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Proceedings of the
Linnean Society
of New South Wales

VOLUME 99
Nos. 437-440

CONTENTS OF PROCEEDINGS, VOLUME 99

PART 1° (No. 437)
(Issued 12th November 1974)

CONTENTS

Annual General Meeting :
Report on the Affairs of the Society for the Year ..
Elections
Balance Sheets
Domrow, R. Miscellaneous mites from Australian vertebrates. 1-48 ..

McLEAN, R. A. The rugose coral genera Streptelasma Hall, Grewingkia
Dybowski and Calostylis Lindstrém from the Lower Silurian of New
South Wales

PEPPERELL, J., and Gricc, G. A Labyrinthodont trackway from the
Mid-Triassic near Sydney, New South Wales : se an

RICHARDSON, L. R. A new troglobitic quadrannulate land-leech from
Papua (Hirudinoidea: Haemadipsidae s.1.)

ROBERTSON, R. N. Sir William Macleay Memorial Lecture, 1974. ‘“ A
Society of Natural Bee I hope they may succeed.” The first
hundred years a a a ie ae i

ScuHicHa, E., and DossE, G. A new species of Typhlodromus Scheuten
(Acarina: Phytoseiidae) from apple in Australia oe ate,

PART 2 (No. 438)
(Issued 12th February 1975)

CONTENTS

ALLEN, G. R. Four new damselfishes (Pomacentridae) from the south-
west Pacific. . :

3ALLANTYNE, B. Powdery mildew on Cucurbitaceae: identity, dis-
tribution, host range and sources of resistance 0% ae

Moork, K. M. The Glycaspis spp. (Homoptera: Psyllidae) associated
with BLucalyptus camaldulensis ss ian ais ous

SEMENIUK, V. Biogenic structures in the Ordovician Bowman Park
Group, New South Wales

Page

36

54

57

69

79

Page

PART 3 (No. 439)
(Issued 5th March 1975)

CONTENTS
Page
DENMARK, H. A., and ScHicHA, E. A new species of Amblyseius Berlese
(Acarina: Phytoseiidae) from apple in Australia .. as aol 45
Doo.tey, J. K., and PAxTon, J. R. A new species of tilefish Vaal
Branchiostegidae) from eastern Australia Sf ; : ios ie doy
Zt.ACEK, A. A., and HARRISON, F. W. The systematic and phylogenetic
position of Palaeospongilla chubutensis (Porifera: Spongillidae) .. 157
THO, C., and BAKER, E. P. Mutagenic effects of ethyl methanesulphonate
on the oat stem rust pathogen (Puccinia graminis f. sp. avenae) .. 166
PART 4 (No. 440)
(Issued 25rd April 1975)
CONTENTS
Page
DENMARK, H. A., and ScuHicHa, KE. A new species of Phytoseius eae
(Acarina : Phytoseiidae) from apple in Australia .. a A iGO
McLEAN, R. A. Silurian rugose corals from the Mumbil area, central
New South Wales aid iy ee ise hs “it ay tel onl
PILLAY, R. A., ToHAN, Y. T., and CHo, K. Y. Nucleic acids in foo ee
of a Chlorella Seer ve Ate se ae : 97
Notes and Exhibits sid “iid “he ies sis Se a eng) 202

Index ue He ap Phe a See vs es it ait 205

The Proceedings of the Linnean Society of New South Wales contain original papers relative
to ‘‘ the Science of Natural History in all its branches ’’, and reports of business transacted at the
meetings of the Society.

NOTES AND INSTRUCTIONS FOR AUTHORS

Original papers may be submitted for consideration for publication in the Society’s
Proceedings. Submission of a manuscript must imply that the contents are original and have not
been published previously in the same, similar, or abbreviated form, and that no similar paper is
being or will be submitted for publication elsewhere until it has been either (a) not accepted for
publication by the Society or (6) withdrawn by the author.

Papers submitted should embody the results of a significant piece of work. Interim reports,
serialized studies and student theses should not be submitted.

Papers are accepted for publication (or otherwise) by the Council of the Society after taking
into consideration advice received from at least one referee. The decision of Council is final.

Once accepted for publication a paper becomes the property of the Society, and copyright is
vested in the Society. Reproduction of the author’s synopsis in abstracting journals is authorized.

A paper may be withdrawn by the author at any time prior to acceptance.

Papers may be submitted by members or non-members of the Society but a paper submitted
by a non-member must be communicated by a member of the Society, and will not ordinarily be
considered for acceptance before July in each calendar year.

SUBMISSION OF MANUSCRIPTS

Manuscripts should be forwarded to the Secretary, The Linnean Society of New South Wales,
157 Gloucester Street, Sydney, N.S.W. 2000.

Text and illustrations must be submitted in duplicate. The illustrations may be duplicated
by any acceptable clear method and the duplicate set need not necessarily be identical in size
with the originals. The author should retain a third copy for checking proofs. If a paper is not
accepted for publication, the original copy of the typescript and illustrations will be returned to
the author, but the duplicate copy may be retained by the Society.

Owing to the ever growing costs of publication, authors are requested to write their papers
clearly and concisely and exercise all reasonable economy in the number and form of illustrations
and tables.

Grants towards the cost of publication should be secured wherever possible from the author’s
own institution or elsewhere ; contributions from authors, especially non-member authors, may
be requested at Council’s discretion, particularly in the case of lengthy papers, or those with
numerous illustrations or tables.

Manuscripts must be submitted in final, fully corrected form. Authors may be liable for the
cost of proof corrections (if allowed) if the number of alterations and corrections is considerable.
Authors are urged to submit their paper to a colleague for constructive criticism prior to final
typing and submission for publication.

MANUSCRIPTS

Papers should be typewritten, double spacing throughout, on one side of quarto or foolscap
paper, with at least one-inch margins all round. The desired position of all text-figures, maps
and tables may be indicated in the margin. All pages should be numbered serially and securely
fastened together. Papers should conform to the following general layout as far as practicable
or appropriate :

(a) Title—The title should be concise but informative. The name(s) of the author(s) and
the name of the institution where the work was carried out should follow the title. The name
of a new taxon of plants or animals should not be included in the title. A short title appropriate
for the running heading of subsequent pages may be suggested. The title, etc., should be on a
separate page at the commencement of the typescript.

(b) Synopsis.—A concise abstract (without abbreviations), preferably consisting of not more
than about 200 words, should be supplied on a separate sheet and in triplicate. It should be
complete in itself and indicate the scope of the work. It will be printed below the title of the

paper ; it will be available for reprinting by abstracting journals.
(c) Table of Contents.—This should give the complete hierarchy of headings and sub-headings

used in the subject-matter of the paper. It should be supplied on a separate sheet for guidance
of the editor and will not ordinarily be printed.

(1) Introduction, followed by the Subject Matter of the paper. Papers may be divided into
sections described by short headings.

(e) Discussion, Conclusions or Summary (if necessary).—This should summarize the conclusions
reached rather than be a précis of the paper as a whole (cf. Synopsis).

Proceedings of the
Linnean Society
of New South Wales

Issued Ilth November, 1974

VOLUME 99
PART |
No. 437

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, 1974-75

President
T. G. Vallance, B.Se., Ph.D.

Vice-Presidents
H. G. Cogger, M.Se., Ph.D. ; L. A. S. Johnson, D.Sc. ;
P. J. Stanbury, Ph.D. ; N. G. Stephenson, M.Sc., Ph.D.

Honorary Treasurer
Joyce W. Vickery, M.B.E., D.Sc., F.L.S.

Secretary
Mrs. Ruth J. Inall

Council
D. J. Anderson, Ph.D. Elizabeth C. Pope, M.Sc., C.M.Z.S.
Barbara G. Briggs, Ph.D. P. J. Stanbury, Ph.D
H. G. Cogger, M.Sc., Ph.D. N. G. Stephenson, M.Sc., Ph.D.
D. W. Edwards, B. Sc. Agr. P. G. Valder, B.Sc.Agr., Ph.D.
Mary M. Hindmarsh, B.Sc., Ph.D.* T. G. Vallance, B.Se., Ph.D.
L. A. S. Johnson, D.Se. Joyce W. Vickery, M.B.E., D.Sc., F.L.S.
D. McAlpine, M.Sce., Ph.D. B. D. Webby, M.Se., Ph.D.
Helene A. Martin, Ph.D.+ G. P. Whitley, F.R.Z.8.
Lynette A. Moffat, B.Sc., Ph.D. A. J. T. Wright, B.Se., Ph.D.t

P. Myerscough, M.A., D.Phil.
* Resigned 22/5/74. + Elected 19/6/74. { Elected 17/4/74.

Councillors Hmeriti
W. R. Browne, D.Sc., F.A.A. A. B. Walkom, D.Sc.

Auditors
W. Sinclair and Company, Chartered Accountants

Linnean Macleay Lecturer in Microbiology, University of Sydney
K. Y. Cho, Ph.D.

Inbrarian
Constance B. McKay, B.A., A.L.A.A.

Associate Editor
Janet Donald

The Society’s Headquarters are in Science House, 157 Gloucester Street, Sydney,
N.S.W. 2000, Australia

NOTES AND INSTRUCTIONS FOR AUTHORS

(f) Acknowledgements, if any.

(9) References.—The author, who is alone responsible for their accuracy, should carefully
check all references against the text, to ensure especially that :
(i) The spelling of authors’ names is consistent, and the dates, volume numbers and page
numbers are correct.
(ii) All authors quoted in the text are included in the list of references.
References should be cited in the text by the author’s name and date, e.g., Bullough (1939)
(Bullough, 1939) according to context, and listed alphabetically under References thus :
Bullough, W. 8., 1939.—A study of the reproductive cycle of the minnow in relation to the
environment. Proc. zool. Soc. Lond., Ser. A, 109 : 79-108.
If more than one reference by the same author(s) published in the same year is cited, use a, b
ete., after the year in both text and list, e.g., Bullough (1939a).
Titles of books must be given in full together with the place of publication, name of publisher
and indication of edition if other than the first.

Latin names of plants and animals (genera, species and their subdivisions) should be underlined
to indicate italics. An initial capital letter must be used for Latin names of orders, families and
similar names, but these should not be underlined.

Abbreviations.—Only standard abbreviations should be used; authors should refer to the
British Standard 1991: Part 1, 1967. Recommendations for Letter Symbols, Signs and
Abbreviations. The abbreviations of names of periodicals should conform to those in the World
List of Scientific Periodicals.

Footnotes should be avoided as far as possible.

Codes of Nomenclature.—The Botanical and Zoological Codes of Nomenclature must be strictly
followed. If a new taxon is described, the institution in which the type material is deposited
must be indicated.

Language.—Papers in languages other than English are unlikely to be accepted for publication.
Foreign authors are urged to seek assistance, if necessary, in rendering the paper in fluent English
prior to submission.

Measurements must be expressed in the metric system as far as practicable, and exclusively
in scientific description.

>

ILLUSTRATIONS AND TABLES
The maximum dimensions available for either text-figures or plates are 20 x 12-5 cm.

Black and white drawings and diagrams for text-figures should be in Indian ink on smooth
white card such as Bristol board, the lines cleanly drawn, and due attention paid to the
consequences of reduction. Graphs should similarly be in Indian ink on white card, or on faint
blue-lined co-ordinate paper. Maps should be in Indian ink on white card or tracing linen ; if
the latter the linen should not be folded. Maps should be of a size and scale that can be
conveniently reduced to the size of a single page.

It is desirable that all black and white illustrations should be drawn 14-2 times the size that
they will appear in print.

Photographs for half-tone plates should be the size required for final reproduction or slightly
larger, printed on glossy paper, of good contrast and best quality. A number of small photographs
can be arranged to form one plate.

Illustrations for colour reproduction cannot be accepted unless the author meets the difference
in cost between black and white and colour reproduction.

Illustrations of all kinds should bear the author’s name and the number of each, lightly
pencilled on the back, and their required orientation clearly indicated.

Legends for illustrations should be typed on separate sheets at the end of the manuscript.
Great care should be taken in stating magnifications of illustrations, making it clear whether
numbers quoted refer to before or after reduction.

Tables.—Tables should be numbered consecutively and referred to specifically in the text
by number. Lach table, together with a heading descriptive of its contents, should be submitted
on a separate sheet. Elaborate tables should be avoided.

GENERAL

Proofs.—A set of galley proofs will be forwarded to the author who should correct and return
them promptly to the Secretary.

Reprints.—Twenty-five reprints are supplied free to authors who are members of the Society-
Additional reprints may be ordered at the authors’ expense, if desired. No free reprints are
supplied to non-members, who may order the number required at their own expense. Orders
for reprints should be placed with the Secretary immediately after the paper has been accepted
for publication. Orders received after the paper has been printed may not be accepted.

Joint Authorship.—In the case of papers by more than one author, one of them must accept
responsibility for all correspondence with the Society regarding the paper.

AUSTRALASIAN MEDICAL PUBLISHING CO. LTD.
71-79 ARUNDEL ST., GLEBE, N.S.W., 2037
1975

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ANNUAL GENERAL MEETING
27th MarcH 1974

The Ninety-ninth Annual General Meeting was held in the Society’s Rooms,
Science House, 157 Gloucester Street, Sydney, on Wednesday, 27th March 1974
at 7.30 p.m.

Dr. P. J. Stanbury occupied the chair. The minutes of the Ninety-eighth
Annual General Meeting (28th March 1973) were read and confirmed.

REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR 1973

Publications

The Society’s Proceedings were published on the following dates during the
year :
1972 Vol. 97, Part 4 30th April 1973
1973 Vol. 98, Part 1 25th June 1973
1973 Vol. 98, Part 2 10th October 1973
1973 Vol. 98, Part 3 16th January 1974

The cost of publishing the Proceedings was as follows :

$ $
Blockmaking _ (Illus- Subscriptions 3,935.21
trations) 836.12 Miscellaneous Sales 682.83
Printing 2,870.10
Postage 103.70
Cost of Reprints : Printing : $960.76; Sales : $762.03

As reported last year, in order to comply with Post Office regulations, a
proportion of every member’s subscription is set aside specifically as a subscription
to the Proceedings ; this amount remains at $5.00 per annum.

It was regretfully decided during the year to increase the annual charge for
the journal from $11.00 to $15.00 for subscribers other than members, due to the
considerable increase in costs since the last variation in 1965/66.

Membership
During the year 13 new members were admitted to the Society and four
resigned. The numerical strength of the Society at the Ist March 1974 was
Ordinary Members, 268 ; Life Members, 29 ; Corresponding Members, 4 ; total, 301.

Monthly Meetings
Addresses were given at the meetings in March, April, June, July and Septem-
ber. A Field Day was held in October and the Notes and Exhibits evening took
place as usual in November. Details of these may be found in the Abstract of
Proceedings, Vol. 98, Part 4, (1973).

Library
Cataloguing of the serials held by the Society has now been completed.
New and amended entries for the serials collection were sent to C.S.I.R.O. in
March 1973 for inclusion in Scientific Serials in Australian Libraries. Many of
these entries have already appeared in amended pages of SSAL.

PROCEEDINGS OF THE LINNEAN Society oF New SoutH WaAtgES, Vou. 99, Part 1

2 REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR

The number of requests for photocopies of reference material not only from
libraries in Australia but from as far afield as Malaysia and New Zealand, has con-
tinued to increase.

Publications of the British Museum held by the Society have been catalogued,
and a subject list sent to all Australian university libraries, museums, herbariums
and relevant divisions of C.S.I.R.O.

A letter was sent to Sydney University requesting information on the progress
made regarding the arrangements and cataloguing of monographs sent to that
University to be held as a Special Collection. The reply received indicated that
the work was in progress but no catalogue cards have yet been received. Cards
were received for a number of serials which had been transferred because they
were not related to the Society’s subject fields.

Work has begun on the Cumulative Index to the Linnean Society’s Pro-
ceedings : 1926to date. It is hoped to publish this on the completion of volume
100.

Acknowledgement was received from the Mitchell Library for maps, portraits,
notes, etc. which relate to the Society’s affairs and are now held by that library.

Library Fund

The Royal Society of New South Wales recently established a Library Fund
to which donations are tax-deductible to the donors. The conditions imposed
by the Commissioner of Taxation permit the fund to be used for such purposes as
removal, rehousing, equipping and maintaining the library.

This Society has been quick to follow this example and has now been granted
the same concession on the same terms, and a Library Fund has now been estab-
lished.

