in
crosses with susceptible varieties 5-95 per cent susceptible adult plants
(Upadhyaya and Baker, 1960). With the two factors for adult plant resistance
contributed by Landhafer the percentage of susceptible F, plants would be
expected to be 1-12 in the cross Victoria x Landhafer.

In all the above cases in F, adult plant segregation good agreement
statistically between observed and expected results was observed on the basis
of these hypotheses presented. In the cross Santa Fe x Victoria the percentage
of susceptible plants would be expected to be 1:49 with the four factors from
Victoria and one from Santa Fe. Hence in the 258 plants tested, approximately
four susceptible plants would have been expected. This deviates markedly

Y. M. UPADHYAYA AND E. P. BAKER 137

from the 14 observed. The results in the cases of this cross were best explained

on the operation of only one of the two factors (Vc, or Ve3) conditioning adult
plant resistance in Victoria, on which basis 15:4 susceptible plants would have
been expected.

These results confirmed the operation of the following factors conditioning
adult plant resistance to crown rust: Two factors in Landhafer (one recessive
in inheritance).—One factor each in Santa Fe and Trispernia, the factors being
allelic and identical with those conferring seedling resistance.—Three factors
in Ukraine, two acting in complementary dominant fashion.—One factor in
Victoria linked with a dominant inhibitor gene and one or two additional adult
plant factors according to the particular cross involved.

F, segregation

Seedling tests of F, progenies from seedling classified F, plants were con-
ducted by taking representative samples from the major F, reaction categories.
The expected F, behaviour in the various crosses was as follows :

F, behaviour

Cross
Resistant Segregating Susceptible
Santa Fe x Landhafer 7 8 1
Landhafer x Trispernia 7 8 I
Santa Fe x Trispernia All resistant
Santa Fe x Ukraine All resistant

Victoria x Landhafer |

Victoria x Santa Fe f{ 40° 4% 52-6% 7:0%

These expectancies could be further categorized and subdivided in certain
cases thus :

Landhafer x Trispernia

Resistant 7, comprising four homozygous for the partially epistatic
Landhafer resistance (‘‘ 1=: ” reaction type) denoted by Ld, one homozygous
for Trispernia resistance (‘1+ ”’ reaction type) denoted by Tr, and two
segregating for Landhafer and Trispernia reaction types (denoted by Ld: Tr),
the hypostatic Trispernia gene appearing due to the heterozygous state of the
Landhafer gene in tnis particular genotype.

Segregating 8, comprising four segregating for the Landhater reaction
type (preponderant), Trispernia reaction type and susceptible plants, two
secregating for Landhafer reaction type and susceptibility, the designation
Ld (fr):S being used to represent both categories, and two segregating for
Trispernia reaction type and susceptibility (designated Tr: 8S).

Santa Fe x Trispernia

All resistant, comprising one homozygous for the Santa Fe resistance
(“; to ““1= ” reaction type), two segregating for the Santa Fe and Trispernia
resistances (‘41+ ” reaction type), designated as S.F.: Tr, and one homozygous
for Trispernia resistance.

Victoria <x Landhafer

Resistant 40-4 per cent, comprising 25-0 per cent homozygous for the
epistatic Landhafer resistance (denoted by Ld), 10-3 per cent segregating for
the Landhafer and Victoria reaction types (symbolized as Ld: Vc), and 5-1 per
cent homozygous for the Victoria resistance (denoted by Ve).

138 STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, III

Segregating 52-6 per cent, comprising 39-8 per cent segregating for Land-
hafer resistance and susceptibility, or Landhafer and Victoria resistances and
susceptibility (designated as Ld (VYe):S), and 12-8 per cent segregating for
Victoria reaction type and susceptibility (denoted as Ve:S).

Susceptible 7-0 per cent, comprising those segregating for susceptibility
and a low proportion of Victoria type resistant plants (symbolized as S: Ve)
and those lines homozygous susceptible (denoted as 8).

Victoria x Santa Fe

Similar categories to those in the Victoria x Landhafer cross.

In the cross Santa Fe x Ukraine no segregation was noted in tests
involving a mixture of races 237 and 237-4. Table 5 presents the F, data
relevant to Santa Fe in crosses with Wandhafer and Trispernia and shows good
agreement between observed and expected results.

TABLE 5

Seedling behaviour of F, lines in crosses involving the resistant oat variety Santa Fe with the
resistant varieties Landhafer and Trisperma tested with Race 203 of Crown Rust

F, Behaviour

Cross Total P value
Res. Seg. Sus.
Santa Fe x Landhafer 105 131 16 252 0:9-0-8
(110-3) (126-0) (15-8)
Homo.S.F. Seg.S.F.: Tr. Homo.Tr.
Santa Fe x Trispernia 28 51 28 107 0-9-0-8
(26-8) (53-5) (26-8)?

1— Expected ratio 7: 8:1 resp.

2— Expected ratio 1:2:1 resp.
Res. = Resistant, Seg. = Segregating, Sus. = Susceptible. Homo. S.F.= Homozygous
for Santa Fe reaction type (;). Seg. S.F.: Tr. = Segregating for Santa Fe reaction
type (;) and Trispernia reaction type (1+). Homo. Tr. = Homozygous for Trispernia
reaction type (1+).
(Expected values in brackets).

Data involving the three other crosses studied, Landhafer x Trispernia
and Victoria in its crosses with Landhafer and Santa Fe, are p:esented in
Table 6. Comparison of expected and observed results within the previously
indicated subclasses of two of the three F., categories, viz. homozygous resistant
and segregating, in these crosses are included in this table. There was good
Statistical agreement between observed and expected results, except in the
cross Santa Fe x Victoria where the agreement was satisfactory only when
the major classes (resistant, segregating and susceptible) were considered but
not further subdivided.

When the F, classification for reaction types was based on F, breeding
behaviour in the three crosses, 75 per cent would be expected to be Landhafer
or Santa Fe types, 18-75 per cent Trispernia or 18 per cent Victoria types, and
the remainder susceptible. From F, data the appropriate factors were separated
as follows where the observed and expected numbers (in brackets) are compared :

Ld. or 8.F. Tr. or Ve. Susceptible

types types
Landhafer x Trispernia 135(136-5) 34(34:2) 13(11-3)
Victoria x Landhafer 63(62-3) 15(14°8) 5(5-8)

Santa Fe x Victoria 293(320-0) 104(76-2) 29(29-8)

139

Y. M. UPADHYAYA AND E. P. BAKER

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140 STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, TI

The observed frequencies in the first two crosses showed no significant
deviations from those expected. In the cross Santa Fe x Victoria, however,
it was clear that an excessively large number of lines showed the Victoria
reaction type on the hypothesis adopted. Reference has already been made
to similar conclusions based on F, data. Unfortunately, as previously indicated,
no studies of F', reaction type where epistasis could be directly assessed were
carried out in this cross in tests involving races to which both parents were
resistant. The greater number of F, plants and of F, lines showing the Santa
Fe type of resistance suggested the epistatic behaviour of the Santa Fe reaction
type over that of Victoria. The general observation in all crosses in general,
that the lower reaction type was epistatic, supports this hypothesis. Certain
genotypes, however, involving the variety Victoria, seemed to have inhibited
the action of the Santa Fe gene. Such behaviour was evident in studies on
correlated tests of identical F, lines involving race 226 (to which both parents
were resistant) and race 286 (to which only Victoria was resistant). The data
are included in Table 12.

Relationship of seedling reaction types to different races and to adult plant field
reactions
(i) F, seedling vs. F, seedling
This correlation was studied only in the cross Landhafer x Santa Fe,
where 321 F, plants were first classified for primary leaf reaction type to race
237 and the leaves subsequently cut off and of these 207 were then inoculated
at the secondary leaf stage with race 226. Perfect correlation of reaction
types was observed in this test.

(ii) F, seedling vs. F, adult plant

This association was studied in all crosses except Victoria x Landhafer.
Classified seedlings were transplanted and tested in the field for subsequent
adult plant behaviour. The resistant class was expected to maintain its
resistance at the adult plant stage in Landhafer crosses but, due to the operation
of an additional factor conditioning adult plant resistance in this variety, the
seedling susceptible class was expected to produce some resistant adult plants.
In the cross Santa Fe x Landhafer, 86 seedlings tested maintained their
resistance ; in the cross Landhafer x Trispernia only one plant giving an
intermediate type of reaction for the heterozygous condition of the Trispernia
gene gave a susceptible field reaction, the other 193 seedlings maintaining their
resistance as adult plants. Jn the susceptible class one plant out of nine in
the former cross and five out of thirteen in the latter cross remained fully
susceptible, the segregation thereby conforming within approved statistical
limits to the expected 3 resistant: 1 susceptible ratio. These resistant
adult plants, in both cases from the susceptible seedling group, varied in reaction
type, one being immune, six resistant, and one moderately resistant in the
cross Santa Fe x Landhafer, the corresponding figures in the cross Land-
hafer x Trispernia being three, four and one, thus showing that the factor
conditioning only adult plant resistance in Landhafer was incompletely
dominant in inheritance.

Since the single factor pairs conditioning seedling as well as adult plant
resistance in both Santa Fe and Trispernia were previously shown to be allelic
(or closely linked) in the seedling stage, no susceptible F, adult plant segregates
were expected in the cross between these two varieties ; none were onpserved
among 192 plants tested.

In tests involving race 226 (to which Ukraine was susceptible) seedling
resistant F, plants in the cross Santa Fe x Ukraine were expected to remain
resistant as adult plants due to the influence of the Santa Fe gene, and susceptible
seedlings were expected to show a 57 resistant: 7 susceptible adult plant.
segregation due to the action of the three factors (two acting in complementary

Y. M. UPADHYAYA AND E. P. BAKER 141

fashion) conditioning adult plant resistance in Ukraine. In the _ cross
Victoria x Santa Fe in tests involving races to which both parents were
resistant, seedlings possessing the Santa Fe reaction type (‘‘;”’ to ‘“ 1=”’) were
expected to remain resistant as adult plants. The group of seedlings possessing
the Victoria reaction type (‘‘ In ’’) and those susceptible were expected to show
some susceptible and resistant adult plants respectively. Previously cited F,
data indicated that in this cross only one factor conditioning solely adult plant
resistance in Victoria was operative. This factor, when linked with two
complementary factors for seedling resistance, was expected to show the
following relationship between seedling and adult plant behaviour in F,:

Seedling reaction types
Adult plant reactions

Santa Fe Victoria Susceptible
type type
Immune—Moderately -——— ant
resistant ae ihe 100% 66:9% 9-5%
Moderately susceptible
—Susceptible oe — 5:0% 18-6%

Data relating to crosses of Santa Fe with Ukraine and Victoria are presented
in Table 7. In all cases good statistical fits to the expected results were clearly
obtained and confirmed the operation of three factors in Ukraine and one major
factor in Victoria for adult plant resistance.

TABLE 7

Relationship between seedling reaction types and adult plant reactions to Crown Rust of F, plants
am crosses of the oat variety Santa Fe with the varieties Ukraine and Victoria

Adult plant Seedling reaction types (Race 226)!
reactions
Santa Fe x Ukraine Santa Fe x Victoria
2 l=, 1 3-4 2 l=, I In 2-—n 3-4
Immune 59 7 — 34 15 47 — 1
Resistant 5 3 3 — — 2 — 3
Mod. Res. 3 2 11 = — — Il —
Total 67 12 14 34 15 29 11 4
(12-7) (35-5) (5-0)
Mod. Sus. and — — 1] —= — — i 8
Susceptible eres ae
(2-3) (2-6) (9-9)
P value 0-7-0: 5? 0:7-0:5

1 Seedling reaction types corrected from F, behaviour.

* Yates’ correction factor applied for small numbers.
Mod. Res. = Moderately resistant, Mod. Sus. = Moderately susceptible.
(Expected values in brackets.)

(iii) F, seedling vs. F, seedling
The F, behaviour of representative samples from each F, class of reaction
type was studied in all crosses except Santa Fe x Ukraine. In certain cases
the particular race to which the F, was tested was used in a mixture with
certain other races for F, tests. In other cases, a mixture not involving the
particular race used in F, was utilized, whilst in one test an identical strain

142 STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, IT

(race 226) was used in tests for both generations. The data pertinent to these
studies are set out in Table 8 and were designed to study the postulated
breeding behaviour of F, genotypes based on reaction types and to investigate
if the same factors were operative against all races.

In the cross Santa Fe x Landhafer one plant each from the highly
resistant class (‘‘;”’ reaction type) and moderately resistant class (‘2—”
reaction type) of F, segregates gave susceptible progenies. The latter plant
would have been expected to be heterozygous on reaction type and hence
segregate in F,. Classification as homozygous susceptible may have been
erroneous, due to the chance absence of a resistant plant, but this would
be highly improbable statistically in the sample of approximately 25 plants
tested in each F, line. One moderately resistant F, plant (‘ 2--” reaction
type) also gave a homozygous susceptible line, but this could occur statistically
at a relatively nigh probability level as pointed out by Upadhyaya and Baker
(1962a). Except for these instances, the first of which was almost certainly
due to an error in classification, labelling or transplanting, all other plants from
the different F, reaction classes behaved as expected, indicating correlated
inheritance to all races with which they were tested, since no mixed reaction
types were observed on the same leaf by the use of inoculum comprising a
mixture of races.

In the cross Santa Fe x Trispernia there was also good agreement between
observed and expected results. The operation of certain modifying genes was
indicated, however, since from the “1” reaction type in F,, 12 plants gave
homozygous highly resistant progenies similar to Santa Fe (“;” to “1=”
reaction type). In the F, classification also certain seedlings had shown
reaction types higher than Trispernia; the progenies of these plants showed
reaction types similar to Trispernia at normal temperatures of about 75°F,
except for one plant which showed segregation for the Santa Fe and Trispernia
reaction types.

In the cross Victoria x Landhafer also good agreement between the F,
reactions to race 237 and F, behaviour to a mixture of races 226, 237 and 237-4
was shown. Onlv four lines—one homozygous resistant for the Victoria
reaction type and three segregating for Victoria type resistance and susceptibility
—were observed from F, plants in the ““1— ” to “1” reaction type category
intermediate between that of Landhafer and Victoria and expected to segregate
for the Landhafer reaction type and susceptibility. This discrepancy may
have been due to difficulty in distinguishing the necrotic reaction type associated
with the Victoria type of resistance on the basis of a single F, plant. Similarly,
a small number of discrepancies were observed in the cross Landhafer x Tris-
pernia. The difficulty in distinguishing a ‘*‘ 2” reaction type with the associated
‘‘ green island’ from a “3” type with the pustule surrounded by chlorosis
resulted in three plants assigned to the former class giving homozygous susceptible
progenies and two to the latter class producing segregating progenies. Apart
from these instances it was clear that the same factors in the two varieties
conditioned resistance to the four races 226, 237, 237-4 and 259.

In the cross Santa Fe x Victoria, F, plants classified for reaction
type to race 203 were tested for their progeny reactions against races 226 and
237 separately. Reactions in I*, were corrected, taking into account behaviour
to both races since small numbers of seedlings were tested to each race separately.
For example, if an F, line was apparently homozygous resistant to one race
in the small sample tested, but segregating to the other, its behaviour was
indicated as segregating. In another test, F, and F, reaction types were noted
when both were tested against the same race—race 226. Agreement between
observed and expected results was not completely satisfactory in this cross and
discrepancies existed in certain instances. From among the 85 F, plants
classified as possessing the Santa Fe reaction type (‘‘;”’), three were found to

Y. M. UPADHYAYA AND BE. P. BAKER 143

TABLE 8

F, behaviour of F, plants classified for seedling reaction type to Crown Rust in crosses involving
the resistant oat varieties Landhafer and Santa Fe

F, Reaction

types of various F, reactions to race(s) Total
erosses to
different races 226, 237, 237-4 and 259
Race 259 Res. Seg. Sus.
Santa Fe x
Landhafer 35 18 1 54
1 1 23 — 24
2— — 3 1 4
3-4 — — 7 g

226, 237, 237-4 and 259

Soltc Slt, eWbee Ube:

Santa Fe x
Trispernia ; 13 12 — 25
1 12 36 2 50
2- —_ 6 20 26
2 — 1 5 6
Itel, ii@leibe, “he, ibel (te) sS> “irs S. S.
Landhafer x ; 46 13 — 1l = = 70
Trispernia 1-2= 1 7 1 46 1 — 56
2- — 4 2 6 3 — 15
2; — — 2 3 21 3 29
3-4 — — — — 2 12 14
226, 237 and 237-4
Race 237
Ld. id.:Ve. Ve. Ld.(Ve.):S. Ve.:S. S.
Victoria xX 3 3 1 — 3 — — 7
Landhafer J--l — 1 1 8 3 — 13
In-2n — — 4 — 4 — 8
3n — — — — 3 — 3
3-4 — — — — — 7(2 S.: Ve.) 7
226 and 237
Race 203 ———_—_——
S.F. S.F.: Ve. Ve. S.F.(Ve.):S. Ve.:S. S.
Santa Fe x g 32 10 I 2 1 — 46
Victoria 1 Il 7 2 43 — — 63
1+, 3-¢ — — — 15 2 — 17
In, 2—n — 6 19 19 26 — 70
2n, 3n — — 1 2 8 — 11
3-4 — — — — 1 16(8 S.Ve.) IV
Race 226 226
g 11 9 2 14 2 1 39
1 — 2 — 13 — — 15
In- 2-n — — 5 — 20 — 25
2n— 3n — — 2 ] 4 1 8
3-4 — — — — 1 10(3 S.: Ve.) 11
Reaction types indicated as follows :—S.F. = Homozygous ;—1= ; Tr. = Homozygous
1+; Ld.= Homozygous;-1= ; Ve.= Homozygous In; S.= Homozygous 3-4 re-

action types; S.F.:Tr., etc. = segregating for 8.F. and Tr. reaction types etc.; Ld.
(Tr.):S., ete. = segregating for Ld. reaction type and susceptibility or Ld. and Tr.
reaction types and susceptibility, etc.; S.: Ve. = segregating for Ve. reaction type and
susceptibility with preponderance of susceptible plants; Res. = Homozygous resistant ;
Seg. = Segregating ; Sus. = Homozygous susceptible.

144 STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, II

be homozygous for the Victoria reaction type (‘‘1n’’), three segregated for
this reaction type and susceptibility, and one was homozygous for susceptibility.
Of 78 plants classified as showing a ‘‘ 1” reaction type, intermediate between
Santa Fe and Victoria, two were homozygous for the Victoria type of resistance,
whilst two segregated for the Victoria reaction type. A total of 28 plants from

TABLE 9

Ff, seedling behaviour to certain Crown Rust races of F', plants classified for adult plant reaction in
crosses involving the resistant oat varieties Landhafer and Santa Fe

F, Field F, Seedling behaviour
Cross reaction or reaction type(s) Total P value
Res. Seg. Sus.

Races 226, 237, 237-4 and 259

Santa Fe x I 44 42 — 86

Landhafer R 1 41 — 42
MR — 7 3 10
MS-S — 4 8 12

Races 226 and 226-2

Santa Fe x I 14 26 7 47
Ukraine R 1 3 2 6
MR — 1 4. 5
Total 15(14:8) 30(29-9) 13(13-3) 58 0:-9-0:8
MS-S — — 2(2-0) 2 =
Race 226
‘S.F. S.F.:Ve. S.F.(Ve.):S. V.c Ve.:S. Ss.
Santa Fe x ;
Victoria I 13 11 27 5 4 — 60
R 1 — 5 1 1 1 9
MR — = 5 3 14 61 28
MS-S — — 1 — 1 7 9
Race 203
Ld. ld.:Ve. Ld.(Ve.):S. Ve. Ve.:S. Ss.
Victoria x
Landhafer I 20 10 28 8 8 4 78
R — — 3 — — — 3
MS = — — — — 2
(1 S.: Ve.)

1 Four lines segregating with preponderance of susceptible plants.
2 Three lines segregating with preponderance of susceptible plants.
For interpretation of symbols S.F., ete., see Table 8.
(Expected values in brackets.)
= Immune, R= Resistant, MR = Moderately resistant, MS = Moderately susceptible,
S = Susceptible.

the 114 in the class showing the necrosis associated with the Victoria reaction
type and classified as having ‘in’, ‘°2—n”’, “2n” or ““3n”’ reaction types
Showed segregation for the Santa Fe genotype, whilst one F, plant from the
‘“2n” to “* 3n” reaction type category was homozygous susceptible. Most of
these aberrant instances were those involving F, tests with race 263 and a closer
association of reaction types between F, and F, seedlings was evident when
race 226 was used in the F, generation.

Y. M. UPADHYAYA AND E. P. BAKER 145

The four plants under the F, ‘“;” reaction type category, producing
susceptible progenies or segregating for Victoria type resistance and susceptibility,
were almost certainly misclassifications. The discrepancies in other cases
indicated that the F, classification was affected by modifying genetic factors
or environmental variations. Further evidence for this was shown since
11 lines were homozygous resistant for the Santa Fe reaction type from F,
plants classified as exhibiting a ‘‘1” or various intermediate reaction types.
Later reported. correlated KF, studies involving different races produced
unexpected results and, as indicated in Table 8, 28 F, plants out of 114 classified
for the Victoria reaction type gave F, lines segregating for the Santa Fe factor
as previously indicated.

(iv) F, adult plant vs. IF, seedling

In this analysis the progenies of plants classified for adult plant reaction
in the field were studied for seedling behaviour and the data are presented in
Table 9.

This analysis involved tests on F, seedlings which had been initially classified
for seedling reaction type and subsequent adult plant behaviour. In the cross
Santa Fe »~ Landhafer three moderately resistant plants gave fully susceptible
progenies. These presumably carried the recessive factor conditioning moderate
resistance in the adult plant stage in Landhafer. Since these would be expected
to comprise 3/64 of the population the number expected was 7-0. The
probability of chance deviation was 0:2—0-1 for this result. Segregating
progenies from the moderately resistant and moderately susceptible to
susceptible classes of F, adult plants indicated that the single factor conditioning
seedling as well as adult plant resistance in Santa Fe and/or Landhafer was
incompletely dominant in the latter stage.

In the F, generation of the cross Santa Fe x Ukraine, the adult plant
segregation conformed to a ratio of 249 resistant: 7 susceptible. Of these
249 resistant plants 64 were expected to produce homozygous resistant, 128
segregating and 57 homozygous susceptible lines. The progenies of the 58
resistant F, plants conformed to this hypothesis statistically, whilst two
susceptible I’, plants gave susceptible progenies as expected.

In tne cross Santa Fe = Victoria, all the F, lines homozygous or
heterozygous for the Santa Fe reaction type were expected to be derived
from the resistant classes of F, plants. Of 63 lines of this type only one was
derived from a susceptible plant and was probably an error in F, classification
labelling. The expected ratio was one homo7ygous:2 segregating progenies
for this type of resistance. Twenty-five homozygous resistant lines were
observed, giving a P value of 0-5—0-2 in the sample of 63. The remaining
F, plants would depend for their resistance on the Victoria genes or would be
susceptible. The expected behaviour of these classes in F,; was as follows on
the basis of the operation of one of the adult plant resistance factors in Victoria :

F, seedling behaviour (in percentages)

F, adult
plant
reaction Resistant Segregating Susceptible (including
(Victoria type) (Victoria type : segregating with pre-
susceptible) ponderance of suscept-
ible plants)
Resistant 20-3 46-5 9-5= 76-3
Susceptible 0-2 4-9 18-6= 23-7
Total 20-5 51-4 28-1

146 STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, OI

Grouping together the field resistant F, groups (immune, resistant and
moderately resistant) the observed and expected frequencies (in brackets)
corresponding to the above table were as follows :

F, adult plant F, seedling behaviour
reaction
Resistant Segregating Susceptible
Resistant .. ie 9(8-7) 19(20-0) 7(4:1)

Susceptible .. ee 0(0-1) 1 (2-1) 7(8:0)

This indicated good agreement between observed and expected results.

In the cross Victoria <x Landhafer, 206 F, plants were classified for adult
plant reaction and the results are as previously set out and explained in relation
to the data of Table 4. The progenies of the three resistant plants and 78 of
the immune reaction class were tested. Of these 81 plants, four gave susceptible
F, lines. From the behaviour of the Victoria genotype one-fourth of the
resistant F, class was expected to produce susceptible F lines, whilst F; behaviour
due to the Landhafer genotype with a second factor acting solely at the adult
plant stage was expected to give one-fifth of lines susceptible from the field
resistant F, class. Hence, on combining the two genotypes one-twentieth of
resistant F', adult plants were expected to give susceptible F, seedling progenies
in a cross between the two varieties. The four such plants observed showed
perfect agreement with the number expected.

Confirmation on the operation of the factor Vc, and its inhibitor was
obtained since several lines in the two crosses involving Victoria showed
segregation for the Victoria type of resistance with a preponderance of susceptible
segregates. The behaviour of these subclasses has already been dealt with in
the F, studies.

TaBLE 10

Correlation of F., seedling behaviour to different races of Crown Rust in crosses involving the
resistant oat variety Landhafer with certain other resistant varieties

F, behaviour

Cross F, behaviour to race -
Res. Seg. Sus. (inel. 8. : Ve.)

203 Race composite 226, 237, 237-4, 259
Santa Fe x Res. 45 — —
Landhafer Seg. — 49 —-
Sus. — — 11
226 Race composite 203, 237, 237-4, 259
Landhafer x Res. 78 i —_
Trispernia Seg. 1 83 —
Sus. — — 14

203 Race composite 226, 237, 237-4
Victoria x Res. 48 1 —
Landhafer Seg. — 61 —
Sus.(imel. 8. : Ve.) — 1 161

1 One line segregated S.: Ve. (Susceptible and Victoria reaction type plants, with suscept-
ible types preponderant) in both tests, two segregated S.: Ve. to race composite only and
one segregated S.: Ve. to race 203 only.

Seg. = Segregating, Res. = Resistant, Sus. = Susceptible
S.: Ve = Segregating for susceptible reaction type and Victoria reaction type. with
preponderance of the former.

Y. M. UPADHYAYA AND E. P. BAKER 147

(v) EF, seedling vs. F, seedling
The relationship of I’, reactions in crosses involving Landhafer and Santa
Fe are presented in Tables 10, J1 and 12. The same seedlings were tested
with one race on the primary leaf and then either with a mixture of races or
a second race on the secondary leaves. From Table 10 which involves data

TaBLeE 11

Correlation of seedling behaviour of F, lines to Crown Rust Race 203 and a composite of Races 226,
237, 237-4 and 259 in a cross between the resistant oat varieties Santa Fe and Trispernia

F, reactions to F, behaviour to composite of Races
Race 203 =

S.F. Sold, 3 dle. cine

S.F. 11(10-9) 2(0-0) ——
Sol, oe, YL 13) : 40) : —
Tr. :S.F1f ee) eo) an)
ir. —="/(16) 6(3-3) 26
y’ values 1:04, P= 0-7-0-5; 2-01, P= 0-5-0-3

1Tr.:S.F.= Segregatimg for 14+ and ; —1 = reaction types, with the former preponderant.
For interpretation of symbols §8.F., etc. see Table 8.
(Expected values in brackets.)

TABLE 12

Correlation of seedling behaviour of F', lines to Crown Rust Race 226 and Races 237, 259 and 286
respectively in a cross between the oat varieties Santa Fe and Victoria

F, behaviour F; behaviour to Race 226
to Race Total
Res. Seg. Sus.
237 ~=Res. 59 6 oe | 65
Seg. 8 69 77
Sus. = 1 22 23
SIMA Sites We, We, Sale) eS. Wess Sowell. S32 We;))
259 ~=Res. (S.F.) 11 — = == = — 11
Seg. — 8 24 — 1 33
Sus. = a 6 — 21 + 31
286 Res. (Vc.) 8 Il 21 — — —- 40
(17-6) (22-0)
Seg. 19 5 1 4] 24 — 90
(29-3) (0-0) (41-4) (25-0)
Sus.? 15 5 — 23 1 — 44
(16-1) (0-0) (22-6) (0-0)
x? values 2-08 — 0-011 nee ==
P= 0-5-0:3 0-95-0-9

1 For interpretation of symbols S.F., etc. see Table 8.
2 Hight limes showed S.: Ve. behaviour.
(Expected values im brackets.)
Res. = Homozygous resistant, Seg. = Segregating, Sus. = Homozygous susceptible.

relating to Landhater crosses in such tests it was clear that there was complete
agreement in reaction types in the cross Landhafer x Santa Fe. Discrepancies
in the cross Landhafer x Trispernia were almost certainly due to delayed
germination of some seeds in tests involving race 203. Reactions to race 203
and to the mixture of races 226, 237 and 237-—+ were correlated in the cross

148 STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, III

Victoria x Landhafer, except for two lines which were found to be segregating
to the mixture of races but which were resistant and susceptible respectively
to race 203. These discrepancies were probably due to errors in classification
and the data indicated that the same factors conditioning seedling resistance
were responsible for resistance to all the races to which both parents were
resistant.

In the cross Santa Fe x Trispernia, F, lines were first tested to race 203
and then to a mixture of races. From the reactions presented in Table 11,
14 lines which segregated when tested with race 203 were fully resistant to
the mixture of races. Similarly six lines which were homozygous for the
Trispernia reaction type to race 203, gave segregating reactions to the race
mixture.

Again, certain lines showed a preponderance of Trispernia reaction type
plants over Santa Fe types in tests with race 203. These facts indicated the
operation of certain modifying factors, which inhibited the expression of the
Santa Ié gene. On the assumption of a pair of such complementary factors
the expected behaviour of the F, Santa Fe reaction class to the mixture of
races was seven homozygous lines homozygous for the Santa Fe reaction
type, 8 segregating for Santa Fe and Trispernia reaction types, and one homozygous
for the Trispernia reaction type. All Trispernia reaction type I’, plants were
expected to produce lines homozygous for such reaction type. The deviations
were not statistically significant on this hypothesis. Minor modifications were
observed in tests against the mixture of races both in F, and F, studies. since
some plants in the homozygous Santa Fe lines showed ‘‘ 1= ” and “ ; ” reaction
types on the same leaf.

In the cross Santa Fe x Victoria, 426 F, lines were tested against race
226. On the secondary leaves of 75 lines reactions to race 259, to which Victoria
was susceptible, were recorded ; similarity, on 174 lines reactions to race 286,
to which Santa Fe was susceptible, were noted. In another test 155 lines were
tested against race 257. Relationship of the reaction types in the three cases
is Shown in Table 12.

In tests involving races 226 and 237, to which both Santa Fe and Victoria
were resistant, certain lines were resistant to one race but segregated to the
other. Delayed germination, whereby certain plants escaped infection, may
have been responsible for this difference. There was a general agreement in
the behaviour to the two races indicating the operation for the same factors
for resistance in each case except for one line which gave a segregating reaction
against race 226 but a susceptible reaction to race 237, and this discrepancy
was probably an error in classification. The general agreement in the reactions
indicated the operation of the same factors in each variety against the two races.

In tests involving race 259 it was expected that lines homozygous for the
Santa Fe reaction type would remain resistant, those segregating for the Santa
Fe reaction type would segregate, whilst those of the Victoria type or susceptible
to race 226 would be susceptible to race 259. There was almost complete
agreement with this hypothesis except for one line, an obvious misclassification
error, which was susceptible to race 226 but segregated in tests involving race
259.

In tests against race 286, fron: data included in Table 12, eight lines from
the homozygous Santa Fe reaction type class to race 226 were homozygous
resistant for the Victoria reaction type, thus clearly indicating the epistatic
behaviour of the Santa Fe reaction type in this cross. On this basis the expected
frequencies in the homozygous Santa Ke reaction type F, lines to race 226 were
20-5 per cent homozygous for the Victoria reaction type, 51-4 per cent of lines
segregating for the Victoria reaction type, and 28-1 per cent of lines susceptible
in tests involving race 286. All F,; progenies showing segregation for the Santa
Fe and Victoria reaction types (designated as S.F.: Ve.) in tests involving race
226 were expected to be homozygous for the Victoria reaction type (designated

Y. M. UPADHYAYA AND E. P. BAKER 149

as Ve.) when race 286 was inoculated onto identical seedlings. From Table
12 it was obvious that the class S.F.: Vc. did not behave as expected, nor did
the homozygous Santa Fe (S.F.) category, where there was an excess of
susceptible lines. However, when the two classes S.F. and S.F.: Vc. were
combined, good statistical agreement was obtained. Therefore the 10-3 per
cent S.F.: Ve. class, giving only the Victoria reaction type, was added to the
20-5 per cent of the S.F. category to race 226 expected to benave similarly.
The satisfactory agreement resulting from this procedure indicated the action
of some modifving gene(s) which resulted in the expression of a reaction type
resembling that characteristic of Victoria im certain lines in the S8.F.: Ve.
category to race 226, despite the absence of the genetic factors conditioning
the Victoria type of resistance. This was also apparent since five lines susceptible
to race 286 were obtained from the 8.F.: Ve. class to race 226. In the other
reaction classes to race 226, results from tests involving race 286 showed good
agreement between observed and expected figures except for two lines, one
in the Victoria reaction class and one in the category segregating for the
Victoria type of resistance.

In the cross Victoria x Landnafer 114 lines were tested to both race 259
and race 286, Victoria being susceptible to the former and Jandhafer to the
latter race. When observed and expected results were compared on the basis
of independent segregation, the chi-square value was 3-76 for four d.f., giving
a P value between 0-5 and 0-3, indicating that the factors for resistance in
the two varieties were independent.

DISCUSSION AND CONCLUSIONS

Segregation studies in crosses between certain members of the group of
varieties comprising Landhafer, Santa Fe, Trispernia, Ukraine and Victoria,
all resistant to the prevalent Australian races of crown rust, established certain
facts which substantiated previous findings but provided additional information.

Firstly, the two factors in Landhafer conditioning adult plant resistance,
one of whicn conferred seedling resistance as well, were independent of the
factors in the varieties Santa Fe, Trispernia and Victoria. Indirect evidence
indicated that the factors were also independent of that responsible for seedling
resistance in the variety Ukraine. The reaction type of Landhafer was epistatic
over those of Trispernia and Victoria.

The factor for seedling as well as adult plant resistance in Santa Fe was
independent of the factors in the variety Victoria and epistatic to them. Some
modifying gene(s), however, resulted in the expression of a reaction type similar
to that characteristic of Victoria by suppressing the Santa Fe gene. The Santa
Fe factor was allelic with the factors for seedling resistance in the varieties
Ukraine and Trispernia. The factors in the three varieties were not identical,
since the gene in Santa Fe conditioned resistance to a larger number of races
than did the allele in Ukraine. The allele in Trispernia exhibited a higher
reaction type than that in Santa Fe or Ukraine. The resistance of Santa Fe
was dominant over that of Trispernia in tests against races 226, 237, 237--4
and 250 but with race 203 the Santa Fe gene was inhibited by the action of
a pair of complementary factors, one contributed by each variety, which
resulted in the expression of the Trispernia reaction type in the F, between
the two varieties.

The cross between Santa Fe and Ukraine also revealed the independence
of the factors for adult plant resistance in the variety Ukraine from the alleles
for seedling resistance. Since the factors in the three varieties Santa Fe,
Trispernia and Ukraine conditioning seedling resistance were allelic, even though
segregation was not studied in crosses of Ukraine with Trispernia and Victoria,
nor in the cross Trispernia < Victoria, it could be assumed from the evidence
of other crosses that the factors responsible for seedling resistance in Santa Fe
(as well as T'rispernia and Ukraine) and Victoria were genetically Independent.

150 STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, TT

The independence of the factors conditioning adult plant resistance in Landhafer
and Ukraine, and the independence in turn of the Ukraine and Victoria adult
plant factors, could not, however, be established in the absence of studies
on the appropriate .crosses.

The concept of allelism of the factors conditioning seedling resistance in
the varieties indicated is proposed, although, as indicated by Luig, McWhirter
and Baker (1958) with higher plants where segregating population sizes are
restricted compared with microorganisms, it is technically difficult to establish
closer linkage of less than a few crossover units. However, as reviewed, the
hypothesis of allelism has been proposed by various North American workers,
and is accepted until evidence is advanced to refute it. In this connection
the fact that race 286, described by Baker and Upadhyaya (1955), and first
found in low proportions on adult plants of the variety Trispernia, proved
susceptible on seedlings of Trispernia, Santa Fe and Ukraine as well as Land-
hafer, is of interest.

F, seedling segregation was studied in crosses of Santa Fe with Landhafer,
Ukraine and Trispernia and in the cross Landhafer x Trispernia against race
286, to which all four varieties were susceptible. No complementary gene
action between these varieties was indicated since no F, seedling gave a
resistant reaction. This also excluded the possibility of the operation of any
inhibitor against this race.

These results confirmed certain conclusious proposed by Litzenberger
(1949), Finkner (1954), Finkner ef al. (1953) and Simons and Murphy (1955).
More factors were, however, identified in the present instance because a larger
number of races were employed in the seedling stage analyses and studies were
were also made on adult plant segregation in the field.

In conformity with the symbols used in describing the factors found in
Victoria (Upadhyaya and Baker, 1958), the genes for resistance revealed in
the current studies are designated thus:

Landhafer—Ld,—for adult plant resistance and seedling resistance to
races 203, 226, 226-2, 230, 237, 237-4 and 259 ; —Ld,—responsible for adult
plant resistance only.

Santa Fe—Sf,—conferring adult plant resistance and seedling resistance
to the same races as Ld,; —Tra—-compiementary to a factor (Trp) in
Trispernia, complementary gene action resulting in the expression of the
Trispernia reaction type and inhibition of the action of Sf, against race 203.

Mutica Ukraine—-Si,/—responsible for seedling resistance to races 237
and 237-4 and allelic to Sf, ; —Mu,—conferring adult plant resistance ; —Mu,
and Miip—coniplementaty genes for adult pl nt resistance.

Trispernia—Sf,’——allelic with Ld, but exhibiting a higher reaction type ;
—Tr;,—complementary to Tra.

In the case of non-allelic genes the lower reaction type was consistently
epistatic in the seedling stage.

Additional genes were revealed in certain of these varieties in crosses with
Bond and will be reported in a subsequent paper.

References
Baker, E. P., and UpapHyaya, Y. M., 1955.
Proc. Linn. Soc. N.S.W., 80: 240-257.
FinknerR, V. C., 1954.—Genetic factors governing resistance and susceptibility of oats to
Puccinia coronata Corda var. avenae F. and L., race 57. Iowa State Coll. Agric. Expt.
Stat. Res. Bull., 411: 1040-1063.
FInKNER, V. C., Atkins, R. E., and Mureuy, H. C., 1955.—Imheritance of resistance to two
races of crown rust in oats. Jowa State Coll. Jour. Sct., 30: 211-228.

Physiologic specialization m crown rust of oats.

Y. M. UPADHYAYA AND E. P. BAKER 151

LITZENBERGER, S. C., 1949.—Inheritance of resistance to specific races of crown rust and stem
rust, to Helminthosporium blight and of certain other agronomic characters of oats. Iowa
Agric. Expt. Stat. Res. Bull., 370: 454-496.

Lute, N. H.,. McWuirrer, K. S.. and Baxer, E. P., 1958.—Mode of immheritance of resistance
to powdery mildew in barley and evidence for an allelic series conditioning reaction. Proc.
Liyn. Soc. N.S.W., 83: 340-362.

Stuons, M. D., and Murpuy, H. C., 1954.—Inheritance of resistance to two races of Puccinia
coronata Cda var. avenae F. and lL. Proc. Iowa Acad. Sci., 61: 170-176.

UpapHyaya. Y. M., and Baxerr, E. P., 1960.—Studies on the mode of inheritance of Hajira
type stem rust resistance and Victoria type crown rust resistance as exhibited in crosses
involving the oat variety Garry. Proc. Linn. Soc. N.S.W., 85: 157-179.

, 1962a.—Studies on the inheritance of net resistance in oats. I.

iheriterice of stem rust resistance m crosses involving the varieties Burke, Laggan,

White Tartar and Anthony. Proc. Linn. Soc. N.S.W., 87: 141-147.

, 19626. —Studies on the inheritance of rust resistance in oats. II. The

mode of inheritance of crown rust resistance in the varieties Landhafer, Santa Fe, Mutica

Ukraine, Trispernia and Victoria in their crosses with susceptible varieties. Proc.

Linn. Soc. N.S.W., 87: 200-219.

THE OCCURRENCE AND COMPOSITION OF MANNA
IN EUCALYPTUS AND ANGOPHORA

RALPH BASDEN
Department of Chemistry, University of Newcastle

[Read 30th June, 1965)

Synopsis

Manna, an exudate from the mjured leaves and branches of certain eucalypts and angophoras,
has a different composition from the sap. The mode of occurrence, composition and the methods
of analysis employed are given. An hypothesis is advanced that the manna is the result of
the action of the enzymes of the saliva of insects on the sugars present in the phloem sap.

INTRODUCTION

The secretion of manna by certain eucalypts was noted early in the 19th
century. Virey (1832) described it in a paper to the Journal de Pharmacie
and Mudie mentioned it in a report on #. mannifera in 1834. In 1843 Johnston
examined this manna and distinguished it from the manna of commerce. The
principal sugar in eucalyptus manna he called melitose (now called raffinose)
whereas commercial manna consists mainly of mannitol. A saccharine exudation
from HE. punctata was examined by Smith (1897) who identified the sugar with
raffinose. He observed that the manna exuded from a wound in the tree was caused
by the larva of a wood-borer. It was noted that where the puncture had not
entirely penetrated the bark, the exudation was white and attracted ants, but
where the puncture had penetrated right through the bark and had entered
the wood, a mixture of manna and kino exuded and was not taken by ants.
Smith found that the manna consisted of raffinose and a small amount of
reducing sugar, that it was exuded from the tree as a syrupy liquid and
crystallized on evaporation. No further reference to eucalyptus manna has
been found except a statement in several books that H. viminalis secretes a
Sugary substance called manna.

In the present paper the term ‘* manna ”’ will be confined to the saccharine
secretion from the trees themselves. It excludes the sugary secretion from
aphids, scale, lerps and other insects. These have been called manna by some
writers, e.g. Penfold and Willis (1961), but are entirely different in composition
from the manna discussed in this paper.

MODE OF OCCURRENCE AND COMPOSITION

Manna has been observed not only on the wood of H. punctata and on the
leaves of H. viminalis and H. mannifera as mentioned above, but the writer
has collected it also from the wood of EH. maculata and from the leaves of HL.
punctata, EH. maculata, E. citriodora, E. tereticornis, Angophora floribunda and
A. costata.

On leaves and twigs it occurs as white nodules of interlaced acicular crystals,
the nodules varying in diameter from 1-5 to 4 mm. and weighing about 0-005
to 0-03 grammes. Occasionally larger nodules occur but, after growing to a
certain size, most of the nodules appear to be dislodged by the motion of the
leaves in the wind or they are dissolved by dew or rain. The largest nodule
found on a leaf was the size of a pea and weighed 0-07 gramme. The secretion
from the trunk or a large branch of a tree may weigh several grammes, but
it generally consists of a number of smaller nodules or tears.

PROCEEDINGS OF THE LINNEAN Soctety or New SoursH Wates, Vol. 90, Part 2

RALPH BASDEN 153

The manna occurs only on the site of a wound inflicted by an insect. In
more than 600 attempts to obtain manna by wounds artificially inflicted by
punching holes in leaves, scarifying twigs, cutting leaves in half, drilling a hole
in the trunk or cutting a blaze in the tree, not one resulted in the generation
of manna. On the other hand, leaves partly eaten by an insect or timber
infested with a borer will frequently produce manna from the wound. The
secretion of manna does not take place immediately a leaf is injured. Several
hours or even days may elapse before exudation commences. In the case of
a leaf the manna always occurs at the severed end of a vein (Fig. 1). On a
terminal shoot the nodule of manna conforms to the shape of the phloem sheath
of the shoot and sometimes invests the truncated end of the shoot. The manna
is secreted from the vein of a leaf at the rate of 0-001 to 0-0025 gramme per
day. There is evidence that in some cases the rate of growth of a nodule varies,

Fig. 1. Leaf of #. maculata showing nodules of manna on end of veins.

the nodule being alternately large and small in diameter. When a nodule is
removed from a leaf, the wound often continues to “ bleed ’’ and form another
nodule. As the manna exudes from the wound it crystallizes below that
already formed. The nodules thus increase by accretion from below and, not
like a stalactite, by having the newer material deposited on the distal end.
The manna is secreted from the leaves and young twigs as a liquid but appears
to crystallize immediately. No free liquid has been observed in the nodules
on the leaves. The secretion from the trunk and large branches of a tree,
however, often remains liquid for some time and may run down the trunk
several centimetres before it crystallizes.

The secretion of manna takes place throughout the year but appears to
be most abundant in the spring and early summer, when the growth of new
leaves is most rapid. Also, it has been noticed that the nodules occur more
frequently on the side of the tree facing the sun. The rate of formation of
manna thus appears to be related to the rate of flow of the sap.

The analysis of several specimens of manna from the leaves and twigs of
angophoras and eucalypts has shown that it is not just dried sap which has
exuded from the wound. The comparison of paper chromatograms of the
sugars of manna and of the juice expressed from an uninjured leaf of the same
age as that from which the manna was taken, shows a marked qualitative and

154 MANNA IN EUCALYPTUS AND ANGOPHORA

quantitative difference in composition. The approximate proportions of the
various sugars present in the manna and the corresponding leaves are shown
in Table 1.

Actually, in most cases of injury to a leaf or to a branch the wound does
not ‘‘ bleed ’”’. Only about one of every hundred injured leaves will secrete
manna. It appears that some special environment or condition is needed to
permit the formation of manna. It is postulated that this condition is the
secretion of an enzyme by the insect, possibly in the saliva, which hydrolyses
the pectins and hemicelluloses of the cells forming, inter alia, galactose and/or
galactose phosphate which, under the influence of another galactosidase or
glycosidase, synthesizes raffinose from sucrose, melibiose from glucose and, in
some cases, Stachyose from raffinose present in the phloem fluid. It may be
suggested that the causative agent is not the saliva of the insect but a micro-
organism introduced indirectly into the wound. That this is improbable is
supported by the observation of Fisher (1945) that inoculation of cuts on
Myoporum platycarpum with three micro-organisms most abundant in its manna
did not cause manna formation.

TABLE |

Comparison of the approximate proportions of sugars in the manna with those of the leaf-sap of
HK. maculata, EH. punctata and A. costata

H. maculata EH. punctata A. costata
Sugar
Manna Leaf Manna Leaf Manna Leaf
Stachyose .. He fe 0 0 2 0 10 0
Raffinose .. aK ie 80 5 80 10 65 10
Melibiose .. a io 10 0 0 0 0 0
Sucrose aE a te 6 85 10 80 20 70
Glucose and Fructose .. 4 AO 8 10 5 20

The presence of galactose in the tissue surrounding the injury to leaves
has been detected, but no galactose has been found in the uninjured part of
the leaf. That this process of synthesis of raffinose is possible is supported
by another investigation (not yet complete) in which the sap of the stems of
EH. maculata is ingested by a scale insect, Hriococcus coriaceus (Mask.). The
secretion of this insect consists of a mixture of some five or six galactosides
varying in complexity from di- to penta-saccharides. In this case the cell
sap has actually passed through the alimentary tract of the insect as distinct
from the manna which is formed in the plant substance itself. It is suggested
that some similar enzyme is involved in both cases. It is relevant to this
investigation to note that Lechevallier (1962) obtained an «-galactosidase from
germinating barley with which she converted sucrose into raffinose.

Since the concentration of sugars in the phloem sap is not constant, being
dependent on the rate of photosynthesis, translocation, metabolic trans-
formations and other processes, so the composition of manna varies. In the
leaf, which is relatively rich in sucrose and poor in glucose, the manna consists
largely of raffinose and contains little or no melibiose. The phloem sap of
the trunk and larger branches, on the other hand, contains a much higher
proportion of glucose and consequently the manna is richer in melibiose. This
sugar is about 20 times as soluble in water as raffinose and hence the exudate
from the trunk and larger branches takes longer to crystallize and is more fluid
than that of the leaves. This accounts for its habit of often running down
the trunk or branch several centimetres before it crystallizes.

The results of the analyses of specimens of manna from different sources
are shown in Table 2. It will be seen that they vary slightly from sample to
sample but in general they consist of about 60% sugars, 16% water (mainly

RALPH BASDEN 155

water of crystallization), and a small amount of ash. The remaining 20%,
has been shown to contain pectin and uronic acids. The manna has a pH of

5-3 and an acid number of 3-75 to 5-1

It should be noted that mannitol, a major constituent of the manna ot
Myoporum platycarpum (Hatt and Hillis, 1947), has not been found in any
Specimen of eucalyptus or angophora manna so far examined.

The individual sugars and their relative proportions in each specimen of
manna were determined by paper chromatography. The identity of the sugars
was deduced by comparison of their spots on the chromatogram with those
of authentic specimens of the respective sugars when subjected to certain tests.
These tests included their Reicose values, the characteristic colours given
with various spray reagents, the preparation and examination of the osazones
(of those sugars which yielded them) and, in the case of tri- and tetra-saccharides,
by identification of the products of hydrolysis with acid and with invertase.

TABLE 2

The composition of specimens of manna from different sources

E. maculata EH. punctata EH. punctata

Leaf Leaf Trunk
Loss on drying .. ae ab 16-6 14-4 17-04
Ash Eh on = a 3-6 1-3 1-6
P.O, re: it oe 0-66 Trace Trace
Total sugars (as glucose) ee 58-8 66-0 61-17
Reducing sugars (as glucose) . . 4:3 4-2 18-72

79-0 81-7 79-81

It is not always possible to determine which insect inflicted a particular
wound from which manna exudes. The secretion of manna does not take place
immediately an insect attacks the plant, so that frequently the insect causing
the wound has gone before exudation commences. However, some of the less
mobile insects have been observed on the site of the manna secretion.. For
example, the blister on a leaf caused by the leaf-miner Philactophaga eucalypt
(Frogg.) has been found with a number of nodules of manna attached to the
margin inside the blister. The larva of the leaf case-moth Hyalarcta hubneri
(Wwd) and of the saw-fly Perga dorsalis (Leach) have both been identified as
causing the secretion of manna. A wound in the trunk of HL. punctata from
which manna was flowing was caused by the larva of a beetle (not identified).

The majority of the specimens of manna, both eucalypt and angophora,
came from the leaves of suckers growing around the stumps of trees that had
been felled. Only rarely do specimens occur on mature trees. The most
abundant source of leaf manna is HL. maculata, followed by EF. mannifera and
HE. punctata. Only very few specimens have been taken from the leaves of
E. citriodora, E. tereticornis and the two angophoras. None of the other eucalypts
in this district has yielded any specimens. The most productive source of
manna from the trunk and large branches is H. punctata.

EXPERIMENTAL

The sugars present in the manna were identified by paper chromatography
and the results confirmed by other tests such as the examination of the osazones
and the products of hydrolysis by acid and by invertase. In the chromato-
graphic examination, Whatman’s no. 1 paper was used. The descending solvent
method was found to give the best results, using as developing solvents A,
butanol-acetone-water 3:4:1 and, when a two dimensional chromatogram
was required, solvent B, butanol: ethyl acetate : acetic acid : water 5:4:2: 2.

156 MANNA IN BUCALYPTUS AND ANGOPHORA

As reference sugars, stachyose, raffinose, sucrose and glucose were mainly
used, but other standard sugars were used on occasions. Table 3 shows the
Retucose Values of the sugars for solvents A and B at 25°.

After about 20 hours’ development in the tank the papers were dried at
105° and treated by the method of Bailey and Bourne (1960). The components
of the manna were identified not only by their Rg, value but also by
the characteristic colours given by the spray. The presence of the fructo-
furanose group in the molecule was detected by spraying with naphtho-
resorcinol : HCl: acetone 1:2: 200. The group was indicated by the develop-
ment of a reddish-orange colour on heating.

TABLE 3
Rglucose values of sugars at 25° for solvents A and B

Sugar Solvent A Solvent B
Stachyose 0-06 0-058
Raffinose 0:24 0-18
Melibiose ee sed 0:33 0:26
Sucrose a ie We: 0-66 0:56
Galactose 0:86 0:90
Glucose 1-00 1:00
Fructose 1-15 1-29

The oligo-saccharides were further identified by hydrolysis with 0:-2N HCl
and also with invertase. Hydrolysis by HCl was carried out at 70° for one
hour and hydrolysis by invertase at 35° for 24 hours.

The determination of the relative proportions of stachyose to raffinose, of
raffinose to sucrose, etc., was made by comparing the area and intensity of
colour of the stains on a chromatogram with those of a number of standard
solutions of sugars of known concentration.

References

Batney, R. W., and Bourng, HB. J., 1960.—Colour reactions given by sugars and diphenylamine-
aniline sprays on paper chromatograms. J. Chrom., 4: 206.

BuakELy, W. F., 1955‘ Key to the Eucalypts’”’, 2nd ed. Forestry and Timber Bureau,
Canberra, p. 154.

FIsHER, EILEEN E., 1945.—Manna formation in Myoporum platycarpum. J. Council Sci. Ind.
Res. (Aust.), 18: 159.

Hatt, H. H., and Hituis, W. E., 1947.—The manna of Myoporuwm platycarpum as a possible
source of commercial manna. J. Council Sci. Ind. Res., 20: 207.

LrecHEVALLIER, D., 1962.—Evolution de l’activité de l’-galactosidase au cours de la germination
et de la maturation de diverses semences. Compt, rend., 255: 3211.

Penrotp, A. R., and Wrttis, J. L., 196].—‘ The Hucalypts”’. Leonard Hill, Interscience,
(London and New York), p. 150.

Smitu, H. G., 1897.—On the saccharine and astringent exudations of the Grey Gum, Hucalyptus
punctata (DC.). J. Roy. Soc. N.S.W., 31: 177.

Virey, J., 1832.—.J. de Pharmacie. 18: 705.

AUSTRALIAN LARVAL CARABIDAE OF THE SUBFAMILIES
HARPALINAEH, LICININAE, ODACANTHINAE AND
PENTAGONICINAE (COLEOPTERA)

B. P. MoOoRE
C.S.I.R.O., Canberra

[Read 30th June, 1965!

Synopsis
Larvae of the followmg Carabidae are described and figured for the first time: Cenogmus
castelnaut Csiki (Harpalinae); Lestignathus cursor HWrichs. and Dicrochile brevicollis Chaud.
(both Licminae); Hudalia macleayi Bates (Odacanthinae); and Scopodes simplex Blackb.
(Pentagonicinae). The subfamily Pentagonicinae and all five of the genera were previously
unknown in the larval state.

This is the second of a projected series of papers (for the first, see Moore
(1964)) dealing with Australian carabid larvae, a group about which singularly
little information has ever appeared in print. The ultimate aim of the series,
namely, the recognition and description of all the principal genera, is likely
to prove a long-term project, in view of the size of the fauna and of the difficulties
associated with the collection of adequate larval material in an essentially arid
environment. However, the piecemeal description of isolated genera, as they
become available, serves the very important immediate purpose of adding to
our knowledge of world carabid larvae as a whole, and so providing a better
perspective for developing the general classification admirably pioneered by
van Emden (1942). Thus, on the basis of even the present very limited material,
it has been possible to add three subfamilies and nine genera to the tally of
those already positively identified in the larval stage. Addition of the tew
important subfamilies remaining as yet unknown (some of which will
undoubtedly fall to the lot of Australian coleopterists) would place the taxonomy
of larval Carabidae on a sound basis and allow it to play its full part in the
understanding of carabid evolution as a whole.

Subfamily HARPALINAE
CENOGMUS CASTELNAUL Csiki (rotundicollis Cast.)
(Figs 1—2)
Mostly pale, whitish ; head light brown, the tips of the mandibles darker.

Head rather large, very transverse, moderately sclerotized ; frontal piece
triangular, almost reaching hind-margin ; epicranial suture very short ; ventral
suture obliterated anteriorly ; nasale truncate, not prominent, lightly cuspidate ;
neck weak but with strong cervical keels ; ocelli present, six on each side ;
postocular furrows feeble ; mandible short and stout, with a basal penicillus ;
retinaculum small ; antenna slender, shorter than the mandible, four-segmented ;
maxilla setose ; inner lobe present as a stout tubercle, fused with the stipes,
unisetose before apex ; maxillary palp three-segmented, the palpiger distinct ;
labium quadrate, palp two-segmented ; ligula small, bisetose, the setae situated
on small tubercles. Pronotum slightly transverse, lightly sclerotized, slightly
broader than head ; legs short and stout, with two subequal terminal claws.
Abdomen with tergites lightly sclerotized, unmargined at sides ; pleurites and
ventrites membranous ; cerci fixed, very short, unsegmented but with setiferous
nodes ; pygopodium stout, slightly shorter than the cerci.

PROCEEDINGS OF THE LINNEAN Society or New SoutrH Watss, Vol. 90, Part 2

158 AUSTRALIAN LARVAL CARABIDAE

Length (including cerci): L,, 10 mm.; Lg, 12-13-5 mm. Head-width :
lbs Ile 100i, 8 Ib, 2. jaan.

Described from one L, and three Ls, Koojan, W.A., 16.vii.61 (L. E. Koch),
taken from soil, in association with many adults. Although the adults were
not reared individually, they appeared in numbers in laboratory trays of Koojan
pasture material, where the larvae in question were the only coleopteron
previously observed. The identification therefore seems secure.

Figs 1-2. Cenogmus castelnaur Csiki, third instar larva (L,). 1, Fore parts. 2, Right cercus
and pygopodium, right side.

In their systematic characters, Cenogmus larvae agree well with the general
description given by van Emden (1942, p. 39) for larval Harpalini (=Harpalinae
in the sense of the present paper) and they appear to come close to the South
American genus Anisotarsus Chaud. Important characters linking the two
genera include the fusion of the inner lobe to the stipes, the small retinaculum
and the weakly marked postocular furrows. The main point of difference
that can be made out concerns the abdominal praeterga which, in Anisotarsus,
are defined by a transverse furrow (van Emden, loc. cit.), but in Cenogmus, are
not differentiated from the corresponding terga.

Subfamily LIcININAE
LESTIGNATHUS CURSOR Erichs.
(Figs 3-5)

Very slender larvae, with exceptionally long appendages. Sclerites dark
brown ; intersegmental membranes and underside mostly pale.

Head small, elongate, strongly sclerotized ; frontal piece reaching hind-
margin ; ventral suture forked anteriorly ; nasale emarginate, unarmed ; neck
weakly marked ; cervical keels weak ; ocelli present, six on each side ; mandible
slender, the terebrum finely and irregularly dentate ; retinaculum long, falcate ;
antenna very long and slender, all four segments elongate ; vesicle well-marked ;
maxilla setose ; inner lobe a well-marked tubercle, with a stout apical seta ;
maxillary palp three-segmented ; palpiger distinct ; labium elongate, the palp
stout, two-segmented ; ligula minute, bisetose. Pronotum elongate, conical,
tapering anteriorly to width of head, strongly sclerotized ; legs long and slender,
with two subequal terminal claws. Abdomen with tergites strongly sclerotized,
margined anteriorly and laterally ; cerci long and slender, smooth, unsegmented
and unarmed but pubescent towards apex, not articulating with the ninth
segment, but each attached to a separate sclerite ; pygopodium slender, tubular,
about one-third the length of the cerci.

B. P. MOORE 159

Length (including cerci): L;, 20-23 mm. Head-width: L,, 1-0-1-1 mm,

Described from two Ls, Tasmania: Waratah and Mount Field, 21—26.1.61
(B. P. Moore), taken in wet forest litter. The Waratah specimen occurred
in company with numerous adults of Lestignathus cursor Hrichs. (length,
13-16 mm.) and L. foveatus Sl. (length, 7-8 mm.), but only the former species
was noted at Mount Field. In view of the size of the larvae and of their
obvious licinine affinities, Lestignathus cursor and four species of Dicrochile
(quadraticollis Cast., goryi Guér., brevicollis Chaud. and minutus Cast.) would
appear to be the only candidate species on the Tasmanian list. However,
larval Dicrochile brevicollis have since been identified from mainland material
(see below) and they prove so distinct from the Tasmanian larvae as to leave
no doubt that the latter belong to the large species of Lestignathus.

Figs 3-5. Lestignathus cursor Erichs., third instar larva (L;). 3, Head. 4, Right hind leg.
5, Right cereus and pygopodium.

DICROCHILE BREVICOLLIS Chaud.
(Figs 6-7)

Upperside largely shining black, except head, which is mostly red ;
ventrites brown, intersegmental membranes pale, whitish.

Head small, quadrate, lightly sclerotized ; frontal piece reaching hind-
margin on a wide front; ventral suture forked anteriorly ; nasale a single
triangular projection ; no obvious neck ; no cervical keels ; ocelli present, six
on each side ; mandible strongly curved, denticulate ; retinaculum large, falcate,
denticulate ; basal penicillus present ; antenna slender, four-segmented, longer
than the mandible ; vesicle minute ; maxilla setose; inner lobe well marked,
with two stout, subapical setae; maxillary palp three-segmented ; palpiger
distinct ; labium trapezoidal; ligula small, bisetose; labial palp stout, two-
segmented. Pronotwm trapezoidal, widest near base, closely adapted to head,
strongly sclerotized ; legs rather short, strongly spinose, with two subequal
terminal claws. Abdomen with tergites and pleurites strongly and completely

160 AUSTRALIAN LARVAL CARABIDAE

sclerotized, the tergites margined anteriorly and laterally ; ventrites moderately
sclerotized ; cerci short, fixed and unsegmented but with setiferous nodes ;
pygopodium short and stout.

Length (including cerci): L,, 11-12 mm. Head-width: L,, 1-0—-1-1 mm.

Described from two L,, Kast Queanbeyan, N.S.W., 13.1.65 (B. P. Moore),
taken from under stones in a dried-up river course, and in company with
numerous adult D. brevicollis. Dicrochile is the only genus of Licininae, with
adults of sufficient size, that would be expected to frequent such a habitat.

Larvae of Lestignathus and Dicrochile differ so widely in general habitus
as to suggest that the two genera belong to separate phyletic lines. Such a
conclusion is in agreement with the latest arrangement of adult Licininae (Ball,

+]
a

0.Smm.

Figs 6-7. Dicrochile brevicollis Chaud., second instar larva (L.). 6, Head. 7, Right cercus
and pygopodium.

1959), where, according to its special mandibular characters, Dicrochile is
placed in an isolated group. Nevertheless, both genera agree well with the
general diagnosis for larval Licininae given by van Emden (1942) and they
both show an important subfamily character not mentioned by that author,
namely, the pronounced anterior forking of the cranial ventral suture. Jeannel
(1942) first drew attention to this distinction from other subfamilies (where
the ventral suture is almost always a simple groove) and he looked upon it as
involving the formation of a true gula. However, Hinton (1963), who refers
to the central segment as the ventral apotome, has shown that it is not
homologous with the gula of the adult: the enclosing sutures merely represent
lines of weakness associated with ecdysis.

Subfamily ODACANTHINAE
EUDALIA MACLEAYI Bates
(Figs 8-9)

Selerites dark brown; head light brownish-testaceous ; underside mostly
pale, whitish.

Head of average size, quadrate, moderately sclerotized ; frontal piece
broadly triangular, not reaching hind-margin ; epicranial suture well marked ;
ege-bursters a row of spinules inside the frontal suture on each side; nasale
and adnasalia rather prominent, together forming an irregularly octodentate
lobe ; neck strongly marked ; cervical keels present ; ocelli present, six on each
side ; mandible rather long, slender, the terebrum serrate ; retinaculum strong,

B. P. MOORE 161

smooth ; basal penicillus present ; antenna four-segmented, as long as mandible ;
vesicle large ; maxilla setose ; inner lobe replaced by a stout seta; maxillary
palp three-segmented ; palpiger large ; labium trapezoidal; ligula triangular,
bisetose ; labial palp slender, two-segmented. Pronotuwm transverse, broader
than head ; apex and base of equal width ; sides regularly curved ; legs rather
long, spinose, with two subequal terminal claws. Abdomen with tergites and
pleurites moderately sclerotized, the latter rather prominent ; tergites margined
anteriorly but scarcely so at sides ; cerci long and slender, with setiferous nodes,
fixed at base but with three articulations ; pygopodium short, tubular.

Figs 8-9. Hudalia macleayi Bates, third imstar larva (L;). 8, Head. 9, Right cereus and
pygopodium.

Length (including cerei): L,, 5-0-5-5 mm. ; L,, 12-14 mm.. Head-width :
L,, 0:47-0:54 mm.; Ls, 1-2 mm.

Described from two L, and eight L;, Murrumbidgee River, A.C.T., x.60,
xi.61 (B. P. Moore), taken amongst gravel at the water’s edge, and in company
with numerous adults. Hudalia macleayi is the only known Odacanthine from
this habitat.

The larvae run smoothly to Colliurini (—Odacanthinae), genus Colliuris
in van Emden’s (1942) key and in the absence of material of this northern
genus, it is impossible to make out suitable separation characters. However,
the evident close agreement in structural characters, between two such
geographically isolated genera, serves to support the general system of classifica-
tion proposed. ;

Subfamily PENTAGONICINAE
SCOPODES SIMPLEX Blackb.
(Figs 10-11)
Upperside mostly dark chestnut-brown ; underside straw-coloured.

Head of average size, quadrate, strongly sclerotized ; frontal piece large,
not reaching hind-margin ; epicranial suture distinct ; ventral suture simple ;
nasale, with adnasalia, sexdentate; neck not apparent; no cervical keels ;
ocelli present, six on each side; mandible rather slender, the terebrum with

Cc

162 AUSTRALIAN LARVAL CARABIDAE

fine, rather blunt teeth ; retinaculum strong, smooth ; basal penicillus present ;
antenna four-segmented, about as long as mandible; vesicle well marked ;
maxilla weakly setose ; inner lobe replaced by a fine seta ; maxillary palp slender,
three-segmented ; palpiger distinct; labium small, trapezoidal; labial palp
slender, two-segmented; ligula small, bisetose. Pronotum subrectangular,
transverse, strongly sclerotized ; legs short, spinose, with two subequal terminal
claws. Abdomen with tergites strongly sclerotized, unmargined ; cerci slender,
moderately long, fixed, but with five articulations and numerous very long
setae ; pygopodium short, conical.

Length (including cerci): Ls, 5-8-6:-2 mm. Head-width: Ls3, 0-66—-0-70
mm.

Figs 10-11. Scopodes simplex Blackb., third imstar larva (L;). 10, Fore parts. 11, Right
cereus and pygopodium.

Described from eight L, and the exuviae from which a pharate adult was
bred, Mount Kosciusko (6,000 feet), N.S.W., 26.11.62, 27.31.65 (B. P. Moore),
taken in the open, in company with numerous adults. The bred individual
failed to free itself completely from the pupal membranes but its characters
are sufficiently developed for positive identification.

This is apparently the first larval Pentagonicine to be recognized and
described ; its characters suggest a rather close relationship with the Odacanthinae
although, in the adult stage, the two subfamilies are usually placed far apart,
largely on account of the state of the anterior coxal cavities (uniperforate in
Odacanthinae, biperforate in Pentagonicinae, teste Sloane, 1923 ; Jeannel, 1948).
However, the value of this character may need to be re-assessed.

According to present data, larvae of the two subfamilies may be separated
thus :—

Neck well marked; cerci with three articulations—ODACANTHINAE (Hudalia, Odacantha).
Neck not apparent ; cerci with five articulations—PENTAGONICINAE (Scopodes).

Acknowledgement
The author is grateful to Mr. L. E. Koch, Curator of Insects in the Western
Australian Museum, for providing material of Cenogmus castelnauwi, of the
Harpalinae—a subfamily where reliably identified larvae are seldom to be
obtained.

B. P. MOORE 163

References

Batu, G. E., 1959.—A taxonomic study of the North American Licinini with notes on the Old
World species of the genus Diplocheila Brullé (Coleoptera). Mem. Amer. ent. Soc., 16:
7-8.

Empen, F. I. van, 1942.—A key to the genera of larval Carabidae (Col.). Trans. R. ent. Soc.
Lond., 92: 1-99.

Hinton, H. E., 1963.—The ventral ecdysial lines of the head of endopterygote larvae. Trans.
R. ent. Soc. Lond., 115: 39-61.

JEANNEL, R., 1942.—Coléoptéres carabiques, 2. Paune de France, 40: 988-989.

, 1948.—Coléopteres carabiques de la région malgache, 2. Faune de l’empire frangais,
10: 375-380.

Moorg, B. P., 1964.—Australian larval Carabidae of the subfamilies Broscinae, Psydrinae and
Pseudomorphinae (Coleoptera). Pacific Ins., 6: 242-246.

Stoans, T. G., 1923.—The classification of the family Carabidae. Trans. ent. Soc. Lond.,
1923: 234-50.

SOME LAELAPID MITES OF SYNDACTYLOUS MARSUPIALS

ROBERT DOMROW
Queensland Institute of Medical Research, Brisbane

[Read 30th June 1965]

Synopsis

Australolaelaps greent, n. sp., is described from the peculiarly Tasmanian Bettongia
cuniculus (Potoroimae, Macropodidae), and Haemolaelaps calypso, n. sp., from Petaurus breviceps
(Phalangerinae, Phalangeridae), which occurs throughout eastern Australia and New Guinea.
The comparative anatomy of the species of T’richosurolaelaps and Australolaelaps is tabulated,
all being parasites of recent syndactylous marsupials in the Australian zoogeographical region.
Both genera are recognized as valid. The former comprises two species-groups, one from
peramelids and one from phalangerids and Hypsiprymnodon, the sole member of an aberrant
macropodid subfamily with traces of its scansorial ancestry. The latter comprises parasites
of the remaining macropodid subfamilies (Potoroinae and Macropodinae).

Among the mite parasites of the peculiarly Australian syndactylous
marsupials is a small group of laelapids with heavily armed coxae and edentate
chelicerae. Through the courtesy of Mr. J. H. Calaby, Division of Wildlife
Research, C.S.I.R.O., Canberra, I recently received a new species of this,
perhaps the most characteristic of all the Australian laelapid groups. As there
has been some uncertainty about the natural grouping of these mites, the
description of the new species (genus Avwstralolaelaps) seems worthwhile. The
opportunity is also taken to tabulate the comparative anatomy of all ten
Species involved, and to suggest a natural classification based both on
morphology and ecology.

A new species of the ulysses species-group, genus Haemolaelaps Berlese,
is also described.

AUSTRALOLAELAPS GREENI, N. Sp.
(Figs 1-6)

Diagnosis: Within the genus Australolaelaps, A. greeni, possessing an
immense hook on coxa II and elongate peritremes, is much closer morpho-
logically to the new combination A. validipes (Domrow)* than to A. mitchelli
Womersley. This is confirmed by ecological data. A. greent and A. validipes
parasitize potoroines, while all known hosts of A. mitchelli are macropodines.
From Table 1 below, the two species from rat-kangaroos may be separated,
in both sexes, by the number of setae on the dorsal shield and, in the female,
by the number of usurped ventral setae on the genitoventral shield.

* The original assignment of this species to Hchinonyssus Hirst (1925) was based solely on
coxa II, but it now seems that the hook on this segment has evolved separately in the two groups.
It should also be poimted out that Hirstionyssus da Fonseca (1948) is a synonym of Echinonyssus,
whose type species (H. nasutus Hirst, a common parasite of Malaysian primates—or insectivores
according to some authorities—of the genus J'upaia, Tupaiidae) is only one step removed (hook
on coxa II stronger, vertex extended) from H. callosciuri and other species with incipient hooks
on coxa II recently described from Eurasian rodents and imsectivores (see Willmann, 1952 ;
Delfinado, 1960; Bregetova and Grokhovskaya, 1961; and Wang, 1962). Hchinonyssus has
not crossed Wallace’s line to the east—H. musculi (Johnston), a cosmopolitan parasite of the
introduced house mouse, is, however, now present m Australia (Domrow, 1961, 1963)—and,
apart from zoogeographical considerations, may be separated (i) from T'richosurolaelaps by the
absence of (a) an armed tritosternal base, and/or (6) usurped ventral setae on the genital shield,
and (ii) from Avustralolaelaps by the absence of (a) usurped ventrals in the female, and (b)
modifications to femur and tarsus II in the male.

PROCEEDINGS OF THE LINNEAN Society oF NEw Sours Wates, Vol. 90, Part 2

ROBERT DOMROW 165

Types: Holotype female, allotype male, 13 paratype females, four para-
type males and two morphotype deutonymphs from the Tasmanian rat-kangaroo,
Bettongia cuniculus (Ogilby) (Potoroinae, Macropodidae, Marsupialia), Green’s
Beach, Tasmania, 6.iv.1964, R. H. Green leg. The holotype, allotype and
one pair of paratypes have been lodged in the Australian National Insect
Collection, C.S.I.R.O., Canberra.

Female: Idiosomal length in mounted, only slightly compressed specimens
always within circumscribed limits 440-473, av. 454 u. Dorsal shield texture-
less, about twice as long as wide, slightly concave vertically and midlaterally,
semicircular in posterior quarter. Margin, though somewhat eroded, distinct
vertically and laterally, leaving some setae from shield series free in adjacent
cuticle. Shield with 33 pairs of setae, those on posterolateral margins some-
what stronger, and one subterminal pair minute; also with few paired pores
in anterior half. Dorsal marginal cuticle with six to eight pairs of setae with
rather stronger shafts than those on shield. Stigmata dorsolaterally located,
with short peritremes showing two parallel series of net-like markings ;
peritremal shields extending forward from tip of peritremes, with eight-shaped
sclerotization evident on focussing more deeply.

Venter. Sternal snield extensive, but textureless, broadly arched and
very weakly defined anteriorly ; posterior margin not identified. Six sternal
setae and four spot-like sternal pores present. Metasternal complex represented
only by obsolescent shieldlets and adjacent setae. Genital shield broad,
truncate posteriorly, textureless except for two weak areolations (muscle
insertions) ; marginal strip less heavily sclerotized than disc of shield ; with
two genital setae, two pores and four usurped ventral setae. Genital cper-
culum strongly raved, encroaching proadly onto sternal area; supported by
two weakly seclerctized apodemes between coxae IV. Anal shield large, twice
as long as wide, of all body shields the most heavily sclerotized ; minutely
eranulate discally and heavily sclerotized laterally, with weak longitudinal
striae ; cribrum present. Anus set well forward, with adanal setae slightly
behind its centre ; postanal seta centrally placed, slightly weaker than adanais.
Only merest indications of metapodal shields. Ventral body cuticle with about
ten pairs of setae, of which some posterolateral pairs are decidedly stronger
than remainder.

legs. Coxal setal formula 2.1.2.1, anterior seta on coxa II obliterated
by hypertrophy of process on anterodorsal margin, which forms immense,
ventrally directed hook, with minute striate ridges basally. Formulae for
remaining segments: trochanters 6.5.5.5; femora 13.11.6.6; genua
13.11.9.9; tibiae 13.10.8.10; tarsi -—.16.16.16 (this parallels Till’s 1963
formulae for Androtaelaps Berlese, including Haemolaelaps Berlese, except for
genu LY, which in A. greent has one fewer setac). Coxa I with rather sharp,
backwardly directed process on anterobasal angle ; I-EV with somewhat blunter
exerescence on posterior aspect. Aiteroventral margin of coxa IV spinulose.
Femora I and II with basally directed setigerous spur dorsally ; femur IV with
somewhat similar, but asetose, elevation. All tarsi rather irregular in outline,
especially on posterior face. Ali leg setae slenderly tapering, two or three on
posterior aspect of tarsi I! and [IT being somewhat expanded and hyaline
basally. Tarsus I with dorsodistal sensory zone, including one distinctly bent
rod. Pulvilli I with shorter stalk and weaker claws than II-IV.

Gnathosomal and inner posterior hypestomal setae subequal, much stronger
than outer posterior and anterior hypostomals. Labial cornicles ill-defined.
Deutosternum with about five denticulations mostly in double file. Tritosternum
with very weakly barbed base and laciniaec. Palpi with five free segments ;
setal formula (trochanter to genu) 2.4.6; tibia probably with 11 setae,
including two dorsodistal rods; tarsus with bifurcate claw and several setae,
one of which is quite long. Chelicerae with basal segment short, and distal
segment slenderly tapering ; digits elongate, weak and edentate ; corona absent.

166

SOME LAELAPID MITHS OF

SYNDACTYLOUS MARSUPIALS

(Each division on the scales equals 100y..)

Fig. 2. Australolaelaps greeni, n. sp.—Venter 9.

Australolaclaps greeni, 2. sp.—Dorsum 9.

Fig. 1.

ROBERT DOMROW 167

Maie: Idiosomal length mere variable than in female; three specimens
418-429 and two 341 and 352 u, all carefully mounted. Dorsal shield more
evenly ovate than in female, slightly wider at humeral level; with all setae
(33 pairs) set ou shield and rather stronger than in female. Remainder of
dorsum essentially as in female.

Venter. Sternogenitoventral shield produced anteriorly between coxae I,
widest between coxae II and III and truncate behind coxae IV. Sternometa-
sternal area with eight setae and four pores as in female. Genitoventral
area with indications of more weakly sclerotized margins, two genital setae
and six usurpec ventral setae (i.e., pair of ventrals marked ‘‘ X ” near genito-
ventral shield of female actually taken onto shield in male). Genital aperture
well in front of SI; internal duct elongate, leading back to between SII and
it. Anai shield discrete, as in female. Ventral body cuticle with only six
or seven pairs of subequal setae.

Legs larger in respect to idiosoma than in female. Coxal setal formula
2.2.2.1, coxa II with well developed pointed process on anterodorsal margin,
but anterior seta normally developed. Coxa I with weak process and IJ and
JIt with slight excrescence on posterior aspect. Coxa IV spinulose on antero-
ventral margin. Setigercus spurs on femora I and IT incipient and femur IV
unarmed above ; femur II with strongly modified, flask-like seta on postero-
ventral aspect. Three distal segments of legs II each with one short seta
posteroventrally with base strongly infiated. Tarsus II produced into strong
spur ventrodistally, causing pulvillus to appear subterminal. Leg setation
otherwise as in female, but somewhat stronger. Ambulacra as in female.

Gnathosoma essentially as in female, but chelicerae with movable digit
coalesced with seemingiy tubular spermatophore: carrier.

Deutonymph: Neither specimen is badly compressed. The smaller (idio-
soma 412 long) is, to judge from coxa IT, prefemale ; tue larger (440..) contains
male so well developed that double setation hinders examination. Dorsally,
ineluding peritremes, as in male, with same 33 pairs of shield setae, although
Shield is even more reduced, at least in anterior half (see dotted line), than in
female (seta marked ‘ Y ” is also off shield in premale).

Intercoxal shield elongate, with usual five pairs of setae. Anal shield as
in adult.

Armature of coxae as in male, except for coxa If of prefemale, which bears
both incipient hook and weak process on anterodorsal margin, but no anterior
seta. Legs, apart from weakly developed setigerous spurs on femora I and
II, unarmed and with setal formulae, including genu IV, as in adult. Sensory
zone and ambulacra as in adult, but pulvilli I less sessile.

Gnathosoma, including chelicerae, as in female.

Notes: The syndactylous marsupials are confined entirely to the Australian
zoogeographical region. They are absent from New Zealand (apart from the
introduced Trichosurus vulpecula), but one genus, Phalanger, extends as far
west as Sulawesi (Celebes). None have crossed Wallace’s Line to the west
(Darlington, 1957). Three distinct superfamilies are involved, the Dasyuroidea,
Perameloidea and Phalangeroidea, but their phylogenetic relationships remain
obscure (Simpson, 1945).

Two major dichotomies are in common use in marsupial classification.
Using the condition of the incisor teeth, they may be divided into Polyproto-
dontia and Diprotodontia ; using the condition of the second and third toes
on the hind foot, they may be divided into the Didactyla and Syndactyla.
However, as Simpson /1945) points out, ‘‘ as might be expected of classifications
based essentially on single characters, these are contradictory and unsatis-
factory.”” Using the former, and confining ourselves to the Australian
zoogeographical region, the Dasyuroidea and Perameloidea are polvprotodont,

168 SOME LABLAPID MITES OF SYNDACTYLOUS MARSUPIALS

JSW

\ \ VA ;
a aie
AD Za « ies
ae —
KG
P

Australolaelaps greeni, n.sp.—Dorsum
Australolaelaps greeni, n. sp.—Venter 3.

Fig. 3
Fig. 4.

ROBERT DOMROW 169

and the Phalangeroidea diprotodont ; using the latter, the Dasyuroidea are
didactylous, and the Perameloidea and Phalangeroidea syndactylous.

The prodlem is, therefore, should the perameloids, comprising the one
(and only) syndactylous polyprotodont family Peramelidae, be associated with
the dasyuroids on dentition, ignoring toe structure, or, vice versa, with the
phalangeroids ? The latter choice would seem to be indicated by the host-
parasite relationships within the emanwelae species-group, genus T'richosurolaelaps,
discussed. below.*

A somewhat similar uncertainty surrounds the classification of the
Phalangervidea, which aie all syndactylous diprotodonts, comprising the three
families Phalangeridae, Macropodidae and the aberrant Phascolomidae (as
only one mesostigmatic mite has been described from wombats, Raillietia
australis Domrow (1961), this last family may be excluded from further
discussion). The anatomy and phylogeny of the macropodids have been
reviewed in detail by Tate (1948). They are usually considered to be descendants
of remote ancestors of modern phaiangerids, but it is difficult to indicate any
one division of the latter that couid have given rise to any of the macropodid
subfamilies, all of which possess varied combinations of primitive and specialized
characters. Excluding two extinct groups, the macropodids may be treated
in the order Hypsiprymnodontinae, Potoroinae and Macropodinae.

The aberrant Hypsiprymnodon, its feet *‘ already profoundly modified for
leaping’ and therefore macropodid in habitus, is, in fact, little removed in
certain characters from the phalangerids—it ‘“‘ alone of all recent macropodid
genera retains the big tce’’, anc friction ridges, typical of scansorial animals,
are still present on its feet (Tate, 1948). It is not unexpected, therefore, that
the mite species peculiar to it is inseparable, even at species-group level, from
the parasites of phalangerids, and forms, with them, the crassipes species-group,
genus Trichosuroluelaps. ‘Che parasites of the remaining, and more typical
macropodid subfamilies, confirm the generally sharp division between the
phalangerids and macrepodids—they form a distinct genus, Awstralolaelaps.

I am most grateful to Mr. Calaby for reviewing the preceding paragraphs
of these notes, and for pointing out to me Ride’s latest classification (1964)
of the Marsupialia, which does not, however, affect my argument.

The species of mites may now themselves be reviewed. Four have been
described from bandicoots (Peramelidae). These form a compact group, and
all were originally assigned to Trichosurolaelaps Womersley (1956). The first
was T. emanuelae Domrow (1958) from Echymipera kalubu in New Guinea,
which was recognized on first examination as being a little atypical and was
accordingly keyed out first, leaving the two species from phalangerids to a
later couplet. It has since been recorded from N.G. bandicoots, including
the type host on many occasions, by Domrow (1961) and Mitchell and Strandt-
mann (1964). The latter authors also described a further three species of

* To consider laelapid mites outside the scope of this paper, additional support for this
choice is found in the host relationships of the marswpialis species-group, genus Haemolaelaps
(see Womersley, 1958 ; Domrow, 1961, 1963). Of the four species whose hosts are known, two
parasitize rat-kangaroos (Potoromae, Phalangeroidea) and two bandicoots (Perameloidea).

I hasten, however, to add that there is also an argument for the former choice. Of the six
species of Mesolaelaps Hirst, four are confined to peramelids and one is host-specifie for
dasyurids (the sixth includes both peramelids and dasyurids, but apparently not phalangeroids,
amongst its numerous hosts). See Domrow (1958, 1962, 1963) and Wilson and Strandtmann
(1963).

The wheel comes full cirele when one considers the ulysses species-group, genus Haemolaelaps
(see Domrow, 1964). Of the four species, one is confined to dasyurids, while the remaiming
three (including the new species described below from Petaurus breviceps) are parasites of
phalangerids !

With many mite species undoubtedly still to be discovered when the rarer Australian
mammals are examined for ectoparasites, this is clearly an approach which may prove profitable
in future studies of marsupial phylogeny.

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2 — — — — + 4p 4 ae a “ & »Sprerys Tepodeqeyl
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< N Vv Vv N N N GE a cl a we "* 2 pplerys [eayueAcloHy
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a = —
is 0/0 s/s S/S S/T S/T S/T ela TT TT. WT = is "* P/d seuorztieg
5 = = a ap ar ap = — = == ai = = © peperq
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DD — = = =f + tr as as a <a P overs ATTeUlpnytsuoT plPerys
= “= — — — = a= — + + oe ‘+ 4 pperys tepun puepg wins.az0c(y
=r = x ra ao aa (aE) a sie a ee a a © snonuts
eVJOS PTEIYS [e19ye[o1sjUy
ce/ce es/es =a E/08 Z6/eé Te/TS — se/es ~~ 9e/O 9e/8E SESE gs/sé ss /S rPIETUS wo sated yeI0g
S eeyouu woaaih sadrpyna VWUOSVLDY pypiys sadissplod wayng wop,MYNn woop anjanunua
om
dnois-setseds sadissnuo dnois-setoods anjanunwa IOJOVIVYO OLULOUOX,
sdnjanjojp.usny snuer) sdnjanjounsoyori,T, SNUEs

sdejexjoreagsny pun sdejeejomsoyporay, fo saweds fo huomun sayunimduop
] WISV I,

ROBERT DOMROW 171

Trichosurolaclaps from New Guinea as follows: 7’. domrowi and T. whartoni,
both from Peroryctes raffrayanus (I am grateful to Dr. N. Wilson, B. P. Bishop
Museum, Honolulu, for confirming that the host of the former species, originally
listed as from 9 ‘“‘ marsupial skunk ”’, was, in fact, a peramelid), and 7. bukeri
from a ‘ bandicoot’’. No related species are yet known from the Australian
mainland peramelid fauna which, apart from a single specinien of Mehymipera
(Vate, 1952), is not known to include the recorded host genera of T'richosurolaelaps.

The two closely related species from phalangerids mentioned in the previous
paragraph are T. crassipes Womersley (1956), the type species, from T'richosurus
vulpecula (Phalangerinae) in eastern Australia and also in New Zealand. where
this possum has been introduced (Domrow, 1961; Mitchell and Strandtmann,

Figs 5-6. Auwstralolaelaps greeni, n. sp.—d, Gnathosoma 2 (ventral, with left palp dorsal) ;
6, Chelicera g.

Figs 7-9. Haemolaelaps calypso, n.sp.—i7, Gnathosoma 2 (ventral, with left palp dorsal) ;
8, Chelicera 9; 9, Tectum 2°.

1964); and 7. striata Domrow (1958) which was described from Pseudocheirus
laniginosus (Phascolarctinae) in Queensland and subsequently recorded (Domrow,
1961, 1964) from P. convolutor, a species confined to Tasmania, and another
phascolarctine, Schoinobates volans, in eastern Australia. Mitchell and Strandt-
mann (1964) note the occurrence of a species of T'richosurolaelaps on Petaurus
breviceps (Phalangerinae) in New Guinea, but do not describe it for lack of
material (this is not the species of Haemolaelaps described below).

Four species have been taken from macropodids. Tvrichcesurolaelaps
harrison’ Domrow (1961, 1962) (the use of the specific name quadratus in the
second last line of p. 80 in the original description is a slip for which I apologize ;
it should read harrisoni, the name upen which I finally decided), was described
from the musk rat-kangaroo, Hypsiorymnodon moschatus, which is restricted to
north Queensland. As noted above, it clearly belongs with the parasites of
phalangerids.

172 SOME LAELAPID MITES OF SYNDACTYLOUS MARSUPIALS

Two species are known from potoroine hosts (rat-kangaroos). <Austraio-
laclaps validypes was described from Potorous tridactylus in Queensland by
Domrow (1855) who subsequently (1958, 1963) extended its range, on the same
host, to New South Wales and Tasmania. A new species, extremely closely
related to A. validipes, is described above from the peculiarly Tasmanian
Bettongia cuniculus.

Womersley (1956) described Australolaclaps mitehelli from a small wallaby
(Protemnodon eugenii, Macropodinae) in South Australia, and this species has
since been recorded in Queensland from the larger P. dorsalis, and also from
a pademelon, Thylogale stiamatica, another macropodine (Domrow, 1961, 1962).
In addition to sharing several other characters, particularly on leg II of the
male, A. miichelli also shows the gross coxal modifications of the two parasites
of potoroines in intermediate form. The three species are, I believe, congeneric.

Fig. 10. Haemolaelaps calypso, n. sp.—Dorsum 2.

The comparative external anatomy of these mites is set out in Table 1,
which gives the characters of the dorsum, venter and leg II. The vertical
division of the table into three sections is that indicated by morphological
characters, and it will quickly be seen that this division is nicely correlated
with host preferences.