The Councils of the two Societies have agreed to devote any moneys accumu-
lated in their respective Library Funds to the costs of alteration of such parts of
the new Science House as are required for library purposes and to equipping the
libraries in their new home.

It is intended to inaugurate a fund-raising scheme during the year, details of
which will be announced later.

Innnean Macleay Fellowship

Dr. Ross A. McLean was appointed Linnean Macleay Fellow as from 1st
January 1974 in the Department of Geology and Geophysics, Sydney University.
His research topic is a general study of rugose coral faunas of Middle and Upper
Silurian age from New South Wales.

Linnean Macleay Lectureship in Microbiology

The appointment continues to be held by Dr. K. Y. Cho. The main research
activities are still concerned with the physiology of the extreme halophilic
bacteria and the regulation of membrane synthesis in Azotobacter vinelandit.

Carotenoids of the extreme halophilic bacterium Sarcine morrhuae have
been characterised by thin layer chromatography, spectrophotometry, and
nuclear magnetic resonance. They are similar to those present in Holobacterium
salinarium, including the presence of the unusual C,, carotenoids.

Some of the possible membrane precursors in Azotobacter vinelandii have
been partially purified and characterised.

Finance

The Honorary Treasurer presented the Balance Sheets and Income and
Expenditure Accounts of the Society’s four Funds and made the following
comments on them :

PROCEEDINGS OF THE LINNEAN Society of New SourH WAtgES, Vou. 99, Part 1

REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YHAR 3

In the General Account balance sheet you will note that a year ago the
accumulated funds stood at $123,446.88. From this has been deducted the
advance of $15,500 on compensation for Science House that we had received by
then. This sum has subsequently been added into the surplus of $409,442.51 on
resumption of Science House. You will remember that we received a total of
$438,000 as compensation for the Society’s share of Science House. From that
sum has been written off both our previous investment of $30,000 in the building
and legal costs incurred in connection with the resumption. The net figure of
$409,442.51 includes our share of the small reserves in the hands of Science
House Management Committee.

Our expenditure during the year has exceeded income by $6,043.53. This
amount is approximately equal to the rent we now pay for our premises and the
Librarian’s salary. Our bank account in this Fund shows a deficiency of $449.33
as at 28th February 1974.

The right hand side shows how the moneys in this Fund are held. Under
fixed assets I point out the third item—the investment of one share of $1 in
Science House Pty. Ltd. Then are listed the various interest-bearing investments.
The second last, Science House, represents our share in the reserves held by Science
House Management.

Under current assets is listed the loan of $407,500 to Science House Pty. Ltd.
for the Science Centre project. While it is hoped that this will ultimately yield a
satisfactory return, immediate financial benefit cannot be anticipated. In early
March $7,500 of this loan was repaid to the Society.

In the Income and Expenditure Account, the printing costs of the Proceedings
seem much lower than last year. This year the bill for the last part was not
received before the balance sheet was drawn up, whereas last year the Account
included the cost of five parts.

Income was received from subscriptions both for membership and for the
Proceedings ; from reimbursement of salaries and fees from Science House Man-
agement ; from interest received from our investments ; from the surplus trans-
ferred from the Fellowships Account ; from rents ; from the sale of copies of the
Proceedings and reprints ; from our share in the surplus from Science House
Management.

The Bacteriology Account shows little change from last year. Our con-
tribution towards the salary of the Linnean Macleay Lecturer in the University
of Sydney has been increased to $2,000.

The Scientific Research Fund has increased through interest and donations
to $5,554.10.

The Fellowships Account has been augmented chiefly by the transfer of
$2,666.68 from its Income and Expenditure Account. Our Fellow resigned early
in the financial year and in accordance with Sir William Macleay’s Will the
remaining salary was capitalised.

Report on Science Centre

At the Annual General Meeting in 1971 members were informed that Science
House (owned jointly by the Linnean Society of N.S.W., the Royal Society of
N.S.W. and the Institute of Engineers) had received notice of resumption from
the Sydney Cove Redevelopment Authority. At that time negotiations were in
progress with the Authority and with the Government of New South Wales in
hopes of finding means whereby the learned Societies could rehouse themselves
on a site within the redeveloped area.

At the Annual General Meeting in 1972 members were advised that the
Societies had requested the Government of New South Wales to provide, as it
had done in 1928, a new site for these Societies to rehouse themselves. The

PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WALEs, Vout. 99, Part 1

4 REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR

Societies had resolved to accept responsibility for the building of a new Science
Centre, as they had in the building of Science House some 44 years ago. The
Government had encouraged the Planning Committee in its efforts and had
indicated that it would give as much support as possible to our scheme. The
Planning Committee, with a membership of representatives appointed from both
Societies, was meeting regularly and engaging in broad initial planning and in
clarification of the position of both Societies in relation to the project. A claim
for compensation for Science House had been made (through Science House
Management Committee, which represents jointly the three owner bodies). It
was the intention of the Linnean and Royal Societies to invest the compensation
money (excluding the amount which had been claimed in respect of disturbance
costs) in a new Science Centre. Sites in the Rocks area were under consideration,
and a firm of property consultants was assisting the Committee in its investi-
gations, together with the solicitors who were the legal advisers in respect of the
present Science House.

You were also advised that the Act of Incorporation of the Linnean Society
of New South Wales of 1884 was unduly restrictive in relation to powers of
investment and since this could hamper operations concerned with a new building
action was being taken to secure a revised Act of Incorporation. It was also
foreshadowed that in order to act jointly in relation to the Science Centre action
would be taken to form a company. Negotiations with State and Federal
governments were continuing.

Having briefly reviewed matters that have been reported to you earlier, I
would now like to record events up to the present.

By the middle of 1972 it was becoming clear that re-establishment within
the Rocks redevelopment area, as earlier envisaged, would not be a practicable
proposition. The Act under which the Sydney Cove Redevelopment Authority
operates stipulates that all land within the area shall be leasehold not freehold.
The normal term of lease would be 50 years. Though in our case it might have
been possible to have this extended to not more than 99 years, upon termination
of the lease all land and improvements on it (i.e. buildings) would revert to the
Government without compensation ; this meant that any money expended by the
Societies on a building would then be lost to them forever. Not being organis-
ations trading for substantial profits, the Societies conld not entertain this
proposition. We suggested that a lease be granted to us at a peppercorn rental
but this concession was not within the powers of the Authority to grant. At this
time and again at a later stage we asked for a government-guaranteed loan, but
our Societies do not fall within existing legislation enabling that to be done and
the Government of New South Wales has not agreed to altering the legislation to
enable it to be done. It was becoming clear that, although we cannot negate
benefits that may yet come, the attitude of the Government was that we had
received a generous amount of compensation and should stand on our own feet.

The Planning Committee therefore determined to seek a freehold property
that would be suitable for the erection of a Science Centre. It accepted the
principle that, to be appropriate for the Societies’ needs, any site selected must
be reasonably central and accessible to public transport.

Between about June 1972 and December 1973, the Planning Committee (or
when the Company was registered its Board of Directors) examined a number
of sites and feasibility studies of them to enable a choice to be made appropriate
to our needs and financial circumstances. As you can appreciate it is by no
means easy to acquire blocks of land in the city or its immediate environs and
valuations are very high. It soon became clear that sites within the central city
zone (between George and Elizabeth Streets) were quite beyond our means, so
thoughts were directed to areas outside but adjacent to the central area.

PROCEEDINGS OF THE LINNEAN Society or New SourH Wass, Vou. 99, Part 1

REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR 1)

It would be unavailing to detail either the propositions we have examined
which for one reason or another came to nothing or the hopes and frustrations
the Committee experienced. Suffice it to say that our property consultants
made offers on our behalt for two separate sites which were regarded as financially
suitable, and up until about August 1973 the Company had strong hopes of
obtaining one of these.

You will recall that about September 1973 a marked change in economic
circumstances took place in the community with a dramatic increase in the
already high rates of interest payable upon borrowings. Your Company’s
proposals were inevitably severely affected by this change since it would be
obliged to borrow heavily for erection of a building. Our property consultants
then expressed the view that within the context in which we had been thinking
we simply did not have a proposition at all. It seemed that matters had come
to a standstill.

For the purposes of making a feasibility study of one of the sites I have just
mentioned a firm of architects, Jackson, Teece, Chesterman and Willis, had been
commissioned to draw up sketch plans of a building so that it could be appro-
priately costed. Learning of our dilemma, this firm, late in October 1973, in-
formed the Societies of an existing building which was for sale and which they
felt could be adopted structurally to the Societies’ needs.

The Directors and their architects and financial advisers turned their atten-
tion to this new proposition of purchasing an existing building on an attractively
central and accessible site, and of progressively renovating it as a Science Centre.

The Directors resolved by majority vote on 14th November 1973 to proceed
with this proposition. Although it was manifest that strict economy and good
housekeeping would be required for its success, an equally good opportunity was
unlikely to occur again in the foreseeable future. Our Secretary had meanwhile
secured an undertaking for a bank loan on terms that under present conditions
must be regarded as unusually favourable. The Directors’ resolution to proceed
with the acquisition and development of the site was confirmed on 12th December
1973 after further study of all the implications.

I cannot conclude my report on the purchase of the site of our new home
without recording our indebtedness to the vendors for their co-operative approach
to all the problems and technicalities that beset the two parties during negoti-
ations leading to the signing of the contract.

I must also pay tribute to the tremendous effort of the Company’s Secretary,
who is also this Society’s Secretary, in furthering the project and by her initiative
and energy finding means of surmounting what at times appeared insuperable
difficulties.

Science House Pty. Ltd.

At the Annual General Meeting 1973 it was reported that the total com-
pensation offered and accepted for Science House amounted to $1-314 million,
and that the Linnean Society’s share of this was $438,000. <A lease agreement
had been negotiated with the new owner, the Sydney Cove Redevelopment
Authority, enabling the previous three owner bodies to continue to occupy and
manage Science House for the time being, in return for a reasonable annual
rental. Since the Institution of Engineers desired to be relieved of the manage-
ment services associated with Science House, which it had undertaken for many
years past, the Linnean Society with the concurrence of its Secretary, Mrs. Inall,
had agreed to accept this responsibility for a fee met jointly by the three bodies.

A Board of Directors for a proposed company had been nominated by the
Royal and Linnean Societies in anticipation of its early registration.

Our solicitors however advised against early registration of the company
under the New South Wales Companies Act (1961) until we had both reached

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutH WALES, VoL. 99, Part 1

6 REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR

a more advanced stage of planning, necessitating the use of the company for
joint action and received written assurance that, just as the Linnean and Royal
Societies were respectively exempt from income tax, so would the company
jointly owned by the two Societies be exempt from income tax.

Satisfactory advice to this effect having been received from the Taxation
Commissioner, it was resolved at the meeting of the Planning Committee on
15th March 1973 that the company be registered as soon as possible, each Society
owning one share of $1.00.

For purely historical interest it may be mentioned that an existing but in-
active registered company with the improbable name of Hodilia Pty. Ltd. was
taken over by the Planning Committee under the guidance of our solicitors. Its
previous directors resigned and the following persons were appointed by the
Councils of the two Societies as its Directors on 12th April 1973 :

Royal Society: Mr. Edric Keith Chaffer
Mr. John William Humphries
Mr. Maurice James Puttock
Professor William Erie Smith
Linnean Society : Professor Derek John Anderson
Dr. Harold George Cogger
Professor Neville George Stephenson
Dr. Joyce Winifred Vickery

Mrs. R. J. Inall was appointed Company Secretary. The registered office of the
Company is 3rd Floor, 157 Gloucester Street, Sydney.

At the first meeting of the new Board of Directors of Hodilia Pty. Ltd. on
26th April 1973, the memorandum changing the name of the Company to Science
House Pty. Ltd. was read. The Directors proceeded with business under this
name which we trust will continue for the indefinite future.

Dr. H. G. Cogger and Professor D. J. Anderson resigned from the Board of
Directors in November 1973, and the Linnean Society appointed Mr. D. W.
Edwards and Dr. P. J. Stanbury to the Board in their places.

Each Society has loaned to the Company the sum of $400,000 totalling
$800,000 free of interest for the time being, for the purposes of the Science Centre
project. This money has been invested on short-term loan and has meanwhile
earned interest that will enable the Company to meet its administrative expenses
and pay $810,000 towards the acquisition of the site and building.

Financial Advisers

You were informed at the last Annual General Meeting that the Science
Centre Planning Committee appointed three honorary financial advisers, Mr. J.
Studdy, Mr. J. Bragg and Mr. H. McCredie. The Company has valued their
advice during the last anxious year, and we are very gratefnl for the service they
have so willingly rendered.

Annual Elections

No nominations of other candidates having been received, the Secretary
declared the following elections for the ensuing year to be duly made :
President: Professor T. G. Vallance.
Members of Council : Dr. H. G. Cogger, Mr. D. W. Edwards, Miss I. C. Pope,
Dr. L. A. Moffat, Mr. G. P. Whitley.
Auditors : W. Sinclair & Co.

PROCEEDINGS OF THE LINNEAN SocieTy oF New SourH WaAtss, Vou. 99, Part |

REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR 7

PRESIDENTIAL ADDRESS

THE ROLE OF SCIENTIFIC SOCIETIES—SOME SUGGESTIONS FOR
THE FUTURE

by P. J. STANBURY

For the past year I have been President of our Society which we claim is
devoted to science. We are a wealthy Society. You know the plans for con-
structing Science Centre at a cost of two million dollars. We are a Society whose
membership has included most of the important natural scientists of this State.
World-renowned visitors have lectured in this room. Yet what benefit has the
wealth and talent of the Society brought to most of its members or to the general
public of New South Wales ?

Attendances at recent meetings have been very poor. Less than 10 per
cent of members have attended a monthly meeting in the last year. Can any
society continue to function under these conditions ?

The Royal Society of New South Wales had trouble with attendance over a
century ago. At that time it was called the Philosophical Society of New South
Wales. It solved its problems by attaining royal patronage and changing its
name.

At that Society’s recent annual dinner Sir Roden Cutler, one of its Patrons,
said :

‘““The rapidly increasing fields of scientific study and the proliferation of
scientific knowledge in recent years almost makes it an impossible task for
[a] society to keep the public informed. Scientific investigations and dis-
coveries in the last 25 to 30 years have both excited public imagination and
at the same time occasionally frightened the public. [A] society’s task is to
bring a balance into people’s assessment of the advantages and limitations of
scientific progress. You need to encourage research and investigation, and
occasionally express a word of warning ”’.

Since I have been Curator of the Macleay Museum at the University of
Sydney it has recovered from a period of sad neglect and is now growing and
expanding. The Macleay Museum grew, as you probably know, from the col-
lections of Alexander, William 8. and Sir William Macleay. Sir William donated
the total collection to the University in 1886. I have always felt his presence
there strongly. Today the Museum is being used and many thousands of people
each year have reason to feel grateful for Sir William’s generosity. In my work
for the survival and restoration of this Museum I have gained some experience in
determining what will raise a body from the dead and give it a new lease on life.

This year the Council of our Society has considered ways and means of
enlivening the meetings. The Future Role of the Society Sub-committee has
been formed. Soon you will receive a circular asking your views on the relative
importance of a number of activities, including the publishing of the Proceedings,
which is the one thing we believe we do well. We shall be trying out a new style
of meeting this year. Your response to these things will determine what we do
next year.

However, I am now concerned with the future of scientific societies in the
long term. How can Sir Roden Cutler’s remarks be brought to fruition? I can
think of two possible ways.

PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WALES, VOL. 99, Part 1

8 REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR

First, I believe that a society like ours could employ a public information
officer who would have the job of advertising the society’s activities and of in-
forming both its members and the public at large of its doings.

At one time no one paid much attention to scientists unless it was to remark
on their supposed eccentricity. However, by the 1930's scientists were becoming
regarded as people who could solve problems. Today scientists are accepted as
a normal part of society. The media on the whole report on the work of a
scientist or scientists with no more or less enthusiasm than on the doings of any
other individual or group in society. Most often the media pay no attention to
scientists at all unless scientists deliberately draw attention to themselves. As
a rule scientists are very hesitant to do this. The ones who do are usually most
adept at raising not only the interest of the public but also large sums of money
to be used on research.

With these considerations in mind I shall list the possible duties of a public
information officer for a scientific society. He should inform, advertise and
stimulate interest both within and outside the society. He would produce
newsletters for the members, which would contain, among other things, discussion
of the issues being considered by the society’s executive body. He would present
accounts of the members’ current research in the newsletters. He would be
responsible for feeding selected information of significance and interest to the
public media for dissemination to a wide audience. He should have the experience
and qualifications to maintain a strong working relationship with those who
control the media. An information officer must be free to speak independently
and should be responsible to the executive body of the society only through the
president. He must gain and maintain the confidence of the members of the
society. Scientists often need to be persuaded of the significance of their work,
and an information officer could assist in this area. The officer should present
the president or the executive body as the public image of the whole society, thus
providing some tangible link between the public and the members.

An information officer should address as many groups as possible both within
and outside the society. He must be an active man, providing liaison among
the executive body, the member scientists, the media and individuals and groups
of the general public. His job is to disspell inertia, apathy and ignorance. He
can do this by employing his knowledge and experience, by using tact, by radi-
ating and rousing enthusiasm and by using for the society’s profit the same
methods of persuasion employed by advertisers and salesmen in the commercial
world. An information officer must use modern methods.

This brings me to my second suggestion for enlarging and enlivening a
society’s activities. This is the use of radio.

It is hardly likely that a society like ours could acquire substantial regular
broadcasting time on the ABC or on commercial radio. However, we could
investigate the alternative types of radio which include adult and extension
programmes, University of the Air private stations, radio stations with low
output which are virtually closed circuit radio, and Frequency Modulation (FM)
radio.

If FM radio is introduced into Australia, it is probable that many community
group radio stations will spring up. I suggest that the Linnean Society could
make good use of an FM group station. We could, for example, broadcast
weekly in the evening, giving news of special interest to scientists ; details of
available grants ; job opportunities ; information on new equipment ; consumer
reports on scientific equipment; news and information for members; talks,
lectures and courses ; advice to conservation groups and other environmental
bodies ; reports on the best places to see flora, fauna and geological phenomena,
and many other activities.

PROCEEDINGS OF THE LINNEAN Socrety or New Soutrn Waxes, Vou. 99, Part 1

REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR 7)

If we found difficulty in compiling enough material for our own weekly
programme, we could join together with other societies for the purpose of the
broadcast. This idea is in line with the concept of the Science Centre that the
Royal and Linnean Societies are striving to establish. The Centre is envisaged
as a building in which many scientific societies may have offices and facilities for
their mutual benefit.

The initial cost of a radio station is $75,000. We should consider how our
Society can best look to the future and how it can best serve the general public.
No longer can scientific societies and scientists keep ostrich-like attitudes. The
future of this country depends on the co-operation of all members of its population.
How can you, as a scientist, best serve the community ?

PROCEEDINGS OF THE LINNEAN SOCIETY oF NEw SoutH WALES, Vot. 99, Part 1

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PROCEEDINGS OF THE LINNEAN Society oF NEw §

MISCELLANEOUS MITES FROM AUSTRALIAN VERTEBRATES. 1-48
ROBERT DomMRow*

[Accepted for publication 2lst November, 1973]

Synopsis
A miscellaneous collection of mite parasites of Australian frogs, reptiles, birds,and mammals
is detailed. The data are largely restricted to new host records and major extensions of
geographical range. The 48 species belong to the following 10 families: Blattisociidae 1,
Ameroseiidae 1, Dermanyssidae 4, Spinturnicidge 1, Sarcoptidae 2, Teinocoptidae 1, Knemido-
koptidae 1, Ereynetidae 1, Cheyletidae 1, and Trombiculidae 35.

Guntheria vegrandis, n. sp. (Trombiculidae), is described from Rattus fuscipes in South
Australia.

Over the years, many series of mites have accumulated beyond the scope of
current studies. Taken individually, these records may be minor, but as a
whole they provide a useful body of confirmatory records and considerable
extensions of known geographical and host ranges. Their publication as the
first of an occasional series will both place these data on record and underline
that the ‘‘ rarity ’’ of many species is due only to the desultory collecting of the
past. Only extensive and intensive collecting over the entire continent will
provide a full picture of the ranges and host spectra of the mites parasitic on
Australian vertebrates.

In the following list, families are in systematic order, and genera and species
in alphabetic order. For brevity, and since this is a largely non-taxonomic
paper, generic documentation is not given, this being available in Neave (1939-66).
All measurements are in micra. Hosts are largely given after Worrell (1963),
Leach (1958) and Ride (1970), a skeleton classification of the hosts listed being :

AMPHIBIANS: Frogs (Hyla).
REPTILES: Snakes (Liasis), lizards (Lygosoma, Varanus).

Birps: Hawks (Falco), owls (Ninox, Tyto), frogmouths (Podargus), king-
fishers (Sauromarptis), swifts (Collocalia), perching birds (Artamus, Chloris,
Climacteris, Eopsaltria, Hylacola, Hylochelidon, Malurus, Meliornis, Meliphaga,
Myzantha, Oriolus, Pachycephala, Pomatostomus, Sericornis, Struthidea).

MAMMALS: Marsupials (Antechinus, Cercartetus, Dasyurus, Isoodon,
Macropus, Potorous, Schoinobates, Thylogale, Trichosurus, Vombatus), rodents
(Melomys, Rattus), bats (Miniopterus, Pteropus), carnivores (Felis), primates
(Homo).

The collectors, indicated by their initials, are: A. Agadjanian, R. Ainslie,
W. H. Andrews, J. H. Arundel, G. J. Barrow, H. Battam, I. Beveridge, E. T.
Bulfin, J. H. Calaby, R. W. Campbell, L. Cannon, C. HE. Chadwick, G. de Chaneet,
K. J. Chipperfield, J. Culley, J. J. Davis, R. Domrow, A. L. Dyce, I. D. Fanning,
Penelope A. Graf, R. H. Green, E. Hamilton-Smith, W. B. Hitchcock, B. Elizabeth
Horner, R. Jasper, B. H. Kay, K. Keith, Elizabeth Knox, Jacky Lowry,
M. Josephine Mackerras, B. McMillan, Elizabeth N. Marks, G. M. Maynes,
J. A. R. Miles, B. C. Mollison, D. E. Moorhouse, D. J. Moss, B. L. Munday, J. B.
Paton, B. Pudney, K. Reye, N. Robinson, D. 8S. Saunders, J. H. Seebeck,
Rosamond Shepherd, H. A. Standfast, P. D. Strong, J. Mary Taylor, R. Tumman,
R. M. Warneke, J. S. Welch, S. H. Wheeler, R. H. Whitehead, G. Wolf, T. O.
Wolfe, J. Wolfenden and Patricia A. Woolley. I thank them all.

* Queensland Institute of Medical Research, Brisbane, Queensland, 4006.

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutTH WALES, VOL. 99, Part 1

16 MITES FROM AUSTRALIAN VERTEBRATES

I am also grateful to Misses Jenny Kiibler and Leanne Jackson for clerical
and technical assistance ; to Associate Professor D. J. Lee, School of Public
Health and Tropical Medicine, Sydney, and Mr. D. C. Lee, South Australian
Museum, Adelaide, for lending material from the Gunther, Hirst, and Womersley
collections; to Mr. M. Nadchatram, Institute for Medical Research, Kuala
Lumpur, for Malayan specimens of Siseca rara; and to Dr. P. H. Vercammen-
Grandjean for confirming my identification of Leptotrombidium robustum.

Family BLATTISOCIIDAE
Genus PROCTOLAELAPS Berlese

1. Proctolaelaps pygmaeus (Miller)

Gamasus pygmaeus Miller, 1859, Z. Naturw., 9: 29.

Garmania nesbitti Womersley, 1956, J. Linn. Soc., 42: 548. New synonymy.

Material. Many females causing annoyance, timber handlers, Brisbane, Qd,
ix.1964, J.J.D.

Three females, Rattus fuscipes, Palmerston Highway, Qd, 3-10.xii.1963,
G.J.B.

Notes. Hughes (1961) recorded this species both in association with mites
of stored food products and as a free-living predator.

Family AMEROSEIIDAE
Genus KLEEMANNIA Oudemans
2. Kleemannia plumosa (Oudemans)
Seiulus plumosus Oudemans, 1902, Tijdschr. Ent., 45: 17.
Material. Several females, Weelawadji Cave, near Eneabba, W.A.,
Ox 972, Je.
Notes. See notes above on Proctolaelaps pygmaeus.

Family DERMANYSSIDAE
Genus STRATIOLAELAPS Berlese
3. Stratiolaelaps miles (Berlese)

Laelaps (Iphis) miles Berlese, 1892, Acari, Myriapoda et Scorpiones hucusque
in Italia reperta, 63: 9.

Cosmolaelaps scumitus Womersley, 1956, J. Linn. Soc., 42: 580. New
synonymy.

Material. Two females, debris in total darkness in drains (inhabited by
bats) in disused railway tunnel, Samford, Qd, 30.iv.1963, R.D. Three females,
dark zone of Church Cave, Wee Jasper, N.S.W., 3.vi.1962, E.H-S.

Two females, Collocalia vanicorensis, Gatop, Finschhafen Subdistrict, New
Guinea, x.1960, B.McM.

Notes. See notes above on Proctolaelaps pygmaeus.

Genus IcHoRONYssUS Kolenati
4. Ichoronyssus minioptert (Zumpt and Patterson)

Neospinolaclaps mimioptert Zumpt and Patterson, 1952, J. ent. Soc. sth. Afr.,
be A159:

Material. Three females, Miniopterus schreibersii, Naracoorte, S.A.,
23.41.1963, E.H-S. One female, four males, and one protonymph, Miniopterus
tristis, lawarere Caves, 09° 29'S, 147° 38’ E, Central District, Papua, 10.vii.1973,
D.E.M. and G.W.

Notes. Although the population of MW. schreibersii in the Naracoorte caves
is isolated and biologically distinct from others in Australia, these specimens,

PROCEEDINGS OF THE LINNEAN Society oF New SourH WALES, VOL. 99, Part 1

ROBERT DOMROW aL

and those of Macronyssus aristippe below, are typical. On the other hand,
Spinturme loricata Domrow (Spinturnicidae) from this host at Naracoorte is
very distinct from S. psi (Kolenati) on other populations (Domrow, 1972).

Genus MAcronyssus Kolenati
5. Macronyssus aristippe (Domrow)
Ichoronyssus aristippe Domrow, 1959, PRoc. LINN. Soc. N.S.W., 83: 228.
Material. Numerous specimens, Miniopterus schreibersii, Naracoorte, 8.A.,
5.xi1.1961, 12.ii and 1.ix.1962, and 23.11.1963, E.H-S. One female, Miniopterus
australis, We Lifou Cave, New Caledonia, 1.1963, W.H.A.
Notes. See notes above on Lchoronyssus minioptert.

Genus OPHIONYSSUS Mégnin
6. Ophionyssus natricis (Gervais)
Dermanyssus natricis Gervais, 1844, Hist. nat. Insectes apt., 3: 223.
Material. Three females, Liasis childrent, Townsville, Qd, 24.vii.1966.
Notes. This species is a serious pest of snakes in captivity (Camin eéé al.,
1964), but the above new host was captured in the wild.

Family SPINTURNICIDAE
Genus MERISTASPIS Kolenati
7. Meristaspis calcarata (Hirst)
Ancystropus (Meristaspis) calcaratus Hirst, 1923, Proc. zool. Soc. Lond.,
1923: 983.
Material. Four females, three males, and one deutonymph (plus material
in spirit), Pteropus tonganus geddiei, Ipota, Eromanga, New Hebrides, 10.viii.1971,
A.A. (The Royal Society and Percy Sladen Expedition to the New Hebrides,
AQT):
Notes. This is a new host record (Domrow, 1972).

Family SARCOPTIDAE
Genus NOTOEDRES Railliet
8. Notoedres muris (Mégnin)
Sarcoptes notoedres var. muris Mégnin, 1877, Revue Mag. Zool., (3) 5: 71.
Material. Many specimens causing mange, Trichosurus vulpecula, Pomonal,
Vie., 31.11.1968.
Many specimens causing severe mange of head and forequarters, [soodon
macrourus, Paddington, Brisbane, Qd, 5.ix.1955, R.D.
Many specimens causing mange on ears and face, Melomys cervinipes, North
Stradbroke Island, Qd, 11.1973, E.K.
Notes. These records indicate that this parasite, whose normal hosts are
murid rodents, especially Rattus norvegicus (see Fain, 1968), can also establish
itself on a variety of native species.

Genus SARCOPTES Latreille
9. Sarcoptes scabier (Linnaeus)
Acarus Siro Secabiet Linnaeus, 1758, Systema Naturae. Holmiae: Laurentii
Salvii, 616.
Material. Many specimens causing mange, Vombatus ursinus, Saint Mary’s,
Mase 24 xd O71 Bale Me,
Notes. See notes above on Notoedres muris.

PROCEEDINGS OF THE LINNEAN Socrety or NEw SoutH WaAtEs, Vow. 99, Part 1

18 MITES FROM AUSTRALIAN VERTEBRATES

Family TEINOCOPTIDAE
Genus TEINOCOPTES Rodhain
10. Teinocoptes domrowi Fain
Teinocoptes domrowt Fain, 1961, Proc. LINN. Soc. N.S.W., 85: 269.
Material. Twenty females, Pteropus scapulatus, Mount Isa, Qd, ix.1966.
Notes. This is a new host record (Domrow, 19625).

Family KNEMIDOKOPTIDAE
Genus KNEMIDOKOPTES Firstenberg
11. Knemidokoptes jamaicensis Turk
Cnemidocoptes jamaicensis Turk, 1950 (sic), Parasitology, 40: 60.
Material. Numerous specimens causing scaly-leg, Chloris chloris, Beaumont,
S.A., 17.xii.1963, J.B.P.
Notes. This is a new host record (Fain and Elsen, 1967).

Family EREYNETIDAE
Genus LAWRENCARUS Fain
12. Lawrencarus hylae Fain
Lawrencarus hylae Fain, 1961, Bull. Annls Soe. r. ent. Belg., 97: 247.
Material. One pallid adult inside mouth near entrances to nasal passages,
Hyla caerulea, Kowanyama, Qd, 14.11.1966, R.D. and P.A.G. (Four of six other
H. caerulea with these data also harboured L. hylae, but the mites have been
lost.)
Notes. The other known specimens from this host bear only the data
Australia, 1880 (Fain, 1961).

Family CHEYLETIDAE
Genus CHELETONELLA Womersley

13. Cheletonella vespertilionis Womersley

Cheletonella vespertilionis Womersley, 1941, Rec. S. Aust. Mus., 7: 61.

Material. Three females, nest of Hylochelidon ariel, Richmond, N.S.W.,
B.MeM.

Notes. This species is probably a nidophile rather than a true parasite—
bats are well kncwn to frequent the disused nests of this bird (Kaiser and
Hoogstraal, 1973).

Family TROMBICULIDAE
Genus ASCOSCHOENGASTIA Ewing
14. Ascoschoengastia lorius (Gunther)
(Figs 1-8)

Neoschongastia lorius Gunther, 1939, Proc. LINN. Soc. N.S.W., 64: 86.

ELuschongastia daria Traub and Audy, 1954, Stud. Inst. Med. Res. F.M.S.,
26: 81. New synonymy.

Material. Holotype and five paratypes, the only specimens remaining
of the original series.

Notes. This little known species from a New Guinea parrot (Lorius) shows
the following scutal measurements :

AW PW SB ASB PSB SD AP AM AL PL Sens Specimen
3 6] 22 20 22 42 22 — 18 35 36 x 10 Holotype

40) 57 21 18 2] 39 20) 36 17 — 36 x 10 Paratype

42 57 22 18 23 4] — 32 17 — 35x11 nS

42 59 22 20 21 4] 20) 34 W7/ 34 38 x 10 op

39 56 2) 18 20 38 22 — ~ 35 35 x 10 AS

4] 58 22 19 2) 40 21 34 17 35 36 « 10 SS

PROCEEDINGS OF THE LINNEAN Society of New SoutH WALES, Vou. 99, Part 1

ROBERT DOMROW 19

8 pe i
ee [pe Sea. a)
o- i =. eG oc

Figs 1-8. Ascoschoengastia lorius larva. 1—2. Dorsal and ventral views of idiosoma
(engorged). 3. Scutum and eyes. 4. Dorsal view of capitulum (cheliceral blade half turned and
base foreshortened). 5. Ventral view of palpal tibiotarsus. 6-8. Specialized setation of legs
I-III. (All drawn from holotype except Fig. 3; each division on scales=100p.)