I therefore accept both Trichosurolaelaps (with two species-groups, one
from peramelids and one from phalangerids and Hypsiprymnodon, with T.
emanuelae and T. crassipes, respectively, as chefs de file), and Australolaelaps
(comprising the parasites of the remaining two macropodine subfamilies), as
valid genera, distinct, as discussed above, from Hchinonyssus (=Hirstionyssus).

Incidentally, this solution, natural on both morphological and ecological
grounds, is also the one that does least violence—only one new combination
is necessary—to the presently accepted classification, not that this is, in itself,
a valid argument for the system I accept.

ROBERT DOMROW 173

HAEMOLAELAPS CALYPSO, 0. Sp.
(Figs 7-11)

Diagnosis: In my key (1964) to the ulysses species-group, genus Haemo-
laelaps, H. calypso, a large species from a phalangerid showing the anterior
seta on coxae II and III expanded and hyaline, and the anal shield only slightly
wider than long, is nearest H. ulysses Domrow (1961). The two species are,
however, abundantly distinct, H. calypso showing decidedly shorter setae on
the dorsal shield, decidedly longer sternal and genital setae, a narrower genito-
ventral shield and the differences in leg armature detailed in the description
below.

Fig. 11. Haemolaelaps calypso, nu. sp.—Venter &.

Types: Holotype female and six paratype females from the sugar glider,
Petaurus breviceps Waterhouse (Phalangerinae, Phalangeridae, Marsupialia),
Bearii, north of Strathmerton, Vic., 20.vi.1964, R. M. Warneke leg. The
holotype and one paratype have been lodged in the Australian National Insect
Collection.

Female: Idiosoma a rounded oval, length within circumscribed limits
968-1012, av. 990 uw in six specimens, seventh slightly snialler, 935u. Dorsum,
apart from narrow marginal strip, entirely covered by single, rounded dorsal
Shield. Shield textureless except for few weak humeral striae ; bearing system
of paired pores and 39 pairs of setae, which are extremely weak, except those
at vertex, humeri and extreme posterior.

Sternal shield slightly wider than long, textureless. Anterior margin
convex, posterior margin concave. Sternal setae strong, reaching well beyond
insertions of subsequent pair. Two pairs of small transverse pores on shield.
Metasternal shields weak, each bearing metasternal seta half as strong as
sternals ; flanked internally by longitudinal pore. Genital shield only very
slightly wider than anal shield ; operculum rayed ; traces of muscle insertions

7A SOME LABELAPID MITES OF SYNDACTYLOUS MARSUPIALS

present between two strong genital setae ; disc with chevron-like striae and
two pores. Anal shield slightly wider than long, with one or two irregular
striae anteriorly and two pores laterally. Anus centrally placed, with small
adanal setae set near its anterior margin ; postanal seta slightly stronger, set
immediately in front of distinct cribrum. Metapodal shields distinct, longi-
tudinal and textureless. Peritremes extending forward almost to level of
anterior margin of coxae I; peritremal shields small, not extended posteriorly
to fuse with exopodal shields IV. Ventral cuticle with five pairs of setae
flanking shields; margins with about 52 pairs of stiff setae.

Leg setation as follows: coxae 2.2.2.1; trochanters 6.5.5.5; femora
13.11.6.6; genua 13.11.9.9; tibiae 13.10.8.10; tarsi -.16.16.16 (excluding
two terminal filaments). This compares exactly with the typical formulae
given by Till (1963) tor Androlaelaps Berlese s./. (including Haemolaelaps),
except that one seta less is present on genu IV (on checking all the species of
the complex, the same is found to be true of H. ulysses, both in the holotype
and an extensive series from Schoinobates volans, while H. penelope and H.
telemachus are typical). Anterior seta on coxae II and III expanded and
hyaline ; seta on coxa IV minute. Trochanters III and IV with three and
four expanded hyaline setae respectively (against none in H. ulysses). Apically
bifurcate setae present on femora only, formula 2.2.1.1. Remaining leg
setation undistinguished. Ambulacra I only half as strong as II-IV, all with
two claws.

Gnathosomal and outer posterior hypostomal setae subequal, considerably
weaker than anterior and inner posterior hypostomals. Gnathobase with
longitudinal hyaline flange anterolaterally ; deutosternum with about five
denticles, mostly in single file. Tritosternum with laciniae strongly bipectinate.
Labial cornicles quite well developed ; hypostomal processes, salivary stylets
and epipharynx as figured. Tectum triangular, with marginal strip smooth
and diaphanous, merging via dendritic line into denser central area as in other
species of the complex. Chelicerae with movable digit bidentate ; fixed digit
unidentate, with small pilus dentilis. Corona present. Palpal setal formula
as detailed by Till (1963) for Androlaelaps, i.e. 2.5.6.14 (trochanter to tibia,
including two dorsodistal tibial rods). One trochanteral seta hyaline and
strongly foliate, obscuring small triangular outgrowth on ventrodistal margin
of segment. Tarsus with few setae and rods in addition to bifid claw.

References

Brecetova, N. G., and GroxHovsKaya, I. M., 1961.—|New genus and some new species of
gamasid mites from North Vietnam and South China]. Rev. Ent. U.R.S.S., 40: 225-232.
Darurinetron, P. J., 1957.—‘* Zoogeography: the geographical distribution of animals. ”’
Wiley, New York, 675 pp.
Detrinapo, M. D., 1960.—Philippime Zoological Expedition 1946-1947. On some parasitic
laelaptid mites (Acarina) of the Philippmes. veldiana (Zool.), 42: 938-114.
Domrow, R., 1955.—A new species of Hchinonyssus Hirst, 1925, from Queensland (Acarina :
Liponyssinae). Proc. Linn. Soc. N.S.W., 80: 133-136.
, 1958.—New and little known Australian Laelaptidae (Acarina). Proc. Linn. Soc.
N.S.W., 82: 352-366.
, 1961.—New and little known Laelaptidae, Trombiculidae and Listrophoridae
(Acarina) from Australian mammals. Proc. Linn. Soc. N.S.W., 86: 60-95.
, 1962.—Mammals of Innisfail. II. Their mite parasites. Aust. J. Zool., 10:
268-306.
, 1963.—New records and species of Austromalayan laelapid mites. Proc. Linn.
Soc. N.S.W., 88: 199-220.
, 1964.—The ulysses species-group, genus Haemolaelaps (Acarina, Laelapidae). Proc.
Linn. Soc. N.S.W., 89: 155-162.
pA Fonssca, 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. -
Hirst, S., 1925.—Descriptions of new Acari, mainly parasitic on rodents. Proc. zool. Soc.
Lond., 1925: 49-62.
MircHEeLy, C. J.. and STRANDTMANN, R. W., 1964.—Three new Trichosurolaelaps (Acarina :
Laelaptidae) with a key to the species. J. med. Hnt., 1: 119-128.
Ripe, W. D. L., 1964.—A review of Australian fossil marsupials. J. roy. Soc. W.A.,47: 97-1381.

ROBERT DOMROW WA

Smupson, G. G., 1945.—The principles of classification and a classification of mammals. Bull.
Amer. Mus. nat. Hist.. 85: 1-350.

Tarr, G. H. H., 1948—Results of the Archbold Expeditions. 59. Studies on the anatomy
and phylogeny of the Macropodidae (Marsupialia). Bull. Amer. Mus. nat. Hist., 91:
233-352.

, 1952.—Results of the Archbold Expeditions. 66. Mammals of Cape York Peninsula,
with notes on the occurrence of rain forest in Queensland. Bull. Amer. Mus. nat. Hist.,
98: 563-616.

Tmt, W. M., 1963.—Ethiopiar mites of the genus Androlaelaps Berlese s. lat. (Acari:
Mesostigmata). Bull. Brit. Mus. (Nat. Hist.) (Zool.), 10: 1-104.

Wane, D. C., 1962.—[Two new mites of the genus Hurstionyssus Fonseca, 1948 (Acarina,
Liponyssidae)]. Acta zool. sin., 14: 411-416.

WILiMany, C., 1952.—Parasitische Milben an Klemsaugern. Zeitschr. Parasitenk., 15: 392-428.

Witson, N., and StTRANDTMANN, R. W., 1963.—Laelaptid mites from the New Guinea bandicoot,
Peroryctes raffrayanus raffrayanus. Pacific Insects, 5: 281-286.

WomersLrey, H., 1956.—On some new Acarina—Mesostigmata from Australia, New Zealand
and New Guinea. J. Linn. Soc. (Zool.), 42: 505-599.

, 1958.—Notes on the Haemolaelaps marsupialis Berl. complex, with the description

of a new species of the genus (Acarina, Laelaptidae). Proc. Linn. Soo. N.S.W., 82: 297-302.

Corrigendum

These PROCEEDINGS, vol. 89, part 1, page 161. line 11, for first “IV” read ~ III”.

COMPARATIVE STUDIES ON THE EXTERNAL ACOUSTIC MEATUS

I. THE MORPHOLOGY OF THE EXTERNAL EAR OF THE ECHIDNA
(TACHYGLOSSUS ACULEATUS)

RICHARD TUCKER
Veterinary School, University of Queensland, Brisbane, Australia

(Plates iv-v)

[Read 30th June, 1965]

Synopsis

The study of the external ear of Tachyglossus aculeatus revealed that echidna has an intra-
muscular pinna which merges with the external acoustic meatus. The latter, a completely
extracranial structure, consists of four chambers, each marked off by a sharp bend in the meatus.
The morphological and functional significance of this arrangement is discussed.

INTRODUCTION

The marked specialization of the external ear of echidna (Tachyglossus
aculeatus), combined with the conspicuous taxonomical position of the
monotremes, makes the study of the acoustic passages of the echidna interesting,
useful, and convenient as a starting point for broader investigations on the
comparative histology of the external acoustic meatus.

The comparative histology of the external acoustic meatus is at present
a virgin field. Concentration on the ear of the human and the dog prompted
by clinical interests left us without morphological perspectives and without
the basis for assessment of adaptations which can only be understood through
comparative investigations. With this in mind, a variety of species were studied.
The present paper is the first of a series of reports on the external acoustic
meatus.

MATERIALS AND METHODS

The external acoustic passages of echidna were investigated by means of
anatomical dissections and by study of unsaturated polyester resin casts.

THE MORPHOLOGY OF THE PINNA

The external opening of the acoustic meatus of the echidna is hidden
completely by surrounding spines and hairs. Topographically, it is located
close to the ventral margin of the region of the coat which bears spines and
the distance between the anterior margin of hair and the opening of the
external ear equals roughly the distance between the anterior end of the maxilla
and the outer margins of the hair (Plate iv, fig. 1). However, the removal
of skin makes the large opening leading into the spacious cavity visible (Plate
iv, fig. 2). More complete dissection reveals that the cavity is in fact a pinna
embedded in the skin and musculature. The pinna is orientated dorso-ventrally
with its cartilaginous margin, bending medially to form a roughly triangular
hiatus covered with the relatively short and dense hairs (Plate v, fig. 5). In
the ventral portion of the pinna the lateral margins of the cartilage merge,

PROCEEDINGS OF THE LINNEAN Soctety oF New SoutrH Watss, Vol. 90, Part 2

RICHARD TUCKER IAT

forming a large goblet-like structure. The median surface of the pinna is smooth
and flat (Plate v, fig. 6). The pinneal cavity passes at once but at a right-angle
into the external acoustic meatus.

THE EXTERNAL AcousTIC MEATUS

The diameter of the external acoustic meatus is about a quarter of that
of the pinneal cavity. The meatus itself is tortuous: at first (in the para-
pinneai portion) it runs anteriorly, then medially and finally turns dorsally
to reach the base of the skull (Plate iv, fig. 4, and Plate v, fig. 6). The meatus
is narrowest at its cranial end and widest in the proximity of the pinna (Plate
v, fig. 7). On the internal surface of its wall the distinct crista is present
(Plate iv, fig. 6). The cranial opening of the meatus is elongated (Plate v,
fig. 8). The sharp angles between the various portions of the external acoustic
passages divide the extracranial acoustic meatus and pinna into four (4) distinct
chambers, the most external being formed by the pinna; this is elongated and
contains hairs. The second chamber, directed anteriorly, is relatively spacious.
The third and fourth are smaller in diameter and develop the internal cartilaginous
crista. The wall of the meatus consists of cartilaginous “‘ ribs’ connected by
@ membrane.

DISCUSSION

The external ears of all mammals and birds have the same general functions.
To these belong: (1) expression ; (2) temperature regulation ; (3) protection
of the middle and inner ear; (4) the isolation and localization of sound ; and
(5) the transmission of sound. These functions are modified from species to
species. Nevertheless, they form a convenient reference frame for discussion
and the assessment of adaptations and modifications within the particular
auditory passages. Therefore, the conditions described above will be discussed
in relation to these functions.

In most mammals the function of expression belongs wholly to the pinna,
the muscular plate of the platysma myoides and to the facial nerve. The
expressive ability of the pinna is governed by its size and the degree of its
mobility. Animals of terrestrial habitat such as the horse, ox, and dog have
large and freely mobile pinnae. Aquatic animals and those with subterranean
or burrowing habits have a pinna of considerably reduced size, while primates
of arboreal habit or descent have rather immobile ears. The musculature
of the external ear develops from the platysma myoides, the cutaneous muscle
of the second branchial arch, and the degree of mobility of adult external ears
depends on the final connection of the pinna with this muscle. The human
ear has lost most of its muscular connections, while the mobile and expressive
ear of most terrestrial animals is connected with the platysma by special
muscular strands—the auricular muscles. These muscles as well as the
platysma are supplied by fibres from the facial nerve so that the expressive
movements of the ears are stimulated by fibres from the same area of the central
nervous system as those connected with the facial expression in man.

According to McKay (1894), in echidna the panniculus carnosus consists
of three layers, and is innervated only by the cervical nerves, the auricular
region being supplied by the second cervical nerve. However, Westling (1889)
describes innervation of subcutaneous cervical muscles by both the facial and
cervical nerves. Also Edgeworth (1935) pictures (after Schulman) the platysma
in echidna together with M. sphincter colli superficialis in the area of the external
acoustic meatus.

The intramuscular position of the pinna in echidna changed also the
character of its movements: pinna is not moved by the muscle; instead, it
is moved with the muscles. It is in fact a muscular cartilaginous insertion
(insertio cartilaginea) comparable in a sense with the tendinous insertions in the

D

178 COMPARATIVE STUDIES ON THE EXTERNAL ACOUSTIC MEATUS. I

rectus system (Tucker, 1955). This insertion is large and isolated. The
existence of different relations of pinna to the musculature (the echidna type
being an extreme case) enables us to distinguish between the three types of
pinna in mammals.

In relation to the platysma the mammalian pinna can acquire an evira-
platysmal position, with no, or only negligible, connections with this muscle—
e.g. Homo; or a supra-platysmal position with only the deep portion of pinna
connected with platysma and supplied by strong auricular muscles, e.g. Bovidae,
Equidae. In echidna, we have the case of an intramuscular pinna connected
with panniculus carnosus. The different types of relationship between the
pinna and the musculature in mammals are shown diagramatically in Text-fig. 1.
There is probably a relative lack of expression in echidna due to the factors
mentioned above.

<— = ¢F

Text-fig. 1. Diagram showing the different types of relation between pinna and the muscles
connected with it ; on the left side, extraplatysmal pmna, im the middle supraplatysmal pinna,
and on the right side intramuscular pinna.

The attempts to explain the conditions in the echidna must develop into
discussion of :

(a) Musculo-pinneal relations, such as invasion of pinna by the panniculus
carnosus in echidna; or of the acceptance of the above described relations as
the primary condition and proceeding to the subsequent separation of pinna
by the recession and differentiation of the cutaneous musculature in other
mammals.

(b) Relating the intramuscular pinna to the way of life of Tachyglossus,
especially to its defence technique which, at the same time, removes all surface
protrusions and develops the powerful musculature for the erection of spines.
Lack of sufficient differentiation in cortex, cranial nerves, platysma, and the
connections between them cause a basic lack of suitable conditions for facial
expression.

The pinna itself is characterized by :

a. The robustness of its cartilage ; b. Lack of morphological differentiation
of its cartilage and of differentiation into separate cartilages ; c. The absence
of any structure which can cause friction during the shift due to muscular
contraction ; d. The fusion of the ventral margins of the cartilage resulting in
the formation of the flask-like pinneal cavity ; and e. The presence of hair on
the internal surface of the pinna.

The robustness of the cartilage and the thickness of its margins are probably
related to the strong muscular insertions onto it; lack of differentiation of
separate cartilages may be related to the absence of independent and complicated
movements. The absence of morphological differentiation of the cartilage
itself is probably a result of the lack of the sound-dispersing function in the ear
of echidna.

The smoothness of the median surface of the pinna and its flatness seem,
however, to form a more general and therefore a more interesting feature. The
median surface of the pinna is smooth in all types of ears, even in the extra-
and supra-platysmal ears which can reduce friction by the lateral bending of

RICHARD TUCKER 179

the pinna. It seems, therefore, that this smoothness of the median surface
is phylogenetically a stable morphological feature, independent of the local
forces and stresses.

The tubular shape of the pinna stresses again conditions indicated by its
embedding in the musculature, and by the presence of hairs, that the external
ear in echidna is not involved in the temperature regulative mechanism. The
presence of the arterio-venous anastomoses in the echidna ear is nevertheless
possible, and needs further investigation.

The most developed specialization in the echidna ear seems to be its
protective adaptations ; external hairs and spines guard the external opening
of the meatus. All of these—hairs, spines, pinna and meatus—are connected
with the cutaneous musculature. The entrance of foreign bodies can be next
prevented by the dense hairs on the internal surface of the pinna. The narrow-
ness and angularity of the meatus form another conspicuous protective
mechanism: it can protect against the entrance of foreign bodies as well as
against high intensity of sound.

In mammals the most common protective device against the high intensity
of sound is the increase in the mobility of the pinna so that it can be turned
away from the direction of intense sounds. Primates can achieve a similar
effect by covering their ears with their hands, but aquatic mammals develop
an especially long and angular meatus which serves the same purpose. In
echidna, the external acoustic meatus is very long and very angular. It is
also essentially an extracranial structure. Because of this extracranial position,
passive mobility and frequent changes of plane, the mutual transmission of
sound from bone to the meatus is negligible. The intramuscular pinna of the
echidna has a poor ability to divert sounds into the meatus. This disability
may be partially compensated by the diminishing diameter of each subsequent
chamber. In a smaller chamber the same intensity of sound gives a stronger
effect or, conversely, a fraction of the number of sound waves passed to a
smaller chamber will give the same acoustic effect.

The functional significance of the crista (Plate v, fig. 6) could not be assessed
adequately on the material at my disposal.

The external ear of echidna was pictured without description by Westling
(1889) and mentioned by Winge (1941). It was also studied in more detail
by Ruge (1898) who considered pinna in echidna to be a derivative of the hyoid
bone. He found a ramification of the hyoid bone merging with the tympanic
cartilaginous ring, which is closely connected with the external acoustic meatus,
the latter merging with the pinna. However, the same author found the above
connections less pronounced in Ornithorhynchus, and intended to prove his
point through a series of comparative embryological investigations which,
however, to my knowledge were never published. Both the descriptions of
Ruge (1898) and Winge (1941) differ from our findings. According to Ruge
(1898) the oval pinna is perforated by the canal, and has a transverse process.
Winge (1941) did not account for the geniculate structure of the external
acoustic meatus. In our material, the pinna was distinctly elongated, no canal
or transverse process was observed, and the external acoustic meatus exhibited
Sharp bendings. The partially cartilaginous and partially membranous structure
of the external acoustic meatus in echidna is noted by both previous authors,
but not discussed, and its presence was confirmed in the present investigation.

Acknowledgements

I wish to express thanks to Mr. E. Hollywood for the photographs, and
to Mrs. L. Endean for help with preparation of the manuscript. Thanks are
also due to the Rural Credits Development Fund for financial assistance with
the work.

180 COMPARATIVE STUDIES ON THE EXTERNAL ACOUSTIC MEATUS. I

References

EpcrwortH, F. H., 1935.—* The Cranial Muscles of Vertebrates”’. Cambridge University
Press.

McKay, W. J. S8., 1894.—The morphology of the muscles of the shoulder-girdle in Monotremes.
Proc. Linn. Soc. N.S.W., 1894, Vol. 9, 2nd Series: 263-360.

RuceE, G., 1898.—Das Knorpelskelet des Ausseren Ohres der Monotremen—ein Derivat des
Hyoidbogens. Morph. Jahrbuch, 25: 202-223. Leipzig.

WESTLING, CHARLOTTE, 1889.—Anatomische Untersuchungen tiber Echidna. Béihang till Kong.
Svenska Vetensk. Akad. Handl., Stockholm, 1889. Afd. IV, Bd. XV, No. 10.

Winecet, H., 1941.—“ Interrelations of Mammalian Genera’. Vol. I, 1941. C. A. Reitzels
Forlag. Edited by Ad. S. Jensen, D.Sc., R. Sparch, D.Sc., and H. Volsoé, Mag. Sc.

EXPLANATION OF PLATES IV-V

Plate iv

Fig. 1. Tachyglossus aculeatus. The topography of the ear. A notch indicates the position
of the opening.

Fig. 2. Tachyglossus aculeatus. External ear, after removal of the skin.

Fig. 3. Tachyglossus aculeatus. The median view of the pinna, in relation to the musculature.

Fig. 4. Tachyglossus aculeatus. The resin cast of the external acoustic meatus, showing its
bending, and relation to the pinna.

Plate v

Fig. 5. Tachyglossus aculeatus. Pinna and external acoustic meatus dissected away from the
muscles (lateral view).

Fig. 6. Tachyglossus aculeatus. Morphology of the pinna and external acoustic meatus. Pinna
completely dissected—note its flatness, close connection with the external acoustic meatus,
and the presence of the cartilaginous crista inside the external acoustic meatus.

Fig. 7.—Tachyglossus aculeatus. The transverse portion of the external acoustic meatus.

Fig. 8. Tachyglossus aculeatus. Entrance of the external acoustic meatus mto the skull.

THE DEVONIAN TETRACORAL HAPLOTHECIA AND NEW
AUSTRALIAN PHACELLOPHYLLIDS

A. KE. H. PEDDER
University of New England, Armidale, N.S.W.

(Plate vi)
[Read 28th July, 1965]

Synopsis

Research in progress on Australian Lower Devonian corals is exposing a gradation between
the Cyathophyllidae and the Disphyllidae. It is therefore recommended that the Disphyllidae
be demoted to subfamily rank within the Cyathophyllidae.

Haplothecia filata (Schlotheim), a disphyllinid from the Frasnian of Germany, is restudied
and the genus Haplothecia upheld. Two phacellophyllids from New South Wales are also
described: the first, Bensonastraea praetor, gen. et sp. nov., is from the Timor Limestone of
probable Hifelian age, while the second, Macgeea touti, sp. nov., occurs in the Loomberah and
uppermost Sulcor Limestones, both late Emsian or early Hifelian in age.

A new term, veprecula(e), is introduced for the type of spinose projections found on the sides
of the septa in Bensonastraea.

INTRODUCTION

Preparation of systematic accounts of several Devonian tetracoral faunas
in New South Wales has necessitated the undertaking of various ancillary
investigations. One of these has been the restudy of the type specimen of
the genus Haplothecia, and an evaluation of its relationship to highly carinate
Australian species referred to Phillipsastrea.

The results of this study, together with the description of two new
Australian species of a related family, form the subject of the present paper.

A NEw MorRPHOLOGICAL TERM

The stability of morphological terms, which almost without exception
characterizes current descriptions of tetracorals, stems mainly from Hill
(1935, 1956). In these works carinae are defined as flanges, or flange-like
elevations, on the sides of a septum formed by thickened trabeculae.

In Bensonastraea, a new genus described below, the septa bear small, but
nevertheless prominent spinose projections which do not conform to the
definition of carinae given above. As far as can be judged from the less than
perfectly preserved available material, these are composed of fibrous skeletal
material and appear to be prolongations of lateral trabeculae. If so, a very
similar trabecular pattern has been figured by Rdzkowska (1953, text-fig. 6)
in ‘ Synaptophyllum’ soshkinae (see McLaren, 1959, for a revision of Synapto-
phyllum. Rozkowska’s species possesses horseshoe dissepiments and is presum-
ably a phacellophyllid).

It is here proposed that spinose projections of the type occurring on the
septa of Bensonastraea be known as vepreculae, singular veprecula, from the
latin meaning a small thorn. Vepreculae may be homologous with the
Synapticulae of scleractinian corals.

PROCEEDINGS OF THE LINNEAN SocieTY OF NEw SourH Watss, Vol. 90, Part 2

182 HAPLOTHECIA AND NEW AUSTRALIAN PHACELLOPHYLLIDS

SYSTEMATIC DESCRIPTION
Family CYATHOPHYLLIDAE Dana, 1846

According to the majority of recent classifications, the Cyathophyllidae
are distinct from the Disphyllidae (= Phillipsastraeidae auct.). In reality,
however, a basis for the recognition of the Cyathophyllidae has been possible
only since Birenheide’s (1963) redescription of Cyathophyllum and related
genera. In view of this and other recent work (Philip, 1962; Pedder, 1966)
on earlier Victorian faunas, the distinction between the families is much less
clear. For example, among the fasciculate forms, species of Tipheophyllum
such as T. ops and T. cognatum completely bridge the gap between the
Cyathophyllidae and Disphyllidae, and among the massive forms this gap is
bridged by species such as Hexagonaria approximans. It is therefore proposed
that the Disphyllidae be relegated to subfamily rank within the Cyathophyllidae.

Subfamily DISPHYLLINAE Hill, 1939
HAPLOTHECIA FILATA (Schlotheim)
(Text-figs 1, 2, 4, 7)
1820. Madreporites filatus Schlotheim (partim), p. 359, var. « only.
1885. Haplothecia filata Schloth. sp. Frech, pp. 68, 69, Pl. 4, figs 7, 7a.
1951. Phillipsastraea filata (Schloth.), Soshkina, pp. 98-100, text-fig. 36,
Pl. 18, figs la, D.

1952. Phillipsastraea filata (Schlotheim), Soshkina, p. 101, Pl. 42, figs 141.
1956. Haplothecia filata (Schloth.), Hill, p. 280, figs 191. la, b.

Type series.—Frech (1885, p. 68) states that Schlotheim’s specimens
included Phillipsastrea hennahi as well as a specimen from the Lias of Wurtem-
berg. The specimen figured by Frech is technically a lectotype and is now
in the Institut fiir Palaontologie und Museum der Humboldt-Universitat,
Berlin, where it is registered Q. Kat. A. 138, p. 1530. Previous to the present
investigation it consisted of two small pieces and four transverse sections,
including the one figured by Frech. Frech’s longitudinal section is now lost
and as no other existed, the writer was permitted to prepare a new one.

The museum label indicates that the specimen was obtained from the
Iberger Kalk, Winterberg bei Grund (Harz, Germany).

Description.—The corallum is cerioid with axes of adjacent corallites 5 to
8 mm. apart. Nothing of the exterior is preserved, but the disposition of the
dissepiments suggests that there would have been a relatively wide calicular
platform.

Neighbouring corallites throughout the corallum are separated by a wall,
typically 2-5 to 3-5 mm. thick, composed of an apparently structureless light-
coloured skeletal material divided by a dark axial plate.

Septa are not embedded in the wall. In the dissepimentarium they tend
to be arranged in parallel groups so that some are almost parallel with the wall
and a few are even contratingent with an adjacent septum ; the arrangement
suggests a thamnasterioid origin. Septal counts range from 12 x 2 to 13 x 2
in the material studied by the writer; however, Frech, who studied further
specimens, gave the maximum as 15 x 2. Major septa may extend to the
axis, but more commonly are just withdrawn from it; the minor are either
confined to the dissepimentarium, where they are scarcely differentiated from
the major, or just project into the tabularium. Prominent yard-arm carinae
are present in the dissepimentarium, but in the tabularium septa are smooth
and thin. The carinae are so well developed that they may touch the wall
where wall and septum are almost parallel. At the carinae, septa are trabeculate,
whereas between them they are composed of an apparently structureless and
lighter coloured material identical with that forming the wall. Frech’s
longitudinal section shows divergence of the trabeculae ; however, this is not

A. E. H. PEDDER 1383

as marked as in normal phacellophyllids and in fact there is no divergence at
all on one side of the newly prepared section. Calcite fibres spread upwards
randomly from near the trabecular axis and are not grouped in discrete
fascicles.

The dissepiments are mostly small and globose and are more or less
horizontally disposed; towards the tabularium they steepen so that the
transition to the tabularium is abrupt. Where septum and wall lie close
together, dissepiments commonly occur between them.

In both longitudinal sections of the lectotype, the tabulae, which are
considerably disrupted by septa, are closely spaced and generally flat.

2

1

Figs 1,2. Haplothecia filata (Schlotheim), lectotype x24. 1, Transverse section. 2, Longi-
tudinal section. The stipplmg between carinae is diagrammatic and represents apparently
structureless skeleton.

Remarks.—Frech (1885, p. 68) proposed the genus Haplothecia solely for
this species. Subsequently the genus received little attention until reassessed
by Lang and Smith (1935, pp. 549, 550), exactly 50 years later. Although
Lang and Smith’s conclusion that it is synonymous with Phillipsastrea has
gained wide acceptance (Stumm, 1949, p. 35; Wang, 1950, p. 220; Soshkina,
1951, p. 98; Schouppé, 1958, p. 233; Soshkina and Dobrolyubova, in Orlov,
1962, p. 336), there is no indication that they examined topotypic material.
Indeed both their description, which minimizes the differences between
Haplothecia filata and Phillipsastrea sensu stricto, and their figures, which
illustrate specimens that are not necessarily Haplothecia, suggest that they
did not.

Thanks to Drs. Jaeger and Forbes it has been possible to compare the
lectotype of Haplothecia filata with a topotypic specimen (Sedgwick Museum
No. H438c, d from Barton Quarry, Devon) of Phillipsastrea hennahi, the type
species of Phillipsastrea. The latter is thamnasterioid to subcerioid, has a
phacellophyllid zone of trabecular divergence as well as a few horseshoe
dissepiments and is, therefore, a phacellophyllid and not a disphyllinid.

The allotment of Haplothecia to the Disphyllinidae must be regarded as
_ provisional. The genus may have evolved from Phillipsastrea but, for the
moment at least, the family Phacellophyllidae is reserved for corals having
a normal phacellophyllid zone of trabecular divergence.

Family PHACELLOPHYLLIDAE Wedekind, 1921
Genus BENSONASTRABA nov.

Name derivation.—Patronym for W. N. Benson, pioneer geologist of the
Great Serpentine Belt of New South Wales and Greek, gotoov = star, with
traditional ending for coral genera.

Type species.—Bensonastraea praetor, sp. nov., see below.

HAPLOTHECIA AND NEW AUSTRALIAN PHACELLOPHYLLIDS

184

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(Schlothem

A. E. H. PEDDER 185

Diagnosis.—Corallum thamnasterioid. Septa vepreculate. Trabeculae
divergent exterior of the tabularium. Dissepimentarium and tabularium
complex. Five zones present in dissepimentarium ; from periphery inwards
these are: broad zone of large and small, normal and lateral dissepiments ;
narrow zone of flat dissepiments; narrow zone of small outwardly-convex
dissepiments ; series of horseshoe dissepiments ; and a narrow zone of inwardly-
convex dissepiments. Tabularium divided into a narrow outer zone of flat
tabellae ; a periaxial zone of outwardly-convex tabellae ; and a central region
of more or less flat tabulae.

Remarks.—The new genus most obviously recalls Keriophylloides and
Billingsastraea. Soshkina (1951, p. 102) erected Keriophylloides for Kerio-
phyllum astraetforme Soshkina (1936, pp. 62-64, figs 71, 72), an Hifelian species
from the northern Urals, but subsequently, with Dobrolyubova as co-author
(in Orlov, 1962, p. 336), merged the genus in Buillingsastraea Grabau (1917,
p. 957). The difficulties in recognizing Billingsastraea have been outlined
previously by the present writer (1964), p. 447); since then a further account
of the genus has been given by Oliver (1964). Oliver’s interpretation of
Billingsastraea is similar to Ehlers and Stumm’s (1953) and, if correct,
Keriophylloides would be distinguished from it by its carinae which appear
to be of the veprecular rather than the yard-arm type, its highly arched
dissepiments giving rise to an exsert calicular rim, and by the dilation of its
Septa immediately exterior of the tabularium.

Bensonastraea is distinguished from both of these genera, and also from
the polyonymous genus Phillipsastrea d’Orbigny (1849, p. 12), by its complex
dissepimentarium. Sulcorphyllum Pedder (1964a, p. 366) and Pseudoacervularia
sensu Rozkowska (1953, p. 49) non Lang, Smith and Thomas (1940, p. 108) in
Some respects resemble the new genus, but are cerioid, have a different
dissepimentarium, and lack vepreculae.

At the present time, only the type species is referred to Bensonastraea which
may prove to be but a local offshoot from the central plexus of the Phacello-
phylldae.

BENSONASTRAEA PRAETOR, gen. et sp. nov.
(Pl. vi, figs 1, 6, 7; text-figs 3, 5, 6)
Name derivation.—Latin, praetor = leader.

Type series.—Holotype, Geological Survey of New South Wales No. 3463,
Timor Limestone (probably Hifelian), Portion 133, Parish of Lincoln, County
Brisbane, N.S.W.; the collector is not recorded.

Diagnosis.—Large Bensonastraea with axes of adjacent corallites 12 to
23 mm. apart and tabularium normally 5 to 6 mm. in diameter. Septal count
mie 2 GO. 2 xX 2.

Description.—The corallum is thamnasterioid and apparently large ; when
first seen by the author, the holotype had already been cut and yet measured
approximately 80 x 65 x 80 mm. Axes of adjacent adult corallites are
Separated by from 12 to 23 mm. and the tabularium is normally 5 to 6 mm.
in diameter. No exterior surface is preserved.

There are 17 x 2 to 21 x 2 septa in adult corallites, although differentiation
into two orders is evident only in the tabularium. Throughout most of the
dissepimentarium septal arrangement is thamnasterioid with the peripheral
ends of the septa being either confluent with, abutted against, or withdrawn
from, a septum of an adjacent corallite ; arrangement is radial in the tabularium.
In the outer region of the dissepimentarium where vepreculae are abundant,
the septa are thin and locally degenerate, being represented by vepreculae
only. In the region of the horseshoe dissepiments, however, they are strongly
dilated and are also asymmetrically (peripheral end blunter) fusiform in transverse

1386 HAPLOTHECIA AND NEW AUSTRALIAN PHACELLOPHYLLIDS

section ; vepreculae are fewer here and generally masked by thick sclerenchyme.
In the tabularium septa are thin, straight or sinuous, and smooth ; the major
commonly extend to within about 1 mm. of the axis, whereas the minor terminate
close to the inner margin of the dissepimentarium. Trabeculae diverge in a
zone opposite the horse-shoe dissepiments.

Five concentric zones may be distinguished in the dissepimentarium. The
outermost of these is the broadest and consists of both large and small
dissepiments ; lateral dissepiments are characteristic of this zone. The next
zone is one of flat, sloping or sagging dissepiments. Inside these there is a
narrow and, in places, discontinuous zone of small outwardly-convex dissepiments,
typically up to three deep. Irregularly superposed horse-shoe dissepiments
form the next zone; these vary considerably in size and some are sigmoidal.
The innermost zone consists of small inwardly-convex dissepiments, typically
two to four deep.

The peripheral part of the tabularium is formed of flat tabellae, inside which
there is a periaxial series of generally upwardly and outwardly convex plates.
Flat, or only gently arched or sagging tabulae occupy the central region of
the tabularium.

MACGEEA TOUTI, sp. nov.
(Pl. vi, figs 2-5, 8-11; text-figs 8-11)
21917. Zaphrentis typlasmoides Dun, p. 218. Nomen nudum et oblitum.
1918. Zaphrentis (2) sp. (sp. et subgen nov. ?); Dun in Benson, pp. 335,
375, 376, text-fig. 3, Pl. 34, fig. 1.
1922. Zaphrentidae (new genus); Benson, p. 143(60).

Name derivation.—Patronym for S. M. Tout who, according to Benson
(1918, p. 322), “‘was the first to bring the Loomberah limestones under
Scientific notice °’.

Type series—Holotype and paratypes 1-4, University of New England
Nos. F8851-8855 respectively, collected by the author from the Loomberah
Limestone (late Emsian or early Hifelian) in Portion 58, Parish of Loomberah,
County Parry, N.S.W. Paratypes 5, 6, University of New England Nos. F8856,
8857 respectively, collected by the author from the uppermost beds of the
Sulcor Limestone (late Emsian or early EHifelian) at the northern end of the
outcrop in Portion 249, Parish of Burdekin, County Inglis, N.S.W.

Diagnosis.—Solitary ceratoid to cylindrical tetracoral with a maximum
known length and diameter 70 and 19 mm. respectively. Septa considerably
dilated and commonly contiguous in a zone immediately exterior of the
tabularium ; trabeculae divergent in this zone. Septal counts 18 x 2 to 24 x 2
at maturity. Dissepimentarium in two parts, an outer of predominantly flat
plates and an inner of small and commonly masked horse-shoe dissepiments.
Tabulae variable, mostly short and arched.

Description.—All available specimens are completely embedded in matrix ;
from thin sections the corallum appears to be ceratoid in early stages and
subcylindrical to cylindrical at maturity. Rejuvenescence occurred rarely.
Specimens up to 70 mm. long and 19 mm. in diameter are known ; however,
diameters in excess of 15 mm. are unusual.

An epitheca is preserved in a minority of specimens ; where present it is
between 0:15 and 0-25 mm. thick and consists of a thin dark axial plate and
an inner lighter layer.

Arrangement of the septa is radial, or faintly pinnate about the presumed
cardinal-counter plane. The major septa extend to near and in some Cases
beyond the axis, and may be rhopaloid or forked axially ; the minor just intrude
the tabularium. Both orders of septa are considerably dilated in the dissepi-
mentarium, appearing fusiform or wedge-shaped in transverse section, and are

A. E. H. PEDDER 187

commonly contiguous just exterior of the tabularium. In the tabularium
they are irregularly bent and variably carinate. Septal counts vary from
18 x 2 to 24 x 2 in adult specimens.

Trabeculae diverge in the normal phacellophyllid manner. Fibre fascicles
are prominent as dark regions in most sections, although individual fibres can
not be discerned.

A series of flat or gently convex plates forms the outer region of the
dissepimentarium ; this may be as much as 2 mm. wide, although it has been
eroded from most specimens at the type localities. A collar of relatively small
horse-shoe dissepiments constitutes the inner part of the dissepimentarium ;

AE

Se
=H

Figs 8-11. Macgeea touti, sp.nov. x3. 8, 11, holotype. 9, paratype 4. 10, paratype 2.

however, in many cases this is largely obscured by dilation of the septa. The horse-
Shoe dissepiments are normally superposed, but in some specimens from the
Sulcor Limestone (e.g. paratype 6), short subsidiary strings of horse-shoe
dissepiments branch from the main collar into the outer part of the dissepi-
mentarium.

The tabularium, which is one-half to three-fifths the total width of the
coral, is composed of numerous tabulae varying considerably in width and
curvature ; locally these may be invested by sclerenchyme.

Remarks.—The relatively wide zone of flat dissepiments, the degree of
dilation of the septa in the zone of trabecular divergence, and the low ratio
between the number of septa and diameter, distinguish Macgeea touti from the
majority of previously described species which are, of course, Givetian or
Frasnian in age.

Macgeea (2) murchisoni (Penecke, 1894, pp. 595, 596, Pl. 7, figs 15-17) is
probably the closest known species. It was originally proposed for a fragment

188 HAPLOTHECIA AND NEW AUSTRALIAN PHACELLOPHYLLIDS

from the Emsian of the Carnic Alps and has subsequently been reported to
be present in the Hifelian of both the type area (Heritsch, 1935, pp. 188, 189)
and Armenia (Soshkina, 1952, p. 84, Pl. 18, fig. 65). Penecke’s species differs
from the new one in having more septa at a given diameter, and broader and
more widely spaced tabulae. Specimens from the Emsian at Chalonnes (Le
Maitre, 1934, pp. 148, 149, P1.5, figs 3, 4) and from the Givetian at Ville De-d’Ardin
(Le Maitre, 1937, pp. 111-113, Pl. 7, figs 3-5, 11, 12; Pl. 8, fig. 7), which have
been referred to Thamnophyllum murchisoni, are quite unrelated to Macgeea touti,
as are the specimens, which have been compared with 7. murchisoni (Firtion,
1957, p. 127, Pl. 5, figs 6, 7), from the Givetian of the Val de Bruche.

A number of species of Pexiphyllum, established by Walther (1928) on
specimens from the Frasnian of Germany, resemble Macgeea touti. Apart
from discrepancies in septal counts, septal dilation in these species is as pro-
minent in the outer as in the inner region of the dissepimentarium, with the
result that transverse sections of eroded specimens do not simulate cog-wheels,
as those of Macgeea touti do.

In a footnote, Glinski (1961, p. 284) has claimed that Macgeea (Webster,
1889, p. 710) is a junior synonym of Péerorrhiza (Ehrenberg, 1834, p. 312).
However, pending publication of figures of the interior of Cyathophyllum
marginatum Goldfuss, the type species of Pterorrhiza, the present author prefers
to retain the name Macgeea for species such as M. toutt.

Acknowledgements

Assistance rendered by a number of geologists is gratefully acknowledged.
Dr. EH. O. Rayner and Mr. H. F. Whitworth of the New South Wales Depart-
ment of Mines, sanctioned the loan of specimens from the Mining Museum,
Sydney. Dr. J. W. Pickett, now also of the New South Wales Department
of Mines, forwarded a description of the holotype of Phillipsastrea hennahi,
while Mr. M. Mitchell, of the Geological Survey of Great Britain, provided
photographs of the same specimen. Dr. C. L. Forbes arranged for loan of
Barton Quarry corals from the Sedgwick Museum, Cambridge, and Dr. H.
Jaeger, Humboldt Universitat, Berlin, loaned part of the holotype of Haplothecia
filata ; he also allowed a further section to be prepared from it.

Field-work, during which the type material of Macgeea touti was obtained,
has been supported by University of New England’s Research Grant No. 225.

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Puinip, G. M., 1962.—The palaeontology and stratigraphy of the Siluro-Devonian sediments
of the Tyers area, Gippsland, Victoria. Proc. roy. Soc. Vict., 75: 123-246, Pls 11-36.

RoézkowsKa, Maria, 1953.—Pachyphyllinae et Phillipsastraea du Frasnien de Pologne.
Palaeont. polon., 5.

ScHLOTHEIM, H. F., 1820.—“‘ Die Petrifactenkunde auf ihrem jetzigen Standpunkte durch die
Beschreibung semer Sammlung. . .erlautert.’’ Gotha.

ScHourPh, ALEXANDER, 1958.—Revision des Formenkreises um Phillipsastraea d’Orb., “‘ Pachy-
phyllum” E. and H., Macgeea (Webst.), ““ Thamnophyllum” Pen., Peneckiella Soshk. und
verwandter Formen. Neues Jb. Min. Geol. Paldont., 106: 139-244, Pls 5 6.

SosHxina, E. D., 1936.—Les coraux Rugosa du Dévonien moyen de L’Oural du Nord. Akad.
Nauk. S.S.S.R., Tr. polar Comm., 28: 15-76, Pl. 1.

, 1951.—Late Devonian rugose corals, their systematics and evolution (im Russian).
Akad. Nauk. S.S.S.R., Paleont. Inst. Trudy, 34.
, 1952.—Key to the Devonian tetracorals (in Russian). Ibid., 39.

Stumm, E. C., 1949.—Revision of the families and genera of the Devonian tetracorals. Mem.
geol. Soc. Amer., 40.

WALTHER, CONRAD, 1928.—Untersuchungen tiber die Mitteldevon—Oberdevongrenze. Z. dtsch.
geol. Ges., 80: 97-152.

Wane, H. C., 1950.—A revision of the Zoantharia Rugosa in the light of their minute skeletal
structures. Phil. Trans. (B), 234: 175-246, Pls 4-9.

WEssTER, C. L., 1889.—Description of a new genus of corals, from the Devonian rocks of Io wa.
Amer. Nat., 23: 710-712.

EXPLANATION OF PLATE VI
All figures x 2

Figs 1, 6, 7. Bensonastraea praetor, gen. et sp. nov., holotype. Figs 2-5, 8-11. Macgeea touti,
sp. nov. 2, 9, paratype 5: 3, 11, holotype; 4, paratype 2; 5, paratype 1; 8, paratype 6;
10, paratype 3.

SOME MITE PARASITES OF AUSTRALIAN BIRDS

ROBERT DOMROW
Queensland Institute of Medical Research, Brisbane

[Read 28th July, 1965]

Synopsis
Thirty-six species of mites from five families are listed from Australian birds.

LAELAPIDAE: New hosts are given for 13 species (Ornithonyssus bursa, O. sylviarum,
Pellonyssus reedi, Mesonyssus kakatuae, M. trichoglossi, M. belopolskiu, M. melloi, M. geopeliae,
Sternostoma cooremani, S. laniorum, S. thienponti, Ptilonyssus cractict and P. thymanzae). Seven
new records for Australia are listed: (Sternostoma tracheacolum, Rhinonyssus himantopus, R.
rhinolethrus, Larinyssus benoiti—a genus also new to Australia, Rallinyssus caudistigmus,
Passeronyssus bradypteri and Ptilonyssus triscutatus). Hight new species are described (Sternostoma
gliciphilae, n. sp., from Gliciphila indistincta, Meliphagidae ; S. zosteropus, n. sp., from Zosterops
lateralis, Zosteropidae ; Ptilonyssus microecae, n. sp., from Muicroeca fascinans and KHopsaltria
capito, Muscicapidae ; P. rhipidurae, n. sp., from Rhipidura fuliginosa, Muscicapidae ; P. dicaei,
n. sp., from Dicaeum hirundinaceum, Dicaeidae ; P. gliciphilae, n. sp., from Gliciphila indistincta,
Meliphagidae ; P. stomioperae, n. sp., from Stomiopera unicolor and Meliphaga flava, Meliphagidae ;
and Hattena panopla, n. sp., from Gliciphila indistincta, Meliphagidae.

SPELEOGNATHIDAE: Sypeleognathopsis benoiti and Neoboydaia merops are newly recorded
from Australia.

CHEYLETIDAE: The genus Neocheyletiella, represented by N. artami, n. sp., from Artamus
cyanopterus (Artamidae), is added to the Australian fauna.

TROMBICULIDAE: New records are provided for Odontacarus australiensis, Leptotrombidium
myzantha and Neoschoengastia posekanyt. Trombicula shiraii, known only from the original
series from Japan, is recorded from a migratory wader on the Great Barrier Reef.

TURBINOPTIDAE: The genus Passerrhinoptes, represented by P. pomatostomt, n. sp., from
Pomatostomus temporalis (Timaliidae), is listed from Australia for the first time.

Recent accessions have included the most interesting variety of bird-
parasitic mites detailed below. For further details on the Australian members
of the families discussed, the reader is referred to Domrow (1964a, b, ¢ ; 1965c)
on laelapids; Domrow (1965a) on speleognathids; Womersley (1941) and
Volgin (1964) on cheyletids ; Womersley (1952) on trombiculids ; and Domrow
(19656) on turbinoptids.

Messrs. D. P. Vernon and J. T. Woods, Queensland Museum, Brisbane,
have checked many bird identifications, and Mr. J. H. Calaby, C.S.1.B.O.,
Canberra, provided some of the material. The collectors, acknowledged by
their initials in the text, are, apart from myself, B. C. Allan, G. J. Barrow,
J. Booth, I. D. Fanning, R. H. Green, H. I. McDonald, M. D. Murray, J. M.
Paton, R. G. Rees, R. V. Southeott and J. S. Welch. I am most grateful to
them all, and to Miss B. Nolan for typing the manuscript.

The holotypes and allotypes of new species have been deposited in the
Australian National Insect Collection, C.S.I.R.O., Canberra ; paratypes, when
available, have been lodged in the collections under the care of Drs. A. Fain
(Prince Leopold Institute of Tropical Medicine, Antwerp), and R. W. Strandt-
mann (Texas Technological College, Lubbock), and myself.

Family LAELAPIDAE
ORNITHONYSSUS BURSA (Berlese)

Host records additional to those listed by Domrow (1963) are extremely
heavy infestations with both females and protonymphs on two Australian
black-shouldered kites, Hlanus notatus Gould (Accipitridae, Falconiformes),

PROCEEDINGS OF THE LINNEAN Soctery oF NEw SourH WALES, Vol. 90, Part 2

ROBERT DOMROW 191

14.vi.1963, I.D.F. and R.G.R. (occasional specimens were taken on several
other bird hosts with the same collection data, but as the risk of field
contamination is high with such an active species, they have not been listed
here); 1 protonymph from a fledgling laughing kookaburra, Dacelo gigas
(Boddaert) (Alcedinidae, Coraciiformes), Brisbane, 17.xii.1964, R.G.R.; many
females from around the vent of a pheasant coucal, Centropus phasianinus
(Latham), Samford, 21.1.1964, R.G.R. and J.S.W., and 1599 from a koel,
EHudynamys orientalis (Linnaeus), Brisbane, 8.11.1965, B.C.A. (both Cuculidae,
Cuculiformes) ; also 729 from a starling, Sturnus vulgaris Linnaeus (Sturnidae,
Passeriformes), Brisbane, 3.xii.1963, R.D. An interesting southerly record of
this, the tropical fowl mite, is 299 biting children, Launceston, Tas., 7.1.1963,
R.H.G.

ORNITHONYSSUS SYLVIARUM (Canestrini and Fanzago)

As all known Australian records of this species are from the far south-east
of the continent (Womersley, 1956a; Domrow, 1963), the following record
from Queensland is of interest: 19 from the welcome swallow, Hirundo neoxena
Gould (Hirundinidae, Passeriformes), Brisbane, 3.xii.1963, R.D. The Australian
H. neozena is a migratory species, which departs for the northern parts of the
continent in the autumn (Cayley, 1963). I have since seen 1399 from nestlings
of the blackbird, Turdus merula Linnaeus (Turdidae, Passeriformes) (introduced
from Europe, and now common in 8.H. Australia), Evendale, Tas., 5.1.1963,
R.H.G. Also 12 from a golden whistler, Pachycephala pectoralis (Latham)
(Pachycephalidae, Passeriformes), Esk, 14.vii.1965, R.D. and J.S.W.

PELLONYSSUS REEDI (Zumpt and Patterson)

Five 992 from the beautiful firetail, Zonaeginthus bellus (Latham)
(Ploceidae, Passeriformes), Waitpinga, S.A., 31.xii.1963, J.M.P., comprise the
second record of this species from Australia. See Womersley (19566) and Till
(1964). An additional synonym is Steatonyssus stenosternus Wang (1963).
I am most grateful to Dr. F. Zumpt, South African Institute for Medical
Research, Johannesburg, and the Director, South Australian Museum, Adelaide,
for the gift or loan of material of this genus.

MESONYSSUS KAKATUAE (Domrow)

One 2 and 14 from the nares of a red-tailed black cockatoo, Calyptorhynchus
banksi (Latham) (Psittacidae, Psittaciformes), Mitchell R., Gulf of Carpentaria,
x1.1964, R.D., comprise a new host record. See Domrow (1964a) and Wilson
(1964).

MESONYSSUS TRICHOGLOSSI (Domrow)

This species, previously recorded from several psittacids in Australia and
New Guinea (Domrow, 1964a; Wilson, 1964), may now be recorded from a
further Australian host: 299 from the nares of a little lorikeet, Glossopsitta
pusilla (Shaw) (Psittacidae, Psittaciformes), Esk, 27.11.1965, R.D. and J.S.W.

The specimens agree with the original description of the typical form,
except that only twelve furled setae are present on the dorsum: four along
midposterior margin of podosomal shield, one on shieldlet at each postero-
lateral angle of podosomal shield, and three on each half of posterior dorsal
shield (two on inner, and one on outer edge).

MESONYSSUS BELOPOLSKII (Bregetova)
Six 99, 103g and 2 nymphs from the nares of a pied heron, Notophoyax
picata (Gould) (Ardeidae, Ciconiiformes), Mitchell River, xi.1964, R.D.,
comprise a new host record. See Domrow (1965c).

192 SOME MITE PARASITES OF AUSTRALIAN BIRDS

MESONYSSUS MELLOI (de Castro)

Seven 99, 43g and 1 nymph from the nares of two domestic pigeons,
Columba livia Gmelin (Columbidae, Columbiformes), Brisbane, 2.xii.1964, R.D.
and J.S.W., comprise a new host record for Australia. See Fain (1962a) and
Domrow (1965c).

MESONYSSUS GEOPELIAE Fain

One 3 from the nares of a peaceful dove, Geopelia placida Gould (Colum-
bidae, Columbiformes), Mitchell R., xi.1964, R.D., comprises a new host record.
See Fain (1964) and Domrow (1965c).

STERNOSTOMA COOREMANI Fain

Two 99 from the nares of a rainbow-bird, Merops ornatus Latham (Mero-
pidae, Coraciiformes), Esk, 28.vili.1964, R.D. and J.S.W., comprise a new host
record. Previous records, all from coraciiforms, are summarized by Domrow
(1965c¢).

STERNOSTOMA GLICIPHILAE, N. Sp.
(Figs 1-5)

Female.—A small, oval mite with idiosoma wider in anterior half, 440-473.
long. Podosomal shield with anterolateral margins ill-defined, but postero-
lateral margins distinct and virtually straight medially. Shield almost entirely
covered by very sharply defined subhexagonal reticulation which is more
heavily sclerotized than remainder of shield, giving the effect of honeycomb.
Areas of muscle insertions nestle among this texture, and the shield further
bears five pairs of minute setae both laterally and medially. Opisthosomal
Shield rectangular, with greater axis longitudinal; texture similar to that of
podosomal shield; with about six minute setae. Dorsal cuticle otherwise
unarmed except for two small, circular stigmata (without peritremes).

Sternal shield discally imitating texture of podosomal shield, but reticula-
tion finer ; encircled by six sternal sete ; margins evanescent. Genital shield
short and broad, with nondescript texture and merest asetose traces of original
genital setae. Anal shield terminal, typically with at least adanal setae and
cribrum. Ventral cuticle typically with six minute setae arranged 4.2, but
minor variations were noted as follows: (i) five setae rather than four in
central group; and (ii) outer posterior pair of setae apparently lacking.

Coxal setal formula 2.1.1.1, with occasional asetose traces of posterior
seta on coxae II and III (aberrations from the typical 2.2.2.1 of the Gamasina
are apparently common in this genus, but they are of less than specific
importance (see Fain, 1957; Fain and Bafort, 1963b ; Furman, 1957 ; Hyland,
1962; Hyland and Clark, 1959; Hyland and Ford, 1961). Leg setation in
general extremely weak, but few longer setae on tarsi IJ-IV. Tarsi I with
several thicker setae dorsodistally. Claws I obsolescent ; claws II-IV strongly
curved, as is usual.

Gnathosomal and hypostomal setae apparently absent. Tritosternum
absent. Palpi with four free segments, line of demarcation between tibia and
tarsus indistinguishable. Palpal setation as figured. Chelicerae typical of
genus.

Discussion.—S. gliciphilae is quite unlike other species of the genus in its
peculiarly textured dorsal shield. It is, in addition, the only species known
from a peculiarly Australian group of birds—only one species of the family
Meliphagidae has crossed Wallace’s Line to the west, see Leach [1958], whose
classification is used in this paper.

Types.—Holotype female and two paratype females from the mucous
membranes at the extreme posterior of the nares of a brown honeyeater,
Gliciphila indistincta (Vigors and Horsfield) (Meliphagidae, Passeriformes), Esk,
16.1.1965, R.D. and J.S.W. Holotype NIC; paratypes RD.

ROBERT DOMROW 193

ak

‘Cay
PeN
y> ‘of

X

ew

RD
JSW

Figs 1-5. Sternostoma gliciphilae,n.sp. Female.—l, Idiosoma (dorsal) ; 2, Idiosoma (ventral) ;
3, Leg III (ventral at left, dorsal at right); 4, Tarsus I (anterior above, posterior below) ;
5, Gnathosoma (ventral, with left palp dorsal). (Hach division on the scales equals 100u.)

STERNOSTOMA LANIORUM Fain

Four 9° from a crested bellbird, Oreoica gutturalis (Vigors and Horsfield)
(Falcunculidae, Passeriformes), Mitchell, S.Q., 25.v.1964, I.D.F., and 192 from
a rufous shrike-thrush, Colluricincla megarhyncha (Quoy and Gaimard) (Pachy-
cephalidae, Passeriformes), mist-netted along the Innisfail-Palmerston Highway,
16.xii.1964, H.I.McD. and G.J.B., comprise new host records (other birds listed
below from this latter locality were also netted, while almost all the others
were shot). All specimens were taken from the nares, and the ventrodistal
setae on tarsi II-IV are blunt in both series (see Fain, 1957, and Domrow,

E

194 SOME MITE PARASITES OF AUSTRALIAN BIRDS

1965c). Also 19 from a leaden flycatcher, Myiagra rubecula (Latham), a d
9°92 from a pale-yellow robin, Hopsaltria capito Gould (both Muscicapidae,
Passeriformes), Innisfail, 3 and 4.vili.1965, R.D. and J.S.W.

STERNOSTOMA THIENPONTI Fain

One 2 from the nares of a black butcher-bird (black phase), Cracticus quoyt
(Lesson and Garnot) (Cracticidae, Passeriformes), Innisfail-Palmerston Highway,
11.xii.1964, H.I.McD. and G.J.B., comprises the second Australian record of
this species, again from a cracticid (see Domrow, 1965c). Also 1999 from a
black butcher-bird (red phase), Innisfail, 1.vi.1965, G.J.B. and H.I.McD.

STERNOSTOMA TRACHEACOLUM Lawrence

This widespread species may now be formally recorded from Australia
(see Domrow, 1965c): 2199 from the trachea of the Gouldian finch, Poephila
gouldae (Gould) (Ploceidae, Passeriformes), Sydney, 19.x.1964, M.D.M. Previous
records are summarized by Fain and Hyland (1962).

STERNOSTOMA ZOSTEROPUS, Nl. Sp.
(Figs 6-12)

Female.—A small, oval mite with idiosoma shaped as in S. gliciphilae,
n. sp., 402 long in little deformed specimen figured, 495u. in second somewhat
compressed specimen. Podosomal shield sharply arched anteriorly, sinuous
laterally and posteriorly ; surface heavily granulate except at extreme margins,
marked by muscle insertions and bearing four pairs of setae both marginally
and medially, in addition to two pairs of lateral pores. Opisthosomal shield
subquadrate, of similar texture to podosomal shield. One specimen shows
two stronger setae both anteriorly and posteriorly, as well as six smaller setae
discally. The other shows two stronger setae posteriorly, two setae and perhaps
four asetose indications of setae discally, while it is flanked anteriorly by an
unpaired, stronger seta. Shield with pore in each posterolateral angle. Dorsal
body cuticle with two distinct setae behind stigmata, which latter have no
peritremes.

Sternal shield weakly granulate except for textureless, but rather well
defined margins; bearing six blunt setae. Genital shield short and broad,
weakly granulate, with muscle insertions and truncate, rayed operculum.
Genital setae obsolescent. Anal shield terminal, with at least adanal setae
and cribrum present. Ventral cuticle with four distinct setae.

Coxal setal formula 2.2.2.1, setae weak, as they are on all segments except
tarsi. Tarsus I with sensory islet dorsodistally showing both longer setae and
rodlets. Tarsi II-IV with setae arranged slightly differently from S. gliciphilae,
showing the pattern typical of Ptilonyssus ; all setae, except one, like elongate
droplets. Claws I obsolescent ; claws II-IV straight along most of their length,
eurved only distally.

Gnathosomal and hypostomal setae apparently absent. Tritosternum
lacking. Palpi with four movable segments, the line of demarcation between
tibia and tarsus virtually invisible; setation as figured. Chelicerae typical
of genus.

Discussion.—S. zosteropus may be separated from all its described con-
geners by its peculiarly straight tarsal claws I-IV.

Types.—Holotype female and one paratype female from the nares of a
erey-backed 'silvereye, Zosterops lateralis (Latham) (Zosteropidae, Passeriformes),
mist-netted at Mt. Jukes, Mackay, vi.1964, R.D. and J.S.W. Holotype NIC;
paratype RD. ;

ROBERT DOMROW 195

Figs 6-12. Sternostoma zosteropus, n. sp. Female.—6, Idiosoma (dorsal) ; 7, Idiosoma (ventral) ;
8, Leg III (ventral at left, dorsal at right); 9, Ambulacrum III (two views); 10, Gnathosoma
(ventral, with right palp dorsal); 11, Opisthosomal shield (variant); 12, Tarsus I (ventral at
left, dorsal at right).

RHINONYSSUS HIMANTOPUS Strandtmann

This widespread parasite of waders may now be listed from Australia :
19 from a red-kneed dotterel, Hrythrogonys cinctus Gould; and 29° and 3¢¢
from a masked plover, Lobibyx miles (Boddaert) (both Charadriidae, Charadrii-
formes), Mitchell R., xi.1964, R.D. All specimens were collected in the nares.
The former series resembles Strandtmann’s original (1951) specimens from
Himantopus (Recurvirostridae, Charadriiformes), and the latter his later (1959)
specimens from Charadrius. I am grateful to Dr. Strandtmann for the loan
of specimens of this species, of which I have since seen 1099, 4g¢ and 1 nymph
from black-fronted dotterels, Charadrius melanops Vieillot, Mitchell R., iv.1965,
R.D., and 1¢ from a spur-winged plover, Lobibyx novaehollandiae (Stephens),
Esk, 16.v.1965, I.D.F. and J.S.W.

196 SOME MITE PARASITES OF AUSTRALIAN BIRDS

RHINONYSSUS RHINOLETHRUS (Trouessart)

This widespread parasite of anseriforms may now be recorded from
Australia: 299 from the nares of a whistling tree-duck, Dendrocygna arcuata
(Horsfield) (Anatidae, Anseriformes), Mitchell R., xi.1964, R.D. It has also
been recorded from the black duck, Anas superciliosa Gmelin, in New Guinea
(Wilson, 1964), but I have as yet no such Australian record.

LARINYSSUS BENOITI Fain
(Figs 13-21)

This genus and species may now be listed from Australia: 3399, 9¢¢
and 2 nymphs from the nares of five Australian pratincoles, Stiltia isabella
(Vieillot) (Glareolidae, Charadriiformes), Mitchell R., xi.1964, R.D. The only
previous record is from Galachrysia, an African glareolid. Dr. Fain has kindly
compared my illustrations with his specimens, and confirmed my identification.

RALLINYSSUS CAUDISTIGMUS Strandtmann

This species, known only from American rallids (Strandtmann, 1948),
may now be recorded from Australia: 699 and 1¢ from the nares of a dusky
moorhen, Gallinula tenebrosa Gould (Rallidae, Gruiformes), Esk, 27.11.1965,
R.D. and J.S.W.

PASSERONYSSUS BRADYPTERI Fain

This species may now be listed from Australia: 499° from the nares of
a rufous songlark, Cinclorhamphus mathewsi Iredale (Sylviidae, Passeriformes),
Hsk, 29.viii.1964, R.D. and J.S.W. The only previous record is from Bradypterus,
an African, sylviid (Fain, 1962b).

PTILONYSSUS TRISCUTATUS (Vitzthum)

This parasite of European and African bee-eaters (see Fain, 1957) may now
be recorded from Australia: 19 from the nares of a rainbow-bird, Merops
ornatus Latham (Meropidae, Coraciiformes), Esk, 29.vii.1964, R.D. and J.S.W.
The dorsum of femur III of this specimen shows an oblique row of three
closely-set setae reminiscent of genu III in Tyranninyssus Brooks and Strandt-
mann (1960). See also Hyland (1961).

PTILONYSSUS CRACTICI Domrow

Seven 99, 1 deutonymph and 1 protonymph from a black-backed butcher-
bird, Cracticus mentalis Salvadori and d’Albertis (Cracticidae, Passeriformes),
Chillagoe, 2.1.1965, G.J.B., comprise a new host record. See Domrow (1964c).
I have since also seen 499 and 1 deutonymph from the nares of a white-winged
triller, Lalage tricolor (Swainson) (Campephagidae, Passeriformes), Mitchell R.,
7.iv.1965, R.D., which I assign to this species. They differ from paratypes
only in showing the adanal setae set just behind the anus, and slight processes
at the anterior edges of coxae II and the palpal trochanters (the former process
is absent, and the latter incipient, in paratypes). Also 499 and 1 protonymph
from a grey butcher-bird, Cracticus torquatus (Latham), Esk, 5.x.1965, R.D. and
J.S.W. Finally, 12 from a laughing kookaburra, Dacelo gigas (Boddaert)
(Alcedinidae, Coraciiformes), Esk, 14.vii.1965, R.D. and J.S.W. This seems
an abnormal host, and it is recognized that ‘“‘ many records of stragglers are
simply curiosities, though their publication should presumably not be suppressed ”’
(Audy, 1956, Bull. Raffles Mus., Singapore, 28: 74).

PTILONYSSUS MICROECAE, 0. Sp.
(Figs 25-26, 28-31)
Female.—An elongate mite with idiosoma about 850-880 long in mounted,
rather compressed specimens. Podosomal shield about one and a half times

ROBERT DOMROW 197

Figs 13-18. Larinyssus benoiti Fain.—13, Idiosoma @ (ventral); 14, Idiosoma 3 (ventral) ;

15, Idiosoma ¢ (dorsal) ; 16, Chelicera 2 (lateral) ; 17, Leg III 9 (dorsal) ; 18, Leg III 9 (ventral).

198 SOME MITE PARASITES OF AUSTRALIAN BIRDS

19

| Nol ge

Figs 19-21. Larinyssus benoitd Fam.—I19, Leg IV 2 (ventral); 20, Leg I @ (ventral) ;
21, Gnathosoma ¢ (ventral, with left palp dorsal).

Figs 22-24. Ptilonyssus thymanzae Domrow. Female.—22, Idiosoma (ventral, M. chrysops) ;
23, Idiosoma (dorsal, WM. chrysops); 24, Whole mite 9 (freehand, M. notata).

Figs 25-26. Ptilonyssus microecae, n.sp. Kemale.—25, Gnathosoma (ventral, with left palp
dorsal, Muicroeca); 26, Chelicera (lateral, Mzcroeca).

Fig. 27. Ptilonyssus rhipidurae, n.sp. Kemale.—Gnathosoma (ventral, with left palp dorsal).

as long as wide, and slightly wider in anterior half (223-228 x 154-161u) ;
with anterior and posterior margins nondescript and subequal, and lateral
margins tending to convexity in anterior half. Shield not strongly outlined,
very minutely granulate, with weakly marked muscle insertions and sixteen
paired setae (in specimen figured, seta marked X is somewhat displaced to the
front). Peritremalia and adjacent setae as in P. rhipidurae, n. sp. Middorsum
with eight setae, of which midanterior pair is set between posterior of two pairs

ROBERT DOMROW 199

of shieldlets. Hysterosoma with entire pygidial shield bearing traces of muscle
insertions, at least one pore and two spinose pygidial setae ; surrounded by six
setae arranged 4.2. All dorsal (and ventral) setae tapering to point somewhat
stronger on posterior half of body.

Sternal shield elongate, very weakly defined and textureless, bearing two
pores and six marginal setae. Genital shield shorter, somewhat flared
posteriorly ; lateral margins not heavily sclerotized, bearing two genital setae ;
disc denser, with granulations and muscle insertions ; operculum rayed. Anal
shield almost twice as long as wide (125 x 74u in holotype, 119 x 64u in
paratype), with anterior margin arched and lateral margins rather straight and
sclerotized ; cribrum present. Anus set well forward, with adanal setae level
with its anterior; postanal seta present. Ventral cuticle with eight setae
arranged 2.6 between genital and anal shields, and latter flanked by four
additional setae.

Leg segments with setation as follows: coxae 2.2.2.1; trochanters 4.4.4.5 ;
femora 9.7.4.5; genua 6.6.6.3; tibiae 7.7.6.6 (5 on one side of one specimen) ;
tarsi —.15.15.15 (excluding two very fine terminal setae). Leg setae resembling
those on coxae, slightly smaller dorsally ; two setae on dorsum of genu III set
in enlarged alveoli; two ventrodistal setae on tarsi II-IV slightly stronger.
Tarsus I with dorsodistal sensory zone. Ambulacra I not greatly modified.
Coxa II without process on anterodorsal margin.

Gnathosomal setae subequal to inner posterior hypostomals, slightly weaker
than anterior hypostomals ; outer posterior hypostomals minute. Deutosternum
with about nine denticles in single file. Chelicerae attenuate in distal two-
thirds, with chelate portion occupying one-thirty-fifth of total length. Palpal
setal formula 1.2.4.8 (including two dorsodistal tibial rods). Palpal trochanter
distinctly salient on inner ventrodistal angle. Tarsus with about eight minute
setae ; claw not detected. Tritosternum absent.

Discussion.—P. microecae recalls P. motacillae Fain, both possessing a
saliency on the palpal trochanter, but the new species may be easily separated
from Fain’s by the shape and setation of the podosomal shield, the absence
of a process on coxa II and (possibly) by the condition of the pygidial shield.

The few Malayan specimens from Poliomyias mugimaki (Temminck) (listed
by McClure, 1963, as Muscicapa mugimaki) recorded by Fain and Nadchatram
(1962) have the podosomal shield exactly as in P. motacillae and possess a process
on coxa II. Further, at least in the specimen I have examined (through the
courtesy of Dr. Fain), the pygidial shield is in a semidivided condition, being
eroded midposteriorly. George (1961) has reported that the pygidial shield,
while normally divided, may occasionally be entire in P. echinatus Berlese and
Trouessart, while the opposite is true of two of thirteen females of P. thymanzae
Domrow (1964c), Myzantha melanocephala (type host), Esk, 29.vii.1964, R.D.
and J.S.W. I would tend to consider these Malayan specimens merely as
variants of the widespread P. motacillae.

Types.—Two females were collected—one (holotype) from the nares of a
Jacky winter, Microeca fascinans (Latham), Esk, S.H.Q., 8.11.1964, R.D., I.D.F.
and J.S.W.; and one (paratype) from a pale-yellow robin, Hopsaltria capito
Gould, Mitchell River, xi.1964, R.D. Both hosts are muscicapids (Passeri-
formes). Holotype NIC; paratype RD.

PTILONYSSUS RHIPIDURAE, 0D. Sp.
(Figs 27, 32-35)

Female.—An elongate mite with idiosoma 693u long in one unengorged
and relatively uncompressed specimen, 781-869. in replete specimens.
Podosomal shield slightly longer than wide (172-178 x 143-156u) ; anterior
margin slightly concave, lateral margins convex and posterior margin weakly
trilobed. Shield minutely granulate, with muscle insertions, including two
posterolateral zones, weakly marked ; with twelve evenly arranged setae on

200 SOME MITE PARASITES OF AUSTRALIAN BIRDS

Figs 28-31. Ptilonyssus microecae, n.sp. Female.—28, Idiosoma (ventral, Hopsaltria) ;
29, Idiosoma (dorsal, Hopsaltria) ; 30, Leg III (ventral, Microeca) ; 31, Leg ITI (dorsal, Microeca).
Figs 32-35. Ptilonyssus rhipidurae, n.sp. Female.—32, Idiosoma (ventral); 33, Leg III
(ventral) ; 34, Leg III (dorsal) ; 35, Idiosoma (dorsal).

ROBERT DOMROW 201

shield, which is also preceded and followed by two setae. Five setae arranged
1.1.3 on each side between shield and peritremalia, which latter are as in
P. dicaei, n. sp., but with poststigmatic shields present. Middorsum with
band of ten setae, of which midanterior pair is between posterior of two pairs
of shieldlets. Hysterosoma with row of six setae and two discrete, subcircular
pygidial shields (each with pore and spinose seta, and flanked posterolaterally
by one or two setae). All dorsal setae, except pygidials, minute rods; setae
on podosomal shield rather smaller than those on cuticle.

Sternal shield elongate, virtually textureless, but fairly well defined, bearing
four pores and flanked by six setae. Genital shield narrow, with muscle
insertions amidst longitudinal fluting ; operculum weakly rayed; two genital
setae and accompanying pores flank shield posterolaterally. Anal shield
almost three times as long as wide (128-143 x 50-54u), strongly arched
anteriorly and slightly concave laterally ; lateral margins strongly sclerotized ;
cribrum present, slightly expanded. Anus set well forward, preceding all three
subequal anal setae. Ventral cuticle with six setae arranged 2.4 between genital
and anal shields, and posterolaterally with 14 to 16 additional setae. Of setae
on ventral cuticle and shields, only genitals are somewhat blunt, while remainder
all taper to sharp point.

Leg segments with setation as follows: coxae 2.2.2.1; trochanters 4.4.4.5 ;
femora 9.7.5.5; genua 6.6.6.3; tibiae 7.7.6.6; tarsi —.15.15.15 (excluding two
extremely fine terminal setae). Setae on ventral face of segments (except on
tarsi) tapering to point, resembling those on coxae; dorsal setae rather weaker.
Some distal setae on tarsi II-[V somewhat bluntened. Tarsus I with dorso-
distal sensory zone. Ambulacra I little modified. Coxa II with process on
anterodorsal margin.

Gnathosomal setae twice as strong as posterior, and three times as strong
as anterior hypostomals, all rather blunt. Deutosternum with about ten
denticles in single file. Chelicerae attenuate in distal half, chelate portion
occupying one-twenty-fifth of total length. Palpal setal formula 1.2.4.(7),
two dorsodistal tibial rods included. Tarsus with about eight setae; claw
not detected. Tritosternum absent.

Discussion.—P. rhipidurae immediately calls to mind P. macclurei Fain,
recorded from Rhipidura albicollis in Malaya and R. leucophrys in Australia
(Fain, 1963a ; Domrow, 1964c). The two species may, however, be readily
separated by the number of anal setae and the condition of the pygidial shield.
Further, in P. macclurei, the setae of the idiosomal venter (coxae included)
are stronger. (I might add that the merest traces of poststigmatic shields
are present in P. macclurei, of which I have since taken two further series from
R. leucophrys at Esk and Brisbane, 8 and 20.ii.1964, respectively.)

Types.—Four females were collected from the nares of a grey fantail,
Rhipidura fuliginosa (Sparrman) (Muscicapidae, Passeriformes), Esk, 25.vii.1964,
R.G.R. and J.S.W. Holotype NIC; paratypes RD and AF.

PTILONYSSUS DICAEI, n. sp.
(Figs 36-44)

Female.—An elongate mite with idiosoma 960 and 1000y long in two
mounted, slightly compressed specimens. Podosomal shield one and a half
times as wide as long (185 x 138 and 196 x 143u); anteromedial margin
slightly concave, anterolateral angles strongly convex ; posterior quarter of
shield much narrower, with outline more irregular and slightly concave
posteriorly. Shield with two pores and twelve setae, all paired (one of two
setae marked ‘‘ X ” lacking on one side of one specimen) ; also two setae, both
vertically and at anterolateral angles, set just off shield. Two closely-set setae
immediately behind, and two groups of four setae between peritremalia and

202 SOME MITE PARASITES OF AUSTRALIAN BIRDS

posterolateral angles of shield. Middorsum with four shieldlets and eight setae
arranged 2.6 in addition to two between posterior shieldlets. Pygidial shield
convex anteriorly and concave posteriorly, with two pores and two spinose
pygidial setae ; surrounding cuticle with eight setae arranged 4.4. Setae on
podosomal shield weaker than remaining dorsal setae, perhaps slightly more
spinose than figured. Both dorsal shields quite well defined, minutely granulate

RD
JSW

Figs 36-44. Ptilonyssus dicaei, n.sp. Female.—36, Idiosoma (dorsal, freehand); 37, Podo-
somal shield, peritremalia and mid-dorsal shieldlets ; 38, Coxae II-IV, and sternal and genital
shields ; 39, Whole mite (freehand, ventral) ; 40, Anal shield; 41, Pygidial shield; 42, Coxa I
(ventral) ; 43, Chelicera (lateral); 44, Gnathosoma (ventral).

and with weak indications of muscle insertions. Each stigma provided with
short peritreme, surrounded by very weak shieldlet. Poststigmatic shields
absent.

Sternal shield elongate, weakly defined, virtually textureless and bearing
SI and four pores; SII and III free in cuticle. Genital shield narrow, not
reaching beyond posterior margin of coxae IV ; granulate, with weakly rayed
operculum and merest traces of muscle insertions ; flanked subposteriorly by
two genital setae. Anal shield twice as long as wide (107 x 54u in specimen
with smaller podosomal shield), with anterior margin weakly defined and fairly
straight ; lateral margins also straight, but more strongly sclerotized ; cribrum

ROBERT DOMROW 203

present. Adanal setae near anterior of anus in specimen figured, but nearer
posterior in second specimen. Postanal seta slightly stronger than adanals.
Ventral cuticle with eight setae arranged 2.6 in front of, and six setae behind
anus.

Leg segments with setation as follows: coxae 2.2.2.1; trochanters 4.4.4.3 ;
femora 9.7.4.5; genua 6.6.5.4; tibiae 7.7.6.6; tarsi —.15.15.15 (excluding two
extremely fine terminal setae closely associated with base of ambulacral stalk).
Setae on ventral face of segments tapering, resembling those on coxae (two at
apices of tarsi [I-IV stronger) ; those on dorsum blunter and very much weaker.
Tarsus I with dorsodistal sensory zone. Ambulacra I more slender than IJ-IV ;
claws I slightly weaker than II-IV, little modified in shape. Coxa II without
process on anterodorsal margin.

Gnathosomal setae slightly stronger than all three pairs of hypostomal
setae, of which inner posteriors are longest and outer posteriors shortest.
Deutosternum with about ten denticles in single file. Chelicerae attenuate
in distal half, with chelate portion occupying one-thirtieth of total length.
Palpal setal formula 1.2.4.8 (including two dorsodistal tibial rods). Tarsus
with about eight minute setae; claw seemingly present under oil-immersion,
but extremely weak. Tritosternum absent.

Discussion.—The Old World and Australian nectar eaters, ‘‘a group of
about 400 species entirely confined to the Old World and scarcely entering
the north-temperate zone even there” (Darlington, 1957), comprise the four
families Dicaeidae (flowerpeckers), Nectariniidae (sunbirds), Meliphagidae (honey-
eaters) and Zosteropidae (silvereyes) (Mayr and Amadon, 1951). All four
families are now known to be parasitized by an apparently closely related group
of species of Ptilonyssus with the genital shield so narrowed that the genital
setae, normally set on the shield itself, are left free in the adjacent cuticle.

Mayr and Amadon place the dicaeids next to the nectariniids, noting that
their distributions are complementary, the former being Oriental-Australian and
the latter African-Oriental, with only one species reaching Australia. One
species of Ptilonyssus, P. cinnyris Zumpt and Till, has been described from
African sunbirds, and may easily be separated from P. dicaet by having the
podosomal shield decidedly longer than wide, with ‘‘a pair of conspicuous
bristles on its posterior border”’, and lacking the pygidial shield (fide Zumpt
and Till, 1955; Fain, 1957). Dr. Zumpt has since kindly loaned me two
paratype females of P. cinnyris, and, while they are much overcleared, they
show, in addition to the two setae noted above, a pair of strong setae on each
side between the podosomal shield and peritremalia. This recalls such species
as P. andropadi Fain, P. calamocichlae Fain, P. chlorocichlae Fain, P. ruandae
Fain, P. prunellae Fain and Bafort, P. pittae Domrow and P. psophodae
Domrow (see Fain, 1957, 1963a; Fain and Bafort, 1963); Domrow, 1964b).

Of the species of Ptilonyssus described from meliphagids, an essentially
Australian family, P. lymozemae Domrow (1965c) shows a setal pattern on the
podosomal shield most closely approaching that of P. dicaei (allowing for the
minor movement of the vertical and extreme anterolateral pairs onto the
shield proper). However, P. lymozemae shows obsolescent, divided pygidial
shields in contradistinction to the fully-formed shield of P. dicaei.

P. ruandae Fain (1956a, 1957) is the only species of Ptilonyssus recorded
from silvereyes, which are common in all three African, Oriental and Australian
regions. This species, recorded both from Africa and Australia, shows a
podosomal shield similar to that of P. dicaei in shape, with two anterolateral
pores and an extremely similar setal pattern, both on and about the shield.
Both species further possess entire pygidial shields and are, I believe, closely
related. P. ruandae, however, has a more starkly cruciform podosomal shield,
and exhibits several pairs of very strong dorsal setae quite absent in P. dicaet.

204 SOME MITE PARASITES OF AUSTRALIAN BIRDS

Types.—Three females were collected from the nares of a mistletoe-bird,
Dicaeum hirundinaceum (Shaw) (Dicaeidae, Passeriformes), mist-netted in
brushland at Mt. Jukes, near Mackay, N.Q., vi. 1964, R.D. and J.S.W. Holo-
type NIC; paratype RD. The third specimen, which was not taken into
account in the above description, is in the care of Dr. Fain.

PTILONYSSUS THYMANZAE Domrow
(Figs 22-24)

Three 2° and 1 protonymph from two yellow-faced honeyeaters, Meliphaga
chrysops (Latham), Samford, 18.1. and 8.v.1964, R.D., I.D.F. and J.S8.W. ;
19 from a Lewin honeyeater, Meliphaga lewint Swainson, Esk, 27.ii.1965, R.D.
and J.S.W.; and 622 and 1 protonymph from a lesser Lewin honeyeater,
Meliphaga notata (Gould), Innisfail-Palmerston Highway, 20.1.1965, H.I.MecD.
and G.J.B. (all Meliphagidae, Passeriformes), comprise new host records. All
specimens have podosomal shields resembling that of the male sex figured by
Domrow (1964c), while the females, especially of the first two series, are
grossly engorged, with lobate body contours as in P. meliphagae Domrow, the
anal and pygidial shields being just to the fore of the ventrally directed,
posterior opisthosomal lobe (Figs 22-23). The midlateral dorsal lobes were
seen to be erect and conical in life in the latter series (Fig. 24). The mouth-
parts of a female from M. chrysops are figured in Domrow (1965c). See also
the above discussion on P. microecae, i. sp.

PTILONYSSUS GLICIPHILAE, 0. Sp.
(Figs 45-51)

Female.—An elongate mite, but idiosomal length unavailable because of
rupture during mounting procedure. Posterior margin of hysterosoma distinctly
bilobed in one specimen. Podosomal shield one and a half times as long as
wide (258 x 165u); anteromedial margin ‘‘M’’-shaped, anterolateral angles
strongly convex ; lateral margins concave, but shield expanding towards convex
posterior margin. Shield with four pores and twelve setae, all paired (one
posterior pore lacking on one side of one specimen) ; flanked midlaterally by
two shieldlets. About six pairs of setae between shield and peritremalia, which
latter are as in P. dicaei, n. sp. Middorsum with four shieldlets and 10 setae
arranged 4.6 (not figured) in addition to two between posterior shieldlets.
Pygidial shield much as in P. dicaei, both flanked and followed by one pair
of setae. Both dorsal shields well defined, shagreened and marked by muscle
insertions.

Sternal shield elongate, weakly defined, virtually textureless, and bearing
SII and two pores; SI and III free in cuticle. Genital shield narrow, not
reaching beyond posterior margins of coxae IV ; with longitudinally arranged
granulations, weakly rayed operculum and muscle insertions; flanked sub-
posteriorly by two genital setae. Anal shield three times as long as wide
(190 x 58u), with anterior margin very strongly, and lateral margins only
slightly convex; disc weakly granulate, but cuticle shagreened laterally ;
elongate cribrum present. Anal setae weak, particularly postanal ; all behind
anus. Two setae (not figured) on ventral cuticle between genital and anal
shields, and latter shield surrounded by setae arranged 11.11 and 12.13, one
of subposterior pairs being quite weak, cf. P. myzanthae Domrow, 1964), also
a parasite of meliphagids.

Leg segments with setation as follows: coxae 2.2.2.1; trochanters 4.4.4.5 ;
femora 9.8.5.5; genua 6.7.7.5 (4 on one side of one specimen) ; tibiae 7.7.7.7 ;
tarsi —.15.15.15 (excluding two extremely fine terminal setae closely associated
with base of ambulacral stalk). Setae on ventral face of segments rather
similar to those on coxae (two at apices of tarsi II-ITV somewhat hypertrophied
basally, with dorsally directed, filamentous apical portion at right angles to

ROBERT DOMROW 205

Shaft proper) ; those on dorsum considerably weaker, especially on legs I and
II. Tarsus I with dorsodistal sensory zone. Ambulacra and claws much as
in P. dicaei. Coxa II with strong process on anterodorsal margin.

Gnathosomal setae slightly stronger than subequal posterior hypostomals ;
anterior hypostomals extremely weak. Deutosternum with about eight minute
denticles in single file. Chelicerae attenuate in distal half, with chelate portion
occupying one-thirtieth of total length. Palpal setal formula 1.2.4.8 (including
two dorsodistal tibial rods). Tarsus with about seven minute setae ; claw not
detected, even under oil-immersion. Tritosternum absent.

RD
JSW

Figs 45-51. Ptilonyssus gliciphilae, n.sp. Female.—45, Coxae, sternal and genital shields ;
46, Gnathosoma (ventral, with right palp dorsal) ; 47, Podosomal shield, peritremalia and mid-
dorsal shieldlets ; 48, Pygidium (dorsal) ; 49, Setae from tarsus IV (freehand) ; 50, Tarsus IV
(ventral) ; 51, Pygidium (ventral).

Discussion.—Two other species of Ptilonyssus with accessory shieldlets
flanking the podosomal shield are known from Australian meliphagids, P.
thymanzae Domrow and P. meliphagae Domrow (1964c), but these have the
podosomal shield, both in its shape and setation, quite different from that of
P. gliciphilae. P. gliciphilae further differs (i) from P. thymanzae by the
position of the adanal setae; and (ii) from P. meliphagae by the contours of
the hysterosoma.

Types.—Two females were collected from the nares of brown honeyeaters,
Gliciphila indistincta (Vigors and Horsfield) (Meliphagidae, Passeriformes), one

206 SOME MITE PARASITES OF AUSTRALIAN BIRDS

mist-netted in mangroves at Chelona, near Sarina, vi.1964; and one shot in
flowering red bottle-brush (Callistemon viminalis), Esk, 15.x.1964, both R.D.
and J.S.W. Holotype (the Chelona specimen) NIC; paratype RD.

PTILONYSSUS STOMIOPERAE, Nl. Sp.
(Figs 52-61)

Female.—An elongate mite with idiosoma 1,045-1,2871 long in four
unengorged, relatively slightly compressed specimens (three from Meliphaga,
one from Stomiopera), 1,386 and 1,529% in two replete specimens from
Stomiopera. Specimens from Stomiopera show a larger podosomal shield
with antero- and posterolateral lobes well developed as in Figure 59. Three
specimens show this format, the shield measuring 446-459 x 366-379n.
Two show the following aberrancies: one vertical seta on shield (Fig. 60) and
one vertical and one body seta ‘“‘X” on shield (Fig. 58). In both these
specimens, the podosomal shield is longer (464), but wider (402u) in the former
and narrower (348) in the latter. A third aberrancy (Fig. 57) involves the
loss of one posterolateral lobe, narrowing the shield to 459 x 324u (measure-
ments overall, as throughout this paper). Specimens from Meliphaga show
the antero- and posterolateral lobes reduced, resulting in a smaller shield,
typically 379-402 x 276-299 in three specimens (Fig. 52). The fourth
specimen is aberrant, with the shield even smaller (370 x 264y.), showing an
increased insularity of the posterolateral lobes, leaving one shield seta marginal
and the other free in the cuticle (Fig. 61). Shield well defined, distinctly
Shagreened and with muscle insertions particularly strongly marked in specimens
from Stomiopera; bearing two usually closely-set setae anteriorly and two
submarginal setae posterolaterally. Shield preceded by two vertical setae,
flanked laterally by three pairs of setae and followed by two setae. Five
additional setae arranged 1.1.3 present on each side between posterolateral
lobes and peritremelia, which latter are contained in weakly sclerotized
shieldlets. Middorsum with band of ten setae, of which midanterior pair is
set between posterior of two pairs of shieldlets. Hysterosoma with about
twelve setae surrounding pygidial shield, which is convex anteriorly and concave
posteriorly, with muscle insertions and at least one pore laterally, and two
pygidial setae posteriorly. In specimens from Meliphaga (Fig. 52), the shield
is wider and somewhat irregular in outline; in specimens from Stomiopera
(Fig. 59), it is narrower and more compact. All dorsal setae, including pygidials,
particularly strong, except for verticals, those on, and one or two pairs flanking
podosomal shield anterolaterally.