15. Ascoschoengastia rattus (Womersley and Heaslip)

Neoschongastia rattus Womersley and Heaslip, 1943, Trans. R. Soe. 8. Aust.,
8. Tie

Material. Three larvae, Trichosurus vulpecula, Kelso, Tas., 16.11.1961,
B.C.M.

Notes. The scutal measurements of two of these specimens are :

AW PW SB ASB PSB SD AP AM AL PL Sens

35 51 17 20 20 40 30 25 12 35 34x 8
33 53 19 20 20 40 28 23 12 36 35 x 8

34 52 18 20 20 40 29 24 12 36 34x 8

This is a new host record (Nadchatram and Domrow, 1964).

Genus EUTROMBICULA Ewing
16. Eutrombicula hirsti (Sambon)
Trombicula hirstti Sambon, 1927, Ann. Mag. nat. Hist., (9) 20: 157.

Material. Several attached larvae, forestry workers, Gin Gin, Qd,
16.11.1971, K.R.

PROCEEDINGS OF THE LINNEAN Society or New SourH Watss, Vot. 99, Part 1

20 MITES FROM AUSTRALIAN VERTEBRATES

Notes. Previous records of this itch mite in Australia have been from
North Queensland (Domrow, 1962b, 1967b).

17. Hutrombicula macropus (Womersley)

Trombicula macropus Womersley, 1936, J. Linn. Soc., 40: 112.

Material. Seven larvae attached to inner surface of knee, Macropus
giganteus, Kowanyama, Qd, 25.x.1966, R. D. LHighteen larvae attached inside
ear, Macropus agilis, Kowanyama, Qd, 25—-26.x.1966, B.D.

Notes. This is the first record of this species since the original series from
Macropus, Darwin, N.T., 1934.

18. EHutrombicula tovellt (Womersley)

Trombicula (? Neotrombicula) tovelli Womersley, 1952, Rec. S. Aust. Mus.,
Ope eElG:

Material. Eleven larvae, Varanus varius, Eidsvold, Qd, 25.xi.1960, M.J.M.
leven larvae, skink, Lower Ballunjui, Lamington Plateau, Qd, 12.11.1964, L.C.
Twenty-two larvae attached in axillae and inguina, Lygosoma fuscum,
Kowanyama, Qd, 14.11.1966, R.D. and P.A.G. Three larvae, Lygosoma chal-
lengert, Springbrook, Qd, 10.11.1955, E.N.M.; nine larvae attached in axillae,
L. challengert, Canungra, Qd, 111.1971, R.D. and R.W.C.

Notes. These new host records confirm the preference of this species for
lizards.

Genus GUNTHERIA Womersley
19. Guntheria alpina (Demrow)

Guntherana alpina Domrow, 1964, Acarologia, 6: 327.

Material. Nine larvae, Rattus fuscipes, Tommy’s Bend, Vic., 10.11.1963,
R.M.W.; five larvae, R. fuscipes, Mount Macedon, Vic., 18.xii.1963, R.M.W. ;
two larvae, R. fuscipes, Rifle Range, Pomborneit, Vic., 17.xii.1964, R.M.W.

Notes. These specimens, all from far south-eastern Australia, confirm the
original record from Mount Kosciusko, N.S.W.

20. Guntheria antipodiana (Hirst)
Schongastia antipodianum Hirst, 1929 (sie), Proc. zool. Soc. Lond., 1929: 175.
Material. Several larvae, Rattus fuscipes, Mount Magnificent, S.A.,
20.vii.1970, R.A.
Notes. These specimens, from the type host, are virtually topotypie.

21. Guntheria cassiope (Womersley)

Schéngastia (Ascoschéngastia) cassiope Womersley, 1952, Rec. S. Aust. Mus.,
UO Si.

Material. One larva attached at base of underside of bill, Meliornis novae-
hollandiae, road from Bulli to Appin, N.S.W., 26.vii.1967, H.B.

Many larvae attached inside ears, Rattus sordidus, Kowanyama, Qd,
iv.1969, R.D. and E.T.B. Twenty larvae attached inside ears, Rattus leucopus,
Dayman Point, Qd, 12-19,vi.1970, R.D. and R.W.C.

Notes. See notes below on Guntheria coorongensis and Leptotrombidium
deliense with reference to the first two series. The third is a new host record.

22. Guntheria coorongensis (Hirst)
Schéngastia coorongense Hirst, 1929 (sic), Ann. Mag. nat. Hist., (10) 3: 565,
Material. Three larvae attached around cloaca, Hylacola pyrrhopygia,
four miles west of Fairy Meadow, N.S.W., 27.xii.1967, H.B. Seventy larvae
attached mostly around cloaca, but also on head and abdomen, and among

PROCEEDINGS OF THE LINNEAN Society of New Sourn Wars, Vou. 99, Part 1

ROBERT DOMROW 21

scapulars and retrices, Sericornis frontalis, near Fairy Meadow, N.S.W.,
27.xi1.1967, 29.1.1968, 24.11.1968, 26.11.1967, 16.11.1968, and 13-15.iv.1968,
H.B.; three larvae, S. frontalis, Cordeaux Dam, N.S.W., 29.11.1969, H.B. Two
larvae, Malurus cyaneus, one mile west of Keira, N.S.W., 21.v.1972, H.B. Nine
larvae attached around cloaca, Meliornis novaehollandiae, four miles west of
Fairy Meadow, N.S.W., 13.11.1967, H.B.

Several larvae, Antechinus stuartit, near Melbourne, Vic., 29.vii.1972, B.S. ;
two larvae, A. stuartii, Powelltown, Vic., 20.viii.1973, I.B.

Seven larvae, Rattus fuscipes, Pearl Beach, N.S.W., 4.1.1955, B.E.H. and
J.M.T.; many larvae, RA. fuscipes, Bat’s Ridge, Vic., 19—25.1.1968, J.H.S.; many
larvae, R. fuscipes, Bemm River and Bonang, Vic., 21—27.11.1968, J.H.S. and
R.T.; several larvae, R. fuscipes, Dargo Road near Mount Saint Bernard, Vic.,
14.11.1967, K.J.C.; several larvae, R. fuscipes, Otway Ranges, Vic., 13.iv.1968,
J.H.S. ; several larvae, R. fuscipes, Heathmere, Vic., 22.vii.1968, J.H.S. ; several
larvae, R. fuscipes, Rifle Range, Pomborneit, Vic., 17.xii.1964, R.M.W. Many
larvae, FR. fuscipes, Carey’s Gully, Mount Lofty, 8.A., vi-vii.1969, J.C. and B.P. ;
several larvae, R. fuscipes, Mount Magnificent, S.A., 20.vii. 970, R.A.; two
larvae, Rk. fuscipes, Kangaroo Island, 8.A., 11.1969, S.H.W. Several larvae,
Rattus lutreolus, Bemm River, Vic., 26.11.1968, R.T.; several larvae, R. lutreolus,
The Grampians, Vic., 28.iv.1968, J.H.S.; several larvae, R. lutreolus, Healesville
Sanctuary, Vic., 20—21.viii.1964, R.M.W.

Notes. Two considerable collections of chiggers from small ground mammals
in north Queensland (Domrow, 19625, 1967b) yielded large numbers of Guntheria
spp., but only one species—G. innisfailensis (Womersley and Heaslip), see Domrow
(1960a)—was recorded from a bird until Brennan (1965) and Brennan and
Amerson (1971) consistently obtained G. domrow? (Brennan) from various waders
in the central Pacific. The present records (and those of G. cassiope above and
G. pannosa below) indicate birds are regular, if minor, hosts for Guntheria spp.

The specimens from South Australia may be regarded as topotypes, the
species having been originally described from Robe.

23. Guntheria derricki (Womersley)
Neoschongastia derricki Womersley, 1939, Trans. R. Soc. S. Aust., 63: 162.

Neoschongastia similis Womersley and Heaslip, 1943, Trans. R. Soc. S. Aust.,
67: 124. New synonymy.

Guntherana (Derrickiella) rec Domrow, 1960, Pacif. Insects, 2: 212. New
synonymy.

Material. Thirteen larvae, Antechinus stuartvi, Condor Creek, A.C.T.,
26.11.1964, P.A.W. Five larvae, Antechinus swainsonii, above Silverband Falls,
The Grampians, Vic., 3.xi.1962, R.M.W.

One larva, Rattus fuscipes, Pearl Beach, N.S.W., 4.1.1955, B.E.H. and J.M.T. ;
10 larvae, R. fuscipes, Bat’s Ridge, Vic., 21.1.1968, J.H.S.

Notes. Domrow (1960a), while noting minor variation, maintained the
above three nominal taxa; more extensive collections, however, indicate
Synonymy. G. similis normally shows DS commencing 2.4.4, though 2.5.4 was
noted once and 2.6.4 13 times. G. derricki normally shows 2.6.6, but this pattern
is not uncommon (I can only say “‘ not uncommon ”’ since these specimens were
deliberately selected) in long series that can only be assigned to G. rex (73 random
specimens of which show 2.8.8 in 21, 2.8.7 in 19, 2.8.6 in 30, and 2.7.6 in three
Specimens).

The present series, the first collected outside Queensland, are also best
assigned here. All are typical of G. derricki except that from Condor Creek.
Here, the DS commence 2.8 in four, 2.9 in two, 2.10 in six, and 2.11 in one

PROCEEDINGS OF THE LINNEAN Society or New SoutH WAtzEs, Vot. 99, Part 1

22 MITES FROM AUSTRALIAN VERTEBRATES

specimen. It is difficult to decide the number of setae in the next row, but 8-10
is a fair approximation. Typical specimens have the following scutal standard
data in micra:

AW PW SB AS BaP SB ras!) ii Ate PAU AC ss wee Sens

66 85 33 26 22 48 17 61 29 90
72 89 37 24 25 49 17 59 29 77
65 81 31 23 23 46 16 58 24 82
64 82 33 24 24 48 14 64 25 88
67 85 34 24 25 49 16 60 29 90 .
67 85 37 24 25 49 16 61 29 87 39x 18

Pil |

67 84 34 24 24 48 16 60 28 86 39 x 18

24. Guntheria kallipygos (Gunther)
Neoschéngastia kallipygos Gunther, 1939, PRoc. LINN. Soc. N.S.W., 64: 83.
Material. Hight larvae attached to reddened, slightly scabby area on inner
surface of thigh, Macropus giganteus, Yan Yean, Vic., x.1972, J.H.A.
One larva, Trichosurus vulpecula, Kelso, Tas., 16.11.1961, B.C.M.

Notes. Previous records of this species are from New Guinea to south-eastern
Queensland (Womersley, 1952; Domrow, 1961).

25. Guntheria newmant (Womersley)

Schéngastia (Ascoschéngastia) newmant Womersley, 1952, Rec. S. Aust. Mus.,
10: 427.

Material. The syntype slide—this contains 16 over-cleared specimens.

Twenty-six larvae, Cercartetus nanus, Yackandandah, Vic., 23.11.1965,
R.M.W.

Five larvae, Antechinus flavipes, Tarago, near Goulburn, N.S.W., 29.11.1970,
N.R.; 14 larvae, A. flavipes, Mundaring Road two miles from Kalamunda, W.A.,
10.iv.1963, P.A.W. Ten larvae, Antechinus stuart, Pearl Beach, 30 miles
north-east of Sydney, N.S.W., 26.vii.1955, B.E.H. and J.M.T.; one larva,
A. stuartii, Paddy’s River, A.C.T., 19.iv.1964, K.K.; five larvae, A. stuart,
Mallacoota, Vic., 23.vili.1965, J.W.

Eight larvae, Rattus rattus, Pomonal, Vic., 2.iv.1968, J.H.S.

Seven larvae attached to ear, domestic cat, Shenton Park, W.A., 25.1.1973,
G. de C.

Notes. Interestingly, the only other known series is also from a domestic
cat (Western Australia, 1931).

26. Guntheria pannosa (Domrow)

Guntherana (Guntherana) pannosa Domrow, 1960, Pacif. Insects, 2: 225.

Material. One larva attached among retrices, Sericornis frontalis, Mount
Keira, N.S.W., 13.1.1968, H.B.

Notes. The only other known specimen of this species is from North
Queensland.

Q°-
is
)
oe |

. Guntheria peregrina (Womersley)
Schingastia (Ascoschéngastia) peregrina Womersley, 1952, Rec. S. Aust.
Musi, 105 220;
Material. Three paratypes.
Twenty-two larvae, Schoinobates volans, Esk-Toogoolawah district, Qd,
8.11.1964, R.D. and I.D.F.

PROCEEDINGS OF THE LINNEAN Society oF New SourH Wass, Vou. 99, Part 1

ROBERT DOMROW 23

Hight larvae, Antechinus bellus, Victoria, Port Essington, N.T., 10.vii.1965,
J.H.C.; 15 larvae, A. bellus, Mount Brockman Range, N.T., 11.vii.1972, T.O.W.

Notes. The only other known series of this species is from North Queensland.
28. Guntheria philippensis (Philip and Woodward)

Neoschongastia philippensis Philip and Woodward, 1946, Am. J. trop. Med.
26: 158.

Material. About 20 larvae, Isoodon macrourus, Nourlangie Camp, South
Alligator River, N.T., 17.viii.1962, J.H.C.

Notes. These specimens, and their scutal measurements, fit the original
description so closely that the presence of this species in Australia can no longer
be doubted (Domrow, 1960a, 1961).

Scutal standard data in micra of G. philippensis

ANWY ® SAP se tSilay | SANS Be ERS Bie GISID)e JAE ANE SUA Dine SET Sens

48 69 19 21 12 33 21 25 42 42 —
53 73 18 21 13 34 22 22 41 — —
49 66 20 22 12 34 22 24 37 46 —
49 68 19 20 ql 31 21 25 43 43 —
43 66 16 20 10 30 21 24 40 —- —
50 75 20 20 12 32 22 24 — 45 25 x 16

49 70 19 21 12 32 22 24 41 44 25x 16

29. Guntheria quatuor Domrow

Guntheria quatuor Domrow, 1972, J. Aust. ent. Soc., 11: 103.

Material. Several larvae, Rattus fuscipes, Bat’s Ridge, Vic., 19—25.1.1968,
J.H.S.; several larvae, R. fuscipes, Kalorama, Dandenong Ranges, Vic.,
18.11.1965, J.H.S.: several larvae, R. fuscipes, Bemm River and Bonang, Vic.,
21—26.11.1968, J.H.S. and R.T.; one larva, R. fuscipes, Rifle Range, Pomborniet,
Vic., 17.xii.1964, R.M.W. Two larvae, Rattus lutreolus, Bemm River, Vic.,
26.11.1968, R.T.

Notes. Although this species was described from Western Australia, the
present series show all three diagnostic characters (entire sensillary insertions in
front of PL, six setae in first row of DS, and three genualae I), and the following
minor deviations from the original description are therefore considered to fall
within the range of intraspecific variation: AM>AL (the opposite is true of the
type series); galeal setae with one-three branches (but occasionally nude as in
the type series) ; dorsal palpal tibial seta nude (in the type series, this seta may
be so lightly branched as to appear virtually nude).

Scutal standard data in micra of G. quatuor

AW PW SB ASB PSB SD SPs CAN Ai Pir Sens
57 81 17 33 16 49 38 57 46 69 35 X 20
59 80 17 33 16 49 41 50 53 72 34 x 18
63 86 20 30 13 43 37 53 49 73 36 X17
60 83 17 32 13 45 39 o7 43 72 36 x 20
61 85 20 33 13 46 37 54 43 Tl 33 x 19
64 86 20 32 14 46 39 58 46 73 36 x 20

ea FPP ihe i SUSE? a LE PAAR IS. SI fal SN Mchuaey =
cE 4 PIE 1 gs anes rea aers 24 F PARAS) ul Sp CLLN Gc Une me te Man ce a) =
BSE eB pies ay ONES I ay Perel NY AMT ey Ip a PE ae
Sage oe tO. 67) as

61 84 18 32 14 46 38 56 48 72 35 x 19

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24 MITES FROM AUSTRALIAN VERTEBRATES

30. Guntheria queenslandica (Womersley)

Neoschongastia queenslandica Womersley, 1939, Trans. R. Soe. S. Aust.,
63: 162.