Sternal shield elongate, with extremely weak granulations and ill-defined
margins ; flanked by two pores and six setae. Genital shield narrow, distinctly
granulate, with muscle insertions and rayed operculum ; flanked subposteriorly
by two setae and attendant pores. Anal shield slightly more than one and
a half times as long as broad (219-233 x 120-125 in three specimens from
Meliphaga and two from Stomiopera ; three other specimens from Stomiopera
are 240-250 x 129-147u); margins evenly rounded anterolaterally and fairly
straight posterolaterally ; cribrum present. Anus well forward, set in front
of all three anal setae. Ventral cuticle with eight setae arranged 2.6 between
genital and anal shields, which latter is flanked posterolaterally by an additional
ten setae. Ventral setae also strong with exception of genitals and anals.

Leg segments with setation as follows: coxae 2.2.2.1 ; trochanters 4.4.4.5 ;
genua 7.6.7.5; tibiae 7.7.7.7; tarsi -.15.15.15 (excluding two extremely fine
terminal setae). Femora variable, 9.8.7 (6 on one side of one specimen) .6 in
series from Meliphaga, and 9.8.8.6 (7 on one side of two specimens) in series
from Stomiopera. Setae on ventral face of segments similar to those on coxae,
but dorsal setae generally weaker. Two ventral setae at apices of tarsi II-IV
with tips suddenly constricted and angulate, ef. P. gliciphilae, n. sp. Tarsus

ROBERT DOMROW 207

“D3 ,,US

AV
SY

i)

RD 5
JSW

Figs 52-61. Ptilonyssus stomioperae, n. sp. Female.—52, Idiosoma (dorsal); 53, Idiosoma
(ventral) ; 54, Gnathosoma (ventral, with right palp dorsal) ; 55, Leg III (ventral) ; 56, Leg III
(dorsal) ; 57—61, Podosomal shield (variants, 59 with inset of pygidial shield). (Figs 57-60
Stomiopera, remainder Meliphaga.)

I with dorsodistal sensory zone. Ambulacra I more slender than IJ-IV.
Coxae II with process on anterodorsal margin.

Gnathosomal setae slightly smaller than inner posterior hypostomals ;
outer posterior and anterior hypostomals smaller still. Deutosternum with
about ten denticles in single file. Chelicerae attenuate in distal two-thirds,
chelate portion occupying one-thirty-fifth of total length. Palpal setal formula
typically 1.3.4.8 (including two dorsodistal tibial rods), but may be one fewer
setae on femur and/or genu. Tarsus with about six minute setae and weakly
bifid claw. Tritosternum absent.

Discussion.—In showing the chelicerae suddenly attenuate distally and
the pygidial shield entire and well developed, P. stomioperae is closest to P.
thymanzae Domrow (1964c) among the species of Ptilonyssus parasitizing meli-

208 SOME MITE PARASITES OF AUSTRALIAN BIRDS

phagids, an essentially Australian group of passeriform birds. The former
species, however, has the podosomal shield wider posteriorly, bearing only
two pairs of setae, while this shield in the latter is wider anteriorly, and bears
four to five pairs of setae. Further, the dorsal setation of P. stomioperae is
decidedly heavier than that of P. thymanzae.

Types.—Ten females were collected from the nares of honeyeaters as
follows: holotype and five paratypes from two white-gaped honeyeaters,
Stomiopera unicolor (Gould), and four paratypes from a yellow honeyeater,
Meliphaga flava (Gould), all mist-netted amidst flowering Callistemon in the
bed of Magnificent Creek, Mitchell River, xi.1964, R.D. Holotype NIC;
paratypes RD, AF and RWS.

HATTENA PANOPLA, 0. Sp.
(Figs 62-64)

Female.—Idiosoma 547u long in slightly compressed specimen. Dorsal
shield reduced, marked by irregular, reticulate striae and lightly punctate.
System of paired pores present on shield, together with 25 pairs of setae, of
which four pairs are behind posterolateral incisions. Broad band of marginal
cuticle with two setae humerally, two posteriorly and four in line closely
following that of posterior margin of shield.

Sternal shield concave between SI, reduced and palely triangular behind
SII, textureless. SI and accompanying pores on shield proper ; SII and pores
borne on minute posterolateral promontories. SIII and pores on shieldlets ;
metasternal setae on shieldlets. Genital shield unexpanded, barely reaching
beyond posterior margins of coxae IV ; bearing two setae and rayed operculum.
Anal shield with anterior margin angularly convex and slightly denser than
remainder of shield, whose surface is slightly reticulate. Posterior margin
roundly convex, entirely occupied by narrow cribrum. Anus centrally placed,
with adanal setae near its anterior margin, and weaker than postanal seta.
Ventral cuticle with four pairs of setae preceding, and one pair following anal
shield. Also with five pairs of distinct pores borne on small plaques. Peri-
tremes extending forward to near level of posterior margin of coxae I, minutely
crenulate at edges. Peritremal shields extended posteriorly to fuse with
exopodal plates IV.

Coxae, some trochanters and gnathobase with rows of spinulose denti-
culations. Distal margins of leg and palpal segments similarly armed. All
leg setae slenderly tapering, formulation as follows: coxae 2.2.2.1; trochanters
6.5.5.5; femora 12.10.7/6.6 ; genua 12.11.9.10; tibiae 12.10.8.10; tarsi —16.16.16
(excluding two terminal filaments). This compares well with Till’s (1963)
formulae for Androlaelaps Berlese s.l. (including Haemolaelaps Berlese), except
that one seta less is present on femora I and II and genu and tibia I. (The
same formula occurs in H. erosa Domrow, where femur III is regularly 7.)
Ambulacra all well developed, but claws obsolescent.

Gnathosomal and hypostomal setae subequal except for smaller outer
posterior hypostomals. Deutosternum with six small denticles. Tritosternum
small, but with two ciliated laciniae. Labial cornicles also small, sharply
pointed. Palpal setal formula (trochanter to tibia) 2.5.6.14 (including two
dorsodistal tibial rods), agreeing with that given by Till for Androlaelaps.
Inner seta on palpal trochanter filamentous, and two on inner face of genu
clavate. Tarsal claw two-tined. Chelicerae stout, with two strongly sclerotized
digits, whose armature is not clear; corona absent.

Discussion.—As Baker and Yunker (1964) have recently reported blattisociine
mites both in flowers and the nares of hummingbirds in America, it is of interest
to note similar records from Australia. Members of this subfamily have been
seen on the pollen-strewn beaks and bare facial skin of several noisy friar-birds,
Philemon corniculatus (Latham) (Meliphagidae, Passeriformes), feeding in

ROBERT DOMROW 209

f
4

————_

Figs 62-64. Hattena panopla, n.sp. Female.—62, Idiosoma (ventral) ; 63, Idiosoma (dorsal) ;
64, Gnathosoma (ventral, with left palp dorsal).

Figs 65-66. Passerrhinoptes pomatostomi, n.sp. Male.—65, Gnathosoma and coxal apodemes I
(ventral) ; 66, Hysterosoma (ventral above, dorsal below).

flowering Eucalyptus at Logan Village, S.E.Q., but these specimens are not
now available for closer study. The opportunity has been taken, however,
to describe a specimen from the nares of another honeyeater.

Using Evans’ key (1957) (see also Chant, 1963), it is a little difficult to
decide if this specimen is a blattisociine or a platyseiine, as the inner palpal
trochanteral seta is filamentous, while the anterior hypostomals are not, etc.
The former choice has been made, as, while the specimen little resembles the
platyseiine genera figured by Evans and Hyatt (1960), it also shows the dorsal

F

210 SOME MITE PARASITES OF AUSTRALIAN BIRDS

shield laterally incised as in some blattisociine genera (Evans, 1958). However,
in Evans’ latter key, the new species will not run to either of the relevant genera,
Leioseius Berlese or Arctoseius Sig Thor. Nor does it appear to belong to Baker
and Yunker’s two genera, Rhinoseius and Tropicoseius.

In some respects, particularly the erosion of the sternal shield and the
shape of the anal shield, the new species appears congeneric with Hattena erosa
Domrow (1963), described from an unidentified bird from Sabah (British North
Borneo). In H. erosa, the dorsal shield (unincised) bears 21 pairs of setae and
the sternal shield one pair; in H. panopla, the corresponding figures are 25
and two.

Types.—Holotype female from the nares of a brown honeyeater, Gliciphila
indistincta (Vigors and Horsfield) (Meliphagidae, Passeriformes), Chelona, Sarina,
vli.1964, G.B. Holotype NIC.

Family SPELEOGNATHIDAE
SPELEOGNATHOPSIS BENOITI Fain

The following records (all adult specimens) are the first of this species from
Australia: one from a black-fronted dotterel, Charadrius melanops Vieillot,
Esk, 29.viii.1964, R.D. and J.S.W. ; one from a red-kneed dotterel, Hrythrogonys
cinctus Gould, Mitchell River, xi.1964, R.D.; and ten from a masked plover,
Lobibyx miles (Boddaert), Mitchell River, xi.1964, R.D. (all Charadriidae,
Charadriiformes).

All three series show the seta on coxa II obsolescent (+-), their coxal
formule being, in turn, 2.+.1.1, 2.4.1.1 and 2.+.1.0. The first specimen
agrees with the description of S. charadricola Fain (1964), except for the presence
of (i) seta on coxa IV ; and (ii) four setae (4B) rather than three (3B) on femur
IV. The second specimen recalls S. benoiti Fain (1955, 1956b, 1963b), possessing
five setae in the first postsensillary row and genital setae arranged 5.4, but
differs from that species in having (i) only six setae (5B.1N) on femur I rather
than 6B.1N ; and (ii) three setae on femur IV rather than four. The third
series agrees entirely with S. charadricola, but normally has four setae on femur
IV rather than three (however, two show 4.3 and one even 3.3). Granting a
considerable range of individual variation in this widespread and weakly
sclerotized group of internal parasites, only one species need be involved, and
I therefore consider S. charadricola a synonym of S. benoiti. This is further
confirmed by a study of individual variation in the dorsal setal pattern of a
series of 19 adults (one damaged specimen omitted) since collected in the nares
of a single black-fronted dotterel (Mitchell River, 17.iv.1965, R.D.). The
number of setae in the first postsensillary row was 4 three times, 5 seven times,
6 eight times and 7 once, the full formula for the lattermost specimen
(2.7.4.3.2.5.2) showing three rows uneven.

NEOBOYDAIA MEROPS (Fain)

Four adults collected as follows are the first records of this species in
Australia: rainbow-bird, Merops ornatus Latham (Meropidae, Coraciiformes),
Esk, 29.viii.1964 and 27.11.1965, R.D. and J.S.W.; and Innisfail-Palmerston
Highway, 1.1965, H.I.McD. and G.J.B. See Fain (1955, 1956c).

Family CHEYLETIDAE
NEOCHEYLETIELLA ARTAMI, ND. Sp.
(Figs 67, 69)
Female.—An oval-bodied mite with idiosoma 366 and 410y long in slightly
compressed specimens. Dorsal shield evenly rounded, but very weakly defined

anteriorly ; narrower and irregular, but clearly demarcated posteriorly ; virtually
textureless and bearing two fine setae on extreme anterolateral margins. Dorsal

ROBERT DOMROW 211

body cuticle with additional ten pairs of softly filamentous setae, all of which
are minutely bipectinate, and two pairs of smooth adanal setae.

Ventral body cuticle with four sternal, two preanal and four adanal setae,
all smooth. Valves of terminal genitoanal aperture each with four smooth
setae.

Legs. Coxal apodemes I and II elongate, all discrete posteriorly ;
apodemes III and IV also discrete, but smaller. Coxal setal formula 2.1.1.1,
all smooth. Remaining leg setation generally bipectinate (some pretarsals
especially so), though shorter setae tend to be smooth, particularly ventro-
distally. Trochanters 1.1.1.1, all ventral. All femora with one seta dorsally ;

Fig. 67. Neocheyletiella artami, n. sp. Kemale.—Dorsum.
Fig. 68. Passerrhinoptes pomatostomi, n.sp. Female.—Dorsum.

I and II also with ventral seta. Genua I and II each with two setae dorsally ;
former also with dorsal rod; genu III with seta ventrally ; genu IV asetose.
Tibia I with one dorsal and three ventral setae ; also with minute rod ; tibia IT
with two setae both dorsally and ventrally ; tibiae III and IV each with one
seta dorsally and two ventrally. Tarsus I with five setae and five rods, four
of latter borne on distinct dorsodistal saliency ; tarsus II with seven setae and
rod ; tarsi III and IV with seven and six setae respectively, arranged as figured.
Claws paired, strongly curved and somewhat swollen basally ; attached to
sclerotized basal apodemes. Pulvilli divided, with tips abruptly bent and
minutely bifid.

Gnathosoma stout, with two setae ventrally on gnathobase and four on
rostrum. Palpal trochanter obsolescent. Femur with one seta dorsally and
two ventrally. Genu and tibia with seta both dorsally and ventrally. Tarsus

212 SOME MITE PARASITES OF AUSTRALIAN BIRDS

ill-defined, with about two setae and rod. All gnathosomal setae smooth except
dorsal setae on palpal femur and genu. Chelicerae styliform, forming J with
articulatory sclerites. Hach arm of peritreme with five segments, sigmoid.

Discussion.—Of the species of Neocheyletiella Baker (1949) (less those forms
with two dorsal shields removed in 1964 by Volgin to Ornithocheyletia), N. artami
recalls N. smallwoodae Baker, but differs (as does Ornithocheyla megaphallos,
». infra), in having an additional pair of setae immediately behind the dorsal
shield. Dr. R. L. Smiley, U.S. Department of Agriculture, Washington, has
kindly compared my specimens with Dr. Baker’s, and confirmed this difference.

Of the 19th century species listed by Baker (1949), only Cheyletus macronycus
Mégnin (1878) seems near to NV. artami. I am grateful to Dr. M. André, Paris,
for the following information: “ Mégnin était Professeur a VEcole Vétérinaire
W@ Alfort, pres Paris. Sa collection est trés probablement restée dans cette Institution
mais, jusqwici, il n'a pas été possible de la retrouver. J’ignore si les échantillons
sont provisoirement égarés ou bien s ils ont disparu définitivement. Hn tout cas
les exemplaires dont vous venez de me faire parvenir les illustrations sont certaine-
ment trés voisins de macronychus et peut étre appartiennent-ils a cette méme
espéce.”’ Subsequent enquiries to Alfort have gone unanswered.

The genus Ornithocheyla was erected by Lawrence (1959), primarily on the
male intromittent organ, for O. megaphallos, a parasite of a waxbill (Ploceidae,
Passeriformes), for the loan of specimens of which I am most grateful to Dr.
Rk. F. Lawrence, Natal Museum, Pietermaritzburg. WN. artami is readily
separated by its unisetose trochanter III, and the presence of a seta dorsally
on tibia III and an additional seta on the ventral face of tarsus III. In
addition, coxal apodemes I and II are free distally, the dorsal setae on femur
and genu I are longer, and genuala I is internal to the adjacent seta.

Types.—Holotype female and paratype female from the dusky wood-
swallow, Artamus cyanopterus (Latham) (Artamidae, Passeriformes), Exeter,
Tas., 9.iv.1964, R.H.G. Holotype NIC; paratype RD.

Family TROMBICULIDAE
ODONTACARUS AUSTRALIENSIS (Hirst)

New host records for larvae of this species are: five from eyelids of one,
and two from another Australian black-shouldered kite, Hlanus notatus Gould
(Accipitridae, Falconiformes), Dalby, 14.vi.1963, I.D.F. and R.G.R. ; five from
a nankeen kestrel, Falco cenchroides Vigors and Horsfield (Falconidae, Falconi-
formes), same data ; nine from a grey-crowned babbler, Pomatostomus temporalis
(Vigors and Horsfield) (Timaliidae, Passeriformes), Condamine, same data ;
three from a black-faced cuckoo-shrike, Coracina novaehollandiae (Gmelin)
(Campephagidae, Passeriformes), Condamine, 6.vii.1963; 27 from a rufous
whistler, Pachycephala rufiventris (Latham) (Pachycephalidae, Passeriformes),
same data; and three from a noisy friar-bird, Philemon corniculatus (Latham)
(Meliphagidae, Passeriformes), Logan Village, 16.vii.1963, R.D., I.D.F. and
R.G.R. See Hirst (1925), Domrow (1956) and Brennan (1959). Also 1 larva
from a Lewin honeyeater, Meliphaga lewint Swainson (Meliphagidae, Passeri-
formes), Innisfail, 2.viii.1965, R.D. and J.S.W.

TROMBICULA SHIRAII Sasa, Kano and Ogata

This species, previously known only from two larvae from the eastern
golden plover, Pluvialis dominica (Miller) (Charadriidae, Charadriiformes) in
Japan, may now be recorded from Australia as follows: 15 larvae from the
bar-tailed godwit, Limosa lapponica (Linnaeus) (Scolopacidae, Charadriiformes),
Heron Is., Great Barrier Reef, 8.i.1964, J.B. Japan and Australia are included
in the range of both hosts. See Sasa et al. (1952) and Sasa and Jameson (1954).

ROBERT DOMROW 213

LEPTOTROMBIDIUM MYZANTHA (Womersley)

Eleven larvae from a green-winged pigeon, Chalcophaps chrysochlora
(Wagler) (Columbidae, Columbiformes), mist-netted at Mt. Jukes, Mackay,
vi.1964, R.D. and J.S.W. ; and five larvae from a pale-yellow robin, Hopsaltria
capito Gould (Muscicapidae, Passeriformes), Innisfail-Palmerston Highway,
11.ix.1964, H.I.McD., have been examined. See Gill et al. (1945), Womersley
(1952), and Womersley and Audy (1957). The last authors say ‘‘ the subgenus
is not indicated in the original description of the larva on p. 71’, but this is
not true of either copy in this Institute. They further wonder if the “ lousy
jack’ is the grey butcher-bird (Cracticus torquatus), but, in my experience, it
is Struthidea cinerea, the apostle-bird (i.e. the first of the birds listed by Gill
et al.), that goes commonly under this name in Queensland. The name stems
from their frequent infestation with mites (presumably tropical fowl mites,
which are popularly called ‘“ sparrow lice’’), and has since been reported to
me to be in use for two other Queensland birds, the grey-crowned babbler
(Pomatostomus temporalis) and the introduced Indian myna (Acridotheres tristis).
Of the several common names for Struthidea and Pomatostomus, ‘‘ apostle-bird ”
and ‘“‘ happy family ” are used interchangeably, while the former is also applied
to the white-winged chough (Corcorar melanorhamphus). All three are
gregarious (Cayley, 1963).

NEOSCHOENGASTIA POSEKANYI Wharton and Hardcastle
This widespread member of a bird-parasitic genus (Wharton and Hardcastle»
1946 ; Sasa and Jameson, 1954) has been once recorded from Australia (Derrick
and Womersley, 1954), and the following material has since been noted: one
larva (ACB635, formerly ACA1334), Wondecla, N.Q., 7.x.1943, R.V.S.; and a
very active colony of 12-15 reddish, newly-hatched larvae on top of burnt
tree-stump, about 2’6” from ground, Samford, 14.xi.1963, R.D. and I.D.F.

Dr. R. V. Southeott, Adelaide, has kindly made available his field notes
on the first specimen. It was taken running over a book on an army field
exercise in rainforest, and was recognized, at x 28, as a trombidiform larva.
It was red in colour and reminded one of Microsmaris Hirst (Erythraeidae,
see Southcott, 1961). Its eyes appeared 1 + 1 and between them were seen
two dots. These dots were undoubtedly the expanded sensillae, which also
appear quite dark in the Samford series, which was mounted directly from
spirit into Hoyer’s medium on the morning of capture. The eyes are rather
2 + 2, but they are borne on each side on a distinct ocular plate, and the
posterior two are quite dwarfed by the convex corneae of the anterior pair.

Family TURBINOPTIDAE
PASSERRHINOPTES POMATOSTOMI, Nn. sp.
(Figs 65-66, 68, 70)

Female.—Idiosoma 750-7704. long in three mounted (but only slightly
compressed) specimens, 836 in fourth flattened specimen. Ovate, with five
blunt extensions anterolaterally above gnathosoma and trochanters I and II
(formermost bearing merest suggestion of dorsal shield); slightly constricted
just in front of coxae III ; cuticle largely textureless, except for striations out-
lining evenly-arranged lobules middorsally, two of which each bear heavy seta
with sclerotized insertion. Posterolateral margins with three pairs of setae
(anterior pair much more evident than remainder) surrounding four pores.
Pair of supracoxal III setae present.

Vulva transverse, flanked by six setae (anterior pair issuing from contiguous
bases in one specimen) ; endogynium absent. Anus longitudinal ; adanal setae
in four pairs. Details of unpaired internal duct near anus not clear.

Legs with five free segments, coxae incorporated into body wall. Apodemes
I fused to form Y, with posterior arm twice as long as anteriors ; II sigmoid ;

214 SOME MITE PARASITES OF AUSTRALIAN BIRDS

III and IV contiguous and virtually complete. Coxal, trochanteral and femoral
setal formulae 1.0.1.0, 1.1.1.0 and 1.1.0.0, respectively. Genua I and II with
two basal setae and distal solenidion; genu III with solenidion; genu IV
unarmed. All tibiae with seta (point of insertion variable) and dorsodistal
solenidion (solenidia I-III three times as long as IV). Tarsi I and II much
compacted, heavily sclerotized; with dark, curved claw issuing dorsally,
together with two and one solenidia, respectively ; each with about six minute
setae ventrally. Tarsi III and IV normally formed, fully half as strong as
corresponding tibiae, each with four slender setae (three dorsal and one ventral).
It seems likely that the thickened (but pale and straight) structures set

Fig. 69. Neocheyletiella artamz, n. sp. Female.—Venter.
Fig. 70. Passerrhinoptes pomatostomi, n.sp. Female.—Venter.

terminally on the ventral aspect of tarsi III and IV are setae rather than claws.
They are quite unlike claws I and II, and much resemble the thickened ventral
seta on tarsus III. All ambulacra stalked and slightly expanded distally ;
IT and II issue beneath claw, III and IV above terminal “ spine ”’.

Gnathosoma as in male.

Male—As in female unless otherwise stated. Idiosoma 660yu long.
Hysterosoma with irregular X-shaped shield, whose posterior arms are the more
heavily sclerotized; notched midposteriorly. Remnants of genital discs
present. Penis support in reversed U; penis elongate, slenderly tapering
throughout its single coil. Anal discs small, diameter 13y. Gnathosoma
minute, with two ventral setae. Palpi displaced ventrally, very weak,
apparently with only one segment and at least one seta. Chelicerae set into
biconcave dorsal emargination; shaft and fixed digit stout, movable finger
very weak and slender.

ROBERT DOMROW 215

Nymphs.—At least two free nymphal stages occur. One (apparently
subadult) has idiosoma 750-760 long, and is similar to female except for lack
of vulva, ambulacra on all tarsi and fully-formed coxal apodemes III and IV.
An earlier stage (640) is similar to subadult, but lacks all trochanteral, tibial
IV and all but two genital setae.

Discussion.—Only one other species of Passerrhinoptes is known, P. andropadi
Fain, which has been recorded from bulbuls (Pycnonotidae) in Africa and babblers
(Timaliidae) in the Orient (see Fain, 1956d, 1960; Fain and Bafort, 1963a ;
Fain and Nadchatram, 1962). Dr. Fain has kindly lent me paratypes of his
species, as well as his Malayan specimen, and P. pomatostomi, while also a
parasite of a babbler, is clearly separable in both sexes by (i) its two heavy
dorsal setae ; and (ii) the proportions of the arms of the fused coxal apodemes I.
Further, in the male, the adanal discs are decidedly larger, and the details of
the opisthosomal shield differ.

Types.—Holotype female, allotype male, three paratype females and four
morphotype nymphs from the nares of a grey-crowned babbler, Pomatostomus
temporalis (Vigors and Horsfield) (Timaliidae, Passeriformes), Esk, 29.viii.1964,
R.D. and J.S.W. Holotype and allotype NIC; paratypes RD and AF.

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with a critical revision of the genera and subfamilies. Aust. J. Zool., 9: 367-610.

ROBERT DOMROW PHU G

STRANDTMANN, R. W., 1948.—The mesostigmatic nasal mites of birds I. Two new genera from
shore and marsh birds. J. Parasit., 34: 505-514.

, 1951.—The mesostigmatic nasal mites of birds Il. New and poorly known species
of Rhinonyssidae. J. Parasit., 37: 129-140.

, 1959.—New records for Rhinonyssus himantopus and notes on other species of the
genus. J. Kansas ent. Soc., 32: 133-136.

Tint, W. M., 1964.—A revision of the genus Pellonyssus Clark and Yunker (Acari: Mesostigmata).
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Votan, V. I., 1964.—[On the taxonomy of predatory mites of the family Cheyletidae VI. The
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Wane, D. C., 1963.—[Records of four species of Steatonyssus Kolenati, 1858 (Acarina,
Liponyssidae) from Fukien, China]. Acta ent. sin., 12: 54-60.

Waarton, G. W., and Harpcastie, A. B., 1946.—The genus Neoschéngastia (Acarinida :
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Witson, N., 1964.—New records and descriptions of Rhinonyssidae, mostly from New Guinea
(Acarina: Mesostigmata). Pacific Insects, 6: 357-388.

WomersteEy, H., 1941.—Notes on the Cheyletidae (Acarma, Trombidoidea) of Australia and
New Zealand, with descriptions of new species. Rec. S. Aust. Mus., 7: 51-64.

, 1952.—The scrub-typhus and scrub-itch mites (Trombiculidae, Acarina) of the
Asiatic-Pacific region. Rec. S. Aust. Mus., 10: 1-673.

, 1956a.—On some new Acarina-Mesostigmata from Australia, New Zealand and New
Guinea. J. Linn. Soc. (Zool.), 42: 505-599.

, 1956b6.—A new genus and two new species of Acarina from northern Australia. PRoc.
Linn. Soc. N.S.W., 80: 214-216.

Womerstry, H., and Aupy, J. R., 1957—Malaysian parasites XXVIII. The Trombiculidae
(Acarina) of the Asiatic-Pacific region: a revised and annotated list of the species in
Womersley (1952), with descriptions of larvae and nymphs. Stud. Inst. med. Res., Malaya,
28: 231-296.

Zumer, F., and Tint, W. M., 1955.—Nasal mites of birds hitherto known from the Ethiopian
Region, with keys and descriptions of nine new species (Acarina: Laelaptidae). J. ent.
Soc. S. Afr., 18: 60-92.

DEVELOPMENT OF THE EGGS AND EARLY LARVAE OF THE
AUSTRALIAN SMELT, RETROPINNA SEMONI (WEBER)

N. EH. Mitwarp
Department of Zoology, University of Queensland

(Plates vii—ix)

[Read 28th July, 1965]

Synopsis

Retropinna semont (Weber) is a small freshwater fish, occurring widely in eastern Australia.
Its eggs, embryonic development, and early larvae are described.

INTRODUCTION

The Australian smelt, Retropinna semoni (Weber), has a wide distribution
in eastern Australia, occurring throughout the great Murray-Darling River
system and also in coastal streams (Munro, 1957). The species grows to a length
of only 10 cm. and is unimportant either commercially or to the angler. How-
ever, In many areas it is extremely abundant and is preyed upon by several
of the larger fishes utilized by man (Butcher, 1945; personal records).

The following account of the eggs, embryonic development, and early
larvae of R. semont is based on studies carried out at the Inland Fisheries
Research Station, Narrandera, New South Wales. The studies were initiated
following evidence of natural breeding in one of the experimental ponds at
this Station.

METHODS

Following the discovery of larval stages of R. semoni in an experimental
pond on September 22, 1961, collections of adults in reproductive condition
were made from the pond later the same day and also on September 23.

A first attempt to obtain fertilized eggs was made by adding milt to ova,
apparently ripe but not strippable, removed from a female by dissection. No
fertilization was achieved.

A second attempt proved successful. On this occasion the ova were readily
stripped from a female, measuring 75 mm. total length, by applying slight
pressure on the abdomen. Milt was expressed from a male, measuring 73 mm.,
in a similar manner, and placed onto the eggs held in a Petri dish. Water was
immediately added to cover the eggs and the whole mixed by gentle shaking.
Approximately 20 minutes later the eggs were washed with several changes
of water and then transferred to a shallow enamel tray. This was immersed
in a deeper tray, through which water was circulated. On the eighth day,
when hatching appeared imminent, the water circulation was stopped to avoid
loss of larvae in the overflow.

The temperature of the water flowing over the eggs was not controlled,
but for the greater part of the hatching period fluctuated within the range
15.5 to 18.0°C. On two occasions, for periods of three to four hours, the
temperature fell to about 13°C.

Measurements were made with an ocular micrometer. The photographs
were taken using transmitted light.

PROCEEDINGS OF THE LINNEAN Society oF New SourH Watss, Vol. 90, Part 2

N. E. MILWARD 219

Efforts were made to rear the larvae in glass and plastic containers in the
laboratory. However, although various kinds of finely ground food were supplied,
no feeding was observed and the longest that any larvae survived was 8 days.

THE HGGS

The eggs when stripped are spherical, with an average diameter of 0.80 mm.
Following fertilization they swell to an average diameter of 0.95 mm. They
sink in fresh water and are strongly adhesive, becoming firmly stuck to the
bottom of the hatching tray. The yolk is pale amber in colour. Initially it
completely fills the egg, but quite a large perivitelline space is formed during
the swelling of the thin capsule. One to several large and many small oil
globules are present within the yolk.

EMBRYONIC DEVELOPMENT

The formation of the blastodisc commences almost immediately upon
fertilization. Cytoplasm, which has hitherto invested the yolk in an invisible
layer, slowly accumulates at the animal pole of the egg (Plate vii, fig. 1). When
concentration of the cytoplasm is complete the formed blastodisc is of a lenticular
shape, the surface of the yolk immediately opposite having flattened (Plate vii,
fig. 2).

The first three or four cleavages occur within 3 hours following fertilization.
These early cleavages, at least up to and including the fourth, take place regularly
throughout the blastoderm, the blastomeres formed being fairly uniform in size
(Plate vii, fig. 3). In a small number of eggs the cleavage rate was slower and
in a few the cleavages were irregular, so that unequal blastomeres were formed.
However, these eggs having retarded and/or obviously irregular development
all died during the first two days. Subsequent mortalities until after hatching
were few and development progressed more or less uniformly in all eggs.

Sixteen hours after fertilization the blastodermal cap has been formed.
It is of a similar lenticular shape to that of the blastodisc but more opaque
(Plate vii, fig. 4).

About 22 hours after fertilization the germ ring has reached approximately
an equatorial position about the yolk. It is slightly thickened and appears to
constrict the yolk sphere as it advances over it (Plate vii, fig. 5). The thickening
of the germ ring is more pronounced on one side than the other, the thickened
portion marking the posterior pole at which the embryonic shield develops.

The embryo is clearly evident and shows marked development at 41 hours.
It extends approximately two-thirds the way around the yolk, and is noticeably
thickened in the cephalic region. The thin blastodermal layer now almost
fully encloses the yolk, except at the blastopore, situated just posteriorly to
the tail end of the embryo, through which there is a slight bulging of the yolk.

About 47 hours after fertilization, the optic vesicles are easily discernible
(Plate vii, fig. 6). The embryo is much thickened along its whole length,
especially in the cephalic region, and dorsally protrudes markedly into the
perivitelline space. Kupffer’s vesicle, a small transparent sphere lying ventrally
near the posterior end of the embryo, has appeared.

Considerable differentiation has occurred in the embryo by 66 hours after
fertilization (Plate viii, figs 7, 8). The eyes are now very clear and the
pupils have developed. The head is further enlarged and the lobes of the brain
are apparent. Auditory capsules are present and more than 30 mesodermal
somites are distinguishable. The embryo almost fully encircles the yolk, which
is slightly constricted around the line of contact. To this point no pigmentation
has been developed.

The embryo more than fully encircles the yolk at 95 hours, the tail slightly
overlapping the head (Plate viii, fig. 9). The heart, which was not distinguished
at 66 hours, is now easily seen. It is situated just under and posterior to the

220 EGGS AND BARLY LARVAE OF THE AUSTRALIAN SMELT

eyes, and pulsates quite regularly. The first pigmentation has now appeared
as series of melanophores on the sides of the body, along portion of the midline
and to a lesser extent along the ventral edge. Otoliths have developed within
the auditory capsules. The first movements, slight twitchings, of the embryo
were observed at this stage.

At about 113 hours the embryo extends approximately one and a quarter
times around the yolk, and the eyes are becoming quite heavily pigmented.
Movements of the embryo are now more frequent, the posterior portion of the
body being detached from the yolk and moving freely.

From this stage until hatching there is a continued increase in the length
of the embryo, so that at 137 hours it encircles the yolk approximately one
and a half times (Plate viii, fig. 10) and at 165 hours about two times (Plate viii,
figs 11 and 12). There is a marked lateral expansion of the head, causing it
to be roughly triangular in shape by 165 hours (Plate viii, fig. 12). The
melanophores along the body increase in number, becoming more uniform and
pronounced, and others develop at about 130 hours over the yolk sac. The
dorsal and ventral fin-folds are clearly evident at about 130 hours.

Towards hatching the embryo moves almost incessantly, the tail twisting
and switching from side to side and at intervals the whole embryo revolves
completely within the egg capsule. The pectoral fins, now evident as transparent
fan-like structures slightly posterior to the auditory capsules, become active and
beat rapidly for increasing periods of time from about 210 hours onwards.

Hatching commenced at 216 hours after fertilization and all the larvae
had emerged from the egg capsules by 225 hours.

THE LARVAE

The newly hatched larvae (Plate ix, figs 13 and 14) are extremely elongate
in form, the average total length being 4.61 mm. The head is inflected down-
wards and is anteriorly rounded, so that the eyes appear ventrally placed. The
mouth is present as a small opening situated below the eyes, but is probably
non-functional for the first day or so after hatching. The auditory capsules
are comparatively large and protrude prominently from the sides of the head.
A small mass of yolk contained in an elongate, ovoid sac is still present at
hatching. A single oil globule is present within the anterior end of the yolk sac.
The hind portion of the alimentary canal is clearly evident as a long, straight
tube and the anal opening is situated two-thirds the way along the body. Both
the dorsal and ventral fin-folds are fairly uniform in height throughout and are
continuous with the caudal fin-fold, which is slightly more expanded and lobate.

On hatching, the larvae congregated near the surface and sides of the hatching
tray. For most of the time they remained fairly passive, normally orientated
horizontally with dorsal side uppermost, but occasionally, particularly if
disturbed, swimming actively with apparently well directed movements.

One day after hatching, the yolk sac and contained oil globule are both
much reduced in size (Plate ix, fig. 15). The head has now pivoted forwards
and upwards, so that the mouth is situated more anteriorly and the eyes lie
slightly more dorsally than in the newly hatched larva. The straightening of
the head contributes to a relatively large increase in length, so that the average
total length attains 5.25 mm.

After two days the yolk has been almost completely absorbed and the oil
globule has disappeared (Plate ix, fig. 16). Up to this time there is evidence
of continued differentiation in the larvae, particularly in the head region. The
jaws are now well developed, the auditory capsules further enlarged, and the
pectoral fins larger, stronger and much easier to see. Compared with the
increase in length over the first day, that during the second is small, the
average total length of the two-day larvae being 5.29 mm.

N. E. MILWARD 221

Following the complete utilization of the yolk on the third day, development
of the larvae almost ceased and there was evidence of emaciation. This was
undoubtedly due to unsuitability of food provided. A slight increase in length
occurred to the fifth day, the average total length of larvae then surviving
being 5.51 mm. No further growth was recorded and the single eight-day
larva measured only 5.30 mm., the shrinkage possibly being due to a natural
consolidating of tissues, but more probably to the larva having to resort to its
own body substance for nourishment.

Acknowledgements

The work was carried out during employment by the Fisheries Branch,
Chief Secretary’s Department, New South Wales. The author wishes to thank
members of the staff of that Department for their co-operation and, in particular,
Mr. J. S. Lake, Officer in Charge of the Narrandera Research Station.

References
ButcHer, A. D., 1945.—The food of indigenous and non-indigenous freshwater fish in Victoria,
with special reference to Trout. Vict. Fish. and Game Dept., Fish Pam. No. 2, (Govt.
Printer, Melbourne).
Munro, I. 8. R., 1957.—Handbook of Australian fishes, No. 7, p. 29. (Published in Fisheries
Newsletter, Dept. Primary Industry, Canberra. Jan. 1957).

EXPLANATION OF PLATES VII-IX

Plate vil

Fig. 1. Egg 17 min. after fertilization. Incomplete blastodisc.—Fig. 2. Egg 1 hr. after
fertilization. Complete blastodisc.—Fig. 3. Egg 3 hr. after fertilization. Sixteen cells.—
Fig. 4. Egg 16hr. after fertilization. Blastodermal cap.—Fig. 5. Egg 22 hr. after fertilization.
Germ ring at equatorial position.—Fig. 6. Egg 47 hr. after fertilization. Blastopore closed,
embryo protruding into perivitelline space, and optic vesicles evident.

Plate vii

Figs 7, 8. Egg 66 hr. after fertilization. Brain lobes, pupils of eyes, auditory capsules, and
somites distinct.—Fig. 9. Egg 95 hr. after fertilization. Embryo completely encircling yolk,
first pigmentation on body.—Fig. 10. Egg 137 hr. after fertilization. Embryo encircling yolk
approx 14 times, eyes heavily pigmented, melanophores over yolk sac.—Figs 11,12. Egg 165 hr.
after fertilization. Very advanced embryo, encircling yolk approx. 2 times.

Plate ix

Figs 13, 14. Newly hatched larva. Average length 4-61 mm.—Fig. 15. One day old larva.
Average length 5:25 mm.—Fig. 16. Two days old larva. Average length 5:29 mm.

THE FIRST ZOEA OF THE SOLDIER CRAB MICTYRIS LONGICARPUS
(GRAPSOIDEA: MICTYRIDAE)

ANN M. CAMERON
Zoology Department, University of Queensland

(Communicated by Dr. HE. J. Reye)

[Read 28th July 1965]

Synopsis
The first zoea of Mictyris longicarpus is described and figured.

M. longicarpus Latreille, 1806, is one of the most characteristic elements
of the estuarine sand flat fauna on the eastern Australian coast. No information
on its larval development is available.

METHODS

Ovigerous female M. longicarpus were collected from Moreton Bay, southern
Queensland, and kept in aerated filtered seawater containing 200,000 i.u. of
penicillin per litre. All experiments were carried out at 25°C but the photo-
period was not controlled. Usually seawater was changed daily, occasionally
after two days.

When hatching was imminent each female was confined to a separate
container, at first an aquarium, later a finger bowl. By this method, many
‘* prezoeae ’? were obtained but all remained sluggishly on the bottom of the
container. The eggs of one female hatched successfully. She was observed to
flex and relax her abdomen rhythmically, sweeping the pleopods outwards and
inwards with the same rhythm. Thus the hatching larvae were swept out and
away from the rest of the egg mass. All the eggs this female was carrying
hatched within a period of ten minutes, and all were first zoeae. They swam
actively and continuously and aggregated in that part of the finger bowl where
maximum illumination prevailed.

Because of the limited success obtained with the method described above,
egg masses were removed from females prior to hatching. Groups of from
100 to 500 eggs were dissected under the microscope and were placed in a
compartmented perspex container with a water depth of 2.5 cm. The
compartments were of two sizes, having surface areas of approximately 20
and 40 cm?. The maximum number of eggs per square cm. of water surface
was 25. A shaking machine vibrating through a very small amplitude at 120
times per minute housed the container, which was covered with a lid of the same
transparent perspex. Successful hatching was obtained repeatedly under these
conditions and eggs were maintained for up to eight days prior to the hatching
of healthy zoeae.

After hatching, the zoeae were divided into groups of from 10 to 200 per
compartment. Any sluggish or dead larvae were removed when the seawater
was changed. Artemia nauplii were added to the compartments but were not
eaten by the zoeae. Melarapha scabra trochophores were similarly rejected.
Zoeae survived for twelve days, eventually dying of starvation.

PROCEEDINGS OF THE LINNEAN Society or NEw SoutH WaAtzgEs, Vol. 90, Part 2

ANN M. CAMBRON 223

DESCRIPTION OF THE FIRST ZOEA

The first zoea of M. longicarpus is illustrated in Figures 1-9. The
cephalothorax bears a rostral spine only. The dorsal and the lateral spines
are absent. Extensive flanges are present on the lateral and postero-lateral
margins of the carapace. There are five abdominal segments and the telson,
all of which have chromatophores. There is a secondary chromatophore on
the first maxilliped but none on the second.

5

Figs 1-9. 1, Lateral view of M. longicarpus first zoea. 2, Posterior view. 3, Antennule.
4, Antenna. 5, Maxillule. 6, Maxilla. 7, First maxilliped. 8, Endopodite of second maxilliped.
The four setae of the exopodite have been truncated. 9, Telson.

The scale lmes for Figs 3-9 are 0-1 mm.

The antennule (Figure 3) has two terminal aesthetes and a terminal seta
about one-fifth the length of the aesthetes. A subterminal spine is also present.
The antenna (Figure 4) has a tapered protopodite which bears two rows of short
spines. The exopodite bears a long spine somewhat swollen proximally. This
antenna is of the B, type of Aikawa (1933, p. 126). The mandible was not
examined. The endopodite of the maxillule (Figure 5) has four terminal setae,

224 FIRST ZOEA OF THE SOLDIER CRAB MICTYRIS LONGICARPUS

one subterminal seta, and one seta projecting from the basal segment. There
are five spines on the basal endite and the coxal endite has four spines.

The unsegmented endopodite of the maxilla (Figure 6) has two terminal
setae and two subterminal setae. There are eight spines on the basal endite
and six on the coxal endite. From the distal margin of the scaphthognathite
four soft bristles project. The first maxilliped (Figure 7) has four swimming
setae on the exopodite. The setation of the five-segmented endopodite is
5-2-1-2-2 (beginning with the distal segment). The second maxilliped
(Figure 8) likewise has four swimming setae on the exopodite, while the setation
of the three-segmented endopodite is 6-1-1. The telson (Figure 9) is of the
B type according to Aikawa’s classification (1929, p. 23). There is a dorsal
spine on each prong of the fork. Inside the fork, there are three pairs of spines
on a median lobe, the outer two pairs being about twice as long as the inner
pair.

DISCUSSION

Starvation inhibits moulting, and the failure to rear the larvae through
the first moult was due evidently to the unsuitability as food of the nauplii
and trochophores provided.

Features of greatest importance in the classification of brachyuran first
zoeae are the nature of the second antenna and telson (Lebour, 1928 ; Aikawa,
1937). Other significant characters are the presence or absence of carapacial
spines, and the setation of the maxillae and maxillipeds. Mictyris longicarpus
first zoea resembles the first zoeae of Macrophthalmus in the nature of the second
antenna and telson (Aikawa, 1937, p. 152, 153). Table 1 compares the setation
of the mouthparts in Mictyris longicarpus and Macrophthalmus (Aikawa, 1929,
p- 28, 31).

TABLE |
Setation of Mouthparts

Species Mx. I Mx. IT Mxpd. [ Mxpd. II
Mictyris longicarpus » 5-1 2—2(4) 5-2—1-2-2 6-1-1
Macrophthalmus spp. . - sg 5-1 2—2(4) 6—3—-1—2-2 6-1-1

Further, Macrophthalmus first zoeae lack lateral carapacial spines as does
the first zoea of Muictyris longicarpus. Despite the very close resemblance
between Mictyris longicarpus first zoea and those of Macrophthalmus, the former
can be distinguished readily by its lack of a dorsal carapacial spine.

References

Aikawa, H., 1929.—On larval forms of some Brachyura. Rec. oceanogr. Wks Jap., 2 (1):
17-55.
, 1933.—On larval forms of some Brachyura, Paper II: A note on indeterminable
zoeas. Rec. oceanogr. Wks Jap., 5 (2): 124-254.
, 1937.—Further notes on brachyuran larvae. Rec. oceanogr. Wks Jap., 9 (1): 87-162.
Lesovr, M. V., 1928.—The larval stages of the Plymouth Brachyura. Proc. zool. Soc. Lond.,
1928: 473-560.

PLANT PARASITIC NEMATODES IN FRUIT TREE NURSERIES
OF NEW SOUTH WALES

HK. J. ANDERSON*
Faculty of Agriculture, University of Sydney, Sydney, N.S.W.

(Communicated by Dr. C. D. Blake)
[Read 28th July, 1965]

Synopsis

Soil samples taken from under 11 kinds of fruit trees in 20 nurseries and from a few bearing
orchards in New South Wales, were examined for plant parasitic nematodes. Spiral nematodes
(Helicotylenchus spp.) were present in all areas and associated with all root stocks examined.
Root lesion nematodes (Pratylenchus spp.) were also widely distributed. Stubby root nematodes
(Lrichodorus spp.) were more prevalent in Gosford than in other districts and were associated
with Citrus spp. The citrus nematode (Tylenchulus semipenetrans Cobb) was found in one
quarter of the samples examined from citrus at Gosford and Sydney, in one sample of fallow
soil and in a young orange planting in the Murrumbidgee Irrigation Areas (M.I.A.) established
from plants imported from Gosford. Stylet nematodes (Lylenchorhynchus spp.) were found
only in the M.I.A., and dagger nematodes (Xiphinema spp.) were found infrequently in the
Gosford, Sydney and M.I.A. districts. The citrus, root lesion and stubby root nematodes were
present sufficiently often to pose a threat to new orchards planted with stocks from infested
nurseries.

INTRODUCTION

A few samples of soil taken from a nursery near Gosford, New South Wales
(N.S.W.), by the author in 1957 yielded many nematodes known to be parasitic
on the roots of plants. Such nematodes were found also in samples of soil
taken from areas then growing either native bush or grasses and from cultivated
areas devoted to forage, fruit and vegetable crops. These collections were
studied by Drs. M. W. Allen and RB. C. Colbran at the University of California,
U.S.A., who confirmed the author’s identifications. Included in these
collections were 15 species in nine genera known, or thought to be plant
parasites. Hither larvae or insignificant numbers of adults of four other genera
known to contain plant parasites were found. Table 1 lists the nematodes
and the plants with which they were associated.

The presence of the genera Meloidogyne, Pratylenchus, Radopholus and
Tylenchulus species, which live within roots, and of AHelicotylenchus and
Rotylenchus spp., which are not always removed when roots are washed,
raised the question whether these nematodes are introduced to new orchard
areas with nursery stock.

METHODS

Samples of soil and small roots were taken to a depth of six to eight inches
after the top one to two inches of soil had been removed. Most samples, each
of about 500 ml., were treated separately, but a few from two or more sites
along a nursery row were bulked before treatment. Nurseries near Gosford,
Sydney, Bathurst, Orange, Griffith, Leeton and Yanco were sampled. In
addition, five young orchards and three “‘ virgin”’ or long fallow areas were
sampled. About 300 samples were collected between October 1 and December
11, 1964.

* Plant pathologist, on leave from the Pineapple Research Institute of Hawaii, Honolulu,
Hawaii, U.S.A.

PROCEEDINGS OF THE LiINNEAN Socrmty ar New SourH Wates. Vol. 90, Part 2

226 PLANT PARASITIC NEMATODES IN FRUIT TREE NURSERIES

TABLE |
Plant parasitic nematodes in soil samples collected in New South Wales in 1957

Nematode Plants with which associated
Criconema sp. orange
Criconemoides mutabile Taylor maize on old grassland
Criconemoides xenoplax Raski peach, cowpea
Criconemoides sp. Casuarina sp.
Helicotylenchus multicinctus Cobb Kikuyu grass, peach, Hucalyptus pilularis
Helicotylenchus nannus (Steiner) banana, Casuarina sp., Cynodon sp., Datura sp., grape,

Andrassy nectarine, orange, pineapple, strawberry

Hemicycliophora sp. banana, maize, Hucalyptus maculata, E. pilularis
Heterodera sp. Casuarina sp.
Meloidogyne javanica (Treub) Chitwood Kikuyu grass
Meloidogyne sp. banana, strawberry
Paratylenchus sp. asparagus, maize, orange, peach, tobacco

Pratylenchus minyus Sher and Allen tomato
Pratylenchus thornet Sher and Allen cauliflower

Pratylenchus sp. lucerne

Radopholus similis (Cobb) Thorne banana, Kikuyu grass, pineapple

Rotylenchus brachyurus Steimer grape

Rotylenchus robustus (de Man) Filipjev Hucalyptus sp. and mixed grasses

Rotylenchus sp. mixed grasses

Trichodorus minor Colbran asparagus, banana, Bermuda grass, Casuarina palulosa,
Datura sp., grape, nectarme, orange, pineapple,
strawberry

Trichodorus porosus Allen Casuarina sp., Eucalyptus sp., nectarine, peach

Trichodorus sp. peach, pineapple, Hucalyptus sp.

Tylenchulus semipenetrans Cobb orange (on sour lemon stock)

Tylenchorhynchus brevidens Allen lucerne

Tylenchorhynchus latus Allen cauliflower

Tylenchorhynchus sp. strawberry

Nematodes were extracted from 250 ml. of soil by either an elutriator
(Seinhorst, 1956) or a flotation method (Jenkins, 1964). These methods yielded
comparable results and were used interchangeably. The nematodes were killed
by heating in a water bath to 125°F and preserved in 4% formalin. For all
observations reported the nematodes were recovered from the soil samples
within 10 days of being collected.

RESULTS

Tables 2 and 3 show marked differences in the distribution of nematodes
between nurseries. Some nurseries grew citrus, others predominantly stone
or pome fruits, while others grew all of these. Thus, it is difficult to separate
the effects of host and location on nematode infestation. Some species appear
to be widely distributed and others more restricted both as to district and host.
The species, in so far as they could be determined, the plants with which they
were associated, and the districts in which they were found are summarized.

Criconemoides teres Raski was found in a sample of “ virgin” soil near
Gosford and in apple, peach and orange (Poncirus trifoliata stocks) nursery
rows near Sydney. COviconemoides sp. larvae also were found in the Gosford
and M.I.A. districts associated with P. trifoliata stocks and in a plum seedling
planting near Sydney.

Helicotylenchus nannus (Golden) Perry was associated with P. trifoliata
stocks, grapefruit, orange, pear and plum plantings and in “ virgin ”’ soil in
the Gosford district, and with all these plants as well as apple, lemon, walnut
and in fallow soil near Sydney. In the Bathurst-Orange district it was found
in plantings of apple and peach seedlings. H. multicinctus (Cobb) Golden was
found near Gosford in a planting of P. trifoliata seedlings and, near Sydney,

E. J. ANDERSON

TABLE 2

Distribution of principal genera of plant parasitic nematodes in fruit tree nurseries
of New South Wales

Number of Samples Infested

A)
Nursery plant S =
and S s
District § >
= 3
ss
= S)
S &
Apple
Sydney 14
Bathurst—Orange — 8
M.1.A.* — 8
Apricot
Sydney — 6
M.I.A. — 5
Cherry
Sydney = 3
Bathurst—Orange — 1
Citrus
Gosford 1 29
Sydney 4 45
M.I.A. 2 2
Grape
M.I.A. — 2
Mulberry
Sydney — 2
Nectarine
Sydney — 2
M.I.A. — 2
Peach
Sydney 4 14
Bathurst—Orange — 6
M.LA. — 2
Pear
Gosford — 2
Sydney — 6
Bathurst—Orange — 2
M.I.A. — —
Plum
Gosford = 2
Sydney 1 10
Walnut
Sydney — 2
M.I.A. — 1

* Murrumbidgee

Irrigation Areas

Paratylenchus

cs

Pratylenchus

bo Ou >

CE

Sr le ee

Qo —

bo

Trichodorus

Tylenchorhynchus

Tylenchulus

Xiphinema

Number
of samples
examined

—

bo

rl |e

bo

15
14

42
49

bo & -1 bo

228 PLANT PARASITIC NEMATODES IN FRUIT TREE NURSERIES

TABLE 3

Principal genera of plant parasitic nematodes in soils from bearing orchards and
oe -

>

virgin” or fallow areas
a)
SS
2 3
~ = ey
3 3 S 3 % S. a Number
Source $ $ 8 = > s = 5 of samples
S > 5 = S g S 3 examined
S) cd > 3S = o~
= iS => S S 3 3 §
S 8 S es 5 Sa
eS S aA Spi eb nes
S q q Q iS & Ss a
Apple
Sydney = 6 — — — — 1 6
Bathurst—Orange — 1 — — — — 3
Orange
Gosford — 2 — 2 2 — 2 — 2
M.1.A.* 1 2 4 10 10 —- 2 7 17
“Virgin” soil or fallow
Gosford 1 4 — 4
Sydney — 7 — 1 — _- 1 — 7
M.L.A. — 1 _— 2 — — — 1 2

* Murrumbidgee Irrigation Areas

in plantings of peach and plum. In addition, Helicotylenchus sp. larvae were
found in a planting of walnut at Sydney, plantings of apple and pear near
Bathurst, in lemon, nectarine and peach plantings and in fallow soil in the
M.I.A. Helicotylenchus spp., therefore, were found more frequently and
associated with a larger number of plants in the Gosford and Sydney districts
than in the M.I.A.

Paratylenchus nanus Cobb was found only in the M.I.A. and was associated
with apple, apricot, peach and P. trifoliata stocks. A few specimens of
Paratylenchus sp. were found near Gosford associated with citrange stocks,
near Sydney with lemon and cherry, and in the M.I.A. with apricot.

The genus Pratylenchus was represented by many species and was widely
distributed. P. coffeae (Zimmerman) Goodey was associated with apple and
peach near Bathurst, with apricot in the M.I.A., and with apple, cherry and
mulberry near Sydney. P. minyus Sher and Allen was associated with plum
at Gosford, apple, lemon, peach and plum near Sydney, with apple at Bathurst,
and with apple, apricot, grape, lemon, nectarine, peach, pear, P. trifoliata
stocks and walnut in the M.I.A. P. penetrans (Cobb) Chitwood and Oteifa was
seen only in a collection from apple in the M.I.A. P. zeae Graham was associated
with apple, cherry, lemon and P. trifoliata stocks and in fallow land near Sydney,
and with peach and pear in the Bathurst-Orange district. Specimens of
Pratylenchus which could not be further identified were found associated with
lemon, P. trifoliata and orange stocks at Gosford, with apple, apricot and cherry
at Sydney, and with apple, apricot and peach in the M.I.A.

Trichodorus minor Colbran was found associated with apple and lemon
near Gosford and with apricot, peach, pear and P. trifoliata in the M.I.A. T.
porosus Allen was associated at Gosford with citrange, lemon, orange, P. trifoliata
and pear stocks, near Sydney with lemon and orange, and in the M.I.A. with
apricot, grape, orange and P. trifoliata. T. teres Hooper was associated with
citrus at Gosford. Specimens of Trichodorus, the species of which could not
be determined, were found at Gosford in grapefruit and plum plantings, at
Sydney in fallow soil, and in the M.I.A. in plantings of lemon and orange.

E. J. ANDERSON 229

Specimens of Tylenchorhynchus which appeared to be T. brevidens Allen were
associated with apple, apricot, grape, peach and walnut in the M.I.A. and in
the same area Tylenchorhynchus sp. was found in a planting of P. trifoliata stocks.

Larvae of Tylenchulus semipenetrans Cobb were found in large numbers in
soil from nursery rows of citrange, lemon and orange seedlings and from an
orange grove near Gosford, in soil from rows of lemon, orange and P. trifoliata
as well as a fallow area near Sydney, and from three young orange plantings
on stocks of rough lemon, orange and P. trifoliata near Yanco in the M.I.A.
The young trees in the latter plantings were said to have been grown from seed
at Gosford.

Xiphinema americanum Cobb was found in small numbers in soil from
rows of apple and peach stocks near Sydney and from grape, P. trifoliata and
walnut in the M.I.A. Xiphinema spp. were found also in soil from apple and
P. trifoliata rows near Sydney, and from apple, apricot and grape in the M.I.A.

A few specimens of larvae or lone adults of Belonolaimus (Sydney: plum),
Hemicycliophora (Gosford: “virgin” soil), Hoplolaimus (Sydney: pear),
Longidorus (Sydney: apple, peach and pear), Meloidogyne (Sydney: peach
and fallow), Rotylenchus (Gosford: P. trifoliata; Sydney: peach and fallow)
and Tylenchus (Sydney: peach; M.I.A.: apple and peach) were found.
These are considered incidental.

DISCUSSION

The failure to find a nematode species in a few samples does not necessarily
indicate that it is not present in the area or that the plant in question is not
a host. However, its absence or rare occurrence in one area or crop as opposed
to another can be taken to indicate that it probably does not exist there in
serious numbers because it has not been introduced, has been introduced only
recently, or the plants in the area are not suitable hosts for its rapid reproduction.
Proper interpretation of the observations presented here requires knowledge
of the host range and the pathogenicity of the nematodes on the various crop
plants. Such information is at present lacking. Many of the nurseries are
infested with weeds which may be better hosts than the crop plant. Thus,
numbers of nematodes per soil sample may not be meaningful unless the
nematode in question is known to be pathogenic on the crop plant. Finally,
the determination of species sometimes is questionable because authentic slides
for comparison were unavailable.

Relatively few nematodes were found in samples from the Bathurst-
Orange district. This district had been unusually wet for several weeks before
the samples were collected. Some sites had been flooded and all were nearly
saturated with water when sampled. Thus, the results from this district may
not be valid.

The genus Helicotylenchus was ubiquitous, occurring in more than half of
the samples and in 12 of the 13 samples from “ virgin” or long fallow soils
examined. Golden (1956) showed that R. buxsophilus was pathogenic on box-
wood, but the significance of this genus in this study is unknown. On pineapple
in Hawaii special nematodes appear to cause limited injury, but they are so
widespread that control measures in the nursery alone would not be justified.

The genus Pratylenchus (root lesion nematodes) is also widespread. One
or more species were found in all four districts and associated with all crops
sampled. P. minyus Sher and Allen was found also in fallowed soil in the
M.I.A. Colbran (1953) showed that P. coffeae causes serious injury to apples
in Queensland, while Seinhorst and Sauer (1956) found P. scribnert and P.
vulnus attacking grapes in Victoria. Because this genus is pathogenic and
widespread, it poses a threat to new orchard plantings and warrants control
measures being considered.

230 PLANT PARASITIC NEMATODES IN FRUIT TREE NURSERIES

Species of the genus Trichodorus (stubby root nematodes) which were
found in three-quarters of the samples from citrus nurseries in the Gosford
district, in fewer nurseries in the Sydney and M.I.A. districts, but again
associated with citrus. Stubby roots symptoms in citrus were seen frequently
in this study. The genus contains known parasitic species and an attempt
should be made to prevent its spread to new citrus planting.

The citrus nematode (Tylenchulus semipenetrans Cobb) has been recognized
as a pathogen in citrus and grapes in Australia for some time (Seinhorst and
Sauer, 1956; Sauer, 1962). In this study it was found in nurseries in the
Gosford and Sydney districts, in an orange orchard near Gosford, and in the
M.I.A. The trees with which it was associated in the M.I.A. were raised at
Gosford. It was not associated with other crops, but was recovered from. a
fallow soil at the border of a nursery near Sydney. It is a serious pest in citrus
and poses a threat to the yield from any planting made with infected nursery
stock. It can be controlled in orchards but at a very much greater expense
per tree than in the nursery. Every effort should be made to eliminate this
nematode from nursery stock.

Xiphinema species (dagger nematodes) found infrequently in the Sydney
and M.I.A. districts are of interest because the genus contains vectors of plant
viruses. These nematodes and Longidorus spp. merit study, but do not appear
to be a threat at present.

Other nematodes, Belonolaimus, Criconemoides, Paratylenchus, Tylencho-
rhynchus and Tylenchus were encountered infrequently and are not likely to
be carried in nursery stocks.

A successful control measure for one of the more serious nematode pests
in nurseries, such as the citrus nematode, is likely to control other nematodes
at the same time. Studies of chemical treatment of the soil, treatment of
lifted root stocks, with either a nematicide or heat or both, before sale, should
be pursued vigorously.

Acknowledgements

I thank Mr. R. McLeod and the Fruit Officers of the Department of
Agriculture for help in collecting samples, Miss Lynette Snelson for help in
recovering the nematodes from the samples and preparing slides, Assoc.
Professor N. H. White and Dr. C. D. Blake for criticism of the manuscript
and, particularly, Mr. and Mrs. Edwin Street whose gift to the University of
Sydney financed this study.

References

CoLBRAN, R. C., 1953.—Problems in tree replacement. I. The root lesion nematode, Pratylenchus
coffeae Zimmerman, as a factor in the growth of replant trees in apple orchards. Aust. J.
agric. Res., 4: 384-389.

GoLpEN, A. M., 1956.—Taxonomy of the spiral nematodes (Rotylenchus and Helrcotylenchus)
and the developmental stages and host-parasitic relationships of R. buxophilus n. sp.
attacking boxwood. Univ. Md. agric. exp. Sta. Bull. A-85: 1-28.

JENKINS, W. R., 1964.—A rapid centrifugation-flotation technique for separating nematodes
from soil. Pl. Dis. Rep., 48: 692.

Saver, M. R., 1962.—Distribution of parasitic nematodes in irrigated vineyards at Merbem
and Robinvale. Aust. J. expt. Agric. and An. Husb., 2: 8-11.

SErnHorst, J. W., 1956.—The quantitative extraction of nematodes from soil. Nematologica,
1: 249-267.

SErInHORST, J. W., and Sauer, M. R., 1956.—Eelworm attacks on vines in the Murray Valley
Irrigation Area. J. aust. Inst. agric. Sci., 22: 296-299.

DIURNAL VARIATION IN THE RELEASE OF POLLEN BY
PLANTAGO LANCEOLATA UL.

J. M. MATTHEWS
Research School of Pacific Studies, The Australian National University

(Communicated by Dr. D. Walker)
[Read 28th July, 1965]

Synopsis

An air sampler was used to record the pollen release of Plantago lanceolata. The pollen
catch indicates a tendency for pollen release to reach a major morning peak which is followed by a
secondary peak two to four hours later. Observations were also made on anther exertion.
Some of the observations indicate two main periods of anther exertion during the day.

INTRODUCTION

Percival (1955) has observed anther dehiscence for Plantago lanceolata at
hourly intervals. She records that anthers are presented from 0400 to 1700 hr.,
with a period of peak presentation from 0700 to 1000 hr. Anthesis can take
place when the relative humidity is 100% and is suppressed at temperatures
below 10°C (Percival, 1955). Hyde and Williams (1946) also observed that
flowering is suppressed below 10°C and that flowering may be profuse at high
humidity (80% or more) although dehiscence may then be delayed. Hyde
and Williams, who made observations of the impact and gravity catch on slides
exposed in a stand of Plantago lanceolata, noted that the catch rose steeply
between 0600 and 0800 hr. or 0800 and 1000 hr., slides were changed at two-
hourly intervals, and then fell more or less steeply (Hyde and Williams, 1946).

A simple air sampler was devised for the detection of pollen release by
Epacris paludosa. The effectiveness of the sampler was checked with Plantago
lanceolata which is known to be anemophilous. Pollen grains were collected
in quantity. It appeared from a single observation that the daily distribution
of pollen released by Plantago lanceolata has two peaks, the first major peak
being followed by a minor peak after a lapse of three hours. A series of
observations was made with the air sampler to investigate this diurnal variation.
Another set of observations was made on anther exertion.

ATR, SAMPLER

A tube 1’3” long and 12” in diameter was constructed of Bristol board.
A slot was cut at the mid point of the long axis for the insertion of a microscope
slide. The tube was attached to a “ Pifco vacette’’ vacuum cleaner by a
plasticene collar, the tube and cleaner being held in laboratory clamps.

The mouth of the tube was adjusted vertically to be at the point midway
between the highest and lowest heads on the plant under observation and 9”
from the nearest flowering head. The sampler was operated for periods of
three-quarters of an hour. The slides were changed at the end of each running
period and the cleaner switched off for a quarter of an hour to prevent over-
heating. The slides were smeared with silicone fluid (AK2000) and were
inserted in the tube with the adhesive surface towards the open end.

A cover slip (14” x 2”) was placed on each slide after its removal from
the tube. The cover slip was sealed with clear nail lacquer and the excess

PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH Watss, Vol. 90, Part 2

232 RELEASE OF POLLEN BY PLANTAGO LANCEOLATA

silicone fluid removed. The slides were than traversed four times under a
microscope at a total magnification of x 100 at intervals of five mm. The
mean number of grains observed per traverse was calculated for each slide.

The efficiency of the sampler is not known, but is probably low (ef. Gregory,
1961). The observed pollen catch should be taken as a relative rather than
an absolute record. Even though Gregory (1961) has pointed out that the
performance of an air sampler should be explored experimentally before use,
the assumption has been made that the mean number of grains observed per
traverse is directly proportional to the amount of pollen released by the plant
during the exposure of the slide.

OBSERVATIONS OF POLLEN RELEASE

Pollen release was recorded with the air sampler on five occasions (Table 1).
On four occasions the observations were made on a second floor balcony of
the H. C. Coombs Building, the Australian National University. At this time
the extensive stands of Plantago lanceolata on disturbed ground in the vicinity
of the H. C. Coombs Building had finished flowering, and a control run of the
sampler established only a negligible background of atmospheric pollen. The
balcony faces to the east and is shaded from direct sunlight, except between
0700 and 1000 hr. On 9.2.65 observations were made on a plant growing
in a garden some three miles to the north of the H. C. Coombs Building.

TABLE 1
Duration of observations of pollen release, and the number of heads on the observed plants

Duration of Number of Number of Number of
Date observations flowering immature mature
heads heads heads

21-1-65 0615-1800 25 16 =

9-2-65 0500-1545 15 17 2
11-2-65 0515-1700 20 11 8
19-2-65 0500-1645 19 2 1]
20:-2-65 0500-2400 19 2 1
21-2-65 0000—2400 19 2 1
22-2-65 0000-0445 19 2 1

The plants used at the H. C. Coombs Building were lifted from the same
garden the previous evening, placed in 6” earthernware flower pots and watered
copiously. On 21.1.65 and 11.2.65 the plant was watered at 0600 hr., but not
thereafter. On 19, 20, 21 and 22.2.65 a drip watering system was set up and
the plant watered continuously.

With the exception of the plant used on 21.1.65, all plants at the H. C.
Coombs Building were shaded from direct sunlight between 0700 and 1000 hr.

Observations of air temperature, humidity (when a whirling hygrometer
was available) and cloud cover were made at hourly intervals during the running
of the air sampler. On 9, 19, 20 and 21.2.65 air movement was also recorded.

Temperature and humidity were recorded on the balcony at the H. C.
Coombs Building five feet from the plant. In the garden, temperature and
humidity were recorded at a shaded station 25 yards away from the plant and
six feet above it.

Air movement was recorded on 19, 20 and 21.2.65 in arbitrary units by
a sensitive anemometer (C. F. Casella and Co., London, catalogue number
684A) adjacent to the plant. To prevent illumination of the plant no observa-
tions of air movement were made during the hours of darkness on 20, 21 and
22.2.65. Air movement was recorded in the garden adjacent to the plant

233

J. M. MATTHEWS

38%
15%

ege
&§ 8 8 8
oa ©
i ee

+ 75%
4 55%

]

+ 800

4 600

+ 400
200
0
10
8
6
4
2

15%

{160
4180
148
128
4108
se
60
40
4 20
0
e
'
15%
455%
35%

NO RECORD

18-2-1965
|
Record of pollen catch from Plantago lanceolata.

Vv SAINT NI
ANSH3AOW HIV $009

11-2-1965

9-2-1965
=<
eéesess
sess 8282

NO RECORD

4007
300

2 vow
8 SAIN Ni t

ANGH3AQH div

Fig. 1.

21-17-1965

—-
20-2-1965

fe i i ey at pote ry | a i tf (ts sy panty Cee eee ner treeel

e de Be. Sareps Seo Sarna S) 189) (8. SR SSeS RCS MTR e Gate Ghat ong leis eee earn meee
3OVYDILN39e Ni ALi Maske stnia ie te Sn ae Wy ie
GSUFAVSI/SNIVSS BIBHAN NYAW BuNiveadn3L div SALVvIGa JOVYSIANSI0 Ni ALIOINN ¥ SLINN NI “sy301

ASUSAVEI/SNIVHO BAGWNN NVaW SuNiVESdW31 div SAI 13a ANGWZAOW BV NI 32ZA09 GN019

e029 20
Gao es 6

NO RECORD

NO RECORD

Record of pollen catch from Plantago lanceolata.

Fig. 2.

J. M. MATTHEWS 235

the addition of two parts per 100 of soap solution. The stain was applied with
a hypodermic needle.

On each occasion four heads were observed (Figure 3). On 3.2.65 each
head was on a separate plant, on 5.2.65 on a single plant, and on 9.2.65 again
on different plants.

Once anthesis commenced on heads 1, 3, 5, 10, 11 and 12 it appeared to
proceed at a fairly constant rate until complete for the day. On heads 2, 4,
6, 7, 8 and 9 anthesis tended to occur in two bursts, the second following the
first after a period of two to five hours.

In order to observe any influence of the sun’s position on anther exertion,
each head was scored in four quadrants orientated SE-NE, NE-NW, NW-SW,
and SW-SH. The numbers of anthers exerted in each quadrant during three
periods are given in Table 2. On 3.2.65 there was a tendency for more anthers
to be exerted in the quadrant facing the sun than in the others. This was
not apparent on the other occasions.

TABLE 2
Numbers of anthers exerted in four quadrants

Quadrants
Date Time
SE-NE NE-NW NW-SW SW-SE
3.2.65 0800-1000 24 19 16 19
1000-1300 4 24 16 19
1300-1700 = — 10 5
5.2.65 0800—1000 30 23 24 28
1000-1300 14 1 8 6
1300-1700 — — — —
9.2.65 0800-1000 38 37 31 33
1000-1300 6 4 — 18
1300-1700 — — — —
DISCUSSION

The pollen catch at Canberra falls mainly within the limits noted by
Percival for anther dehiscence (Canberra Time is Eastern Standard Time).
On no occasion did the air temperature fall below 10°C. The late summer of
1965 was a particularly dry period with marked diurnal fluctuations in
temperature and humidity and with mainly clear skies.

On the six days when pollen release was observed the first peak occurred
between 0600-0700 hr. and 1100-1200 hr. (Table 3). With the exception of
21.2.65, the major peak was followed by a second peak (Table 4). The time
intervals between the two peaks vary from two to four hours (Table 5).

TABLE 3
Time of day, relative humidity and air temperature for first peak pollen catch

Date Time Relative Air
Humidity Temperature °C
21.1.65 0900-1000 — 18
9.2.65 0800-0900 30 24
11.2.65 1100-1200 40 22
19.2.65 0600-0700 — 22
20.2.65 1000-1100 30 31
21.2.65 1000-1100 37 27

236 RELEASE OF POLLEN BY PLANTAGO LANCEOLATA

TABLE 4
Time of day, relative humidity and air temperature for second peak pollen catch

Date Time Relative Air
Humidity Temperature °C

21.2.65 1300-1400 — 21

9.2.65 1100-1200 25 28

11.2.65 1600-1700 37 24

19.2.65 1000-1100 — 30

20.2.65 1500-1600 20 33

Neither air movement nor cloud cover appears to influence pollen catch
directly.

Hyde and Williams (1946) did not comment on two peak periods of pollen
release in Plantago lanceolata. However, their impact and gravity catches
record fluctuations in the pollen released from a large number of plants in a
stand, rather than from a single plant. Even so, their record (Hyde and
Williams, 1946) shows multiple peaks for impact catch on two out of nine days.
Hyde and Williams (1945) have noted that two grasses, Festuca rubra and
Holcus lanatus, as a rule flower slightly in the morning and more profusely in
the afternoon and that this is reflected in the pollen catch on gravity and
impact slides by a minor morning peak followed later in the afternoon by a
major peak.

The record of anther exertion is ambiguous ; it is possible that the staining
technique may have interfered. Nevertheless anthesis tended to take place
in two bursts on six of the 12 observed heads (cf. Centaurea nigra: Percival

TABLE 5
Duration of interval between first and second peaks

Interval between.
Date first and second
peak in hours

21.1.65 3

9.2.65 9
11.2.65 4
19.2.65 3
20.2.65 4

1950). There is some indication that the position of the sun influences the
course of anther exertion in Plantago lanceolata (cf. Papaver orientale: Percival,
1950).

On 5.2.65 and 9.2.65 anthesis had begun at or before 0600 hr. On
3.2.65 the flowering heads were saturated by a heavy dew. The plants were
growing along the foot of a garden fence which shaded them from the early
morning sun. However, it was noted that sunlight fell on heads 2 and 4
between 0700 and 0800 hr. through cracks in the fence ; anther exertion began
at this time. Heads 1 and 3 were in the shade of the fence until 0900 hr.,
when anthesis began.

CONCLUSIONS

Pollen release in Plantago lanceolata may reach two peaks during the day
but there is some variation in this behaviour. The data are not sufficiently
detailed to allow the detection of causal factors.

Speculations can be made about the nature of the circumstances which
lead to diurnal variation in pollen release. If it is supposed that anthers are
exerted at regular intervals during the day and that early morning conditions

J. M. MATTHEWS 231

are unfavourable to pollen release, then a relatively large release of pollen is
to be expected when the inhibition is removed. Thereafter pollen release should
fall to a constant rate. However, the frequently observed double peak of pollen
release and the less certain parallel behaviour in anther exertion do not support
this hypothesis. It would seem more likely that the first main peak represents
the attainment of favourable conditions for pollen release and that the magnitude
of this peak reflects the accumulation of exerted anthers in which dehiscence
has not, or has only partly, taken place. The second peak would then reflect
the second burst of flowering, although it appears to be a more common
phenomenon than the observation of anther exertion would substantiate.

Acknowledgements

Acknowledgement is made to Professor O. H. K. Spate, in whose Depart-
ment this work was carried out. Dr. Donald Walker gave invaluable assistance
with the planning of the observations and the operation of the air sampler,
and has read a draft of this paper.

References

Grecory, P. H., 1961.—~* The microbiology of the atmosphere’’. Leonard Hill Interscience.
London and New York; 96—107.
Hyove, H. A., and Wituiams, D. A., 1945.—Studies in atmospheric pollen. II. Diurnal variation
in the incidence of grass pollen. New Phytologist, 44: 83-94.
, , 1946.—Studies in atmospheric pollen. ILI. Pollen production and pollen
incidence in the ribwort plantain (Plantago lanceolata L.). New Phytologist, 45: 271-7.
PrrRoivaL, M., 1950.—Pollen presentation and pollen collection. New Phytologist, 49: 40-63.
, 1955.—The presentation of pollen in certain angiosperms and its collection by Apis
mellifera. New Phytologist. 54: 353-68.

THE VERTEBRATE FAUNA OF “ GILRUTH PLAINS ”,
SOUTH-WEST QUEENSLAND

M. G. BRooKER! and G. CAUGHLEY?

(Communicated by Dr. Mervyn Griffiths)

[Read 28th July 1965]

INTRODUCTION

This paper presents a list of the vertebrates identified on a defined area
in the semi-arid zone of south-west Queensland. Despite the apparently harsh
temperature and rainfall conditions, more than 200 species were observed during
a three-year period.

In addition, an attempt has been made to give some indication of the
density and seasonal distribution of the birds and mammals.

ENVIRONMENT

‘‘Gilruth Plains” is situated near Cunnamulla, latitude 28° S (approxi-
mately), longitude 146° (approximately). It is a 40,000 acre C.S8.I.R.O. field
station concerned with research into wool production. The climate is semi-arid
with a 15-inch mean annual rainfall.

We consider that ‘“ Gilruth Pains” should give a representative picture
of the fauna of this zone because :

(i) The vegetation is diverse and contains examples of most associations
typical of this semi-arid region. ‘‘ Gilruth Plains” is primarily a mosaic of
mulga-box (Acacia aneura—Eucalyptus populnea), gidgee-buddah (Acacia cam-
bagei—Eremophila mitchelli), ironbark-spinifex (Hucalyptus melanophloia—Triodia
spp.) and Mitchell grass (Astrebla spp.) with small areas of coolibah (Hucalyptus
microtheca), pine (Callitris columellaris), and swamp.

(ii) Seasonal conditions during the period of observations represented the
normal climatic pattern with a range from severe drought to lush pasture
conditions. Permanent surface water was always available in bore drains and
small dams while large areas of temporary swamp formed after heavy rain.

METHOD OF RECORDING

The following list of vertebrates was compiled during the period October
1960 to October 1963.

While the authors did devote some time to searching specifically for
vertebrate fauna, especially for birds and mammals, a large number of the
Species were either observed when the authors were engaged in other field work
or collected by members of the staff of ‘“‘ Gilruth Plains ”’.

The fish species were identified by Mr. G. P. Whitley of The Australian
Museum. Species of Amphibia and Reptilia for which there are housed specimens
(marked H on list) were identified by Mr. H. G. Cogger of The Australian
Museum. The location of housed specimens is shown on the list thus: Hg,
Specimen(s) housed at Gilruth Plains; Hm, Specimen(s) housed at The
Australian Museum.

1 Division of Animal Genetics, C.8.I1.R.O., National Field Station, “ Gilruth Plains ’”’,
Cunnamulla, Queensland.

2 School of Biological Sciences, University of Sydney, Sydney. Present address: Forest
and Range Experimental Station, Rangiora, New Zealand.

PRCOEEDINGS OF THE LINNEAN Society oF New SoutH Watss, Vol. 90, Part 2

M. G. BROOKER AND G. CAUGHLEY 939

The Australian Reed-Warbler (Acrocephalus australis) and Gould’s Wattled
Bat (Chalinolobus gouldii) were identified by Mr. Warren Hitchcock and Mr.
John Calaby respectively, of C.S.I.R.O. Division of Wildlife Research, Canberra.
The remaining frog and reptile species were authoritatively determined, although
housed specimens of these species are not available.

All other species of birds and mammals were identified by one or both

of the authors. The 9th Edition of ‘‘ An Australian Bird Book” by J. A.
Leach was used as a reference for bird identification.

Where possible, each bird or mammal species is placed in one of four broad
density classes depending on how often an observer is likely to see it.

The classes are defined as follows: A, At least once per day ; B, Once per
10 days; C, Once per 100 days; D, Once per 1000 days.

Additional symbols used were: M, Species is migratory and is thus present
for only part of each year. The density class of an M species refers only to
part of the year; S, When the presence of species appears to be governed by

seasonal conditions ; E, Exotic species.

FisH

Madigania unicolor (Native Grunter) Hg
Gambusia affinis (Mosquito Fish) EH Hg
Carassius auratus (Goldfish) EK Hg

FrRoes

Cyclorana australis WHmg

C. alboguttatus

C. platycephalus Hm

C. cultripes Hm

Notaden bennetti Hmg
Limnodynastes fletcheri WHmg
Hyla rubella Hmg »
H. caerulea Hm

Uperoleia marmorata Hg

TORTOISE

Chelodina longicollis (Long-necked Tortoise)

SNAKES

Pseudechis australis (Mulga Snake) Hmg

Demansia nuchalis—carinata group. (Brown
Snake) Hmg

Denisonia suta (Whip Snake) Hm

Brachyurophis australis (Australian Coral
Snake) Hg

LiIzaRDS
Geckonidae

Diplodactylus tessellatus (Variegated Gecko)
Hmg

D. strophurus

D. wiliamsi Hmg

Lucastus damaeus (Bearded Gecko)

Nephrurus laevis (Smooth Knob-tailed
Gecko) Hg

Heteronota binoet (Bynoe’s Gecko) Hmg

Rhynchoedura ornata (Beaked Gecko) Hm

Gehyra variegata (Dtella) Hmg

Oedura marmorata (Velvet Gecko) Hg

Agamidae

Amphibolurus barbatus (Jew Lizard) Hmg
A. muricatus (Tree Dragon) Hmg

Scincidae

Tiliqua scincoides (Blue Tongue) Hg

Trachysaurus rugosus (Shingle-back) Hg

Hgernia striolata (Arboreal Skink) Hmg

Ablepharus bouton Hmg

A. lineo-ocellatus Hm

A. timidus Hmg

Sphenomorphus lesueurit (Lesueur’s Skink)
Hmg

Rhondona punctatovittatum Hmg

Pygopodidae
Lialis burtonii (Common Snake Lizard)
Hmg
Pygopus nigriceps

Varanidae

Varanus gouldi (Gould’s Goanna) Hg
V. tristis (Black Goanna) Hg

Brrps

Dromaius novae-hollandiae (Emu) A
Podiceps poliocephalus (Hoary-headed Grebe)
CS

P. ruficollis (Little Grebe) BS

Pelecanus conspicillatus (Pelican) CS

Phalacrocorax carbo (Black Cormorant) CS

P. melanoleucos (Little Pied Cormorant) CS

P. sulcirostris (Little Black Cormorant) CS

Ardea novae-hollandiae (White-faced Heron)
A

A. pacifica (White-necked Heron) A

Hgretta alba (White Egret) DS

E. garzetta (Little Egret) DS

Threskiornis molucca (White Ibis) CS

T. spinicollis (Straw-necked Ibis) BS

Plegadis falcinellus (Glossy Ibis) DS

Platalea flavipes (Yellow-billed Spoonbill)
BS

P. regia (Royal Spoonbill) CS

Cygnus atratus (Black Swan) CS

Chenonetta jubata (Maned Goose) B

Anas gibberifrons (Grey Teal) A

A. rhynchotis (Blue-winged Shoveler) DS

A. superciliosa (Black Duck) BS

Malacorhynchus membranaceus (Pink-eared
Duck) B

240 VERTEBRATE FAUNA OF GILRUTH PLAINS, S.W. QUEENSLAND

Aythya australis (White-eyed Duck) CS

Circus assimilis (Spotted Harrier) C

Accipiter cirrocephalus (Collared Sparrow-
Hawk) B

Hieraaetus morphnoides (Little Eagle) C

Aquila audax (Wedge-tailed Eagle) A

Haliastur sphenurus (Whistling Eagle) A

Milvus migrans (Fork-tailed Kite) A

Hlanus notatus (Black-shouldered Kite) D

Falco berigora (Brown Hawk) B

F'. cenchroides (Nankeen Kestrel) A

F. longipennis (Little Faleon) B

Coturnix pectoralis (Stubble-Quail) BS

Turnix velox (Little Quail) AS

Grus rubicundus (Brolga) B

Tribonyx ventralis (Black-tailed Water-Hen)
CS

Fulica atra (Coot) DS
Hupodotis australis (Bustard) C
Lobibyx novae-hollandiae (Spur-winged Plover)

Zonifer tricolor (Banded Plover) BS

Charadrius melanops (Black-fronted Dotterel)
A

Hrythrogonys cinctus (Red-kneed Dotterel)
CS

Hrolia acuminata (Sharp-tailed Sandpiper) D
Rostratula benghalensis (Australian Painted

Snipe) DS

Himantopus leucocephalus (White-headed
Stilt) A

Recurvirostra novae-hollandiae (Red-necked
Avocet) CS

Stiltia isabella (Australian Pratincole) C

Burhinus magnirostris (Southern Stone-Cur-
lew)

Larus novae-hollandiae (Silver Gull) DS

Chlidonias hybrida (Marsh Tern) CS

Geopelia cuneata (Diamond Dove) A

G. placida (Peaceful Dove) A

Phaps chalcoptera (Forest Bronzewing) A

Histriophaps histrionica (Flock Pigeon) D

Ocyphaps lophotes (Crested Pigeon) A

Kakatoe galerita (White Cockatoo) D

K. leadbeatert (Pink Cockatoo) A

K. roseicapilla (Galah) A

Leptolophus hollandicus (Quarrion) A

Aprosmictus erythropterus (Red-winged Parrot)
A

Barnardius barnardi (Ring-neck Parrot) A

Psephotus haematogaster (Blue Bonnet) A

P. varius (Mulga Parrot) A

Melopsittacus undulatus (Budgerygah) A

Cuculus pallidus (Pallid Cuckoo) A

Chalcites basalis (Horsfield Bronze-Cuckoo)
C

Cacomantis pyrrhophanus (Fan-tailed Cuckoo)
D

Scythrops novae-hollandiae (Channel-billed
Cuckoo) C

Tyto alba (Barn Owl) C

Ninox novae-seelandiae (Boobook) B

Hurostopodus guttatus (Spotted Nightjar) B

Aegotheles cristata (Crested Owlet-Nightjar)
B

Podargus strigoides (Tawny Frogmouth) B

Hurystomus orientalis (Dollar-Bird) D

Apus pacificus (Fork-tailed Swift) C

Dacelo gigas (Laughing Kookaburra) A

Halcyon pyrrhopygius (Red-backed King-
fisher) A

H. sanctus (Sacred Kingfisher) B

Merops ornatus (Rainbow-Bird) AM

Mirafra javanica (Horsfield Bushlark) B

Hirundo neoxena (Welcome Swallow) C

Hylochelidon ariel (Fairy Martin) A

H. nigricans (Tree Martin) A

Pteropodocys maxima (Ground Cuckoo-Shrike)
A

Coracina novae-hollandiae
Cuckoo-Shrike) A

C. robusta (Little Cuckoo-Shrike) B

Lalage sueurti (White-winged Triller) A

Rhipidura fuliginosa (Grey Fantail) AM

R. leucophrys (Willie Wagtail) A

Seisura inquieta (Restless Flycatcher) A

Microeca fascinans (Jacky Winter) A

Petroica goodenovit (Red-capped Robin) A

P. cucullata (Hooded Robin) A

Acrocephalus australis (Australian Reed-
Warbler) D

Pachycephala rufiventris (Rufous Whistler) A

Colluricincla harmonica (Grey Shrike-Thrush)
A

Oreoica gutturalis (Crested Bellbird) A

Pomatostomus temporalis (Grey-crowned Bab-
bler)

Epthianura aurifrons (Orange Chat) CS

H. tricolor (Crimson Chat) BS

Gerygone fusca (Fuscous Warbler) A

Smicrornis brevirostris (Brown Weebill) A

Acanthiza chrysorrhoa (Yellow-tailed Thorn-
bill) A

A. nana (Little Thornbill) B

A. uropygialis (Chestnut-tailed Thornbill) A

Aphelocephala leucopsis (Eastern Whiteface)
A

(Black-faced

Malurus assimilis (Purple-backed Wren) A
M. leuconotus (White-winged Wren) A

M. melanotus (Black-backed Wren) C
Cinclorhamphus cruralis (Brown Songlark) A
C. mathewsi (Rufous Songlark) B

Anthus australis (Pipit) A

Artamus cinereus (Black-faced Wood-Swallow)

A. leucorhynchus (White-breasted Wood-
Swallow) BS

A. minor (Little Wood-Swallow) CS

A. personatus (Masked Wood-Swallow) CS

A. superciliosus (White-browed Wood-
Swallow) AS

Climacteris affinis
Creeper) C

C. picumnus (Brown Tree-Creeper) A

Neositta chrysoptera (Orange-winged Sittella)
B

(White-browed Tree-

Dicaeum hirundinaceum (Mistletoe-Bird) A
Pardalotus substriatus (Red-tipped Pardalote)

A
P. rubricatus (Red-browed Pardalote) C
Melithreptus brevirostris (Brown-headed
Honeyeater) 5B
Meliphaga penicillata (White-plumed Honey-
eater)
M. virescens (Singing Honeyeater) A
Myzomela nigra (Black Honeyeater) CS
Plectorhyncha lanceolata (Striped Honeyeater)
A

Grantiella picta (Painted Honeyeater) DS
Gliciphila indistincta (Brown Honeyeater) A
Myzantha flavigula (White-rumped Miner) A

Proc. LINN. Soc. N.S.W., Vol. 90, Part 2 PLATE Iv

Tachyglossus aculeatus.

1
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=
:
a
x
i
"
i 7%
Tea nee
i i
py
ne

Proc. Linn. Soc. N.S.W., Vol. 90, Part 2 PLATE

Tachyglossus aculeatus.

Ye eee ‘
ie ie ‘ f fail . ‘ ae
=F a { \ ae gaa oe
ae) re ea ?
¥ pre of ‘ q q 2 an
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. A
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PLATE VI

Proc, Linn, Soc, N.S.W., Vol. 90, Part 2

2-5, 8-11. Macgeea tout,

1, 6, 7. Bensonastraea praetor,

; ; : aM Gnd, Ph hey me a a won ite Rw ni

ty SR Ty ; ; ir

ME

pe oi rae if
pela ne

Proc. Linn. Soc, N.S.W., Vol. 90, Part 2 PLATE VO

Development of egg of Retropinna semoni (Weber).

rn rad > : apr ex: ' ———- — Bp
| ty Oe WR Dol rh OR
ee / i” af ‘a el ere ™ i : )

ee
ty PAD

tee

7 won| ie
pS Tae

+5

Proc. Linn. Soc. N.S.W., Vol. 90, Part 2 PLATE VII

Rees Scene

ee

__. He 12

Development of egg of Retropinna semoni (Weber).