Material. Numerous specimens, domestic cat, Colo Vale, N.S.W.,
15.viii.1960, A.L.D.

Notes. This is a new host record, but Domrow (1960a) recorded the species
from this locality on rabbits.

31. Guntheria shieldst (Gunther)
Neoschingastia shieldsi Gunther, 1941, Proc. LINN. Soc. N.S.W., 66: 158.
Material. One larva, Cercartetus nanus, Yackandandah, Vic., 23.11.1965,
R.M.W.
Two larvae, Antechinus stuartii, near Melbourne, Vic., 29.viii.1972, B.S.

Notes. Previous records of this species are from New Guinea and Queensland
(Domrow, 1960a). :

32. Guntheria smitht (Womersley)
Neoschongastia smitht Womersley, 1939, Trans. R. Soc. S. Aust., 63: 164.

Material. Ten larvae, Rattus leucopus, Dayman Point, Qd, 16.vi.1970,
R.D. and R.W.C.

Notes. This is a new host record (Domrow, 1962b, 1967b).

33. Guntheria vegrandis, n. sp.
(Figs 9-15)

Diagnosis. G. vegrandis belongs in the subgenus Derrickiella Audy and
Domrow (see Domrow, 1960a), and keys out near G. dumosa (Womersley, 1952).
The latter species, however, shows stout, stiffly barbed dorsal setae, a convex
posterior scutal margin, subglobose sensillae, and tarsala I set distally, almost
level with the subterminala. The new species has a biconvex posterior scutal
margin as in G. coorongensis, but is readily separable in that the dorsal setae
commence 2.10.10.10 rather than 2.6.6.6.

Types. Holotype and nine paratype larvae attached inside ears, Rattus
fuscipes, Carey’s Gully, Mount Lofty, S.A., vi and vii.1969, J.C. and B.P.
Holotype and three paratypes in Australian National Insect Collection ; six
paratypes in Queensland Institute of Medical Research.

One larva (not a type), Antechinus stuartui, Powelltown, Vic., 23.vii.1973,
EB:

Larva. Palpi of usual proportions, with three-pronged tibial claws.
Setation B.bnnb.5B+T; subterminala absent. Capitular setae two, well
branched. Galeal setae nude. Chelicerae unarmed except for usual tricuspid
cap; bases appearing foreshortened in all specimens, but without obvious
posterolateral expansions.

Body with annulate cuticle. Dorsal setae arranged

2.10(11,12).10(11,12).10(11) +28.
Intercoxal setae 2+2. Ventral setae 60.

Sentum with anterior margin sinuous, lateral margins slightly concave,
and posterior margin biconvex. Surface lightly punctate except behind AM and
SB. PL>AM AL, with PL and AL set on distinct corners, and AM well behind
margin. Sensillae spiculate, clavate, and set slightly in front of PL. LHyes
2+2, but weak, particularly posterior pair.

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ROBERT DOMROW 25

t i
Le Mh 4 i 7
iN
fh > a i A \\
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\
aan. Gra.
| iy ff n pee fe \
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; A
ere |
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, nN iN fe ® )
(7 To ae ip yn ke |
nN ier rh} q AF dh ath RK
VA k 4 ; \ Pg
A i : ie N Li gf oF |
we : ; ®
# >
nm Ay
N il }
c {
rf \4 |
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Zin j F
va j |
A \ A p |
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4 \ je. f
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fi. ater |
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i RR if
FAY YIN i"
fie
Na Teva
oe
Ween —SA—xq_q_uu
ara AS soap BO)

ne Rag ceatas
AEN BBO, C36

Ss Vi
SSS

Figs 9-15. Guntheria vegrandis larva. 9-10. Dorsal and ventral views of idiosoma (en-
gorged). 11. Dorsal and ventral views of capitulum (cheliceral base foreshortened). 12. Scutum
and eyes. 13-15. Specialized setation of legs I-III. (Each division on the scales = 100 uw.)

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26 MITES FROM AUSTRALIAN VERTEBRATES

Scutal standard data in micra of G. vegrandis

AW PW SS Be SAS Bases bres) PAM. CANES Pay Sens
74 94 28 29 20 49 33 44 41 67 35 x17
74 95 29 29 20 49 32 48 — 66 34x 18
73 90 29 27 20 47 30 45 38 67 —
71 90 27 48 a 72 37x 16
74 93 30 31 19 50 32 48 — 72 —

74 90 29 31 20 51 35 50 42 72 —

73 92 29 29 20 49 32 47 40 69 35X17

Leg I with pretarsala, subterminala, parasubterminala, tarsala and micro-
tarsala; two tibialae and microtibiala; three genualae and microgenuala.
Leg II with pretarsala, tarsala and microtarsala ; two tibialae ; genuala. Leg III
with tibiala; genuala. Coxae 1.1.1 (two paratypes show two setae on coxa III
on one side ; the specimen from Antechinus shows two setae on both coxae III) ;
trochanters 1.1.1; basifemora 1.2.2; telofemora 5.4.3; genua 4.3.3; tibiae
8.6.6; tarsi not counted (of these barbed setae, none are unduly long and out-
standing). All tarsi with two simple claws and empodium.

Notes. The specific name is a Latin adjective (vegrandis, not very heey
and refers to the diminutive eyes.

Genus LEPTOTROMBIDIUM Nagayo et al.
34. Leptotrombidium akamushi (Brumpt)

Trombidium akamush Brumpt, 1910, Précis de Parasitologie. Paris: Masson
& Cie, 506.

Material. Several larvae, Rattus exulans, Nendo Island, Santa Cruz, Solomon
Islands, J.A.R.M.

Notes. These typical specimens confirm the known limit of this essentially
Asian vector of scrub typhus (Womersley, 1952). The disease occurs still further
south-east in the New Hebrides, where the vector is inferred to be L. deliense,
the following species (Audy, 1961).

Fortunately, Philip’s proposal (1961) now before the International Com-
mission on Zoological Nomenclature to validate this name is not compromised
by Vercammen-Grandjean’s (1969) selection of a ‘‘ lectotype ’—though his
formal designation on page 100 reads ‘*‘ holotype ’’—in circumstances that demand
a neotype (see Articles 74 and 75).

35. Leptotrombidium deliense (Walch)
Trombicula deliensis Walch, 1922, Geneesk. Tidjschr. Ned.-Indié, 62: 552.

Material. Two larvae attached inside ear, Rattus sordidus, Belbora,
Kowanyama, Qd, iv.1969, R.D. and E.T.B.

Notes. An unexpected record. Previous Australian material was all from
the eastern side of Cape York Peninsula, normally in tropical vegetation (particu-
larly rain forest) with over 60 inches of rain per year. Kowanyama is on the
western side of the Peninsula, but watered by the Mitchell River, which rises
far to the east. The annual rainfall is only about 45 inches, almost all of which
falls in a circumscribed ‘‘ wet’ (December—March) ; the “ dry’ (April-November)
is characterized by heat and very low humidity (Domrow, 1967a). However,
both infested rats were collected in tall grass close to dense undergrowth under a
clump of mango trees along the banks of permanent pools at Belbora. <A
situation not unlike this was noted at Rocky Creek, Qd, described by Barrow,
Domrow and Derrick (1963).

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ROBERT DOMROW 27

36. Leptotrombidium myzantha (Womersley)

Trombicula (Leptotrombidium) myzantha Womersley, 1952, Rec. S. Aust.
Sn WOK ile.

Material. Six larvae, Falco cenchroides, Kowanyama, Qd, iv.1969, R.D.
Numerous larvae, Ninox novaeseelandiae, Kowanyama, Qd, iv.1969, R.D.,
A.U.D. and H.A.S. Numerous larvae, Ninox connivens, Kowanyama, Qd,

iv.1969, R.D., A.L.D. and H.A.S. Numerous larvae, Tyto alba, Kowanyama,
Qd, iv.1969, R.D., A.L.D. and H.A.S.

Numerous larvae, Podargus strigoides, Kowanyama, Qd, iv.1969, R.D.,
A.U.D. and H.A.S.; six larvae, P. strigoides, Mudginberri, N.T., 31.iii.1971,
A.L.D. and H.A.S. Numerous larvae, Podargus papuensis, Kowanyama, Qd,
iv.1969, R.D., A.L.D. and H.A.S.

Ten larvae, Pomatostomus temporalis, Condamine, Qd, 12.1.1966, R.D.,
D.J.M. and J.S.W. Two larvae, Pachycephala rufiventris, Charleville, Qd,
18.1.1966, R.D., D.J.M. and J.S.W. Three larvae, Meliphaga flava, Kowanyama,
Qd, 25.x.1965, R.D. Five larvae, Myzantha melanocephala, Condamine, Qd,
12.1.1966, R.D., D.J.M. and J.S.W. ‘Ten larvae, Struthidea cinerea, Condamine,
Qd, 12.1.1966, R.D., D.J.M. and J.S.W.

Notes. Previous records (Gill, Moule and Riek, 1945; Domrow, 1966) are

from a wide variety of passeriform birds. The present series, however, are
largely from non-passeriform predators (one diurnal kestrel and five species of
nocturnal owls and frogmouths). Almost all of more than 30 predators examined
were infested, the number of chiggers per bird ranging up to 1,800. There was a
light admixture (5-10%) of a second bird-specific chigger, Neoschoengastia
americana (see below), the infestations doubtless having been acquired while
taking prey in dense grassland under open sclerophyll forest.

Scutal standard data in micra of L. myzantha

AWieleRWir SBC] ASB) PSB) SD AP VAM Ab) Pil; Sens

70 79 23 29 16 45 30 33 40 56 58

37. Leptotrombidium robustum (Gunther)
(Figs 16-22)
Trombicula robusta Gunther, 1941, Proc. Linn. Soc. N.S.W., 66: 157.
Material. Wolotype and one paratype, the only specimens remaining of

the original series from Bulolo, New Guinea (on the same slide, with holotype
not differentiated and host not indicated).

Fourteen larvae, Sauromarptis gaudichaud, Vanimo, New Guinea, 25.iv.1965,
W.B.H.

One larva, Podargus strigoides, Mudginberri, N.T., 31.ii1.1971, A.L.D. and
H.A.S.

Notes. This little known species belongs to a group of bird chiggers which
engorge heavily, leaving the vertically orientated scutum almost impossible to
examine. Thus only one of the type series and one of the 14 from Vanimo
are measurable.

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28 MITES FROM AUSTRALIAN VERTEBRATES

(~

ee (
as
~/ q
iM nn ne ert
{ \

Figs 16-22. Leptotrombidium robustum larva. 16-17. Dorsal and ventral views of idiosoma
(engorged). 18. Dorsal and ventral views of capitulum. 19, Scutum and eyes. 20-22. Specialized
setation of legs I-III. (Each division on the scales = 100 yp.)

PROCEEDINGS OF THE LINNEAN Society or New Sours Wates, Vou. 99, Part 1

ROBERT DOMROW 29

The Mudginberri specimen was taken with L. myzantha (see above), but shows
DS 2.10.8.6.4.2.2 and VS 17.4.4.2. This compares better with the counts for
specimens from Vanimo, which show DS 2.10—-13.8-12.8-10.4-6.2.2 (av.
2.12.10.8.4.2.2), VS 12-15.4.4.2. The usual count for DS in L. myzantha is
only 2.8.6.6.4.2.2. The scutal measurements of this specimen also compare
better with undoubted specimens of L. robustum than with L. myzantha.

Scutal standard data in micra of lL. robustum

PAW Vitaiesk Wie an Ss) eS SADR SS. a58!19) ANTEX PANIES UNIS a BIB a Srey ov) Locality

73 81 25 31 13 44 30 42 48 52 50 Bulolo
73 Tel 23 34 13 47 31 48 49 57 51 Vanimo
77 81 25 34 16 50 33 49 49 58 64 Mudginberri

74 80 24 30 14 47 31 46 49 56 55

I have placed this and the preceding species as conservatively as possible,
noting that the nymph as well as the larva of Womersley’s species is very similar
to those of Leptotrombidium (see Domrow, 1957, 1960b ; Nadchatram, 1967).

The plethora of taxa proposed for various members of this complex—
Toritrombicula Sasa, Hayashi and Kawashima, Vorcana Audy, Tsutsugamushia
Asanuma, Cotrombidium Verecammen-Grandjean, Neacariscus Vercammen-
Grandjean, Whartonacarus Vercammen-Grandjean, and Toriacarus Vercammen-
Grandjean and André (the lattermost with the same nominal type-species as
Cotrombidium)—prompts some comment on recent trends in trombiculid
taxonomy.

Complicated tables of genus-group taxa have been proposed, based essentially
on the sensory setation of the larval appendages, with no attention to post-larval
stages, and all characters equally weighted. This rigidity, coupled with a lack
of attention to convergent (adaptive) characters and parallelism, has led to taxa
containing species mismatched zoogeographically.

Further, it is implicit in this method that taxa may be validly defined by
differences. But even Linnaeus saw that one cannot first choose a set of
characters, and then decide that all species showing them are congeneric. It
is the genus that dictates the characters, and the taxonomist must first recognize
natural groups of species and then decide which characters diagnose the genus.

The former approach leads to a certain logicality, though two unfortunate
aspects are the prediction of hypothetical groups and their subsequent pro-
liferation as extremely small taxa. Any resulting system may be useful in that
it is practical, but the natural (phylogenetic) classification we still lack will be
based on degrees of similarity between taxa.

Genus NEOSCHOENGASTIA Ewing

38. Neoschoengastia americana (Hirst)

Schongastia americana Hirst, 1921, Ann. Mag. nat. Hist., (9) 7: 37.

Neoschongastia americana solomonis Wharton and Hardcastle, 1946, J.
Parastt., 32: 292. New synonymy.
Paraschéngastia thomasi Radford, 1946, Proc. zool. Soc. Lond., 116: 262.
New synonymy.

Neoschingastia entomyza Womersley, 1952, Rec. S. Aust. Mus., 10: 265.
New synonymy.

Material. One paratype of N. entomyza.

Numerous larvae, Ninox novaeseelandiae, Kowanyama, Qd, iv.1969, BR.D.,
A.L.D. and H.A.S. Numerous larvae, Ninox connivens, Kowanyama, Qd,
iv.1969, B.D.. A.U.D. and H.A.S.

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30 MITES FROM AUSTRALIAN VERTEBRATES

Numerous larvae, Podargus papuensis, Kowanyama, Qd, iv.1969, R.D.,
A.L.D. and H.A.S.

One larva, Pomatostomus temporalis, Condamine, Qd, 12.1.1966, R.D., D.J.M.
and J.S.W. One larva, Artamus minor, Kowanyama, Qd, 27.iv.1969, A.L.D.
Fifteen larvae, Pachycephala rufiventris, Charleville, Qd, 18.i1.1966, R.D., D.J.M.
and J.S.W. One larva, Struthidea cinerea, Condamine, Qd, 12.1.1966, R.D.,
D.J.M. and J.S.W.

Notes. In all clear specimens, including the paratype, the long outstanding
seta on tarsus III is distinctly ciliated, the sternal setae are arranged 2+-2, and
the DS 2.8.6.4.6.4.2. A few specimens from Charleville show only six setae in
the first dorsal row, but are ciearly conspecific (see Brennan, 1951). Likewise,
Malayan material with intercoxal setae 2+-4 (Domrow and Nadchatram, 1960)
is retained here.

Scutal standard data in micra of N. americana

AW PWS Br ASB PS Bar oS Des 7A a Advis AN PL Sens Locality
43 62 26 22 26 48 33 35 46 45 29x16 Kowanyama
46 67 == 23 34 57 35 34 42 42 — ie
40 64 25 19 32 51 29 33 — 48 — Ss
45 61 28 16 34 50 DY 34 43 43 3017 As
43 65 22 25 Del 52 30 — 35 50 — Charleville
41 60 29) 18 27 45 32 28 37 46 17 in
43 67 25 21 25 46 27 —- 37 45 _ Condamine
43 64 25 21 29 50 30 33 40 46 3017

39. Neoschoengastia posekanyi Wharton and Hardeastle

Neoschéngastia posekanyt Wharton and Hardcastle, 1946, J. Parasit., 32:
302.

Material. Fifteen larvae attached to sternum, Hopsaltria australis, Bulli,
N.S.W., 2.11.1968, H.B. Four larvae attached to abdomen, Sericornis lathami,
Bulli, N.S.W., 1.1.1968, H.B. Seven red-orange larvae attached in opening
of swellings on skin near cloaca, Sericornis frontalis, Bulli Pass, N.S.W.,
31.xii.1966, P.D.S. Four red-orange larvae attached in opening of swelling on
skin caudal to sternum on brood pouch, Pachycephala pectoralis, Bulli Pass,
N.S.W., 1.1.1967, P.D.S.; one larva attached to abdomen, P. pectoralis, four
miles west of Fairy Meadow, N.S.W., 26.1i1.1967, H.B. One red-orange larva
attached in opening of swelling on skin caudal to sternum on brood pouch,
Meliphaga lewinti, Bulli Pass, N.S.W., 1.1.1967, P.D.S.; 18 larvae attached
near cloaca, M. lewinii, Bulli, N.S.W., 1.1.1968, H.B. Five deep orange larvae
attached to bare patch of skin on abdomen, Meliornis novaehollandiae, near
Mount Keira, N.S.W., 4.i1.1967, H.B. Twelve larvae, Oriolus sagittatus, Esk,
Qd, 13.11.1968, R.D. and B.H.K.

Notes. These are all new host records (Domrow 1966).

Genus NEOTROMBICULA Hirst
40. Neotrombicula antechinus (Womersley)
Trombicula antechinus Womersley, 1954, Trans. R. Soc. S. Aust., 77: 69.

Material. Several larvae, Antechinus stuartii, Mount Nebo, Qd, 3.viii.1966,
R.H.W.

Several larvae, Rattus fuscipes, Mount Glorious, Qd, ix and x.1955, R.D.

Notes. Secutal measurements (including a seta from the first dorsal row)
of this topotypic material indicate that the shallowness of the posterior scutal
margin (PSB 19) figured by Womersley is real, and the species is therefore
maintained.

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ROBERT DOMROW 31

Scutal standard data in micra of N. antechinus

AW, SEW) SB ASB PSB: SD” AP OAM AL, PL Sens DS

70 86 27 31 21 52 30 54 — 52 93 53
64 87 29 32 19 51 32 45 — 50 95 45
62 81 29 32 17 49 30 45 40 — 96 49
64 83 27 32 19 51 30 45 — 51 — 49
63 82 26 32 19 51 32 — 38 43 — 46

61 82 27 33 20 53 33 46 43 46 90 45

64 84 28 32 19 51 31 47 40 48 94 48

41. Neotrombicula comata Domrow
Neotrombicula comata Domrow, 1961, PRoc. LINN. Soc. N.S.W., 86: 82.

Material. One larva attached to abdomen, Sericornis frontalis, Mount
Keira, N.S.W., 24.ix.1967, H.B. One larva, Climacteris leucophaea, Mount
Keira, N.S.W., 24.ix.1967, H.B.

Many larvae, Macropus parma, Doyle’s River State Forest, 45 miles north-
west of Wauchope, N.S.W., viii.1973, G.M.M.

One larva, Dasyurus maculatus, Bessiebelle, Vic., 28.vii.1964, R.J. Five
larvae, Antechinus stuartii, near Melbourne, Vic., 29.viii.1972, R.S. One larva,
Antechinus swainsonit, Sherbrooke Forest, Dandenong Ranges, Vic., 4.vii.1966,
R.M.W.

Hight larvae, Rattus fuscipes, Heathmere, Vic., 22—24.vii.1968, J.H.S.

Notes. 'The only previous record is from a bandicoot (Perameles) in northern
New South Wales. The present specimens agree closely with the original
description, the DS commencing 2.12-16.10-14. Average scutal measurements
are given below.

Scutal standard data in micra of N. comata

AW PW SB ASB PSB SD AP) AM © Al PL Sens Host
79 98 30 35 20 55 37 S. frontalis
78 103 30 37 20 57 40 65 55 73 — C. leucophaea
719 106 38 — — — — D. maculatus
82 107 39 39 20 60 41 62 58 64. — A. stuartii
78 101 38 40 20 60 39 — 58 66 — A. swainsonii
74. 93 31 38 19 57 40 51 54 54 lll R. fuscipes

42. Neotrombicula derricki (Womersley)
Trombicula derricki Womersley, 1954, Trans. R. Soc. S. Aust., 77: 67.

Material. Three paratypes and several other specimens with the same
data.

Notes. This species is distinct from other Australian species of Neotrombicula
with one mastitibiala and two mastitarsalae on leg III (the other five are also
listed in this paper) in showing two genualae I (both internal and external distal
rather than internal distal alone), a long outstanding seta on telofemur III,
and strongly branched sensillae.

43. Neotrombicula gemini Domrow

Neotrombicula gemini Domrow, 1971, J. Aust. ent. Soc., 10: 112.

Material. Twenty-nine larvae, Rattus fuscipes, Heathmere, Vic.,
21—24.vii.1968, J.H.S.; three larvae, R. fuscipes, Forrest, Vic., 2.x.1967, J.H.S.
and R.T.

Notes. These specimens confirm the original record. Variation from DS
commencing 2.8.8 is minimal; in 40 specimens, four showed 2.8.7, four 2.9.8
and one 2.8.9.

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32 MITES FROM AUSTRALIAN VERTEBRATES

44. Neotrombicula novaehollandiae (Hirst)
Trombicula novaehollandiae Hirst, 1929, Proc. Zool. Soc. Lond., 1929: 172.
Material. Two syntypes.

One larva attached near cloaca, Sericornis trontalis, Mount Keira, N.S.W. ,
24.ix.1967, H.B.

Eleven red larvae around eyes and mystaciai vibrissae, and in ears, Macropus
rufogriseus, Maydena, Tas., 15.1x.1961, B.C.M. Four larvae, Macropus parma,
Doyle’s River State Forest, 45 miles north-west of Wauchope, N.S.W., viii.1973,
G.M.M. Twenty-four red larvae attached as for M. rufogriseus, Thylogale
billardiervi, Maydena, Tas., 4.1x.1951, B.C.M. Nine larvae attached behind
scrotum, Potorus tridactylus, Cobargo, N.S.W., 8.ix.1968, A.L.D.

Hight larvae, Trichosurus vulpecula, North Midlands, Tas., 2.vii.1962, R.H.G.

Five larvae, Dasyurus maculatus, Bessievelle, Vic., 29.vii.1964, R.J.; nine
larvae, D. maculatus, Piper’s River, Tas., 10.vi.1973, B.L.M. Eleven larvae,
Dasyurus viverrinus, Deddington, Tas., 27.vil.1972, R.H.G. Thirty-three larvae,
Antechinus flavipes, Mount Robinson, 8.A., 29.vi.1970, R.A. Six larvae,
Antechinus stuartii, near Melbourne, Vic., 29.vili.l972, R.S. Thirteen larvae,
Antechinus swainsonti, near Gembrook, Vic., 13.11.1967, D.S.S.; nine larvae,
A. swainsoni, Sherbrooke Forest, Dandenong Ranges, Vic., 4.vii.1966, R.M.W.

Thirty-five larvae, Rattus fuscipes, Mount Robinson, 8.A., vi.1970, R.A. ;
32 larvae, R. fuscipes, Mount Magnificent, S.A., 20.vii.1970, R.A.; two larvae,
R. fuscipes, Heathmere, Vic., 21.vii.1968, J.H.S.; 24 larvae, R. fuscipes, Forrest,
Vic., 1-2.x.1967, J.H.S. and R.T. ; 31 larvae, R. fuscipes, Cordeaux Dam, N.S.W.,
10.vii.1966, C.E.C. Two larvae, Rattus lutreolus, Forrest, Vic., 2.x.1967, J.H.S.
and R.T.

Notes. This material is doubly valuable: the localities cover much of
south-eastern Australia (including Tasmania), and the specimens from South
Australia are virtually topetypes. The scutal standard data (average values)
decrease towards the north, but clearly only one species is involved.

Scutal standard data in micra of N. novaehcllandiae

AWE SPW Sib ASS SERS Bias Dp pales ss SAtV ee AU ee ea Sem Sy teas Locality

74 95 29 37 26 62 34 57 53 76 101 73 South Aust-
ralia

7 95 29 36 26 62 34 57 54 68 106 64 Tasmania

71 94. 30 38 24 62 34 57 57 Wy 2 1KCO}8) 70 Victoria

74 99 31 3 24. 63 37 63 52 Thay i LLLP 74 Cobargo

69 88 26 33 24 ‘537/ 31 43 46 55 95 54 Cordeaux Dam

68 87 ail 33 23 56 31 44 46 55 97 57. Doyle’s River

The specimens from Gembrook, while showing a certain delicacy of structure
and scutal measurements rather smaller than other Victorian specimens, are also
assigned here.

AW PW SB ASB PSB SD AP “AM AL Pi Sens eis

66 86 27 34 22 56 38 45 46 50 — 58
72 88 28 35 2] 56 40 46 43 56 101 57
67 86 26 35 22 57 37 50 43 58 —_— 58
72 37 26 34 23 57 37 53 — 58 98 62
66 86 25 32 20 52 34 53 45 61 — 61
69 88 27 39 22 57 37 45 45 53 95 53
69 87 26 34 22 56 37 49 44 56 98 58

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ROBERT DOMROW 33

45. Neotrombicula thylogale (Womersley)

Trombicula thylogale Womersley, 1954, Trans. R. Soc. S. Aust., 77: 71.

Material. Three paratypes and several other specimens with the same
data.

Notes. Scutal measurements of this material indicate that the shallowness
of the shield (AP 25) figured by Womersley is real, and the species is therefore
maintained.

Scutal standard data in micra of N. thylogale

AW PW SB ASB PSB SD AP AM AL PL _ Sens DS

64 80 26 32 21 53 25 — — — — 50
66 82 28 31 25 56 25 32 40 43 — 46
67 85 30 31 25 56 25 36 —= 46 95 48
64 81 27 30 22 52 25 35 — 49 — 50
67 84 28 30 22 52 25 34 41 50 —- 46

65 81 27 30 22 52 25 35 40 48 90 51

66 82 28 31 23 54 25 34 40 47 92 48

Genus SisEcA Audy
46. Siseca rara (Walch)

Trombicula rara Walch, 1924, Trans. V bienn. Congr. fur east. Assoc. trop.
Med. (Singapore, 1923), 593.

Material. Two larvae attached in axillae, Lygosoma tenuis, Colearra Creek,
near Yeppoon, Qd, 31.viii.1965, H.A.S.; 14 parchment-coloured larvae attached
in axillae, L. tenuis, Manifold Station, near Rockhampton, Qd, 4.ix.1965, H.A.S.

Notes. This species has not previously been recorded from Australia,
but the scutal measurements of the above specimens agree with Womersley’s
(1952) larger values for material from the Philippines and New Guinea (it should
be noted that this author’s material from Malaya and Queensland has since been
described as two distinct species, S. subrara Audy and S. southcotti Womersley
and Audy, see Domrow, 1962a).

Scutal standard data in micra of S. rara

AW PW SB ASB PSB SD AP AM AL PL _ Sens

86 §=102 62 17 45 62 36 25 23 36 —

92 110 64 19 52 71 36 — 24 38 —
86 =:108 61 19 = 45 64 32 — — — 61
87 105 61 19 46 65 35 — 20 34 —

79 97 55 20 44 64. 39 24 20 32 58
85 102 58 20 47 67 33 27 22 34 —
85 §=6108 59 21 48 69 35 27 22 36 —

86 §=6103 60 19 47 66 35 26 22 35 60

47. Siseca vandiemeni Domrow

Siseca vandiemeni Domrow, 1962, J. ent. Soc. Qd, 1: 23.

Material. Twenty-six larvae, Lygosoma ocellatum, North Scotdale, Tas.,
21.1.1962, R.H.G.

Notes. This is a new host record (Domrow, 1962a).

Genus TROMBICULA Berlese
48. Trombicula quadriensis Womersley and Heaslip
(Figs 23-29)
Trombicula quadriense Womersley and Heaslip, 1943 (sic), Trans. R. Soc.
S. Aust., 67: 79.

Material. One paratype from Cairns, Qd.
Three larvae, Trichosurus vulpecula, D’ Aguilar Range, Qd, 4.iv.1957, R.D.

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34 MITES FROM AUSTRALIAN VERTEBRATES

Figs 23-29. Trombicula quadriensis larva. 23-24. Dorsal and ventral views of idiosoma
(engorged). 25. Dorsal and ventral views of capitulum. 26. Scutum and eyes. 27-29. Specialized
setation of legs I-III. (Hach division on the scales=100u.)

One larva, Antechinus stuartii, near Melbourne, Vic., 29.viii.1972, B.S. ;
seven larvae, A. stuartii, Powelltown, Vic., vii-vili.1973, 1.B.

Notes. Previous records of this species as originally understood are all from
North Queensland (Domrow, 1962b, 1967b). Standard scutal measurements
are:

AW PW SB ASB PSB SD AP AM AL PL Sens Locality

57 82 22 18 20 38 23 29 29 4] —~— Palmerston
56 3 22 20 20 40 Oe, 32 29 37 53 ig
55 80 20 20 19 39 22 26 26 40 — i

3 66 16 20 19 39 22 — 26 32 — D’Aguilar
49 3 18 2) 20 4) 21 — 25 32 — m,
53 8] 17 24 20 44 22 25 30 43 — Melbourne

52 78 19 20 20 40 22 28 28 38 53

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ROBERT DOMROW 35

References

Aupy, J. R., 1961.—The ecology of serub typhus. Jn May, J. M. (ed.), Studies in Disease Ecology.
New York: Hafner, 389-432.

Barrow, G. J., Domrow, R., and Derrick, E. H., 1963.—Rocky Creek, an outlying focus of
scrub typhus in north Queensland. Australas. Ann. Med., 12: 166-170.

BRENNAN, J. M., 1951.—Two new species of Neoschéngastia with a key to the species of the world
(Acarina: Trombiculidae). J. Parasit., 37: 577-582.

, 1965.—A small collection of chiggers (Acarina : Trombiculidae) from the north central
Pacific. J. Parasit., 51: 888-892.

, and Amerson, A. B., 1971.—Six new species and additional records of chiggers from
the central Pacific (Acarina: Trombiculidae). J. Parasit., 57: 1311-1317.

Camin, J. H., CLarKke, L. K., Goopson, L. H., and ScHuytER, H. R., 1964.—Control of the snake
mite, Ophionyssus natricis (Gervais), in captive reptile collections. Zoologica, N.Y., 49 :
65-79.

Domrow, R., 1957.—Malaysian parasites. XXII. Four Trombicula nymphs (Acarina, Trom-
biculidae). Stud. Inst. med. Res. F.M.S., 28: 113-120.

, 1960a.—The genus Guntherana (Acarina, Trombiculidae). Pacif. Insects, 2: 195-237.
, 1960b.—Malaysian parasites. XL. Nymphs of Leptotrombidium and Trombiculindus
(Acarina, Trombiculidae). Stud. Inst. med. Res. F.M.S., 29: 163-171.
, 1961.—New and little-known Laelaptidae, Trombiculidae and Listrophoridae (Acarina)
from Australasian mammals. Proc. Linn. Soc. N.S.W., 86: 60-95.
, 1962a.—The genus Szseca in Australia (Acarina: Trombiculidae). J. ent. Soc. Qd,
1: 21-24.
—, 1962b.—Mammals of Innisfail. II. Their mite parasites. Avwst. J. Zool., 10: 268—306.
, 1966.—Some mite parasites of Australian birds. Proc. Linn. Soc. N.S.W., 90:
190-217.
, 1967a.—Rhinonyssine nasal mite infestations in birds at Mitchell River Mission during
the wet and dry seasons. Proc. Linn. Soc. N.S.W., 91: 211-219.
, 1967b.—Mite parasites of small mammals from scrub typhus foci in Australia. Aust.
J. Zool., 15: 759-798.
, 1972.—Acari Spinturnicidae from Australia and New Guinea. Acarologia, 13:
552-584.
, and Napcuatram, M., 1960.—Malaysian parasites. XLIII. Neoschongastia ir Malaya
(Acarina, Trombiculidae). Stud. Inst. med. Res. .M.S., 29: 185-193.

Fain, A., 1961.—Observations sur les acariens de la sous-famille Lawrencarinae (Ereynetidae :
Trombidiformes). Bull. Annls Soc. r. ent. Belg., 97: 245-255.