7

7 — . : ie” - in -
PO Lo A al

iy eee

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Proc. Linn. Soc. N.S.W., Vol. 90, Part 2 IBIAS IDX

Larvae of Retropinna semoni (Weber).

i
ney
My
nui

M. G. BROOKER AND G. CAUGHLEY 241

Acanthagenys rufogularis (Spiny-cheeked
Honeyeater) A

Entomyzon cyanotis (Blue-faced Honeyeater)

Philemon citreogularis (Little Friar-Bird) BS
P. corniculatus (Noisy Friar-bird) AS
Passer domesticus (House Sparrow) DE

Struthidea cinerea (Apostle-bird) A
Grallina cyanoleuca (Magpie-Lark) A
Chlamydera maculata (Spotted Bower-bird) A

MAMMALS
Tachyglossus aculeatus (Echidna) C

Macropus rufus (Red Kangaroo) A

M. robustus (Wallaroo) D

M. canguru (Grey Kangaroo) A

Sminthopsis macrura-crassicaudata group
(Fat-tailed Marsupial Mouse) C

Chalinolobus gouldit (Gould’s Wattled Bat)

Vulpes vulpes (Fox) AE

Felis catus (Cat) AE

Mus musculus (Mouse) BE

Capra hircus (Goat) CH

Sus scrofa (Pig) CE

Orytolagus cuniculus (Rabbit) AE

Zonaeginthus guttatus (Diamond-Firetail) D

Taeniopygia castanotis (Zebra Finch) A

Steganopleura bichenovii (Banded Finch) A

Oriolus sagittatus (Olive-backed Oriole) B

Corvus bennetta (Little Crow) B

C. cecilae (Crow) C

C. coronoides (Raven) A

Cracticus nigrogularis (Pied Butcher-bird) A

C. torquatus (Grey Butcher-bird) A

Gymnorhina tibicen (Black-backed Magpie) A

Corcoraxz melanorhamphus (White-winged
Chough) B

ADDENDUM

The following animals were seen on “ Gilruth Plains” but were not
identified with sufficient confidence to justify placing them on the main list.
They should not be quoted as locality records until their presence has been
confirmed.

Acanthophis antarcticus (Death-Adder) Phascogale tapoatafa (Brush-tailed Phas-

Falco peregrinus (Peregrine Falcon) cogale

Acanthiza lineata (Striated Thornbill) Trichosurus vulpecula (Brush-tailed Possum)

Pomatostomus ruficeps (Chestnut-crowned Chalinolobus morio (Chocolate Wattled Bat)
Babbler)

DISCUSSION

The vertebrate fauna of this relatively small area consists of at least three
fish, nine frogs, one tortoise, four snakes, 23 lizards, 151 birds and 12 mammals,
giving a total of more than 200 species. It is apparent that this climatic zone
is in NO Sense inimical to vertebrates. It would be of great interest to compare
the size of this fauna with that of an area of comparable size in the arid or
temperate zone, but we know of no similar surveys in these regions.

Acknowledgements

The authors are indebted to Messrs. G. P. Whitley and H. G. Cogger of
The Australian Museum, Sydney, and Messrs. W. Hitchcock and J. Callaby of
C.S.I.B.O. Division of Wildlife Research, Canberra, for identifying specimens,
to Mr. C. H. S. Dolling, Officer in Charge, ‘*‘ Gilruth Plains ”’, for his continuous
encouragement of the work, and to Mr. M. Schrader of Cunnamulla for critical
reading of the manuscript.

FURTHER OBSERVATIONS ON THE LIFE HISTORIES
OF LITTORAL GASTROPODS IN NEW SOUTH WALES

D. T. ANDERSON
School of Biological Sciences, University of Sydney

(Plate x)

[Read 29th September, 1965]

Synopsis

The spawn and early development are described for: (1) Xenogalea labiata (Perry)
(Cassididae), in which several females appear to contribute capsules, each containing numerous
small eggs, to a large common egg mass; (2) Bedeva hanleys (Angas) (Muricidae), which lays
dome-shaped capsules in which many eggs are consumed as nurse eggs and only a few embryos
hatch as well developed crawling juveniles; (3) Morula marginalba (Blainville) (Thaididae),
which lays bluntly rounded capsules containing numerous eggs hatching as planktotrophic
veligers ; (4) Nassarius (Alectrion) particeps (Hedley) (Nassariidae), which lays numerous stalked
triangular capsules each containing a single egg developing to a crawling stage before hatching ;
(5) Stphonaria denticulata Quoy and Gaimard (Siphonaridae), which lays gelatmous egg strings
containing numerous eggs hatching as planktotrophic veligers.

Each species is briefly discussed in relation to other species of its family, and a summary
is given of development in N.S.W. littoral prosobranchs.

INTRODUCTION

A number of authors have recently described the spawn and early
development of Australian littoral gastropods (H. Anderson, 1958; D. T.
Anderson, 1959, 1960, 1961, 1962, 1965; MacIntyre, 1961; Murray, 1962a,
19626, 1963, 1964). The present paper reports further observations on this
subject for the mesogastropod Xenogalea labiata (Perry) (Cassididae), the
neogastropods Bedeva hanleyi (Angas) (Muricidae), Morula marginalba (Blain-
ville) (Thaididae) and Nassarius particeps (Hedley) (Nassariidae), and the
pulmonate limpet Siphonaria denticulata Quoy and Gaimard (Siphonariidae).
Of these species, the spawn and development of X. labiata, M. marginalba and
N. particeps have not hitherto been described. The egg capsules of B. hanleyt
were figured by Hedley (1916) and Roughley (1925) and the egg strings of S.
denticulata by Dakin (1953; Plate 55), but development of the eggs of these
species has not been investigated.

MATERIALS AND METHODS

Materials for the studies described in this paper have been gained from
several sources. The egg mass of X. labiata was collected by Miss I. Bennett
of the School of Biological Sciences, University of Sydney, from a sub-littoral
rock face at Fairlight, N.S.W., in November, 1963. It was preserved shortly
after collection, and observations have been made only on the egg mass itself
and on the single developmental stage which it contains. The egg mass of
N. particeps was also collected by Miss I. Bennett, at Long Reef, N.S.W., in
October, 1962. This mass was maintained in aerated seawater in the laboratory
and a number of observations made on the development of the embryos.

Capsules identified as those of B. hanleyi by comparison with the descrip-
tions given by Hedley (1916) and Roughley (1925) were collected on numerous
occasions during the winter months of 1962 and 1963 on the undersurfaces of
loose rocks in tidal pools at Bradley’s Head, on the north shore of Port Jackson.
Their absence from this locality during the remainder of the year indicates

PROCEEDINGS OF THE LINNEAN SocteTy oF NEw SoutH WALzEsS, Vol. 90, Part 3

D. T. ANDERSON 243

that the species is a winter breeder. These capsules were also maintained in
aerated seawater in the laboratory, and observations made on the development
of the embryos.

Morula marginalba was taken spawning on the underside of mid-littoral
rocks at Long Reef, N.S.W., in January, 1963. The egg capsules were collected,
maintained in aerated seawater in the laboratory and studied at intervals until
the embryos hatched. Repeated searching at this and other localities where
Morula is common has failed to provide further capsules, so that the general
breeding habits are not yet clear. The ovaries of females, however, do not
contain ripe eggs during the winter months.

Observations made at frequent intervals in the spring and summer of
1961/62 and 1962/63 at Harbord, N.S.W., and Long Reef, N.S.W., showed that
Siphonaria denticulata breeds at least from September to March in these
localities. Numerous egg masses were found attached to rock surfaces in the
habitat of the adults, and animals were frequently seen in the act of spawning.
Masses collected on various occasions were maintained in aerated seawater in
the laboratory and studied at intervals until the embryos hatched.

Drawings of spawn, embryos and larvae investigated were made with the
aid of a camera lucida. The photograph of Plate x was taken by the Department
of Illustration, University of Sydney.

RESULTS
Xenogalea labiata

The spawn of X. labiata (Plate x) is a large, irregular, sponge-like mass
consisting of several thousand egg capsules. The entire mass, when collected,
had dimensions of about 30 cm., and observations made at the time of spawning
suggest that it is the common product of several females spawning together.
When fresh, the mass was pinkish-orange in colour, the colour being imparted
by the eggs contained in the capsules.

The capsules themselves are colourless and translucent and are cemented
together as shown in Figure 1. Each capsule is roughly rectangular in shape,
about 2-5mm. long and 1:5mm. broad, with a thin irregular base plate
attached to the capsules below it in the mass. The capsule contains a colour-
less albumen in which float several hundred eggs, each about 160 u in diameter,
filled with pale orange yolk. All the eggs in the capsule develop simultaneously,
proceeding through a yolk-filled gastrula stage (Fig. 2). Development beyond
this stage was not observed.

Bedeva hanleyr

The spawn of B. hanleyi consists of a group of about 10-20 separately
attached, dome-shaped capsules, each about 3mm. in diameter, whitish in
colour and semi-transparent, with a transparent oval apical membrane (Fig. 3).
Inside the capsule lie between 50 and 70 eggs, floating in a colourless albumen.
A horny transparent base plate completes the capsule structure.

The eggs are opaque white, very yolky, and ovoid in shape, with a long
diameter of 250 yu. Only about 15 eggs develop in each capsule, the remainder
serving as nurse eggs for the developing embryos. A yolky early veliger stage,
with an inconspicuous velum, simple colourless shell, small foot, and large,
yolky visceral mass develops in about 4 days and begins to move slowly through
the jelly in the capsule. Within a further 4 days, enlargement of the velum
occurs, accompanied by elongation and ciliation of the foot and outgrowth of
a convex, ciliated oral hood. The stomodaeum also becomes well developed,
preliminary to the onset of the ingestion of nurse eggs. Further development
during the next 7 days results in broadening of the velum and slight subdivision
into 4 velar lobes, formation of black eyespots on either side of the oral hood,
and the beginning of spiral coiling of the shell (Fig. 4). As the velar lobes

244 LIFE HISTORIES OF LITTORAL GASTROPODS

enlarge, each becomes wrapped around a nurse egg which rests against its
concave posterior face. This, however, appears to be a simple consequence of
the crowded conditions within the capsule and not a factor in nurse egg
absorption.

Direct feeding on the nurse eggs now begins and is completed in about
5 days, during which the veligers retain a large active velum and glide rapidly
through the albumen of the capsule. Some growth occurs during this phase,
but the major part of the ingested yolk becomes stored in the visceral mass.
Subsequent growth at the expense of the stored yolk proceeds at a slightly
variable rate in different embryos in the capsule for a further 11 days. Gradual
velar resorption is accompanied by elaboration of the head, foot and visceral
mass and growth of the coiled shell, which becomes brown-pigmented (Fig. 5).

Figs 1-2. Xenogalea labiata. 1, egg capsules; 2, gastrula stage.

The fully developed juveniles, about 44 weeks old, escape in rapid succession
from the capsule through an aperture formed by breakdown of the apical
membrane. The newly hatched, crawling juvenile (Fig. 6) has a shell length
of about 900 xu.

Morula marginalba

The spawn of M. marginalba consists of groups of about 20 low, rounded
capsules. Each capsule (Fig. 7) has a tough, translucent wall, normally whitish
in colour but sometimes tinged with purple, with a transparent thin oval area
in the middle of one side through which hatching later occurs. The capsule
is attached to the rock by an irregular, transparent, horny base plate and is
filled with a colourless albumen in which float between 100 and 200 spherical,
yellow, yolky eggs 220 in diameter.

The eggs in the capsule develop simultaneously, passing through a yolky
trochophore stage to an early veliger stage (Fig. 8) within 2 days of oviposition.
In the early veliger, the velum is still rudimentary, but a ciliated oral hood
and stomodaeum are well developed, the foot rudiment is conspicuous and the
yolky visceral mass is covered dorsally by a simple transparent shell.

D. T. ANDERSON 245

During the next 3 days, the veliger becomes well developed (Fig. 9) and
begins to move actively through the capsule albumen by means of its velar
cilia. Torsion occurs and the shell enlarges and begins to develop a spiral
form. The foot elongates, becomes ciliated, develops paired statocysts and
secretes an operculum. The velar lobes expand and develop paired brown
eyespots. The visceral mass remains yolky and undifferentiated, however, and
no withdrawal of the head and foot into the shell is observed.

During the ensuing week, the veliger enlarges further and becomes highly
active and much more differentiated (Fig. 10). The oral hood is reduced in
size, but the velar lobes become larger and brown-pigmented around their
margins. At the base of the right lobe, a tentacle grows out, tipped with a
fan of stiff cilia. The foot also enlarges and becomes brown-pigmented on its
ventral face. Further growth and coiling of the shell is accompanied by the

Figs 3-6. Bedeva hanleyi. 3, egg capsule; 4, 15-day veliger; 5, 25-day veliger; 6, newly
hatched juvenile.

formation of numerous yellowish spots on the shell surface, together with the
deposition of brown pigment at the umbo and around the margin (Fig. 11).
The visceral mass develops a pulsating larval heart and a coiled gut in which
the stomach is black-pigmented and contains yolk particles rotated by ciliary
action. The yellowish yolk is now confined to the apex of the visceral mass.

Due to the increased pigmentation of the developing veligers, egg capsules
12 or more days old are brown in colour, in contrast to the yellowish colour of
fresher capsules.

Between 12 and 18 days after oviposition, little further change is observed
in the veligers in the capsule, apart from a gradual reduction in the amount
of yolk in the visceral mass. The veligers continue to swim actively in the
capsule jelly, however, and at about 18 days the thin window in the side of
the capsule breaks down and the veligers escape to a free swimming existence.
Their development after hatching was not followed. :

Nassarius particeps

The egg mass of N. particeps used in the present investigations was akan
at the time of oviposition. Two females were associated with it, and both

246 LIFE HISTORIES OF LITTORAL GASTROPODS

proved to have numerous ripe oocytes in their ovaries. Whether both were
contributing capsules to the egg mass could not be decided with certainty, but
the extent of the mass suggested possible cooperative spawning. The mass
consisted of several hundred small, stalked, triangular capsules attached by

Figs 7-11. Morula marginalba. 17, egg capsule; 8, 2-day veliger; 9, 5-day veliger; 10, 12-day
veliger; 11, shell of 12-day veliger.

confluent base plates as a single layer directly to the under surface of a rock
in the lower littoral, and covered an area of many square cms.

Each egg capsule of N. particeps (Fig. 12) is about 2-5 mm. in height
colourless and translucent, and filled with a colourless albumen in which floats
a single large, yellowish, spherical egg about 700 in diameter. In spite of
the size of the egg, cleavage is total and the first two divisions are only slightly

D. T. ANDERSON Q47

unequal. Development proceeds through a highly modified veliger phase in
which the velar lobes, although ciliated, remain relatively small and produce
only slow rotation of the embryo inside the capsule. Development of the head,
foot and shell are well advanced after 7 days (Fig. 13), although the large
visceral mass is still yolk-filled and little differentiated. By 12 days, reduction
of the velum is in progress and the yolk reserves are beginning to lessen.
Development was not followed beyond this point but there seems little doubt
that NV. particeps hatches as a crawling juvenile, probably about 1 mm. in length.

Siphonaria denticulata

The egg mass of S. denticulata is an irregular, gelatinous, spiral coil about
5 em. long, creamy-white in colour and firmly glued to the rock surface in the
habitat of the adults. The jelly is colourless and the numerous closely-packed

300

I3

Figs 12-13. Nassarius particeps. 12, egg capsule; 13, 7-day veliger.

Figs 14-17. Siphonaria denticulata. 14, embryo after 3rd cleavage, from animal pole;
15, 3-day veliger; 16, 5-day veliger; 17, newly hatched veliger.

eggs embedded in it are whitish and translucent. Hach egg is about 100» in
diameter and is enclosed in an ovoid, transparent egg capsule about 200 » long
and 160 u broad.

The eggs cleave in a typical spiral manner, the first two divisions being
equal, the third unequal (Fig. 14). Blastula and gastrula phases are passed
through in 24 hours and a simple yolky trochophore is completed 48 hours
after oviposition. During the third day of development, the operculate foot,
bilobed velum and globular shell of an early veliger develop (Fig. 15). Growth
and internal differentiation now set in and the veliger begins to rotate vigorously
within its capsule. The ciliated gut becomes conspicuous as the yolk reserves
are finally resorbed (Fig. 16) and hatching of the veligers as actively swimming
planktotrophic larvae (Fig. 17) occurs on the sixth day after oviposition.

Hatched veligers fed on the diatom Nannochloris continued to swim and
grow for a further 4 days, becoming more differentiated internally and developing
a slight spiral twist to the shell. Laboratory culture beyond this stage was
not achieved.

248 LIFE HISTORIES OF LITTORAL GASTROPODS

DISCUSSION
Xenogalea labiata
Very little is known of spawning in cassidid mesogastropods. The only
two previous descriptions of egg capsules appear to be those of Erlanger (1893)
for Cassidaria echinophora and Lo Bianco (1899) for C. echinophora and Cassis
sulcata. These brief notes suggest that the egg mass of X. labiata is typical
for the family, but it is not yet known whether the apparent cooperative spawning
of X. labiata is a characteristic of cassidids.

Embryonic and larval development in the Cassididae remain to be described,
but the number and dimensions of the eggs of X. labiata suggest that this species
hatches as a pelagic planktotrophic veliger. In contrast, it appears from the
work of Erlanger (1893) and Lo Bianco (1899) that the eggs of Cassidaria
echinophora are larger (280. in diameter) and that only a few develop, the
remainder serving as nurse eggs, to hatch at an advanced stage, probably as
crawling juveniles.

Bedeva hanleyi

The present work confirms the descriptions given by Hedley (1916) and
Roughley (1925) of the egg capsules of B. hanleyi and also shows that develop-
ment includes feeding on nurse eggs and ends in hatching from the capsule as
a crawling juvenile. Hatching at the crawling stage is characteristic of
Muricidae (Anderson, 1960; MacKenzie, 1961; Murray, 1963, 1964) and, as
pointed out by Thorson (1946), dome-shaped capsules of the type laid by B.
hanleyt have been described for several species of Trophon, the genus to which
B. hanleyi is alternatively referred. Feeding on nurse eggs has not previously
been recorded in this genus, but is well known for other muricids (e.g. species
of Murex, Neptunea, Nucella: Lebour, 1937; Ankel, 1937, 1938; Thorson,
1935, 1946 ; Natarajan, 1957; Golikov, 1961) and may yet be found in other
species of Trophon or Bedeva.

Morula marginalba

Spawning and development in Thaididae have been described for a number
of species of Thais (e.g. Burkenroad, 1931; Chari, 1950; Butler, 1953;
Natarajan, 1957), all of which produce egg capsules of the general type
exemplified by M. marginalba, though the shape of the capsule and the position
of the hatching membrane vary in different species. Dakin (1953) has also
figured massed capsules of the same general type for Dicathais orbita. Only
one previous reference has been made to the egg mass of a species of Morula,
however, that of Ostergaard (1950) for M. (= Drupa) dumosa and the gelatinous
egg mass described is so aberrant for the family that it seems likely to be a
misidentification.

The mode of development of the numerous encapsulated eggs in WM.
marginalba, and the form of the pelagic planktotrophic veligers at hatching,
are closely similar to those of several species of Thais (e.g. T. haemastoma :
Burkenroad, 1931; Franc, 1948; Butler, 1953; 7. bufo: Natarajan, 1957; T.
javanica: Natarajan, 1957; 7. species B, T. species C: Natarajan, 1957).
The genus Thais, however, also includes species with larger eggs which hatch
at a later stage of development and are only briefly pelagic (e.g. 7. tssoti:
Natarajan, 1957) and species which develop directly to hatching as crawling
juveniles (e.g. 7. emarginata: Dehnel, 1955). It cannot be assumed, therefore,
that the mode of development in M. marginalba is characteristic of all species
of Morula.

Nassarius particeps

In several species of Nassarius, the females spawn bottle-shaped capsules
containing numerous small eggs which hatch as pelagic planktotrophic veligers

D. T. ANDERSON 249

(Lebour, 1937 ; Thorson, 1946; Natarajan, 1957; Scheltema, 1961, 1962). A
similar mode of spawning and development is also described by Amio (1957)
for Tritia festivus. In a number of Indo-Pacific nassariids, however, the spawn
comprises numerous small triangular capsules with confluent base plates, each
containing a single egg. It is to this group that NV. particeps belongs. In N.
costata and N. thersites, the capsules are stalkless, the eggs are relatively small
(200 u and 250yu in diameter respectively), and hatching occurs as pelagic
veligers which probably adopt a planktotrophic existence (Natarajan, 1957).
In NV. liviscens, the capsules have short stalks, the eggs are larger (320 u in
diameter), and development is more direct, with hatching occurring as a veliger
in which the velum is relatively small and the visceral mass large and yolky.
Development and metamorphosis to a crawling juvenile are completed in this
Species during a brief pelagic lecithotrophic phase (Amio, 1957). In WNW.
suturalis the capsules have long stalks, the eggs are large (probably about 400 p
in diameter), and development appears to be direct, with hatching occurring
as a: crawling juvenile (Risbec, 1935). WN. particeps, with its long-stalked
capsules and very large eggs, shows the most extreme adaptation to direct
development yet known for nassariids with this type of spawn.

Siphonaria denticulata

The egg mass and development of S. denticulata, hatching as a small
planktotrophic veliger, are similar to those of several species of intertidal
siphonariid limpet (Morton, 1955 ; Knox, 1955; Voss, 1959). In Kerguelenella
stewartiana and Siphonaria kurracheensis the egg capsules contain a greater
volume of albumen and the embryos hatch as crawling juveniles in a more
typical pulmonate manner (Thorson, 1940; Knox, 1955), but K. stewartiana
is a Sub-Antarctic littoral species from Stewart Island and S. kurracheensis is
a semiterrestrial species occurring in the Persian Gulf.

CONCLUSIONS

Studies of prosobranch reproduction in the northern hemisphere have
Shown that planktonic larvae are of common occurrence among temperate
species (Thorson, 1946, 1950). Along the New South Wales coast, many more
Species need to be studied before any corresponding generalization can be made
for the 52 species of littoral prosobranch listed by Dakin (1953) as common
to this vicinity. At the same time, the species whose development has been
recorded are now sufficient in number to warrant grouping into the developmental
types distinguished by Thorson (1946, 1950), as follows :—

(a) Viviparous species—none yet described.

(b) Species with a non-pelagic development—5. Bembicium melanostoma
(Littorinidae), Glossaulax aulacoglossa (Naticidae), Cymatilesta spengleri (Cyma-
tidae), Bedeva hanleyi (Muricidae), Nassarius particeps (Nassariidae) (Anderson,
1959, this paper; H. Anderson, 1958; Murray 1962), 1964).

(ec) Species with a very short pelagic life (a few hours to a few days)—5.
Cellana tramoserica (Patellidae), Notoacmaea petterdi, Patelloida alticostata,
Chiazacmaea flammea (Acmaeidae), Melanerita melanotragus (Neritidae) (Anderson,
1962, 1965).

(d) Species with a long pelagic, planktotrophic veliger life—11. Bembiciwm
nanum, Bembicium auratum, Melaraphe unifasciata, Nodilittorina pyramidalis
(Littorinidae), Velacumantus australis (Potamididae), Conuber conicum, Conuber
stranget (= sordida), Conuber melastoma (Naticidae), Cypraea caput-serpentis
(Cypraeidae), Xenogalea labiata (Cassididae), Morula marginalba (Thaididae)
(Anderson, 1961, 1962, this paper; MacIntyre, 1961; Murray, 19625, 1964 ;
Ostergaard, 1950).

Thus 11 out of 21 species whose developmental type is known almost
certainly have a long planktonic larval life, a proportion which accords with

250 LIFE HISTORIES OF LITTORAL GASTROPODS

expectations based on northern hemisphere studies. Further investigation,
however, may yet alter this picture, since of the remaining 31 common species,
many are archaeogastropods, unlikely to have a long planktonic life, and several
others belong to families for which non-pelagic development is characteristic.

ACKNOWLEDGEMENTS

It is a pleasure to acknowledge my debt to Miss I. Bennett, who generously
provided part of the material used in this study, to Dr. D. F. McMichael for
advice on matters of taxonomy, and to Dr. G. Thorson for invaluable help in
tracing relevant literature. The work was supported by a research grant from
the University of Sydney.

References

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livescens (Phillipi). J. Shimonoseki Coll. Fish., 6: 271-80.

ANDERSON, D. T., 1959.—The reproduction and early life history of the gastropod Cymatilesta
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, 1960.—The life histories of marine prosobranch molluscs. J. Malacol. Soc. Aust.,
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, 1961.—The reproduction and early life history of the gastropod Bembicium nanum
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, 1962.—The reproduction and early life histories of the gastropods Bembicium auratum
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and Melanerita melanotragus (Smith) (Fam. Neritidae). Proc. Linn. Soc. N.S.W., 87:
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, 1965.—The reproduction and early life histories of the gastropods, Notoacmaea petterdt
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(Fam. Acmaeidae). Proc. Linn. Soc. N.S.W., 90: 106-114.

ANDERSON, H., 1958.—The gastropod genus Bembicium Phillipi. Aust. J. Mar. Freshw. Res.,
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ANKEL, W. E., 1937.—Der feiner Bau des Kokons der Purpurschnecke Nucella lapillus (L.)
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, 1938.—Die Laichkapsel der Purpurschnecke und die Entwicklung ihrer Jungtiere.
Aus der Natur, 15 Jahrg., Nov. 1938, Heft 8: 269-74.

BURKENROAD, M. D., 1931.—Notes on the Louisiana conch, Thais haemastoma. Ecology, 12:
656-64.

Butter, P. A., 1953.—The Southern oyster drill. Proc. Nat. Shellfish Ass., 44: 65-75.

Cuari, V. K., 1950.—Breeding habits of Thais bufo (Lamarck). J. Bombay Nat. Hist. Soc.,
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Daxtn, W. J., 1953.—“‘ Australian Sea Shores”. Sydney, Angus and Robertson.

DEHNEL, P. A., 1955.—Rate of growth of gastropods as a function of latitude. Physiol. Zool.,
28: 115-44.

ERLANGER, R. von, 1893.—Beitrage zur Kenntnis des Baues und der Entwicklung einiger
marinen Prosobranchier. Zool. Anz., 16: 1-6.

Go.tikov, A. N., 1961.—Ecology of reproduction and the nature of the egg capsules of some
gastropods of the genus Neptunea (Bolten). Zool. Zh. 40: 997-1008.

Heptery, C., 1916.—Studies on Australian Mollusca. Part XIII. Proc. Linn. Soc. N.S.W.,
41: 680-719.

Knox, G. A., 1955.—The development of Kerguelenella stewartiana (Powell) (Gastropoda,
Siphonariidae). Pacific Sci., 9: 85-9.

Lesour, M. V., 1937.—The eggs and larvae of British prosobranchs, with special reference to
those living in the plankton. Journ. mar. biol. Ass. U.K., 22: 105-66.

Lo Branco, §S., 1899.—Notizie biologiche riguardanti specialmente il periodo di maturita
sessuale degli animali del golfo de Napoli. Mutt. zool. Sta. Neapel., 13: 448-573.

MacIntyre, R. J., 1961.—Larval development of Velacumantus australis. Proc. Linn. Soc.
N.S.W., 86: 183-4.

MacKenziz, Jr., C. L., 1961.—Growth and reproduction in the oyster drill Hupleura caudata
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Morton, J. E., 1955.—The evolution of the Ellobiidae, with a discussion of the origin of the
Pulmonata. Proc. zool. Soc. Lond., 125: 127-68.

Murray, F. V., 1962a.—Gelatinous masses spawned by some naticids (Sand-snails). Avust. J.
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, 1962b.—Notes on the spawn and early life history of two species of Conuber Finlay
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, 1963.—Some observations on the egg capsules and embryos of Torvamurex territus
(Reeve 1845) with field notes by Margaret H. Goldsmith. J. Malacol. Soc. Aust., 7: 21-5.

, 1964.—The spawn of some Australian marine prosobranch molluscs. Aust. Nat.
Hist., 14: 405-8.

D. T. ANDERSON 251.

NaTaraJaAn, A. V., 1957.—Studies on the egg masses and larval development of some proso-
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ScHELTEMA, R. S., 1961.—Metamorphosis of the veliger larvae of Nassarius obsoletus (Gastropoda)
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, 1962.—Pelagic larvae of New England intertidal gastropods. 1. Nassarius obsoletus
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, 1940.—Studies on the egg masses and larval development of gastropods from the
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EXPLANATION OF PLATE X

The animal and part of the egg mass of Xenogalea labiata.

OBSERVATIONS ON THE FINE STRUCTURE OF THE MERISTEM
OF ROOT NODULES FROM SOME ANNUAL LEGUMES

P. J. DART* and F. V. MERCERT
(Plates xi-xxv)

[Read 29th September, 1965}

Synopsis
The fine structure of the meristematic zone of the root nodules of subterranean clover,
barrel medic and purple vetch was examined with thin sections of KMnO, and OsO, fixed tissue.
The nodule meristematic cell has basically the same ultra-structure as other types of meri-
stematic cells, as described in the literature. The differentiation of the cells produced by the
meristem to form the cells invaded by the rhizobia is also described. The fine structure of the
nodule husk cells is compared with those of the nodule meristem.

INTRODUCTION

The root nodules of subterranean clover and barrel medic and purple vetch
are formed by the differentiation of cells produced by an apical meristem.
The cells which differentiate basally in relation to the zone of cell division
form the region of the nodule which becomes filled with bacteroids, while cells
differentiating terminally and laterally form the husk or nodule cortex, and
in this region further differentiation forms the vascular system of the nodule.
Development of cells in the bacteroid zone of the nodule has been described
(Dart and Mercer, 1963a, 1963) ; 1964).

MATERIALS AND METHODS

Plants of subterranean clover (Trifolium subterraneum lL. var. Clare)
inoculated with the effective Rhizobium trifolti str. TAI, and barrel medic
(Medicago tribuloides Desr. str. 173) inoculated with the Rhizobium meliloti
strain SU277.1, an effective strain, or SU237, were grown in sand culture in
a greenhouse. This latter strain forms nodules which are red for only 3-5
days. A description of these nodule types has been given previously (Dart
and Pate, 1959). Nodules from Vicia atropurpurea Desf. (purple vetch) formed
by the effective Rhizobium strain SU331 were also examined. For the effective
strains, slices of 1-4 week old nodules were examined ; but for the SU237 strain,
slices were taken from nodules both before they became pigmented and during
the pigmented phase. The nodule slices were fixed in KMnO, or OsO,, stained
in uranium acetate, and embedded in araldite. Thin sections were examined
in a Siemens Elmiskop I or II.

Full details of techniques have been described previously (Dart and Mercer,
1963).

OBSERVATIONS AND DISCUSSION
The low power electron micrographs (Pls xi; xii, a; xiii) show the general
fine structural features of the nodule meristematic cells. There is a relatively
large nucleus usually containing one nucleolus, mitochondria, proplastids,
Golgi bodies, endoplasmic reticulum occasionally connected with the nuclear
membrane, a ground cytoplasm with many ribosome-like particles, and
occasionally ‘‘ spherosomes’”’? and unidentified vesicular organelles. As can

* Linnean Macleay Fellow of the Society.
+ Macquarie University, Ryde, N.S.W., and the Plant Physiology Unit, School of Biological
Sciences, University of Sydney and C.S.I.R.O., Division of Food Preservation, Ryde, N.S.W.

PROCEEDINGS OF THE LINNEAN Society or New SoutH Wass, Vol. 90, Part

P. J. DART AND F. V. MERCER 253

be seen, the basic structure is similar to the ultrastructure of other types of
meristematic cells (e.g. root apex Whaley et alii, 1960; Falk, 1962; and stem
apex Buvat, 1958). This confirms the suggestion made from light microscope
observations that ‘‘ the meristematic cell of the nodule corresponds to a
meristematic cell of any growing region’’ (Fred, Baldwin and McCoy, 1932).
Intercellular spaces are not usually present in the meristem, but develop as
the cells differentiate (Pl. xi).

Mitochondria

In KMn0O, fixation, the mitochondria are mostly spherical to ellipsoid in
shape, and have the usual structure with cristae arising from the inner limiting
membrane and a homogeneous matrix between the cristae. Mitochondria
from meristematic and newly differentiated cells usually contain a prominent
‘vacuole’ in this matrix, superficially similar to the Rhizobium nucleoid, and
it is tempting to associate this region with mitochondrial deoxyribonucleic
acid (e.g. see Nass and Nass, 1963; Nass et alti, 1965; Bell and Miuhlethaler,
1964 ; Gibor and Granick, 1964). Occasionally some mitochondria have figure-
of-eight shapes, a narrow constriction separating the two lobes—suggesting
division by constriction (Pl. xiv, b; Pl. xxiii, a). Similar origins for new
mitochondria have been proposed by other workers (e.g. Whaley et alii, 1960).
No large promitochondrion bodies which segment to form mitochondria (Bal
and De, 1961; Manton, 1962; Vesk, Mercer and Possingham, 1965) were found
in the nodule meristem. Mitochondria in the differentiated uninvaded cells
usually are smaller with fewer cristae than those of the meristem.

Proplastids

These organelles are prominent features of the nodule meristematic cell,
and are scattered through the cytoplasm. They are usually bounded by two
membranes which enclose a poorly developed internal membrane system and
a matrix or stroma. The limiting membrane generally stains more deeply
in KMnO, fixation than any of the other cell membranes. The internal
membranes are formed by invagination of the inner limiting-membrane (PI.
xvii, ¢; Pl. xviii, a-c) or from a ‘“‘ prolamellar body ”’.

In leaf tissue a prolamellar body has been implicated in the formation of
the internal plastid membranes (e.g. Muhlethaler and Frey-Wyssling, 1959).
In the nodule plastids the ‘ bodies ’ are much smaller and the membrane bounded
compartments less organized than those found in leaf tissue (Pl. xviii, a-c).

In the proplastids of the nodule, small (~ 60 A diameter), electron-dense
granules are found. These are distributed through the stroma of the proplastids
before, and at the beginning of, starch formation (Pl. xvi, a; Pl. xviii, e) but
in young plastids with small starch grains they may be arranged in groups
(Pl. xv, b; Pl. xvii, a, b). The particles resemble the phytoferritin described
by Hyde et ala (1963) as occurring in young bean and pea plant plastids.
Bergersen (1963) has also observed small electron-dense granules in soybean
nodule plastids. The particles are much more readily resolved in OsO, fixation
(Pl. xv, 6; Pl. xvi, a) for in KMnO, fixation the electron-density of the plastid
stroma tends to mask the granules. In older plastids with large starch grains
the particles can occasionally be found. The particles are sometimes organized
into a tight array from which tubular (?) profiles appear to originate (Pl. xvii,
a, b). Similar ‘tubules’ often run between the normal, internal plastid
membranes and the inner limiting membrane of the plastid (Pl. xvii, a, d, e;
Pls xix ; xx; xxi b) but are much less electron dense than the normal plastid
membranes. Often there are several of these profiles running roughly parallel
to each other, superficially resembling a mitochondrion in outline, although
the tubule diameter is much smaller than the profile of a mitochondrion crista
in cross section. Serial sections show that the tubules, with the enclosing
dense-staining plastid membranes are often organized into a separate, oval-

254 FINE STRUCTURE OF THE MERISTEM

shaped ‘‘ compartment ”’ at the edge of the plastid. The membrane bounding
the ‘‘ compartment ’”’ in the plastid stroma is not continuous in all sections
with the plastid limiting membrane which forms the rest of the ‘‘ compart-
ment” boundary. The “ compartment ’’ would appear to be formed by an
invagination and folding back of the inner-limiting membrane of the plastid
(RI Sippxcisxas = eexexe) -

The proplastids in the differentiating cells adjacent to the meristem often
have bud-like protrusions (Pls xiv, ¢; xv, d; xvii, e), usually with several
membranes running across the ‘ bud ”’.

The nodule proplastids appear to arise in two related ways—by segmenta-
tion of a large membrane bounded body (Pl. xiv, a, b) and by constriction
division of an existing proplastid (Pls xili; xxiv, b). A similar situation has
been described by Vesk, Mercer and Possingham (1965) for the proplastids of
leaves of Zea mays. Plastids in the nodule meristematic cells rarely contain
starch and have few internal membranes, but synthesis and development of
these accompanies cell enlargement and vacuolation (Pls xi, a; xii, b).

In the bacteroid-filled cells plastids become elongated and filled with
elongate starch grains (Dart and Mercer, 1963a, 1964) while in the adjacent
noninvaded cells the plastids are oval-shaped, and contain three or four large
roughly circular, starch grains, but much more plastid stroma remains than
in the plastids of the invaded cells (Pl. xvii, ¢).

As with the mitochondria, proplastids in the nodule meristem often contain,
in the stroma, small electron empty ‘ vacuoles’ crossed by very fine fibrils
(Pl. xiv, a). The similarity between these areas and the bacterial nucleoid has
been remarked on by others (e.g. Ris and Plaut, 1962), the implication being
that these are the deoxyribonucleic acid-containing regions in the plastid (see
Gibor and Granick, 1964; Gunning, 1965).

Golgi Bodies

Several Golgi bodies are usually present in each thin section of the
meristematic cell and each consists of a varying number of flattened discs.
Occasionally in KMnO, fixation, the membrane bounding each dise can be
resolved into a unit membrane (Robertson, 1960). Often the rims of the dises
are enlarged into vesicular structures which form a sequence ranging from a
slightly inflated periphery to large sacs, some of which apparently bud off to
form small single membrane bounded vesicles in the cytoplasm, as described
for other cells (Whaley et alii, 1960, 1964). As in other plant cells (e.g. Whaley
and Mcllenhauer, 1963), these Golgi vesicles appear to be associated with cell
plate and primary wall formation.

Ground Cytoplasm

In KMn0O, fixation the ground substance is a homogeneous granular matrix,
while in OsO, fixation electron-dense, 120-150 A diameter ribosome-like
particles are found, dispersed through the cytoplasm and associated with the
endoplasmic reticulum (Pls xii, a@; xiii; xv, a; xvi, a). Surface views of the
reticulum (PI. xiii) show the ribosomes may be organized in a spiral or a linear
array with some 8-12 ribosomes per unit (polyribosome ?). The relative
concentration of ribosomes in the cytoplasm decreases as the cells differentiate
and vacuolation begins. A marked increase in ribosome numbers (mostly
free in the cytoplasm) occurs following infection thread invasion and subsequent
release and dispersal of the Rhizobium cells through the host cytoplasm. The
endoplasmic reticulum system is rather sparse in the meristematic cells and
is mostly plate-like and generally granular. The amount of endoplasmic
reticulum appears to be even less in differentiating cells (Pl. xii, b) before
Rhizobium invasion. Small spherosome-like bodies (see Frey-Wyssling et alii,
1963 ; Drawert and Mix, 1962) are sometimes found in the nodule meristematic
cells but are more frequently seen in the differentiating cells. These bodies

P. J. DART AND F. V. MERCER 255

have an electron-dense granular composition, and are surrounded by a dense
staining membrane. Sometimes an electron-empty region is found in the
centre of these bodies (Pl. xviii, e). Similar bodies have been found in maize
and rye root meristem cells (Whaley et alii, 1960; Fabergé and Lewis, 1962).
Occasionally other unidentified inclusions resembling a sack of tiny vesicles
are found (Pl. xxii) in both KMnO, and OsO, fixation, as well as small vesicles
with a single membrane enclosing a homogeneous ground substance (Pl. xxii, a).
The former are closely associated with the endoplasmic reticulum and similar
bodies are found next to the cell wall enclosed by the plasma membrane (PI.
xxil, b, d). This suggests that the bodies may be involved in transport of
materials from the endoplasmic reticulum to the cell wall. Jensen (1963) has
recently reported that similar bodies in cotton synergids are specialized
endoplasmic reticulum vesicles. Occasionally membrane profiles of what is
presumably endoplasmic reticulum contain a dense inclusion (Pl. xxii, a).

Nucleus

The interphase nucleus occupies a major proportion of the volume of the
nodule meristematic cell. In KMn0O, fixation the nucleus is clearly bounded
by two unit membranes that are usually closely appressed to each other and
OsO, fixation shows that the outer membrane is studded with ribosomes.
Nuclear pores and larger gaps are occasionally present and there are often
several connections between the nuclear envelope and endoplasmic reticulum
(Pl. xi).

In early interphase cells, gaps much wider than the normal nuclear pore
are often present in the nuclear envelope. Similar large gaps occur in newly
invaded, differentiated nodule cells. In both these cell types ribosome synthesis
is active at this stage as well as the synthesis of new cytoplasmic proteins.
Woodard et ali (1961) have shown that in pea root meristem cells there is a
rapid synthesis of ribonucleic acid in early interphase. The large gaps in the
nuclear envelope would permit rapid transfer of ribosomes to the cytoplasm
if in fact ribonucleic acid and protein are organized into ribosomes in the
nucleolus (see Bonner and Huang, 1962). Little structure is observable in
the interphase nucleoplasm with KMnO, fixation but occasionally denser
eranulation, presumably corresponding to chromatin material, can be seen,
while the nucleolus appears as a more electron-dense area usually circular in
outline. In OsO, fixation followed by uranyl acetate staining considerable
structure can be seen in the nucleus. Several areas of dense staining materials
with an overall granular appearance are present. These dense, presumably
euchromatin areas have an irregular outline and are bounded by an electron-
lucent nucleoplasm containing dispersed fibrillar material (Pls xii, a; xv, 4a).

The nucleolus is clearly defined in OsO, fixation with uranium acetate post
staining. It is basically more electron-dense than the surrounding nucleoplasm
and chromatin. Occasionally the nucleolus is haloed by an area free of
electron-dense material (Pl. xii, a) but, as Lafontaine (1958) observed, there
are often places where chromatin and nucleolus merge together (Pl. xv, a).
The nucleolus itself contains tightly packed 130-150 A diameter granules.
The nucleolus often has a relatively electron-lucent core and in the granular
cortex there often appears to be filamentous material ~ 100 A wide.

Cell Plate Formation in the Meristem

Following cell division the nuclear membrane often has large gaps near
the region of cell plate formation. The new cell plate begins as a collection
of vesicular material lined by endoplasmic reticulum and phragmosomes,
between the two telophase nuclei. Vesicle coalescence, and consolidation of
the electron-lucent material within the new vesicle aggregates, mark the
beginning of wall synthesis which continues centripetally (Pl. xi, b). In the
initial stages of new wall synthesis there are several intercellular endoplasmic

256 FINE STRUCTURE OF THE MERISTEM

reticulum connections. The pattern of cell division parallels closely that
outlined by Porter and Caulfield (1958) and Porter and Machado (1960) for
onion root-tip cells, and Whaley et ala (1960) for maize root meristematic cells.
Towards the end of cell plate formation, evenly dispersed electron-dense material
is deposited at the centre of the new wall. As in cell plate formation this zone
(middle lamella?) then consolidates and expands centripetally.

Differentiation of the nodule meristematic cell

After division has ceased the newly-formed cells undergo differentiation to
form the husk and bacteria-filled zone of the nodule. The continuous
differentiation of the cells and the continuous infection of the differentiated
cells maintain a zone, generally three cells wide, between the meristem and
the zone of infection.

Vacuolation of the cytoplasm is the first change, associated with differ-
entiation of the nodule cells. Various theories have been proposed for the
origin of vacuoles in plant cells, and in electron micrographs of the nodule
meristem, profiles of vacuoles can be found which are consistent with most of
these views. The most usual *‘ method” of vacuole formation observed in
the nodule cells was that described by Miihlethaler (1958) where a phase difference
becomes apparent in the cytoplasm (PI. xxiii, a) and as this region expands a
tonoplast is synthesized de novo at the interface in several parts before joining
to form the tonoplast observed around fully-developed vacuoles. Buvat (1957,
1958, 1960) and Poux (1962a and b) proposed that vacuoles are initiated by
expansion of the two membranes of the endoplasmic reticulum. Membrane
profiles consistent with this can be found in differentiating nodule cells (Pls
Xxii, @; xxill, ¢) but these may well be plasmolysis figures of vacuoles rather
than stages in vacuole formation. Small vesicles are sometimes found associated
with the plasmalemma suggesting that pinocytosis might be occurring. Weiling
(1961) has suggested that subsequent expansion of pinocytotic vesicles forms a
vacuole. In some meristematic cells there are small irregular-shaped, membrane-
bounded bodies with phase differences characteristic of vacuoles (Pl. xi). These
are presumably the “‘ pro-vacuoles ”’ that Whaley et alii (1962) and Leech et alia
(1963) suggest are transformed into true vacuoles. Marinos (1963) claims that
the tonoplast in barley shoots is derived from a swelling of the outer Golgi
body cisterna but no profiles suggestive of this have been found in the
nodule meristem.

The vacuoles in the nodule usually contain a sparsely distributed, electron-
dense material. Occasionally dense granular bodies are found in the vacuole
and sometimes sharply defined differences in electron-density (phase difference?)
exist within the vacuole.

The tonoplast is not always preserved after KMnO, fixation, but better
preservation is obtained with OsO, fixation. The tonoplast can be resolved
into a unit membrane structure (as defined by Robertson, 1960) with a dark-
light-dark profile of overall dimension 90-100 A. Occasionally after OsO,
fixation electron-dense material adheres to the vacuole side of the tonoplast.

Some of the cells which differentiate adjacent to the meristem remain
uninvaded by Rhizobium cells. A proportion of these uninvaded cells, which
usually have a thin layer of peripheral cytoplasm intact, degenerate just before
adjacent cells become infected. This involves a loss of the cytoplasmic matrix
and disorganization of the usual organelles, leaving the plasmalemma and most
of the tonoplast intact. In these cells the membranes are readily resolved
(Pl. xvi, b)—possibly due to a lack of background cytoplasm obscuring the
structure, but could also conceivably be due to a change in the membrane
structure itself, induced during the cell degeneration. Occasionally small
electron-dense lines cross between the two dense lines of the tonoplast membrane
giving the membrane a banded appearance similar to the ‘ globular’ structure
observed in mitochondrial and some cytoplasmic membranes by Sjéstrand

P. J. DART AND F. V. MERCER 257

(1963). These ‘degenerate’ uninvaded cells are thought to be a defence
mechanism response of the host cell to restrict invasion by Rhizobium. Alter-
natively, these non-living cells may be functioning as vascular or conducting
tissue as has been postulated for degenerate cells in pea cotyledon tissue (Bain
and Mercer, 1965).

In meristematic cells, but more frequently in the differentiated, recently
invaded cells, the plasmalemma often invaginates, enclosing a system of tightly
coiled membrane-bounded tubules and vesicles (Pl. xxi, 6). These structures
resemble the lomasomes observed in fungi (Girbardt, 1961 ; Moore and McAlear,
1961 ; Peyton and Bowen, 1963). Invaginations of the plasmalemma are also
found, with only a few membrane-bounded vesicles between the plasmalemma
and the cell wall similar to the structures observed by Grun (1963) in Solanum
root meristem cells and by us in barrel medic and subterranean clover root
meristem cells (Dart and Mercer, unpublished observations). In some cells a
single membrane fragment is sometimes found immediately outside the
plasmalemma in the cell wall material (Pl. xvi, b) and occasionally membranous
elements are found deeper in the wall layers (Pl. xxi, b). These membrane
fragments may be remains from the deposition of material during cell wail
thickening (Wardrop, 1964). An incorporation of small, single membrane-
bounded vesicles, with the vesicle membrane fusing with the plasma membrane,
also appears to be involved in wall development (Pl. xxii, b). In other places
the vesicles themselves appear to be incorporated in the wall (Pl. xvi, 5).

Plasmodesmata are frequently observed between meristematic cells,
becoming less so as the cell differentiates with associated cell wall growth.
Some of these plasmodesmata are branched (Pl. xviii, e) and in some the
plasmalemma is observed to evaginate and line the structure so that the
plasmalemmas of adjacent cells are contiguous. Some plasmodesmata-like
structures which penetrate the cell wall are completely bounded in the wall
by a membrane-like structure. These might also be Frey-Wysslings “ wall
papillae ” (1962). Only an outer dense zone with an adjacent electron-empty
zone can be resolved, presumably because the inner dense line of the membrane
(assuming it is a unit membrane) merges with the electron-dense material
enclosed by the ‘‘membrane” (Pl. xxi, a).

Nodule Husk Cells

Quite a distinct difference is apparent between the cells of the nodule
meristem and the large, vacuolated, ‘‘ protective ’’ cells that enclose the nodule.
The husk cells have a very large central vacuole, and a thin layer of cytoplasm
containing a few small mitochondria with few cristae, Golgi bodies and segments
of endoplasmic reticulum (Pls xxiv; xxv). Plastids are few in number and
large in size. Starch formation increases with distance from the meristem.
The vacuoles usually contain more stainable material than the vacuoles of cells
about to be invaded by infection threads (compare Pl. xxv and PI. xii, 5).

The nucleus lies in the thin layer of cytoplasm adjacent to the cell wall
and often has a wrinkled appearance (Pl. xxv, c). Plasmodesmata connections
between the husk cells are prevalent—and usually occur in groups (Pl. xxv),
possibly corresponding to apit field. Some of the husk cells lose their cyto-
plasmic contents, leaving a granular material attached to the cell wall in places.

CONCLUSION
There are no basic differences in fine structure between the nodule
meristems of subterranean clover, barrel medic or purple vetch. It can be
seen that the ultrastructure of the meristematic cell is very similar to the basic
ultrastructure of the root meristematic cell. It seems that meristems have a
similar subcellular organization and pattern of activity whether they are
“normal” structures or whether they arise as a response to invasion by

Rhizobium.

B

258 FINE STRUCTURE OF THE MERISTEM

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EXPLANATION OF PLATES XI-xXxXV

Plate xi.

a. Panorama of the meristematic zone of a barrel medic nodule (SU237). Vacuole formation
and cell enlargement has commenced in some cells. The single arrow points to a ‘ provacuolar
body ’ in a meristematic cell which appears to have a ‘ tail ’ of endoplasmic reticulum. The
double arrow points to an ‘unknown body’ which is bounded by a single membrane.
n—nucleus, v—vacuole. KMnO, fixation. x 11,000.

b. Cell plate formation in a meristematic cell of a barrel medic nodule (SU237). Endoplasmic
reticulum and small vesicles (Golgi vesicles ?) are closely associated with the new wall.
The Golgi bodies (g) appear to be budding off small vesicles. The double arrow points to a
major discontinuity in the wall. The small mitochondria have prominent ‘ vacuoles’
(e.g. arrow). p—phragmosome (?). KMn0O, fixation. x 35,000.

Plate xii.

a. Meristematic cell of a 7-day-old barrel medic nodule (SU237). The nucleus contains sparsely
distributed fibrillar material but around the dense nucleolus (Nu) there is a ‘ halo’ virtually
free of fibrils. In the ground cytoplasm there are free ribosomes and some ribosomes
attached to endoplasmic reticulum. The proplastids (p) have shrunk during preparation.
OsO, fixation. x 12,500.

b. Panorama showing the vacuolating cells adjacent to the meristem in a barrel medic nodule
(SU237). Most of the plastids contain starch grains. A few sparsely distributed, plate-like
endoplasmic profiles are present. KMnO, fixation. x 11,000.

260 FINE STRUCTURE OF THE MERISTEM

Plate xiii.

Portion of a meristematic cell from a barrel medic nodule (SU237) showing the distribution of
ribosome-like particles, endoplasmic reticulum, Golgi bodies and mitochondria. Surface
views of the endoplasmic reticulum show the ribosomes often grouped (polyribosomes ?)
into whorled, rosette arrangements (e.g. arrows). The free ribosomes are also grouped
in units (e.g. circles). The plastid (p) at the bottom of the figure appears to be dividing
by constriction. The inset shows some of the ribosome-like particles in more detail : some
of them are attached to endoplasmic reticulum. OsO, fixation. x 20,000. Inset x 40,000.

Plate xiv.

a. Large proplastid bodies in a newly-invaded cell. One appears to be dividing by constriction
(arrow). ‘The proplastids contain small electron empty regions (e.g. double arrow) con-
taining fine fibrils which are reminiscent of a bacterial nucleoid (n). 35,000.

b. Irregularly-shaped proplastids are apparently segmenting (see double arrow) in a non-
invaded meristematic cell. One of the mitochondria has a figure-of-eight profile suggestive
of division by constriction (single arrow). w—cell wall. KMnO, fixation, barrel medic
nodules (SU237). x 20,000.

c. Shows a small ‘bud’ on a plastid from a barrel medic nodule (SU277.1). 40,000.

Plate xv.

a. Interphase nucleus and part of the cytoplasm of a differentiating cell adjacent to the
meristem of a barrel medic nodule (SU277.1). The nucleus contains a prominent nucleolus
(nu) and several smaller more diffuse electron dense areas (c—presumably euchromatin).
OsO, fixation. x 20,000.

b. Plastid from a vacuolated, non-invaded cell of a subterranean clover nodule fixed in OsQO,.
The plastid contains a large, central starch grain and several closely packed arrays of
phytoferritin-like particles (e.g. arrow). In the adjacent cytoplasm several ribosome-like
particles (r) are present, along with a Golgi body (g). x 50,000.

Plate xvi.

a. Plastids from a meristematic cell of a subterranean clover nodule fixed in OsO,. The
plastids contain numerous phytoferritin-like particles and several larger, osmiophilic
bodies (0). Ribosome-like particles (r) are mostly free in the cytoplasm. x 60,000.

b. Degenerate non-invaded cell in a barrel medic nodule (SU277.1). The tonoplast (t) and
plasmalemma (pl) are resolved into a dense-light-dense profile. An invagination of the
plasmalemma (arrow) contains several circular membrane profiles apparently embedded
in the cell wall and possibly the remains of vesicular packages of material incorporated
in the wall. The double arrow indicates another membrane profile running parallel to
the plasmalemma and between it and the cell wall. KMnO, fixation. x 70,000.

Plate xvii.

a. Proplastid filled with phytoferritin-like particles. These are aggregated in one portion of
the plastid (single arrow). The double arrow points to three small tubular elements attached
to the limiting plastid membrane. x 60,000.

b. Another aggregation of the small, electron-dense, particles with some fine tubules running
between the aggregation and the plastid limiting membrane. x 35,000.

c. Plastid from a non-invaded cell. The plastid containing four large starch grains and two
areas where an internal plastid membrane is joined to the limiting membrane by fine tubules
(arrows). In Fig. d another plastid has been sectioned closer to the edge and shows several
of the tubules running between internal plastid membranes and the limiting membrane.
7a-d KMnO, fixation, barrel medic nodules (SU277.1). 7c and d. x 40,000.

Plate xviii.

a. and b. are serial sections of a plastid from a vacuolating barrel medic (SU237) nodule cell.
An array of small membrane-bounded compartments in the plastid stroma resembles a
‘prolamellar body’. In Fig. ¢ a similar body (arrow) can be seen with a well-developed
internal plastid membrane attached. The double arrow indicates the junction of an internal
plastid membrane with the peripheral membrane. The internal membrane changes from
a plate-like form to a tubule at the junction.

d. Shows a‘ bud’ on a plastid from a barrel medic (SU277.1) nodule (division by constriction ?).
Small, electron-dense, phytoferritin-like particles are present in the plastid stroma.

P. J. DART AND F. VY. MERCER 261

e. Shows a large proplastid from a vacuolated non-invaded nodule cell. The plastid stroma
contains numerous phytoferritin-like particles and the ‘ bud’ on the right contains numerous
circular profiles—apparently small, membrane-bounded tubules cut in cross section. A
branched plasmodesmata is shown in more detail in the inset. The plasmalemmas of
adjacent cells are contiguous and line the plasmodesmata. s—spherosome-like body.
x 50,000. Inset x130,000. Figs a-d x 40,000. KMnO, fixation, barrel medic nodules.
(d—SU277.1; a, b, c, e, f—SU237).

Plates xix—xx.

Plates xix and xx are serial sections (in sequence) of parts of two plastids from a recently invaded
cell in a purple vetch nodule.

Plate xix, f is oriented about 90° to Plate xix, figs a-e. The figures illustrate an arrangement
(arrows) of small tubules and plastid membranes. Plate xx, 7 is the same section as Plate
xx, f showing the location of the tubules within the plastid. Adjacent mitochondria (m)
show that the membranes of the plastid inclusion have a different appearance from the
mitochondria cristae (e.g. arrow). KMnO, fixation. Pl. xix, a—h, Pl. xx, a-h x 40,000;
Pl. xx, 7 x 20,000.

Plate xxi.

a. Shows numerous plasmodesmata-like fragments in the cell wall of an uninvaded, vacuolated
cell of a subterranean clover nodule. One of the fragments (arrow) is apparently completely
bounded by a membrane—presumably the plasmalemma. KMn0O, fixation. x 60,000.

b. Recently-invaded cell in a barrel medic nodule (SU237). The plasmalemma invaginates
to enclose a lomasome-like body (1) containing a tightly coiled system of membranes.
Membrane envelope synthesis is almost completed around an adjacent Rhizobium cell.
The inset shows the lomasome-like body at higher magnification. KMnO, fixation.
x 40,000. Inset x 100,000.

c. Portion of three differentiating cells adjacent to the meristem. Narrow tubular elements
are present in the cell wall adjacent in the middle lamella region and adjacent to a small
intercellular space. The arrow indicates where a cristae of a mitochondrion has been
sectioned tangentially showing the circular plate-like profile of the cristae. The endoplasmic
reticulum is closely associated with the cell wall (double arrow). Barrel medic nodule.
KMn0O, fixation. x 60,000.

Plate xxii

a. Newly invaded cell and two adjacent uninvaded cells in a barrel medic nodule (SU237)
with their intercellular space filled with an electron-dense material. The double arrow
points to a profile which could be interpreted as the origin of a vacuole by expansion of
endoplasmic reticulum. The single arrow points to bodies containing small vesicles and
at (i) a similar body appears to be fused to the cell wall. A dense body (c) is apparently
enclosed by endoplasmic reticulum. Another unidentified inclusion is present (b), and it
consists of a single enclosing membrane and a homogeneous matrix. A similar body is
indicated by the double arrow in Fig. a. Yet another unidentified organelle (u) is present
in this cell. KMn0O, fixation. x 40,000.

b. In 6} two of the bodies containing small vesicles (arrows) lie close to the cell wall and at (i) one
has fused with the wall. The double arrow indicates a small bulge of the cell wall partly
bordered by similar material to that enclosed by the arrowed bodies suggesting that this
may be a later stage of incorporation of wall material to that at (i). Three similar bodies
are adjacent to the cell wall (i) in Fig. d. Im Fig. 6 endoplasmic reticulum profiles and
small single-membrane-bounded vesicles are present close to the wall in much greater
“concentration ’ than in the rest of the cytoplasm suggesting that they also may have a
role in cell wall development.

c. Portion of a meristematic cell from a barrel medic nodule showing the unidentified bodies
(ub) with the single limitmg membrane enclosing a number of small vesicles. Similar
bodies can also be found after OsO, fixation. g—Golgi cisternae. a—d, KMnO, fixation.
6 x 25,000; c¢ x40,000; d x 50,000.

Plate xxiii.

a. A ‘phase difference’ (v) is apparent in the cytoplasm of a cell from a barrel medic nodule
meristem (SU237). It is suggested that this is the first stage in vacuole formation. The
arrow indicates a mitochondrion with a figure-of-eight profile suggestive of division by
constriction. x 30,000.

6. A cytoplasmic bridle crosses the vacuole of a differentiating cell in a barrel medic nodule
(SU237). The bridle contains a mitochondrion and some endoplasmic reticulum (er).
KMnO, fixation. x 20,000.

262

C.

FINE STRUCTURE OF THE MERISTEM

Profile of a vacuole (v) in a subterranean clover nodule with a constricted region where the
tonoplast resembles an endoplasmic reticulum profile. The arrow points to other
membrane-bounded elements which may be vacuole or expanded endoplasmic reticulum.
KMn0O, fixation. x 25,000.

Plate xxiv.

Young husk cells from a barrel medic nodule (SU237) showing that they contain a similar
complement of organelles to the uninvaded, vacuolated cells basal to the meristem. The
plastids are relatively large and mitochondria small. The cell wall in places has conspicuous
blebs (arrows) which may be the site of incorporation of new wall material. At the top
left of the figure an oblique cut through the wall shows several plasmodesmata in cross
section (e.g. circle). KMnO, fixation. x 10,000.

Plastid from a barrel medic nodule (SU277.1), uninvaded cell, with two small starch grains
and two narrow constrictions suggestive of division. KMn0O, fixation. x 40,000.

Plate xxv.

The cell wall between two husk cells is crossed by several large plasmodesmata. In an
adjacent cell (d) the vacuole has collapsed and the cell is degenerating. The arrow points
to a coiled membrane fragment. Subterranean clover nodule. Xx 12,800.

Shows the thin layer of cytoplasm in some husk cells from a barrel medic nodule (SU237).
The cell walls are crossed by several plasmodesmata. KMnO, fixation. x 10,000.

Cortex region of a barrel medic nodule (SU237), showing the nucleus (n) and several mito-
chondria closely appressed to the cell wall. Plasmodesmata are conspicuously grouped
in the cell wall. x 12,800.

CERIOID STRINGOPHYLLIDAE (TETRACORALLA) FROM DEVONIAN
STRATA IN THE MUDGEE DISTRICT, NEW SOUTH WALES

A. J. T. WRIGHT

Iinnean Macleay Fellow of the Society in Palaeontology
Department of Geology and Geophysics, University of Sydney

(Plate xxvi)

[Read 29th September, 1965]

Synopsis
Melrosia rosae, gen. et sp. nov. and Melasmaphyllum mullamuddiensis, gen. et sp. nov. are

described from near Mudgee. This is the first record of cerioid members of the Stringophyllidae
and the relationships and stratigraphic significance of the occurrences are discussed.

STRATIGRAPHIC INTRODUCTION

Study of the previously poorly-known Devonian tetracoral faunas of an
area near Mudgee, New South Wales, has revealed that two cerioid species
belonging to the Stringophyllidae are developed at two different stratigraphic
horizons. From their respective associated faunas it appears that the occurrence
of Melasmaphyllum gen. nov. is stratigraphically lower than that of Melrosia
gen. nov.

Melasmaphyllum is known, as yet, only from its type locality within the
Sutchers Creek Formation!, the youngest Devonian beds in the Queens Pinch
area which is located about 12 miles south-east of Mudgee. Other tetracorals
identified from this formation include Pachyphyllum auct., Dendrostella,
Dohmophyllum, Tryplasma and Pseudamplexus, and brachiopods found in shales
interbedded with the limestones of the formation include ? Dolerorthis,
Phragmophora s.s., Leptostrophia and Adolfia. Tryplasma and Pseudamplexus
were thought by Hill (1957, p. 43 and p. 49) to be absent from beds younger
than Emsian ; also, Williams (1953, p. 40) considered Leptostrophia to be absent
from beds younger than Lower Devonian, and Dolerorthis is not yet recorded
from rocks younger than early Devonian (Boucot, 1960; Philip, 1962). On the
other hand Pachyphyllum, Dendrostella, Dohmophyllum, Phragmophora and, to
a lesser degree, Adolfia are suggestive of a Middle Devonian age by comparison
with known extra-Australian stratigraphical ranges. The age of the assemblage
is thus uncertain.

Comparisons of the fauna from the Sutchers Creek Formation with faunas
described from elsewhere in Eastern Australia are rather inconclusive. The
tetracoral assemblage is very similar to that from the Sulcor Limestone in the
presence of Tipheophyllum, Xystriphylium, Phillipsastraea s.1., Trapezophyllum
and Pseudamplexus (Hill, 19426). It could be suggested on the basis of Brown’s
(1942) correlation of the Sulcor and Loomberah Limestones and Pedder’s
(1964, p. 437) “‘ Hifelian or Givetian ”’ age for the Loomberah that the Sutchers
Creek Formation is Middle Devonian. The faunas from the Buchan district
(Hill, 1950; Talent, 1956) have no obvious similarities with faunas of the
Sutchers Creek Formation although the latter appears to be younger than the
fauna from the Kilgower Member of the Wentworth Group described by Talent
(1963) as Emsian.

1 The new stratigraphic names introduced herein will be formally defined and fully described
in a future publication.

PROCEEDINGS OF THE LINNEAN Soctety oF NEw Sours Watss, Vol. 90, Part 3

264 DEVONIAN CERIOID STRINGOPHYLLIDAE

Melrosia is known at present from only the type locality where it is plentiful
in limestones exposed in the core of an anticline near ‘‘ Melrose’ homestead,
about five miles east-south-east of Mudgee. These limestones are correlated
with well-bedded limestones of the Mount Frome Limestone! on the south-west
slopes of Mount Frome to the north of ‘‘ Melrose’. The presence of Endophyllum,
stringophylliids and Pachyphyllum auct. at ‘“ Melrose”? and Mount Frome
suggests a Middle Devonian age by comparison with overseas ranges.

All type material has been deposited at the Department of Geology and
Geophysics, University of Sydney (S.U.G.D.). No prefix is given for the
numbers of fossil specimens housed there, but in explanations to text-figures
the letter W precedes numbers referring to thin sections upon which figures
are based. Grid references refer to the Dubbo 1: 250,000 topographic sheet.

The author wishes to acknowledge all assistance given during this work,
especially from Professor Dorothy Hill who drew attention to the similarities
between Melasmaphyllum and Xitphelasma, and from Professor Hill and Dr.
B. D. Webby who criticized the manuscript.

SYSTEMATIC PALAEONTOLOGY

Phylum COELENTERATA

Class ANTHOZOA

Order TETRACORALLA

Family STRINGOPHYLLIDAE Wedekind, 1922.

This family is distinguished principally by lonsdaleoid dissepiments,?
concave or flat tabular floors, and septa consisting of stout, contiguous or
separate, monacanthine trabeculae. The only compound forms previously
described have been fasciculate species for the reception of which Birenheide
(1962) erected Stringophyllum (Soctophyllum), which Pedder (1964, p. 444)
considered to be a distinct genus. Solitary species have been placed in a
number of genera of which only Stringophyllum Wedekind, 1922, and
Neospongophyllum Wedekind, 1922, were recognized by Engel and Schouppé
(1958) and Birenheide (1962), the latter considering Neospongophyllum a
subgenus of the type genus; the two forms are principally distinguished by
the prominent lonsdaleoid dissepimentarium in Neospongophyllum. Vollbrecht-
ophyllum Taylor, 1951 (pro Schizophyllum Wedekind, 1925, non Verhoeff, 1896)
was held by Taylor to be distinct from Stringophyllum in having peripherally
discontinuous major septa. Any such attempts to differentiate a third group
in addition to Stringophyllum and Neospongophyllum must be regarded with
suspicion until specific variation of these taxa is adequately documented.
Loipophyllum Wedekind, 1925, and Vollbrechtophyllum were treated as junior
synonyms of Neospongophyllum by Engel and Schouppé (op. cit.).

Some of the species assigned to Grypophyllum Wedekind, 1922, including
the type species, can be likened to Stringophyllum in that their minor
septa are often short, aborted or even discontinuous (Middleton, 1959, p. 143
et seq.). It is quite possible that when lineages of species become evident
some of these species at present placed in Grypophyllum will prove to be
genetically related to Stringophyllum and some to Acanthophyllum Dybowski,
1873; Birenheide (1961, p. 128) has already suggested the latter, and Engel
and Schouppé (1958) were so impressed by the similarities between Grypophyllum
and Stringophyllum that they considered them as type genera of subfamilies

2 McLaren (in McLaren and Norris, 1964, p. 5) advocated the use of the term ‘“ wand-
blasen ”’ to replace the term “lonsdaleoid dissepiments”’. There does not seem to be any
fundamental difference in the mode of secretion of lonsdaleoid dissepiments and post-septal
dissepiments, as McLaren suggests. Therefore the only advantage of McLaren’s usage is the
ease of distinction between dissepiments preceding both major and minor septa and those preced-
ing only the minor septa. The change in terminology is no more explanatory of the sequence of
structural elements than the system recommended by Hill (1935) and has not been adopted herein.

A. J. T. WRIGHT 265

of the Stringophyllidae. The tabulae developed in Stringophyllum indicate a
close relationship with Acanthophyllum, whereas the flat or weakly concave
tabulae often seen in Neospongophyllum and Sociophyllum suggest a different
origin, possibly from a Spongophyllum-like ancestor. Melrosia appears to be
closely related to Xystriphyllum Hill, 1939, possibly most closely to X. dunstani
(Etheridge, 1911) which commonly has lonsdaleoid dissepiments. Melasma-
phyllum is similar to Spongophyllum Edwards and Haime, 1851, but possesses
axially discrete trabeculae ; among possible ancestors of Melasmaphyllum are
many of the species ascribed to Spongophyllum, as well as Neomphyma
pseudofritscht Soshkina.

Hill (1957, p. 43) lists as “the earliest Stringophyllum, S. carnicum
(Charlesworth, 1914)’, from probably lower Devonian strata in the Carnic
Alps. Wang (1948, p. 18, and 1950, p. 215), Engel and Schouppé (1958, p. 96)
and Spassky (1964, table 16, p. 104) also considered that stringophyllids
appeared in the Lower Devonian. The only apparent evidence given by Wang
and Engel and Schouppé concerns Australia, so this age determination was
probably based on misconstrued estimates of the age of the Moore Creek
Limestone (Hill, 1942b) and some of the North Queensland limestones (Hill,
1942a); Hill considered these probably Middle Devonian. So there appears
to be little evidence to suggest that stringophyllids first appeared in the Lower
Devonian.

The only report of cerioid Stringophyllidae appears doubtful; Smith
(1945, p. 55) likened a Canadian cerioid form to Spongophyllum semiseptatum ;
but the Canadian form is probably not a stringophyllid, as Birenheide (1962),
p. 65) has remarked. Other comparable cerioid tetracorals include Spongo-
phyllum, Donia Soshkina, 1951, and Hndophyllum abditum Edwards and Haime,
1851; all possess peripherally discontinuous septa but none exhibits discrete
trabeculae in the axial region. Apart from the axially discontinuous septa,
Melasmaphyllum and Melrosia are identical with Spongophyllum and Xystriphyllum
respectively. Donia was considered by Soshkina (1951) and Pedder (1964) to
be a disphyllid and this opinion seems justified by the nature and arrangement
of the dissepiments as well as the septal structure. H. abditum is quite distinct
in having domed tabular floors. Kozlowiaphyllum Rhukin, 1938 (see Soshkina,
1962, p. 335) is a Silurian cerioid form possibly related to Spongophyllum ;
further information is needed to clarify this genus. The apparent nature of
the septa of Australophyllum praeclarum Crickmay (1962, p. 6, Pl. I], figs 4-5)
suggests that it may well belong in the Stringophyllidae, despite the fact that
the septa are not discontinuous axially ; this form may be close to Melasma-
phyllum.

Genus MELROSIA, novum
Type Species: Melrosia rosae, sp. nov.

Diagnosis: Cerioid tetracorals with septa consisting of monacanthine
trabeculae which may be discrete axially. Major septa long; minor septa
are almost completely suppressed. Lonsdaleoid dissepiments weakly developed
peripherally. Tabular floors concave, often with an axial depression in the
close-set tabulae.

Remarks: This genus, at present monospecific, differs from Melasma-
phyllum, gen. nov., in general septal, dissepimental and tabular features. Apart
from the cerioid habit it is very close to the normal Stringophyllum style in
having long, axially discontinuous septa with characteristic bilateral symmetry
about the plane containing the cardinal and counter septa.

Derwation of Names: The genus takes its name of feminine gender from
the property ‘‘ Melrose ”’, on which it occurs ; the species is named for my wife
whose encouragement has been invaluable.

266 DEVONIAN CERIOID STRINGOPHYLLIDAE

MELROSIA ROSAE, sp. nov.
Pl. xxvi, fig. 1; Text-figs 1-3

Diagnosis: Melrosia with 25 to 29 major septa at an average diameter
of 9mm.; two to five rows of steeply inclined dissepiments are present, the
outer ones often being lonsdaleoid ; tabularium from one-half to three-fifths of

corallite diameter.

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Text-figs 1-3. Melrosia rosae, gen. et sp. nov.; la. Transverse section of holotype 21104,
W1128, x2-4. 16. Longitudinal section of one corallite from holotype 21104, W1129, x2-4.
2a. Transverse section of paratype 21105, showing lateral bud, W1121, x2-4. 26. Longitudinal
section of paratype 21105, W1122, x1-6. 3. Transverse calical section of paratype 21106,

W1147, «2-4.

Description: Coralla are apparently hemispheroidal, and growth form
cerioid ; corallites are generally irregularly hexagonal and up to 11 mm. in
diagonal diameter. Mode of increase is uncertain, but apparently peripheral ;
buds mature gradually, having an early stage where dissepiments are few and

septa short (Text-fig. 2a).

A. J. T. WRIGHT 267

From 26 to 31, but generally about 28, major septa extend almost to the
axis, where they are occasionally represented by isolated monacanths ; minor
septa extend to about half their length, and are seldom continuous, generally
appearing as crests on the dissepiments or more rarely as discrete monacanths.
Peripherally major septa are strongly dilated, forming a stereozone about
0-5 mm. thick ; dilation is much less in minor septa, but lessens in both orders
towards the axis and is weak in the tabularium. Bilateral symmetry is
moderately developed about the plane containing the two short protosepta
(Pl. xxvi, fig. 1). Trabeculae are moderately inclined both peripherally,
diverging slightly towards the axis, and are about 0-2 mm. in diameter.

Dissepiments occur as two to five rows of steeply inclined, flattened plates,
a few of which function as lonsdaleoid dissepiments (first order ‘‘ wandblasen ”’) ;
commonly some act as bases (second order ‘‘ wandblasen’”’) of minor septa
which appear as stout, isolated trabeculae between the much more continuous
major septa.

Tabulae may be complete or incomplete ; tabular floors may be regularly
and moderately concave or may form broad, gently axially-inclined rims with
a broad, deep axial depression ; tabularium occupies from one-half to three-
fifths of corallite diameter with steeply- or gently-inclined peripheral areas
abutting sharply on, or conforming broadly with slope of, dissepiments.

Remarks: The discontinuous nature of the septa indicates that the
affinities of this species are with Stringophyllum rather than Xystriphyllum.

The species Xystriphyllum dunstani and Spongophyllum cyathophylloides
Etheridge, 1911 (see Hill, 1939) from Clermont, Queensland, may well be
phylogenetically connected with this species although neither possesses axially
discontinuous septa.

Material: Holotype, 21104 (W1128-9), Pl. xxvi, fig. 1, Text-figs la-d ;
paratype, 21105 (W1121-2), Text-figs 2a-b; paratype, 21106 (W1147-8),
Text-fig. 3; Other material: 21107 (W464—5); 21108 (W1149-50); 21109
(W1140) ; 21110 (W469-70) ; 21111 (W460-1) ; 21112-21117, 21118 (W466-7)
and 21119 (W462-3).

Type Locality: S.U.G.D. locality number Mu/IV/60, grid reference
26279658 ; immediately north of quarry, north-west of ‘‘ Melrose”? home-
stead ; in portion 54, parish of Bumberra, county of Philip.

Typical Formation: Mount Frome Limestone.

Genus MELASMAPHYLLUM, novum
Type Species: Melasmaphyllum mullamuddiensis, sp. nov.

Diagnosis: Cerioid tetracorals with very large, prominent lonsdaleoid
dissepiments. Septa consist of monacanthine trabeculae which are often
separated axially ; minor septa mostly limited to short ridges just emerging
from septal stereozone. Tabulae generally flat, rarely concave.

Remarks: This genus is distinct from Melrosia, gen. nov., and can in some
respects be considered a cerioid counterpart of Neospongophyllum. It appears
at present to be monospecific.

Xiphelasma Smith and Lang, 1931, is rather similar to Melasmaphyllum ;
Hill (1956a, F312) treated Xiphelasma as a junior subjective synonym of
Storthygophyllum Weissermel, 1894, which is ‘like Tryplasma but cerioid and
with a narrow zone of dissepiments”’ (Hill, loc. cit.). Storthygophyllum
(= Xiphelasma) differs from Melasmaphyllum in possessing rhabdacanthine
trabeculae as well as in detailed nature of septa, tabulae and dissepiments.

Two Russian species assigned to Neomphyma Soshkina, 1937, are similar
to Melasmaphyllum ; the Gedinnian NV. pseudofritscht Soshkina (1962, p. 335,
Pl. 19, fig. 1) and the Ludlovian NV. rosiformis Zheltonogova (1961, p. 81, Pl.

268 DEVONIAN CERIOID STRINGOPHYLLIDAE

S20, figs 3a—b) are structurally similar to Melasmaphyllum, as is Spongophyllum
originalis Kraevskaya (1955, p. 214, Pl. 41, figs 3a—b) but all three differ from
Melasmaphyllum in not possessing discrete trabeculae axially. The same
applies to some cerioid members of Spongophyllum with large lonsdaleoid
dissepiments. Melasmaphyllum may be genetically related to these species.

Text-figs 4-6. Melasmaphyllum mullamuddiensis, gen. et sp. nov. Transverse sections of
holotype, 21103, all x 2-4. 4. Proximal part of corallum, W999. 5. Median part of corallum,
W869. 6. Distal part of corallum, W1179.

Derivation of Name: The name of the genus is partly based on the Greek
word (neuter gender) for ‘“‘ black spot”; the species is named after nearby
Mullamuddy Creek.

269

A. J. T. WRIGHT

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Melasmaphyllum mullamuddiensis, gen. et sp. nov.

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Longitudinal sections

xt-figs 4-10
major septa long, occasionally with weak
Up to six daughter corallites

9a, b. Distal part of corallum, W836.
septa represented by only short trabecular crests.

Pl xxvi, ties 2 Te
Melasmaphyllum with from 19 to 24 major septa strongly

MELASMAPHYLLUM MULLAMUDDIENSIS, Sp. Nov.

8. Distal part of corallum, W870.

Text-figs 7-9.
of holotype, 21103, illustrating in places mode of increase. All x 2-4. 7. Proximal part of corallum,

W1000.

Diagnosis :
dilated peripherally and generally discontinuous axially and with an average

corallite diameter of about 6 mm. ;

bilateral symmetry ; minor
Lonsdaleoid dissepiments long and wide, generally in one row. Tabulae

complete and distant, flat or weakly concave.

270 DEVONIAN CERIOID STRINGOPHYLLIDAE

parricidally produced by axial increase ; peripheral increase rare. Tabularium
from one-third to three-fifths of corallite diameter.

Description : Corallum hemispheroidal and apparently originally reached
30cm. in diameter. Corallites are mostly polygonal; in distal parts of
corallum there may be irregular subtriangular spaces between walls where
three corallites approach each other, and more rarely with narrow spaces
between weakly curved walls; maximum diagonal diameter is about 8 mm.,
average diameter in longitudinal section is 5 to 6 mm. ; corallites have four to
seven sides, generally five or six walls which are mostly straight.

From 19 to 24 major septa are sometimes wholly developed but generally
are strongly interrupted peripherally by lonsdaleoid dissepiments and where
developed axially are represented only by large monacanths. Minor septa are

Text-fig. 10. Melasmaphyllum mullamuddiensis, gen. et sp. nov. Serial transverse sections
of 21103B/8, illustrating budding, x4. Distance of sections above a. measured in millimetres :
b. 0-8; c. 1-1; d. 1-3; e. 1:7; f. 2:3; g. 3-2; h. 3-9; 2. 5-6; 7. 7-5. Where the limits of
drawings are regular, this represents epitheca.

very strongly suppressed and may be completely absent; peripherally they
are sometimes distinct as low ridges between larger major septa, and where
developed elsewhere are seen only as isolated monacanthine crests on dissepi-
ments. Septa are mostly straight and dilated peripherally to form a thick
stereozone up to 0-5 mm. thick, but on the average coat a thin epitheca to a
thickness of 0-2mm. Bilateral symmetry is weakly developed; cardinal
septum not distinct. Despite poor preservation monacanthine trabeculae
about 0-3 mm. in diameter can be seen diverging within the plane of a septum
so aS to be separated along the axial edge; peripherally the inclination of
trabeculae is generally at about 45° to a horizontal plane, but occasionally
subhorizontal ; towards the axis they may be almost vertical.

In mature corallites, the tabularium varies in width from one-third to
three-fifths of the total diameter. Tabulae are generally complete, and are

A. J. T. WRIGHT 271.

mostly flat or may be occasionally weakly convex, weakly concave, or delicately
scalloped ; from 16 to 22 irregularly to regularly spaced tabulae are spaced
in a length of 10mm. Usually there is one series of long, steeply-inclined
lonsdaleoid dissepiments which may extend about half-way around the corallite
and up to one-third of the maximum diameter.

Increase is almost invariably axial and parricidal, only one possible instance
of peripheral increase having been observed (Text-fig. 5). No interruption is
seen in the vertical continuity of tabular structures as daughters are not
produced high on calical walls, above the space formerly occupied by the parent
polyp. In increase a complex thickening develops on a tabula and eventually
extends by means of furcation to the periphery, producing up to six daughters
which attain adult characteristics in a length of about 6mm.; this outward
spread may be irregular with the axial plate and one or more septa being fused
while the plate is still only a thickening or may proceed more or less symme-
trically (Text-fig. 10). This undulating tabular thickening is at first a simple
non-planar sheath but eventually it extends distally along ridges in its surface.
Apparently from the proximal edges of the apices of ridges in this thickening
a thin median plate analogous to normal epitheca develops and persists distally
throughout all daughters.

Remarks: The validity of the erection of a new taxon on a single specimen
may well be queried. The monotype consists of a large colony, probably of
several hundred corallites exhibiting no more than a reasonable amount of
phenotypic variation. Despite the absence of further material, which thorough
collecting has failed to yield, there is no reason to consider this colony a
pathological variant or an end member of a variable species ; in the associated
fauna of compound tetracorals none of the species of Hexagonaria, Phillipsastraea
s.l., Pachyphyllum s.l., Trapezophyllum, and Xystriphyllum can be considered
similar to Melasmaphyllum. It seems reasonable to conclude that this corallum
was a successful if bizarre mutation living in association with the other distinct
colonial forms.

Spongophyllum rosiforme Yoh, 1937, is similar to Spongophyllum elongatum
Schliiter, 1880, the type species of Stringophyllum (Sociophyllum) Birenheide,
1962. Although Birenheide (1962, p. 72) refers S. rosiforme to Spongophyllum,
axially isolated trabeculae which it exhibits indicate stringophyllid affinities.
The Middle Devonian species from Kwangsi is very similar to WM. mullamuddiensis
apart from phaceloid habit and regularly concave tabulae.

The multipartite parricidal axial increase seems to be close to that seen
in Hexagonaria quadrigemina (Goldfuss): Smith (1945, p. 46, Pl. 14, figs
5a—b) but the segregating structure extends in that species from the ends of
the major septa to the axis rather than from the axis to the walls as in M.
mullamuddiensis.

Material: Only the holotype corallum 21103 is known; weathered into
three pieces, two have been sectioned as follows whereas 21103A remains
whole :—21103B: Text-fig. 10, based on serial cellulose peels of 21103B/8 ;
W999 (Text-fig. 4); W1000 (Text-fig. 7); 21103C: W835 (Pl. xxvi, fig. 2) ;
W836 (Text-figs 9a—b) ; W869 (Text-fig. 5) ; W870 (Text-fig. 8) ; W1178 ; W1179
(Text-fig. 6); W1180.

Type Locality: S.U.G.D. locality number Mu/IV/38, grid reference
26379535 ; in portion 152, parish of Broombee, county of Wellington.

Typical Formation : Sutchers Creek Formation.

References
BIRENHEIDE, R., 1961.—Die Acanthophyllum—Arten (Rugosa) aus dem Richtschnitt Schonecken
—Dingdorf und aus anderen Vorkommen in der Hifel. Senck. leth., 42: 77-146, Pls 1-7.

, 1962.—Revision der kolonielbildenden Spongophyllidae und Strimgophyllidae aus
dem Devon. IJbid., 43: 41-99, Pls 7-13.

272 DEVONIAN CERIOID STRINGOPHYLLIDAE

Bovcot, A. J., 1960.—Lower Gedinnian Brachiopods of Belgium. Mem. geol. Inst. Louvain,
21: 283-324, Pls 10-18.

Brown, I. A., 1942.—The Tamworth Series (Lower and Middle Devonian) near Attunga, N.S.W.
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Crickmay, C. H., 1962.—‘‘ New Devonian Fossils from Western Canada’’. Published privately,
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ENGEL, G., and ScHourprk, A. von, 1958.—Morphogenetisch—taxonomische Studie zu der
devonischen Korallengruppe Stringophyllum, Neospongophyllum und Grypophyllum. Palaént.
Z., 32: 67-114, Pls 8-9.

ETHERIDGE, R., Jr., 1911.—The Lower Palaeozoic corals of Chillagoe and Clermont, Part 1.
Pub. geol. Surv. Qd., 231: 3-8, Pls A-D.

Hitz, D., 1935.—British terminology for rugose corals. Geol. Mag., 72: 481-510.

———,, 1939.—The Middle Devonian rugose corals of Queensland, I. Douglas Creek and
Drummond Creek, Clermont District. Proc. roy. Soc. Qd., 50: 55-65, Pls 4-5.

, 1942a.—The Middle Devonian rugose corals of Queensland, III. Burdekin Downs,
Fanning R., and Reid Gap, North Queensland. Jbid., 53: 229-268, Pls 5-11.

, 1942b6.—The Devonian rugose corals of the Tamworth District, N.S.W. J. Proc.
roy. Soc. N.S.W., 76: 142-164, Pls 2-4.

, 1950.—Middle Devonian corals from the Buchan District, Victoria. Proc. roy. Soc.
Vict., 62: 137-162, Pls 5-9.

, 1956.—Rugosa in “ Treatise on Invertebrate Paleontology’, Part F (ed. Moore,
R. C.) F234-324. Univ. Kansas Press, Lawrence.

, 1957.—The sequence and distribution of Upper Palaeozoic coral faunas. Aust. J.
Sct., 19: 42-61.

Krarvskava, L. N., 1955.—Devonian Rugosa (pars) in “ Atlas of important fossil animals
and plants of Siberia’. Siberian geol. Dept., 1: 206-218, Pls 32-42. Moscow (in Russian).

McLaren, D. J., and Norris, A. W., 1964.—Fauna of the Devonian Horn Plateau Formation,
District of Mackenzie. Bull. geol. Surv. Can., 114: 1-74, Pls 1-17.

Mippieton, G. V., 1959.—Devonian tetracorals from south Devonshire, England. J. Paleont.,
33: 138-160, Pl. 27.

PEppeER, A. HK. H., 1964.—Correlation of the Canadian Middle Devonian Hume and Nahanni
formations by tetracorals. Palaeontology, 7: 430-451, Pls 62-73.

Puitre, G. M., 1962.—The palaeontology and stratigraphy of the Siluro-Devonian sediments
of the Tyers Area, Gippsland, Victoria. Proc. roy. Soc. Vict., 75: 123-246, Pls 11-36.

ScHLUTER, C., 1880.—Versammlung des naturhistorischen Vereins fur Rheinland und Westfalen
in Bonn am 3. Oct., 1880, 147-8 (not seen).

SmitH, S., 1945.—Upper Devonian corals of the Mackenzie River Region, Canada. Spec. Pap.
geol. Soc. Amer., 59: 1-126, Pls 1-35.

, and Lane, W. D., 1931.—Silurian corals—The genera Xiphelasma gen. nov. and
Acervularia, Schweigger, with special reference to T'ubiporites tubulatus, Schlotheim and
Diplophyllum caespitosum, Hall. Ann. Mag. nat. Hist., (10), 8: 83-94, Pls 2-3.

SosHxina, E. D., 1937—Upper Devonian Tetracorals of the Urals. Trav. Inst. paleozool. Acad.
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, 1951.—Upper Devonian Tetracorals, their systematics and evolution. Trav. Inst.
paleont. Acad. Sct. U.R.S.S., 34: 1-122, Pls 1-24 (in Russian).

, 1962.—Rugosa (pars) in “ Osnovi Palaontologii’’ (ed. Sokolov, B.S.), 286-345,
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(in Russian).

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, 1963.—The Devonian of the Mitchell and Wentworth Rivers. Mem. geol. Surv.
Vict., 24: 1-118, Pls 1-78.

Taytor, P. W., 1951.—The Devonian Tetracorals of the Plymouth Limestone. Trans. roy.
geol. Soc. Cornwall, 18: 161-214, Pls 1-5.

Wana, H. C., 1948.—The Middle Devonian rugose corals of Eastern Yunnan. Contr. geol. Inst.,
Univ. Peking, 33: 1-45, Pls 1-5.

, 1950.—A revision of the Zoantharia Rugosa in the light of their minute skeletal
structures. Phil. Trans. roy. Soc. Lond., 234, B: 175-246, Pls 4-9.

WEDEKIND, R., 1922.—Zur Kenntnis der Stringophyllen des oberen Mitteldevon. S. B. Ges.
ges. Naturw. Marburg, 1: 1-16.

, 1925.—Das Mitteldevon der WHifel. Hine biostratigraphische Studie. II. Teil.
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A. J. T. WRIGHT 273

You, 8. S., 1937.—Die Korallenfauna des Mitteldevons aus der Provinz Kwangsi, Stidchina.
Palaeontographica, 87, A: 45-75, Pls 4-9.

ZHELTONOGOVA, V. A., 1960.—Devonian Rugosa (pars) in “ Palaeozoic Biostratigraphy of the
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D57 (in Russian).

EXPLANATION OF PLATE XXVI.

Fig. 1. Melrosia rosae, gen. et sp. nov. Transverse section of holotype. 21104, W1128, »~ 4.

Fig. 2. Melasmaphyllum mullamuddiensis, gen. et sp. nov. Transverse section of holotype.
21103, W835, x 6.

AN EMBRYOLOGICAL STUDY OF FIVE SPECIES OF
BASSIA ALL. (CHENOPODIACEAE)

GWENNETH J. HINDMARSH
Department of Botany, The University of New England

[Read 27th October, 1965]

Synopsis
Development of male and female gametophytes and embryogeny of Bassia bicornis, B.
brachyptera, B. dwaricata, B. paradoxa and B. patenticuspis is described.

Plate crystals of calctum oxalate were found in the perianth and ovary wall of all species,
and anther filaments of some species.

The anther, which is tetrasporangiate, becomes four to five layered due to irregular divisions
in the tapetum. Both amoeboid and secretory tapetum types occurred. Cytokinesis of the
microspores is simultaneous and the mature pollen grain is three-celled.

The ovary contains a basal, campylotropous, bitegmic ovule which is crassinucellate. The
hypodermal archesporial cell cuts off a parietal cell and the megaspore mother cell undergoes
normal megasporogenesis. The chalazal megaspore of a linear tetrad develops into the mono-
sporic eight-nucleate embryo sac of the Polygonum type.

Embryogeny conforms to the Chenopodiad type and the mature embryo is elongate and
spiral. The endosperm is at first nuclear, but later becomes cellular and is digested by the
developing embryo until only a cap remains over the tip of the radicle. In the seed the food
storage region is the perisperm.

INTRODUCTION

Representatives of the family Chenopodiaceae are native to Australia in a
wide variety of habitats, which include xerophytic and halophytic situations.
In semi-arid areas, which are too dry for grasses, many genera such as Atriplex,
Bassia, Chenopodium, Kochia, and Rhagodia are valuable fodder plants. The
genus Bassia, according to Black (1948), consists of 60 species, of which about
50 are Australian endemics and the remainder occur in Europe and Asia. No
indigenous members of the family have been studied embryologically and,
although some overseas species of genera represented in Australia have been
examined, no previous work of this nature has been carried out on Bassia.

MATERIALS AND METHODS

The material used in this investigation was collected in the field by Mr.
E. Hoult in May, 1963, or from plants grown in the glasshouse from seeds
collected in western New South Wales (Table 1).

Conventional paraffin sections were cut at 9-14 » and stained with Delafield’s
Haematoxylin and Johansen’s Safranin; supplementary examinations were
made by dissections and squashes of fresh and preserved material.

The drawings, unless otherwise indicated, are of B. paradoxa and comparative
studies were made with the other species.

MorRPHOLOGY

Bassia paradoxa is a small shrub with narrow to linear, thick, alternate
leaves which, together with the stems, bear a white woolly indumentum (Fig. 1).
The hairs are multicellular and uniseriate (Figs 2, 3). The flowers, which are
also hairy, are sessile, and 8-20 are united in each dense axillary cluster (Figs
4, 9), although according to Black (1948) clusters of 6-10 flowers are usual.
Bisulputra (1960) has shown that the cluster is morphologically a condensed

PROCEEDINGS OF THE LINNEAN SocrETY oF NEw SoutH Watss, Vol. 90, Part 3

GWENNETH J. HINDMARSH 275

dichasium. In the remaining species examined, the flowers occurred singly in
the leaf axils due to the suppression of the lateral buds of the dichasium
(Bisulputra, 1960). The development of the hairs, the perianth tube and the
spines is variable (Figs 8-14) but is specifically distinct (Black, 1948).

The ovary is monocarpellary and unilocular with a solitary basal
campylotropous ovule (Fig. 5). The elongated style terminates in two elongated
stigmatic branches, although in some cases it is trifid due to the presence of

TABLE |
Tist of species and locations from which material was collected

Species Location Collected from
glasshouse
B. paradoxa (R. Br.) 31m. W. Broken Hill May °63—July °64
K. v, M.
B. bicornis (Lindl.) 60m. E. Quilpie, Q’ld. May “63—Nov. 63
F. v. M.
B. brachyptera (F. v. M.) 40m. KE. Broken Hill =
R. H. Anderson
B. divaricata (RB. Br.) 70m. N. Broken Hill ee
F. v. M
B. patenticuspis — Nov. *64—Feb. ’65

R. H. Anderson

a smaller third branch. Its surface is finely papillose and its open stylar canal
communicates directly with the loculus (Fig. 6). The five stamens are
obdiplostemonous with dorsifixed tetrasporangiate anthers and are at first
enclosed within the perianth tube, but with elongation of the filament after
maturity of the pollen grains they are displayed outside the woolly mat of
hairs surrounding the flowers ee 7). Introrse dehiscence: occurs by means
of longitudinal slits.

TABLE 2
Figures in microns indicate the average length of the larger crystals

Species Top of ovary Base of ovary Perianth Anther
B. paradoxa 20 5 9 —
B. bicornis 25 15 15 15
B. dwaricata 12 4 10 4
B. brachyptera 14 13 11 8
B. patenticuspis 17 20 17 —
Crystals

Plate crystals were present in the perianth and ovary wall of all species
studied (Figs 15, 16) and were identified as calcium oxalate by their solubility
in 2N hydrochloric acid and their insolubility in 20% acetic acid. Although
their shape was constant they varied in length from 2 to 30u, the average
size for the larger ones varying between species and according to their location
within the flowers (Table 2). The larger crystals were solitary and almost
completely filled the cells, while in other instances there were up to four smaller
ones present (Figs 17, 18). In B. bicornis, and to a lesser extent in the other
species, the crystal-containing cells were larger than those lacking them (Figs

276 EMBRYOLOGICAL STUDY OF SPECIES OF BASSIA

14

Figs 1-8. Morphology of B. paradora. 1, 4, 7, 8, Flower clusters of different ages; 2. 3,
Hairs; 5, The gynoecium; 6, Stylar canal.

Figs 9-14. Fruits: 9, B. paradoxa; 10, 11, B. patenticuspis; 12, B. bicornis; 13, B.
divaricata ; 14, B. brachyptera.
Figs 3, 4, 6-8, 11, 12, m L.S.; remainder whole mounts.
(c, crystals; pg, pollen grains; vs, vascular strand.)
Figs 1, 7, x3; 2, 3, x220; 4, 9-14. x7; 5. x30: 6, x130: 8, x13.

GWENNETH J. HINDMARSH 277

17, 19). In the course of floral development, the crystals were first observed
when the ovule was at the megaspore mother stage. They appeared in the
inner hypodermal layer of the perianth lobes and tube, connecting with a broad
erystal-bearing layer at the base of the ovary. Similarly, crystals were deposited
in the upper hypodermal cells of the inner wall of the ovary wall and followed
up the hollow style but with reduced size and frequency. A relationship was
noted in the ovary between the absence of the crystals in the hypodermis and
the radial elongation of the overlying inner epidermal cells (Figs 20, 21), while
cells above and below these areas, which contained crystals in the hypodermis,
were smaller although still glandular in appearance (Fig. 22). By the time of
anthesis, the crystal layers had become thickened through further deposition
of crystals in the second to the fourth sub-hypodermal cell layers where they
persisted into the fruiting stage.

In B. bicornis, B. brachyptera, and B. divaricata a small number of crystals
were also found in the central cells of some anther filaments but their distribution
followed no definite pattern.

THE MICROSPORANGIUM

The undifferentiated anther is at first ovoid in cross section, but becomes
rectangular due to radial expansion resulting from localized cell divisions. The
cells are initially of similar size and non-vacuolate with prominent nuclei, after
which four hypodermal, uniseriate rows differentiate as archesporial cells (Figs
23, 24), and initiate the formation of the four sporangia.

(a) Wall Formation

Each archesporial cell divides periclinally to form an inner primary
sporogenous cell and an outer parietal cell which undergoes anticlinal divisions
and, together with the adjacent cells from the ground tissue, forms the primary
parietal layer, which encloses the sporogenous tissue (Figs 25, 26). Periclinal
division of the primary parietal layer forms the secondary parietal layer and
the outer endothecial layer (Figs 27, 28), whose cells divide only anticlinally.
A periclinal division of the secondary parietal layer then gives rise to a single
middle layer and to the tapetum, which may become irregularly two-layered
due to further periclinal cell divisions (Figs 29-31). At this stage the micro-
sporangium wall consists of four or five layers, the epidermis, the endothecium,
the middle layer, and the tapetum which may be irregularly two-layered. The
development of the wall layers is the same as that outlined diagrammatically
in Themeda australis (Woodland, 1964, Fig. 7).

A four-layered wall is the usual condition in members of the Chenopodiaceae ;
however, in Beta vulgaris (Artschwager, 1947) more than four layers have been
described while, according to Miller, Kline and Weber (1959), in Chenopodium
ambrosioides either one or two middle layers are present.

The pressure exerted by the expanding sporogenous tissue stretches all
the wall layers tangentially and, although some anticlinal divisions occur, these
cease when vacuolation sets in prior to the differentiation of the wall layers.

After the initial stretching of the tapetal cells rapid cytoplasmic synthesis
occurs, and the cells increase in size and become glandular in appearance, with
prominent nuclei and dense cytoplasm. When the adjacent microspore mother
cells enter Prophase I many of the tapetal nuclei undergo an apparently normal
mitotic division, in which the spindles are obliquely orientated within the cells
(Figs 32, 33), and the cells become binucleate (Fig. 34), although fusion may
occur and result in the formation of a single large polyploid nucleus (Fig. 35).

The tapetal cells first show signs of breakdown just prior to the release
of the microspores from the tetrads (Fig. 36). This is indicated by the con-
traction of the protoplasm, which is followed by disorganization of the bounding

278 EMBRYOLOGICAL STUDY OF SPECIES OF BASSIA

Figs 15-22. Calcium oxalate crystals. 15, Distribution of crystals in B. paradoxa; 16,
Distribution of crystals in B. divaricata; 17, L.S. of B. paradoxa perianth; 18, B. patenticuspis,
surface view of perianth crystals; 19-22, B. bicornis: 19, L.S. of perianth; 20, T.S. of flower ;
21, Ovary wall as in Fig. 20; 22, T.S. of wall at top of the ovary.

Figs 23-26. Development of the microsporangium. Figs 23, 25 in L.S.
(a, archesporium; c, crystals; cf, crystals in the anther filament; co, crystals in the ovary
wall; cp, crystals in the perianth ; e, epidermis; ie, inner epidermis of ovary wall; 0, ovary ;
oe, outer epidermis of the ovary wall; , primary parietal layer; s, sporogenous tissue.)
Figs 15, 16, 20, «20; 17, x320; 18, 19, 24-26, «540; 21, 22, x220; 23, x130.

GWENNETH J. HINDMARSH 279

membrane (Fig. 37). At a later stage, when the pollen grains enter the ‘ signet
ring’ configuration, it is usual in B. paradowa for the inner walls of the tapetal
cells to degenerate and so allow the protoplasts to form a continuous mass around
the periphery of the loculus. As a result, the pollen grains come into contact
with, and lie between, the remaining portions of the tapetal cells (Fig. 38). At
this point, the nuclei show signs of degeneration but maintain their identity as
dense areas in the tapetal cytoplasm until it is absorbed. Finally, the tapetum
is represented by small oval globules on the inner walls of the endothecium
(Fig. 39), which disappear before dehiscence (Fig. 44). In B. brachyptera, B.
divaricata, B. patenticuspis, aS well as B. paradoxa, the contents of the tapetal
cells break down and are absorbed in situ; the tapetum conforms to the
secretory or glandular type. In some anthers of B. paradoxa, however, after
dissolution of the cell walls, the protoplasts become amoeboid and move into
the loculus between the recently liberated tapetal microspores, forming a
periplasmodium (Fig. 43). The liberated tapetal nuclei maintained their
identity, and underwent mitotic divisions, indicating that this was a ‘true’

TABLE 3

Types of tapetum in the species of Bassia studied

Amoeboid
Secretory
Periplasmodium No Periplasmodium
B. paradoxa o ab =
B. bicornis — = +
B. brachyptera =e — =
B. dwvaricata + — —
B. patenticuspis + — —

periplasmodium, although in some sections of B. paradoxa the tapetal cells
appeared to become detached and lie between the developing pollen grains,
forming an apparently ‘false’ periplasmodium (Fig. 40). However, on
examination of serial sections it was seen that these cells were still in contact
with the endothecium, and the pollen grains were merely lodged between the
degenerating cells of the glandular tapetum (Fig. 38). An additional variation
in B. bicornis, and of occasional occurrence in B. paradoxa, is the formation
of an amoeboid tapetum which does not form a periplasmodium (Table 3).
The protoplasts, which maintain their contact with the endothecium, become
enlarged and vacuolated, and protrude into the loculus between the developing
pollen grains, where they are absorbed (Fig. 41).

The occurrence of both a secretory and an amoeboid tapetum in the same
species is unusual and, in some instances in B. paradoxa, both types were found
in sporangia of adjacent anthers within the same flower, with a higher proportion
of aborting microspores being found in the loculi with a secretory tapetum
(Figs 42, 43). The only previous record in which both tapetal types occur is
in the male-sterile plants of Beta vulgaris (Artschwager, 1947) where the
periplasmodium is thought to delay pollen abortion, since in those micro-
sporangia with a secretory tapetum the microspores degenerate while still
retained in the tetrads. In fully fertile flowers of the same species Artschwager
(1927) had previously reported only a secretory tapetum. In the present
investigation, about 60% of microspores were degenerating in microsporangia
with a secretory tapetum (Fig. 42), as against only about 10% in those with
an amoeboid tapetum (Fig. 43).

Variation of tapetal behaviour is common within the family and, according
to Mahabale and Solanky (1954a) in Arthrocnemum indicum, although it is of

280 EMBRYOLOGICAL STUDY OF SPECIES OF BASSIA

Figs 27-36. Development of the microsporangium. Figs 27, 30, 33, in L.S., remainder in T.S.
Figs 34, 35, Tapetal cells.

(d, region of dehiscence ; e, epidermis ; en, endothecium; m, middle layer; mme, microspore
mother cell; mt, microspore tetrad; s, sporogenous tissue ; sp, secondary parietal layer; ss,
secondary sporogenous tissue; ¢, tapetum.)

Figs 27-30, x 540; 31, 32, 36, x 310; 34, 35, x 780.

GWENNETH J. HINDMARSH 281

the secretory type, the “walls of the tapetal cells break down and the
protoplasts coalesce to form a continuous mass at the periphery of the pollen
chamber”’ which is similar to that reported in Chenopodium ambrosioides
(Mahabale and Solanky, 1954b), Kochia scoparia (Mahabale and Solanky,
1953b) and Suaeda fruticosa (Mahabale and Solanky, 1953a). In Chenopodium
album (Bhargava, 1936), however, although the tapetum is amoeboid it does
not form a periplasmodium, and a similar condition is indicated by the
illustration of Mahabale and Solanky (1954c¢) in Chenopodium murale, although
the authors state that a periplasmodium forms.

The middle layer shows signs of stretching and of being crushed when
the microspore mother cells are formed (Figs 31, 32), and at the microspore
tetrad no remains are visible (Fig. 36).

The endothecial cells contain many starch grains (Fig. 45) which disappear
after microsporogenesis, and the cells then become vacuolated, followed by the
deposition of dimorphic ‘fibrous’ wall thickenings. The cells from the
connective region to about a third way around each sporangium are thickened
in a scalariform manner (Fig. 46), while those extending to near the region
of dehiscence bear numerous ‘ fibrous’ rods which are united at their bases
and extend out along the tangential wall (Figs 47, 48). The first six or seven
cells on either side of the point of dehiscence do not bear any thickenings and
become highly vacuolated before breaking down completely at dehiscence
(Fig. 40).

The epidermal cells become vacuolated simultaneously with the rounding
off of the microspore mother cells prior to meiosis (Fig. 33), and all except
those external to the ‘non-fibrous’ cells of the endothecium are greatly
stretched tangentially. At dehiscence they form a dead layer of cells over
the endothecium, while those in the region of dehiscence increase in size at
the same time as the thickenings are laid down in the endothecium (Fig. 48).

Introrse dehiscence takes place by a longitudinal slit in each side of the
anther at the junction of the outer wall and the intersporangial septum (Figs
48, 49).

(b) Microsporogenesis and Male Gametogenesis

The primary sporogenous cells undergo two transverse and one vertical
division to form the secondary sporogenous cells which then divide obliquely
to form the microspore mother cells (Fig. 32), which round off and enter meiosis
(Figs 50-52) followed by simultaneous cytokinesis. The occurrence of secondary
spindles is common at telophase II (Figs 53, 54). The microspore tetrads are
tetrahedral and isobilateral (Figs 56, 57), but in B. bicornis some decussate
tetrads were found (Fig. 55). Separation of the microspores is accomplished
by centripetal furrows and they are at first angular when liberated by the
gelatinization of the enclosing wall of the microspore mother cell (Figs 58, 59).
The appearance of a vacuole in the dense cytoplasm represents germination
into the one-nucleate pollen grain or male gametophyte, and as this vacuole
increases in size the nucleus becomes displaced laterally and the pollen grain
assumes the ‘ signet ring’ configuration (Fig. 61).

Cytoplasmic synthesis reduces, and finally obliterates, the vacuole, and
nuclear division is followed by the formation of the small generative cell (Figs
62, 63), which undergoes a further division to form the two male gametes
(Figs 64, 65). It is in this three-celled condition that the pollen grain is shed.
The deposition of exine is first apparent after the microspores are released
from the tetrad and it thickens irregularly as the pollen grain increases in size
to give the granular appearance of the mature polyforate pollen grain (Fig.
41).

EMBRYOLOGICAL STUDY OF SPECIES OF BASSIA

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GWENNETH J. HINDMARSH 283

THE MEGASPORANGIUM
(a) Development of the Ovule

The primordium of the ovule develops at the base of the ovary when the
microsporangium wall is two-layered, and two integumentary primordia appear
as folds at the base of the nucellus simultaneously with the differentiation of
the archesporial cell (Figs 66, 67). The integuments are at first two-layered,
but further cell divisions occur in the inner integument and extend it beyond
the slower growing outer one which has no part in the formation of the micropyle
(Figs 69, 84). This form of integumentary development is commonly found
in this family. A prominent air space was observed in the chalazal region
between the integuments. During the differential growth of the ovule, which
assumes a campylotropous form, the funiculus becomes elongated and results
in the micropylar end of the ovule coming to lie across the base of the funiculus
(Figs 68-70). A single vascular strand of annular and spiral vessels differentiates
in the funiculus and passes to the chalazal region of the ovule (Fig. 84).

(b) Megasporogenesis

A single hypodermal archesporial cell makes its appearance at the apex
of the ovule (Fig. 71) and divides periclinally to form an outer primary parietal
cell and a megaspore mother cell, which is consistent with other records for
this family. Mahabale and Solanky (1954a) quote Billings (1934) as having
reported parietal cell formation in the ovule of Atriplex hymenelytra and, while
this may well be true, it should be pointed out that Billings’ statement referred
to the behaviour of the anther archesporium. The parietal cell divides both
anticlinally and periclinally and, together with similar divisions in the over-
lying nucellar epidermis, forms the massive nucellus of the crassinucellate
ovule (Figs 72-74).

The megaspore mother cell undergoes meiosis accompanied by cytokinesis
and gives rise to a dyad followed by a linear tetrad of megaspores (Figs 73,
74). In the Chenopodiaceae, the chalazal megaspore is invariably functional
and, with vacuolation, increases in size at the expense of the three non-functional
megaspores until it occupies the place which was filled previously by the tetrad
(Figs 75, 76).

(c) Female Gametogenesis

The one-nucleate embryo sac is embedded deeply within the nucellus and,
following nuclear division, passes into the two-nucleate stage in which the
nuclei are separated by a central vacuole (Fig. 77). Both nuclei then divide
simultaneously to form a four-nucleate embryo sac in which the central vacuole
is retained (Fig. 78), and a third post-meiotic mitosis gives rise to an unorganized,
eight-nucleate embryo sac in which small vacuoles separate the nuclei (Fig. 79).
Cytokinesis follows rapidly to form seven cells, of which the central endosperm
cell is binucleate (Fig. 80). In its development, the embryo sac is, therefore,
of the monosporic, Polygonum or ‘ normal’ type which has been reported in
all other species investigated in this family.

Legends to figures on opposite page.

Figs 37-43. Types of tapetum. 37-39, Secretory tapetum; 40, ‘False’ periplasmodium ;
41, B. bicornis—amoeboid tapetum, but no periplasmodium forms ; 42, Secretory tapetum with
degenerating microspores ; 43, Periplasmodium.

Figs 44-49. Maturation of the anther wall and dehiscence ; 45, Endothecial cell; 46, Scalari-
form thickening; 47, Rod thickening.
Figs 50-65. Formation of the microspores and the male gametes.

(d, region of dehiscence ; dm, degenerating microspores; e, epidermis; ee, enlarged epidermal
cells; en, endothecium ; ent, endothecium with thickenings; g, germ pore; gc, generative cell ;
m, microspores; mg, male gametes; p, periplasmodium ; pg, pollen grains; sg, starch grains ;
t, tapetum; in, tapetal nuclei; tr, tapetal remains; vc, vegetative cell.)

Figs 37, x 780; 38, 40-43, «320; 39, x540; 44, x90; 45-47, 50-65, x360; 48, 49, x53.

284 EMBRYOLOGICAL STUDY OF SPECIES OF BASSIA

Figs 66-70. Development of the ovule. Figs 71-81. Megasporogenesis and embryo sac
development.
(a, archesporium ; an, antipodals; as, air space; d, dyad; dm, degenerating megaspores ;
e, epidermis; eg, egg; es, embryo sac; f, funiculus; fm, functional megaspore; 7, inner
integument ; m, micropyle ; mmc, megaspore mother cell; mt, megaspore tetrad; o7, outer
integument ; pt, parietal tissue; sn, secondary nucleus: sy, synergids.)

Figs 66-70, x80; 71-81, 540.

GWENNETH J. HINDMARSH 285

The three antipodal cells are short lived and do not persist after fertilization.
The binucleate endosperm cell consists of the original central vacuole surrounded
by a thin layer of cytoplasm containing the polar nuclei. Just before fertiliza-
tion, the chalazal polar nucleus migrates to the micropylar pole of the cell,
where it becomes closely associated and then fuses with the micropylar polar
nucleus to form the secondary nucleus (Figs 80, 81). The remaining three
nuclei of the embryo sac form the egg apparatus which consists of two synergids
and the egg (Fig. 80). The synergids, which are situated below the micropyle,
are wedge-shaped, and their nuclei are apically situated, while a large vacuole
occupies the base of each cell. The synergids commence degeneration as the
pollen tube makes its way into the embryo sac, but their remains are still
visible at the two-celled stage of the pro-embryo (Fig. 83). The egg cell is
overlaid by the synergids and is at first non-vacuolate, but a large vacuole
forms in the micropylar region of the cell at maturity.

The mature embryo sac becomes elongated in its chalazal region and it
is enclosed by the persisting parietal and nucellar tissues (Fig. 81).

FERTILIZATION

Large numbers of germinating pollen grains were found on the stigma
(Fig. 5) and, although they were monosiphonous, branching frequently occurred
on the stigmatic papillae to give the appearance of bisiphonous grains (Fig. 82).
The pollen tubes grow down the surface of the papillae and between the loosely-
packed cells of the stigmatic branches, finally reaching the hollow stylar canal
which is in direct communication with the loculus of the ovary. At the base
of the stylar canal, the pollen tubes encounter the funiculus of the ovule which
is in contact with the upper wall of the ovary, and follow its course to the
micropyle (Fig. 83). In some instances pollen tubes passed directly from the
stylar canal to the outer integument and then to the micropyle over the surface
of the integuments, thereby passing the funiculus and taking, apparently, a
longer route. After penetrating the micropyle, the pollen tube makes its way
between the elongated cells of the nucellus which encloses the embryo sac and,
although more than one pollen tube reaches the ovule, each branching freely
and becoming entangled with each other, only one appears to liberate male
gametes and effect double fertilization. The pollen tubes do not persist into
embryogeny.

POST-FERTILIZATION CHANGES

(a) EHndosperm

Division of the primary endosperm nucleus precedes that of the zygote
and is not accompanied by wall formation, so that the endosperm is of the
nuclear type (Fig. 84). After further divisions, the free nuclei are distributed
around the proembryo and the periphery of the embryo sac in cytoplasmic
Strands in which starch grains are also visible (Fig. 85). Nuclear divisions
continue until the embryo sac is almost filled with endosperm (Fig. 86), and
cell formation is initiated in the micropylar region when the embryo has become
spherical (Fig. 87). Wall formation proceeds slowly and free nuclei are still
visible in the chalazal region until after the initiation of the cotyledons (Figs
88, 89). The embryo digests the endosperm as it increases in size and finally
only a small cap remains over the apex of the radicle (Fig. 90).

(b) Embryogeny

The zygote enlarges but does not divide until after several free endosperm
nuclei have formed (Figs 84, 91). A transverse division forms the two-celled
proembryos, or first cell generation, which consists of an upper cell, ca, and a
basal cell, cb (Figs 85, 92). These cells divide simultaneously forming the
superposed cells /, l’, m, and cz of the second cell generation (Fig. 93). Each

286 EMBRYOLOGICAL STUDY OF SPECIES OF BASSIA

SY,

30

Figs 82, 83. Growth of the pollen tubes, drawn from whole mounts. Figs 84-90. Endosperm

development.

(ce, cellular endosperm ; ec, endosperm cap; em, embryo; en, endosperm nuclei; es, embryo

sac; ne, nuclear endosperm; pe, perisperm; pg, pollen grains; sc, stylar canal; sg, starch

graims ; sp, stigmatic papilla; sy, synergids; tw, pollen tubes; vs, vascular strand ; z, zygote.)
Figs 82, x220; 88, 86-88, x130; 84, 85, x540; 89, x50; 90, x20.

GWENNETH J. HINDMARSH

bo
ora
AQ

Figs 91-107: 91-103, Embryogeny and changes in shape and dimensions of the embryo :
104-107, The mature seed; 104, T.S. of seed wall; 105, Tangential section of cells of the
inner integument; 106, Tangential section of cells of the outer integument; 107, T.S. of seed.
Lettering of the embryo follows the system of Soueges.
(c, cotyledon; em, embryo; 7w#, imner integument; oz, outer integument: pe, perisperm ;
&, Suspensor; 2, zygote.)

Figs 96-100, x 220; 101-103, 107, «20; 104-106, x 330.

288 EMBRYOLOGICAL STUDY OF SPECIES OF BASSIA

of these divides to give the eight-celled proembryo of the third cell generation,
the lowest cell, ci, dividing transversely into n and n’, while the other three
cells divide vertically (Fig. 94). In the fourth cell generation n and n’ both
undergo a transverse division to give rise to h, k, 0, and p (Fig. 95), which is
followed by quadrants being produced in tiers m and l’ and, at a later stage,
1 (Fig. 96). The embryo proper develops from J, l’, m, and h, while k, 0, and
p form the uniseriate suspensor which is elongate and usually consists of six
superposed cells, which results from further divisions in these cells (Fig. 98).
However, in B. patenticuspis further divisions may form eight superposed cells.
The cells of the suspensor when fully formed are vacuolate and the basal one
enlarges slightly, remaining in contact with the nucellus (Figs 87, 99). The
hypophysis cell, h, divides vertically into two juxtaposed cells (Fig. 96), and
a similar division follows in the inner cells, and a transverse division in the
outer cells of the tiers J, l’, and m (Fig. 97). The cells of tier h divide longitudinally
to form the hypophyseal quadrant, which then divides horizontally (Figs 98,
99). The embryo proper is now spherical in form due to the enlargement of
cells in tiers J, l’, and m. After further divisions, the embryo becomes heart-
shaped and the primordia of the cotyledons differentiate (Fig. 100). The
embryo sac meanwhile has continued its growth into the chalazal region of
the ovule, becoming spirally curved to accommodate the enlarging embryo
which, at maturity, is elongate and spiral (Figs 101-103). This type of embryo
development conforms to the Chenopodiad type of Soueges (1920).

POLYEMBRYONY

No case of polyembryony was observed in the species investigated, but
Cole (1895) reported this to be the normal condition in Beta rubra, in which a
single seed produced as many as four plants. Favorsky (1928) also described
polyembryony resulting from nucellar budding in Beta vulgaria, but Artschwager
and Starrett (1933) reported that it did not occur in their material.

THE MATURE SEED

The fully developed integuments are two-layered except in the micropylar
region where they are four to six cell layers in thickness. During development
the cells of the outer layer of the inner integument gradually lose their contents
and disappear, while the inner layer remains intact into the seed stage (Figs
104, 105). Both the cell layers of the outer integument persist and deposition
of tannin occurs in their cells, while small areas of thickening develop on the
tangential walls of the outer cells and project into the lumina (Figs 104, 106).
As the endosperm is digested, starch grains are deposited in the nucellus so
that in the mature seed the food storage region is the perisperm (Figs 90, 107),
which is characteristic of the Centrospermales.

Acknowledgements

The author wishes to express her thanks to members of the Botany Depart-
ment and especially to Associate Professor G. L. Davis for her guidance during
this investigation.

References

ARTSCHWAGER, E., 1947.—Pollen degeneration in male sterile sugar beets, with special reference
to the tapetal plasmodium. J. Agric. Res., 75: 191-197.

—, and Srarrett, R. C., 1933.—The time factor in fertilization and embryo develop-
ment in the sugar beet. J. Agric. Res., 47: 823-843.

Buareava, H. R., 1936.—The life history of Chenopodium album Linn. Proc. Ind. Acad. Sci.,
BA: 179-200.

Biutines, F. H., 1934.—Male gametophyte of Atriplex hymenelytra. Bot. Gaz.. 95: 477-484.

BisuLtputTrA, T., 1960.—Anatomical and morphological studies in the Chenopodiaceae. I.
Inflorescence of Atriplex and Bassia. Aust. J. Bot., 8: 226-242.

Brack, J. M., 1948.—* Flora of South Australia”. Part II.

Coxe, J. F., 1895.—Polyembryony. Nature, 51: 558.

GWENNETH J. HINDMARSH 289

Favorsky, N., 1928.—Materialien sur biologie und embryologie der suckerrube. Trudy Nsuch.
Inst. Selek, 2: 1-18.

JOHANSEN, D. A., 1950.— Plant Embryology’. (Chronica Botanica Co. Walthon, Mass.).

Manasate, T. S., and Sonanky, I. N., 1953a.—Studies a the Chenopodiaceae. I. Embryology
of Suaeda fruticosa FKorsk. Jour. Univ. Bombay, 21 (5): 81-92.

, 1953b.—Studies in the Chenopodiaceae. III. Embryology of Kochia

scoparia Schard. Jour. Univ. Bombay, 22 (3): 18-25.

, 1954a.—Studies in the Chenopodiaceae. II. Embryology of Arthrocnemum

indicum Mog. Proc. Ind. Acad. Sct., 39B: 212-222.

—, 1954b6.—Studies in the Chenopodiaceae. IV. Embryology of Chenopodium

ambrosioides Linn. Jour. Univ. Bombay, 22 (5): 31-42.

, 1954c.—Studies in the Chenopodiaceae. V. Embryology of Chenopodium
murale Linn. Jour. Univ. Bombay, 23 (3): 25-37.

Mitimr, H. A., Kirne, H. J., and Weser, A. V., 1959.—The development of the androecium,
the gynoecium, and the embryo of Chenopodium ambrosioides L. 9th Cong. Internat. Bot.,
2: 263-264.

Sovurcss, R., 1920.—Development de l’embryon chez le Chenopodiwm bonus-henricus L. C. R.
Acad. Sci. Paris, 170: 467—469. (Quoted in Johansen).

Woop.LAnD, Poh S8., 1964.—The floral morphology and embryology of Themeda australis (R. Br.)
Stapf. Aust. Jour. Bot., 12: 157-172.

STUDIES OF NITROGEN FIXING BACTERIA. IX

STUDY OF INOCULATION OF WHEAT WITH AZOTOBACTER
IN LABORATORY AND FIELD EXPERIMENTS

Y. T. TcHAN and D. L. JACKSON
(Plates xxvii-xxvill)
[Read 27th October, 1965]

Synopsis

Azotobacter used as inoculum on wheat seed can multiply during the germination of the seed.
The organic substances exuded by the seed provide the necessary carbon source to support the
growth of Azotobacter. However, multiplication was subject to competition from other micro-
organisms present on the seed coat and in the soil. On agar media or in sand culture, Azotobacter
was capable of multiplying in the presence of other micro-organisms if combined nitrogen were
not added into the media. In soil, the numbers of Azotobacter increased during the early
germination of wheat seeds but were reduced at later stages.

In field trials our experimental results do not confirm claims that Azotobacter moculation
increases crop yields.

INTRODUCTION

Agricultural experiments with Azotobacter have been carried out since 1902
(Gerlach and Vogel, 1902). Since this date, a considerable amount of data has
accumulated ; review papers by Cooper (1959) and Rubenchik (1960) have
summarized the earlier work. More recently several papers have been published
dealing with the use of Azotobacter as an inoculum and the development of
the organism in the rhizosphere (Brown et al., 1962; Helmeczi, 1963 ;
Katznelson and Strzelezyk, 1961; Macura, 1963 ; Maliozewska, 1961; Némec
and Pecina, 1964; Pochon, 1963 ; Panosyan, 1964; Rakhno and Ryys, 1963 ;
Rovira, 1963; Samtsevich, 1963; Starkey, 1961 ; Strzelezyk, 1961; Vancura,
1964).

The literature gives the impression that seed inoculation by non-symbiotic
micro-organisms is inconsistent in its effects on plant yields. No logical general
explanation has yet been proposed.

In view of the possible importance of Azotobacter inoculation to yield
increase in the wheat industry of Australia, and since no such experiments
on New South Wales wheat soils were available, the authors have examined
the effects of Azotobacter on the growth of wheat under laboratory and field
conditions.

METHODS

The effects of Azotobacter were examined in laboratory trials using sterile
media, sand and soil cultures, and unsterilized cores of soil, and in a number
of field trials. Wheat, Triticum aestiuum L. var. ‘Gabo’ was used in the
laboratory trials; var. ‘ Festival’ was used in the field trials. However, the
wheat used in some earlier experimental work done in France was of unknown
variety.

(a) Laboratory trials

Organisms used: Azotobacter chroococcum strains NG241, Veg. B, Veg. 2,
Sydney peggy, wheat root (a strain of Azot. chroococcum isolated from wheat
rhizosphere), IP1, IP2; Azotobacter macrocytogenes ; Azotobacter vinelandii
(obtained from Delft, Holland) ; Azomonas agilis ; Azomonas insigne.

PROCEEDINGS OF THE LINNEAN SocitnETy oF NEw SoutH WALES, Vol. 90, Part 3

Y. T. TCHAN AND D. L. JACKSON 291

(A) Winogradsky nitrogen free mineral medium: K,HPO,, 1¢;
NaCl, 0:5¢; MnSO,4H,O, 0:-01g; MgSO,,7H,O, 0-5¢;
FeSO,,7H,O, 0-01 g¢; tap water, 1,000 ml.

(B) Ammonium nitrate: 1° solution.

(C) Winogradsky mineral agar medium: Solution (A) with 1-5%
agar.

(D) Winogradsky sucrose agar medium: Medium (C) with 1%
sucrose.

(E) Sand: Coarse river sand, washed with tap water.

(F) Soil cores were collected in tins driven into the soil to a depth
of four inches. Samples were taken from a_ long-cultivated
wheat field at Tamworth, N.S.W., (approx. 34 crops) and from -
an adjacent area that had never been cropped.

Seed sterilization techniques: Seeds were washed in detergent (5%
teepol), then in 1% calcium hypochlorite for 10 minutes. The hypochlorite
was removed by a series of 10 rinses in sterile distilled water.

Azotobacter counting technique: The number of Azotobacter was determined
by the method described previously by Tchan (1952).

(b) Field experiments

During the 1962 season, field trials of Agotobacter inoculation of wheat
were carried out in the vicinity of Tamworth, N.S.W.

A 2 x 3 x 2 factorial design was used to examine the effects of Azotobacter
and the interactions due to added phosphorus and nitrogen fertilizers.

TABLE |

Chemical analyses of soil samples from the land sown to the factorial
trial and from adjacent uncultivated land

Cultivated Soil Adjacent
(Trial Area) Uncultivated Soil
Q—4’” AUER 0—4” Aa
pH (‘sticky point ”
method) .. a: 6-0 6-5 6-0 6-3
Eaxtractable phosphorus
Truog (p.p.m.) ¥: 0 ) 0-4 0
Olsen (p.p.m.).. 3 0-4 2-6 10-9 1-9
Bray (p.p.m.) .. : 3-7 1-4 4-5 2-1
Organic Carbon
Tinsley (%) .. a 0-99 0-78 1-69 1-16

The trial was carried out on a solodized Red-Brown Earth soil (Dr2-23—
Northeote, 1960) that had been under cultivation for about 20 years, with a
rotation based on several years of grazing lucerne (Medicago sativa L.) followed
by some years of wheat cropping. 1962 was the first year of cropping following
_@ period under lucerne. Table 1 shows a comparison of chemical analyses of
the cultivated soil and of samples taken from an adjacent area that had been
cleared and grazed, but had never been cultivated.

The decline in organic carbon and extractable phosphorus due to cultivation
is typical of the differences found in similar comparisons in this district.

The factors and levels used in the trial were: (1), Azotobacter inoculation :
not inoculated, inoculated ; (2) Superphosphate: 0, 80 and 160 Ib/acre; (3)
Sulphate of ammonia: 0, 50 Ib/acre.

292 STUDIES OF NITROGEN FIXING BACTERIA. IX

The treatments were fully randomized in each of two replicates.»

The inoculation treatment was applied by steeping 5 lb. of seed in 40 oz.
of water containing 20 ml. of a 7-days-old Azotobacter (strain IP,) suspension
(10° cells/ml.) for 30 minutes. After soaking, the grain was spread out in a
Shaded place. The uninoculated seed was steeped in the same way, in water
without the Azotobacter suspension.

The seed was sown from a commercial cultivator-drill within a couple of
hours from inoculation or soaking. The seed and fertilizer were separate up
to the time of sowing, but were brought into contact in the delivery tube of
the drill during sowing and were sown together in the drill row in the soil.
The sowing rate was 47 lb/acre.

The plots were nine rows wide (4’8”) by 130 ft. long. At harvest the length
was trimmed to 124’6” and the middle five rows were harvested to give the
yield from 1/120 acre.

In addition, comparison trials were sown at nine other sites in the district,
these consisted of uninoculated and inoculated plots, and were designed to
measure the responses to inoculation over a range of farm soils.. From these
nine and the factorial trial ten sets of data were available for comparison.

RESULTS AND DISCUSSION

(1) Multiplication of Azotobacter in the presence of wheat seedlings in agar
medium.

Wheat seeds (of an unknown French variety) were sterilized as previously
described. This simplified the interpretation of the results by eliminating the
interaction of the microflora of the wheat rhizosphere. The sterile seeds were
inoculated with one drop of a suspension of a culture of Azotobacter strain IP,
(agar culture) and transferred to Winogradsky’s mineral agar medium. The
seeds were then allowed to germinate at room temperature. After one week
the wheat roots were found to be covered with Azotobacter (Plate xxvil, a, ), c).
This was reisolated and found to be identical with the original inoculum. The
experiment was repeated and the results were reproduced. This result indicated
that the culture of Azotobacter used was capable of multiplication in the
rhizosphere of a wheat seedling without added organic substances.

After these preliminary encouraging results, other strains of Azotobacter
chroococcum (including IP,) and other species of Azotobacter and Azomonas as
listed above (under Methods, p. 290) were tested in Australia in a similar
manner with local wheat varieties. The results were all negative. There
was no growth of Azotobacter or Azomonas in the rhizosphere of the wheat
seedling including the strain of Azotobacter chroococcum (wheat-root) originally
isolated from a wheat rhizosphere.

In spite of the reproducible results of the early experiment it was impossible
to repeat the colonization of the rhizosphere of wheat by Azotobacter. Close
examination of the plates showed that the sites where the seeds were deposited,
before their displacement due to germination, contained numerous small
colonies of Azotobacter and Azomonas (Plate xxviii). This suggested that
during germination, at least under these artificial conditions, enough organic
substances had been exuded to support limited growth of Azotobacter in situ.
Also, it appears that if the seeds are inoculated, under such conditions, an
increase in Azotobacter surrounding the seedling at the early stage of germination
may be expected without establishing a true rhizosphere association. Such
growth could provide some growth factors or plant hormone-like substances
(known to be excreted by Azotobacter) to influence the growth of wheat. There-
fore, it was decided to examine the effects of Azotobacter on. wheab grown to
a more advanced stage in sand culture and in soil. Dae:

Y. T. TCHAN AND D. L. JACKSON 293

(2) Multiplication of Azotobacter in contact with wheat seeds in plant tube
cultures.

To obtain more quantitative information a sand culture technique was
used. 10 ml. of Winogradsky mineral solution was added to 50g. of air dried
sand in 3-5cm. diameter tubes. The tubes were then sterilized at 15 lb. for
20 minutes.

The tubes were inoculated with 5ml. of suspension of Azotobacter
chroococcum strain IP, containing 500 cells per ml. (a total of 2,500 cells).
Immediately after inoculation, 35 ml. of mineral medium was added to the
control tube (total volume of 50 ml.) and ten-fold dilutions carried out to
estimate the initial number. The other tubes were seeded with six wheat
grains either killed (by boiling the seed in water), surface sterilized, or unsterile.
In the nitrogen treatment 0:15 ml. of 1% ammonium nitrate was added to
the tubes.

At seven and 21 days, the number of Azotobacter per tube was estimated
and the plant growth was measured by length in ecm. The results are
summarized in Table 2.

In the absence of added nitrogen there was an increase in Azotobacter in
all cases. The killed seeds, the surface sterilized and unsterilized seeds all
provided an organic exudate for the Azotobacter multiplication. Where
nitrogen was added, multiplication again occurred in the presence of the killed
and surface sterilized seed but not in the case of unsterilized seed.

The multiplication of Azotobacter did not influence the growth of seedlings.
At seven days level the interpretation is difficult since the length of the
seedlings is influenced by the initial germination energy. For all practical
purposes, the difference cannot be regarded as having any importance.

The above experiment indicated that the utilization of the organic matter
from exudates by Azotobacter in the case of unsterile seed is subject to
competition from micro-organisms carried by the seed coat.

(3) Multiplication of Azotobacter in soil in the presence of wheat seedlings

In soil, the micro-flora could also influence the multiplication and survival
of Azotobacter in seed inoculation experimentation. Also the amount of
available mineral nitrogen can rarely reach 0-01°% (100 p.p.m.). Therefore
the competition may not be as severe as in the sand experiment with added
NH,No,

The experiment was repeated using soil from a wheat field in place of
sand. Calcium carbonate was added to produce a near neutral pH. The soil
was not sterilized and the ammonium nitrate treatment was omitted. The
results are included in Table 2. A

The seven day count showed an initial increase in Azotobacter, but at 21 days
the numbers had fallen to less than the original inoculation.

These results suggest that limited growth of Azotobacter is possible during
the early stages of germination.

(4) Effects of Azotobacter in pot trials

Soil cores were collected from cultivated and uncropped soil at Tamworth,
N.S.W. Calcium carbonate was added to the surface to bring the pH
approximately to neutrality.

A 23 Azotobacter x phosphorus x nitrogen trial was made. The pre-
liminary result indicated that wheat seed without fertilizer responded very
slightly but not significantly to Azotobacter inoculation. With fertilizer treatment,
there was no response to inoculation. The detailed results are not reported
here.

STUDIES OF NITROGEN FIXING BACTERIA. IX

TABLE 2

Multiplication of Azotobacter in the presence of wheat seeds (initial
number of Azotobacter introduced as inoculum 0-25-10*)

SAND CULTURE NH,NO, not added

7 days incubation

No. of Azotobacter
per tube
Mean value of seed-
ling length
inoculated
uninoculated* ..

21 days incubation

No. of Azotobacter
per tube Ss
Mean value of seed-
ling length
inoculated.
uninoeculated* ..

7 days incubation

No. of Azotobacter
per tube dis
Mean value of seed-
ling length
inoculated
uninoculated* ..

21 days incubation
No. of Azotobacter
per tube ;
Mean value of seed-
ling length
inoculated
uninoculated* ..

Killed

Wheat Grains

Sterilized

40-4-104

31-3-104

41-

21-
20-

104

cm.

-104

Unsterilized

NH,NO, added (final concentration 0-01%)

20-104

37-104

SOIL CULTURE (unsterilized)

7 days incubation

No. of Azotobacter
per tube
Mean value of seed-
ling length
inoculated
uninoculated

21 days incubation

No. of Azotobacter
per tube
Mean value of seed-
ling length
inoculated
uninoculated

2-7-104

26

We

9

19
23

-104

-104

104

0-1-107

12
10-6

0-6-1074

26
27

2-2-104

13
13

0-002-107

27
25

* No Azotobacter was detected at any stage of seedling growth.

Y. T. TCHAN AND D. L. JACKSON 295

A 2 x 2 trial with eight replications was set out to ascertain the effect
of inoculation in cultivated and uncultivated soil. The wheat seeds for the
inoculation treatment were soaked in a suspension of Azotobacter strain IP,
for one hour. Seeds for the uninoculated treatment were soaked in water for
the same time. The soil cores were manipulated with a minimum disturbance
possible to the soil structure. Each pot received six seeds and was later
thinned to four seedlings which were allowed to grow for 74 days in glasshouse
conditions. The pot arrangement and the results were as indicated in Table 3.

Statistical analysis showed that there were no significant differences of
plant weight between the inoculated and the uninoculated treatments.

TABLE 3

Inoculation experiment in cultwated and uncultivated wheat soil. (Dry weight in g.
of 4 wheat plants)

Uncultivated soil Total Mean Cultivated soil Total Mean
Inoculated I 1:07 0-94 0:88 0-60 0-56 0-56 0-87 0-88
7:19 0-898 5°84 0-73
Inoculated IT 0-870 1:02 0-97 0-84 0-71 O-70 O-78 0:78
Uninoculated I 0-7 0-95 0-87 0-75 0-62 O-78 1:06 O-71
6:34 0-906 4-80 0-685
Uninoculated II 0-90 0°83 1:34 — 0-56 0-62 0-45 —

(5) Study of Azotobacter inoculation in field trials

To complete the investigation, field trials as described above (under
Methods, p. 291) were sown in May, 1962. Inspections of the plots during
the growing season and visual scoring for growth revealed no response to the
Azotobacter inoculation.

At harvest the plots were well grown and free from weeds. There was
no damage due to hail, disease, frost or lodging. A summary of the grain
yields is presented in Table 4. The accuracy of the results is indicated by
the low coefficient of variation (4:9%).

TABLE 4

Field Factorial trial. Grain yields in bushels per acre.
(Means of two replicates)

Superphosphate 0 80 160
lb/acre

Sulphate of
Ammonia 0 50 0 50 0 50
lb/acre

Means

Azotobacter
Not inoculated .. 19-7 22-3 26-6 27-6 30-0 29-
Inoculated. . a5 21-4 19-3 27-8 26-5 29-9 29-

Differences due to
Azotobacter
inoculation .. i 1-7 —3:0 +1:2 —l-1 —0-1 +0-5 —0-1

In the absence of fertilizers the difference in yield due to inoculation was
an increase of 1-7 bushels per acre (on means of two plots). Over the whole
trial, however, the mean effect of inoculation was a yield reduction of 0-1
bushels per acre. Neither of these results was statistically significant.

The results indicated that there was an Azotobacter x phosphorus x
nitrogen interaction. Nitrogen appeared to be the dominant factor in the

296 STUDIES OF NITROGEN FIXING BACTERIA. IX

interaction. In the absence of nitrogen, the responses to inoculation at 0, 80
and 160 lb/acre of superphosphate were + 1-7, + 1-2 and — 0-1 bushels per
acre. In the presence of added nitrogen the corresponding responses were
— 3:0, —1-1 and + 0-5 bushels per acre. None of the interactions were
statistically significant.

The experimental data with sand and soil under laboratory conditions
(Table 2) indicated that the multiplication of Azotobacter in the presence of
unsterilized seed was influenced by the available nitrogen. After a week in
sand cultures the number of Azotobacter had dropped below the inoculum level
where the available nitrogen was high. In the absence of added N, the number
of Azotobacter had significantly increased. The situation was not very different
after three weeks. The number of Azotobacter was still substantially higher in
the no nitrogen treatment.

In the soil under laboratory conditions, with no fertilizer added, the number
of Azotobacter increased only ten-fold during the first week and dropped well
below the inoculated number in three weeks. It would not be unreasonable
to speculate that in the presence of nitrogenous fertilizer Azotobacter inoculated
with the seed would not increase but probably decrease and it could not
exercise any influence on the wheat growth. The negative response at 50
Ib/acre of sulphate of ammonia of —3-0 (no superphosphate) and of —1-05
(at 80 lb/acre of superphosphate) in the inoculated trial can not be explained
on a microbiological basis.

Comparison plots

Inoculated and uninoculated plots were sown at nine other sites in the
vicinity of Tamworth, N.S.W. Together with data from the factorial trial
these gave ten sets of data. Two of the sites were on Black Earth soils; the
others were on Solodized Red-Brown Earths. The data are summarized in
Table 5.

TABLE 5
Grain yield from comparison plots. Bus/ac.

Soil Group Inoculation treatment Response

Uninoculated Inoculated

Red-Brown Earth 19-7 21-4 +1:7
14-0 12-1 —1-9
8-5 7:8 —0-7
27-9 25-8 —2-1
8-6 8-0 —0-6
19-6 20-7 =-1-1
15:8 23-4 +7-6
14-0 14-6 -+0-6
Black Harth 41-1 34:1 —7:0
17-8 17:7 (eal
Total 187-0 185-6 —1:-4
Mean —0-14
The mean response, — 0-14 bushels per acre, shows no benefit from

inoculation. In two cases, however, the responses were marked. In the first,
there was an apparent response in favour of inoculation (+ 7-6 bushels per
acre) ; an inspection of the harvest records suggests that soil variation was the
main reason for this apparent response. In the second, the difference was
against inoculation (— 7-0 bushels per acre) ; no explanation can be given.

From the above data no apparent difference existed between the inoculated
and uninoculated treatments.

Y. T. TCHAN AND D. L. JACKSON 29

XQ

CONCLUSIONS

It has been suggested that plant hormone-like substances or growth factors
excreted by Azotobacter could be beneficial to the higher plant (see Starkey,
1961, and Pochon, 1963). Under laboratory conditions Azotobacter is capable
of multiplication by utilizing the organic matter excreted during the germination
of the seed. Therefore, such beneficial influence of Azotobacter should be
noticeable in the early phase of plant growth. Our experiment in the laboratory
and in the field failed to show such response by wheat seedling. Also, under
our experimental conditions, the multiplication of Azotobacter during germination
of the seed occurs only when the competition of other micro-organisms was
not severe. When the available combined nitrogen is high Azotobacter can not
compete successfully for the available organic matter. In the pot trials and
field experiments, no significant beneficial effect could be obtained. Such
conclusion may only apply to our experimental data ; however, when favourable
ecological conditions are prevalent it could not be excluded that a possible
significant influence of yields could be obtained by Azotobacter inoculation.
To determine the most suitable ecological factors in this regard a very elaborate
programme is needed. This would include the study of soil factors, climatic
conditions, the investigation of interrelationship of micro-organism and higher
plant, and genetic studies of biotic partners.

Acknowledgements

We are indebted to Professor J. M. Vincent for reading the manuscript
and helpful criticism. Thanks are due to Dr. J. Pochon of Pasteur
Institute for his hospitality to one of us (Y.T.T.) in his laboratory during the
preliminary period of this work. The authors would like to thank Misses R.
Webb, D. Shaw, P. Mowatt and B. Dein for their help with this work. Financial
assistance from the Wheat Industry Research Committee of New South Wales
and from the Rural Credits Development Fund is gratefully acknowledged.

References

Brown, M. E., Burtineuam, S. K., and Jackson, R. M., 1962.—Studies on Azotobacter species
in soil. I. Comparison of media and techniques for counting Azotobacter in soil. Plant
and Soil, xvii: 309-319; IL. Population of Azotobacter in the rhizosphere and effects of
artificial moculation. Plant and Soil, xvii: 320-332.

CoorrrR, R., 1959.— Bacterial fertilizers in the Soviet Union. Soils and Fertilizers, xxii: 327-333.

Hewtmeczi, B., 1963.—Possibilities of Azotobacter inoculations to maize. Agrokém Talajt.,
11: 481-492 (Hun. f.) (Mezégazd Akad. Drecen, Hungary). (Seen Extract from Soils and
Fertilizers, xxvi: 180 [1963)).

GerRLAcH, M., und VocEL, J., 1902.—Stickstoffsammelnde Bacterien. Zenir. f. Bakt., Abt. I,
8: 669.

; , 1902.—Weitere Versuche mit stickstoffbinden den Bakterien. Zentr. f.
Bakt., Abt. I1, 9: 817.

Kartznetson, H., and SrrRzeLczyK, H., 1961.—Studies in the interaction of plants and free-
living nitrogen-fixing micro-organisms. I. Occurrence of Azotobacter in rhizosphere of
crop plants. Canad. J. Microb., i: 437-447.

Macura, J., 1963.—Interactions between micro-organisms and plant in the rhizosphere. Proc.
of a Symposium on Relationships between Soil Micro-organisms and Plant Roots, 26-33.
Prague, 1963.

MauiszEwska, W., 1961.—Inoculation of plants with Azotobacter. Roezniki Nauk Rolniczych,
91-D: 103-235.

Nemec, A., and Prcina, L., 1964.—Experiments on the application of Azotobacter preparations
in the bacterial inoculation of potato. Ust. Vét. Inf. MZLVV (Rostl. Vyroba), 36: 715—
727 (Cz.r.e.). (Seen Extract from Soils and Fertilizers, xxvii: 47 [1964)]).

Norrucorr, K. H., 1960.—A factual key for the recognition of Australian soils. C.S.I.R.O.
Division of Soils. Divisional Report 4/60.

Pocuon, J., 1963.—Interactions entre les micro-organisms telluriques et les plantes. Microbiol.
Hispan., 16: 117-122.

Panosyan, A. K., 1963.—Influence of some factors on effectiveness of Azotobacteria. Izv.
Akad. Nauk armyan SSR, 16, No. 7: 11-26 (R. Arm.). (seen Extract from Soils and
Fertilizers, xxvii, 46: 1964).

Raxuno, P. K. H., and Ryys, O. O., 1963.—Use of Azotobacter preparation. Mikrobiologiya,
32: 558-561 (R). (seen Extract from Sols and Fertilizers, xxvi, 6: 425 [1963]).

298 STUDIES OF NITROGEN FIXING BACTERIA. IX

Rovrra, A. D., 1963.—Microbial inoculation of plants. I. Establishment of free-living nitrogen
fixing bacteria in the rhizosphere and their effects on maize, tomato and wheat. Plant
and Soil, xix: 304-314.

RUBENCHIE, L. I., 1963.—Azotobacter and its use in agriculture. (translated from Russian).
Published by the Israel Programme for Scientific translations, Jerusalem.

SamTsEvicH, S. A., 1962.—Preparation, use and effectiveness of bacterial fertilizer im the
Ukrainian §8.8.R. Microbiologiya, 31: 932-933. (seen Ext. from Soils and Fertilizers,
xxvi: 427 [1963)).

Starkey, R., 1961.—Relationship of micro-organisms to plant growth: The Rhizosphere.
Micro-organisms of the Rhizosphere: facts and speculations. Recent advance in Botany :
601-604.

STRZELCZYK, E., 1961.—Studies on the interaction of plants and free-living nitrogen-fixing
micro-organisms. II. Development of antagonists of Azotobacter in the rhizosphere of
plants at different stages of growth in 2 soils. Canad. J. Microb., 9: 507-513.

TcoHan, Y. T., 1952.—Studies of N-fixing bacteria. I. A note on the estimation of Azotobacter
in the soil. Proc. Linn. Soc. N.S.W., lxxvu: 89-91.

Vancura, V., 1964.—Root exudates of plants. I. Analysis of root exudates of barley and
wheat in their initial phases of growth. Plant and Soil, xxi: 231-248.

EXPLANATION OF PLATES XXVII-XXVIII.

Plate xxvii.

la. Wheat seed germinated on agar medium. Note the colonies of Azotobacter surrounding
the roots (arrows).

Ib. Detail of a root and root-hairs. Note the growth of Azotobacter in the rhizosphere (dark
areas).

Ie. Detail of root tip and root-hairs with Azotobacter.

Plate xxvili.

If. Wheat seed germinated on agar medium. Micro-colonies of Azotobacter (arrows) using the
exudate of the seed as organic matter. Note the absence of Azotobacter in the rhizosphere.

STUDIES ON THE GENETIC NATURE OF RESISTANCE TO
PUCCINIA GRAMINIS VAR. TRITICI IN SIX VARIETIES
OF COMMON WHEAT

N. H. Lure and I. A. WATSON
Faculty of Agriculture, University of Sydney

[Read 27th October, 1965]

Synopsis

The mode of inheritance of resistance to stem rust was determined for six varieties of 7’.
vulgare. In the case of Eureka the type of F, segregation obtained in crosses with susceptible
varieties was influenced by temperature and by the strain used. The resistance of Eureka to
the laboratory strain 103—H—2 was controlled by two independent factors, Sr6 and an in-
completely dominant factor which was not temperature sensitive.

In each case a different factor conditioned the seedling resistance of Gabo to the strains
126—Anz-—6, 21—Anz—2, 103—H-—2 and A20. Three of these factors were also present in Charter,
but only one (Srll) m Yalta. The gene Srll in Gabo, Charter and Yalta was differentially
transmitted in crosses between these three varieties and the susceptible Mentana and Chinese
Spring, and also, apparently, in crosses with Federation and Morocco. Srll was closely linked
with the factor for leaf rust resistance in Mentana. Another factor, Srgs2, in Gabo and Charter
conferred resistance to strains 103-H—2 and 111—E—2, but was ineffective against field strains.

Seedling resistance of Mentana to 21—Anz—2 was controlled by a single factor, Sr8, which,
together with another factor, conferred resistance to NR-7.

Kenya 117A was found to possess three factors for seedling resistance of which one, Sr9b,
also conferred field resistance, while the other two had only a modifying effect. In the seedling
stage Sr9b was dominant with 222—-Anz-1, 2, 4, 6 but incompletely dominant with other strains.

INTRODUCTION

The task of the plant breeder who is concerned with developing varieties
of wheat resistant to Puccinia graminis Pers. var. tritici (Eriks. and E. Henn.)
has been made difficult by the existence of many pathogenic strains of the
parasite. Especially in recent years, the rapid spread of virulent strains of
the 21 and 34 standard series has been a setback to breeders in New South
Wales and Queensland, as these strains rendered susceptible the commonly
used sources of resistance. All the six varieties used in this study are no longer
resistant to the stem rust strains occurring in the field, while a few years ago
the resistances of Hureka and Kenya 117A were still effective against most
field strains.

The present study was undertaken to study the mode of inheritance of
resistance in six varieties of Triticum vulgare to some of the new strains of stem
rust, and to correlate results obtained by Australian and overseas workers,
which were at variance. A special effort has been made to study the effect of
minor or modifying genes as these could be of some use in future breeding work.

REVIEW OF LITERATURE

For the purpose of this review the types of resistance carried by the three
varieties, Gabo, Charter and Yalta are considered together and the other
varieties seriatim.

Hureka

In 1941 Macindoe reported that a single factor in Eureka conferred resistance
at lower temperatures ; this factor was non-allelic to the single factor in the
varieties Charter and (Gaza x Bobin?), the latter being a sisterline to Gabo.

PROCEEDINGS OF THE LINNEAN Society or New Soutu Wates, Vol. 90, Part 3

300 RESISTANCE TO PUCCINIA GRAMINIS VAR. TRITICI

Watson and Waterhouse (1945) showed that the resistance factor in Eureka
derived from Kenya W 743* C6040 was inherited independently of the single
factors for resistance in Kenya W 744 C6041 and Kenya W 745 C6042. Knott
and Anderson (1956) have designated this factor Sr6. Studying the mode of
inheritance of resistance, they postulated that Sr6 is dominant with race 56
but recessive with race 15B. Recent work by Green ef al. (1960), using many
North American and Australian strains, indicated that Sra: (a designation used
by Australian workers for the resistance gene in Eureka) and Sr6 are the same
and this was also apparent when all F, plants from a cross between Eureka
and the isogenic Sr6 line proved resistant to races 56 and 15B. Peterson and
Campbell (1953) located Sr6 on chromosome XX and this has been confirmed
by other workers. The effect of temperature on the breakdown of rust
resistance in varieties carrying Sr6 has been described by Green and Johnson
(1954) and Forsyth (1956).

A second gene for resistance in Hureka was reported by Athwal (1955)
who used race 42 from India. This gene was not temperature sensitive. Race
42 was interesting as it also did not attack the varieties Bencubbin, Mentana,
Dundee and Uruguay which are susceptible to Australian strains. Athwal
concluded from his studies that in each of these varieties a single factor con-
trolled resistance to race 42 and that the factor in Eureka was allelomorphic
with the factor in Beneubbin. This gene in Eureka was inherited independently
of two factors in Gabo and a single factor in Mentana. The factors for resistance
in Mentana and Dundee were independent of each other: Dundee is one of
the parents of Eureka. Athwal also considered that the factor for resistance
to race 42 in Eureka was allelic or closely linked with the single factor in
Uruguay and with one of the two factors in Kenya 117A which confer resistance
to race 42.

Gabo, Charter and Yalta

Watson (1941) showed that Kenya 745 (the parent of Charter and Yalta)
possessed a single incompletely dominant factor for resistance, and that seedling
resistance and field resistance were highly correlated. In the same year
Macindoe obtained a similar result with Charter using North American race
34. When crosses between the resistant variety (Gaza x Bobin?) and
susceptible varieties were studied with race 34, a single major factor controlled
resistance in the seedling and adult plant stage. Seedling tests with race 19,
however, indicated at least one additional factor for resistance in Charter and
(Gaza x Bobin?).

On the basis of F, data Watson and Waterhouse (1949) classified many
wheat varieties into three groups each carrying a major gene present in Kenya
743, Kenya 744 or Kenya 745 respectively. Into the latter group they also
placed Gabo, Charter and Yalta.

In 1948, however, Sears and Rodenhiser suggested that two linked,
dominant complementary factors were responsible for resistance in crosses
between the resistant Timstein and Chinese Spring monosomic lines. Mono-
somic analysis located these factors on chromosome X, and they were designated
Sri1 and Sri2 (Knott, 1959). Similar conclusions were reached by other
workers who used the varieties Gabo and Lee in crosses with monosomic lines
of Chinese Spring. Lee originated from a Hope x Timstein cross, and Watson
and Stewart (1956) have shown that the varieties Timstein C.I. 12347 and Lee
carry the same resistances aS Gabo. They concluded that Timstein probably
was introduced into the U.S. as a sisterline of Gabo. More recently, a line
from a cross Steinwedel x 7’. timopheevi and carrying the resistance of the latter
has been named Timvera (Watson and Luig, 1958).

* Refers to the University of Sydney Wheat Accession Register.

N. H. LUIG AND I. A. WATSON 301

Work by Luig (1960) demonstrated that the Gabo-type resistance was due
to a single factor and that this factor was differentially transmitted whenever
Mentana was used as the susceptible parent. Sears and Loegering (1961) found
evidence for a pollen-killing gene in Chinese Spring located on chromosome X.

Watson (1943) found a second factor in Kenya 745 which operated against
four out of 22 standard races against which the single incompletely dominant
factor reported earlier (Watson, 1941) gave protection. Both factors seemed
to be independent of each other. As mentioned above Athwal (1953) explained
the segregation in a cross between susceptible Federation and Gabo to Indian
race 42 on the basis of two dominant independent factors, and both these
factors were found to be inherited independently of single factors for resistance
to this race in Mentana, Bencubbin and Eureka.

Kenya 117A

In 1953 Athwal reported a single incompletely dominant factor for
resistance in Kenya 117A to four strains of stem rust, while two factors were
operative in this variety against Indian race 42. Subsequently Athwal and
Watson (1954) found that the single incompletely dominant factor in Kenya
117A was allelic to the factor Srgp; in Kenya 744. Seedling and field resistance
were correlated, but appeared also to be influenced by modifying factors.

More recently Knott and Anderson (1956) reported that Kenya 117A C.I.
13140 carries three independent genes Sr7, Sr9 and Sr10 for resistance to races
56 and 15B. From tests of isogenic lines of Marquis carrying Sr7, Sr9 and
Sr10 with 29 North American and eight Australian strains Green et al. (1960)
concluded that Sr9b is the same as Srp, this gene being operative against
all eight Australian strains. The lines having Sr7 and Srl0 were only
moderately resistant or were susceptible to the eight strains.

Aslam and Ausemus (1958) explained the inheritance of seedling resistance
to race 15B in a cross of Mida with Kenya 117A C.I. 12568 on the basis of
two or three genes for resistance in Kenya 117A and two genes for susceptibility
in Mida. That their results did not agree with the previous findings was
probably due to a different strain of Kenya 117A being used. Watson and
Stewart (1956) stated that Kenya 117A C.I. 12568 and Australian Kenya 117A
W 1347 were dissimilar in their reaction to many strains of stem rust, with
the former being the more susceptible. Kenya 117A W 1347, however, was
indistinguishable from Kenya 117A C.I. 13140. Omar (1959) reported that the
seedling resistance of Kenya 117A to 15B was inherited independently of field
resistance to many races of rust.

Sears, Loegering and Rodenhiser (1957) located Sr9 on chromosome XIII,
and Knott (1959) found that Sr7 was on chromosome VIII.

Mentana
Although Mentana has been used as an extra-differential variety in several
countries, the earlier mentioned work of Athwal (1953) with Indian race 42 is
the only report on the inheritance of resistance that has been found during
the course of this search of the literature.

MATERIALS AND METHODS

A short description of the six main varieties used in the present study
follows.

Hureka II W 1325. A selection from a cross (Kenya C 6040 x Florence
F,) x Dundee. (Macindoe, 1948).

Gabo W 1422. Bred at Sydney University from a cross (Bobin 39 x Gaza
(T. durum) ) xX Bobin 39. (Athwal, 1953).

Charter W 1371. Bred by Dr. S. L. Macindoe from a cross Kenya C 6040
x Gular. In 1948, however, Macindoe stated that the cross was probably

302 RESISTANCE TO PUCCINIA GRAMINIS VAR. TRITICI

made with Kenya C 6042 and not with Kenya C 6040, as Watson had found
no segregation for resistance in crosses between Charter and Kenya C 6042
(Macindoe, 1948).

Yalta W 1373. Originated from a cross (Kenya C 6042 x Pusa 4) x
Dundee made by Dr. 8. L. Macindoe. The spike is square and has short awns,
the glumes are pubescent and white, the grain is white.

Mentana W 1124. Athwal and Watson (1957) stated that Mentana W 1124
was different from Mentana Genetic Stock No. 1028. Professor Ugo de Cillis,
Italy, has identified Mentana W 1124 as ‘“‘ Ciro Menotti ”’, also known as Rachael
or Awnless Mentana. Ciro Menotti resulted from a cross Akagomuki x (Rieti
<x Wilhelmina Tarwe) made in 1917. The spike is short awned, semi-compact,
somewhat clubby at the tip; the glumes are glabrous and brown, the grain
is white.

Kenya 117A W 1347. Accessioned under C.I. 13140 in the U.S.A. The
spike is tip-awned, mid-dense, the glumes are glabrous and white and the
grain is red.

The following varieties were used in particular crosses as susceptible
parents :

Federation W107. Bred by W. Farrer from a cross between a selection
of Improved Fife and Yandilla.

Little Club W1. One of Stakman’s 12 differential hosts. Very susceptible
in the seedling and adult plant stage to all Australian strains of stem and leaf
rust of wheat, but semi-resistant to hybrid strains from crosses between P.
graminis var. tritici and P. graminis var. secalis.

Morocco W 1103. Pedigree unknown. This awned pubescent variety was
chosen for this study because of its high susceptibility to all Australian field
strains of stem and leaf rust.

Chinese Spring W 1806. Obtained from Dr. E. BR. Sears. Susceptible in
the seedling stage to all Australian field strains of stem and leaf rust, but
resistant to all leaf rust strains as adult plants.

Bobin W 39. The pedigree is unknown as Bobin W 39 is distinct from
the commercial Bobin which resulted from a cross between Thew and Steinwedel.

Gular W 1101. Originated from a cross between Wagga 13 and a selection
from Marshalls No. 3. Gular is one of the parents of Charter.

Insignia W 1989. Originated from a cross Ghurka x Ranee. It does not
carry the gene Sr11 which has been incorporated into its derivatives Insignia
48 and Insignia 49.

Kenya C 6042 W 745. An unnamed crossbred introduced from Kenya
Colony.

T156.48.1.2.1 W 2691. Developed from an F, plant from cross (Little
Club x (Gabo? x Charter) ). More susceptible than Little Club to strains of
P. graminis var. secalis.

Mona W 1168. Cross between Plowman’s No. 3 (Bunyip selection) and
Canberra.

The material under study comprised the following crosses :

(a) Reciprocal crosses in all 30 possible combinations between the
six abovementioned varieties.

(b) Backerosses involving the abovementioned crosses.

(c) Reciprocal crosses between the six varieties and the susceptible
Federation.

(d) Crosses involving resistant F, lines for the purpose of obtaining
lines with single genes for resistance.

(e) Crosses between certain varieties to solve special problems which
had arisen during the course of these studies.

N. H. LUIG AND I. A. WATSON 303

Inoculations were made and reaction types recorded as described by
Stakman and Levine (1922). Rust studies in the field were conducted at
Castle Hill Research Station under an artificial epiphytotic.

Designation and description of strains used

In a recent paper Watson and Luig (1963) have proposed a new system
of nomenclature for strains of stem rust occurring in the geographical region
of Australia and New Zealand. Basically the scheme is the same as Watson
and Luig (1961) proposed for leaf rust. According to the new system the
numbers preceding the regional designation ‘“‘ Anz” refer to the international
set of differential varieties, and the numbers following ‘‘ Anz” to the six
extra-differential varieties, which are numbered in a standardized way :
McMurachy (Sr6) —1, Yalta (Srl11) —2, W 2402 (Sr9b) —3, C.I. 12632
(W 1656) — 4, Renown W 2346 — 5, Mentana W 1124 (Sr8) — 6. For example,
if an isolate which on the international set conforms to 21 attacks McMurachy
and Yalta, but is avirulent on the other four supplementals the designation
is Strain 21—Anz-1, 2.

Twelve strains of stem rust were used to test the material under study.
They represent stock cultures and were frequently checked for contamination.
A short description of the strains and their origin is given below:

21—Anz—0. Accession Number 57043. A field strain, mainly occurring in
Tasmania, New Zealand, and in the southern part of the Eastern Australian
Wheat Belt. First identified in 1954 (Watson, 1955), its origin is unknown.

21—Anz-2. Accession Number 57072. Several years ago the most pre-
valent strain in Australia. Was first detected in 1956 and could have arisen
as a mutation from 21—Anz-0, or as the result of somatic recombination between
21—Anz-0 and a strain capable of attacking the Yalta type of resistance.

21—Anz-2, 6. Accession Number 59137. Not widespread. Origin unknown.

126—Anz-6. Accession Number 56196. This is probably identical with the
rust first isolated by Waterhouse in 1926 and determined as race 34. It has
now been replaced by more virulent strains in the field. Origin unknown.
A yellow colour mutant found in the stock culture of 126—Anz—6 was also used.
It proved to be identical with the stock culture in its reaction types on differential
varieties.

126—Anz-1, 6. Accession Number 7316. First identified from Eureka at
Narrabri in 1942. This strain is now nearly extinct, but was widespread in
the years following 1942 when it first heavily damaged crops of Eureka. Could
have originated as a mutation from 126—Anz-—6. (Watson and Singh, 1952).

126—Anz-2, 6. Accession Number 55350. First reported by Watson,
1955. Its spread in the field has been limited. It is closely related to
126—Anz-6 and 222—Anz—2, 6 and could have arisen as a mutation or as a
somatic hybrid. .

222—Anz-1, 2, 4, 6. Found at Hawkesbury Agricultural College, N.S.W.,
on genetic material carrying Sr6 and Triticum timopheevi resistance, and
possibly resulted from a mutation of strain 222—Anz—-1, 2, 6. It is distinctly
lighter in colour than the other Australian field strains and has also a prolonged
incubation period.

Nk-7. Obtained as a somatic hybrid between Yellow NR-2 and Red
111—E-2 (Watson and Luig, 1958b).

103—H-2*. Obtained as a somatic hybrid between Yellow NR-—2 and Red
P. graminis var. secalis (57241) in 1958 (Watson and Luig, 1959). Previous
designation M9-a.

* The letter H refers to a laboratory strain of hybrid origin, while E indicates a strain of
foreign origin.

304 RESISTANCE TO PUCCINIA GRAMINIS VAR. TRITICI

111-H-2. Obtained from Minnesota, U.S.A. A very avirulent strain of
P. graminis var. tritici, which could have arisen from an intervarietal cross
between P. graminis var. tritici and P. graminis var. secalis. Described by
Watson (1957).

A.20. Obtained from selfing a culture of P. graminis var. secalis (57241)
(Watson and Luig, 1962). This strain is virulent on Black Winter Rye, but
gives also a high reaction on the wheat varieties Yalta and Hureka at temperatures
above 75°F.

The rust reaction of eight varieties used in this study to 11 strains of P.
graminis is given in Table 1.

TABLE |

Reaction of the six varieties under study and of Federation and Little Club to the
eleven selected strains of P. graminis*

Variety 21-0 21-2 21-256 126-6 126-1,6 126-2,6 222-1,2,4,6 NR-7 103-H-2 111-H-2' A20

Kureka at

60-65°F 5 3 > 3 3+ 5 3 5 3 3 3
HKureka at

75-80°F 3+ 3+ 3+ 3 3+ 3 3+ 3e gills Silae 3en
Gabo a Be 3¢ i i Bie 3¢ BiG i oi = 2
Charter l= 3+ 3+ 31= c= 3-4 3 3 i i=
Yalta 3sl= 3+ 3+ l= l= 3+ 3 3+ 3e 71+ +35¢ 3e
Mentana 2 2 3 38+ 3+ 3+ 3 : 3 s :
ikea, 1lyN =e Re BB pan Qn me i sie Ds a :
Federation 3 3 3+ 3+ BE 3+ 3+ 3 3=c gtN 1
Little Club 3+ 38+ 3+ 3+ 3+ 3+ 3+ 3+ 2-- 3+ 25

* At temperatures 70°—-75°F unless otherwise stated.

EXPERIMENTAL RESULTS

A. Inheritance of resistance to stem rust
in the varieties under study

1. Hureka W 1325

(a) Inheritance of resistance to strains 21—Anz—2, 126—Anz—6, 126—-Anz-2, 6
and NR-7.

F, Studies

At temperatures of 60°-65°F Eureka is resistant to all except two of the
Strains used in this study. The virulent ones are 126—Anz-1, 6 and 222—Anz—
1, 2, 4, 6. As mentioned in the literature review and indicated in Table 1,
the resistance of Hureka to most of the former strains is ineffective at high
temperatures. F, seedlings of crosses between Eureka and the susceptible
varieties Little Club, Mentana, Federation and Yalta were tested at tempera-
tures of 60°-65°F. The results are shown in Table 2 and indicate dominance
of resistance to strains 126-Anz-—6, 126-Anz—2, 6 and NR-7 and incomplete
dominance where 21—Anz—2 was used.

F, and F, Studies
Seedlings of five F, families from a cross between Eureka (resistant) and
Yalta (susceptible) were tested with strain 21-Anz—2 and seedlings of one of
these families were tested with strain NR-7. The tests were carried out at
temperatures of 60°-65°F. When analysed statistically the data were homo-
geneous and indicated a single dominant factor for resistance in Eureka
(Table 3).

N. H. LUIG AND I. A. WATSON 305

In 1958, F, seedlings from a cross between Gabo (susceptible) and Eureka
were tested with strain 126—Anz—2, 6 at day temperatures of 60°-65°F. Out
of a total of 204 plants, 48 were susceptible (‘“‘ 3+ ” reaction) and the remainder
gave a resistant reaction of a ‘‘;1 —” type. The F, seedlings were tagged
according to their reaction type and transplanted into the field at Castle Hill.
Later in the year a severe stem rust epiphytotic developed which was mainly
caused by 21—Anz—2. All plants which had given a resistant reaction in the
seedling stage were resistant or semi-resistant in the field, while the plants

TABLE 2

Reaction types of F, seedlings of crosses between Hureka
and susceptible varieties when tested with strains 21—Anz-2,
126—Anz—6, 126—Anz—2, 6 and NR-7

Susceptible Parent Strain used F, Reaction
Little Club... .. 126—Anz-6 een
Federation .. .. 126—Anz-6 estan
Mentana fox .. 126—Anz-6 aise ti
Federation .. .. 126—Anz-2, 6 et
Yalta .. a .. 126—-Anz-2, 6 eae
Federation .. .. NR-7 Shas
Federation .. .. 21—Anz-2 ee
Yalta .. Ae .. 21—Anz—2 “xX+”

which had been susceptible as seedlings reacted similarly in the adult stage.
The resulting F, generation was tested with strains 126—Anz—2, 6, 21—-Anz—2
and NR-7 (Table 4). A similar result was obtained for each F, line and from
the data it was concluded that a single, dominant factor in Eureka controlled
reaction to these three strains.

The results from studies of crosses involving Eureka with Mentana, Little

Club, Charter and Federation are given in Tables 3 and 4 and provide further
evidence for a single dominant factor hypothesis.

In the foregoing it has been noted that when F, plants of the cross (Gabo
< Eureka) were studied at Castle Hill in 1958, the one single, dominant factor

TABLE 3

Segregation of F, plants of crosses involving Eureka and susceptible varieties when tested with
strains 21—Anz—2, 126—Anz-6, 126—Anz—2, 6 and NR-7

Susceptible Cross No. Strain Number of seedlings P-value
parent used (3:1)
Resistant Susceptible
Yalta .. A os .. 1156.22.) 21—Anz—2 111 41 0:70-0:50
2 21—Anz—2 133 55 0-20—-0-10
8 21—Anz-2 97 27 0-50-0-30
4 2\—Anz-—2 127 55 0-20-0-10
5 21—Anz—2 161 60 0:50-0:30
1 NR-7 143 47 0-95—-0- 90
Total .. ee ae 772 285 0:20-0:10
Gabo .. 7 sie .. I1156.9.6 126—Anz—2, 6 156 52 1-00
Mentana ‘. oS .. 1158.420.1 126—Anz—6 89 32 0:90-0:80
Federation ! x .. I1158.21.8 126—-Anz-6 107 34 0:90-0:80
Chane 50 .. 1156.24.38 21—Anz—2 146 60 0-20-0-10
Grand Total. . Sec 1,270 463 0:10-0:05

306 RESISTANCE TO PUCCINIA GRAMINIS VAR TRITICI

controlled resistance both in the seedling and adult plant stage. Field studies
on F, plants of crosses between Eureka and the susceptible varieties Little
Club and Federation at Castle Hill in 1960, however, suggested that resistance
in Eureka was recessive. In both seasons the field inoculum comprised mainly
21—-Anz-2. Of the 147 F, plants of cross 1158.21.8 (Kureka x Federation),
35 were classified as resistant and 112 as susceptible, while of the 87 F, plants
of cross 1158.84.4 (Little Club x Eureka), 21 were resistant and the remainder
susceptible. Classification of adult plants in 1960 was made on the basis of
parental reaction: Little Club and Federation were fully susceptible (more
than 60% infection) and Eureka was resistant, being less than 15% infected
and showing only small pustules.

TABLE 4

Reaction of F'; lines of crosses between Eureka and three susceptible varieties when tested with strains
21-Anz—2, 126—Anz-—6, 126—Anz—2, 6 and NR-7

Susceptible Cross No. Strain F, Behaviour P-value
parent used. (il-g 2 3 1)
Resist. Segreg. Suscept.
Yalta .. iS: .. 1156.22.11 21—Anz—2 39 75 47 0-50—0-30
Gabo .. ae .. I156.9.6 126—Anz-2, 6
21—-Anz-—2 45 106 48 0-70-0-50
NR-7
Charter is .. 1156.24.38 21—Anz-—2 39 68 40 0:70-0:50
Little Club .. .. 1158.84.4 126-Anz-6 19 44 24 0-70-0-50
Total .. ee Bi 142 293 159 0-70-0-50

When seedling tests with strain 126—Anz-6 were carried out on the
progeny of the F, plants from the two crosses it was found that the F, plants
resistant in the field gave only resistant progeny. Approximately two-thirds
of the susceptible F, plants gave segregating progeny and one-third gave
homozygous susceptible progeny. The combined field and glasshouse data
from the cross 1156.9.6 (Gabo x EHureka) and from the two above crosses
(1158.21.8 and I158.84.4) indicate that plants heterozygous for Sr6 were
resistant in the field in 1958 but susceptible in the 1960 season. This was
probably due to environmental influences, mainly temperature differences.

(b) Inheritance of resistance in Eureka to strain 103-—H-2.

The results of seedling tests (Table 1) showed that Eureka was resistant
to strains 103—H-2 and 111—E-2 at high temperatures whereas the resistance
to certain other strains became ineffective. This suggested that the resistance
to 103-H-2 might be due to a gene(s) other than Sr6 and studies were carried
out to investigate this possibility.

At day temperatures of above 80°F, 78 F, seedlings of the cross Little
Club x Eureka were tested with 103—H-2 and the results are presented in
Table 5.

Little Club, used as the susceptible parent, gave evidence of some resistance
to this strain and this could account in part for the variations obtained in both
the intermediate and susceptible classes. Our unpublished work shows that
many other varieties in addition to Little Club have genes which operate against
strains like 103—H-2 which have arisen from inter-varietal crosses in P. graminis.
From the broad classification used, however, the data suggest a single factor
for high resistance in Eureka.

N. H. LUIG AND I. A. WATSON 307

TABLE 5

Segregation of F, plants of the cross (Little Club x Eureka) when tested with strain 103—H—-2
at temperatures above 80°F

F, Segregation

-Parent Parental P-value
reaction Resistant Intermediate Susceptible (Ul 3 2-8 1D)
(O° 3 Be) (“2—, 2, 3—¢”’) (“2++,37)
Little
Club “24++4+,3+ce” 14 42 24
0 -30-0-20
Eureka BO) Se

Correlated studies on F, lines of the cross Gabo x Eureka, using strains
126—Anz-2, 6 and 103—H—2 further indicated that Sr6 was not operative against
103—H-2 at high temperatures. The resistance of Eureka to the latter was
due to a second major gene tentatively designated Srg.

2. Gabo W 1422, Charter W 1371 and Yalta W 1373
The varieties Gabo, Charter and Yalta will be discussed together as the
data from crosses involving these varieties show that they have common
factors for resistance to certain strains.

(a) Intercrosses between Gabo, Charter and Yalta.

As indicated in the literature review, Gabo, Charter and Yalta were found
to possess the same gene (or genes) for resistance to strain 126—Anz-6. To
confirm this, F, and F, generation material of intercrosses between these three
varieties was studied (Table 6) and, as expected, no susceptible segregates
were found.

TABLE 6

Reaction of F, plants and breeding behaviour of F lines of intercrosses
involving Gabo, Charter and Yalta when tested with strain 126—Anz-6

F, Segregation Upper limit of
Cross — SSS —— recombination
Resistant Susceptible at -05 level
Gabo x Yalta On 788 = 12-65%
Gabo x Charter .. 169 = 26-62%
Charter x Yalta .. 226 = 23-16%
EF, Behaviour
Resist. Segreg. Suscept.
Gabo x Yalta 2 178 — = 1-72%
Charter x Gabo .. 93 — — 3-26%,
Charter x Yalta .. 119 — == 2-54%

(b) Crosses between Gabo, Charter and Yalta and susceptible varieties
(i) Inheritance of resistance in Gabo, Charter and Yalta to strains 126—Anz-6
and 126—Anz-1, 6.
F, Studies

F, seedlings of crosses between the three resistant varieties Gabo, Charter
and Yalta and susceptible varieties gave a resistant reaction type (‘‘;1 + ”)
to strain 126—Anz-6, indicating that resistance is completely dominant in the
F, generation.

308 RESISTANCE TO PUCCINIA GRAMINIS VAR. TRITICI

F, and F, Studies

F, and F, generation material from crosses between the resistant varieties
Gabo, Charter and Yalta and the susceptible varieties Kureka and Bobin W 39
were tested with strain 126—Anz-1, 6. Segregation of a single dominant factor
pair for resistance in each cross was indicated (Tables 7 and 8). Likewise,
F, segregation results of crosses between Gabo and Charter and the susceptible
Federation, Insignia and Little Club suggested a single dominant factor for
resistance.

Data from some F, families of crosses (Gabo x Morocco) and (Morocco
x Yalta), however, did not fit a single factor hypothesis and in the crosses
(£156.48.1.2.1 W 2691 x Gabo) and (Yalta x Federation) and reciprocal,
deviations from a three to one ratio were so great that another explanation

TABLE 7

Results of tests with strain 126—Anz-1, 6 on F, populations from crosses between
the three resistant varieties Gabo, Charter and Yalta and susceptible varieties

Parents Cross No. F, Segregation P-value
and Family (3: 1)
Resist. Suscept.
Gabo x Hureka fs 1156.9.2 335 113 0:95-0:90
Eureka x Gabo Be 1156.20.1 127 48 0:50-0:30
Total .. 2: 462 161 0:70-0:50
Gabo x Federation .. 1T58.42.5 92 32 0:90-0:80
Federation x Gabo .. 1158.32.2 126 47 0:70-0:50
Federation x Gabo 6 310 101 0:90-0:80
it Do} r?-)| See on 528 180 0:80-0:70
Gabo x Morocco ai 1159.345.1 208 73 0:80-0:70
Gabo x Morocco ey ne, 129 59 0:05-0:02
Gabo <x Morocco 3 65 15 0:20-0:10
Gabo x Morocco 4 112 44 0:50-0:30
Total .. gs 514 191 0-20
Insignia x Gabo a T159.358.1 203 82 0- 20-0: 10
Insignia x Gabo me 22, 150 50 1:00
Total .. ae 353 132 0:30-0:20
1156.48.1.2.1 x Gabo .. 1161.90.5 83 62 <0-001
1156.48.1.2.1 x Gabo .. 11 lll 49 0:20-0:10
Total .. eS 194. 111 <0:001
Mona x Gabo .. is 1159.383.1 156 50 0:90-0:80
Charter X Boeke a 1156.40.1 89 31 0:90-0:80
Charter M Federation . . 1158.19.3 279 99 0:70-0:50
Charter x Federation . . 4 34 12 0:90-0:80
Totaly. as 313 Ill 0- 70-0: 50
Little Club x Charter 1158.84.3 408 125 0:50-0:20
Yalta x Eureka ad 1158.40.1 59 22 0-70—0-50
Yalta x Hureka ate oe 96 37 0-50—-0-30
Eureka x Yalta an 1156.22.6 186 61 0:95-0:90
Eureka x Yalta i 8 58 19 0:95
Eureka x Yalta tae 1158.419.5 126 44 0:80-0:70
Total .. 23 525 183 0:70-0:50

N. H. LUIG AND I. A. WATSON 309

TABLE 7.—Continued.

Results of tests with strain 126-Anz-1, 6 on F, populations from crosses between the
three resistant varieties Gabo, Charter and Y alta and susceptible variettes—Continued

Parents Cross No. F, Segregation P-value
and Family (@ 2 1)
Resist. Suscept.