, 1968.—Etude de la variabilité de Sarcoptes scabiei avec une révision des Sarcoptidae.
Acta zool. path. antverp., 47: 3-196.

, and Ensen, P., 1967.—Les acariens de la famille Knemidokoptidae producteurs de gale
chez les oiseaux (Sarcoptiformes). Acta zool. path. antverp., 45: 3-142.

Gitt, D. A., Moutz, G. R., and Riex, R. F., 1945.—Trombidiosis of sheep in Queensland. Aust.
vet. J., 21: 22-31.

Hucues, A. Margaret, 1961.—The Mites of Stored Food. London: Her Majesty’s Stationery
Office.

Kaiser, M. N., and Hoocsrraat, H., 1973.—Bat ticks of the genus Argas (Ixodoidea : Argasidae).
9. A. (Carios) daviesi, new species, from Western Australia. Ann. ent. Soc. Am., 66: 423-428.

Leracu, J. A., 1958.—An Australian Bird Book. Melbourne: Whitcombe and Tombs.

Napcuatram, M., 1967.—Notes on the genus Torttrombicula (Sasa et al.) with designation of
neotype, description of nymph and redescription of larva of Tort. densipiliata (Walch),
and descriptions of two new species from southeast Asia (Acarina: Trombiculidae). J. med.
Ent., 4: 401-415.

, and Domrow, R., 1964.—The intranasal species of Laurentella (Acarina, Trombiculidae).
J. med. Ent., 1: 29-39.

Nave, S. A., 1939-66.—Nomenclator Zoologicus. Zoological Society of London.

Puiu, C. B., 1961.—Akamushi (Trombidium) Brumpt, 1910 (Class Acarina) : proposed validation
under the plenary powers. Z.N. (S.) 400. Bull. zool. Nomencl., 18: 140-142.

Ripe, W. D. L., 1970.—A Guide to the Native Mammals of Australia. Melbourne: Oxford
University Press. :

VERCAMMEN-GRANDJEAN, P. H., 1969.—Htablissement d’un lectotype pour Leptotrombidium
(Leptotrombidium) akamusht (Brumpt, 1910) (Acarina: Trombiculidae). Acarologia, 11 :
94-103.

Womers.Ley, H., 1952.—The scrub-typhus and serub-itch mites (Trombiculidae, Acarina) of the
Asiatic-Pacifie region. Rec. S. Aust. Mus., 10: 1-673.

WoreeE LL, E., 1963.—Reptiles of Australia. Sydney: Angus and Robertson.

CORRIGENDA
Acarologia, 13: 557. In couplet 1 of key, amend setal formulae to read 1—-3/2-0 and 1-3/I-0.

Proc. Linn. Soc. N.S.W., 88: 218. In line 6, read ‘*‘ eared’ for *“‘ earless’’. Proc. Linn. Soc.
N.S.W., 93: 314. In line 6, read “‘ textureless ”’ for *‘ textures”: -

PROCEEDINGS OF THE LINNEAN Society or New Souru WaLEs, VoL. 99, Part 1

THE RUGOSE CORAL GENERA STREPTELASMA HALL, GREWINGKIA
DYBOWSKI AND CALOSTYLIS LINDSTROM FROM THE LOWER
SILURIAN OF NEW SOUTH WALES

R. A. McLEAN*

(Plates I and m1)
[Accepted for publication 12th December, 1973]

Synopsis
Streptelasma recisum sp. nov. is described from the Late Lower or Early Middle Llandoverian
Bridge Creek Limestone of the Four Mile Creek area, central New South Wales. Grewingkia
parva sp. nov., G. neumani sp. nov. and Calostylis panuarensis sp. nov. are described from the
Brown Mudstone Horizon near Angullong, central New South Wales, this unit being probably
stratigraphically equivalent to the Bridge Creek Limestone. Possible affinities between the
genera Grewingkia Dybowski and Calostylis Lindstrom are discussed.

INTRODUCTION

The rugose corals described occur in horizons of the Panuara Group south-
west of Orange in central New South Wales. A summary of the stratigraphy
of these horizons (Bridge Creek limestone at Four Mile Creek and Brown
Mudstone Horizon at Angullong) together with evidence for their late Lower
or early Middle Llandoverian age are given by McLean (1974). Their distribution
is illustrated in Text-fig. 1.

This paper presents the first description of the genera Streptelasma Hall,
Grewingkia Dybowski and Calostylis Lindstrém from Australia. The species
‘* Streptelasma”’ australe (Foerste, 1888) from the Upper Silurian of the Yass
district, New South Wales, is not a representative of Streptelasma as that genus
is currently defined (see discussion below).

SYSTEMATIC PALAEONTOLOGY

The morphological terminology used is that employed by Hill (1956) and
Neuman (1969).

All numbers of specimens in the University of Sydney Palaeontological
Collections bear the prefix SUP. Where more than one section has been prepared
from the one specimen, the numbers bear the suffix a, b, ete.

Text-figures have been prepared from tracings of photographs.

Family STREPTELASMATIDAE Nicholson
in Nicholson and Lydekker, 1889
Diagnosis (based on Hill, 1956, p. F268). Solitary or fasciculate coralla
generally with short minor septa and with septal stereozone often developed.
Dissepiments lacking and tabulae usually domed, complete and incomplete.

Subfamily STREPTELASMATINAE Nicholson
in Nicholson and Lydekker, 1889
Diagnosis. Septal stereozone may or may not be developed. Septa
lamellar, complete throughout ontogeny.
Discussion. The generic composition of this subfamily is considered here
as that given by Hill (1956, pp. F268-F269), although Coelostylis Lindstrém

ie Department of Geology and Geophysics, University of Sydney, New South Wales, 2006.

PROCEEDINGS OF THE LINNEAN Society or New SoutH Waxzs, Vou. 99, Part 1

R. A. MCLEAN 37

has been shown to possess monacanthine septa and has been transferred to the
family Tryplasmatidae Etheridge (Neuman, 1967). Genera subsequently
described and considered here as representatives of the Streptelasmatinae include
Porfirieviella Ivanovskiy, 1963 ; Helicelasma Neuman, 1969 ; Borelasma Neuman,

5
Bridge Creek_e
Limestone

&
&
CG

PANUARA

Mandurama

Horizon

\ Pd 1
ANGULLONG

Text-fig. 1. Distribution of the Bridge Creek Limestone and Brown Mudstone Horizon,
South-east of Orange, New South Wales.

1969 ; Crassilasma Ivanovskiy, 1962; and Kenophyllum Dybowski, 1873. The
relationships of these latter genera to the type genus Streptclasma Hall, 1847,
are discussed below.

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38 LOWER SILURIAN RUGOSA FROM NEW SOUTH WALES

Genus STREPTELASMA Hall, 1847

1847 Streptelasma Hall, p. 69

1873 Calophyllum Dybowski, p. 75

1873 Streptelasma ; Dybowski, p. 384

21901 Zaphrentis; Lambe, p. 118 (non Rafinesque and Clifford, 1820)
21901 Zaphrentis ; Lambe, p. 119
1927 Dybowskia Wedekind, p. 17
21928 Zaphrentis; (part.) Twenhofel, p. 114
21928 Streptelasma; Troedsson, p. 110

1933 Dybowskia ; Scheffen, p. 7

1937 Streptelasma; (part.) Cox, p. 2

1940 Brachyelasma Lang, Smith and Thomas, p. 28

1955 Brachyelasma; Nikolaeva, p. 22

1956 Brachyelasma; Kaljo, p. 71

1958 Brachyelasma; Kaljo, p. 102

1958 Brachyelasma; Reiman, p. 36

21960 Brachyelasma; Pestana, p. 868
1963 Brachyelasma ; Ivanovskiy, p. 42
?1965a Brachyelasma ; Ivanovskiy, p. 104
1969 Streptelasma; Neuman, p. 8
1971 Streptelasma; Scrutton, p. 207
21971 Brachyelasma; Lavrusevich, p. 51
non 1970 Streptelasma; Fligel and Saleh, p. 287

Type species. S. corniculum Hall, 1847, Trenton Limestone, New York
(2? Middle Caradocian).

Diagnosis (based on Neuman, 1969, p. 8). Solitary streptelasmatid coral
with cylindrical, ceratoid or trochoid corallum with a convex cardinal side.
Major septa in brephic and neanic stages long, thin or moderately dilated, normally
fused into a weak axial structure. Major septa in ephebic stage thin, com-
paratively short, normally not forming axial structure. Tabulae of complete,
convex type, often provided with complementary plates.

Discussion. Until recently, knowledge of the internal structure of the genus
Streptelasma Hall, 1847 (type species S. corniculum), had been based on Canadian
material described by Lambe (1901). Subsequent workers have used these
descriptions as a basis for ascribing other specimens to the genus. Lambe’s
material is characterized by early growth stages having thickened major septa
extending to the corallite axis. However, Hall’s original type material had
never been studied in thin section until the work of Neuman (1969), who found
that the internal structure showed thin septa extending to the axis in the early
stages, and that later the thin major septa became withdrawn from the axis.
It thus became necessary to redefine Streptelasma as it was understood at that
stage, and since the genus Brachyelasma Lang, Smith and Thomas, 1940, shows
the same septal arrangement as Hall’s syntypes of S. corniculum revealed on
sectioning, Brachyclasma was included by Neuman as a junior subjective synonym
of Streptelasma.

Neuman (1969) proposed the new genus Helicelasma (type species H. simplex
Neuman, 1969) for forms included in Streptelasma under its old interpretation.
Species now ascribed to the genus Streptelasma are generally those interpreted
by earlier authors as belonging to Brachyelasma. Included in the family
Streptelasmatidae are a number of genera that are obviously very closely related
to Streptelasma. Distinctions between these genera can normally only be made
by study of the character of their early ontogenetic stages. Such studies have
generally been done only by recent workers (e.g. Kaljo, 1956, 1958; Reiman,
1958; Ivanovskiy, 1963; and Neuman, 1969). Hence, of the many streptel-
asmatid species described in the past, only a relatively small proportion may be

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R. A. MCLEAN 39

ascribed to a particular streptelasmatid genus with certainty. The genera
showing closest similarities to Streptelasma are listed in Table 1, together with
their distinguishing features.

TABLE |
Distinguishing Features of Streptelasma and Related Genera

_ Genus Neanic Stage Ephebic Stage
Streptelasma Hall, 1847... Septa show little dilation ; Major septa, thin, withdrawn
axial structure may be from axis
present
Porfirieviella Ivanovskiy, 1963 As in Streptelasma Major septa thin, reaching

axis and may form weak
axial structure

Dinophyllum Lindstrém, 1882 Septa moderately dilated Septa may show moderate
dilation, reach axis and
typically form vortex

Helicelasma Neuman, 1969 Septa strongly dilated, usually As in Porfirieviella

in contact ; noaxialstructure
Borelasma Neuman, 1969 .. As in Helicelasma As in Streptelasma
Crassilasma Ivanovskiy, 1962 As in Helicelasma Septa strongly dilated, reach-

ing or almost reaching axis

Kenophyllum Dybowski, 1873 Septal structure is apparently similar to that of Crassilasma
but tabulae are absent at all stages.

The relationships of Streptelasma, Porfirieviella and Dinophyllum are
uncertain. As mentioned above, Streptelasma typically shows septa withdrawn
from the axis in the ephebic stage. However, in some species, the septa are only
slightly withdrawn (e.g. S. estonicum Dybowski as described by Kaljo, 1958,
and Neuman, 1969). Hence such forms are not widely different from those
having septa extending to the axis as in Porfirieviella, e.g. P. stokesi (Kdwards
and Haime) as described by Ivanovskiy (1963). The major features distin-
guishing Dinophyllum from Porfirieviella appear to be formation in Dinophyllum
of an axial vortex by the major septa and the presence of weak dilation of the
septa. Ivanovskiy (1970a, p. 121) has in fact suggested that Porfirieviella
differs only from Dinophyllum in lacking an axial vortex and that the genera
should be considered synonymous. This feature is again demonstrated to be
gradational in Streptelasma by Neuman (1969), the formation of an axial vortex
being limited of course by the fact that the septa do not reach the axis in that
genus. However, twisting of the axial ends of the major septa is exhibited by
some species of Streptelasma, e.g. S. primum Wedekind and S. linnarssoni
(Lindstr6m) as described by Neuman (1969). The early ontogenetic stages of
Dinophyllum (as in D. involutum Lindstrém, described by Minato, 1961) show
long major septa, weakly dilated and similar to some species of Streptelasma
(e.g. S. corniculum Hall; see Neuman, 1969). Future study of a larger variety
of material may reveal that differences between these two genera and Streptelasma
are merely gradational, in which case they should all be considered synonymous.

In a restudy of the composition of the genus Streptelasma, it is evident
that many species, although similar to Streptelasma in their ephebic stage, are
not well enough known in their earlier growth levels to be classified in that genus
with certainty. !

Brachyelasma fervida Kaljo, 1958, from the Oandu Horizon (Caradocian)
of Estonia, possesses long, thin major septa in both ephebie and ? neanic stages

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40 LOWER SILURIAN RUGOSA FROM NEW SOUTH WALES

(Kaljo, 1958, Pl. I, figs 5, 6) but in the ephebic stage some septa reach the axis
and there appears to be a weak axial structure of septal lobes and lamellae rather
reminiscent of the genus Grewingkia Dybowski. Kaljo (1958, p. 104) also stated
that major septa reach the axis. Ivanovskiy (1965b) listed B. fervida as a species
of Porfirieviella, but Neuman (1969) included it in his revised genus Streptelasma
(no mention was made by Neuman of Porfirieviella). The incomplete nature of
Kaljo’s material makes it difficult to classify the species with certainty. Similarly,
B. oanduensis Kaljo, 1956, from the Keyla Horizon (Caradocian) of Estonia has
long, thin major septa in the ? ephebic stage (Kaljo, 1956, Pl. I, fig. 10), but was
included in his revised Streptelasma by Neuman (1969). It too was included in
Porfirieviella by Ivanovskiy (1965b) but lacks sufficient illustration of its growth
stages for definite classification. Ivanovskiy (1963) described the new Siberian
Platform species Brachyelasma siluriense and B. fossulatum from the Upper
Llandoverian, and B. nikiforovae from the Upper Caradocian. While all possess
short, thin major septa in the ephebic stage, no figures were given of the earlier
stages and their inclusion in Streptelasma cannot be certain. Ivanovskiy (1970b)
later included all three species in the Upper Llandoverian B. sibiricum Nikolaeva
from the Siberian Platform, but the synonymy of B. siluriense at least appears
doubtful. More material, particularly of the younger stages, is necessary before
a definite classification of these forms is possible. Similarly, Brachyelasma
concavifundatum Ivanovskiy, 1965a, from the Upper Llandoverian of the Siberian
Platform, lacks illustration of its early growth stages. However, it has thin,
short major septa in the ephebic stage. Ivanovskiy (1970b) listed it as a sub-
species of B. sibiricum.

Two new species of Brachyelasma were described from Tadzhikistan by
Lavrusevich (1971). B. agbaschiricum from Horizon A (Upper Ordovician)
has not been studied in its early growth stages. It has strongly dilated, short
major septa in the ephebic stage, however, and may be a representative of
Streptelasma, although the septa are thicker than is typical of that genus.
B. digitiforme from Horizon H (Upper Llandoverian) also has been described only
from ephebic sections, but these appear typical of Streptelasma, and the species
when fully studied should probably be included in that genus.

Although no North American representatives of Streptelasma, as now defined,
have been described (except the type species), several species show some
similarities to that genus. Zaphrentis affinis Billings, 1865 (Lambe, p. 118,
Pl. VU, fig. 6) from the Ellis Bay Formation (? Lower Llandoverian) of Anticosti
Island shows thin major septa withdrawn from the axis in the ? ephebic stage,
but earlier ontogenetic stages have not been described. Also Zaphrentis patens
Billings, 1865, from the Jupiter Formation (Upper Llandoverian) of Anticosti
Island (Lambe, 1901, p. 119, Pl. VIII, fig. 2) has thin, short major septa at the
calice, but early ontogenetic stages are unknown. Zaphrentis anticostiensis
Twenhofel, 1928, from the Gun River-Jupiter Formations (Middle-Upper
Llandoverian) of Anticosti Island has thin, major septa somewhat withdrawn
from the axis at the calice but extending to the axis below this level (Twenhofel,
1928, p. 114, Pl. I, figs 6-8). Unfortunately no sections of early growth stages
have been figured, but it is likely that the species belongs to Streptelasma.
3rachyelasma bassleri Pestana, 1960, from the Johnson Spring Formation
(? Trenton) of California has thin major and minor septa strongly withdrawn
from the axis in ephebic sections, but the early ontogenetic stages are not described
and the species cannot be assigned to Streptelasma with certainty.