Yalta x Federation .. 1158.41.2 54 9 0-05—0- 02
Yalta x Federation .. 3 115 43 0:70-0:50
Yalta x Federation .. 4 64 20 0-80
Federation x Yalta .. 1158.33.1 26 12 0:50-0:30
Federation x Yalta .. aD, 36 3 0:02-0:01
Federation x Yalta 3 54 18 1-00
Federation x Yalta 5 63 10 0-05—0- 02
Federation « Yalta 6 42 22 0-10—0-05
Federation x Yalta ad 21 6 0-80-0- 70
Federation x Yalta 8 47 9 0:20-0:10
Federation <x Yalta .. 9 56 26 0-20-0-10
Federation x Yalta .. .10 118 39 0: 98—0-95
Federation x Yalta .. ll 102 18 0:02-0:01
Federation x Yalta ole 71 21 0:70-0:50
Federation x Yalta 13 244 62 0:10-0:05
Federation < Yalta 14 87 26 0:70-0:50

Total. .. he 1,200 344 0-02-0-01

Heterogeneity (3-49: 1): y?= 30-674; df= 15; P-value= 0-01-0-001

Yalta aie 1159.355.

Morocco x 2 115 36 0-80—0-70
Morocco x Yalta Nee o 159 64 0:30-0:20
Morocco x Yalta mee 4 236 71 0:50-0:30
Morocco x Yalta 5 229 56 0:05-0:02
Morocco x Yalta 6 96 38 0:50-0:30
Moroceo x Yalta a 103 22 0:10-0:05

4toyell =e a 938 287 0- 30-0: 20

had to be found. Moreover, the test for heterogeneity of the latter reciprocal
cross gave a significant y? value.

When Chinese Spring and Mentana were used as the susceptible parents
in crosses with three resistant varieties, the F, ratios again did not fit a single
factor hypothesis (Tables 9 and 10). While the F, data of crosses involving
Gabo and Charter proved to be homogeneous when analysed statistically this

TABLE 8
Reaction of F lines of three crosses when tested with strain 126—Anz—l, 6

F, Segregation
Cross P-value
Resist. Segreg. Suscept. Total (2)

Eureka x Yalta 3 43 37 79 45 161 0:70-0:50
Gabo x Eureka Be 49 103 47 199 0- 90-0: 80
Charter x Bobin 39 16 34 16 66 0-98—0-95

Total .. id 102 216 108 426 0-90—0- 80

was not so of reciprocal crosses of Yalta with Mentana and the y? value from
the cross involving Yalta with Chinese Spring also suggested heterogeneity.
The F, ratios varied according to the F, plant from which they came, and
apparently no maternal influences were present in the reciprocal crosses.

It has been suggested that this departure from Mendelian segregation in
crosses involving Gabo, Charter and Yalta is due to differential transmission

310 RESISTANCE TO PUCCINIA GRAMINIS VAR. TRITICI

of gametes containing the alleles for rust reaction (Luig, 1960; 1964). The
data which provided the first conclusive evidence to this were obtained mainly
from correlated F, studies of stem rust and leaf rust resistance. Mentana is
resistant to strain 68—Anz-1, 2, 3 of leaf rust, Puccinia recondita Rob. ex Desm.,
and this resistance was found to be closely linked with the stem rust resistance
of Gabo, Charter and Yalta. By testing F, lines with the two pathogens
(Table 11) it was possible to establish the following: (i) a very close linkage
in the repulsion phase between leaf rust and stem rust reaction, so that among

TABLE 9
Segregation of plants of Ff, populations from crosses between the three resistant varieties Gabo,
Charter and Yalta and the susceptible varieties Chinese Spring and Mentana when tested with
strain 126—-Anz-6

Cross No. F, Segregation Ratio P-value
Parents and amily — (Res. : Sus.) (3:1)
Resist. Suscept.
Chinese Spring x Gabo 1161.445.1 163 80 2-0: 1 0-01—0-001
Chinese Spring x Gabo 2 144 75 Iho®) 3 i 0-01-0-001
Chinese Spring x Gabo 3 133 77 log I <0-001
Chinese Spring x Gabo 4 214 116 the} 3 J <0-001
Total 654 348 ILe@) 3 I <0-001
Chinese Spring x Yalta 1161.444.1 24. 13 1-8: 1 0-20-0-10
Chinese Spring x Yalta .2 11 18 0-6: 1 <0-001
Chinese Spring x Yalta A 26 20 Wess J 0-01—0-001
Chinese Spring x Yalta 9) 47 53 0:9:1 <0-001
Chinese Spring x Yalta 6 21 12 1-9: 1 0:20-0- 10
Chinese Spring x Yalta 7 12 10 Io} g I 0-05—0- 02
Chinese Spring x Yalta 8 48 25 tlo@)¢ J 0-10-0-05
Total 189 151 leg} g i <0-001
Heterogeneity: y*?= 11-885; d.fr.= 6; P-value = 0-05-0-02
Gabo x Mentana .. I156.7.4 107 51 Bell 3 I 0-05—-0- 02
Gabo x Mentana a8 5 83 31 Moly eI 0-70-0- 50
Mentana x Gabo Ula Galrl 85 27 Bells il 0-90-0-30
Mentana x Gabo a 2 129 60 202) 2 I 0-05—-0-02
Mentana x Gabo .. I161.436.1 218 90 2-4: 1 0-10-0-05
Mentana x Gabo oa 2 194. 95 2-0: 1 0-01—0-001
Mentana x Gabo oe 3 118 66 Iho} ¢ It <0-001
Total 934 420 HoH 3 Ih <0-001
Charter x Mentana 1156.37.1 171 80 Bo ths ih 0-02-0-01
Charter x Mentana .... 2 260 111 2-3:1 0-05-0-02
Charter x Mentana is 7 69 39 20) 8 Ih 0-05—-0-02
Total 500 226 2-2: 1 <0-001

786 F, lines none was homozygous resistant to both rusts; (ii) a difference in
the segregation ratios according to whether Gabo, Charter or Yalta was used
as the parent in crosses with Mentana.

The significance of these results in relation to differential transmission of
gametes has already been reported (Luig, 1964).
(ii) Inheritance of resistance in Gabo to strain 21—Anz—2.

The gene Sri1 carried by Gabo, Charter and Yalta is not operative against
21—Anz—2. Gabo gives a “ 3-c, 3c” type of reaction when tested with this
strain while Charter and Yalta are fully susceptible.

When F, and F; generation material of the cross (Gabo x Charter) was
tested with 21—Anz—2, the results suggested that the difference in the reaction

N. H. LUIG AND I. A. WATSON dll

TABLE 10
Reaction of F, families of cross Yalta x Mentana and reciprocal to strain 126—Anz—-6

F, Segregation xr? P-value Xe P-value
Family Ratio (33 3 1) (3: 1) (seo) oil) (ile2) 9 ih)
Resist. Suscept.

Yalta x Mentana 1156.25

“1 10 8 1-3:1 2-722 -10—-05 0-004 -95—-90
-3 25 11 2-3:1 0:593 -50—- 30 2-516 -20—-10
“4 34 16 Poll adh 1-307 -30—-20 2-767 -10—-05
-5 81 43 1:9:1 6:194 -02—--O1 4-075 -05—- 02
“7 122 129 0-9:1 93-260 <-001 6-091 -02—--01
-8 142 109 1198} 6 I 45-452 <-:001 0-006 -95—-90
+9 119 122 1:0:1 84-383 <:001 4-738 -05—- 02
-10 129 82 1-6:1 21-626 <-:001 1-981 -20--10
Total 662 520 ow 3 il 22-178* 01—-001
Mentana x Yalta 1156.4
“1 79 98 -0-8:1 87-053 <:001 9-858 -01—- 001
-2 119 48 Dosy8 Al 1-248 -30—-20 15-103 <:001
-3 160 158 1:0:1 103-350 <:001 4-679 -05—- 02
“4 109 87 Ileoa} 3 Il 39-293 <:001 0-041 -90—- 80
-5 99 47 1-1:1 4-027 -05—- 02 7:829 -01—-001
-6 134 132 ilo) IL 86-020 <:001 3-836 -10—-05
:7 $l 47 a7 3. 9-375 -01—: 001 2-513 -20—-10
-8 92 53 IleZ ¢ Ih 10-320 -O1—- 001 2-985 -10—-05
Total 873 670 1-3:1 46-844* <-001
Grand
Total 1,535 1,190 1:29: 1 69 -022* <-001

* Heterogeneity y? (1-29: 1) = 69-022; P-value (15d.f.)= <-001.

TaBLeE 11

Reaction of F' lines of the crosses (Gabo x Mentana), (Mentana x Charter), (Mentana x Yalta)
and (Yalta x Mentana) to strains 126—Anz—6 of stem rust and 68—Anz-1, 2, 3 of leaf rust

Reaction to stem rust
Gabo x Mentana

Resistant Segregating Susceptible

Resistant — _— 62
Reaction to leaf rust Segregating — 121 3
Susceptible 40 — —

Mentana x Charter
Resistant — — 52
Reaction to leaf rust Segregating — 88 =
Susceptible 37 1 =

Mentana x Yalta
Resistant — — 62
Reaction to leaf rust Segregating — 84 2
Susceptible 24 1 1

Yalta x Mentana
Resistant — — 107
Reaction to leaf rust Segregating 1 93 2
Susceptible 4 1 25

312 RESISTANCE TO PUCCINIA GRAMINIS VAR. TRITICI

types of these two varieties was due to a single factor pair possessed by Gabo
(Table 12). Provisionally, this factor is designated Sr@.

(iii) Inheritance of resistance in Gabo and Charter to strains 103-H-2,
111—-E-2 and A20.

As shown in Table 1, Gabo and Charter are highly resistant in the seedling
stage to 103—H-2, while Yalta gives a semi-susceptible reaction. Apparently
the gene Srl1l does not operate against this strain. To investigate the nature
of the resistance to 103—H-2, F, and F, generation material from crosses between
the three varieties was studied. F, seedlings of crosses of W2691 (“3”
reaction type) x Gabo and Little Club (“‘2 + +” reaction type) x Charter
were resistant (‘‘; 1” reaction type) thus indicating dominance of resistance.

TABLE 12
Reaction of F, plants and breeding behaviour of F lines to strain 21—Anz—2 of
cross (Gabo x Charter)

F, Segregation

P-value
Resistant Intermediate Susceptible Total (1:2: 1)
« 3 =, 3c 2) (ss 3c, 3 ale c 2) (ee 3 aL oo)
29 62 21 112 0:30-0:20
F, Behaviour
Resistant Segregating Susceptible Total
5 13 3 21 0-50—-0-30

F, and F, Studies

From a cross (Gabo x Charter), 169 F, seedlings were tested with 103—H-2
but no susceptible segregates were found. Likewise, only highly resistant
plants were found when 97 F, families of this cross were tested. The presence
of the same factor pair for resistance to strain 103-H-—2 in both varieties is
thus suggested. If it is assumed that the genes in Gabo and Charter are non-
allelic and dominant, an upper limit of recombination of 3-06% at the 0:95
level of probability could be calculated according to the formula 0-05 =

2\n
1—p—7Z] where p is the recombination value and n the number of F,

lines tested.

When F, populations from a cross Gabo x Yalta were tested with 103—H-2
a single factor difference was obtained. 104 F, seedlings gave a reaction as
high as that of Yalta or were fully susceptible, while the remaining 306 plants
resembled the resistant parent (Table 13).

TABLE 13
Reaction to strain 103—H-2 of F, seedlings of the crosses (Gabo x Yalta) and (Charter x Yalta)

Parents F, Segregation
or Parental Probability
Cross reaction Resistant Susceptible (3 Res.: 1 Sus.)
(74 : 1 = 99 (79 if + bi) (T4 XE +, 8c 2° ce 3 o>

Gabo ca geese — ae

Yalta eS aCe

1156.10 on 287 19 80 24 -90—: 80
Charter eg oa ae

Yalta Bip SOG.”

1156.41.7 ... 124 18 30 9 -30—- 20

N. H. LUIG AND I. A. WATSON 313

F, plants were tagged according to their reaction type and grown to maturity.
Subsequently their breeding behaviour to 103—H-—2 was studied (Table 14).

The correlated results indicated that a single major factor pair was
responsible for the resistance in Gabo. Variations in reaction type in the
resistant and susceptible group are probably due to the action of a minor gene
(or genes) in Yalta and/or Gabo.

Further evidence for a single dominant factor for resistance to strain
103—H-2 was obtained from F, studies of cross Charter x Yalta (Table 13).
Studies using monosomic lines of Chinese Spring indicate that this single factor
in Gabo and Charter is located on chromosome XVII (Baker and Luig, un-
published ; McIntosh, unpublished).

TABLE 14

F, Breeding behaviour of F, plants from cross 1156.10.3 (Gabo x Yalta) representing different
reaction types, when tested with strain 103—H-2

F, Behaviour

Resistant Segregating Susceptible Total
“s1—” 43 92 — 135
Reaction types “yT+4” — 2 6 8
in F, “X+, 3c” — 1 38 39
ce 3 39 ees ae. 8 8
Total 43 95 52 190
xy” for a 1:2:1 ratio= 0-852 P-value = 0-70-0-50

Studies were also carried out with 111—E-2 and A20 on F, generation
material of the crosses (Charter x Gabo) and (Gabo x Yalta). Charter and
Gabo are fully resistant to these strains, while Yalta at temperatures above
75°F gives a “ 3-n”’ reaction with 111-E-2 and a “3” reaction with A20.
No susceptible segregates were obtained from 97 F, lines of the cross (Charter
x Gabo) when tested with the two strains, and this suggested that Charter
and Gabo have genes for resistance in common, as was found when testing
the same lines with 103-—H-2. When 42 F, lines of the cross 1156.10.3 (Gabo
x Yalta) previously tested with 103-H-—2 were inoculated with 111—E-2 and
with A20 it was evident that the same dominant factor in Gabo was operative
against 103-—H-2 and against 111—E-2, and that the high resistance to A20
was due to a different single factor. Tentatively these two factors have been
designated Sres and Sree, respectively.

3. Kenya 117A W 1347
F, Studies

The F, seedlings from crosses between Kenya 117A and _ susceptible
Federation and reciprocal were inoculated with six strains of stem rust and
the results are shown in Table 15. The reaction types varied from a “3 =c”
to a ‘“‘3 + c¢” according to the strain used and indicated incomplete dominance
of resistance.

F, Studies

When F, populations of crosses between Kenya 117A and susceptible
varieties were tested with strains 126-Anz-6, 126—Anz-2, 6, 21—Anz—2,
222—Anz-1, 2, 4, 6 and NR-7, varying segregation patterns were obtained.
It was evident that the resistance of Kenya 117A was conditioned by more
than one factor. As tests were not all carried out under similar environmental

314 RESISTANCE TO PUCCINIA GRAMINIS VAR. TRITICI

conditions it was not possible to make accurate comparisons between segrega-
tion ratios obtained with different strains, but there seemed to be fewer
susceptible segregates when testing with strains 21-Anz—2 and NR-7.

F, Studies
(i) Inheritance of resistance in Kenya 117A in a cross with susceptible Yalta.

Of cross 1156.35.2 (Kenya 117A x Yalta) 202 seedlings were transplanted
into the field and seed was harvested from 181 plants. When the resulting
181 F, families were tested with strain 21—-Anz—2, 176 were either resistant or
segregated with resistant and susceptible plants and five were susceptible. A
ratio of 63: 1 is proposed (P-value = 0-20-0-10) on the basis of segregation
of three independent factor pairs. The detailed results of testing these lines,
together with the proposed genotypes, are outlined in Table 16. In two of
the eight classes the numbers obtained did not conform to the expected ratio.

TABLE 15

Reaction of F, seedlings of crosses between Kenya 117A and the susceptible varieties Yalta and
Federation when tested with strains 21—Anz—2, 126—Anz—6, 126—Anz—2, 6, 222—Anz—1, 2, 4, 6

and NR-7
Parents Reaction of parents and of F, plants
or :

Cross Strain used 21-2 126-6 126-2, 6 222-1, 2, 4, 6 NR-7
Kenya 117A = 2) == im 2=n 2 = aa
Federation 3 + 3+ 3+ 3 3+
Yalta oe oe até rors — — — —

Kenya 117A x Federation 3 — 3¢ —— d= Co =

Federation <x Kenya 117A 3—e — 3¢ — 3c

Yalta x Kenya 117A 3—e — — = =

There were approximately 20 lines in excess in one class (‘‘ Segregating
Resistant and Susceptible’) and about the same number was lacking in the
other class (‘‘ Segregating Resistant, Intermediate and Moderately Susceptible ’’).
This is probably due to plants homozygous for the third factor being classified
as susceptible.

The inheritance of resistance of Kenya 117A was also studied by testing
the same F, families of cross 1[156.35.2 with strain 126—-Anz—2, 6 (Table 16).
Results were in most cases similar to those observed when testing with 21—Anz—2.
This similarity in behaviour is thought to be due mainly to the segregation
of Sr9b, a gene which confers resistance to both strains. The other two factors
postulated to be effective against 21—-Anz—2, seemed to confer a moderate
resistance to 126—Anz—2, 6. One of these factors, presumably Sr10 (Green
et al., 1960), gave a ““3—e, 3c” reaction type when homozygous, while the
other factor, presumably Sr7 (Green et al., 1960), produced a “3 +c”
reaction type. It is probable that in some instances the moderate susceptibility
of lines carrying Sr7 only was mistaken for full susceptibility, and this would
explain the discrepancy of observed and expected numbers in some segregation
classes. Additive effects of Sr7 and Sr10 in combination with each other and
with Sr9b were also in evidence.

Kenya 117A has noticeably more resistance to 222—Anz-1, 2, 4, 6 than
to strains 21—Anz—2, 21—Anz—2, 6, 126—Anz-6, 126—Anz—2, 6 and NR-7. But
comparatively more susceptible or semi-susceptible F, plants and F, lines were
found when crosses of Kenya 117A with two susceptible varieties were tested
with 222—-Anz-1, 2, 4, 6. Tests on 128 F, families from a cross of Kenya 117A
with Yalta (“3 reaction type) showed that Kenya 117A carries one gene
conferring a high degree of resistance, and a second minor gene conditioning
a moderately susceptible reaction (‘‘ 3c”) to 222-Anz-1, 2, 4, 6 (Table 16).

N. H. LUIG AND I. A. WATSON 5 5)

Correlated F; data indicated that the major gene was the same (Sr9b) as that
which gave protection against 21—Anz—2, 21—Anz-—2, 6, 126—Anz-—6, 126—Anz-—2, 6
and NR-7, but it was evident that against 222—Anz-1, 2, 4, 6 the gene con-
ferred a much higher degree of resistance. Sr9b conditioned a type “;2 =”
reaction to 222—-Anz-1, 2, 4, 6 when homozygous and a type ‘‘ 2 — ”’ reaction
when heterozygous. With the other five strains Sr9b was incompletely
dominant.

The minor gene in Kenya 117A for resistance to 222—Anz-1, 2, 4, 6 is
probably Sri0. This gene was also operative against 126—Anz—2, 6 and gave
a semi-resistant ‘“‘3—en”’ reaction to 21—Anz—2. The third gene reported
in Kenya 117A, Sr7, apparently does not condition a reaction on its own to
222-Anz-1, 2, 4, 6 which could be easily distinguished from the reaction type
of Yalta ; it might, however, act as a modifier in combination with Sr9b or Sr10.

Finally, F; families of the cross 1156.35.2 of which sufficient seed was at
hand were tested with NR-7. This strain originated as a somatic hybrid from
two North American strains and one of the objectives for using it was to con-
firm the previous assumption (Green et al., 1960) that the three factors which
condition reaction types in segregating material of cross (Kenya 117A x Yalta)
are identical with the three genes Sr7, Sr9b and Srl0. Only 68 F, families
were tested with NR-7, but it was evident from the results that the three
factors which conditioned reaction to the other three strains were also operative
against NR-7.

In an attempt to isolate F, lines which carry only one of the three factors
for resistance in Kenya 117A, seedling tests with the above-mentioned four
strains were simultaneously carried out on selected lines of cross 1156.35.2 at
temperatures ranging from 60° to 70°F. Three lines were found which were
homozygous for each of the three genes, and one line which apparently carried
both Sr7 and Sr10. The reaction types produced by these lines together with
those of their parents, Yalta and Kenya 117A, are shown below:

Proposed Line or Strain used
genotype variety —
21-2 126-2, 6 222-1, 2, 4, 6 NR-7 21-2, 3*
Sr7 246 3¢ 3+¢ 3+¢ 3 +¢
Sr9b 256 2— 2—e 5D 2+ 3. +
Srl0 244 3 — cn 3—e 3c 3—e¢
Sr7, Srl0 2,121 3 = cn 3 = © 3—e 3=e
— Yalta 3 + 38+ 3+¢ 3+ 3+
Sr7, Sr9b, Srl10 Kenya 117A Y= in Q= im :2= D — = ¢C

* This strain was obtained from the 1960 Cereal Rust Survey. It is virulent on the isogenic
Marquis-line W2402 (obtained from Dr. D. R. Knott) and on the variety Festival, both of which
earry Sr9b.

(ii) Inheritance of resistance in Kenya 117A in a cross with susceptible Mentana.

The mode of inheritance of resistance in Kenya 117A to the Australian
field strain 126—Anz-—6, which has been used extensively in inheritance studies
by other workers, was also studied. Yalta, used as the susceptible parent in
the cross I156.35.2, is resistant to 126-Anz—6 and therefore the resistance of
Kenya 117A to 126—Anz-—6 could not be studied in this cross. Cross 1156.5.3
(Mentana x Kenya 117A) was used instead as Mentana is susceptible to
126—Anz-6 as well as to 126—Anz-2, 6, 21—Anz-2, 6 and 222—Anz-1, 2, 4, 6.
Mentana gives only a semi-resistant ‘‘2”’ type of reaction to 21—Anz-—0 and
21—Anz-2.

When 126 F, lines of cross [156.5.3 were inoculated with 126—Anz-—6, the
results indicated segregation of two independent and incompletely dominant

316 RESISTANCE TO PUCCINIA GRAMINIS VAR. TRITICI

TABLE 16
Distribution of F lines of the cross [156.35.2 (Yalta x Kenya 117A) for reaction to strains
21—Anz-2, 126—Anz—-2, 6 and 222—Anz—-1, 2, 4, 6, and proposed genotypes for the segregation
classes

F, Rust behaviour and proposed genotypes

Strain Total
R Seg. R, I Seg. MS Seg. Seg. I, S)
I & MS I & MS R&S MS &«&S
21—Anz-2 44 21 6 iG 5 74 19 5 181
(Expected numbers) (45-3) (39-6) (5:7) (11-3) (2-8) (50-9) (22-6) (2-8)
126—Anz-2, 6 34 12 6 2 4 54 15 4 131
(Expected numbers) (32-8) (28-7) (4-1) (8-2) (2-0) (36-8) (16-4) (2-0)
Proposed AABBCC AaBBCC aaBBCC aaBBCe aabbCC AaBbCc aaBbCe aabbce
genotypes and AABBCe AaBBCce aaBBcec aaBbCC AaBbce aaBbece
fraction of total AABBce AaBBce AabbCe aabbCe
AABbCC AaBbCC Aabbee
AABbCe <AabbCC
AABbce
AAbbCC
AAbbCc
AAbbecec
16/64 14/64 2/64 4/64 1/64 18/64 8/64 1/64
F,; Rust behaviour and proposed genotypes
Strain —_—_ Total
R Seg. MS Seg. Seg. NS)
R & MS R&«&s MS & 8S
222-Anz-1, 2, 4, 6 30 12 9 54 15 8 128
(Expected numbers) (32) (16) (8) (48) (16) (8)
Proposed AABB AaBB aaBB AaBb aaBb aabb
genotypes and AABb Aabb
fraction of total AAbb
4/16 2/16 1/16 6/16 2/16 1/16

Explanation of abbreviations: R —resistant; I —intermediate; MS—moderately susceptible ;

S —susceptible; Seg.—segregating.

Proposed genes: A—Sr9b; B—Sr10; C—Sr7.

factor pairs (Table 17). The two factors, however, were not equal in the degree
of resistance they conferred on the F, plants. The major factor, which is
later shown to be Sr9b, imparted a high resistance (type “‘ 2—” reaction)
when homozygous, while the minor factor conditioned a reaction type ranging
only from “3—n” to “3c”. This variation in reaction type of F, lines
thought to possess the minor gene in the homozygous state could be due to
the action of a modifying factor, and/or environmental influences.

Strains 126—Anz—2, 6 and 21—Anz-—2, 6 were also used to inoculate these
F, families. The inheritance of resistance in Kenya 117A to the former in

TABLE 17

Distribution of F, lines of cross [156.5.3 (Mentana x Kenya 117A) for reaction to strain 126—Anz-6,
and proposed genotypes for the segregation classes

Behaviour of F, lines and their proposed genotypes

Segreg. Segreg. Segreg. Segreg.
oe Qa Hee 6 Di 29 oe im vee oe 3—e 29 ce Oy aah ie oe j= ae ee 3+ 29 Total
“6 = 29 ee 3—e br) oe 3+ 9° oe 3 29
Act ual
numbers 25 11 15 3 44 20 8 128
Expected
numbers 24 8 16 8 48 16 8 128
Prop osed
genotypes AABB AAbb AaBB aaBB AaBb aaBb aabb
AABb Aabb

y* for a 3:1:2:1:6:2:1 ratio= 5-688 P-value (6 d.fr.) = 0-50—0- 30

N. H. LUIG AND I. A. WATSON Bl 7)

eross 1156.35.2 has been reported above and Green et al. (1960) have shown a
close similarity in the reaction types produced by strains 126—Anz—6 and
126—Anz-—2, 6 on isogenic Marquis lines carrying genes Sr7, Sr9b or Sr10. As
expected, lines tested with 126—Anz—6 and 126—Anz—2, 6 gave the same reactions
to each strain, suggesting that the major factor for resistance to 126—Anz—6
was Sr9b, while the minor factor was presumably Sr10. Mentana itself gives
a ‘*‘ 3” type reaction to these two strains and it is possible that F, lines carrying
only Sr7 were not significantly more resistant than Mentana.

When the reaction types on F; lines to strains 126—Anz—6 and 21—Anz-—2, 6
were compared two at a time it was evident that several lines carried more
resistance to the latter strain than to the former. This was not unexpected,
as 21—Anz—2, 6 probably originated from 21—Anz—2 and thus would be similarly
constituted in regard to most genes for pathogenicity. When studying cross
1156.35.2 it was found that the factor Sr7 conferred a higher resistance to
21—Anz-—2 than to 126—Anz—2, 6. Therefore the presence of Sr7 presumably
accounted for the higher resistance of some F, lines to 21—Anz—2, 6 than to
126—Anz-6.

FIELD STUDIES

During the years 1958-1960 segregating material of the two above-
mentioned crosses and of other crosses involving Kenya 117A was studied
under field conditions at Castle Hill Research Station, strain 21—Anz-—2 con-
tributing most of the inoculum. Classification of adult plants was made on
the basis of percentage of rust and size of pustules. Plants with less than 30%
infection of stem and with pustules small to medium in size were classified as
resistant, and plants showing mainly large confluent pustules and/or carrying
more than 30° infection were classified as susceptible.

The mature plant reactions of 181 F, plants of cross 1156.35.2 (Yalta x
Kenya 117A) were recorded and the behaviour in the field of the progenies
of these plants was studied in the F, and F, generations. Kenya 117A was
resistant in the field, showing only small pustules on stems less than 10%
infected, while Yalta was extremely susceptible (summer sowings of Kenya
117A, however, showed a much higher degree of infection). Nine F, plants
of the cross I156.35.2 susceptible in the seedling stage to 21—Anz—2 were
susceptible as adult plants. However, 94 of 172 resistant F, seedlings were
also susceptible in the adult plant stage and the remainder were resistant.

The results of the F, field tests were compared for each line with F,
seedlings tests with 2i1—Anz—2 and it was evident that many F, lines resistant
or segregating to 21—-Anz—2 as seedlings were susceptible as adult plants. These
findings can be explained by the incomplete dominance of gene Sr9b and by
assuming that the other two genes, Sr7 and Srl0, become ineffective in the
adult plant. There was, however, evidence which suggested that these latter
genes acted as modifiers in combination with Sr9b. The above results would
indicate that selection for resistance of Kenya 117A to 21—Anz—2 in the F,
generation is of only limited value in breeding procedures.

When seedling reactions of F; lines to 126-Anz—2, 6 and NR-7 were
compared with the behaviour of these lines under field conditions, the same
lack of correlation as in the case of 21—Anz—2 was found. This was expected,
as many lines reacted similarly to the three strains.

There was, however, good correlation between seedling reaction to
222—Anz-1, 2, 4, 6 of F, lines and their behaviour to stem rust in the field.
The comparative data are shown in Table 18. In classifying seedling reaction
only the major factor for resistance (Sr9b) was considered and it can be seen
that all except two of the 32 families classified as susceptible to 222—Anz-1,
2, 4, 6 in the seedling stage were also susceptible in the field to a combination of
strains consisting mainly of 21—-Anz—2. The two families which were classified

318 RESISTANCE TO PUCCINIA GRAMINIS VAR. TRITICI

as segregating in the field probably comprised susceptible plants and plants
which had escaped infection to a large degree. These results indicate that
no gene (or genes) of Kenya 117A other than Sr9b can confer field resistance
to 21-Anz-2. Of the 30 F, lines which in seedling tests appeared to be homo-
zygous for Sr9b, 25 lines were found to be resistant in the field, a further
indication that Sr9b is the main factor for field resistance in Kenya 117A.
However, 21 out of 66 families, apparently heterozygous for this gene, were
classified as susceptible in the field. It is possible that resistant adult plants
were present in the 21 lines, but were not classified as such.

TABLE 18

Distribution of F, lines of the cross Yalta x Kenya 117A for
reaction to a collection of strains in the field, being mainly
21—Anz—2, and for seedling reaction to 222—Anz—l, 2, 4, 6

Seedling Behaviour in the field
reaction
to 222— Resistant Segregating Susceptible
Anz-l, 2, 4, 6
Resistant 30 25 3 2
Segregating 66 4 4] 21
Susceptible 32 = 2 30
Total 128 29 46 53

In 1958, F, plants of cross 1156.5.3 (Mentana x Kenya 117A) were also
studied for their reaction in the field. Mentana was moderately susceptible
in the field, showing approximately 40 to 60% infection on stems with pustules
of medium size. Of the 122 F, plants, 21 were found to be resistant and 101
were classified as susceptible. In some cases, however, difficulties were
experienced in making the classification. As mentioned before, the progenies
of these F, plants were tested in the seedling stage with strains 126—Anz-6,
126-—Anz-—2, 6 and 21—Anz—2, 6, and when the results of the seedling tests were
compared for each F, line with the mature plant reaction of the corresponding
F, plant, it was evident that the same, single, major factor for resistance was
operating in the seedling and adult plant stage (Table 19). Of the 21 resistant
F, plants all, except one, carried the gene Sr9b either in the homozygous or
heterozygous condition. Thus the findings on the nature of mature plant
resistance in Kenya 117A in this cross are in agreement with those for cross
1156.35.2

TABLE 19

Reaction in the field of F, plants of the cross Mentana x Kenya 117A and seedling reaction of F;
lines raised from these plants when tested to strains 126—Anz-—6, 126—Anz—2, 6 and 21—Anz-2, 6

Behaviour of Proposed No. of Field reaction of
F, lines genotype* F, lines F, plant
of F, plant
Resistant Susceptible
All 2—n, 2—c, 2— ip ss AABB 25 9 16
AABb
All 2— : AAbb 11 3 8
Segregating Sasi igen Bulg: 3c AaBB 15 3 12
Segregating 2—, 3=c, 3+ sue AaBb 44 5 39
Aabb
All 3—e, 3e aes ae aaBB 3 — 3
Segregating 3—c, 3+ as is aaBb 16 — 16
All 3+ 3 oes vis aabb 8 1 7
Total ae are ae 122 21 101

*A— Sr9b; B= Srl0

N. H. LUIG AND I. A. WATSON 319

In 1960, 91 F, plants of cross 1158.73.3 (Kenya 117A x Federation) were
studied for their behaviour to stem rust in the field. Of these plants 25 were
resistant or semi-resistant, the remaining 66 were semi-susceptible or susceptible.
A ratio of one resistant to three susceptible is indicated (P-value = 0-70-0-50),
and this is further evidence for a single major factor for adult plant resistance
in Kenya 117A. The semi-resistant and semi-susceptible plants carried
approximately 20 to 30 and 40 to 60% infection, respectively, and this variation
could be due to the action of the modifying genes Sr7 and Sr10 in combination
with Sr9b in the heterozygous condition.

4. Mentana W 1124
(i) Inheritance of the resistance in Mentana to strain 21—Anz—2 of stem rust.

F, Studies
When tested with strain 21—Anz—2 Mentana gave a _ semi-resistant
““;2, 3—c” reaction type. F, seedlings from crosses of Mentana with sus-
ceptible varieties gave a type ‘‘3—ce, 3c” reaction with this strain thus
indicating incomplete dominance of resistance.

F, and F, Studies

The segregation of reaction in F, plants to strain 21—Anz—2 in crosses of
Mentana with Yalta and Federation are given in Table 20. A single, incompletely
dominant factor pair for resistance in Mentana is indicated, as the data fit a
ratio of one resistant: two intermediate: one susceptible. The resistant F,
seedlings gave a type “;2, 3—c” reaction similar to that of Mentana, the
intermediate exhibited a ‘‘3c” reaction type, and the susceptible were as
susceptible as Yalta or Federation.

TABLE 20
F, segregation to strain 21—Anz—2 of crosses involving Mentana and susceptible varieties

Susceptible Cross No. F, Segregation P-value P-value
parent and family —_— (3: 1) (ils 4 2 Il)
Resist. Inter. Suscept. (R +1:S)
Yalta I156.4.1 19 4] 22 0:70 0:90-0:80
Federation I158.86.1 65 112 52 0-50—0-30 0:50-0:30
Federation 1158.86.2 68 133 61 0-70-0-50 0:90-0:80
Total 152 286 135 0:50-0:30 0-70-0-50

Two crosses between Mentana and susceptible varieties were studied in the
F, generation for their reaction to 21—Anz—2 (Table 21). The data fit a
1:2:1 ratio and thus provide further evidence for monofactorial segregation
in crosses between Mentana and susceptible varieties. Recent work has
indicated that this factor is the same as Sr8 (Watson and Luig, 1963).

TABLE 21

Segregation of F, lines of crosses between Mentana and susceptible Federation and Yalta for
reaction to strain 21—Anz—2

F, Segregation P-value
Cross SS (3B ei)
Resistant Segregating Susceptible Total
Federation x Mentana 12 12 8 32 0:30-0:20
Yalta x Mentana 14 31 11 56 0-70—0-50

Total 26 43 19 88 0-70-0-50

320 RESISTANCE TO PUCCINIA GRAMINIS VAR. TRITICI
(ii) Inheritance of resistance in Mentana to strain NR-7.

F, Studies

Mentana is fully resistant to this strain and shows a ‘“‘;” reaction type.
Yalta is susceptible and Federation and Chinese Spring moderately susceptible
(‘3 + en” and “3” type reactions respectively). F, seedlings of crosses
between Mentana and the latter three varieties when tested with NR-7 gave
“X, 3—c” reaction types, thus indicating that resistance was incompletely
dominant.

F, and F, Studies

The F, and F, data from a cross between Yalta and Mentana are set out
in Table 22, and they can be best explained on the basis of two independent,
incompletely dominant factors for resistance present in Mentana. A high
correlation between F, and F; reaction was obtained in spite of the incomplete
dominance of resistance indicated by F, tests. Hight distinct F, behaviour
patterns were observed and genotypes were assigned to them by assuming
that Mentana possesses two factor pairs for resistance. One factor pair con-
ditions an ‘‘ X =” reaction in the homozygous and an ‘“‘ X +” reaction in
the heterozygous state, while the second produces a moderately resistant
reaction of a “2, 3—c” type in the homozygous and a semi-susceptible
“3c” reaction type in the heterozygous state. The statistical analysis shows
close agreement between the observed numbers of plants and those expected
on this hypothesis.

TABLE 22

Correlated data showing reaction to strain NR-7 of F, plants of the cross Yalta (susceptible) x
Mentana (resistant) and of fF, lines raised from the F, plants

Behaviour and reaction in F3;

Number Number
Reaction of F, Segreg. Segreg. Segreg. Segreg. of F,
in Ia plants Bele Giotien), Go5 ego apa, Maes io Samm pe can. o5 lines
and 0 SS BG and and
COX = 2 66 3-¢c ” 66 Bae ” 66 3+ >
«or 3 ES = a hays i su 1 ne 1
asl e— la 9 7 = 1 1 == = = = 9
nasil 22 il 14 2 1 2 1 == = 21
RD ae 51 — 4 2 8 25 3 7 — 49
08 Be 24 = — = 1 12 — 6 4 P23)
“3+” 11 —— = = = 5 = 2 2 9
Total 120 8 18 5 1 44 4 16 6 112
Suggested CeDd
genotype CCDD CCDd CCdd CeDD Cedd ceeDD ecDd ecdd
Expected number
of F? lines 7 14 7 14 42 7 14 7 112
; x2 = 4-336 P-value (7 d.fr.) = 0:80-0-70

Further evidence for two factor pairs for resistance to NR-7 in Mentana
was obtained in a cross with susceptible Charter (Table 23). The F, and F,
segregations of this cross followed very closely those observed in the cross
where Yalta was the susceptible parent, and again a non-significant y?-value
for the eight segregation classes was obtained.

In order to obtain lines which carry only one of the two postulated major
genes for resistance to NR-7, two lines from the cross (Yalta x Mentana) were
selected: ‘1841’ apparently homozygous for a factor pair giving a ‘““3—e¢”
reaction, and ‘1843’ which appeared to carry a factor pair conditioning a
‘““X =” reaction. A cross was made between these two lines and each line
was also crossed with the moderately susceptible Federation.

The F, plants of these crosses when tested with NR-7 gave a reaction
similar to that of F, plants from crosses between Mentana and susceptible
varieties. In the following year F, populations from these crosses were tested

N. H. LUIG AND I. A. WATSON 321

TABLE 23

Correlated data showing reaction to strain NR—17, of F, plants of the cross Charter (susceptible) x
Mentana (resistant) and of F, lines raised from the F, plants

Behaviour and reaction in Fs
SEE SRE INO

Reaction of F, Segreg. Segreg. Segreg. Segreg. of F;
in ins plants “ce 1= 2”? ee 39 to ee x= 93 66 : 39 to “é 8 39 to 66 2 ae 66 ) 29 to ce pee 29 lines
ee x= 39 ee 3-¢ 29 ce 38+ 9 ee 3-¢ 99 €e 38+ 39
aie Bee 30 12 14 2 2 — —= = oad 30
“1+” 36 — 7 2 17 7 1 2 — 36
$5 Xoo 81 — 2 2 6 45 6 10 == al
7 Be 6 —— —_ — = 1 = 4 6
6e 3 29 i, == — = eee ax Ae 4 5
Total 160 12 23 6 25 54 7 13 8 148
Suggested CeDd
genotype CCDD CCDd CCdd CeDD Cedd ecDD ceeDd ecdd
Expected number
of F; lines 9-25 18-5 9-25 18-5 55-5 9-25 18-5 9-25 148
98 = Sil P-value (7 d.fr.) = 0:50-0-30

with NR-7 and 21—Anz-2 (Table 24). A single factor segregation was indicated
in cross (‘1841 x Federation) with both strains. F, seedlings of cross
(“1843 ” x Federation) segregated in a ratio of approximately three resistant
to one susceptible when tested with NR-7, but were all susceptible to strain
21-Anz-2. F, tests with strain NR-7 on cross ( 1843” x ‘“ 1841’) gave
a Similar segregation pattern to that obtained previously in F, populations
from crosses between Mentana and susceptible varieties. These findings suggest
that lines “‘ 1841 ” and ‘‘ 1843 ” together possess the full resistance of Mentana
to NR-7 and that each line carries a single factor pair for resistance to it.
‘1841 ” carries the factor pair Sr8 which also gives protection from 21—Anz—2
in the seedling stage, and ‘‘ 1843 ” possesses the factor pair which confers the
higher type of resistance to NR-7 (“;1 + 3—n” reaction) but no resistance
to 21-Anz-2. This factor is tentatively designated Sry.

TABLE 24

Reactions of F, seedlings of crosses involving the two resistant lines “ 1841” and “‘ 1843” when
tested with strains 21—Anz—2 and NR-7

F, Segregation and reaction
Parents Strain Ratio P-value
used. 8 Opell Om EXGR | (2 Cle te Ca ORC mee Ch you at=
pom X+ 3c gte

1843 X Federation 21-Anz-2 — cr
Bey Cha ee

1843 x Federation NR-7 — — — 16 — 1:2:1 0:20-0:10
1841 x Federation 21—Anz-2 — = —= — — — 47 69 40 1:2:1 #£0-30-0-20
1841 x Federation NR-7 — — 56 91 54 1:2:1 £0-50-0-30
1843 x 1841 NR-7 6 18 20 9 25 23 13 15 LO sa! 0:70-0:50

Proposed genotype of F, plant CCDD CCDd CeDD CCdd CeDd Cedd ecDD eceDd eccdd

B. The linkage relationship of genes controlling reaction to stem rust

As indicated in the foregoing, several factors for resistance to stem rust
were found in the six varieties under study. The possibility of linkage between
these factors was considered and the appropriate tests were carried out. Results
from F,, and F, generation material of crosses between Hureka and the varieties
Gabo, Yalta and Charter are shown in Tables 25 and 26. Hureka carries a
single gene for resistance (Sr6) to strain 126—Anz—6 on chromosome XX (2D)
and Gabo, Yalta and Charter all carry the gene Sri1l for resistance to strain
126—Anz-6 on chromosome X (6B). In certain instances Srll is differentially
transmitted, but this was not the case in crosses with Eureka. The statistical
analysis showed that the data agreed with the hypothesis of two dominant
independent factors (Tables 25 and 26).

F

322 RESISTANCE TO PUCCINIA GRAMINIS VAR. TRITICI

TABLE 25
Segregation in F, populations of crosses between resistant varieties

Strain F, Segregation Expected
Cross used. ratio P-value
Resistant Susceptible

Gabo x Eureka. . .. 126—-Anz—6 135 6 15:

1 0:50-0:30
Eureka x Yalta.. .. 126—Anz-—6 215 14 15: 1 0:95-0:90
Charter <x Eureka .. 126—-Anz-—6 469 30 15:1 0:90-0:80
Kenya 117A x Eureka 21—Anz-2 304 2 255:1 0:80-0:70
Gabo x Eureka.. .. l10Q3—H-2* 244 13 15:1 0:50-0:30

* The testing was conducted at temperatures above 80°F.

No evidence of linkage was found between the gene Sr6 in Eureka and
the genes for resistance to 21—-Anz—2 in Kenya 117A (Table 25). It has been
postulated that three factors operate in Kenya 117A against 21—Anz-2, of
which two, Sr7 and Sr9b, have been located by previous workers on chromo-
somes VIII (4B) and XIII (2A) respectively.

The close linkage of Sr1l with a factor for leaf rust resistance in Mentana
was discussed earlier. In Table 26 it is shown that Srll in Yalta and the
factor for resistance to strain 103-H—2 in Eureka, tentatively designated Srm,
were inherited independently of each other. No linkage was found between
Srg; and the factor for resistance (Srge) to 103—H-2 in Gabo (Table 25).

TABLE 26
Reaction of F'; lines of the crosses (Eureka x Yalta) and (Gabo x Eureka) when tested with strains
126—Anz-1, 6 and 126—Anz—2, 6 and of the cross (Hureka x Yalta) when tested with strains
126—Anz—1, 6 and 103—H-2

Reaction to 126—Anz-—l, 6 P-value
Kureka x Yalta 50% (4 d.fr.)
Resist. Segreg. Suscept. Total
Reaction Resistant 10 20 9 39
to Segregating 19 32 24 75
126—Anz-—2, 6 Susceptible 8 27 12 47
Total 37 79 45 161 2-597 0-70—0-50
Reaction to 126—Anz-—l, 6 P-value
Gabo x HKureka x? (4 d.fr.)
Resist. Segreg. Suscept. Total
Reaction Resistant 10 25 10 45
to Segregating 25 55 26 106
126—Anz-—2, 6 Susceptible 14 23 11 48
Total 49 103 47 199 0-903 0-95-0-90
Reaction to 103—H—2* P-value
Eureka x Yalta x? (4 d.fr.)
Resist. Segreg. Suscept. Total
Reaction Resistant 2 2 1 5
to Segregating 5 4 4 13
126—Anz-—1, 6 Susceptible 3 9 3 15
Total 10 15 8 33 2-689 0-70—0-50

* The testing was conducted at temperatures above 80°F.

N. H. LUIG AND I. A. WATSON 323

The two independent factors for resistance to NR-7 in Mentana, Sr8 and
Sryi, did not show any linkage with Sril in the cross (Yalta x Mentana).

The three genes for stem rust resistance in Kenya 117A appeared also to
be inherited independently of the following genes: Srll and Srge in the
crosses (Gabo x Kenya 117A) and (Kenya 117A x Gabo); and Sr8 in the
eross (Kenya 117A x Mentana).

A study was also made of the segregation of two morphological characters,
brown chaff colour and glume pubescence, in relation to stem rust reaction.

TABLE 27

Probable genotypes of six varieties of wheat regarding genes for resistance to stem rust

Variety Genotype
Eureka Sr6 Srp
Gabo Srll Srgi Srge Sres
Charter Srll Srge Srgs
Yalta Srll
Kenya 117A Sr7 Sr9b =Srl0
Mentana Sr8 Sry

The single factor for pubescent glume in Yalta was inherited independently
of the following genes for stem rust resistance: Sr6, Sr8, Sr9b, Srl1, Sree
and Sryi. The factor for pubescent glume has been located on chromosome
XIV (1A) (Sears, 1953).

The single factor for brown chaff in Eureka was inherited independently
of Srll and of Sr6. Unrau (1950) has located a single gene for brown chaff
colour on chromosome I (1B).

The following Tables 27 and 28 summarize the results of these studies as
to the genetic constitution of the six varieties and as to the nature of the
resistance conferred by the different genes.

TABLE 28

Nature of resistance conferred by eleven genes possessed by six wheat varieties

Gene Dom.
or
Rec.*
Sr6*** D
Srz i d
Srll D
Srgi r
Srg 2 D
Srg3 D
Sr7 d
Sr9b d
Srl0 d
Sr8 r
Sry r

Possessed.
by
Variety

Eureka

EKureka

Gabo
Charter
Yalta

Gabo

Gabo
Charter
Gabo
Charter
Kenya 117A
Kenya 117A

Kenya 117A

Mentana
Mentana

Controls resistance to strain**

21—Anz-—0, 21—Anz-2, 21—Anz-2, 6,
1038—H-2, 111—E-2

103—H-—2

21—Anz-0, 126—Anz-—6, 126—-Anz-—l, 6

21—Anz-—0, 21—Anz—2, 126—Anz-—6
103—H-2 and 111—EK-2

A20

21—Anz-—0, 21—Anz—-2, 21—Anz-2, 6

126—Anz-—6, NR-7,

21—Anz—0, 21—Anz—2, 21—Anz-2, 6, 126-Anz—6, 126—Anz-—l1, 6,
126—Anz-—2, 6, NR-7, 222-Anz—1, 2, 4, 6
21—Anz-—0, 21—Anz—2, 21—Anz—2, 6, 126-Anz—6, 126—Anz-—1, 6,

126—Anz—2, 6, NR-7
21—Anz-—0, 21—Anz—2, NR-7
NR-7

* D = dominant; d= incompletely dominant; r= recessive.
** Includes only the eleven strains listed in Table 1.
*** 'Temperature-sensitive.

324 RESISTANCE TO PUCCINIA GRAMINIS VAR TRITICI

DISCUSSION

The results of the studies on the nature of resistance to stem rust in six
wheat varieties reported herein show that single independent factors were
operating to individual strains. In certain cases the segregation ratios were
distorted by a differential transmission rate for the gametes carrying genes
located on chromosome X (6B). The segregation ratios were also altered by —
genes which had a cumulative effect in modifying the dominant or recessive
condition and by temperature effects changing the expression of the factors
for resistance.

The latter was the case with the gene Sr6 of Eureka, which becomes in-
effective at temperatures above 80°F. Hence when Eureka is crossed with
susceptible varieties and the progenies are studied at low temperatures
(60° to 65°F) an F, ratio of three resistant to one susceptible is obtained with
strains 21—Anz—2, 126-Anz-6 and NR-7. At higher temperatures of 65° to
70°F the heterozygous class can be distinguished from the resistant class when
testing with 21—Anz—2. The intermediate, heterozygous F, seedlings still give
a necrotic reaction, but the type “;1 =” reaction has changed into a type
“1 +4+,3—n” reaction. A similar segregation pattern is obtained with
strains 126-Anz-6 and NR-7 at temperatures of 70° to 75°F. At these
temperatures, however, it is no longer possible to decide between intermediate
and susceptible seedlings when testing with 21—-Anz-2. With this strain the
initial ratio of three to one has thus changed into a one to three ratio, with
the resistant plants giving the same reaction at 70° to 75°F as the intermediates
at temperatures of 65° to 70°F. At temperatures above 75°F, F, seedlings
are all moderately susceptible or susceptible to 21-Anz—2, and at temperatures
above 80°F the gene Sr6 is no longer effective against 126—-Anz-6 or NR-7.
Hence when testing at temperatures of 70° to 75°F with strains 21—Anz-2,
126—Anz-6 and NR-7 it can be assumed that the gene Sr6 is recessive with
21—Anz—2 and incompletely dominant with the latter two strains. Such has
been suggested by Knott and Anderson (1956), who found that Sr6 was
dominant with race 56 but recessive with race 15B. The reaction types recorded
on Eureka to seven strains listed in Table 1, at various temperatures, suggest
that genetic material in which Sr6 segregates would behave similarly to strains
21—Anz-0, 21—Anz—2 and 21—Anz—2, 6. With the other four strains, 126—Anz-—6,
126-Anz-1, 6, 126—Anz—2, 6, and NR-7, the same single factor is operating,
but with a somewhat higher degree of resistance.

No differential transmission rate exists for alleles at the Sr6 locus. Hureka
possesses no other factors for resistance to Australian field strains. However,
when an avirulent laboratory strain, 103—H—2, which had its origin in a somatic
cross between P. graminis var. tritici and P. graminis var. secalis, was used,
segregation of a second, incompletely dominant factor pair was evident.
Whether this factor is identical with the second factor for resistance in Eureka,
as reported by Athwal (1955), cannot be ascertained. Athwal worked with
race 42 from India, and he found that the varieties Bencubbin, Mentana, Dundee,
Uruguay and Gabo, which are susceptible to several Australian strains, were
resistant to race 42. Strain 103-H-2 is also non-pathogenic on these five
varieties. Hence, it is likely that the same factor in Eureka conditions
resistance to 103—H—2 and to race 42.

The gene Srll, possessed by Gabo, Charter and Yalta, has been shown
to be differentially transmitted in crosses between each of these three varieties
and certain other varieties including Chinese Spring. This finding explains
why several investigators using the latter variety as the susceptible parent
postulated two, linked, dominant complementary factors for this type of
resistance, while other workers who used different susceptible varieties reported
a single factor pair. Further results on, and several aspects of, differential
transmission have been discussed elsewhere (Luig, 1961; 1964).

N. H. LUIG AND I. A. WATSON 325

While it has been known for many years that Gabo, Charter and Yalta
carry the same resistance (Sr11) to particular Australian field strains (Watson
and Waterhouse, 1949), it was also noticed that when these strains were present
in the field Gabo was not as severely attacked as Charter, and Charter was less
affected than Yalta (Waterhouse, 1952). It was thought that the early maturity
of Gabo enabled this variety to escape infection to a certain extent and, while
this may be so, the present study indicates that minor factors can be important
once the major factor for resistance is rendered ineffective. Gabo carries one
minor factor, tentatively designated Srg,, for resistance to Australian field
Strains. This factor gives a ‘““3—c” type of reaction in the seedling stage.
Charter and Yalta do not possess this minor factor, but it is present in Bobin
39 and Gular.

To strain 103-H-2, seedling resistance of Gabo was governed by a single
factor, Sree, which is distinct from the two previously mentioned factors in
this variety. Srg: is also present in Charter, but not in Yalta. Whether Sree
is identical with one of the two factors for resistance found in Gabo by Athwal
(1953) cannot be ascertained as Athwal used Indian race 42, as mentioned
earlier.

When strain A20 of P. graminis var. secalis was employed, a fourth factor
for resistance, Srg3, was found in Gabo and this factor was also present in
Charter but not in Yalta. Thus, when testing with strains 126—Anz-—6, 103—H-2
and A20, no segregation for susceptibility occurs in crosses between Gabo and
Charter, but this is in each case due to a different resistance factor. The type
of seedling reaction of Gabo and Charter to these three strains, however, is
very similar. Yalta carries only Srll and is susceptible in the seedling stage
to 103—H-2 and A20.

Results on the resistance of Kenya 117A in the seedling and adult plant
stage are in agreement with those reported by earlier investigators (Watson
and Waterhouse, 1949; Athwal, 1953; Athwal and Watson, 1954; Knott and
Anderson, 1956). During the present investigations it was found that the
major gene for seedling resistance in Kenya 117A, Sr9b, was the same as that
which gives resistance in the mature plant stage. The other two factors,
presumably Sr7 and Sr10, had only a modifying influence under Australian
field conditions. The modifying effect of these genes was evident when the
field reactions of F, seedlings from crosses of Kenya 117A and Gamenya* with
susceptible varieties were compared. Gamenya apparently carries the factors
Sr9b and Srg; only, the latter factor derived from Gabo and having no influence
on field reaction. Because Gamenya does not carry Sr7 or Srl0, F, seedlings
from crosses with Gamenya are moderately susceptible in the field, while those
from crosses with Kenya 117A are intermediate. It was also found that with
most strains the genes of Kenya 117A were incompletely recessive in seedlings
and in adult plants. The effect of these genes was additive rather than
epistatic.

The three genes, when homozygous, produced different reaction types
when tested with different strains. This result is in agreement with those of
Green et al. (1960). Sr9b was very effective against strain 222—Anz-1, 2, 4, 6
at all temperatures, but lines carrying either Sr7 or Sr10 were moderately
susceptible when tested with this strain. At temperatures of 65° to 70°F lines
possessing Sr9b only were more resistant to 126—Anz-—2, 6 than to 21—Anz-2
and NR-7, but became increasingly susceptible at higher temperatures. Lines
_ which carried either Sr7 or Srl10 were more resistant to 21—Anz—2 than to
126—Anz—2, 6 and NR-7 to which they gave identical reaction types. The
data suggested that Sr7 gives less protection than Sr10 under Australian
conditions. Srl0 would be a valuable gene in combination with other genes,
but it is ineffective on its own. As the seedling reaction produced by this

* Bred from a cross Gabo x [(Gabo®> x Mentana) x (Gabo? x Kenya 117A)].

326 RESISTANCE TO PUCCINIA GRAMINIS VAR. TRITICI

gene is easily discernible from a susceptible type there should be no difficulty
in incorporating it into varieties. Although ineffective in adult plants in the
field, and hence useless on its own for breeding, Sr10 is apparently operative
in the seedling stage against all Australian strains of stem rust. Plants
possessing Sr9b, by contrast, react differentially and are rendered ineffective
by several strains now well established in the field, e.g., 17 — 2, 3, 21 — 2, 3,
21 —1, 2, 3, 21— 2, 3, 4, 21— 2, 3, 6 and 116 — 2, 3 (Watson and Luig,
1963).

These studies further demonstrate that the strains of stem rust used in
this study can be grouped as follows when plants having specific genes from
Kenya 117A are inoculated with them :

Group I : 21—Anz-—0, 21—Anz—2, 21—Anz-—2, 6
Group II : 126—Anz-6, 126—Anz-1, 6, 126—Anz-2, 6
Group III: 222—Anz-1, 2, 4, 6

Group IV: NR-7

Group V_: 21—Anz-2, 3

The results from studies on the inheritance of resistance in Mentana to
two strains of stem rust can be interpreted on the basis of two apparently
independent factor pairs. Against most Australian field strains, ike 21—Anz—2,
a single factor, Sr8, conferred resistance in the seedling stage, this resistance
being of a chlorotic ‘“‘2—” reaction type. The same factor was operative
against NR-7, but Mentana also possessed a second factor which conditioned
a necrotic type ““ X =” reaction to this laboratory strain. The combined
effect of these two factors, Sr8 and Sry;, was to make Mentana practically
immune at low temperatures (‘‘0;” reaction type).

Acknowledgements

The authors would like to thank Miss Wendy Ball and Mr. W. Hamlyn
for their technical assistance. Financial assistance is also acknowledged from
The Wheat Industry Research Council and the Rural Credits Development
Fund of the Commonwealth Bank.

References

Astam, M., and AusEmus, EH. R., 1958.—Genes for stem rust resistance in Kenya Farmer wheat.
Agron. Journ., 50: 218-222.
ATHWAL, D. S., 1953.—Gene interaction and the inheritance of resistance to stem rust of wheat.
Ind. J. Genet. and Pl. Breed., 13: 91-103.
, 1955.—The resistance of some wheat varieties to physiologic race 42 of Puccinia
graminis tritict. Ind. J. Genet. and Pl. Breed., 15: 80-87
, and Watson, I. A., 1954.—Inheritance and genetic relationship of resistance possessed
by two Kenya wheats to races of Puccinia graminis tritici. Proc. Linn. Soc. N.S.W.,
79: 1-14.

; , 1957.—Inheritance of resistance to wheat leaf rust in Mentana, a variety

of Triticum vulgare. Proc. Linn. Soc. N.S.W., 82: 245-252.

Forsytu, F. R., 1956.—Studies on the nature of rust resistance in cereals. Report, Third Int.
Wheat Rust Conf., Mexico D.F., Mexico, March 18-27, 1956, pp. 130-131.

GREEN, G. J., and Jounson, T., 1954.—Effect of high temperature on the reaction of adult
wheat plants to stem rust. Proc. Canad. Phytopath. Soc., No. 22: 13-14 (Abst.). Cited
P.B.A., 25: 1902.

, Knott, D. R., Watson, I. A., and Puastey, A. T., 1960.—Seedling reactions to stem
rust of lines of Marquis wheat with substituted genes for rust resistance. Can. J. Plant
Sci., 40: 524-538.

Kwott, D. R., 1959.—The inheritance of rust resistance. IV. Monosomic analysis of rust
resistance and some other characters in six varieties of wheat including Gabo and Kenya
Farmer. Can. J. Plant Sct., 39: 215-228.

, and ANDERSON, R. G., 1956.—The inheritance of rust resistance. I. The inheritance
of rust resistance in ten varieties of common wheat. Can. J. Agr. Sci., 36: 174-195.

Luie, N. H., 1960.—Differential transmission of gametes in wheat. Nature, 185: 636-637.

——., 1961.—_The inheritance of disease resistance in common wheat. Thesis (Ph.D.),
The University of Sydney.

, 1964.—Heterogeneity in segregation data from wheat crosses. Nature. 204: 260—
261.

N. H. LUIG AND I. A. WATSON 327

Mactnpog, S. L., 1948.—The nature and inheritance of resistance to stem rust of wheat, Puccinia
graminis tritici, possessed by several resistant parents. Thesis (Ph.D.), 1941, University
of Minnesota. Dept. Agric. N.S.W. Sci. Bull. 69.

Omar, A. A. M., 1959.—Inheritance of reactions to race 15B and some other races of stem rust
of wheat. Hgypt. J. Bot., 1: 1-17. Cited P.B.A., 29: 373.

PETERSON, R. F., and CAMPBELL, A. B., 1953.—Aneuploid analyses of the genes for stem rust
resistance and head density in MecMurachy wheat. Report, Int. Wheat Stem Rust Conf.,
Winnipeg, Canada, Jan. 5-7, 1953. 133 pp. (Mimeo.) Cited P.B.A., 24: 253.

Sears, E. R., 1953.—Nullisomic analysis in common wheat. Amer. Naturalist, 87: 245-252.

, and LorcErine, W. Q., 1961.—A pollen-killing gene in wheat. Genetics, 46: 897
(Abst.).
, and RopDENHISER, H. A., 1957.—Identification of chromosomes carrying
genes for stem rust resistance in four varieties of wheat. Agron. Jour., 49: 208-212.

StaxkMAN, EH. C., and Levine, M. N., 1922.—The determination of biologic forms of Puccinia
graminis on Triticum sp. University of Minnesota, Tech. Bull. 8.

Unrav, J., 1950.—The use of monosomes and nullisomes in cytogenetic studies of common
wheat. Scr. Agric., 30: 66-89.

WATERHOUSE, W. L., 1952.—Australian rust studies. IX. Physiologic race determinations and
surveys of cereal rusts. Proc. Linn. Soc. N.S.W., 77: 209-258.

Watson, I. A., 1941.—Inheritance of resistance to stem rust in crosses with Kenya varieties
of Triticum vulgare Vill. Phytopath., 31: 558-560.

, 1943.—Inheritance studies with Kenya varieties of Triticum vulgare Vill. Proc. Linn.
Soc. N.S.W., 68: 72-90.

, 1955.—The occurrence of three new wheat stem rusts in Australia. Proc. Linn.
Soc. N.S.W., 80: 186-190.

, 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 Luie, N. H., 1958.—Timvera—a Steinwedel x Triticum timopheevi derivative.
Agron. Jour., 50: 644.

———, ———., 1958b.—Somatic hybridization in Puccinia graminis var. tritici. PRoc.
Liyn. Soc. N.S.W., 83: 190-195.

; 1959. —Somatice hybridization between Puccinia graminis var. triticc and
Puceinia graminis var. secalis. Proc. Linn. Soc. N.S.W., 84: 207-208.

, 1961.—Leaf rust on wheat in Australia: a systematic scheme for the
classification of strains. Proc. Lryn. Soc. N.S.W. , 86: 241-250.

——_—, ————., 1962.—Selecting for virulence on wheat while inbreeding Puccinia graminis
var. secalis. Proc. Linn. Soc. N.S. W., 87: 39-44.

———, 1963.—The classification of Puccima graminis var. tritici in relation to
breeding resistant varieties. Proc. Linn. Soc. N.S.W., 88: 235-258.
, and Sineu, D., 1952.—The future for rust nagigen wheat in Australia. J. Aust.
Inst. Agric. Sci., 18: 190-197.
, and Strwart, D. M., 1956.—A comparison of the rust reaction of wheat varieties
Gabo, Timstein, and Lee. Agron. Jour., 48: 514-516.
, and WatERHOUSE, W. L., 1945.—A third factor for resistance to Puccinia graminis
tritict. Nature, 155: 205.
———.,, 1949.—-Australian rust studies. VIII. Some recent observations on wheat
stem Trust in Wastralia, Proc. Linn. Soc. N.S.W., 74: 113-131.

THE DISTRIBUTION OF SUBMERGED AQUATIC ANGIOSPERMS
IN THE TUGGERAH LAKES SYSTEM

F. R. HIGGINSoN
School of Biological Sciences, Botany Building, University of Sydney

[Read 24th November, 1965]

Synopsis
The submerged communities of Tuggerah Lakes are dominated by the aquatic angiosperms,
Zostera capricornt Aschers and Ruppia spiralis Dumort. The distribution of these species has
been mapped and an attempt made to determine the ecological factors underlying their
distributional pattern.

INTRODUCTION

The Tuggerah Lakes, a system of maritime coastal lakes covering a total
area of 24 square miles, are located approximately 60 miles north of Sydney
on the central coast of New South Wales (Fig. 1). Physiographically, they
appear similar to other coastal lakes or lagoons of Hastern Australia, and have
apparently been formed by longshore currents building a series of sand bars
across an irregularity or indentation in the coastline (Hutchinson, 1957). This
has resulted in eastern foreshores of a coastal sand-dune character, whereas
the foreshores on the western perimeter are of typically sandstone-derived soils.

Lake level and salinity, two important aspects of the submerged environ-
ment, are considerably influenced by the influx of water from streams, and
evaporation from the lake itself. In dry seasons, little fresh water enters the
lake from the streams, and with continued evaporation the salinity may rise
as high as 3:1%. Conversely, in times of heavy precipitation, the salinity
may be as low as 0:5%. Table 1 illustrates these effects in Lake Budgewoi.

By far the most important type of current in the lakes is that caused by
winds, the most effective being from the south-east and north-east, in which
directions the lakes are relatively exposed. The influx of streams also causes
localized turbulence, due to the mixture of water of different densities ;
however, this is generally of secondary importance. There is no tidal effect,
as a result of an extremely small oceanic connection and a relatively large
amount of water entry from creeks and streams.

A combination of shallow depth and abundant supply of nutrients from
the catchment area has allowed the development of a very extensive macro-
flora in the lakes. Table 2 lists the major plants of the submerged communities.
Of these species, Ruppia spiralis and Zostera capricorni occupy at least 80%
of the colonized areas, and therefore most of the following discussion will be
concerned with their distribution.

PREVIOUS STUDIES

Many workers have published accounts of the distribution of submerged
plants in freshwater lakes (Misra, 1938; Pearsall, 1920; Penfound, 1953;
Pond, 1905) but there are few published accounts of the ecology of brackish
water environments, particularly under Australian conditions.

Ferguson Wood (1959a, 1959b), working in Lake Macquarie, found that
Zostera is usually confined to water of marine character, although it may at
times encroach into less saline waters, and is found from low tide mark to a

PROCEEDINGS OF THE LINNEAN Society or New SoutH Watss, Vol. 90, Part 3

F. R. HIGGINSON 329

depth of 20 feet. He also observed that it appears to require good illumination
and can persist in currents of several knots. Ruppia, on the other hand, is
generally found in waters of low salinity, but does extend into more saline
areas where it is associated with Zostera. Ruppia is also observed to require
good illumination, and to be intolerant of strong currents.

Ferguson Wood (1959a) includes adequate descriptions of the important
marine angiosperms; therefore, no attempt will be made to cover their
taxonomic aspect in this paper.

FIGURE 1.

Y
’ MUNMORAH ,
33° 12

4
BUDGEWOI! .
Toukley. Norah Head,

TUGGERAH .

12
yy
3

Fig. 1. Bathymetric map of the Tuggerah Lakes system (depth contours in feet), showing
locations mentioned in text. The system can be seen to consist of three lakes: Tuggerah,
Budgewoi and Munmorah.

THE ENTRANCE.

1 mile
Scale :-

ANGIOSPERM DISTRIBUTION—OBSERVATIONS

The distribution of the three major submerged angiosperms in Tuggerah
Lakes is shown in Figure 2. This map is based on that compiled by the
Electricity Commission of N.S.W. from aerial photographs (November, 1962),
supplemented by skindiver transect: observations in 1963. The distribution of
Halophila ovalis is seen to be restricted to shallow, sandy areas to a depth of
approximately three feet.

From general observations at collecting sites, the only characters of the
water which vary significantly are salinity and light intensity (determined by
depth and turbidity). Although salinity decreases with distance from the
ocean entrance it does not appear to affect the distribution of Zostera within

330 SUBMERGED AQUATIC ANGIOSPERMS IN TUGGERAH LAKES SYSTEM

the lake. For example, Zostera is found 400 yards from the ocean at the
Entrance where salinity approaches that of sea-water (3-4% 8S), and also in
Munmorah Lake, 10 miles from the ocean, where the salinity may be as low
as 0°3%. Likewise, the distribution of Ruppia is not greatly affected by
salinity variations as it has been found over a salinity range of 0-02% to 3-1%,
although it does not appear to grow as well in strongly saline waters.
Current has a noticeable effect upon plant distribution, Ruppia being more
prevalent in lentic situations such as sheltered bays, whereas Zostera can

FIGURE 2.

=== ws
=

(

1 mile
Scale =

Fig. 2. Map of Tuggerah Lakes system showing distribution of submerged angiosperms.

withstand relatively strong currents, for example, the currents of the ocean
entrance and in the constriction between Lake Budgewoi and Tuggerah Lake

at Toukley (see Fig. 2).

The plants are generally found at an average depth of five feet, and very
rarely at depths exceeding nine feet. Ruppia grows quite well in water from
two to eight feet, whereas Zostera may occasionally be found in depths up
to 13 feet.

Although it has been observed that depth and current have a marked
effect upon the distribution of Ruppia and Zostera, these factors alone are
clearly not adequate to explain differences in distribution which exist at the
same depth, or in similar flow areas. Thus other factors must be examined,
the most important being the character of the lake bed.

F. R. HIGGINSON Dol

THE SEDIMENT FACTOR
Methods and Resulis
Sediment samples were collected at 50 sites within the Tuggerah Lakes,
observations on plant growth being taken at each site. A ‘ torpedo” core
sampler was used to collect the surface three inches of the sediment.

The sediment samples were analysed for:

(i) % organic matter by a dichromate wet oxidation method (Tinsley, 1950) ;

(ii) % total nitrogen by the Kieldahl method ;

(iii) % total calcium

(iv) % total potassium | Extraction by Hall’s method (Piper, 1942) and

(v) % total magnesium -estimation of individual cations by Atomic

(vi) % total iron absorption spectroscopy.

(vii) Mechanical analysis (size distribution of particles) by the Bouyoucos
hydrometer method, using the International convention of particle
diameter limits.

All analytical results are expressed on the basis of oven-dry matter (105°C).

The sediment samples were divided into five groups depending on the
type of plant cover, and the mean results for each group presented in Table 3.

Discussion

The mechanical analysis shows that there are three major types of
sediments: sandy, clayey and intermediate. All the sandy sediments occur
close to the lake perimeter, whereas the clayey sediments occur in the centre
or in sheltered places. This zonation of sediment type can be explained if
we assume that initially the lake floor was sandy, and that subsequently large
quantities of soil material have been brought into the lake by fluvial erosion.

TABLE |
The effect of precipitation upon lake-water level and salinity in Budgewoi Lake (Toukley)

Surface water Precipitation* Lake level at Salinity% + Salinity% tf

Month temperature on catchment sampling site lake water sea water
(°C.) (points) (ft.)

1963 May 15-0 1377 8-0 0-84 3-37
June 12-5 811 7-5 0-54 3-28
July 11-0 216 7:5 0-99 3:34
Aug. 12-0 703 8-0 1-11 3-28
Sept. 15-5 153 7-0 1-84 3-44
Oct. 20-0 142 6:8 2-31 3-44
Nov. 21-0 227 6-5 2-54 3-46
Dec. 21-5 443 6-3 2-79 3-49

1964 Jan. 21-5 186 8-0 2-97 3-50
Feb. 22-0 114 8-0 3-10 3-50
March 21-0 658 8-0 2-63 3-50
April 19-5 424 8-0 2-32 3-49
May 16-5 227 6:5 2-79 3-50

* Precipitation measured in a standard 8” gauge at Munmorah.
} Salinity values calculated from chlorinity titrations, according to Harvey (1957).
t Sea water values inserted for comparison; samples collected at Norah Head.

The smaller particles of this erosional material would tend to be deposited in
sheltered positions or in areas of greater depth, whereas the coarse material
would be deposited near the stream mouths. Consequently, the substratum
would become composed of coarse materials in shallow water, and of progressively
finer particles as depth increases.

The results show that Zostera tends to favour the sandy sediments and
Ruppia the clayey sediments, the intermediate sediments having a mixed

332 SUBMERGED AQUATIC ANGIOSPERMS IN TUGGERAH LAKES SYSTEM

community (see Table 3). Differences, however, in the size distribution of
particles of the substratum do not seem sufficient to explain differences in
associated vegetation unless this can be considered in terms of their possible
effects on plant growth.

In all but a few of the 50 soil samples, an increase in clay content was
associated with an increase in organic matter regardless of vegetation. There-
fore Ruppia, as well as being associated with sediments of higher clay content,
is normally associated with higher organic matter levels than Zostera. Where

TABLE 2
Inst of the major planis of the submerged communities in the Tuggerah Lakes

Classification Species name

i. Angiospermae Zostera capricornt Aschers.
Ruppia spiralis Dumort.
Halophila ovalis (R.Br.) Hook. f.

uu. Chlorophyceae Chaetomorpha linum (Muller) Kutz.
EHnteromorpha clathrata (Roth.) Grev.
Cladophora Kutz. sp.

ili. Charophyceae Chara

iv. Phaeophyceae Cystophyllum muricatum (Turn.) J. Ag.
Dictyota dichotoma var. implexa (Desf.) Gray.

v. Rhodophyceae Polysiphonia mollis Wook. and Harv.
Gracilaria verrucosa (Huds.) Papenfuss.

vi. Cyanophyceae Lyngbya majuscula Harv.

sandy sediments are deposited, peaty matter accumulates on the surface of
the substratum and often contains recognizable fragments of plant material,
decay and incorporation being extremely slow. Where clayey sediments are
deposited, the organic litter is rapidly decomposed and incorporated as humus.
This may be an explanation for the increase in mineral content as the percentage
of clay rises.

Results show that sediments high in organic matter are also high in total
content of estimated minerals except calcium. Ruppia, therefore, is associated
with substrata of high fertility and Zostera with those of lower fertility. The
calcium content is approximately the same in all colonized substrata, the highest
calcium level being in non-vegetated sands where there are accumulations of

TABLE 3*
Average results of sediment analysis from five vegetation classes

Vegetation of ta A % % % % %
class Coarse Clay Organic N Ca K Mg Fe
sand matter
Zostera growth only 71:5 13-9 2-31 0-11 0-26 0-10 0-11 0-42
Ruppia growth only 25-4 42-3 5:37 0-21 0:27 0-44 0-52 2°11
Zostera and Ruppia

together rs OG 36:8 6-02 0-27 0-35 0-29 0-50 1-62
Bare clays Bie Ae Ol 60-8 8-14 0-28 0-66 1-02 0-75 3-59

Bare sands (occasional
Halophila) .. a0 8357) 9-1 0-71 0-06 0-80 0-05 0-09 0-12

* Results condensed from Higginson (1963).
~ Coarse sand particle diameter limits: 2-0-0-2 mm.
{ Clay particle diameter limits: less than 0-002 mm.

F. R. HIGGINSON 333

marine skeletons. Plant colonization is observed- to be associated with a
reduction in total calcium content of the sediment, and this may be adequately
explained by conversion of calcium carbonate to soluble forms with subsequent
leaching from the substratum. The increase in carbon dioxide content of the
water, resulting from organic matter breakdown in the sediment, would enable
such a conversion to take place.

Average results of sediment analysis (Table 3) show that an increase in
clay content is associated with an increase in nitrogen content. Hence it is
apparent that a relationship exists between the mechanical properties and
the chemical composition of the substrata. The form of available nitrogen
in the substratum is important. Nitrates are normally absent from submerged
soils (Misra, 1938), nitrogen being available as ammonia. Much of the organic
nitrogen is ultimately converted to ammonium ions, and a large proportion
is adsorbed onto the colloid exchange complex of the sediment replacing other
cations. The cations exchanged would tend to go into solution and be more
or less concentrated in the mud where they diffuse into the aqueous system
or become available for plant absorption.

TABLE 4

Tne interaction of depth and sediment type wpon the distribution of submerged angiosperms in
Tuggerah Lakes

Vegetation Average depth Coarse sand Clay
ft. oF oY

Usually no growth, but with occasional

Halophila ovalis (sterile sands) 2-0 83-7 Goll
Zostera capricorn only .. ce igs 5:4 71:5 13-9
Zostera and Ruppia growing together .. 5-0 37-6 36-8
Ruppia spiralis only a Be a 6-5 25-4 42-3
No growth (bare muds) .. ls He 10-0 9-1 60-8

CONCLUSION

An increase in clay content of the sediment is associated with an increase
in organic matter, nitrogen and all estimated minerals except calcium. There
is, however, no evidence indicating that the greater fertility of sediments of
high clay content is due to chemical rather than physical characters, and it is
probable that, as in terrestrial soils, the two groups of factors cannot be
dissociated.

It seems clear from the results in Table 3 that there is a close relationship
between the nature of the sediment and the type of vegetation growing on it.
The fact that there are distinct zones of sediments with different physical
structure, and that these differ in chemical composition, justifies the assumption
that zonation of vegetation is a result of differences in sediment conditions.

On the basis of evidence presented, it appears likely that an interaction
of depth and sediment type can adequately explain the distribution of submerged
aquatic angiosperms in Tuggerah Lakes. A summary of investigations, presented
in Table 4, illustrates this conclusion.

Acknowledgements

The author wishes to acknowledge the assistance of the late Professor
R. L. Crocker in the early stages of this project, for without Professor Crocker’s
help, the project would not have commenced.

For their frequent assistance in sampling and for their observations, I
am indebted to Mr. B. Clough and members of the Projects Division of the
Electricity Commission of N.S.W.

334 SUBMERGED AQUATIC ANGIOSPERMS IN TUGGERAH LAKES SYSTEM

References

Evecrriciry Commission or N.S.W., 1962.—Investigation of the weed problem in Tuggerah
Lakes. Projects Division Report, October, 1962.

Frercuson Woop, E. J., 1959a.—Some East-Australian sea-grass communities. Proc. LInn.
Soc. N.S.W., 84: 218-226.

, 1959b.—Some aspects of the ecology of Lake Macquarie with regard to an alleged
depletion of fish: VI, Plant communities and their significance. Aust. Journ. Marine
and Freshwater Research, 10: 322-340.

Harvey, H. W., 1957.—* The Chemistry and Fertility of Sea Waters.”’ (Cambridge University
Press, Cambridge.)

Hieeinson, F. R., 1963.—B.Se.Agr. Honours Thesis. Botany Department, University of
Sydney.

Hurcatnson, G. E., 1957.—** A Treatise on Limnology ’’, Vol. I. (J. Wiley and Sons, New
York.)

Misra, R. D., 1938.—EHdaphie factors in the distribution of aquatic plants in the English lakes.
J. Hcol., 26: 411-451.

PEARSALL, W. H., 1920.—The aquatic vegetation of English lakes. J. Hcol., 8: 163-201.

PEenFounbD, W. T., 1953.—Plant communities of Oklahoma lakes. Ecol., 34: 561—583.

Prerer, C. S., 1942.—“‘ Soil and plant analysis.’ (Hassell Press, Adelaide.)

Ponp, R. H., 1905.—The biological relation of aquatic plants to the substratum. Repts
U.S. Comm. Fish., 29: 483.

TinsLtEy, J., 1950.—The determination of organic carbon in soils by dichromate mixtures.
Trans. 4th Int. Conf. Soil Sci., 1: 161-164.

NUMERICAL METHODS IN TAXONOMY

R. C. JANCEY*
Department of Biological Sciences, University of Sydney

[Read 24th November, 1965]

Synopsis

Some of the advantages of a numerical approach to taxonomy are indicated, also the
compatibility of the technique with phylogenetically based taxonomy. Two main avenues
for the application of computer techniques are described—the simplification of individual
relationships and the detection of group structure. Finally, a means of combining the results
of these two techniques is described.

The development of electronic computers has led to the introduction in
recent years of numerical methods to the taxonomic process. That the advent
of such methods has been the subject of some criticism cannot be denied, and
it is the author’s hope that this contribution will serve to dispel some mis-
apprehensions, and to indicate some of the facilities offered by differing forms
of numerical analysis.

Perhaps the foremost objection of many taxonomists to the introduction
of numerical methods is their doubt that any automatic process could replace
the extremely complex and flexible mental comparison of individuals and
attributes which forms the vital part of the taxonomic process. The second
objection is that the use of numerical methods precludes any phylogenetic
basis for the final classification, and is hence an essentially retrogressive step.
With regard to the first objection, assurance may be given that computers
are indeed capable of reproducing the results of mental classifications made
by taxonomists, so long as they are provided with the same or comparable
information. A number of methodological investigations of numerical tech-
niques have been carried out in which data supplied by monographers have
been subjected to numerical analysis, the resulting output being fully in accord
with the taxonomic decisions arrived at independently by the monographer
(Rogers and Fleming, 1964). In an investigation of the genus Phyllota
(Leguminosae) the author demonstrated by numerical methods a group
structure almost identical with one advocated by Bentham more than a century
ago, although in this case the characters used were almost certainly quite
different (Jancey, 1965).

The second objection to the use of numerical methods, that they preclude
a phylogenetic classification, is quite unfounded though widely held. Such a
situation may well be the result of a misunderstanding since, although a number
of numerical taxonomists hold strong views on the place of phylogenetic
considerations in classification, this in no way makes numerical techniques
and a phylogenetic classification necessarily incompatible. Assuming that the
term phylogenetic classification implies the interpretation and modification of
the groupings of present day phenotypically similar organisms in the light of
known or inferred evolutionary trends, then a number of observations may be
made concerning the methods by which such a classification can be achieved.
In the mental taxonomic process it is possible to keep evidence of evolutionary
trends in mind, and to modify taxonomic relationships even as they are being

* Author’s present address: The Department of Quantitative Taxonomy, New York
Botanical Garden, Bronx 58, New York, U.S.A.

PROCEEDINGS OF THE LINNEAN SociETY oF NEw SoutH WALEs, Vol. 90, Part 3

336 NUMERICAL METHODS IN TAXONOMY

constructed, or, alternatively, to construct first an essentially phenotypic
classification and then modify this in the light of such other evidence as may
be available. Both these possibilities are available with numerical techniques ;
in the case of concurrent consideration of evolutionary evidence, such data
would have to be converted into a subjective numerical form, its relative
influence on the final result being entirely in the hands of the taxonomist
during the coding process. If it should be asked how one estimates the
importance of a hypothetical trend, relative to a given piece of phenotypic
data, it may be pointed out that such a subjective estimate must be made,
at least subconsciously, in the mental process, and that an attempt to arrive
at a visible estimate of such degrees of relative importance could in itself be
illuminating. Such a mixing of the factual with the hypothetical is the basis
for the objection of numerical taxonomists to the inclusion of evolutionary
data in the analyses themselves. The second approach indicated above is
perhaps the more satisfactory ; the consideration of phylogenetic data after
completion of a purely phenotypic grouping would result in the final phylogenetic
classification being the same, but it would then be possible to see precisely
what changes in a purely phenotypic grouping had been made by the taxonomist
in order to achieve a more phylogenetic relationship, thus opening the way
for a more informed discussion of the significance of such changes.

Apart from their acceptability as techniques for performing the sorting
and group-forming processes of taxonomy, numerical analyses offer a number
of additional benefits. Information concerning the homogeneity and relative
similarity of the groups is available from most analytical methods, thus enabling
the purely taxonomic decisions regarding the status of the groups to be based
on rather more precise evidence than usual. The analytical techniques involved
are mathematically defined and reproducible, thus the repetition of an analysis
with new or differently defined characters is capable of providing additional
evidence concerning the validity of the original choice of characters, or
classification arrived at, since the computational procedure itself remains
constant.

The basis of numerical methods

The taxonomic process is essentially the translation of observations made
on individuals into statements of similarity and hence of group structure. The
use of mathematical techniques in taxonomy has been largely confined in the
past to their secondary applications of describing and substantiating taxa
which have been established by subjective processes. Techniques of this type,
e.g. Analysis of Variance, Discriminant Functions, still require the prior
establishment of groups by some means or other before they can be applied.
It is only with the advent of electronic computers that it has become practicable
to carry objective translations of information concerning individuals into state-
ments of group structure. While such analyses almost all start by computing
some measure of similarity between all possible pairs of individuals, they differ
greatly in the way in which this information is used to detect group structure.

One of the first results of the use of numerical methods of data analysis
is an increased realization of the multidimensional nature of taxonomic
relationships. A single dimension is sufficient to describe the relationships of
two points. If a third point is added, a statement of its relationship to the
first point will necessarily fix its position relative to the second point, a position
which may well not represent its true relationship. This is a difficulty which
may be overcome by the addition of a second dimension. Clearly, the addition
of a fourth point may require the addition of a third dimension, so that in general
terms it may be said that n—1 dimensions will be needed to describe fully all
possible relationships of » points. This statement will be obviously as true
for taxa or individuals as for points, though it must be emphasized that this
is the maximum number of dimensions which may be needed, particular cases

R. C. JANCEY 337

may well require fewer, the simplest situation being a straightforward clinal
variation with n points arranged in a straight line.

It should be pointed out before proceeding further that not all analytical
methods employ the multidimensional Euclidean space foreshadowed above.
Indeed, the advantages of an entirely non-metric space for the detection of group
structure are considerable (Rogers and Fleming, 1964). It is felt, however,
that the concept of similarity between individuals or groups is intuitively
considered in terms of real spatial relationships, and that for this reason the
concept is worthy of retention and consideration in greater detail, even though
it involves excursions beyond three dimensions.

a. b
Cc d
e f

Fig.l. The representation of correlated characters by oblique axes.

a, Six individuals arranged according to their mutual phenotypic similarities, as revealed
by a single character. 6, As in a, but with two perfectly correlated characters. c, The same
six individuals, now showing their phenotypic similarities as revealed by two highly, but not
perfectly correlated characters (the cosine of the angle enclosmg the points is equal to the
correlation between the characters). d, As in c, but with two orthogonal axes now replacing
the two oblique axes of the correlated characters. The positions of the points remain unchanged
relative to each other. e, Three individuals described in terms of four correlated characters
(the four axes are not necessarily confined to two dimensions). f, As in e, but re-expressed
without loss of information in terms of two orthogonal axes (the maximum number needed to
exvress the relationships of three individuals).

Information as collected by the taxonomist is expressed in terms of a
number of reference variables, i.e. the characters observed and recorded for
each specimen, the variables being more or less correlated. Thus the phenotypic
relationships of a collection of individual plants for which 2 characters have
been recorded may be thought of as being described in terms of x oblique axes
(oblique because of the character correlations), the axes being located in a space
of at most n—1 dimensions where n equals the number of individuals included
in the analysis. Thus the relationships of the individual specimens could equally
well be represented by n—1 orthogonal axes as by the «w oblique ones. If
is less than n—1 then « represents the maximum number of dimensions required
to represent the information available, the extent to which this number of
dimensions can be reduced depending on the extent to which the characters
are correlated (see Fig. 1).

While n-1 dimensions represent the maximum number of dimensions
needed for complete description of the population, in practice far fewer

G

338 NUMERICAL METHODS IN TAXONOMY

dimensions are needed to contain the information available. Indeed, a further
reduction in the number of dimensions may be achieved with a level of distortion
which would be quite acceptable in the interests of simplicity of description.
Thus one of the main objects of a taxonomic computer programme is to take
a population whose interrelationships are described in terms of a large number
of correlated characters, and to re-express the interrelationships in terms of
a relatively small number of dimensions, while allowing the taxonomist to
nominate the level of distortion, if any, which is acceptable. At the same
time, and unlike classical taxonomic methods, the processes to which the data
have been subjected are completely definable. There are then four further
steps in the taxonomic process, all of which will have been simplified by the
re-expression of the characters. Firstly, an examination of the spatial
relationships of individuals for evidence of group structure; secondly, the
assignment of individuals to groups; thirdly, an evaluation of the relation-
ships between the groups; and finally, the setting up of characters, or linear
compounds of characters (cf. discriminant functions) to discriminate between
the groups.

The technique of factor analysis is particularly well adapted to performing
the first part of this process. It is a technique first used by Spearman (1904
et seq.) to describe the results of a large number of different tests of human
ability in terms of a relatively small number of special aptitudes, e.g. manual,
visual, numerical, etc., each special aptitude being described by a linear
compound of the original tests. This is clearly the same as the first part of
the taxonomic process, and by a slight extension, can be used as such. The
analysis is based on the formation of a correlation or similar matrix from the
original characters, from which is extracted a series of vectors or factors
compounded from the characters. Since these factors when extracted from
the matrix are made up of varying contributions from the original characters,
it is possible to re-state the population relationships in terms of factor scores
rather than characters (for a full account of factor analysis, see Harman, 1960).
The relative information content of the factors depends on the method used
for extracting them from the matrix, and for taxonomic purposes particular
requirements for information distribution apply. The prime purpose of factor
analysis of taxonomic data, as has been stated, is to reduce as far as possible
the number of dimensions used in taxonomic description, so that as small as
possible a number of meaningful factors is desirable. The Principal Axes
method of factor analysis is such that the residual variance of the matrix is
minimized with the extraction of each factor, thus the first factor extracted
will contain the most information, and although in the Principal Axes method
the number of factors extracted is equal to the order of the original correlation
matrix, the information content of succeeding factors falls off rapidly and
becomes non-significant. In graphic terms, the analysis examines a population
described in terms of a number of oblique axes set in a multidimensional space,
and computes the one axis best able to describe the spatial relationships of
the population, the axis being composed of a linear compound of the original
characters used. The analysis then investigates the position of the axis best
able to represent the spatial relationships undescribed by the first axis. By
definition, these axes and the succeeding ones must be orthogonal to each other.
Knowing the contributions of the original characters to each factor, it is possible
to re-express the data in terms of factors. By expressing the relationships
of the individuals in terms of the first three factors only, a loss of information
is incurred, but because of the rapid fall off in information content of the factors
this is not usually serious, but has the advantage that limitation to three factors
enables the spatial relationships of the individuals to be expressed graphically
using isometric graph paper.

While factor analysis does not, in itself, delimit groups, it does present
data in a far more comprehensible form as a basis for the establishment of such

R. GC. JANCEY 339

eroupings by other means. Methods are available which do make an objective
demonstration of group structure, notably those of Goodall (1953), Michener
and Sokal (1957), Sneath (1957), Williams and Lambert (1959), and Rogers
and Fleming (1964). A fuller account of these methods will be found in Sokal
and Sneath (1963), but for the purposes of this discussion it is sufficient to say
that while considerable differences exist between the respective techniques,
they all depend essentially on the calculation of some measure of association
between all possible pairs of individuals, based on the characters measured.
Such a measure of association can take many forms, being usually based on
the ratio of character matches to mismatches between pairs of individuals,
since such a measure is particularly well suited to data in a presence or absence,
or limited class form. Having computed some measure of association, groups
may be established by a synthetic process, the agglomeration of individuals
possessing mutually high levels of association, discontinuities in the agglomerative
process indicating group structure. The actual delimitation of groups may be
performed automatically in response to some parameter involving the dis-
continuities—essentially a relationship between variation within the group and
that of the whole population.

The techniques of Goodall (1953) and of Williams and Lambert (1959)
are rather different in that they are analytic processes designed primarily for
ecological use, whereby the population is subjected to successive divisions into
the most homogeneous sub-groups. Such methods are particularly adapted to
two-state character data, and provide a monothetic classification with
hierarchical ordering of groups.

The methods of detecting group structure which have been described
above do not by themselves give any obvious indications of the inter-
relationships of the groups demonstrated. Levels of similarity at which groups
form from individuals submitted to analysis are usually shown in the form
of a dendrogram. Such diagrams have the advantage of illustrating clearly
the discontinuities between groups. They cannot, however, represent in graphic
form the similarity relationships between all pairs of individuals. Such a
representation is not possible in two dimensions for the reasons described
previously. A pictorial representation approximating to group inter-relation-
ships may be obtained by the combination of factor analysis with one of the
techniques of group detection described. The centres of gravity of the groups
can be calculated in terms of three-dimensional space from the factor scores
of individuals on the first three axes of a factor analysis. Knowing the
individual co-ordinates of members of a group, a value for the standard
deviation from the mean can be calculated for the group on each axis. It is
thus possible to construct a perspective diagram of the group relationships
on isometric graph paper, in which the groups are represented as ellipses drawn
at one standard deviation from the mean of the group on each factor axis, the
ellipses serving to indicate both the spatial relationships and the amount of
variation found within and between the groups. It might be argued that more
real information could be obtained from a perspective diagram showing the
positions of all the individuals on which the analysis was based. Such a
diagram is impracticable, since the illusion of three dimensions is lost when
a large number of points need to be shown, and in addition no advantage
would have been gained from the objective discrimination of groups made
previously. An example of a perspective diagram of the former type is shown
in Jancey (1966).

Conclusions

Numerical methods of data analysis are considered by the author to
represent a valuable new technique available to the taxonomist. It is
unfortunate that some taxonomists have looked upon the technique as an
isolated field of endeavour, together with chemotaxonomy and cytotaxonomy,

340 NUMERICAL METHODS IN TAXONOMY

and bearing no close relation to taxonomy as practised in the herbarium.
While for purely practical reasons classification must continue to be based
largely on morphological data, it would seem unreasonable to ignore any
additional information concerning the living organisms which might be
available. Similarly, a technique which enables the maximum amount of
information to be extracted from a mass of raw data by a defined process
would seem to be worthy of consideration by all taxonomists. The nature
of the results yielded by numerical methods should be emphasized, since they
are a frequent source of misunderstanding. The computations do not produce
classical taxa, but group the individuals for which data was provided. Precise
information is provided concerning the membership, distinctness, and diagnostic
characters of the groups produced, but the status of any group in terms of
orthodox taxonomic nomenclature, and its relationship to other taxa, are
entirely in the hands of the taxonomist, the only difference being that he is
provided with rather more information than usual on which to base his decision.

References

Goopatt, D. W., 1953.—Objective methods for the classification of vegetation. (1). Austr.
Journ. Bot., 1: 39-63.

Harman, H., 1960.—** Modern Factor Analysis.” (University of Chicago Press, Chicago.)

Jancry, R. C., 1966.—An investigation of the genus Phyllota (DC.) Benth. Proc. Linn. Soc.
N.S.W., 90: 341-375.

, 1965.—The application of numerical methods of data analysis to the genus Phyllota

(DC.) Benth. Aust. Journ. Bot. (in press).

MIcHENER, and SoxaL, R. R., 1957.—A quantitative approach to a problem in classification.
Evolution, 2: 130-162.

Roaers, D. J., and Fremine, H., 1964.—A computor program for classifying plants (II). Biro.
Science, 14: 9: 15-28.

SneatH, P. H. A., 1957.—The application of computors to taxonomy. Journ. Gen. Microbiol.,
17: 201-226.

Soka, R. R., and Sneatu, P. H. A., 1963.—“ Principles of Numerical Taxonemy.” (W. H.
Freeman and Co., San Francisco and London.)

SPEARMAN, C., 1904.—General intelligence objectively determined and measured. Amer. J.
Psychol., 15: 201-293.

Witiiams, W. T., and Lampert, J. M., 1959.—Multivariate methods in plant ecology (1).
Association analysis in plant communities. J. Hcol., 47: 83-101.

AN INVESTIGATION OF THE GENUS PHYLLOTA (DC.)
BENTH. (LEGUMINOSAE)

R. C. JANCEY*
Department of Biological Sciences, The University of Sydney

(Plates xxix—xxx)
[Read 24th November, 1965]

Synopsis

The generic status of Phyllota has been retained, pending a more extensive examination
of the taxonomic relationships of Pultenaea with Phyllota, Dillwynia, Aotus, and possibly other
related members of the Podalyrieae. At the moment the generic distinction between Phyllota
and Pultenaea rests almost wholly on the differing form and texture of the bracts, a distinction
of comparable magnitude existing between east coast species of Phyllota with leafy bracteoles
and persistent petals, and the remaining species, in which the bracteoles are predominantly
small, linear, and in some species scarious or coriaceous. A detailed study, using quantitative
techniques, demonstrated a number of phenotypically differentiated groups within P. phylicoides.
As a result of this, specific rank has been restored to P. grandiflora, P. squarrosa, and P. humifusa.
A number of other groups retain the identity of P. phylicoides. The need for an investigation
of the breeding behaviour of these groups is shown, also the need for an investigation of the
ecological facters connected with their distribution, in particular an investigation of possible
variation in the mineral composition of the sandstones of the Sydney district. Lectotypes are
named for Phyllota luehmannii and Phyllota pleurandroides.

INTRODUCTION

The genus Phyllota, which is endemic to Australia, was first described by
A. P. de Candolle in his Prodromus Regni Vegetabilis, as a section of the genus
Pulienaea. It was established as a genus in its own right by Bentham in 1838.

In the present study, the genus was investigated from several viewpoints,
firstly an evaluation of its taxonomic relationships with other genera, in
particular with the genus Pultenaea, an association first recognized by
de Candolle, but also with the genera Aotus Sm. and Dillwynia Sm. Secondly,
an investigation of the taxonomic relationships within the genus, particularly
in the Sydney region, where considerable variation occurs in a relatively small
area, and finally an attempt to correlate the variation in the Sydney district
with environmental factors.

The distribution of the genus (see Fig. 1) extends around the southern
part of Australia from Bundaberg on the Queensland coast, through New
South Wales, Victoria, Tasmania, South Australia and the south-west of
Western Australia. Figure 1 is based on herbarium records and, particularly
in the case of Western Australia, the distribution shown may be incomplete.

The genus is confined largely to a sandy substrate, either in the form of
fixed sand dunes or of sandstone, depending on the species. With the exception
of two Western Australian species, the genus is also confined to temperate
regions with a rainfall in excess of 20 inches per annum (Burbidge, 1960).

Since it appeared that infraspecific variation in the Sydney region was
much greater than elsewhere, it was decided to devote particular attention
to this area, where, in recent years, all variations have generally been included

* Now at The Department of Quantitative Taxonomy, New York Botanical Garden, Bronx
58, New York, U.S.A.

PROCEEDINGS OF THE LINNEAN Soctety or New SoutH Wates, Vol. 90, Part 3

342 INVESTIGATION OF THE GENUS PHYLLOTA

within the species Phyllota phylicoides (Sieber ex DC.) Benth. (e.g. Thompson,
1961). In the past these variants have been recognized as being of specific
rank, there being eight specific epithets presently included as synonyms of
P. phylicoides. It is proposed to deal with the investigation carried out in
the Sydney region in a first section, and to consider later the taxonomy of
the genus as a whole.

—
7 7 see

.
Projection (onual with two standard! puralles

135 Longitude East 130 of Greeomch 35 Lo

Fig. 1. Distribution map of the genus Phyllota

VARIATION IN THE SYDNEY REGION
Introduction

This section of the study formed the basis of an investigation using
numerical taxonomic methods. The numerical aspects of the investigation
have been reported elsewhere (Jancey, 1966). In the course of this investigation
a number of groups were established on the basis of phenotypic sunilarity.
In the present account it is proposed to describe the relationships of these
groups in terms of geographic distribution, breeding behaviour, and cytology,
in addition to phenotypic characters, and, in the light of the total information
available, establish the taxonomic rank, if any, to which they may be entitled.
While it is not proposed to introduce any discussion of numerical techniques
into this account, for convenience of reference the variants in the Sydney region
will be referred to by the group numbers used in the investigation previously
referred to.

Methods
Collection of specumens

All the specimens of P. phylicoides from the National Herbarium of New
South Wales and the Queensland Herbarium were examined in detail, and a.

R. C. JANCEY 343

Series of measurements on soaked material were also made on all the specimens
from the National Herbarium of New South Wales with the object of selecting
an area for investigation which would include all the sites of major variation
within the species. A further object of this examination was the selection
of phenotypic characters to be recorded on living material.

As a result of these preliminary investigations coupled with observations
in the field, the area selected for study was one bounded by Lake Macquarie
in the north, Jervis Bay in the south, and Lithgow in the west. It was felt
desirable to collect specimens as evenly as possible from the area under
investigation, since this would yield additional information concerning the
geographic and ecological range of the variants, apart from the location of
any possibly transitional forms. The method chosen was to take specimens
at one-mile intervals along roads and fire trails in the area, with the precaution
of making the collection 100 yards or more from the road to avoid plants
distributed by passing traffic. Detailed measurements were made on 313
specimens, and additional field trips made and specimens examined to establish
more precisely the distributional limits of the groups.

Recording of data

Data recorded at the time of collection included height of specimens, the
formation in which they were growing (heath, dry sclerophyll forest, etc.), soil
type, amount of shading, exposure to wind, and the presence of other plants
where it was felt that these were indicative of a change of habitat.

On returning to the laboratory, a number of records of characteristics
were made on the fresh material, and are listed below. Use was made of a
number of subjective scales for recording characters not amenable to objective
measurement ; illustrations of scale values are included where applicable. All
measurements of floral parts were made on flowers which had just opened,
and could hence be considered to be in a comparable state of development.

Data recorded in the laboratory may be summarized as follows:

Inflorescence length

The longest inflorescence of any specimen was stripped of its flowers, and
the length of the inflorescence axis measured in millimetres. While there was
considerable variation within individuals, there appeared to be some constancy
in the maximum inflorescence length which an individual could attain.

Number of flowers per inflorescence

As could be expected, this character was correlated with inflorescence
length. The correlation was not a necessary one, however, since independent
variation of these characters occurred. It served, in conjunction with the
other two inflorescence characters, to demonstrate affinities which could
otherwise be described only in extremely subjective terms.

Number of flowers per millimetre

A character derived from the two previous inflorescence observations.
It showed a high correlation with both, but demonstrated differences which
would have otherwise been difficult to describe, and was hence treated as an
independent variable.

Bracteole length and breadth

Both these two characters were measured to the nearest half millimetre.
Since all floral measurements were made on flowers which had just opened,
the characteristic enlargement of bracteoles after the death of the petals did
not constitute a source of variation between specimens.

344 INVESTIGATION OF THE GENUS PHYLLOTA

Calyx and bracteole indumentum

For flowers of the same age on the same plant this character was quite
constant. The epidermal hairs responsible were similar to those associated
with the bullae on the leaves. In flowers of the age examined, the hairs were
still present, obviously so in the case of the calyx, but the relationship between
bracteole and calyx indumentum was always the same. A subjective assessment
of this indumentum was made on a 1-4 scale of abundance (see Pl. xxix).

Bracteole colour

Bracteole colour appeared from preliminary observations to be far less
dependent on environmental factors than stem colour, and considerably more
constant within individuals. It was preferred consequently as a character.
It was difficult to assess, and may, like calyx colour, have represented the
outcome of a number of contributing forces. It was recorded on a subjective
scale, with green represented by 1, and varying through yellow to almost
wholly red at 4.

Calyx colour

This was recorded in a similar way to bracteole colour. The colour of the
calyx and bracteoles was frequently similar but not always so. For this reason
both characters were retained.

Calyx length

This character was measured by dissecting out the calyx tube at its point
of attachment to the receptacle, and measuring the length of the tube plus one
of the posterior calyx lobes to the nearest half millimetre.

Standard length

Tt was found that within the species P. phylicoides, the petals did not vary
in relative lengths, consequently standard length was taken as representative
of corolla length. After dissecting the flower, the total length of the standard
was measured, including the claw. The length of the standard was extremely
constant within individuals, such variation as occurred, some 0:5 mm. within
individuals, was almost certainly due to the difficulty experienced in flattening
reflexed standards prior to measurement.

Standard pigmentation

Three separate patterns were found on the standards of living specimens.
The three pigmented areas were recorded on subjective 1—4 scales, the four
values being illustrated in Figure 2.

Leaf spacing
The number of leaves produced within the 10mm. of axis immediately

behind an inflorescence was counted. This character was somewhat susceptible
to environmental factors.

Leaf weight per millimetre of length

The character was measured by determining the air-dried weight of 20
leaves immediately below an inflorescence, and dividing this weight by 20 times
the average leaf length as found in the previous character, thus giving a value
for leaf weight per millimetre of length. Given the assumption that all leaves
of all individuals were of equal density, this character would have provided
a comparable measure of the cross sectional area of the leaves, a character which
was seen in the preliminary investigation to have a marked constancy within
individuals, and an equally marked variation within individuals. No attempt

R. C. JANCEY 345

oe

a

Be

Fig. 2. Illustrating the values from 1 to 4 on the subjective scale for pigmentation of the
standard. Upper, middle and lower zones are shown in a, b, c respectively. The composite
form is shown in d, which also illustrates the alternative form taken by the lower edge of the
middle zone.

346 INVESTIGATION OF THE GENUS PHYLLOTA

was made to confirm the assumption of equal density, since the differences in
cross sectional area were so large and so consistent as to be quite distinct despite
any possible variation in leaf density.

Leaf length

Measures of leaf length were based on the average of 10 fresh leaves taken
from immediately below the inflorescence, the leaves being measured to the
nearest millimetre.

Leaf tips

Leaf-tip shape varied from obtuse and rounded, bearing a very small,
deciduous, black mucro, to an acuminate structure in which the greater part
of the tapering leaf tip was yellow, merging into a still tapering black mucro.
This character was remarkably constant within individuals, and proved to be
of considerable taxonomic worth. The character was recorded subjectively on
a 1-4 scale since no objective measure could readily be applied (see Pl. xxix).
Frequently associated with a value of 4 on this scale was a recurved state of
the leaf tip. This feature was also recorded.

Decurrence of leaf bases

There was a variation in the extent to which leaf bases were decurrent
on the stem, some individuals showing bases decurrent for three to four milli-
metres, others on leaf abscission leaving no more than a small tubercle. There
was considerable variation within individuals, but the subjective decision was
made that this was sufficiently less than the variation between individuals,
when assessed on a 1-4 scale, to be of possible taxonomic value.

Leaf bulla number

Leaf bullae were found to be the persistent bases of epidermal hairs; the
number of bullae on the adaxial surface of the leaf was assessed on a 1-4 scale
(see Pl. xxix).

Leaj bulla size

The size of leaf bullae was effectively constant on any given specimen,
but showed considerable variation between specimens. While there appeared
to be a strong negative correlation between this and the previous character,
it was shown not to be a necessary correlation by one group of individuals and,
at least in this one case, leaf bulla number is of taxonomic significance.

Indumentum of style

Variation of stylar indumentum fell into two classes; in all cases hairs
were of the same type, in one class covering the ovary and lower part of the
style, while in the other class, apart from covering the ovary and lower part
of the style, they continued along the upper edge of the style, around the hook,
and almost reached the stigma.

Continued growth of the inflorescence asis

A marked division existed between individuals in which the axis bearing
the inflorescence died back to the apex of the inflorescence, after having produced
no more than 1-10 mm. of fresh growth, and those in which at least some of
the axes continued to grow on unimpaired (see PJ. xxx).

A number of characters used previously in the taxonomy of this complex
were rejected after a preliminary survey of living material, on the grounds that
the character concerned was essentially constant within the species under
investigation. Characters coming within this category were the shape of calyx
lobes, their length relative to that of the calyx tube, the relative length of the

R. C. JANCEY OAT

petals and their shapes, and also the shape of the style. Other characters
were rejected because of excessive susceptibility to environmental factors ;
these included stem and leaf indumentum.

Fig. 3. Perspective iulustration of the relationships of morphologic groups in three dimensions

Morphologic groups established within P. phylicoides

Figure 3 shows a perspective illustration in three dimensions of the
phenotypic relationships of the groups. It is not proposed to consider the
numerical techniques by which the groups were delimited in this taxonomic
account, but it may be said briefly that the three axes represent the first three
axes of a principal axes factor analysis of the correlation matrix of characters.
Thus each axis is composed of varying contributions from the characters used

348 INVESTIGATION OF THE GENUS PHYLLOTA

in the study. The groups have been discriminated by another multidimensional
analytical technique, but for clarity of expression have been represented in
terms of factor scores of their members. This was achieved by calculating
the mean on each axis for the members of a group, calculating the standard
deviation and drawing the ellipse at one standard deviation from the mean.
The acceptability of the groups from the biological point of view obviously
differs considerably and will be considered later in the text. While it is possible
to relate the axes as drawn to particular combinations of characters, it would
Seem more useful in the present account to consider the diagram purely as an
illustration of the overall phenotypic relationships of the groups. For an
account of the quantitative aspects of this study, see Jancey (1966). Discrimi-
nating features of the morphology of the groups will be considered in conjunction
with the non-morphologic data which follows.

Other data relevant to the taxonomy of Phyllota in N.S.W.

It is proposed to consider the geographical distribution of the groups,
followed by ecological and other factors which may be concerned in group
distribution and relationships; finally, in the light of these observations, the
taxonomy of the N.S.W. groups of Phyllota will be considered.

Geographical Distribution of the Groups

From Figure 4, it will be seen that group 1 is of extremely limited
distribution. Herbarium material suggested that similar individuals might
be found in the Bundanoon-Penrose area, but field examination failed to reveal
any such individuals. It is felt that if this group occurs elsewhere it must
be of extremely local distribution to have escaped detection. It will be seen
from the geological map (Fig. 11) that no other form of Phyllota occurs in the
same area, and that a continuous area of apparently suitable mineral composition
extends to link the group with other forms of Phyllota, in particular groups
2, 3 and 4. This question will be discussed further in connection with the
ecological and systematic relationships of the forms.

The distribution of group 2 seen in Figure 4 is around the edges of the
Sydney Basin; it appeared more prolific in the area to the north of the
Hawkesbury River, elsewhere being locally abundant, but rather discontinuous
in its distribution. The distribution of group 3 shown in Figure 5 is similar
to that of group 2 in general terms, but differs in local habitat. This is
particularly interesting in the light of their very close morphologic relationship.

Group 4, which from the perspective diagram is also seen to belong to the
central complex of groups, has been collected from a limited area of the southern
ramp in the region of Mt. Keira (see Fig. 6). Additional material not included
in the distribution map would suggest that the group extends southwards into
the Bundanoon area.

The relationship of group 5 to the other central groups is less close than
would appear from Figure 3, since an additional character is available for the
discrimination of the group, that of possessing wholly terminal inflorescences.
The terminal position of the inflorescence is caused by the die-back of the very
short continuation of the axis after flowering. It will be seen from Figure 6,
that the group is largely confined to the region south of Botany Bay, having
its centre in the Royal National Park; a small number of individuals from
other areas have been assigned to the group. Of these, one individual from
the near north shore of Sydney, and the two from the Blue Mountains, are
similar to those found in the main distribution area of group 5, their isolated
position possibly being due to distribution by man. The remaining isolated
individuals do not show the terminal inflorescences characteristic of group 5,
and may now be better assigned to groups 2 or 3, an assignment which would
be more in keeping with their distribution.

R. C. JANCHY 349

Group 6 is confined to the north shore of Sydney Harbour (see Fig. 7).
The two individuals from the Royal National Park could be assigned subjectively
with greater confidence to group 5 in the light of data concerning terminalization
of the inflorescence. It will be seen from a comparison of the distribution of
this group (Fig. 7), with the combined distribution of groups 2 and 3 (Figs 4
and 5), that their areas are contiguous, but that there is virtually no overlap.

Very closely associated with each other, but quite distinct from other groups,
eroups 7 and 8 differ only in the characters dealing with red pigmentation of
the standard, group 7 having dark red standards and those of group 8 being

»o LITHGQow

O RICHMOND

Q Picton

rad af WOLLONGONG

B) WOLLONGONG
a Moss VALE
Q Moss VALE
°
NS)
JERVIS BAY
m_ MILES 5
TERVIS GAY ° $ 16 3+
MILES 4
fo) & 16 2. 4

Fig. 4. Distribution map of group 1 (P. humifusa in the taxonomic treatment), and of group 2
(part of P. phylicoides in the taxonomic treatment). Group 1 is indicated by crosses, and group
2 by solid dots.

Fig. 5. Distribution map of group 3 (P. phylicoides in the taxonomic treatment).

almost wholly yellow. The distribution of these two groups (see Figs 8 and 9)
overlaps in the region of Blackheath to Mt. Victoria; indeed plants with yellow
and red standards have been found growing next to each other in this region.
Group 7 contains two individuals isolated geographically from the rest of the
group, one at Mt. Keira, and the other south of Nowra. Group 8 contains
one individual from the Nowra region, but is otherwise confined to the Upper
Blue Mountains. Examination of these three specimens confirmed that their
assignment to groups 7 and 8, like the assignment of two individuals from the
Same region to group 5, was almost certainly a misclassification as a consequence
of the small number of individuals from the region. It will be seen from the
combined distribution of groups 7 and 8, that the groups are confined to the

350 INVESTIGATION OF THE GENUS PHYLLOTA

higher parts of the Blue Mountains. Comparison with Figures 4 and 5 would
suggest that the groups are geographically isolated from groups 2 and 3.
However, the anomalous records for group 5 individuals at Wentworth Falls
and also two Herbarium records for plants of the group 2, 3 type in the same
area would suggest that the geographic isolation may not be complete. The
ease with which plants of the group 2, 3 type can be found in the foothills of
the Blue Mountains, and their absence despite careful searching higher in the
ranges would suggest that their distribution in the latter area must be very
discontinuous and limited. The north-south distribution of groups 7 and 8
is less easy to establish than their east-west distribution. Figures 8 and 9

Dp LITHGOW
BROKEN BAY Bb LITHGow
BROKEN BAY
SYDNEY
SYDNEY
PORT HACKING a KANArGAA WALLS
u PICTON
F
q Picton
+
-
+
of WOLLONGONG
D) WOLLONGONG
bp Moss vase
al ~
JERVIS BAY ZTERVIS BAY
MILE
MILES 6 = 7
° & 16 2¢
° 8 16 a4

Fig. 6. Distribution map of group 4 and group 5 (both are part of P. phylicoides in the taxonomic
treatment). Group 4 is indicated by crosses and group 5 by solid dots.
Fig. 7. Distribution map of group 6 (part of P. phylicoides in the taxonomic treatment).

show the distribution to be confined to the highest parts of the Blue Mountains,
and from further observations it may be said that the northerly distribution
extends at least as far as the inner end of the Wolgan Valley, north of Clarence,
but has ceased before the Capertee Beds east of Rylstone are reached. This
absence of Phyllota is paralleled to the east by the absence of groups 2 and 3
going north along the Colo-Putty-Singleton road (see Figs 4 and 5). The
southerly extension of groups 7 and 8 appears to be limited by the available
high ground of suitable mineral composition. South of the Blackheath-
Katoomba area there is little continuous high sandstone ground, apart from
the neck running out to Mt. Solitary. This area has not been examined for
the presence of Phyllota, but individuals of group 8 have been collected from
Kanangra Walls, where they were found growing on Permian sandstones.
Their presence in this area would imply either great age to the discreteness
of group 8, followed by no evolutionary change, or alternatively, that even

a ee

R. C. JANCEY 351

assuming a route via Mt. Solitary, colonization and genetic interchange over
considerable distances of unsuitable country were possible. No Phyllota has
been found on the non-sandstone rocks surrounding Kanangra Walls. To the
east of the walls, groups 2, 3 have been found at Oakdale, though at a very
much lower altitude. South of Kanangra, the first sandstone examined lies
south of the Wollondilly River, to the east of Mt. Bullio, where the altitude
is also much lower. Here no Phyllota is found at all, though some 10 miles
to the east lies the limited area containing group 1.

Group 9 follows the distribution of group 6 (see Figs 10 and 7). The small
group of individuals from group 9 in the region of Appin, are rather different
from the. bulk of the group found north of Sydney Harbour, being rather less

De %e LITHGOW
® oe
Woe ete

BROKEN BAY

D KATooMgA

o LiTHQGOW

amt vicToRIA BROKEN GAY 37 | SYDNEY

Si¢% KAtoor BA aK ANANGRA WALKS

SY DNEY
a PicTon

DRANANGRA WALLS

o PICTON
D) WOLLONGONG,

O BARGO ”

WOLLONGONG

Jervis BAY

A
Servis GAY
MILES 8

8. Distribution map of group 7 (part of P. squarrosa in the taxonomic treatment).
ig. 9. Distribution map of group 8 (part of P. squarrosa in the taxonomic treatment).

robust, and possessing rather fewer flowers per inflorescence. However, these
distinctions will be considered in more detail in the section dealing with the
taxonomy of the New South Wales forms of Phyllota. The two individuals
found on the Hume Highway south of Bargo correspond very well to the
character values for group 9. Their presence in this area could possibly have
been the result of human distribution, but if so, the distribution must have
taken place in early colonial days or before, as a specimen of the group 9 type
was collected by MacArthur from Bargo, and is at present in the Kew Herbarium.
Failure of the plants to spread may have been due to unsuitable substrate.
Herbarium records show specimens of the group 9 type to be growing immediately
south of Botany Bay in the region of Como. Careful searching has failed to
reveal any living examples in this area, an absence which may be due to urban
development.

352 INVESTIGATION OF THE GENUS PHYLLOTA

Factors affecting Distribution and Variation

Geological and Ecological

Figure 11 shows the distribution of all the individuals used in the analysis,
Superimposed on the areas of sandstone in the region. It will be seen that,
while no records fall outside the sandstone areas shown on the map, there are
large areas of sandstone in which no Phyllota of any group is found. While
no quantitative study has been made of this problem, it may be said that the
presence or absence of Phyllota is to some extent associated with small changes
in altitude. In view of the small changes in altitude involved, this would
suggest stratigraphic changes, and hence possible changes of a mineralogical
nature. Changes in iron content of the Hawkesbury Sandstone certainly occur

o THGow

PoRT HACKING

pf woLrongong

n Moss VALE

~
TERVIS BAY

MILES 10
° 8 16 24

Fig. 10. Distribution map of group 9 (P. grandiflora in the taxonomic treatment).

(David, 1950), and it would seem reasonable to suppose that other metals at
least would be present in varying abundance. Cases where discontinuities of
Phyllota occur without changes in altitude would not necessarily refute this
basis for variation since, in the case of dipping strata, movement over the surface
would bring about the same changes as changes of altitude in the case of
horizontally bedded strata. A particularly clear case of the correlation of
strata with distribution is found in the region north of Wiseman’s Ferry, where,
although Phyllota is apparently absent from the slopes of the sandstone plateau,
it may be found at the top of the plateau. Equally flat, although slightly
lower, areas of the plateau are however once again devoid of Phyllota. Similarly,
individuals of group 5 disappear from the Princes Hghway just south of the
Woronora Dam turnoff as the highway climbs up towards the Bulli Pass, and,
although not included in Figure 6, the distribution of group 5 continues around
the lower contour as far as the western limit of the exposed sandstone to the
north-east of Appin.

353

R. C. JANCEY

NARRABEEN GROUP,
GOSFORD FORMATION

TRIASSIC
==

HAWKESBURY SANDSTONE

ERNE Bay

MILES PERMIAN SANDSTONES
(UN DIFFERENTIATED)

Fig. 11. Map of total distribution relative to areas of sandstone. The westernmost areas shown
as Hawkesbury Sandstone are, according to Standard (1963), Narrabeen Sandstone but are

mineralogically different from that in the Gosford area.

H

354 INVESTIGATION OF THE GENUS PHYLLOTA

The geological basis for discontinuous distribution as described above is
supported by the observation that Phyllota, at least in the Sydney region, is
not unduly sensitive to such environmental factors as shading or exposure,
and appears able to survive under a range of water availabilities. Plants of
the same group have been found in very moist shaded situations, and in cracks
in rocks with scarcely any soil and fully exposed to the sun, in the same area.
Several facts tend to discount the importance of geological factors on distribution.
Firstly, even single groups of Phyllota are found growing on a relatively wide
range of sandstone types, thus groups 2 and 3 are found on the Permian, Nowra
and Hawkesbury Sandstone, and on the mineralogically much richer Narrabeen
Sandstone in the Gosford region. Secondly, it is found that while Phyllota is
characteristically confined to sandstone, it is capable of extending for short
distances on to shale soils, where small cappings of such soils remain in areas
of sandstone, or where lenses of shale occur within the sandstone strata. Finally,
Phyllota of group 8 will grow both on Hawkesbury Sandstone, as in the
Katoomba area (though this is considered to be Narrabeen Sandstone by
Standard, 1963), and also on the mineralogically richer Permian Sandstone of
the Kanangra Walls region. Evidence such as this, that Phyllota is able to
grow on such sandstones, even extending into the margins of the shale, would
make it seem more probable that if distribution were being affected
mineralogically, then, at least within sanstone soils, it would be by paucity
of supply or by imbalance, rather than by general superfluity.

Cytology and barriers to interbreeding

Since the forms of Phyllota growing in the Sydney region showed a variety
of types of genetic isolation, a number of investigations were carried out with
the object of establishing the possibilities which existed for gene interchange
between the groups.

A number of inflorescences were covered with muslin bags on representative
plants from all the groups. The bags were attached prior to the opening of
the flowers, and the plants revisited later in the year when seed could have
been expected to have been set. The muslin-covered inflorescences were
harvested, together with other inflorescences from the same plants which had
not been so covered. In no case did any of the flowers covered with muslin
set seed, from which it would appear that self-pollination is unlikely to occur
in the field. Seed production from cross-pollination was variable, however,
and at a low level, rather less than 5% of the exposed flowers producing seeds.

Isolation due to different flowering times

Absolute flowering times varied from year to year, but in the observation
of three flowering seasons, the relative flowering times of the groups remained
reasonably constant. The first groups to flower were 2, 3 and 5, doing so
simultaneously during early August. These were followed by group 6 about
three weeks later. In October or early November group 9 began flowering
north of Sydney Harbour. This coincided with the end of the flowering season
of group 6 although sufficient overlap occurred to permit of possible cross-
pollination. With very few exceptions groups 2, 3 and 5 had ceased flowering
by this time, so were isolated from direct genetic exchange with group 9. A
possible bridge in time might exist however in the form of group 6, whese
flowering time coincided with groups 2, 3 and 5 and with group 9. This would
present no problem in the case of groups 2, 3, 6 and 9 since their geographical
distributions are contiguous, but with the exception of the individuals mentioned
in connection with the descriptions of geographical distribution, groups 5 and
6 are isolated from each other by both Sydney Harbour and Botany Bay.
The members of group 9 occurring in the Appin region are in geographic contact
with group 5. However, the members of group 9 in the Appin region, apart
from the slight differences from the rest of group 9 already mentioned, also

R. C. JANCEY 355

differ in flowering time, since they do not flower until much later, usually in
January, and are thus quite isolated temporally from the group 5 individuals
with which they are in contact. Late flowering was also found to be
characteristic of groups 1, 4, 7 and 8, all of which flowered in late December
or January onwards. Group 4 and the Appin members of group 9 possessed
a Oe period of similar length to that of the groups already described,

, groups 2, 3, 5, 6 and 9, the precise length appearing to be dependent on
ane season, put extending over about eight weeks. Thus group 4 and the Appin
members of group 9, by commencing flowering in the beginning of January,
experienced a temporal isolation from groups 2, 3 and 5 with which they may
possess geographic contact (as was found to exist certainly between group 5
and the Appin members of group 9), while at the same time they are isoJated
geographically from the groups with contemporaneous flowering periods, i.e.,
groups 1, 7 and 8.

TABLE 1
Flowering periods, and geographic contacts of growps

Flowering period Geographic contact
Group with groups
Aug. Sept. Oct. Nov. Dec. Jan. Feb. Mch.

wZ
ie}
5
®

~)

i)
2
BB
loon

, 3 a

eee 2, 3 and 5.
possibly 2, 3 and 5.
, 39 and 6

2, 3, 4 and 5.

»

OOWDAAAIE WHE !
1
NNanNnwnwbd

(Appin)

The flowering period of the remaining groups, 1, 7 and 8, while beginning
in January, differed from the other groups described in being more extended,
continuing until the autumn, but with fewer and fewer flowers being produced
per inflorescence as the season progressed, a characteristic which is particularly
marked in group 1, where one-flowered inflorescences were frequently produced
late in the season. Groups 7 and 8 are virtually identical, apart from the
pigmentation of the standard, and group 1 exists in geographic isolation.

The possibility that flowering periods were determined by environmental
factors was investigated by transplanting members of the various groups to
sites in the grounds of Sydney University. It was established that flowering
occurred at times characteristic of the group, rather than in response to local
climatic conditions. There was a tendency for flowering to be two to three weeks
later than plants of similar groups in their natural habitats, also the flowering
seasons were much shorter with far fewer flowers produced by all plants than
would have been expected under natural conditions.

Information from field observations which would support the view that
flowering times are controlled genetically rather than purely environmentally
comes from two sources. Groups 2 and 3, which have an extremely wide
distribution, flower at virtually the same time regardless of their position, and
group 1 flowers at the same time as groups 7 or 8, although growing at a very
much lower altitude, and in a much lower rainfall area.

Cytological Hxamination
All cells examined were found to have a chromosome number of 2n=14.
No differences were observed in the morphology of the chromosomes. The
need for artificial breeding experiments is clear but, owing to the time required

356 INVESTIGATION OF THE GENUS PHYLLOTA

to produce flowering progeny, and the difficulties experienced in raising seedlings,
no attempts at artificial crossing were made. Voucher specimens were placed
in the herbarium of Sydney University, and are listed in the formal taxonomic
section of this work.

Growth Habit

Apart from the differing propensities for terminalization of the inflorescence
aS mentioned earlier, a number of further habit differences exist between the
groups which for various reasons it was not possible to utilize in the construction
of the model of phenotypic groups.

Branching pattern

The extent to which plants branched was characteristic of the different
groups, and appeared to be related in some way to the tendency towards
inflorescence terminalization. Groups 2, 3 and 5 branched most profusely,
the branches emerging and remaining at an angle of about 45 degrees to the
parent axis. Such branching tended to be, but was not invariably, associated
with the formation of inflorescences. The resulting appearance tended to be
of a rather compact bushy shrub. Groups 4, 6 and 9 showed less tendency
towards branching and, associated with this, a more vigorous growth of existing
stems, and a greater mean plant height, at least in the case of groups 6 and 9.
Groups 7 and 8, although showing virtually no tendency towards inflorescence
terminalization, and branching relatively infrequently, nevertheless showed a
distinct limitation in the height to which they would grow, even in sheltered
situations. Joined with them in the matter of height limitation was group 4,
though to a rather lesser extent. The most extreme case of height limitation
was found in group 1, in which the production of the characteristically few
flowered inflorescences appeared in no way to interfere with the continued
growth of the stem. Plant height seldom exceeded three to six inches, however.

Rooting system

Some explanation of plant height variations not apparently relatable to
branching characteristics may be found in the form of the plant at and below
eround level. In setting up the transplant experiments described previously,
stolon-like structures were discovered while digging up plants in the field.
Since the presence of runners had been used as a diagnostic character in the
case of a South Australian species of Phyllota (Willis, 1957), their occurrence
among the groups under discussion was investigated. Horizontal underground
structures linking two or more aerial stem systems were found in groups 2, 3,
4, 7, 8 and 9. Microscopic examination showed that these possessed the
anatomical structure of a root. The extent to which the root system developed
suckers varied considerably among the groups, beg most marked in groups
7 and 8, considerably less extensive in groups 4 and 9, but with the suckering
roots much thicker in the latter though occurring rather less frequently.
Suckering was very infrequent among groups 2 and 3, though short horizontal
reots, about six inches long and rather thicker than the normal rooting system,
were encountered frequently, even in seedlings about 12 months old. No
examples of suckering were found in groups 5 or 6, though group 6 showed
occasional examples of short horizontal roots similar to those found in groups
2, 3. No such roots were found in group 5, nor were any other indications
of a tendency to produce suckers. Group 1, while showing no tendency to
produce root suckers, achieved a similar, though lower, growth habit to groups
7 and 8 through an extensive prostrate stem system, from which arose short
erect branches, bearing in turn a few even shorter lateral branches (see Pl. xxx).
Continued growth of some aerial shoots caused them to become procumbent,
finally forming part of the prostrate stem system. Apart from the presence
of leaf bases, anatomical investigation confirmed the distinction of group 1 in

R. C. JANCEY 357

possessing a prostrate stem system, as opposed to the suckering root system
of some other groups. The free and extensive suckering of groups 7 and 8,
and to a lesser extent of group 4 (see Pl. xxx, ¢ and a), would appear to be
related in some way to the limited height achieved by their aerial parts, a
situation paralleled in some degree by group 1. The stimulation of suckering
by burning off the aerial parts during bush fires is a possibility, but since bush
fires are prevalent throughout the areas of Hawkesbury Sandstone, it is not
felt that this would account for the observed differences. The plant shown
in Plate xxx was collected from an area believed not to have been burnt for
some years, yet numerous young suckers were emerging. Examination of
group 5 plants from areas known to have been burnt showed no trace of suckering,
while among plants of groups 6 and 9 growing within a few feet of each other,
plants of group 9 bearing suckers were easily found, while none were found
on plants of group 6.

The Taxonomy of Phyllota in New South Wales

As was stated previously, the forms of Phyllota, presently referred to
P. phylicoides, which occur outside the area covered by the investigation are
remarkably uniform, and correspond in fact to groups 2, 3 in Figure 3. Some
herbarium material from sites in Queensland might be referable to group 5
but, in the absence of non-terminal inflorescences, the distinction between
the groups rests mainly on inflorescence characters which can be established
only by destruction of the inflorescence. In any case, at least in the light of
present knowledge, the entities recognized on the basis of information from
the area under investigation include the whole range of variation found within
what has been recognized hitherto as the species P. phylicoides.

Taxonomic status of the groups

The groups which have been described are, for the most part, easily
recognizable in the field by a trained observer. A considerable amount of
additional data exists supporting the status of the groups, but in many of the
groups there is a lack of suitable discriminatory characters on which the
confident allocation of some herbarium material might be made.

The difficulty described above presents the problem of purpose in erecting
formal taxa. Barriers to gene interchange appear to exist between all groups
with the exception of groups 2, 3 and 7, 8. The barriers, as described in an
earlier section, apparently being distance, differing flowering times or, at least
in the case of groups 6 and 9, genetic incompatibility. In this latter case, the
two groups are found within six feet of each other, and the flowering periods
overlap sufficiently to permit the possibility of frequent cross pollination. The
absence of intermediate forms is so complete that no difficulty was experienced
in distinguishing flowering individuals of the two groups at a glance; it was
scarcely more difficult in the vegetative condition. Clearly information of this
sort is not a satisfactory basis for decisions concerning barriers to gene inter-
change, and should be substantiated by controlled breeding experiments. The
length of time involved, however, placed such a programme beyond the scope
of this investigation. The probability of partial or complete barriers to gene
flow between the groups is relevant to their taxonomic status but, even were
complete genetic isolation demonstrated, the utility of erecting taxa not readily
discriminated on morphologic grounds seems doubtful. A more appropriate
vehicle for describing a number of the morphologically similar groups would
appear to be the deme system of terminology (Gilmour and Heslop-Harrison,
1954), in which groups of plants characterized, for example, by geographic
distribution or potentialities for gene interchange are described in such terms,
without any implication of orthodox taxonomic rank.

Considering the group relationships in the light of all the available
information, it is clear that the distinction least capable of substantiation is

358 INVESTIGATION OF THE GENUS PHYLLOTA

that between 2 and 3. Their separation was the result of small consistent
differences on a number of characters. The distinction was so slight as to be
undetectable by subjective means, and even now cannot be confidently
discriminated at sight. Such a distinction is clearly of little utility for taxonomic
purposes, particularly in view of the common flowering time and distribution
of the two groups. Consequently their distinction will not be maintained.

The combined groups 2 and 3 have, as their closest related form, group 5.
Discrimination between these two groups in morphologic terms rests on the
number of flowers per inflorescence, number of flowers per mm. and the number
of leaves per 10mm. respectively. A further distinction is given by the
invariable inflorescence terminalization in 5 as opposed to the continued growth
of the stem in 2, 3. Other distinctions lie in the production of occasional root
suckers in 2, 3 and, rather less infrequently, short, thickened horizontal roots
resembling abortive attempts at sucker production, as compared with the
complete absence of anything approaching sucker formation in 5. Finally,
the group distributions may be considered; apart from a small number of
herbarium specimens from Queensland whose identity with 5 was doubtful,
the group is confined to an area south of Sydney Harbour and, as far as can
be determined, is isolated geographically from the possibility of gene interchange
with 2, 3. Whether any incompatibility barriers also exist is not known, but
the effective isolation might well be sufficient to account for the observed
morphologic differences. While the magnitude of these differences is not
sufficient to make a formal taxonomic distinction either necessary or practicable,
recognition may nevertheless be given to 5 by describing it as a phenogamodeme.
This term indicates at one time the spatial, temporal, and genetic possibility
of interbreeding within the group and also the existence of phenotypic
distinction from other groups.

The status of 4 is to some extent analogous to that of the members of 9
occurring in the Appin area. Both are represented by small numbers of
individuals, and occur in a limited geographical area. In both cases the large
flowers occurring in lax inflorescences suggest affinities with 7,8 and 9. However,
the inflorescences of Appin individuals are both shorter and contain fewer
flowers than other members of 9. Those of 4 are quite typical of 7 and 8.
Despite the similarity of inflorescence with 7 and 8, in almost all other characters,
4 shows greatest affinity with the central complex of groups as may be seen
in Figure 3. As a consequence of this, it has been decided to unite 4 with 2, 3
and hence with 5 for taxonomic purposes. Individuals at Appin can best be
left as members of 9 since, although differences exist, they are associated most
closely with this group and, in the absence of further information concerning
their origin, cannot be separated reasonably from it. Both group 4 and the
Appin members of 9 may be given some recognition in the deme terminology
aS phenotopodemes, though absence of sufficient evidence precludes the use of
the distinction gamodeme.

Groups 6 and 9 show greater morphologic distinction from each other and
from group 2, 3 than other groups described so far. They differ also in having
superposed distributions, though that of group 9 is more restricted in area.
Both groups 6 and 9 tend to be characterized by taller growing plants than
group 2, 3, though this may be obscured by the age of the plants. As in the
groups described already, flower and inflorescence characters are of particular
discriminatory value. Inflorescence length is least in group 2, 3, followed
by group 6, and finally group 9 has the longest inflorescence. Inflorescence
density serves to distinguish groups 2, 3 and 9 with lax inflorescences, from
group 6 in which the number of flowers per millimetre is much greater, though
not equal to that found in group 5. A suspected connection between inflorescence
density and tendency towards inflorescence terminalization was not borne out
by group 6, since, although possessing denser inflorescences than group 2, 3,
it shows a considerably greater tendency towards vigorous continued growth

R. C. JANCHEY 359

of the stem after inflorescence production. Above average length of inflorescence,
coupled with great density, results in group 6 having the highest value of any
group for the number of flowers per inflorescence. Apart from the characters
already described, group 9 is distinguished from group 2, 3 and from group 6
by high values for weight per millimetre of leaf length; indeed, this character
distinguishes group 9 from all groups other than groups 7 and 8, where other
distinctions apply. The absence of root suckers in group 6 provides a further
link with group 5, while at the same time distinguishing it from group 9 in which
such suckers are frequently found, and group 2, 3 in which they are occasionally
present.
TABLE 2
Summary of major discriminating characters

Presence of Characteristic of groups

Procumbent ens 3
Consistent and prolific production of root suckers.
Consistent terminalization of inflorescence

Large flowers

Lax inflorescence. .

Long inflorescence

Reeurved leaf tips

Numerous flowers in dnvilomee@einaa

Massive leaves

e2)

.

»

BCID NS

’

This qualitative table is by no means exhaustive, and should be interpreted in conjunction
with the text. Quantitative values have been omitted in the interests of simplicity, but may
be found in the formal taxonomic descriptions.

Information concerning breeding behaviour on the north shore of Sydney
Harbour may be summarized as follows: Temporal isolation precludes direct
gene flow between group 2, 3 and group 9, though group 6 represents a possible
temporal bridge. In the absence of direct experimental evidence, no final
conclusions on gene flow can be reached; it may be said, however, that in
view of the distributions and flowering times of the groups, the absence of
intermediate forms, particularly between groups 6 and 9, would suggest inter-
group sterility.

In the light of the above considerations, group 9 will be restored to specific
rank. The status of 6 is less clear; while this group shows sufficient differenti-
ation from groups 2 to 5, when these are considered as individual groups, to
merit infra-specifie distinction, the increase in range of character variation
resulting from the combination of 2 to 5 is such that confident discrimination
between this combined group and 6 can no longer be achieved. Group 6 will
be merged, therefore, with these groups for formal taxonomic purposes, though
it undoubtedly represents a phenotopodeme of rather greater distinction. In
the absence of spatial or temporal breeding isolation, a decision regarding its
status aS a gamodeme must be reserved until direct experimental evidence is
available.

The two groups occurring in the Upper Blue Mountains, 7 and 8, show
little morphologic differentiation from each other, except in pigmentation of
the standard. Geographic distributions are not identical, but show considerable
overlap. Since flowering times are identical, little can be said concerning the
breeding behaviour of the two groups other than that there is no obvious barrier
to interbreeding. The contrary evidence provided by the absence of inter-
mediate forms is of rather less weight than in other groups, since the phenotypic
distinction is confined to characters which might have a very similar genetic
origin. While the differences in pigmentation enable the two groups to be
recognized as separate phenodemes, it is not felt to be a distinction meriting
formal taxonomic status. Consequently, these two groups will be considered
together; in this combined form ready discrimination from other groups is

360 INVESTIGATION OF THE GENUS PHYLLOTA

still possible. In qualitative terms it may be said that the group differs from
the ‘central complex’ of Figure 3 in possessing lax inflorescences of larger
flowers (though see previous remarks concerning group 4), and the much greater
leaf weight per mm. It is distinct from group 9 in its lesser plant height,
number of flowers per inflorescence, rather smaller flowers and narrower
bracteoles. Characters separatng 7 and 8 from all other groups are the
consistently recurved and acuminate leaf tips, and the abundance of root
suckers produced by unspecialized roots. With the exception of two apparently
anomalously located individuals from other groups, geographic isolation would
appear to be complete from all other groups and, even if the two individuals
mentioned should be representative of others as yet undetected, breeding
isolation would still be maintained by virtue of the differing flowering times.
In the light of the above considerations, the combined groups 7 and 8 will be
restored to specific rank.

The remaining group, group 1, shows considerable morphologic differentiation
from other groups. It differs from the combined groups 7 and 8 in its smaller
flowers, absence of acuminate recurved leaf tips, and in having a lower value
for leaf weight per mm. It can be distinguished from 9 by size of floral parts,
pigmentation of standard, inflorescence length and flower number, length and
weight per mm. of leaves, plant height and number of leaf tubercles. From
the ‘central complex’ of groups, group 1 differs in respect of plant height,
inflorescence length, bracteole breadth, pigmentation of standard, and leaf
length. Group 1 differs from all other groups in its low creeping habit and
prostrate stem, and also in geographical distribution. In view of the degree
of differentiation indicated above, it is felt that group 1 should be restored
to specific rank.

Thus in this area, specific rank is given to the following four assemblages :
LOOM UR DSU (3 tA. 162

The Relationship of Specific Epithets previously applied to Phyllota
in New South Wales, to the Groups described

In view of the considerable synonymy of the New South Wales species
of Phyllota it is proposed to consider the specific epithets available, the groups
to which they refer, and hence the synonymies of the species, prior to the formal
descriptions of the species occurring in New South Wales as determined by
this investigation. ,

PHYLLOTA HUMIFUSA A. Cunn. ex Benth. in Ann. Wien. Mus., I]: 77 (1838).

Features of the holotype of this species which are of importance in relating
the type specimen to the groups described in the preceding section are as
follows: Stems prostrate, thin and wiry. Leaves 6 mm. long, 0-5 mm. wide,
apex recurved with a small mucro. Inflorescence few—flowered (ca. 3), loose,
not terminal. Bracteoles linear-lanceolate, 3mm. long, 1mm. wide. Calyx
hirsute with appressed hairs, ca. 4mm. long (lobes 2mm.). Ovary villous,
style glabrous.

Holotype: Wombat Brush, Argyle County, N.S.W. A. Cunningham
No. 8 (K).

These characters, combined with the locality of the type specimen, leave
no doubt as to the identity of P. humifusa with group 1.

PHYLLOTA SQUARROSA (Sieber ex DC.) Benth. in Ann. Wien. Mus., IL: 77
(1838).
Basionym: Puiltenaea squarrosa Sieber ex DC. Prod., II: 113 (1825).
Leaves divergent, recurved, with distinct recurved, acuminate yellow tips,
leaves ca. 1-3 mm. long. Flowers few in spike, axis growing on while stil in
flower. Standard ca. 9-5 mm. long.

R. J. JANCEY 361

Holotype: Sieber No. 406 (GEN). The isotype at Kew bears Sieber’s
original label, and is localized as Blue Mountains; there are also two sheets
bearing this number at MEL. This specific epithet will be applied to the
combined groups 7 and 8.

Two names are available for the species represented by group 9: Phyllota
pilosa Benth. and Phyllota grandiflora Benth. Both species were erected by
Bentham in Ann. Wien. Mus., IL: 77 (1838), but included by him in Phyllota
phylicoides in Fl. Austr., IL: 95 (1864). The latter epithet will be adopted,
Since it describes the more striking and constant feature of the species.

PHYLLOTA PILOSA Benth.

There are three sheets of Huegel’s collection at Vienna, two of which
correspond to group 9; the remaining sheet differs in a number of respects
from both the two preceding sheets and Bentham’s description of the species,
corresponding more closely to the type of P. comosa. In particular this last
Sheet possesses rather more slender, erect leaves, the hairs over the whole plant
being more appressed. The flowers are smaller and the calyx much less pilose.
It corresponds most closely to NSW 7226, Gordon West, M. Tindale, except
for slightly longer, yellowish hairs.

P. GRANDIFLORA Benth.

Only one sheet bearing this name exists at Vienna, and none at Kew.
The specimen corresponds to Bentham’s diagnosis of P. grandiflora, but this
does not differ sufficiently from that of P. pilosa to differentiate the two species ;
thus for P. grandiflora, ‘.. . foliis supra tuberculoso-scabris subtus pube-
scentibus ...’, and ‘. . . bracteolisque pilosis flore brevioribus . . .’, while
for P. pilosa, ‘.. . foliis tuberculoso-scabris muticis, novellis calicibusque
pilosis, floralibus flores aequantibus ...’; the type specimens, with the
exception referred to in the case of P. pilosa, all being referable to group 9.

P. comosa (Sieber ex DC.) Benth. in Ann. Wien. Mus., IL: 77 (1838).
Basionym : Pulienaea comosa Sieber ex DC. Prod., IL: 113 (1825).

Holotype: Sieber No. 407. Locality, Nov. Holl. Two sheets of this
material are at Vienna, and one at Kew, all bearing Sieber’s original label, and
also two at MEL. The Kew isotype is similar to the specimen NSW 7226
Gordon West, M. Tindale. The vigorous continued growth of the axis is
characteristic of groups 6 and 9, while the size of the floral parts satisfactorily
establish P. comosa as a binomial associated with group 6. This is also in
accord with the isotypes at MEL.

P. ASPERA (Sieber ex DC.) Benth. in Ann. Wien. Mus., IL: 77 (1838).
Basionym: Pultenaea aspera Sieber ex DC. Prod., IL: 113 (1825).

Holotype : Sieber No. 408. Locality, Nov. Holl. There are two sheets of
this material at Vienna, one of which was acquired via Reichenbach falls
possibly in 1889. Since Bentham described the species while in Vienna, it
would seem possible that his diagnosis of P. aspera was derived from the other
sheet, previously at Vienna; this deduction would be in accord with the fact
that Bentham’s description ‘. . . bracteis glabris . . . . calycibus vix pube-
Scentibus . . .’ fits the specimen at Vienna. The sheet at Kew bearing Sieber’s
number 408 is also labelled Wm. Mac Arthur, No. 13, Pultenaea asperata (sic) ;
no precise location is given, but another specimen of this is labelled Bargo
Brush, Mac Arthur. It does not agree with Bentham’s diagnosis with respect
to the bracteoles and calyx quoted above, but resembles the specimen W. F.
Blakely, The Valley, Hornsby, NSW 36368. These observations would place
the Kew sheet in group 9, in contrast to the specimen seen by Bentham in
Vienna, and the three sheets at MEL, which have been identified with group 6.

362 INVESTIGATION OF THE GENUS PHYLLOTA

P. BILLARDIERI Benth. in Ann. Wien. Mus., 1: 77 (1838).

The holotype of this binomial is at Vienna, having been collected by
Labillardiere, with no record of the locality. The predominantly glabrous
nature of the plant ‘.. . ramulis vix puberulis, foliis glabris . . . . bracteolis
glabris . . . . calycibus glabrisculis . . .’ would all seem to prohibit reference
of the name to group 9, all the members of which tend to be moderately to
markedly hirsute, at least in the calyx, and never wholly glabrous. The size
of the leaves, 10-0 mm. long, 1-5 mm. broad, pedicels 1mm. long, bracteoles
nearly ovate, with a breadth of 3mm., and an overall flower length of 9 mm.,
which would correspond to a standard length of 8-5-9-0 mm., all tend to exclude
the possibility of reference to group 9. A photograph of the holotype excluded
the possibility of groups 1 and 7, 8, though group 1 was in any case excluded
on the basis of the measurements. The diagnosis ‘ Spica oblonga subterminali ’
excludes group 5 from consideration, thus leaving the conclusion that this
epithet properly belongs with group 2, 3 or group 6, though insufficient data
are available to differentiate between these two groups.

P. PHYLICOIDES (Sieber ex DC.) Benth. in Ann. Wien. Mus., IL: 77 (1838).
Basionym: Pultenaea phylicoides Sieber ex DC. Prod., 11: 113 (1825).

Holotype: Sieber No. 405. Locality, Nov. Holl. There are three sheets
of this collection, all bearing Sieber’s original number, two at Vienna (one via
Reichenbach fil.) and one at Kew. There are three sheets at MEL and a second
sheet at Kew, which does not bear Sieber’s original label, but is marked ‘ ex
Herb. Mus. Vind.’. It bears no flowers, but is mounted with a specimen labelled
‘14, Pultenaea, Sydney, Hooker 1845’, which corresponds to group 5. The
Kew isotype possesses leaves of length 13mm., which are not mucronate.
This feature is characteristic of members of group 5, the leaves being obtuse
and with an extremely small black mucro which is very deciduous; this
characteristic appears in Bentham’s diagnosis as ‘. . . foliis obtusis’. The
diagnosis also states ‘ spicis brevibus terminalibus ’, a distinctive characteristic
of group 5 since any growth of the axis following production of an inflorescence
rapidly dies back, leaving the inflorescence effectively in a terminal position.
Finally, the numerous and small flowers (length overall 6mm.) are both
characteristic of group 5, which is in accord with the MEL isotypes.

P. BAUERI Benth. in Ann. Wien. Mus., IL: 77 (1838).

Now at Kew and ex Herb. Mus. Vind., the holotype is mounted on the
same sheet as another specimen collected by the U.S. Exploring Expedition,
under Wilkes, at Sydney, this second specimen also being labelled P. baueri.
The small leaves with minute mucro, and the small flowers in a subterminal
inflorescence, identify this name with group 2, 3. It consequently becomes
Synonymous with P. phylicoides.

P. sturtTH Benth. in Fl. Austr., Il: 95 (1864).

Holotype: C. Sturt, South Australia (K). Personal examination of the
holotype of this species identified it with groups 2, 3 although it is described
by Bentham as being between P. phylicoides and P. barbata. It is surprising
that no similar collections have been made in South Australia subsequent to
that of Sturt; possibly the location refers in fact to southern New South
Wales, though it is unlikely that Sturt collected in this region. There is,
however, no doubt as to the identity of the specimen with groups 2, 3, and
hence synonymous with P. phylicoides.

TAXONOMIC DISCUSSION
Intrageneric relationships of Phyllota Benth.

The species of Phyllota occurring on the east coast of Australia are
distinguished from the remaining species of the genus in a number of respects.

R. C. JANCEY 363

Adnation of the petals and stamens is well marked in the New South Wales
species, usually all 10 stamens being firmly united to the petals along the length
of the claws. In addition, the claws of one or both wings are not uncommonly
fused to that of the standard. In the remaining species, adnation is much
less common, being confined to 5 or fewer of the stamens attached to the base
of the petal claws, connation of the petal claws not occurring at all.

A distinction which may bear some relation to adnation of stamens is seen
in the persistence of petals and stamens after flowering in the east coast species.
This persistence was such that the remains were still present at maturity of
the legume, while in other species petals and stamens were deciduous soon
after enlargement of the legume began.

Final distinguishing features of the east coast species are seen in the shape
and texture of the bracteoles. The lanceolate, herbaceous bracteoles, common
to all these species with the exception of P. humifusa, are not found in the
remaining species. Many possess scarious or coriaceous bracteoles, e.g. P.
diffusa, P. remota, and P. pleurandroides, while those of the Western Australian
species are very similar to foliage leaves, except in P. lwehmannii, the bracteoles
of which were found to approach tne expanded leafy form found in the east.

P. barbata and P. gracilis in Western Australia represent another distinct
group of species, being distinguished by barbulate styles, and acute, narrowly
lunate keels. The relationship of these species is obscure, due to the lack of
material of P. gracilis, a species represented solely by the type collection.

Apart from the larger discontinuities in the genus described above,
interspecific distinctions within the genus appear, at least from a subjective
approach, to be at approximately the same level of significance, with size,
shape and texture of the bracteole being most useful in taxonomic discrimination,
followed by size and shape of the standard.

The relationship of Phyllota to the genera Pultenaea, Dillwynia, and Aotus
Discriminating characters

Leaves: A satisfactory distinction exists between Phyllota and Dillwynia
in that the leaves of Dullwynia, while being narrow-linear, are, without
exception, involute as opposed to revolute. The leaves of Aotus and of some
species of Pultenaea are similar to those of Phyllota.

Stipules : The presence or absence of stipules is not a reliable distinguishing
character between the genera quoted, though it has been used as such in the
past. All species of Phyllota have been found to possess minute stipules,
stipules of the same size being found in at least some species of Dillwynia and
Aotus. While Pultenaea has been characterized frequently as having more or
less obvious stipules, in many species they are minute, and in others quite
absent.

Bracis : Clear intergeneric distinctions are afforded by the form and texture
of the bracts, these being small, brown and scarious in Pultenaea, Dillwynia
and Aotus, also deciduous in the latter two genera. In Phyllota, however, the
foliage leaves subtending the flower are unaltered, except in two instances:
P. pleurandroides shows what may be an approach to the formation of
differentiated bracts in the virgate clusters of leaves with altered bases which
surround the flower ; on the specimen of P. georgii, now referred to P. luehmannii,
the floral leaves, and those immediately below the inflorescence, no longer show
the characteristic revolute form, but are nearly flat, with recurved margins.

Bracteoles: The range of bracteole form in the genus Phyllota has already
been discussed. The presence, in at least some of the species, of minute scarious
bracteoles removes this character as a source of intergeneric distinction, at
least from Pultenaea, since such bracteoles are also characteristic of the latter
genus. Dillwynia possesses similar bracteoles, but they are remote from the
calyx and deciduous, while Aotus is without bracteoles.

364 INVESTIGATION OF THE GENUS PHYLLOTA

Petals: No satisfactory distinctions may be made on the basis of petal
shapes or relative sizes, due to the variation within genera, the biggest distinction
existing between Phyllota and Dillwynia, where the orbicular to almost reniform
standard of Dillwynia is only approached by some species of Phyllota.

Stamens: Adnation of stamens to the petals is confined to the genus
Phyllota, though it has not been possible to establish the complete absence
of this phenomenon in the other three genera. It has been possible to show,
however, that there is considerable variation in the extent to which adnation
occurs within the genus Phyllota, varying from fusion of stamens to petals and
also of the petal bases themselves in P. phylicoides and the other New South
Wales species, to the situation found in P. diffusa where the adnation is so
sight as to be virtually undetectable, and certainly comparable with the
situation occurring in at least some members of the genus Pultenaea.

Style: As a result of the intrageneric variation associated with stylar
characteristics, and also the range of intraspecific variation associated with
this character, little discriminatory value attaches to it, except in the case of
Dillwynia which differs from the other genera in possessing a thicker, more
truneate style, and Aotws in which the style is generally filiform.

Seed: All the genera under discussion possess two ovules per ovary, of
which characteristicaly only one develops into a mature seed. The genera
are also united in possessing reniform seeds of similar size ranges. The
strophiole is a distinguishing character of limited value, being wholly absent
from Phyllota, wholly present as far as is known in Dillwynia and, with some
exceptions, present in Pultenaea and absent in Aotus.

The intergeneric relationships described above may be summarized as
follows :

The genus Pultenaea, by virtue of its greater size and more diverse nature,
appears to act as a central link joining the other three genera considered. Thus,
in the form of its bracteoles, Phyllota appears to be linked to Dillwynia and
Aotus via Pultenaea. <A similar series may be seen in the form of the bracts,
though there is much greater discontinuity in this case, in which no species
of Phyllota shows any leaf modification comparable with that found in Pultenaea.

In leaf form Phyllota and Aotus appear to be alike, and linked to Dillwynia
via Pultenaea. The small size or absence of stipules in Phyllota, Dillwynia
and Aotus would unite them with relation to the greater part of the genus
Pultenaea, though without creating any discontinuity, since similar forms are
also found in the latter genus. Since stamens are free in all the genera except
Phyllota, the positions of Phyllota and Pultenaea are the reverse of those obtaining
in the case of stipules.

Leaves.
Phyllota
Pultenaea —————__ Dillwynia
Aotus
Stipules.
Phyllota
Aotus ———— Pultenaea
Dillwyma
Bracts. ————— _ Aotus
Phyllota — ——— Pultenaea
Dillwynia
Bracteoles.
Phyllota =——— Pultenaea —————— Dillwynia ——————— Aotus
Stamens. ———— Dillwynia
Phyllota — —— Pultenaea
————— Aotus

R. C. JANCEY 365

Status of the genus Phyllota

The relationships of the genus have been considered in the previous section ;
whether these relationships are sufficiently distant to warrant generic rank is
debatable. It will be seen that the association is particularly close between
Phyllota, Aotus and Pultenaea. The distinction between Phyllota and Aotus is
acceptable at the generic level in that the two genera differ quite clearly in
the form of their bracts and of their bracteoles. These structures bear a
relationship to each other which leaves these two distinguishing characteristics
with rather less weight than might have been carried by a more dissimilar pair
of characters, but when considered in conjunction with a number of correlated
but less constant characteristics, for example, those concerned with the shape
of the standard and of the style, there is a dissimilarity which is of satisfactory
generic rank. The distinction between Phyllota and Pultenaea is less satis-
factory ; founded originally on differences in stipules, bracts, bracteoles, stamens
and seeds, discrimination can now be based only on the bracts. It would
seem doubtful if generic distinction could properly be based on one such
character, without support from a number of other, more or less constant
character differences. That the distinction is narrow is emphasized by some
Western Australian species of Pultenaea, e.g. P. dasyphylla (Turez.) C. A. Gardn.,
P. lycopodioides (S. Moore) and P. capitata (Turez.) C. A. Gardn., none of wnich
possess strophioles, and whose only apparent common difference from Phyllota
luehmannii rests in the recurved rather than revolute form of the leaves, a
distinction which, it will be recalled, was largely bridged by the specimen of
P. luehmannii originally attributed to P. georgi.

While there is no doubt that such doubtful generic limitations are
unsatisfactory, the generic status of Phyllota has been maintained in this study,
since it is felt that an evaluation of the generic status of Phyllota should form
part of an overall review of the Podalyrieae, or at least those members of the
tribe already mentioned and known to be allied to Pultenaea.

Formal Descriptions of Taxa

Characters used in formal descriptions have been largely confined to those
of value in intrageneric discrimination, those of generic distinction being con-
sidered in the section dealing with the status of the genus.

Of the specimens examined, only those belonging to collections known to
have been Jodged in European herbaria, or which are readily identifiable by
accession numbers to Australian herbaria, have been included in the citations
of specimens following the formal descriptions.

TAXONOMY
PuHyYLuotTA (DC.) Benth.

Fl. Austr., IL: 93 (1864); Engler et Pranti, Nat. Pflanz. Fam., Ill, 3:
210 (1894); Benth. et Hook., Gen. Pl., I: 470 (1865); Moore et Betche,
Handb. Fl. N.S.W., 135 (1893); Ewart, Fl. Vic., 640 (1931); Black, Fl. S.
Aust., IL: 442 (1948); Thompson in Contrib. N.S.W. Nat. Herb., Fl. N.S.W..,
No. 101: 45 (1961).

As Pultenaea sect. Phyllota DC., Prod., IL: 113 (1825); Curtis, Stud. Fl.
Tasm., 1: 132 (1956).

Shrubs, with stems terete, pubescent at least in the upper parts, echinate
with decurrent leaf bases. Leaves alternate, simple entire, linear, the margins
revolute ; stipules minute or absent. Flowers axillary, solitary or crowded
towards the ends of the branches, sometimes appearing terminal by death of
the distal axis; pedicels 1-5 mm. long; bracts identical with, or scarcely
differing from, foliage leaves; bracteoles 1-15mm. in length, scarious,
coriaceous or frequently herbaceous, inserted at the base of the calyx. Calyx
with the two upper lobes broader than the lower, and connate higher up.

366 INVESTIGATION OF THE GENUS PHYLLOTA

Corolla: petals all clawed, the standard ovate to orbicular, equal to or
somewhat exceeding the others. Stamens adnate to petals or scarcely so in
Some species. Ovary sessile, pubescent to villous, ovules 2, on short funicles,
the style dilated or thickened at the base, incurved and subulate above, the
stigma small and terminal. Legume inflated, twice as long as the calyx at
maturity, containing 1-2 seeds. Seed reniform, not strophiolate.

A genus of 10 species, endemic in Australia.

Key to the Species of Phyllota

A. Style bearded upwards on inner edge
B. Flower 12mm. long, pedicel less than 1:5 mm. long .. . barbata

- gracilis

re) el

B’. Flower 5mm. long, pedicel 4mm. long, exceeding the calyx

A’. Style not bearded on inner edge
C. Bracteoles herbaceous, linear or lanceolate
D. Stem, bracteoles and calyx yellow tomentose, flowers in dense heads,
keel purple, petals and stamens deciduous after flowering .. .. P. luehmanni

D’. Stem glabrous to pubescent, bracteoles and calyx glabrous to villous,
if flowers in dense heads then keel not purple, petals and stamens
persisting until maturity of legume
Flowers in lax spikes towards the ends of the branches, leaf tips
acuminate and recurved, numerous root suckers formed .. .. P. squarrosa

EK’. Flowers scattered, in lax spikes or dense terminal heads, leaf tips
never acuminate and recurved, root suckers absent or few from a
thickened root-stock
F. Procumbent shrub with purple-red corolla; flowers ta or few
together, bracteoles linear ise P. humifusa
F’. Erect shrub with flowers in dense ayilees at or Viommnds ie ends ai
the branches
G. Leaves massive, 1:25-2:25 mm. broad, standard 10-15 mm. lone.

calyx densely villous .. P. grandiflora
G’. Leaves slender, 0-75-1- Se nina. istoaa: iacataeel 5- lL 5 darn long,
calyx glabrous or pubescent .. : 5 P. phylicoides
C’. Bracteoles scarious or coriaceous, not green
H. Bracteoles oblong-ovate, as long as the calyx, keeled, mucronate... P. remota

H’. Bracteoles ovate, less than 2mm. long, not keeled
I. Flowers solitary or in pairs, in virgate clusters of leaves along the

stem; leaves recurved and acuminate. . ae whe au .. P. pleurandroides
I’. Flowers in spikes towards the ends of the branches; leaf tips not
recurved, obtuse .. a Se by A se is .. P. diffusa

PHYLLOTA PHYLICOIDES (Sieber ex DC.) Benth.

In Ann. Wien. Mus., IL: 77 (1838); Benth., Fl. Austr., IL: 95 (1864) ;
Moore et Betche, Handb. Fl. N.S.W., 135 (1893) ; Thompson in Contrib. V.S.W.
Nat. Herb., Fl. N.S.W., No. 101: 45 (1961).

Nomenclatural synonym: Pultenaea phylicoides Sieber ex DC., Prod., II:
113 (1825). BASIONYM.

Taxonomic synonyms : Phyllota bauert Benth. in Ann. Wien. Mus., 11: 77
(1838); Phyllota billardieri Benth. in Ann. Wien. Mus., Il: 77 (1838);
Phyllota comosa (Sieber ex DC.) Benth. in Ann. Wien. Mus., II: be (1838) ;
Phyllota aspera (Sieber ex DC.) Benth. in Ann. Wien. Mus., IL: 77 (1838) ;
Phyllota sturtu Benth. Fl. Austr., IL: 95 (1864).

A shrub 90cm. (15-165 em.) high, with stems terete, pubescent at least
in the upper parts. Leaves linear, 10mm. (5-5-19 mm.) long, 1mm. (0-75-
1-25 mm.) broad, bullate, obtuse to acute (1-3 on the subjective scale) ; stipules
minute. Flowers 23 (11-83), crowded together into leafy spikes 13 mm.
(8-45 mm.) long towards the ends of the branches. Pedicel 1mm. (0-5-1-5
mm.) long. Bracts identical in appearance with foliage leaves; bracteoles
lanceolate, 7:-5mm. (4:0-11:5mm.) long, 1:-7mm. (0-:5-3-5mm.) broad,
herbaceous with scattered or numerous short appressed silky hairs, borne on
the base of the calyx. Calyx 5-3mm. (3-5-9-0 mm.) long, almost glabrous

R. C. JANCEY 367

or with scattered to numerous short appressed silky hairs; lower lobes
acuminate, longer than, or equal to, or shorter than the tube; upper lobes
broader connate higher up and less acuminate. Corolla: keel equal in length
to standard, broadly lunate to semi-circular, obtuse, yellow to yellow-green ;
wings equal in length to standard, oblong to semi-circular, laciniate at base,
obtuse and rounded, sometimes almost acute, yellow; standard 8mm.
(5-11-5mm.) long, ovate, obtuse and rounded, yellow or yellow with red
markings. Stamens 10, some or all adnate to petals at base, both persistent
after flowering. Ovary villous, style dilated or thickened at base, incurved
or subulate above, pubescent with short appressed silky hairs below the curve.
Legume 1-2 seeded, 1-2 times as long as the calyx. Seed reniform.

Distribution: Coast and tablelands of New South Wales and Queensland
to Bundaberg.

Habitat : Sandstone heath and dry sclerophyll forest.

Chromosome number: 2n=14, voucher specimens R. C. Jancey No. 2;

R. Carolin No. 3933; BR. C. Jancey No.3; V. Sands sine num.; R. C. Jancey
No. 4; R. C. Jancey No. 5 (SYD).

Typification: Holotype: Sieber No. 405 GEN. Isotypes K, WIEN.

Selected specimens examined: New South Wales: Neutral Bay, J. B.
Cleland, 9/1910, (AD 96311325); Port Jackson, R. Schomburgk, 8/1896,
(AD 96311334); Shoalhaven, W. Bauerlen, No. 396, 9/1883, (MEL); Kurra-
jong, Miss Atkinson, No. 13, —, (MEL); Richmond River, Mrs. Hodgkinson,
1874, (MEL); Long Bay, Miss C. Cowle, 1907, (MEL); Sandy Cape, R. Brown,
—, (MEL); FI. Nov. Holl., Sieber, No. 407, (MEL); FI. Nov. Holl., Sieber,
No. 408, (MEL); Mitchell’s Expedition of 1836, Mitchell 291, 8/1836, (MEL) ;
Nov. Holl., Lambert, —, (MEL); Nov. Holl., Sieber, No. 405, —, (MEL) ;
Caloundra, L. J. Brass, 10/1934, (CANB 24244); French’s Forest, G. H. Clarke,
9/1920, (CANB 4535); Middle Harbour, G. H. Clarke, 12/1920, (CANB 4534) ;
Oxford Falls, K. Mair, 26/8/1953, (NSW 36444); Wahroonga, L. A. 8S. Johnson,
23/6/1945, (NSW 36445); Terrey Hills, M. Tindale, 12/8/1961, (NSW 55356) ;
Dural, D. C. Cross, 5/9/1945, (NSW 15668); Cheltenham, N. C. Ford, 4/7/1945,
(NSW 36421); Castlecrag, M. Tindale, 1/8/1948, (NSW 7227); Mount Colah,
G. Chippendale, 18/8/1953, (NSW 36398); Berowra, R. H. Cambage, No. 499,
9/1901, (NSW 36429); Asquith, F. J. Thomas, 24/8/1951, (NSW 36434) ;
Currockbilly, J. L. Boorman, 2/1910, (NSW 36393); Snowball, F. A. Rodway,
No. 11734, 12/1940, (NSW 36395); Cooma, J. L. Boorman, 12/1915, (NSW
35391); Nerriga, F. A. Rodway, No. 13467, 3/1944, (NSW 36394); Como,
J. L. Boorman, 9/1916, (NSW 36412); Gymea Bay, A. Cahill, 10/1938, (NSW
36409) ; National Park, Anderson and Boorman, 9/1921, (NSW 36413);
Glenbrook, A. A. Hamilton, 10/1914, (NSW 36432); Blaxland, Blakely and
Chisholm, 10/1929, (NSW 36426); Bundanoon, H. E. Ellen, 3/1917, (NSW
36448); Pigeon House Ra., Nerriga, E. F. Constable, 10/1957, (NSW 45273) ;
Jervis Bay, F. A. Rodway, 10/1931, (NSW 36403); Wentworth Falls, W. F.
Blakely, 11/1938, (NSW 36480); Box Point to Barber’s Creek, J. H. Maiden,
10/1896, (NSW 36450); Appin, J. H. Maiden, —, (NSW 15662); Wiseman’s
Ferry, J. L. Boorman, 4/1908, (NSW 36433); Gosford, Blakely and Shiress,
1/1927, (NSW 15665); Woy Woy, Blakely and Buckingham, 10/1939, (NSW
15667); Somersby, G. Chippendale, 8/1953, (NSW 36418); Maroota, W. F.
Blakely, 9/1929, (NSW 36439); Corindi, E. J. Constable, 11/1956, (NSW 4221) ;
Bulladelah, J. Garden, 10/1951, (NSW 36379). Queensland: Sunnybank nr.
Brisbane, C. T. White, No. 985, 9/1921, (NSW 36390); Sunnybank, L. A. 8.
Johnson, 6/1951, (NSW 36385); Mt. Gravatt, L. A. S. Johnson, 6/1951, (NSW
36386); Moreton Island, C. T. White, 9/1907, (NSW 36388); Stradbroke
Island, H. S. McKee, No. 8725, 9/1961, (NSW 56450); Burrum, M. E. Watson,
10/1929, (BRI 036876) ; South Brisbane Cemetery, F. M. Bailey, 3/1875, (BRI
036889) ; Apsley, C. T. White, No. 6133, 7/1929, (BRI 036875) ; Coolum, Miss

368 INVESTIGATION OF THE GENUS PHYLLOTA

M. S. Clemens, 4/1945, (BRI 036871); Mt. Gravatt, C. T. White, No. 7409,
3/1931, (BRI 036868); The Blunder, nr. Brisbane, C. E. Hubbard, No. 3584,
8/1930, (BRI 036863); Plunkett, C. E. Hubbard, No. 3784, 8/1930, (BRI
036862); Tingalpa, D. A. Goy, No. 134, 9/1936, (BRI 036860); Maryborough,
Miss M. 8. Clemens, 9/1948, (BRI 036857); Capalaba, L. Pedley, No. 426,
8/1959, (BRI 024332); Keppel Bay, C. T. White, No. 8032, 9/1931, (BRI
036882); Bundaberg, J. Keys, No. 336, —, (BRI 036887).

PHYLLOTA GRANDIFLORA Benth.
In Ann. Wien. Mus,, IL: 77 (1838).

Taxonomic synonym: Phyllota pilosa Benth. in Ann. Wien. Mus., Il: 77
(1838).

A shrub 90cm. (60-150 cm.) high, stems pubescent at least in the upper
parts. Leaves linear, 13 mm. (6-17-5 mm.) long, 1-25—2-25 mm. broad, densely
minute-bullate, obtuse or acute (1-3 on the subjective scale); stipules minute.
Flowers 23 (6-59) in lax leafy spikes towards the ends of the branches. Pedicel
1-2 mm. long. Bracts identical in appearance with foliage leaves; bracteoles
lanceolate, 11mm. (8:5-15-0mm.) long, 2-7mm. (2-0-4:0mm.) broad,
herbaceous, with scattered to numerous appressed silky hairs, borne on the
base of the calyx. Calyx 8:2mm. (6-:0-10:0mm.) long with numerous
appressed silky hairs; lower lobes acuminate, equal to or shorter than the
tube; upper lobes broader, connate higher up, acuminate. Corolla: keel
equal in length to standard, broadly lunate to semi-circular, obtuse, yellow
occasionally tinged with green; wings shorter than or equal to the standard,
semi-circular, laciniate at the base, obtuse and rounded, yellow; standard
12-4mm. (10-5-15-:0 mm.) long, broadly ovate, obtuse and rounded, yellow
or yellow with red marking. Stamens 10, some or all adnate to petals, both
persistent after flowering. Ovary villous, style dilated and thickened at base,
incurved or subulate above, pubescent with short silky appressed hairs, often
extending along upper edge around the hook. Legume 1—2 seeded, 1-2 times
aS long as the calyx. Seed reniform.

Distribution : Between Sydney Harbour and the Hawkesbury River, also
Appin and Bargo regions (see Fig. 10).

Habitat : Sandstone heath and dry sclerophyll forest.

Chromosome number : 2n=14, voucher specimen R. C. Jancey, No. 7 (SYD).

Typification: Holotype: Loc. non cit. F. Bauer, WIEN.

Specimens examined: New South Wales: Parramatta, W. Woolls, —,
(MEL); Parramatta, W. Woolls, —, (MEL); Elanora Heights, V. May,
10/1934, (NSW 36366); Port Jackson, F. J. Sargood, 10/1911, (NSW 36362) ;
Berowra, W. F. Blakely, 10/1940, (NSW 36369) ; Narrabeen, M. Mills, 10/1940,
(NSW 36365); Narrabeen, J. J. Fletcher, 8/1887, (NSW 36364); The Valley,
Hornsby, W. F. Blakely, 11/1939, (NSW 36368); Hornsby, E. Betche, 12/1886,
(NSW 36436); ‘‘ West Australia”’, Maxwell, —, (NSW 36359); St. Ives,
Blakely and Anderson, 9/1936, (NSW 36357); Manly, E. Cheel, 10/1898, (NSW
36363); Hornsby, W. F. Blakely, 10/1914, (NSW 36371); Cheltenham, Ll. A.
S. Johnson, 10/1945, (NSW 36370); Field of Mars, H. Deane, —, (NSW 36367) ;
National Park, W. F. Blakely, 11/1938, (NSW 36361); Cataract Dam, J. H.
Maiden, 11/1906, (NSW 36360).

PHYLLOTA SQUARROSA (Sieber ex DC.) Benth.
In Ann. Wien. Mus., IL: 77 (1838).
Nomenclatural synonym : Pultenaea squarrosa Sieber ex DC., Prod., IL: 113
(1825). BASIONYM.

A shrub 30cm. (15-60 cm.) high, suckering freely from the roots, stems
pubescent at least in the upper parts. Leaves linear, 9mm. (6-6-13-8 mm.)

R. C. JANCEY 369

long, 0:75-1:25 mm. broad, minutely bullate, acuminate and recurved (4 on
the subjective scale), stipules minute. Flowers 8 (3-12) together towards the
ends of the branches in lax leafy spikes 8mm. (2-18 mm.) long. Pedicel
0:75-1:75 mm. long. Bracts identical in appearance with foliage leaves ;
bracteoles lanceolate, 8-25 mm. (5-5-13-0 mm.) long, 1-25 mm. (1-0-2-5 mm.)
broad, herbaceous, glabrous or with scattered to numerous short appressed
silky hairs, borne on the base of the calyx. Calyx 7-5mm. (5-5-9-0 mm.)
long, almost glabrous or with scattered to numerous short appressed silky hairs ;
lower lobes acuminate, equal to or longer than the tube; upper lobes broader,
connate higher up, acuminate. Corolla: keel equal in length to standard,
semi-circular, obtuse, yellow occasionally tinged with red; wings shorter than
or equal to the standard, broadly lunate, obtuse and rounded, yellow ; standard
10-3 mm. (8-0-12- 0 mm.) long, broadly ovate, obtuse and rounded, yellow to
red. Stamens 10, some or all ‘adnate to petals, both persistent after flowering.
Ovary villous, style dilated and thickened at base, incurved or subulate abowe,
pubescent with short appressed silky hairs to below the hook. Legume 1-
seeded, 1-2 times as long as the calyx. Seed reniform.

Distribution : Upper Blue Mountains, central tablelands.

Habitat: Sandstone heath and open dry sclerophyll forest.

Chromosome number : 2n=—14, voucher specimen R. C. Jancey, No. 6 (SYD).
Typification: Holotype: Blue Mountains, Sieber, No. 406 (GEN).

Selected specimens examined: New South Wales: Clarence, F. H. Rodway,
—, 1908, (AD 96311374); Mt. Tomah, W. Woolls, —, (MEL); Blackheath,
Althofer, 8/1945, (MEL); Nov. Holl., Sieber, No. 406, (MEL); Bell, L. A. §.
Johnson, 5/1951, (NSW 36470); Eskbank, A. A. Hamilton, 1/1915, (NSW
36485); Mt. Victoria, J. H. Maiden, 12/1896, (NSW 36452); Mt. Victoria, J.
L. Boorman, 12/1917, (NSW 36457); Bell, A. A. Hamilton, 1/1915, (NSW
36458); Mt. Wilson, ie H. Maiden, 4/1896, (NSW 36459); Clarence hae
WE: Blakely, 11/1938, ( (NSW 36463); Newnes Junction, Blakely and Bucking-
ham, 11/1938, (NSW 36483); Mt. 'Piddington, Mt. Victoria, Blakely and
Buckingham, 1/1939, (NSW 15660) ; Mitchell's Ridge, Blakely and Buckingham,
1/1939, (NSW 36472); Katoomba, W. Forsyth, 12/1899, (NSW 15659) ;
Katoomba, J. H. Camfield, 12/1908, (NSW 36469) ; Blackheath, J. H. Maiden,
1/1905, ( (NSW 36471) ; Narrow Neck, G. Ontppendale, 1/1951, (NSW 36474);
nr. Bald Trig., Clarence, Were Blakely, 1/1939, (NSW 36462).

PHYLLOTA HUMIFUSA Benth.
Fl. Austr., 11: 95; Moore and Betche. Handb. Fl. N.S.W., 135 (1893).

A prostrate shrub 8-15 cm. high, with stems pubescent at least in the upper
parts. Leaves linear, 4:5mm. (3-0-8:0mm.) long, 0-25-0-75 mm. broad,
minutely bullate, obtuse to acute (1-3 on the subjective scale) ; stipules minute.
Flowers 8 (2-15) in lax leafy spikes, 7mm. (1-14 mm.) long, towards the ends
of the branches. Pedicel 0-5-1-25 mm. long. Bracts identical in appearance
with foliage leaves; bracteoles identical in appearance with foliage leaves,
3mm. (2-4:5mm.) long, 0-5mm. (0:25-0-6mm.) broad, glabrous or with
scattered short appressed silky hairs, borne on the base of the calyx. Calyx
5:0mm. (4:0-5-5 mm.) long, almost glabrous or with scattered short appressed
silky hairs; lower lobes acuminate, shorter than or equa! to the tube; upper
lobes broader, connate higher up and acute. Corolla: keel equal in length to
standard, semi-circular, obtuse and rounded, yellow to yellow-red ; standard
7-5mm. (7-0-8-0 mm.) long, broadly ovate, obtuse and rounded, red to deep
red. Stamens 10, some or all adnate to petals, both persistent after flowering.
Ovary villous, style dilated and thickened at base, incurved or subulate above,
pubescent with short appressed silky hairs to below the curve. Legume 1-2
seeded, 1-2 times as long as the calyx. Seed reniform.

i

370 INVESTIGATION OF THE GENUS PHYLLOTA

Distribution : Southern tablelands of New South Wales, between Mittagong
and Mt. Bullio (see Fig. 4).

Habitat: Deep sandy shale soil in dry sclerophyll forest and in open
sparsely grass covered areas.

Chromosome number : 2n=14, voucher specimen R. C. Jancey, No. 1 (SYD).

Typification : Holotype; Wombat Brush, Argyll County, A. Cunningham,
No. 8 (K).

Specimens examined: Mittagong to Bullio, H. Cheel, 11/1919, (NSW 36449) ;
Penrose, Blakely and Buckingham, 11/1939, (NSW 36447).

SPECIES OF PHYLLOTA NOT OCCURRING IN NEw SouTtTH WALES

All observations on interstate material were based on herbarium specimens
on Joan from the various State herbaria. A list of material examined, and its
origin, is included in the description of each species.

Methods

Since the material in question was on loan from other herbaria, some of
the measurements which had been carried out on material collected personally
could not be repeated, owing to the amount of destruction involved. In other
cases, Specimens were so small that it was felt that any further disintegration
would be undesirable and, consequently, such specimens were only examined
superficially under a dissecting microscope.

Measurements were made on material soaked in a mixture of detergent
and water, scored values referring to the same scales as those used in connection
with the New South Wales material. To avoid undue destruction of loaned
material, the whole range of material was first examined, then measurements
made on a limited number ot individuals. In many eases, variation in floral
parts was negligible, and in these cases single values have been quoted. Where
this was not the case, or in the case of other more variable structures, values
are quoted for the range of variation, and one for an individual apparently
representative of the mean for that particular character. A number of
observations other than those employed in the case of the New South Wales
taxa were introduced after preliminary examination of the interstate species.

PHYLLOTA BARBATA Benth.

In Hueg. Enum. 33 (1837), and in Ann. Wien. Mus., IL: 78 (1838), and
in Fl. Aust. IL: 94 (1864).

Taxonomic synonyms: Pultenaea andrews Gardn. ex Blackall and Grieve
‘How to know West Australian Wildflowers’, 234 (1953). NOM. NUD. ET
ILLEGIT.

A shrub with stems terete, pubescent at least in the upper parts. Leaves
linear, 8mm. (6-10 mm.) long, scabrous, with revolute margins, obtuse and
rounded, some also bearing a minute deciduous black mucro (1 on the subjective
scale); stipules minute. Flowers scattered along the branches, sometimes
crowded into leafy spikes towards the ends of the branches. Pedicel 0-5-1-0
mm. long. Bracts identical in appearance with foliage leaves; bracteoles
linear, 8mm. long, 1mm. broad, almost identical in appearance with foliage
leaves, borne on the base of the calyx. Calyx 5mm. long, almost glabrous
to heavily tomentose; lower lobes acuminate, longer than, or equal to, or
Shorter than the tube; the upper lobes broader connate higher up and less
acuminate. Corolla: keel 12mm. long, tapering to an acute point, red;
wings 7-8 mm. long, cuneate-oblong in their upper parts, yellow red; standard
12 mm. long, elliptic, yvellow-red. Stamens 10, some adnate to petals at base,
both deciduous after flowering. Ovary villous; style villous below and
barbulate in the distal half on the upper edge with persistent white hairs.
Legume 1-2 seeded, as long as or longer than the calyx. Seed reniform. |

R. C. JANCEY 371

Distribution : Coastal south-western area of Western Australia.
Habitat: Sandy heath.

Chromosome number: 2n=14. Voucher specimen, V. Sands, No. 638/19/4.
Determined by V. Sands (unpublished).

Typification : Holotype.

Discussion: P. barbata is distinguished from all other species of Phyllota
with the exception of P. gracilis Turez. by the bearding of the distal half of
the style. From P. gracilis it differs in the length of the peduncle, and in the
flowers which are almost sessile in P. barbata but borne on a pedicel 4 mm. long
in P. gracilis. These latter two species also differ in leaf length. Only one
specimen of P. andrewsti was available for examination (Gardner, 2219). Having
been collected by the author of the manuscript name, and bearing his determi-
nation, it may be taken, however, as an authoritative example of the intended
taxon. The specimen examined was bearded on the inner edge of tue style,
as is characteristic in P. barbata and P. gracilis, whereas, in ‘ How to Know
West Australian Wildflowers ’, P. andrewsii is distinguished by two characters,
one of which is the absence of bearding. The position of the inflorescence,
however, was found to be in agreement with the text quoted, being terminal
in the specimen of P. andrewsti, as opposed to the rather lax and interrupted
spikes characteristic of P. barbata. Since the distinction proposed between the
species was based, at least in the case of the specimen examined, on one character,
it is not proposed at this stage to give formal status to the name proposed by
Gardner. There seems little doubt that the range and degree of morphologic
differentiation in this section of the genus in Western Australia is not as yet
fully represented in herbarium collections.

Specimens examined : Cape Riche, R. T. Lange, No. 13, 3/1958, (PERTH) ;
Albany, W. EH. Blackall, 12/1937, (PERTH); Mt. Manypeaks area, S. P. Pfeiffer,
No. 12, —, (PERTH); Narrikup, R. T. Lange, No. 13, 3/1958, (PERTH) ;
Cheyne Beach, J. M. Storr, No. 3900, 5/1959, (PERTH); King George’s Sound,
B. T. Goadby, No. 92, 2/1899, (PERTH); Sand heath south of Stirling Range,
W. EH. Blackall, 4/1939, (PERTH); Nr. West Mount Barren, C. A. Gardner,
10/1928, (PERTH); Nr. Albany, Maxwell, No. 33, 7/1858, (MEL); King
George’s Sound, A. Hugham, No. 22, 1869, (MEL); South West Australia,
Mills, 6/1861, (MEL); Cape Riche, Preiss, No. 846, 1843, (MEL); Sand Plains,
Wilson’s Inlet, Oldfield, No. 766, —, (MEL); Bremer River, Webb, 1884,
(MEL); Phillip’s River, C. R. Andrews, 10/1903, (NSW 36498); King River
Road, F. Staer, 2/1911, (NSW 36519); King George’s Sound, Rev. R. Collie,
1898, (NSW 36513); Western Australia, Drummond, No. 84, —, (NSW 36514) ;
Torbay, A. E. Sheath, 1/1903, (NSW 36515); Warejinup, —, —, (NSW 36517) ;
South West Plantagenet, E. Pritzel, No. 207, 1/1901, (NSW 36518); Western
Australia, Drummond, No. 85, (4th collection), 1848, (NSW 36512).

PHYLLOTA GRACILIS Turez.

In Bull. Soc. Nat. Mose., XXVI: 1267 (1853); Benth, Fl. Austr., II: 94
(1864).

Nomenclatural synonym: Pultenaea gracilis (Turez.) Gardner, Enum. Pl.
Austr. Oce., 59 (1930).

A shrub, with stems terete, pubescent with short white hairs (densely so
in the upper parts); scarcely rugose with decurrent leaf bases (1 on the
subjective scale). Leaves linear, minute, 1-3 mm. long, 0-5 mm. broad, with
a scattered pubescence, revolute margins, obtuse and rounded bearing a
deciduous black mucro (1 on the subjective scale); stipules minute. Flower
solitary, near end of branch. Pedicel4mm.long. Bracts identical in appearance
with foliage leaves; bracteoles linear-lanceolate, 1-5 mm. long, herbaceous,
sparsely pubescent, borne on the base of the calyx. Calyx 3 mm. Jong, densely
villous with pale yellow simple hairs; lower lobes acute, equal to or longer

302 INVESTIGATION OF THE GENUS PHYLLOTA

than the tube; upper lobes acute, equal to or shorter than the tube. Corolla
not seen dissected; 5mm. long from receptacle to apex of keel; keel dark
red; wings dark red; standard dark red. Ovary not seen; style bearded
-in distal half. Legume not seen. Seed not seen.

Distribution : Not known.

Habitat: Not known.

Chromosome number : Not known.

Typification: Holotype: Swan River, Western Australia, Drummond,
No. 91, ca. 1845. KW. Isotypes MEL, K.

Discussion: While this species shows affinities to P. barbata im the bearded
style and boat-shaped keel, it shows striking morphologic distinction in the
very much smaller bracteoles, the lesser overall length of flower, very much
longer pedicel and much shorter leaves. The differentiation from P. barbata
is such as to merit specific rank.

Specimen examined: Swan River, Western Australia, Drummond, No. 91,
ca. 1845, (MEL).

PHYLLOTA LUEHMANNED F. Muell.
Fragm. Phytogr. Austral., X: 33 (1877), as Phyllota luehmannii.

Nomenclatural synonym: Pultenaea luehmanni (F. Muell.) Gardner, Hnum.
Pl. Austr. Occ., 59 (1930).

Taxonomic synonym: Phyllota georgv Hemsl. in Hook. Ic. Pl., t. 2778
(1905). Pultenaea luehmannii var. georgii Gardn. ex Blackall and Grieve ‘ How
to Know West Australian Wildflowers’, 234 (1953). NOM. NUD. ET ILLEGIT.

A shrub with stems terete, covered with a pale gold tomentum at least
in the upper parts. Leaves linear, 6 mm. (4-10 mm.) long, scarcely scabrous,
bearing a sparse pale golden tomentum, with revolute margins; leaf tips
acuminate recurved (4 on the subjective scale); stipules minute. Flowers
crowded into spikes at the ends of the branches becoming terminal by die-back
of the axis. Pedicel 0-5-1-0mm. long. Bracts identical in appearance with
foliage leaves, occasionally with somewhat less revolute margins; bracteoles
linear-lanceolate, 4—6 mm. long, 0-5 mm. broad, herbaceous, with a pale golden
tomentum, borne on the base of the calyx. Calyx 6mm. long, densely
tomentose with a pale golden tomentum ; lower lobes acuminate, about equal
in length to the tube; the upper lobes broader connate higher up and less
acuminate. Corolla: keel 8mm. long, obtuse, purple; wings equal to or
(more usually) 1mm. shorter than the keel, cuneate-oblong in upper parts,
yellow ; standard orbicular, 11 mm. long, yellow with a red marking at the
base. Stamens 10, some slightly adnate to the petals, both deciduous after
flowering. Ovary densely villous; style villous below, glabrous in the distal
half, Legume 1-2 seeded, as long as or longer than the calyx. Seed reniform.

Distribution : The Victoria Desert region of Western Australia.

Habitat : Sand plain.

Chromosome number: 2n=14, also n=7. Voucher specimen V. Sands, No.
639/1/5 (SYD). Determined by V. Sands (unpublished).

Typification: Syntypes of Phyllota luehmanni F. Muell.: Near Waring,
Western Australia, F. Mueller; Elder Exploring Expedition, Victoria Desert
Camp 58, R. Helms sine num., 21/9/91. Lectotype: Elder Exploring Expedi-
tion, Victoria Desert Camp 58, R. Helms sine num., 21/9/91. MEL. Isolecto-
types: K., AD

Phyllota georgui Hemsl. Holotype: Railway between Cunderdin and Dedari,
G. H. Thistleton-Dyer, K.

Discussion : There appear to be no features distinguishing P. georgii from
P. luehmannu, at least among the specimens examined. The short, obtuse

R. C. JANCEY ites

leaves quoted in ‘ How to Know West Australian Wildflowers ’ as characteristic
of P. luehmannit var. georgii were found to be so variable within specimens
as to be unsatisfactory for discriminatory purposes.

Specimens examined: Karalee, C. A. Gardner, 9/1934, (PERTH); East-
west railway, C. French, —, (MEL); No. 15 Pumping Station, Yerbillon, M.
Koch, No. 2892, 10/1923, (MEL); Victoria Desert Camp 58, Elder Exploring
Expedition, R. Helms, 9/1891, (MEL); Karoling, Elder Exploring Expedition,
R. Helms, 11/1891, (MEL); Victoria Springs, Young, 10/1875, (MEL).

PHYLLOTA PLEURANDROIDES F. Muell.
In Trans. Phil. Soc. Vic., 1: 38 (1833); Benth., Fl. Austr., IL: 96 (1864).

A shrub, suckering freely from the roots. Stems terete, more or less
pubescent. Leaves scattered to virgate, linear, 8mm. (6-10mm.) long,
0-75-1-25 mm. broad, bullate, tip acuminate and recurved; stipules minute.
Flowers scattered among the virgate clusters of leaves. Pedicel 0-5-1-:0 mm.
long. Bracts similar in appearance to foliage leaves, somewhat narrowed and
villous towards the base, as are other leaves of the virgate clusters ; bracteoles
ovate-obtuse, 1-5 mm. long, 1 mm. broad, coriaceous, pubescent, borne on the
base of the calyx. Calyx 4mm. long, villous with white hairs; lower lobes
acute, equal in length to the tube; upper lobes broader and connate higher
up. Corolla: keel 6mm. long, obtuse, yellow-red ; wings slightly exceeding
the keel, oblong in upper parts; standard broadly ovate, 5mm. long, yellow.
Stamens 10, some adnate to petals at base of claw, both deciduous after
flowering. Ovary villous; style villous below, glabrous in upper parts.
Legume 1-2 seeded, 1-2 times as long as the calyx. Seed reniform.

Distribution : South-eastern parts of South Australia, and south-western
Victoria.

Habitat: Deep sandy soil on sand ridges of sand plain.

Chromosome number : Not known.

Typification : Syuntypes: Kangaroo Island, F. Mueller, Herb. W. Sonder :
Grampians, Wilhelm, F. Mueller, 1857: Mount Abrupt, F. Mueller, MEL, K.
It is proposed to select Mount Abrupt, F. Mueller, MEL as lectotype of this
species. Isolectotype: Mount Abrupt, F. Mueller, K.

Selected specimens examined: South Australia: Bool Lagoon-Lucindale,
D. Hunt, No. 796, 5/1962, (AD 96227095); Southern Mt. Lofty Range, nr.
Mt. Compass, —, 1/1882, (AD 96311376); South Australia, —, —, (AD
96311378) ; Kangaroo Island, —, No. 1217, 3/1884, (AD 96311375); Lower
Mt. Lofty Range, nr. Strathalbyn, HE. C. Black, 2/1944, (AD 96311330) ;
Malinong, 45km. south of Murray Bridge, R. D. Sharrad, No. 13, 8/1960,
(AD 96149180); Encounter Bay, J. B. Cleland, 11/1924, (AD 96311377) ;
Kangaroo Island, Mt. Pleasant, —, 1/1883, (AD 96311380); Bordertown, D.
Hunt, No. 748, 3/1962, (AD 96220087); Mt. Abrupt, F. Mueller, —, (MEL):
Square Waterhole, O. Tepper, No. 30, 7/1882, (MEL); Gawler Ranges, Dr.
Sullivan, —, (MEL); Lacepede Bay, Herschel and Babbage, —, (MEL) ;
Penola, Rev. Tenison-Woods, No. 15, —, (MEL); N.W. of Lake Albacutya,
C. French, 10/1887, (MEL) Victoria: Grampians, H. B. Williamson, 4/1904,
(MEL); Grampians, J. W. Audas, 11/1920, (MEL); Mt. Zero, C. Wilhelmi,
2/1857, (MEL); Wimmera, D’Alton, No. 16, 1890, (MEL); West of Wimmera,
D’ Alton, 7/1892, (MEL); Shire of Dimboola, F. M. Reader, 1/1893, (MEL) ;
Keith, R. L. Crocker, 9/1943, (CANB 11633).

PHYLLOTA REMOTA J. H. Willis
in Vie; Naot., XXII: 191 (1957).

A shrub, with stems terete, tomentose at least in the upper parts, rugose
with decurrent leaf bases (2 on the subjective scale). Leaves linear, 8 mm.
(5-10 mm.) long, distant, occasionally becoming virgate, tomentose when young,

374 INVESTIGATION OF THE GENUS PHYLLOTA

papillose with age, revolute margins, acute to obtuse, sometimes bearing a
minute black mucro (2 on the subjective scale); stipules minute. Flowers
scattered, solitary or occasionally in pairs. Pedicels 0-:5-1-0mm. long.
Bracts identical in appearance with foliage leaves; bracteoles ovate, 4 mm.
long, scarious, keeled and with a mucronate apex, almost enveloping the calyx.
Jalyx 3-4 mm. long, glabrous to villous; lower lobes acute, more or less equal
to the tube; upper lobes broader and connate higher up. Corolla: keel
6 mm. long, obtuse, yellow; wings slightly exceeding the keel, oblong in upper
parts ; standard broadly ovate, 5mm. long, yellow. Stamens 10, some adnate
to petals at base, both deciduous after flowering. Ovary villous; style villous
below, glabrous in upper parts. Legume 1—2 seeded, twice as long as the calyx.
Seeds reniform.

Distribution : South-eastern South Australia and south-western Victoria.
Habitat: In shallow sandy soil between sand ridges of mallee heath.
Chromosome number: Not known.

Typification: Holotype: Keith, R. L. Specht and P. Rayson, 1954, (MEL).

Specimens examined: Boston Point, Spencer’s Gulf, Wilhelmi, —, (MEL) ;
Lillimur, nr. Wimmera, A. J. Hicks, 9/1954, (MEL); Eyre Peninsula, 85 km.
north of Port Lincoln, R. L. Specht, No.. 2602, 11/1960, (AD 96109031); Dark
Island, 14km. east of Keith, Specht and Rayson, 2/1950, (AD 96311331) ;
11 km. east of Meningie, on Lake Albert, M. C. R. Sharrad, No. 486, 12/1959,
(AD 96150850); Keith, Specht and Rayson, 1954, (MEL).

PHYLLOTA DIFFUSA (Hooker f.) F. Mueller

Hragms 58) (lse7):

Nomenclatural synonym: Pultenaea diffusa Hooker, f., Fl. Tasm., I: 91,
t. 14 (1860); Benth., Fl. Austr., IL: 119 (1864); Curtis, Stud. Fl. Tasm.,
pars 1, 132 (1956).

A small diffuse shrub, much branched and ascending, 10-30 cm. high, and
spreading 30-50 cm. Stems terete, glabrous to pubescent with short appressed
hairs. Leaves linear, 7 mm. (5-10 mm.) long, bullate, acute (3 on the subjective
seale) ; stipules minute. Flowers scattered along the stem, solitary or in pairs,
sometimes crowded towards the ends of the branches. Pedicel 2-2-5 mm. long.
Bracts identical in appearance with foliage leaves; bracteoles lanceolate,
1-75 mm. long, 0-75 mm. broad, scarcely herbaceous, glabrous or a few scattered
short silky hairs, borne on the base of the calyx. Calyx 3-4 mm. long, almost
glabrous or with a few scattered silky hairs; lower lobes acute, shorter than
or equal to the tube ; upper lobes broader connate higher up, obtuse. Corolla:
keel equal in length to the standard, broadly lunate to semi-circular, obtuse,
yellow-red ; wings equal in length to the standard, oblong to obovate, obtuse
and rounded, yellow; standard 6-8 mm. long, orbicular, yellow. Stamens 10,
almost wholly free, deciduous with the petals after flowermg. Ovary pubescent
to villous, style dilated or thickened at base, incurved or subulate above,
pubescent with short appressed silky hairs to below the hook. Legume 1-2
seeded, 1—2 times as long as the calyx. Seed reniform.

Distribution : Tasmania, endemic. Local near the east coast and in the
extreme north west.

Habitat: Sandy heaths.

Chromosome number: Not known.

Typification: Holotype: Loc. non cit. J. D. Hooker, (K).

Specimens examined: Coast Rd. to George’s Bay, A. Simson, No. 1325,
11/1878, (MEL); St. Paul’s River, nr. Broadshead, —, 1/1858, (MEL); Coast
nr. Scamander River, A. Simson, 11/1878, (MEL); South Port, Stuart, —,
(MEL); South Port, —, No. 1515, 1/1856, (MEL); George’s Bay, lL. Rodway,
6/1900, (NSW 36497); Tasmania, A. H. S. Lucas, 1910-1930. (NSW 36499).

i
¥

R. C. JANCEY 375

Acknowledgements

The writer is greatly indebted to Dr. R. C. Carolin of the University of
Sydney, for his help and advice during the course of this investigation, for
detailed descriptions of type specimens lodged in European herbaria, and also
for reading the manuscript. Thanks are also expressed to the Directors of
the following institutions for the loan of herbarium specimens: The Depart-
ment of Agriculture, Government of Western Australia; The State Herbarium
of South Australia; The National Herbarium of Victoria; The National
Herbarium of New South Wales; Queensland Herbarium.

References

BursipGe, N. T., 1960—The Phytogeography of the Australian Region. Austr. J. Bot.,
8: 75-211.

Davin, Sir T. W. E., 1950.—The Geology of the Commonwealth of Australia. (Edward Arnold
and Co., London.).

Gitmovur, J. S. L., and Hestop-Harrison, J., 1954.—The Deme terminology and units of micro-
evolutionary change. Genetics, 27: 147-161.

Jancry, R. C., 1966.—Numerical methods in taxonomy. Proc. Linn. Soc. N.S.W., 90: 335.

———., (in press).—The application of numerical methods of data analysis to the taxonomy
of the genus Phyllota (Sieber ex. DC.) Benth. Austr. J. Bot.

STANDARD, J., 1961.—A new study of the Hawkesbury Sandstone: Preliminary findings. Proc.
Roy. Soc. N.S.W., 95: 145-146.

THomeson, J., 1961.—Contrib. N.S.W. Nat. Herb., Fl. N.S.W., No. 101: 45.

Wittis, J. H.. 1957.— Vascular flora of Victoria and South Australia. Vic. Nat., 73: 191.

EXPLANATION OF PLATES XXIX-XXX
Plate xxix
A, Illustrating from left to right, the subjective values from 4 to 1 for calyx and bracteole
indumentum. B, Leaves showing values of 1 to 4 on the subjective scale for size of leaf
bullae. C, Leaves showing values of 1 to 4 on the subjective scale for number of leaf bullae.
(In both B and C cases it will be seen that many of the epidermal hairs are still present,

while these are lost with increasing age-of the leaf, the bullae remain.) D, Four leaf tips of
P. phylicoides showing examples of the values from | to 4 on the subjective scale for leaf tip shape.

Plate xxx

A, Growth habit of group 1. B, Growth habit of group 4. C, Growth habit of groups 7 and 8.
(Scales equal 12 inches.)

ABSTRACT OF PROCEEDINGS

ORDINARY MONTHLY MEETING
31st MARCH, 1965

Dr. D. T. Anderson, President, in the chair.

The minutes of the last Monthly Meeting (25th November, 1964) were
taken as read and signed.

The following were elected Ordinary Members of the Society: Mr. K. R.
Ayers, Fairfield, N.S.W.; Mr. R. Basden, M.Ed., B.Se. (Lond.), F.R.A.C.L.,
A.S.T.C., Hamilton, N.S.W.; Miss Phillipa A. Croucher, B.Se., Canberra,
A.C.T.; Mr. D. J. McGillivray, B.Se.For. (Syd.), Dip.For. (Canb.), Castle Hill,
N.S.W.; Mr. D. R. Murray, B.Se., Brookvale, N.S.W.; Dr. F. H. Talbot,
M.Se., Ph.D., Australian Museum, Sydney; and Dr. R. Tucker, B.V.Se.,
Dr.Vet.M., University of Queensland, St. Lucia, Queensland.

The Chairman announced that library accessions amounting to 42 volumes,
472 parts or numbers, 13 bulletins, 18 reports and 14 pamphlets, total 559,
had been received since the last meeting.

The Chairman announced that the Conservation Photographic Exhibition
which the Society has arranged is now on view at the Australian Museum.
The Exhibition, which consists of photographs and diagrams, illustrates the
need for nature conservation, particularly in New South Wales. Members are
invited to view and to draw the attention of others to this Exhibition. The
Society particularly wishes to acknowledge the help given by various members
of the staff of the Australian Museum, Dr. Eric Bird (A.N.U.) and Mr. R.
Schodde (C.S.I.R.O.) for help in the production.

PAPER READ
(By title only, an opportunity for discussion to be given at the April
Ordinary Monthly Meeting)

1. Chromosome numbers in some Australian leafhoppers (Homoptera :
Auchenorrhynecha). By M. J. Whitten. (With an Appendix by J. W. Evans,
Australian Museum, describing a new genus and species of Hurymelidae.)

ORDINARY MONTHLY MEETING
28th APRIL, 1965

Dr. D. T. Anderson, President, in the chair.

The minutes of the last Monthly Meeting (31st March, 1965) were read
and confirmed.

LECTURETTE

A lecturette, illustrated by colour transparencies, entitled ‘“‘ Some aspects
of coastal morphology in New South Wales ”’, was delivered by Mr. J. R. Hails,
Geography Department, University of Sydney.

Dr. T. E. Woodward, M.Se. (N.Z.), Ph.D. (Lond.), D.I.C., University of
Queensland, St. Lucia, Queensland, was elected an Ordinary Member of the
Society.

The Chairman announced that the Council had elected the following office-
bearers for the 1965-66 session: Vice-Presidents: Miss Elizabeth C. Pope,

ABSTRACT OF PROCEEDINGS 377

Mr. G. P. Whitley, Professor J. M. Vincent and Dr. T. G. Vallance ; Honorary
Treasurer: Dr. A. B. Walkom ; Honorary Secretaries: Drs. A. B. Walkom and
W. R. Browne.

The Chairman announced that library accessions amounting to 10 volumes,
111 parts or numbers, 2 bulletins, 6 reports and 1 pamphlet, total 130, had
been received since the last meeting.

The Chairman reminded members and their friends of the Conservation
Photographic Exhibition now open for inspection at the Australian Museum,
College Street, Sydney.

The Chairman reminded members that there will be no Ordinary Monthly
Meeting in May.

PAPERS READ

1. A note on blood preferences of Anopheles farauti. By Margaret Spencer.

2. Malaria in the D’Entrecasteaux Islands, Papua, with particular
reference to Anopheles farautt Laveran. By Margaret Spencer.

3. The distribution of the Notonectidae (Hemiptera) in south-eastern
Australia. By A. W. Sweeney.

4, The reproduction and early life histories of the gastropods, Notoacmaea
petterdi (Ten.-Woods), Chiazacmaea flammea (Quoy and Gaimard) and Patelloida
alticostata (Angas) (Fam. Acmaeidae). By D. T. Anderson.

5. The histology and anatomy of the reproductive system of the littoral
gastropod, Bembicium nanum (Lamarck) (Fam. Littorinidae). By lynne
Bedford.

Dr. I. V. Newman, a Trustee of the Muogamarra Sanctuary, appealed to
the younger members of the Society of both sexes to join the Sanctuary’s
Volunteer Fire Brigade. The extent of the devastation caused by the recent
bushfires would have been greatly lessened had more fire fighters been available.
Applications should be made to the Hon. Secretary, Miss Monaghan (Tel.
Evenings only, 44 2624), Box 2770, G.P.O., Sydney.

ORDINARY MONTHLY MEETING
30th June, 1965

Dr. D. T. Anderson, President, in the chair.
The minutes of the last Monthly Meeting (28th April, 1965) were read and
confirmed.

The following were elected Ordinary Members of the Society: N. E.
Milward, B.Sc.(Hons.), M.Sc., Queensland ; Mrs. I. M. Straughan, B.Sc.(Hons.)
Queensland ; and B. D. Webby, Ph.D., M.Sc., Sydney University.

The Chairman announced the death of Miss Florence Sulman, M.B.E., on
15th June, 1965, aged 89 years. Miss Sulman had been a member of the Society
since 1911.

The Chairman announced that library accessions amounting to 26 volumes,
269 parts or numbers, 6 bulletins, 2 reports and 1 pamphlet, total 304, had been
received since the last meeting.

PAPERS READ

1. Australian larval Carabidae of the subfamilies Harpalinae, Licininae,
Odacanthinae and Pentagonicinae (Coleoptera). By B. P. Moore.

2. Some Laelapid mites of syndactylous marsupials. By R. Domrow.

318 ABSTRACT OF PROCEEDINGS

3. The occurrence and composition of manna in Hucalyptus and Angophora.
By Ralph Basden.

4. Comparative studies on the external acoustic meatus. 1. The morphology
of the external ear of the echidna (Tachyglossus aculeatus). By Richard Tucker.

5. Studies on the inheritance of rust resistance in oats. III. Genetic
diversity in the varieties Landhafer, Santa Fe, Mutica Ukraine, Trispernia and
Victoria for crown rust resistance. By U. M. Upadhyaya and E. P. Baker.

LECTURETTE

A lecturette, illustrated by slides and exhibits, entitled ‘ Trace fossils :
their classification and palaeoecological significance’, was delivered by Dr.
B. D. Webby, Lecturer in Palaeontology, University of Sydney.

ORDINARY MONTHLY MEETING
28th JULY, 1965

Dr. D. T. Anderson, President, in the chair.

The minutes of the last Monthly Meeting (30th June, 1965) were read and
confirmed.

Professor W. Stephenson, University of Queensland, Brisbane, was elected
an Ordinary Member of the Society.

The Chairman announced that library accessions amounting to 18 volumes,
170 parts or numbers, 8 bulletins, 5 reports and 10 pamphlets, total 211, had
been received since the last meeting.

The Chairman reminded members that there will be no August Ordinary
Monthly Meeting.

The Chairman announced that a lecture on Radio-carbon Dating in the
Quaternary will be given by Professor H. Godwin on Wednesday, 8th September,
1965, at 3 p.m., in the Chemistry No. 1 Lecture Theatre, University of Sydney.

The Chairman announced that a Conversazione will be held on Saturday,
18th September, 1965, from 2 to 5.30 p.m., in the Zoology Department of the
University of New South Wales. Members engaged in scientific research who
have material or apparatus considered suitable for exhibition are invited to
communicate with Mr. R. Strahan, Department of Zoology, School of Biological
Sciences, University of New South Wales.

The Chairman also announced the proposed formation of a Mycological
Society of New South Wales. Interested persons or representatives of institutions
please communicate with Professor N. H. White, Department of Agriculture,
University of Sydney.

PAPERS READ

1. The Devonian tetracoral Haplothecia and new Australian phacello-
phylliids. By A. HE. H. Pedder.

2. Some mite parasites of Australian birds. By R. Domrow.

3. Development of the eggs and early larvae of the Australian smelt,
Retropinna semont (Weber). By N. E. Milward. (Communicated by Mr. G. P.
Whitley.)

4. The vertebrate fauna of “ Gilruth Plains”’, South-west Queensland.
By M. C. Brooker and G. Caughley. (Communicated by Dr. Mervyn Griffiths.)

ABSTRACT OF PROCEEDINGS 379

5. The first zoea of the soldier crab, Mictyris longicarpus (Grapsoidea :
Mictyridae). By Ann M. Cameron. (Communicated by Dr. E. J. Reye.)

6. Plant parasitic nematodes in fruit tree nurseries of New South Wales.
By EH. J. Anderson. (Communicated by Dr. C. D. Blake.)

7. Diurnal variation in the release of pollen by Plantago lanceolata L. By
J. M. Matthews. (Communicated by Dr. Donald Walker.)

LECTURETTE

A lecturette, entitled ‘“‘ Ecology of Ruppia and Zostera’, was delivered by
Mr. R. Higginson, B.Se.Agr., School of Biological Sciences, University of Sydney.

SPECIAL GENERAL MEETING
29th SEPTEMBER, 1965

The recommendation from the Council, of which the required notice had
been given to members, that Rules V and VI be altered to read as follows, was
read :

Rule V.—No person declared elected under Rule IV shall be admitted
to any of the privileges of membership until he has signed a written
acceptance of membership and paid his first Annual Subscription.

Rule VI.—The Annual Subscription shall be three pounds ten shillings
(seven dollars) and shall become due in advance on the first day of March
in each year, or, in the case of New Members, immediately on election ;
provided that Ordinary Members elected after the first day of October
in any year shall have the option of paying the Annual Subscription
either for that year or in advance for the following year. Any Ordinary
Member who has paid the Annual Subscription for forty years shall be
exempt from payment of further subscriptions.

The President explained that these changes in the Rules, if adopted, would
provide for (a) abolition of the entrance fee of one guinea, (b) increase of the
annual subscription from two guineas to three pounds ten shillings (seven dollars),
and (c) abolition of the provision for subscription for Life Membership.

A letter from Mr. A. F. Batley, a member absent from Australia, was read,
in which he stated, inter alia, that the Society at present is living well within its
annual income, and suggested that members would be interested to know if there
is a Specific purpose in the proposed increase of subscription, other than the
increased cost of publication.

In explanation the Hon. Treasurer reminded members that from 1886 to
the end of 1950 the Society had a full-time paid Secretary. At the beginning
of 1951 the Council was faced with the position that the income of the Society
was no longer sufficient to permit employment of a Secretary and publication
of the Proceedings. To alleviate this position two members of the Council
undertook to carry on in an honorary capacity the work of the Secretary (set
out in Rule XLIX) in the hope that gradual accumulation of surplus income
would, in due course, make it possible for the Society to consider the appointment
of a full-time Secretary. This stage appears to have been reached, and recent
events have emphasized the need for the Council to give the matter serious
consideration.

The Hon. Treasurer then proposed that the recommendation of the Council,
as set out in the notice to members, be approved. This was seconded, and was
carried unanimously.

380 ABSTRACT OF PROCEEDINGS

ORDINARY MONTHLY MEETING
29th SEPTEMBER, 1965

Dr. D. T. Anderson, President, in the chair.

The minutes of the last Monthly Meeting (28th July, 1965) were read and
confirmed.

LECTURETTE

An illustrated lecturette on Opossums was delivered by Dr. M. P. Marsh,
School of Biological Sciences, University of Sydney.

The Chairman announced that Dr. N. G. Stephenson, Mr. L. A. S. Johnson
and Dr. Erik Shipp have been elected members of Council in place of Professors
B. J. Ralph, W. I). Waterhouse and I. A. Watson, who had resigned.

The Chairman announced that the Council is prepared to receive applications
for Linnean Macleay Fellowships tenable for one year from ist January, 1966,
from qualified candidates. Each applicant must be a member of this Society
and be a graduate in Science or Agricultural Science of the University of Sydney.
The range of actual (tax-free) salary is, according to qualifications, up to a
maximum of £1,600 per annum. Applications should be lodged with the
Honorary Secretary, who will give further details and information, not later
than Wednesday, 3rd November, 1965.

The Chairman announced that library accessions amounting to 39 volumes,
295 parts or numbers, 8 bulletins, 7 reports and 8 pamphlets, total 355, had been
received since the last meeting.

PAPERS READ

1. Observations on the fine structure of the meristem of root nodules from
some annual iegumes. By P. J. Dart and KF. V. Mercer.

2. Cerioid Stringophyllidae (Tetracoralla) from Devonian strata in the
Mudgee district, New South Wales. By A. J. T. Wright.
8

3. Further observations on, the life histories of littoral gastropods in New
South Wales. By D. T. Anderson.

SPECIAL GENERAL MEETING
27th OCTOBER, 1965

Dr. D. T. Anderson, President, in the chair.

The minutes of the Special General Meeting of 29th September, 1965, were
read and confirmed.

The recommendation of the Council that Rules V and VI be altered, as
approved at the Special General Meeting of 29th September, 1965, was confirmed,
and carried unanimously.

ORDINARY MONTHLY MEETING
27th OCTOBER, 1965

Dr. D. T. Anderson, President, in the chair.

The minutes of the last Monthly Meeting (29th September, 1965) were read
and confirmed.

ABSTRACT OF PROCEEDINGS 381

LECTURETTE

A lecturette entitled ‘“‘Some Problems of Evolutionary Theory” was
delivered by Dr. Paul Ehrlich, Stanford University, Stanford, U.S.A.

The lecturer raised many controversial issues and much vigorous discussion
ensued.

Miss Alison Kay Dandie, Women’s College, Newtown, N.S.W., and Dr.
David Michael Griffin, M.A., Ph.D. (Cantab.), Sydney University, 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, 1966,
from qualified candidates. Hach applicant must be a member of this Society
and be a graduate in Science or Agricultural Science of the University of Sydney.
The range of actual (tax-free) salary is, according to qualifications, up to a
maximum of £1,600 per annum. Applications should be lodged with the
Hon. Secretary, who will give further details and information, not later than
Wednesday, 3rd November, 1965.

The Chairman announced that library accessions amounting to 25 volumes,
227 parts or numbers, 31 bulletins, 9 reports and 8 pamphlets, total 300, had been
received since the last meeting.

The Chairman drew the attention of members to the portrait of Linnaeus, a
recent gift from the Linnean Society of London. This painting from the Roslin
portrait, presented to the Linnean Society of London in 1814 by Joseph Sabine,
one of the original Fellows of the Society, hung for many years in the meeting
room of the Society behind the President’s chair.

PAPERS READ
1. An embryological study of five species of Bassia All. (Chenopodiaceae).
By Gwenneth J. Hindmarsh.

2. Studies of nitrogen-fixing bacteria. IX. Study of inoculation of wheat
with Azotobacter in laboratory and field experiments. By Y.T.Tchan and D. L.
Jackson.

3. Studies on the genetic nature of resistance to Puccinia graminis var.
tritict In six varieties of common wheat. By N. H. Luig and I. A. Watson.

ORDINARY MONTHLY MEETING
24th NOVEMBER, 1965

Mr. G. P. Whitley, Vice-President, in the chair.

The following were elected Ordinary Members of the Society : Derek John
Anderson, Ph.D., Drummoyne, N.S.W., and David Robert Selkirk, Mosman,
N.S.W.

The Chairman announced that library accessions amounting to 15 volumes,
133 parts or numbers, and 9 pamphlets, total 157, had been received since the
last meeting.

PAPERS READ
1. Numerical methods in taxonomy. By R. C. Jancey.
2. An investigation of the genus Phyllota Benth. By R. C. Jancey.

3. The distribution of submerged aquatic Angiosperms in the Tuggerah
Lakes system. By F. R. Higginson.

382 ABSTRACT OF PROCEEDINGS

NOTES AND EXHIBITS

Dr. T. G. Vallance drew attention to two significant tercentenaries in Natural
Science which have occurred during 1965. In 1665, Dr. Robert Hooke
(1635-1705), encouraged by the newly-established Royal Society, issued
Micrographia. Sir Geoffrey Keynes has recently commented “. . . that Hooke
was almost, if not quite, the most prolific inventive genius that has ever lived and
that at least one of his books, the Micrographia, is among the most important
books ever published in the history of science”’’. Hooke’s scientific career
was outlined briefly and a copy of the original issue of Micrographia was exhibited
together with a modern facsimile reproduction. Dr. John Woodward (1665-1728)
was a much less distinguished scientist than Robert Hooke but he has some claim
to our attention for his work in pioneering the concept of an organic origin for
fossils and through his bequest to the University of Cambridge which led to the
establishment of a Chair of Geology—the first of its kind in the world. Copies
of the first (1695) and second (1702) editions of Woodward’s An Essay toward(s)
a Natural History of the Earth were exhibited.

Mr. A. Mahmood, a visitor (introduced by Dr. I. V. Newman), exhibited
two electronmicrographs of phloem tissue of Pinus radiata as seen in transverse
section, revealing the parental primary wall. Looking centrifugally in the first
photograph, each of the three cells showed the secondary wall, the primary wall,
and broken pieces of ‘‘ parental wall’ of uneven thickness on the outer side of
the radial face of the primary walls of the cells. The intercellular substance
between the tangential faces of the primary walls is seen as a dark line, but its
identity as such is obscure where the cells are rounded at their corners due to
tissue differentiation. In the second photograph two cells are seen, more
enlarged. The stretching of the ‘‘ parental wall’ is more pronounced opposite
the cell corners where the intercellular space has appeared. The black material
lining the intercellular space may be intercellular substance which was displaced
from its original position due to the splitting of the two primary walls as described
by some workers. It looked as if the secondary wall is composed of more than
one layer. The cell cavity showed plasmalemma, mitochondria, golgi bodies
and vesicles. The consequences of formation of new cell wall with each division
were discussed.

Mr. M. V. Ramji, a visitor (introduced by Dr. I. V. Newman), exhibited
slides illustrating a study of embryogeny of Stellaria media showing the following
features: (1) the four terminal cell tiers of a proembryo of six cells produce the
globular embryo while the basal cells produce the stalk with the suspensor below :
(2) the cotyledons, the lateral protoderm, pith and procambium of embryonal
axis arise from the second, third and fourth original tiers of cells and not from the
terminal tier; (3) the terminal tier gives rise essentially to the shoot apex ;
(4) the terminal hemisphere of the embryo cannot be considered to be the first
Shoot apex of the plant as is usually done in many books, since the shoot apex
arises exclusively from the terminal tier ; (5) the cotyledons cannot be considered
as the first leaves of the plant.

Miss J. L. Jacobs exhibited, on behalf of Mr. R. Selkirk, a specimen from
Kiandra, N.S.W. Many of these specimens are found associated with epiphyllous
and leaf-parasitic fungi on the surface of small myrtaceous leaves from lignite at
Kiandra. They are visible as small dots under a dissecting microscope, becoming
visible only after maceration of the leaves to leave clear cuticular preparations.
The specimen is apparently fungal in nature but, out of thousands so far examined,
no form of mycelium, spores or usual fungal structures is exhibited. Three
mycologists have so far examined it, but it remains as much a mystery as ever,
Other fossil fungi occurring with it are tentatively assigned to Asterinaceae.
Meliolaceae and Trichopeltaceae, a number of Cookson’s types (PRoc. LINN.
Soc. N.S.W., 72: 207-214, 1947) being present.

ABSTRACT OF PROCEEDINGS 383

Mr. R. K. Bamber exhibited a specimen and photographs of fossil wood
found in Mount Royal State Forest, New South Wales, on the roadside following
road construction. The specimen is highly silicified, being almost completely
soluble in hydrofluoric acid. The cell formation is fairly well preserved and
under incident light a great amount of detail is visible on the broken faces.
From the pattern of the cells it is estimated that the tree from which the specimen
originated was at least 74 inches in diameter. The fossil wood has only tracheids
and ray parenchyma. Growth rings and resin canals are absent. The tracheids
have an average diameter of 83y radially by 72 tangentially and an average
length of 8mm. The radial walls are profusely pitted with from 2 to 4 rows of
alternate bordered pits. The ray parenchyma is multiseriate, ranging from
2 to 5 cells wide and from 6 to 43 cells high. All the ray parenchyma cells are
procumbent. Pitting in the ray parenchyma could not be observed. This
structure appears generally to be similar to that described in the literature as
cordaitean wood. It is almost identical with a specimen from Wallarobba
described as Pitys? sussmileht by A. B. Walkom in 1928 (PRoc. Linn. Soc.
N.S.W., 53: 255-269). The specimen also shows some similarity to a fossil
wood from Upper Devonian rocks at Mansfield, Victoria, described by I. C.
Cookson (Proc. Roy. Soc. Vict., 1937, vol. 50: 182-189).

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LIST OF MEMBERS.

LIST OF MEMBERS.
(15th December, 1965.)

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

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

*Allman, Stuart Leo, B.Sc.Agr., M.Sc., 99 Cumberland Avenue, Collaroy, N.S.W.

Anderson, Derek John, Ph.D., School of Biological Sciences, Botany Building, Sydney
University.

Anderson, Donald Thomas, B.Sc., Ph.D., School of Biological Sciences, Department of
Zoology, Sydney University.

Anderson, Mrs. Jennifer Mercianna Elizabeth, B.Se.Agr., Macleay Museum, Sydney
University.

Anderson, Robert Henry, B.Sc.Agr., Kareela Road, Chatswood, N.S.W.

Ardley, John Henry, B.Sc. (N.Z.), School of Public Health and Tropical Medicine,
Sydney University.

*Armstrong, Jack Walter Trench, “Cullingera’, Nyngan, N.S.W.

Ashton, David Hungerford, B.Se., Ph.D., 92 Warrigal Road, Surrey Hills, E.10, Victoria.

Aurousseau, Marce!, B.Se., 229 Woodland Street, Balgowlah, N.S.W.

Ayers, Kenneth Robert, 9 Livingstone Street, Burwood, N.S.W.

Bailey, Peter Thomas, B.Sc., C.S.I.R.O., Division of Wildlife Research, P.O. Box 109,
City, Canberra, A.C.T.

Bain, Miss Joan Maud, M.Sc., 18 Onyx Road, Artarmon, N.S.W.

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

Ballantyne, Miss Barbara Jean, B.Sc.Agr., N.S.W. Department of Agriculture Private
Mail Bag No. 10, Rydalmere, N.S.W.
Bamber, Richard Kenneth, F.S.T.C., 113 Lucinda Avenue South, Wahroonga, N.S.W.
“Barber, Professor Horace Newton, M.S., Ph.D., F.A.A., School of Biological Sciences,
Department of Botany, University of N.S.W., P.O. Box 1, Kensington, N.S.W.
Barber, Ian Alexander, B.Sc.Agr., School of Biological Sciences, Department of Zoology,
Sydney University.

Barlow, Bryan Alwyn, B.Sc., Ph.D., c/- Waite Agricultural Research Institute, Private
Bag, Glen Osmond, South Australia.

Basden, Ralph, M.Ed., B.Sc. (Lond.), F.R.A.C.I., A.S.T.C., 183 Parkway Avenue,
Hamilton, N.S.W.

Batley, Alan Francis, A.C.A., 123 Burns Road, Wahroonga, N.S.W.

Baur, George Norton, B.Sc., B.Se.For., Dip.For., 3 Mary Street, Beecroft, N.S.W.

*Beadle, Professor Noel Charles William, D.Se., Uni-ersity of New England, Armidale, 5N,
N.S.W.

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

Beattie, Joan Marion, D.Se. (née Crockford), 2 Grace Avenue, Beecroft, N.S.W.

Bedford, Geoffrey Owen, B.Sc., c/- C.S.1.R.O., Division of Entomology, P.O. Box 109, City,
Canberra, A.C.T.

Bedford, Miss Lynette, B.Sc., School of Biological Sciences, Department of Zoology,
Sydney University.

Bennett, Miss Isobel Ida, Hon. M.Sc., School of Biological Sciences, Department of
Zoology, Sydney University.

Bertus, Anthony Lawrence, B.Sc., Biology Branch, N.S.W. Department of Agriculture,
Private Mail Bag, No. 10, Rydalmere, N.S.W.

Besly, Miss Mary Ann Catherine, B.A., School of Bioiogical Sciences, Department of
Zoology, Sydney University.

Bishop, James Arthur, School of Biological Sciences, Department of Zoology, Sydney
University.

Blackmore, John Allan Philip, LL.B. (Syd. Univ.), 25 Holden Street, Ashfield, N.S.W.

Blake, Clifford Douglas, B.Sc.Agr., Ph.D., Faculty of Agriculture, Sydney University.

Blake, Stanley Thatcher, D.Sc. (Q’ld.), Botanic Gardens, Brisbane, Queensiand.

Bourke, Terrence Victor, B.Sc.Agr., c/- Department of Agriculture, Stock and Fisheries,
Popondetta, Papua.

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

Brewer, Ilma Mary, D.Se., 7 Thornton Street, Darling Point, Sydney.

Briggs, Miss Barbara Gillian, Ph.D., 13 Findlay Avenue, Roseville. N.S.W.

Browne, Ida Alison, D.Se. (née Brown), 3638 Edgecliff Road, Edgecliff, N.S.W.

Browne, William Rowan, D.Sc... F.A.A., 363 Edgecliff 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.

LIST OF MEMBERS. 385

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

Burgess, The Rev. Colin E. B. H., Parks and Gardens Section, Department of the
Interior, Canberra, A.C.T.

Burgess, Ian Peter, B.Sc.For., Dip.For., The Forestry Office, Coff’s Harbour, N.S.W.

Cady, Leo Isaac, P.O. Box 88, Kiama, N.S.W.

Campbell, Keith George, D.F.C., B.Sc.For., Dip.For., M.Sc., 17 Third Avenue, Epping,
N.S.W.

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

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

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

Carolin, Roger Charles, B.Sc., A.R.C.S., Ph.D., School of Biological Sciences, Department
of Botany, Sydney University.

Casimir, Max, B.Sc.Agr., Entomological Branch, N.S.W. Department of Agriculture,
Private Mail Bag, No. 10, Rydalmere, N.S.W.

*Chadwick, Clarence Earl, B.Se., Entomological Branch, N.S.W. Department of
Agriculture, Private Mail Bag No. 10, Rydalmere, N.S.W.

Chambers, Thomas Carrick, M.Sc. (N.Z.), Ph.D., Botany School, University of Melbourne,
Parkville, N.2, Victoria.

Chippendale, George McCartney, B.Sc., Lindsay Avenue, Alice Springs, Northern
Territory, Australia.

Christian, Stanley Hinton, Malaria Research Unit and School, Kundiawa, Eastern High-
lands, Territory of Papua and New Guinea.

*Churchward, John Gordon, B.Sc.Agr., Ph.D., “Shady Trees’, Clarktown Avenue, Mount
Bliza, Victoria.

Clark, Laurance Ross, M.Sc., c.o. C.8.1.R.O., Division of Hntomology, P.O. Box 109, City,
Canberra, A.C.T.

Clarke, Miss Lesley Dorothy, 4 Gordon Crescent, Hastwood, N.S.W.

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

Cleland, Professor Sir 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, M.Sc., Australian Museum, College Street, Sydney.

Colless, Donald Henry, Ph.D. (Univ. of Malaya), c.o. Division of Entomology, C.S.I.R.O.,
P.O. Box 109, City, Canberra, A.C.T.

Common, Ian Francis Bell, M.A., M.Se.Agr., C.S.I.R.O., Division of Hntomology,
P.O. Box 109, City, Canberra, A.C.T.

Copland, Stephen John, M.Sc., 15 Chilton Parade, Warrawee, N.S.W.

Costin, Alex Baillie, B.Se.Agr., C.S.I.R.O., Division of Plant Industry, P.O. Box 109,
City, Canberra, A.C.T.

Craddock, Miss Elysse Margaret, 36 Lyons Road, Drummoyne, N.S.W.

Crawford, Lindsay Dinham, B.Sc., c/- Victorian Plant Research Institute, Department
of Agriculture, Burnley Gardens, Melbourne, Victoria.

Crook, Keith Alan Waterhouse, M.Se., Ph.D. (New England), Department of Geology,
Australian National University, G.P.O. Box 197, Canberra, A.C.T.

Croucher, Miss Phillipa Audrey, B.Sc., Department of Zoology, Australian National
University, P.O. Box 4, Canberra, A.C.T.

Dandie, Miss Alison Kay, Women’s College, Carillon Avenue, Newtown, N.S.W.

Dart, Peter John, B.Sc.Agr., c/- Mr. H. F. Dart, 53 Smith Street, Wollongong, N.S.W.

Davies, Stephen John James Frank, B.A. (Cantab.), C.S.I.R.O., Private Bag, Nedlands,
Western Australia.

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

Deuquet, Camille, B.Com., 56 Avenue Eug. Ysaye, Brussels 7, Belgium.

Dobrotworsky, Nikolai V., M.Sc., Ph.D., Department of Zoology, University of Melbourne,
Parkville, N.2, Victoria.

Domrow, Robert, B.A., B.Se., Queensland Institute of Medical Research, Herston Road,
Herston, N.9, Brisbane, Queensland.

Dorman, Herbert Clifford, J.P., A.S.T.C. (Dip.Chem.), Dip.Soc. Stud. (Sydney), Rodgers
Street, Teralba 2N, N.S.W.

Douglas, Geoffrey William, B.Agr.Sci., c/- The Keith Turnbull Research Station, Private
Bag, Frankston, Victoria.

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LIST OF MEMBERS.

Durie, Peter Harold, M.Sc., C.S.I.R.O., Veterinary Parasitology Laboratory, Yeerongpilly,
Brisbane, Queensland.
Dyce, Alan Lindsay, B.Sc.Agr., 48 Queen’s Road, Asquith, N.S.W.

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

Endean, Robert, M.Sc., Ph.D., Department of Zoology, University of Queensland, St.
Lucia, Brisbane, Queensland.

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

Evans, Miss Gretchen Pamela, M.Se., Box 92, P.O., Canberra, A.C.T.

Facer, Richard Andrew, ‘Moppity’, Parsonage Road, Castle Hill, N.S.W.

*Fairey, Kenneth David, Box 1176, G.P.O., Sydney.

Filewood, Lionel Winston Charles, c/- Department of Agriculture, Stock and Fisheries,
Konedobu, Papua.

Florence, Ross Garth, M.Sc.For., Ph.D., Forest Research Station, Beerwah, Queensland.

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

Gardner, Mervyn John, B.Sc.For., Dip.For., Forestry Office, Wauchope, N.S.W.

*Garretty, Michael Duhan, D.Sc., Box 763, Melbourne, Victoria.

Green, John William, B.Se. (Adel.), Department of Botany, Australian National
University, Box 197, P.O., Canberra City, A.C.T.

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

Griffin, David Michael, M.A., Ph.D. (Cantab.), School of Agriculture, Sydney University.

*Griffiths, Mrs. Mabel, B.Sc. (née Crust), 50 Amourin Street, Brookvale, N.S.W.

Grifiths, Mervyn Edward, D.Se., Wildlife Survey Section, C.S.I1.R.O., 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.

Hadlington, Phillip Walter, B.Sc.Agr., 15 Annie Street, Hurstville, N.S.W.

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

*Hansford, Clifford Gerald, M.A., Se.D. (Cantab.), D.Se. (Adel.), F.L.S., P.O. Box 456,
Uvongo Beach, Natal, South Africa.

Hardy, George Huddleston Hurlstone, 68 Cliff Avenue, Northbridge, N.S.W.

Hartigan, Desmond John, B.Sc.Agr., 75 Northwood Road, Northwood, Lane Cove, N.S.W.

Hennelly, John Patten Forde, B.Sc., Highs Road, West Pennant Hills, N.S.W.

Hewitt, Bernard Robert, B.A. (Qld.), B.Sc. (Syd.), M.Sc. (N.S.W.), A.R.A.C.1., Depart-
ment of Chemistry, University of Malaya, Kuala Lumpur, Malaya.

Hewson, Miss Helen Joan, B.Se. (Hons.), School of Biological Sciences, Department of
Botany, Sydney University.

Higginson, Francis Ross, B.Sc.Agr. (Hon.), 3 Benson Street, West Ryde, N.S.W.

Hill, Miss Dorothy, M.Sc., Ph.D., Department of Geology, University of Queensland,
Brisbane, Queensland.

Hindmarsh, Miss Gwenneth Jean, B.Sc., Botany Department, University College of
Townsville, Pimlico, Townsville, Queensland.

*Hindmarsh, Miss Mary Maclean, B.Sc., Ph.D., 4 Recreation Avenue, Roseville, N.S.W.

*Holder, Miss Lynette Anne, B.Sc., 48 Rutiedge Street, Eastwood, N.S.W.

*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., M.A. (née Maxwell), 2440 Longest Avenue,
Louisville, Kentucky, U.S.A.

Humphrey, George Frederick, M.Sc., Ph.D., C.S.I.R.O. Marine Biological Laboratory,
Box 21, Cronulla, N.S.W.

Ingram, Cyril Keith, B.A., B.Ec., 7 Ramsay Road, Pennant Hills, N.S.W.

Jackes, Mrs. Betsy Rivers, B.Sc., Ph.D. (Univ. Chicago) (née Paterson), 120 Wellington
Street, Aitkenvale, Hermit Park, North Queensland.

Jacobs, Miss Janice Lorraine, B.Sc., School of Biological Sciences, Department of Botany,
Sydney University.

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

Jacobson, Miss Constance Mary, M.Sc., Ph.D., School of Biological Sciences, Department
of Zoology, Sydney University.

James, Sidney Herbert, M.Sce., 54 Holmfirth Street, Mt. Lawley, Western Australia.

Jancey, Robert Christopher, M.Sc., Ph.D., 54 Phoebe Street, Balmain, N.S.W.

Jefferies, Mrs. Lesly Joan, 14 Denman Street, Hurstville, N.S.W.

Jenkins, Thomas Benjamin Huw, Ph.D., Department of Geology and Geophysics, Sydney
University.

LIST OF MEMBERS. 387

Jessup, Rupert William, M.Sc., 6 Penno Parade North, Belair, South Australia.

Jobson, Arthur Edgar, 3 Wellington Road, East Lindfield, N.S.W.

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

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

Jolly, Violet Hilary, M.Se., Ph.D., Metropolitan Water, Sewerage and Drainage Board,
314 Pitt Street, Sydney.

Jones, Edwin Llewelyn, B.A., P.O. Box 196, Leeton 6S, N.S.W.

Jones, Leslie Patrick, Department of Animal Husbandry, Sydney University.

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

Judd, Howard Kenniwell, Minnamurra Falls Forest Reserve, Box 14, P.O., Jamberoo,
N.S. W.

Keast, James Allen, M.Sc., M.A., Ph.D. (Harvard), Professor of Vertebrate Zoology,
Queen’s University, Kingston, Ontario, Canada.

Kerr, Harland Benson, B.Sc.Agr., Ph.D., Summer Institute of Linguistics, P.O. Ukarumpa,
E.H.D., Territory of New Guinea.

Kesteven, Geoffrey Leighton, D.Se., co. Division of Fisheries and Oceanography
C.S.1.R.0., P.O. Box 21, Cronulla, N.S.W.

Kindred, Miss Berenice May, B.Sc., The Institute for Cancer Research, 7701 Burholme
Avenue, Fox Chase, Philadelphia, Pa., 19111, U.S.A.

King, Miss Dianne, B.Se., 9 Royce Avenue, Croydon, N.S.W.

Langdon, Raymond Forbes Newton, M.Agr.Sc., Ph.D., Department of Botany, University
of Queensland, George Street, Brisbane, Queensland.

Lanyon, Miss Joyce Winifred, B.Sc., Dip.Ed., 35 Gordon Street, Eastwood, N.S.W.

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.

Levy, Miss Margery Olwyn, B.Sc., Dip.Ed., Katoomba High School, Martin Street,
Katoomba, N.S.W.

Littlejohn, Murray John, B.Sc., Ph.D. (W.A.), Department of Zoology, University of
Melbourne, Parkville, N.2, Victoria.

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

Luig, Norbert Harold, Ph.D., c.o. Faculty of Agriculture, Sydney University.

Lyne, Arthur Gordon, B.Sc., Ph.D., C.S.I.R.O., Ian Clunies Ross Animal Research
Laboratory, P.O. Box 144, Parramatta, N.S.W. y

Macdonald, Colin Lewis, 20 Ordnance Avenue, Lithgow, N.S.W.

Macintosh, Professor Neil William George, M.B., B.S., Department of Anatomy, Sydney
University.

Mackay, Miss Margaret Muriel, B.Se. (Hons.) (St. Andrews), M.Sc. (Syd.), M.I.Biol., J.P.,
Department of Botany, University College of Townsville, P.O. Box 499, Townsville,
North Queensland.

Mackerras, Ian Murray, M.B., Ch.M., B.Sc., C.S.1.R.O., Division of Entomology, P.O. Box
109, City, Canberra, A.C.T.

*Mair, Herbert Knowles Charles, B.Sc., Royal 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., School of Biological Sciences, Department
of Botany, Sydney University.

Martin, Mrs. Hilda Ruth Brownell, B.Sc. (née Simons), c/- Mrs. H. F. Simons, 43
Spencer Road, Killara, N.S.W.

Martin, Peter Marcus, M.Sc.Agr., Dip.Ed., School of Biological Sciences, Department of
Botany, Sydney University.

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

McCulloch, Robert Nicholson, M.B.E., D.Sc.Agr., B.Sc., Cattle Tick Research Station,
Wollongbar, N.S.W.

*McCusker, Miss Alison, M.Sc., Botany Department, University College, Box 9184,
Dar es Salaam, Tanzania.

McDonald, Miss Patricia M., B.Sc., Dip.Ed., 33 Holdsworth Street, Neutral Bay, N.S.W.

McGarity, John William, M.Sc.Agr., Ph.D., Agronomy Department, School of Rural

Science, University of New England, Armidale 5N, N.S.W.

McGillivray, Donald John, B.Se.For. (Syd.), Dip.For (Canb.), P.O. Box 107, Castle
Hill, N.S. W.

McKee, Hugh Shaw, B.A., D.Phil. (Oxon.), Service des Eaux et Foréts, B.P. 285, Noumea,
New Caledonia.

McKenna, Nigel Reece, Department of Education, Konedobu, Papua.

LIST OF MEMBERS.

McMichael, Donald Fred, B.Sc., M.A. (Harvard), Ph.D. (Harvard), Australian Museum,
College Street, Sydney.

McMillan, 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.

McWilliam, Miss Paulette Sylvia, 4 Kingslangley Road, Greenwich, N.S.W.

Mercer, Professor Frank Verdun, B.Sc., Ph.D. (Camb.), 24 Trafalgar Avenue, Roseville,
N.S. W.

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.

Millerd, Miss Alison Adéle, M.Sce., Ph.D., C.S.1.R.O., Division of Plant Industry, P.O.
Box 109, City, Canberra, A.C.T.

Millett, Mervyn Richard Oke, B.A., 18 Albion Road, Box Hill, E11, Victoria.

Milward, Norman Edward, B.Sc. (Hons.), M.Sc., Department of Zoology, University
of Queensland, St. Lucia, Brisbane, Queensland.

Moore, Barry Philip, B.Sc., Ph.D., D.Phil., C.S.I.R.O., Division of Entomology, P.O. Box
109, City, Canberra, A.C.T.

Moore, Kenneth Milton, Cutrock Road, Lisarow, N.S.W.

Moore, Raymond Milton, D.Sc.Agr., 94 Arthur Circle, Forrest, Canberra, A.C.T.

Moors, Henry Theodore, B.Sc., Department of Geology and Mineralogy, University of
Melbourne, Parkville, N.2, Victoria.

Morgan, Mrs. Eva, M.Sce., 4 Haberfield Road, Haberfield, N.S.W.

Morrison, Gordon Cyril, 21 Urunga Street, North Balgowlah, N.S.W.

Moss, Francis John, 37 Avenue Road, Mosman, N.S.W.

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

Muirhead, Warren Alexander, B.Sc.Agr., C.S.1.R.O., Irrigation Research Station, Private
Mail Bag, Griffith, N.S.W.

Mungomery, Reginald William, c/- Bureau of Sugar Experiment Stations, 99 Gregory
Terrace, Brisbane, Queensland.

Murray, David Ronald, B.Sc., 14 Consul Road, Brookvale, N.S.W.

Murray, Professor Patrick Desmond Fitzgerald, M.A., D.Se., Department of Zoolegy,
University of New England, Armidale 5N, N.S.W.

Nashar, Mrs. Beryl, B.Sc., Ph.D., Dip. Ed. (née Scott), P.O. Box 28, Mayfield, N.S.W.

Newman, Ivor Vickery, M.Sc., Ph.D., F.R.M.S., F.L.S., School of Biological Sciences,
Department of Botany, Sydney University.

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

*Noble, Norman Scott, D.Sc.Agr., M.Se., D.I.C., Unit 16, 25 Addison Road, Manly, N.S.W.

Noble, Robert Jackson, B.Sc.Agr., Ph.D., 32a 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.Se., B.Se., F.R.E.S., Department of
Zoology, University of New England, Armidale 5N, N.S.W.

O’Gower, Alan Kenneth, M.Se., Ph.D., 20 Gaerloch Avenue, Bondi South, N.S.W.

O’Malley, Miss Doreen Theresa, 100 Hargrave Street, Paddington, N.S.W.

Osborn, Professor Theodore George Bentley, D.Sc, F.L.S., 34 Invergowrie Avenue,
Highgate, South Australia.

Oxenford, Reginald Augustus, B.Sc., 107 Alice Street, Grafton 3C, N.S.W.

Packham, Gordon Howard, B.Sc., Ph.D., 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.

Payne, William Herbert, A.S.T.C., A.M.I.H.Aust., M.A.P.E., 250 Picnic Point Road,
Picnic Point, N.S.W.

Peacock, William James, B.Sc., Ph.D., C.S.I.R.O., Division of Plant Industry, P.O.
Box 109, Canberra City, A.C.T.

Pedder, Alan Edwin Hardy, M.A. (Cantab.), Department of Geology, University of New
England, Armidale 5N, N.S.W.

Perkins, Frederick Athol, B.Sc.Agr., 93 Bellevue Terrace, Clayfield, Brisbane, Queensland.

Philip, Graeme Maxwell, M.Sc. (Melb.), Ph.D. (Cantab.), F.G.S., Department of Geology,
University of New England, Armidale, 5N, N.S.W.

Phillips, Miss Marie Elizabeth, M.Sc., Ph.D., Parks and Gardens Section, Department
of the Interior, Canberra, A.C.T.

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

Pryor, Professor Lindsay Dixon, M.Se., Dip.For., Department of Botany, School of
General Studies, Australian National University, Box 197, P.O., City, Canberra,
A.C.T.

1960

1962
1929

1951

1952
1955
1957
1953

1957

1961

1964
1946

1958

1932

1964

1960

1932

1962
1919
1965
1950
1964
1948

1930
1947
1959
1955
1952
1953
1943
1945
1937

1960

1932
1956

1965

1949

1935
1962

1965

1961
1952

1955
1962

1962

LIST OF MEMBERS. 389

Racek, Albrecht Adalbert, Dr.rer.nat. (Brno, Czechoslovakia), School of Biological
Sciences, Department of Zoology, Sydney University.

Rade, Janis, M.Sc., Flat 28a, 601 St. Kilda Road, Melbourne, S.C.3, Victoria.

Raggatt, Sir Harold George, K.B.E., C.B.E., D.Se., 60 Arthur Circle, Forrest, Canberra,
A.C.T.

Ralph, Professor Bernhard John Frederick, B.Sc., Ph.D. (Liverpool), A.A.C.I., School
of Biological Sciences, University of New South Wales, P.O. Box 1, Kensington,
N.S. W.

Ramsay, Mrs. Helen Patricia, B.Sc. (née Lancaster), 8 Namoi Street, Epping, N.S.W.

Reed, Mrs. Beryl Marlene, B.Sc. (née Cameron), 145 Duffy Street, Ainslie, A.C.T.

Reed, Eric Michael, 145 Duffy Street, Ainslie, A.C.T.

Reye, Eric James, M.B., B.S. (Univ. Qld.), Entomology Department, University of
Queensland, St. Lucia, Brisbane, Queensland.

Reynolds, Miss Judith Louise, c.o. Department of Entomology, University of Illinois,
Urbana, Illinois, U.S.A.

Richards, Miss Aola Mary, M.Sc. (Hons.), Ph.D. (N.Z.), School of Biological Sciences,
University of New South Wales, P.O. Box 1, Kensington, N.S.W.

Richardson, Barry John, 12 Bowden Street, Parramatta North, N.S.W.

Riek, Edgar Frederick, B.Sc., Division of Entomology, C.S.1.R.O., P.O. Box 109, City,
Canberra, A.C.T.

Rigby, John Francis, B.Sc., U.S. Geological Survey Coal Geology Laboratory, Orton Hah,
Ohio State University, 155 S. Oval Drive, Columbus, Ohio, 43210, U.S.A.

*Robertson, Rutherford Ness, F.R.S., B.Sc., Ph.D., F.A.A., Professor of Botany, University
of Adelaide, Adelaide, South Australia.

Rothwell, Albert, D.P.A., 11 Bonnie View Street, Cronulla, N.S.W.

Salkilld, Barry William, Dip.Soc.Stud. (Univ. Syd.), 53 Wareemba Street, Abbotsford,
N.S.W.

*Salter, Keith Eric Wellesley, B.Sc., School of Biological Sciences, Department of Zoology,
Sydney University.

Sands, Miss Valerie Elizabeth, M.Sc., Whakatara, Northland, New Zealand.

*Scammell, George Vance, B.Se., 7 David Street, Clifton Gardens, N.S.W.

Selkirk, David Robert, School of Biological Sciences, Botany Building, Sydney University.

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

Shaw, Miss Dianne Margaret, B.Sc., Faculty of Agriculture, Sydney University.

Shaw, Miss Dorothy Edith, M.Sc.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.

Simons, John Ronald, M.Se., Ph.D., 242 Kissing Point Road, Turramurra, N.S.W.

Slack-Smith, Richard J., c/- Fisheries Department, Perth, Western Australia.

Slade, Milton John, B.Sc., 20 Dobie Street, Grafton, N.S.W.

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

Smith-White, Spencer, D.Sc.Agr., F.A.A., 51 Abingdon Road, Roseville, N.S.W.

Southcott, Ronald Vernon, M.B., B.S., 13 Jasper Street, Hyde Park, South Australia.

Spencer, Mrs. Dora Margaret, M.Se. (née Cumpston), No. 1 George Street, Tenterfieid,
N.S.W.

Staff, Ian Allen, B.Sc., Dip.Ed., Department of Botany, Life Sciences, Southern Illinois
University, Carbondale, Illinois, U.S.A. 4

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

Stephenson, Neville George, M.Sc. (N.Z.), Ph.D. (Lond.), School of Biological Sciences,
Department of Zoology, Sydney University.

Stephenson, Professor William, B.Se. (Hons.), Ph.D. (Durham, Eng.), Diploma of
Education, member Aust. Coll. Educ., Fellow Zoological Society, Department of
Zoology, University of Queensland, St. Lucia, Brisbane, Queensland.

Stevens, Neville Cecil, B.Sce., Ph.D., Department of Geology, University of Queensland,
St. Lucia, Brisbane, Queensland.

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

Strahan, Ronald, M.Sce., Department of Zoology, School of Biological Sciences, University
of New South Wales, P.O. Box 1, Kensington, N.S.W.

Straughan, Mrs. Isdale Margaret, B.Sc. (Hons.), Department of Zoology, University of
Queensland, St. Lucia, Brisbane, Queensland.

Strong, Philip David, 166 Mary Street, Grafton, N.S.W.

Sullivan, George Emmerson, M.Se. (N.Z.), Ph.D., Department of Histology and
Embryology, Sydney University.

Sutherland, James Alan, B.A., B.Sc.Agr., “Skye”, Galloway Street, Armidale, 5N, N.S.W.

Sweeney, Anthony William, Malaria Institute, Public Health Department, Rabaul,
Territory of Papua and New Guinea.

Swinbourne, Robert Frederick George, Box 210, Post Office, Alice Springs, Northern
Territory, Australia.

390
1965

1940
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1956
1960
1949
1944
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1921
1965

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1917
1930
1940
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1961
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1909
1946

1947
1930
1911
1936
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1927
1941

1964
1965
1963
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1926
1962
1960

1954
1954

1960

1952

LIST OF MEMBERS.

Talbot, Frank Hamilton, M.Sc., Ph.D., Australian Museum, P.O. Box A285, Sydney
South, N.S.W.

Taylor, Keith Lind, B.Sc.Agr., c/- C.S.I.R.O., Stowell Avenue, Hobart, Tasmania.

Tchan, Yao-tseng, Dr., és Sciences (Paris), Microbiology Laboratory, Faculty of Agri-
culture, Sydney University.

Thompson, Mrs. Joy Gardiner, B.Sc.Agr. (née Garden), 21 Middle Head Road, Mosman,
N.S.W.

Thomson, James Miln, D.Sc. (W.A.), Department of Zoology, University of Queensland,
St. Lucia, Brisbane, Queensland.

Thorne, Alan Gordon, B.A., 82 Ben Boyd Road, Neutral Bay, N.S.W.

Thorp, Mrs. Dorothy Aubourne, B.Se. (Lond.), Ph.D., “Sylvan Close’, Mt. Wilson, N.S.W

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

Tindale, Miss Mary Douglas, D.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., c/- Australian Museum, P.O. Box A285,

_ Sydney South, N.S.W.

Tucker, Richard, B.V.Se., Dr.Vet.M., Department of Veterinary Anatomy, University
of Queensland, Mill Road, St. Lucia, Queensland.

Tuma, Donald James, B.Se., c/- C.S.I.R.O., Division of Fisheries and Oceanography,
P.O. Box 21, Cronulla, N.S.W.

Valder, Peter George, B.Sc.Agr., Ph.D. (Camb.), School of Biological Sciences, Depart-
ment of Botany, Sydney University.

Vallance, Professor Thomas George, B.Sc., Ph.D., Department of Geology and Geophysics,
Sydney University.

Veitch, Robert, B.Sc., F.R.H.S., 24 Sefton Avenue, Clayfield, Brisbane, Queensland.

Vickery, Miss Joyce Winifred, M.B.E., D.Sc., Royal Botanic Gardens, Sydney.

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

*Voisey, Professor Alan Heywood, D.Sc., School of Earth Sciences, Macquarie University,
171-177 Epping Road, Eastwood, N.S.W.

Walker, Donald, B.Sc., M.A., Ph.D., F.L.S., 18 Cobby Street, Campbell, Canberra, A.C.T.
Walker, John, B.Sc.Agr., Biological Branch, N.S.W. Department of Agriculture, Private
Mail Bag No. 10, Rydalmere, N.S.W.
Walkom, Arthur Bache, D.Sc., 5/521 Pacific Highway, Killara, N.S.W.
Wallace, Murray McCadam Hay, B.Sc., Institute of Agriculture, University of Western
‘ Australia, Nedlands, Western Australia.
Ward, Mrs. Judith, B.Sc., c/- H.E.C., Gowrie Park, Tasmania.
Ward, Melbourne, Gallery of Natural History and Native Art, Medlow Bath, N.S.W.
Wardlaw, Henry Sloane Halcro, D.Sce., F.R.A.C.I., 71 McIntosh Street, Gordon, N.S.W.
Waterhouse, Douglas Frew, D.Sc., C.S.I.R.O., Box 109, Canberra, A.C.T.
*Waterhouse, John Teast, B.Sc., School of Biological Sciences, University of New South
Wales, P.O. Box 1, Kensington, N.S.W.
Waterhouse, Professor Walter Lawry, C.M.G., D.Se.Agr., M.C., D.I.C., FE.L.S., F.A.A.,
30 Chelmsford Avenue, Lindfield, N.S.W.
Watson, Professor Irvine Armstrong, Ph.D., B.Sc.Agr., Faculty of Agriculture, Sydney
University.
Webb, Mrs. Marie Valma, B.Sc., 8 Homedale Crescent, South Hurstville, N.S.W.
Webby, Barry Deane, Ph.D., M.Sc., Department of Geology and Geophysics, Sydney
University.
Webster, Miss Elsie May, 20 Tiley Street, Cammeray, N.S.W.
Whaite, Mrs. Joy Lilian, 357 Auburn Street, Goulburn 2S, N.S.W.
Wharton, Ronald Harry, M.Sc., Ph.D., C.S.1.R.O., 677 Fairfield Road, Yeerongpilly,
Queensland.
White, Andrew Hewlett, B.Sc. (Syd.), L. A. Cotton School of Geology, University of New
England, Armidale, N.S.W.
White, Mrs. Mary Plizabeth, M.Sc., 7 Ferry Street, Hunter’s Hill, N.S.W.
*Whitley, Gilbert Percy, F.R.Z.S., Australian Museum, College Street, Sydney.
Whitten, Maxwell John, Department of Botany, University of Tasmania, P.O. Box 252C,
Hobart, Tasmania.
Wildon, David Conrad, B.Sc.Agr., School of Biological Sciences, Department of Botany,
Sydney University.
Williams, John Beaumont, B.Sc., University of New England, Armidale, 5N, N.S.W.
Williams, Mrs. Mary Beth, B.Sc. (née Macdonald), 902D Rockvale Road, Armidale 5N,
N.S.W.
Williams, Neville John, B.Se., c/- Mr. Brian Williams, Albury North High School,
Albury, N.S.W.
Williams, Owen Benson, M.Agr.Se. (Melbourne), c/- C.S.I1.R.O., The Ian Clunies Ross
Animal Research Laboratory, P.O. Box 144, Parramatta, N.S.W.

1950
1959

1947
1965

1964

1964
1965

1949

LIST OF MEMBERS. 391

Willis, Jack Lehane, M.Sc., A.A.C.1., 26 Inverallan Avenue, Pymble, N.S.W.

Wilson, Frank, Officer-in-Charge, C.S.I.R.O., Division of Entomology, Sirex Biological
Control Unit, Silwood Park, Sunninghill, Ascot, Berks., England.

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

Woodward, Thomas Emmanuel, M.Sc. (N.Z.), Ph.D. (Lond.), D.I.C., Department of
Entomology, University of Queensland, St. Lucia, Brisbane, Queensland.

Wright, Anthony James Taperell, B.Sc., Geology Department, Victoria University of
Wellington, P.O. Box 196, Wellington, C.1, New Zealand.

Yaldwyn, John Cameron, Ph.D. (N.Z.), M.Se., Australian Museum, College Street, Sydney.
Young, Graham Rhys, 8 Spark Street, Earlwood, N.S.W.

CORRESPONDING MEMBER.

Jensen, Hans Laurits, D.Sc.Agr. (Copenhagen), State Laboratory of Plant Culture.
Department of Bacteriology, Lyngby, Denmark.

392

LIST OF PLATES
PROCEEDINGS, 1965

I.—Five species of Australian Chthamalidae.

II.—Some tropical Chthamalus spp. and their habitats.
III.—Chromosome numbers in some Australian leafhoppers.
IV.—V.—Tachyglossus aculeatus.

VI.—Bensonastraea praetor and Macgeea touti.
VII.—VIII.—Development of egg of Retropinna semoni (Weber).
IX.—Larvae of Retropinna semont (Weber).

X.—Xenogalea labiata: shelland part of egg mass.

XI.—X xX V.—Fine structure of the meristem of root nodules from some annual
legumes.

XXVI.—1. Melrosia rosae; 2. Melasmaphyllum mullamuddiensis.
XX VIT.—X X VITI.—Wheat seed germinated on agar medium.

LIST OF NEW GENERA AND NEW SPECIES
VoL. 90

New Genera

Page Page
Aloeurymela (EKurymelinae) aS. Melrosia (Stringophyllidae) .. 265
Melasmaphyllum
(Stringophyllidae) .. .. 267
New Species
gearyi (Aloeurymela) a 50) rosae (Melrosia) .. re .. 266
mullamuddiensis

(Melasmaphyllum) .. .. 269

Page

Abstract of Proceedings ..

Acoustic meatus, comparative ‘studies
on the external :

Anderson, D. T., elected President, A
Further observations on the life
histories of littoral gastropods in
New South Wales, 242—The repro-
duction and early life histories of
the gastropods, Notoacmaea petterdi
(Ten.-Woods), Chiazacmaea flammea
(Quoy and Gaimard) and Patelloida
alticostata (Angas)

Anderson, E. J., Plant parasitic nema-
todes in fruit tree nurseries of New
South Wales .. a oe vi

Angiosperms in the Tuggerah Lakes
system, the distribution of sub-
merged aquatic

Annual General Meeting. .

Anopheles farauti, a note on blood

preferences of

Anopheles farauti Laveran, malaria in
the D’Entrecasteaux Islands, Papua,
with particular reference to a5

Australia, South-western, the distribu-
tion of the Notonectidae in ;

Australian and some Indomalayan
Chthamalidae, a review of.. :

Australian birds, some mite parasites of

Australian larval Carabidae of the sub-
families Harpalinae, Licininae,
Odacanthinae and Pentagonicinae

Australian leafhoppers, chromosome
numbers in some ..

Australian Phacellophyllids, the Devonian
tetracoral Haplothecia and new
Australian smelt, Retropinna semoni
(Weber), development of the eees

and early larvae of the ..

Bacteria, studies of nitrogen fixing .

Baker, E. P., see Upashyaye: Y.M.,
and Baker, E. P. ..

Balance Sheets for the year “ending
28th February, 1965

Bamber, R. K., see Notes and Exhibits

Basden, R., The occurrence and com-
position of manna in LHucalyptus
and Angophora

Bassia All., an embryological study of
five species of

Bedford, Lynne, The ‘histology and
anatomy of the reproductive system
of the littoral gastropod, Bembicium
nanum (Lamarck) :

176
176

106

152
274

95

393

Page
Bembicium nanum (Lamarck), the
histology and anatomy of the

reproductive system of the littoral
gastropod a

Brooker, M. G., and Caughley, ce The
vertebrate fauna of “ Gilruth
Plains’, South-west Queensland. .

Cameron, Ann M., The first zoea of the
soldier crab, Mictyris longicarpus. .

Carabidae, Australian larval, of the
subfamiles Harpalinae, Licininae,
Odacanthinae and Pentagonicinae

Caughley, G., see Brooker, M. G., and
Caughley, G...

Cerioid Stringophyllidae " (Tetracoralla)
from Devonian strata in the Mudgee
district, New South Wales

Chromosome numbers in some Australian
leafhoppers

Chthamalidae, a review Rot Australian
and some Indomalayan

Comparative studies on the external
acoustic meatus. I. The morph-
ology of the external ear of the
echidna (Lachyglossus aculeatus)

Conservation Photographic
Exhibition ae 376,

Conversazione :

Dart, P. J. and Mercer, F. V., Observa-
tions on the fine structure of the
meristem of root nodules from
some annual legumes ‘

Development of the eggs and early larvae
of the Australian smelt, Fak opiane
semont (Weber) o6

Devonian tetracoral Haplothecia ‘and. new
Australian Phacellophyllids

Distribution of submerged aquatic Angio-
sperms in the Tuggerah Lakes
system

Distribution of the Notonectidae in
South-eastern Australia

Diurnal variation in the release of
pollen by Plantago lanceolata L.

Domrow, R., Some laelapid mites of
syndactylous marsupials, 164—
Some mite parasites of Australian
birds

Elections .. :
Embryological study of five Avot of
Bassia All...

95

238

222

157

. 263

78
10

. 176

377

. 378

252

218

5 Ue

328
87

. 231

. 190

4, 376-378, 380-381

274

394 INDEX
Page Page
Exhibits—see Notes and Exhibits Manna in LHucalyptus and Angophora,
First zoea of the soldier crab, Mictyris the occurrence and composition of 152
longicarpus .. . 222 Matthews, J. M., Diurnal variation in
Further observations on the life histories the release of Sa by ees
of littoral eee in New South lanceolata L. 231
Wales .. 2. 242 Members, List oF. 384
M
Gastropods, littoral, in. New South sh Wales, ES a Tee i sy Dart, Ee J. 2 and
further observations on the life Milward, N. E., Development “6 ihe
histories of .. 242
de (v eggs and early larvae of the
Gastropods, Notoacmaea “petter @ (Ten.- Australian smelt, Retropinna semoni
Woods), Chiazacmaea flammea (Quoy (Weber) 218
and Gaimard) and Patelloida alti- Mite \ io;

: parasites of Australian birds . 190
eee tae sare oe 106 Mites of syndactylous marsupials, some
ye fens : laelapid : 164

Gilruth Plains”, South-west Queens- Moore, B. Ey Australianilercyali@rnbicdes
land, the vertebrate fauna of . 238 Che Thao iline Harpalinae
Godwin, Eee H., Peer on Radio- at Licininae, Oia Nel
carbon Dating in the Quarternary Pentagonicinae 4 . _. 187
Higginson, F. R., The distribution of Mudgee district, New South Wales,
submerged aquatic Angiosperms in Cerioid Stringophyllidae (Tetra-
the Tuggerah Lakes system .. 328 coralla) from Devonian strata im
Hindmarsh, Gwenneth J., An embry- the... 263
ological study of five species of Muogamarra Sanctuary Volunteer Fire
Bassia All.” 2974 Brigade, appeal to younger members
Histology and anatomy of the repro- of the Society to join 377
ductive system of the littoral Mycological Society of New South Wales,
gastropod, Bembicium nanum proposed formation announced .. 378
(Lamarck) 95 Nematodes, plant parasitic, in fruit tree
Indomalayan Chthamalidae, a review nurseries of New South Wales .. 225
of Australian and some . 10 Note on blood preferences of fn
Tayostieatian of the Oy Phyllota 5S nha a pee bea 128
Benth.. : 341 oves and Ixnibits :
Bamber, R. K.—Specimen and
Jackson, D. L., see Tchan, Y. T., and photographs of fossil wood found
Jackson, D. L. in Mount Royal State Forest,
Jacobs, Janice L., see N: tes and lixhibiss New South Wales 383
Jancey, R. C., An investigation of the Jacobs, Janice L. (on behalf of Mr.
genus Phyllota Benth. 341—Numeri- R. Selkirk)—Specimens from
cal methods in taxonomy .. . 335 Kiandra, New South Wales . 382
E i Mahmood, A. (introduced by Dr.
Laelapid mites of syndactylous marsu- I. V. Newman)—Two electron-
pials.. Sean 20 -. 164 micrographs of phloem tissue of
Leafhoppers, _ Australian, chromosome Pinus radiata : 382
numbers in some ; eo Us Ramji, M. V. (introduced by Dr. I.
Lecturettes. . : 2, 376, 378-381 V. Newman)—Slides illustrating
Library Accessions 2, 376-378, 380-381 a study of embryogeny of
Linnaeus, portrait of, presented to Stellaria media .. 382
Society by Linnean Rovighy. of Vallance, T. G. _ "Attention drawn
London 381 to two significant tercentenaries
Tipnes Meee, Fellowships aR 4 in Natural Science which occurred
eappointmen oF oo in 1965 (Dr. Robert Hooke in
resignation of Mr. P. J. Dart, 3— 1665 issued Micrographia; and
applications invited for 1965, 3— birth of Dr. John Woodward)
appointment for 1965, 3—applica- —exhibition of copies of first and
_ tions invited for 1966 380, 381 second editions of Woodward’s
ee gueelony port of D Be ee An Essay toward(s) a Natural
10108y eport oO r. History of the Earth : 382
L — bh Te A 7 Ns ‘St di 4 Notonectidae, the distribution GE thes
uig, an. avson, pies in South-eastern Australia . .. 87
on the genetic nature of resistance Numerical methods in taxonomy . 335
to Puccinia graminis var. tritict in ; ;
six varieties of common wheat . 299 Obituary Notices : : : ,
C. Baehni, 5 ; Miss Vera Irwin-Smith,
Macleay, Sir William, Memorial Lecture, 5-6 ..
fourth, 1964, delivered : 2 Observations on the fine structure of
Mahmood, A., see Notes and Exhibits the meristem of root nodules from
Malaria in the D’Entrecasteaux Islands, some annual legumes é . 252
Papua, with particular reference to Occurrence and composition of manna in
Anopheles farauti Laveran . . . 155 Hucalyptus and Angophora. . . 152

INDEX

Page

Papua, malaria in the D’Entrecasteaux
Islands, with particular reference to
Anopheles farauti Laveran .

Pedder, A. E. H., The Dewanbera agian
coral H. aplothecia and new Australian

Phacellophyllids
Phyllota Benth., an investigation of the
genus ..

Plant parasitic nematodes in fruit tree
nurseries in New South Wales
Plates, List of
Pollen, diurnal wenn | in the) release
of, by Plantago lanceolata L.
Pope, Elizabeth C., A _ review of
Australian and some Indomalayan
Chthamalidae (Presidential Address)
Presidential Address Ae 1,
President’s Introductory Remarks
Proceedings, change of format . :
Puccinia graminis var. triticr ‘in six
varieties of common wheat, studies
on the genetic nature of resistance
to

Ramji, M. V., see Notes and Exhibits

Report on the Affairs of the Society

Reproduction and early life histories of
the gastropods, Notoacmaea petterdi
(Ten.-Woods), Chiazacmaea flammea
(Quoy and Gaimard) and Patelloida
alticostata (Angas) .. :

Review of Australian and some Indo-
malayan Chthamalidae

Root nodules from some annual legumes,
observations on the fine structure
of the meristem of .

Rules, alterations to TEhrlles, Vv and VI 379,

Rust resistance in oats, III, Studies on
the inheritance of

Science House ..

Soldier crab, Mictyris longicarpus, the
first zoea of the “is

Special General Meetings, 29th September
and 27th October, 1965 379,

155

181

. 341

. 225
. 392

. 231

292
380

395

Page

Spencer, Margaret, A note on blood
preferences of Anopheles farauti, 128
—Malaria in the D’Entrecasteaux

Islands, Papua, with particular
reference to Anopheles farauti
Laveran

Studies of nitrogen fixing bacteria, IX

Studies on the genetic nature of resistance
to Puccinia graminis var. tritici in
six varieties of common wheat

Studies on the inheritance of rust resist-
ance in oats. III. Genetic diversity in
the varieties Landhafer, Santa Fe,
Mutica Ukraine, Trispernia and
Victoria for crown rust resistance

Sulman, Florence, reference to death ..

Sweeney, A. W., The distribution of the
Notonectidae in South-eastern Aus-
tralia ..

Taxonomy, numerical methods in 6

Tchan, Y. T., and Jackson, D. L., Studies
of nitrogen fixing bacteria. IX.
Study of inoculation of wheat with
Azotobacter in laboratory and field
experiments

Tucker, R., Comparative studies on the
external acoustic meatus. I.

Tuggerah Lakes system, the distribution
of ge as eaves a aie
in the . eo

Upadhyaya, Y. M., aad ane EK. P.,
Studies on the inheritance of ES
resistance in oats. IIT.

Vallance, T. G., see Notes and Exhibits
Vertebrate fauna of “ Gilruth Plains ”’,
South-west Queensland é

Watson, I. A., see Luig, N. H., ca
Watson, I. A.

Whitten, M. J., Chromosome numbers i in
some Australian leafhoppers

Wright, A. J. T., Cerioid Stringophyllidae
(Tetracoralla) from Devonian strata
in the Mudgee district, New South
Wales .. ‘

115
290

. 299

129
377

87

335

290
176

328

5 WAY

238

78

263

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Proc. Linn. Soc. N.S.W., Vol. 90, Part 3 PLATE X

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Proc. Linn. Soc. N.S.W., Vol. 90, Part 3 PLATE XI

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Proc. Linn. Soc. N.S.W., Vol. 90, Part 3 PLATE XII

Fine structure of the meristem of root nodules from some annual legumes.

Proc. Linn. Soc. N.S.W., Vol. 90, Part 3 PLATE XII

Fine structure of the meristem of root nodules from some annual legumes.

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Proc. LINN. Soc. N.S.W., Vol. 90, Part 3 PLATE XIV

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Fine structure of the meristem of root nodules from some annual legumes.

PLATE XVI

Proc. Linn. Soc. N.S.W., Vol. 90, Part 3

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Fine structure of the meristem of root nodules from some annual legumes

PLATE XVII

Proc. Linn. Soc. N.S.W., Vol. 90, Part 3

Fine structure of the meristem of root nodules from some annual legumes.

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Proc. Linn. Soc. N.S.W., Vol. 90, Part 3 PLATE XVIII

Fine structure of the meristem of root nodules from some annual legumes.

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Proc. LINN. Soc. N.S.W., Vol. 90, Part 3 PATH xx

Fine structure of the meristem of root nodules from some annual legumes.

Proc. Linn. Soc. N.S.W., Vol. 90, Part 3

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Fine structure of the meristem of root nodules from

some annual legumes.

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Proc. Linn. Soc. N.S.W., Vol. 90, Part 3 PLATE XXI

Fine structure of the meristem of root nodules from some annual legumes.

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Proc. Linn. Soc. N.S.W., Vol. 90, Part 3 PLATE XXII

Fine structure of the meristem of root nodules from some annual legumes.

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Fine structure of the meristem of root nodules from some annual legumes.

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Proc, Linn, Soc, N.S.W., Vol. 90, Part 3 PLATE XXIV

Fine structure pf the meristem of root nodules from some annual legumes.

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Proc. Linn. Soc. N.S.W., Vol. 90, Part 3 PLATE XXVII

Wheat seed germinated on agar medium.

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Proc. Linn. Soc. N.S.W., Vol. 90, Part 3 PLATE XXIxX

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