Two species from the? Llandoverian of Iran recently described by Fligel and
Saleh (1970), Streptelasma ruttneri and S. shirgeshtensis, have sufficiently well-
described early growth stages to permit classification. S. ruttnerr has long,
thin, major septa extending to the axis in ephebic sections, while the septa are
strongly dilated and also reach the axis in the neanic stage. It is probably a

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R. A. MCLEAN Al

representative of Helicelasma Neuman, 1969. S. shirgeshtensis on the other
hand has short, thin septa in the ephebic stage, while in the neanic stage they are
long and moderately dilated. This species should probably be included in the
genus Borelasma Neuman, 1969.

The only form from Australia previously described as belonging to
Streptelasma is ‘*‘ 8.” australe (Foerste, 1888), represented in the *‘ Phacops bed ”’
of the Yass district, New South Wales (Hill, 1940, p. 410). Link and Druce
(1972, p. 7) listed ‘* Streptelasma ”’ australe from the Rainbow Hill Marl Member
(Link, 1970) at Yass, apparently the horizon referred to by Hill. The age of this
unit is regarded as late Ludlovian (Link, 1970; Link and Druce, 1972). From
the descriptions of Hill (1940) it is apparent that ‘‘ S.” australe possesses dilated
septa in its early ontogenetic stages, together with some development of an
axial structure. Hence it is not a representative of Streptelasma as defined by
Neuman and its systematic position needs to be resolved.

Range. Middle Caradocian of Estonia, New York, ? California; Upper
Caradocian of Estonia, ? Siberian Platform; Ashgillian of Sweden, Norway,
Estonia, Urals, ? Tadzhikistan, ? Anticosti Island, Greenland; Lower
Llandoverian of Estonia, Venezuela and N.S.W.; Upper Llandoverian of the
Siberian Platform, ? Tadzhikistan, ? Kazakhstan, ? north-east U-.S.S.R.,
? Anticosti Island.

Streptelasma recisum sp. nov.
Plate 1, figs 1-6
Derivation of name. Latin recisus=reduced, referring to the nature of the
septa in the ephebic stage.

Material. Holotype SUP 45167. Paratypes SUP 45168-45175. Preserved
in a colony of the tabulate coral, Priscosolenia sp., Bridge Creek Limestone.
Late Lower or Early Middle Llandoverian.

Diagnosis. Small Streptelasma with average calice diameter 8-10 mim.
Septa in neanic stage thin, short (reaching up to 0-5 corallite radius) and few
in number (average 26-30). Minor septa very weakly developed ; stereozone
absent. Tabulae horizontal or sagging axially.

Description. Corallum trochoid-ceratoid, weakly curved. Specimens
incomplete but dimensions are of the order of at least 25mm in height and
maximum observed diameter of approximately 15mm. Average corallite
diameter 8-10mm towards base of calice. Epitheca thin and _ peripheral
stereozone lacking at all stages of growth.

In neanic stage (Plate I, figs 3, 6) septa are thin and extend almost to corallite
axis. No axial structure formed but a few septal lamellae may be present in
axial region. ? Cardinal septum shorter than other septa in neanic stage (Plate I,
fig. 6), but owing to poorer preservation this cannot be confirmed in later growth
levels. In neanic stage, at diameters of 5mm and 7:5 mm, septa number 24
and ? 25 respectively (SUP 45172, 45171). In the ephebic stage, septa are
withdrawn from axis, with a common length of about 0-5 of radius of corallite
(Plate 1, figs 1, 5). In ephebic sections available, at corallite diameters of
7:5-9-5 mm, septa number 26-30. Minor septa are difficult to distinguish and
if present occur only as very fine ridges on corallite wall. High in calice, septa
are reduced greatly in length (approximately 0-2 total corallite radius).

Tabulae mainly complete, moderately downflexed peripherally, horizontal
or sagging axially, particularly in late stages of growth. Average spacing of
tabulae 0-4—0-6 mm in small specimen (SUP 45175) and 0-5-1-2 mm in large
form (SUP 45174). Complementary plates lacking.

Remarks. An interesting feature in regard to the growth form of S. reciswm
sp. nov. is its relationship to the massive tabulate coral Priscosolenia in which
the described material grew commensally. Small flexures in the rugosan

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42 LOWER SILURIAN RUGOSA FROM NEW SOUTH WALES

epitheca are common, indicating mutual adjustment of both corals during growth.
Invaginations of the epitheca of S. recisum into the Priscosolenia colony are
evident, possibly for support of the rugosan form (Plate 1, fig. 3, particularly).
The corallites of the Priscosolenia colony have also been considerably disturbed
and distorted by the presence of the Streptelasma (Plate 1, fig. 4).

The only described species of Streptelasma to show any strong similarities
to S. recisum sp. nov. is 8S. sibiricum (Nikolaeva, 1955) from the Upper
Llandoverian of the Siberian Platform. It resembles the New South Wales
form in having comparable corallite dimensions, thin septa almost reaching
the axis in the neanic stage and major septa of similar number (up to 35) and
length (up to 0-5 of corallite radius) in the ephebic stage. It is also similar in
having the minor septa lacking or very short in the ephebic stage, tabulae of
similar shape and little or no peripheral stereozone. However, a clear distinction
between the two forms may be made, since S. sibiricwm has rather more dilated,
thin, wedge-shaped septa in the ephebic stage and possesses a very prominent
cardinal fossula with very short cardinal septum in the late neanic or carly ephebic
stage (Ivanovskiy, 1963, Pl. IX, fig. iB). No definite fossula has been observed
in any of the Bridge Creek material. There is evidence of very short minor
septa in the material of Nikolaeva (1955, Pl. XLVIII, fig. 2a).

The form described by Twenhofel (1928) as Zaphrentis anticostiensis from
the ? Middle-Upper Llandoverian of Anticosti Island shows quite strong
similarities to S. recisum, but lacks illustration and detail of the early ontogenetic
stages and so cannot be closely compared with it. Similarities include septal
number (23 at corallite diameter of 8 mm), the minor septa reduced to ‘‘ spines
or nodes’, very thin major septa in the ephebic stage extending half or more
than half-way to the axis, and the shape of the tabulae. Twenhofel stated
(p. 114) that the major septa reach the axis below the level of the calice. A
cardinal fossula with a short cardinal septum on the calice floor is also reported,
although this is not illustrated by Twenhofel.

Genus GREWINGKIA Dybowski, 1873
1873 Grewingkia Dybowski, p. 384
1901 Streptelasma; Lambe, p. 109 (non Hall, 1847)
1927 Kiaerophyllum Wedekind, p. 17
1933 Kiaerophyllum ; Scheffen, p. 16
1937 Streptelasma; (part.) Cox, p. 10
1948 Streptelasma (Kiaerophyllum) ; Wang, p. 102
1950 Streptelasma (Kiaerophyllum) ; Wang, p. 213
1956 Grewingkia; Hill, p. F268
1956 Grewingkia; Dunean, Pl. 21, figs 4a, b
1958 Grewingkia; Reiman, p. 34
1958 WStreptelasma (Kiaerophyllum) ; Kaljo, p. 25
1960 Brachyelasma; Tcherepnina, p. 387 (non Lang, Smith and Thomas,
1940)
1969 Grewingkia; Pestana, p. 868
1961 Streptelasma (Grewingkia) ; Kaljo, p. 62
1961 Rectigrewingkia Kaljo, p. 62
1961 Cyatholasma Ivanovskiy, p. 120
1963 Grewingkia; Nelson, p. 338
1965a Cyatholasma; Ivanovskiy, p. 76
1965a Grewingkia ; Ivanovskiy, p. 77
1965 Grewingkia; Kaljo and Klaaman, p. 420
1969 Grewingkia; Neuman, p. 33
1970a Grewingkia ; Ivanovskiy, p. 121
1970 Grewingkia; Fligel and Saleh, p. 291
1971 Grewingkia; Wavrusevich, p. 49

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R. A. MCLEAN 43

Type species. Olisiophyllum buceros Bichwald, 1856, Upper Ordovician,
Estonia. (Type horizon unknown.)

Diagnosis (abbreviated from Neuman, 1969, p. 33). Solitary streptelasmatid
coral with cylindrical, ceratoid or trochoid corallum. Septa moderately or
heavily dilated early in ontogeny, major septa long, forming narrow axial
structure. Later in ontogeny major septa relatively short and thin, axial
structure broad, composed of numerous irregularly intertwined septal lobes and
lamellae. Calicular boss present or absent. Tabulae few or numerous, complete
or incomplete, with or without complementary plates.

Discussion. The genus Grewingkia has been interpreted in many different
ways by previous workers. Representatives described as belonging to Kiaero-
phyllum (Wedekind, 1927 ; Scheffen, 1933) were considered as being a subgenus
of Streptelasma Hall by Wang (1948). Hill (1956), however, recognized the
Synonymy of Kiaerophyllum and Grewingkia and considered the latter to be
distinet from Streptelasma. Kaljo (1958) applied the classification adopted by
Wang but later (1961) recognized the synonymy suggested by Hill. He still,
however, considered Grewingkia to be a subgenus of Streptelasma. Kaljo (1961)
also erected a new genus, Rectigrewingkia (see discussion below), to include the
- species G. anthelion Dybowski and G. lutkevitshi Reiman (possibly with G. eminens
Hichwald and G. formosa Dybowski, both needing revision). Neuman (1969)
in a thorough revision of Grewingkia considered differences between the two genera
to be gradational and restored Rectigrewingkia to the genus Grewingkia. Further-
more, Neuman revived the classification of Hill (1956) and gave Grewingkia
the status of a full genus. This classification has been adopted by the author of
this paper.

Densigrewingkia Neuman, 1969, and Lobocorallium Nelson, 1963, are similar
to Grewingkia in possessing a reticulate axial structure. Densigrewingkia may
be distinguished by its concave cardinal side and the considerable amounts of
sclerenchyme in its axial structure, particularly in the early ontogenetic stages.
Lobocorallium differs in having a prominent cardinal fossula and a tri-lobed
corallum with the lobes corresponding to the cardinal and alar septa. As men-
tioned above, Rectigrewingkia Kaljo, 1961, is considered by Neuman (1969),
and in this paper, to be a synonym of Grewingkia. The major distinguishing
feature of Rectigrewingkia, according to Kaljo, is the presence of rounded septal
lamellae in the axial structure, as compared with intertwined lobes and lamellae
in species of Grewingkia. This appears to be only a gradational feature and is
not considered important enough for formation of a separate genus. Further
detail on the distinction of these genera may be obtained in Neuman (1969).

The genera Bighornia Dunean, Bodophyllum Neuman, Dalmanophyllum
Lang and Smith and Ditoecholasma Simpson apparently all possess a solid axial
structure of fused septal lobes and lamellae, together with complete, lamellar
Septa peripherally. Further study of these forms is required ; they may possibly
be synonymous (Ivanovskiy, 1965a; and Neuman, 1969). Their solid axial
structure serves to distinguish them from Grewingkia in any case.

The species G. dentiseptata Lavrusevich, 1971, from Horizon B of the
Zeravshan-Gissar region of Tadzhikistan was considered as Lower Llandoverian
by Lavrusevich (1971). However, Horizon B is correlated with the Porkuni
Horizon of Estonia (Lavrusevich and Menakova, 1971), and the latter is generally

considered as Upper Ashgillian (Mannil, 1966; Roomusoks, 1970). Ivanovskiy
(1965b) also considered the fauna of Horizon B to be more typical of the Upper
Ordovician. G. dentiseptata shows no close similarities to the New South Wales
representatives of the genus.

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44 LOWER SILURIAN RUGOSA FROM NEW SOUTH WALES

kange. Lower Caradocian of ? Scotland ; ? Middle Caradocian of California ;
Upper Caradocian of Estonia, south-west Siberia, Manitoba; Ashgillian of
Ireland, Norway, Sweden, Estonia, Tadzhikistan, Quebec, Michigan; Llando-
verian of north-east Iran; Lower or Middle Llandoverian of New South Wales.

Grewingkia parva sp. nov.
Plate 1, figs 7-10; Text-fig. 2a, b
Derivation of name. Latin parvus =small.

Material. Holotype SUP 45151. Paratypes SUP 45152-45155, 63274.
Brown Mudstone Horizon, Angullong district. Late Lower or Early Middle
Llandoverian.

SS We

DY
mee

Text-fig. 2. Grewinykia parva sp. nov. a. SUP 4515la, holotype, transverse section, x 4.
5. SUP 45152, paratype, longitudinal section, x 4.

Diagnosis. Small, subecylindrical Grewingkia with a prominent calicular
boss and narrow peripheral stereozone. Corallite diameter up to 14mm and
maximum septal number ranges from 60 to 70. Minor septa long, up to half
length of major septa, the latter typically showing dilated lobes at their axial
extremities. Axial structure compact, approximately half corallite diameter.
Tabulae mainly incomplete, weakly concave peripherally and strongly arched
through axial zone.

Description. Subeylindrical corallum gradually tapering proximally
although proximal end not preserved in any specimens. Epitheca thin, although
all specimens have most of epitheca lacking. Average corallite diameter
11-14 mm with a height of at least 30mm. Calice rather shallow with steep
sides comprising stereozone and epitheca only (Plate 1, fig. 9). Depth of calice
4mm in preserved corallite height of 23mm (total height probably at least
30-35 mm) in only specimen showing this feature (SUP 45152). Calicular boss

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R. A. MCLEAN 45

prominent, with a height of 3mm and diameter approximately 6mm (SUP
45152). Calice floor flat around axial boss.

Peripheral stereozone narrow, with a width of 1-1-5 mm in ephebice sections,
consisting of dilated septa and indeterminate ? sclerenchyme tissue. Septa
of two orders, thin, with average number 64-70 (possibly 78 in SUP 45154),
typically showing small, rounded projections irregularly developed on sides of
septa. Major septa also show larger dilated lobes at their axial extremities.
Minor septa extend well beyond peripheral stereozone, reaching a length of
approximately half that cf major septa. Axial structure has mainly consistent
diameter throughout visible ontogeny (approximately the distal 23 mm) and is
about half the diameter of corallite. Structure composed of tightly anastomosing
septal lobes and lamellae, generally of smaller diameter than septa themselves.

Tabulae mainly complete with generally constant spacing of about 0-7 mm.
Individual tabulae strongly arched in axial reticulate zone, conforming in shape
with profile of calicular boss. In peripheral area tabulae are weakly sagging
and may be faintly traced curving distally in peripheral stereozone. Comple-
mentary plates may be present in peripheral regions.

Remarks. Owing to lack of sufficient well-preserved material, the early
ontogenetic stages of this species could not be studied. However, enough
material was available to allow some comparisons with other representatives of
the genus. None of the described species of Grewingkia are closely similar to
G. parva sp. nov. However, some relationships with G. anguinea (Scheffen,
1933) from Division 5A (Lower Ashgillian) of Norway as described by Neuman
(1969) may be seen in its corallite size, septal number and development of long
minor septa, narrow stereozone, weak dilation of septa, convex tabulae with
slightly concave peripheral portions and an axial structure of comparable
dimensions. It clearly does not have the ceratoid growth form of G. anguinea,
although this growth form only appears obvious in the more proximal regions, which
have not been found preserved in any of the Angullong specimens. G. anguinea
also differs in having a deeper calice with far less steeply inclined walls, a low,
broad calicular boss, as a result of less convex tabulae, no broadly dilated septal
lobes at the axial extremities of the major septa and a less dense axial complex
of septal lobes and lamellae.

G. contexta Neuman, 1969, from the Boda Limestone (Ashgillian) of Sweden,
also Shows some similarities to the local form in having similar corallite dimensions,
subcylindrical growth form (but ceratoid also), and a prominent calicular boss.
Like G. parva, it also shows a narrow stereozone, generally long minor septa late
in ontogeny and comparable size and appearance of the axial structure. However,
it too may be clearly distinguished by showing a greater number of septa (90-100
average), lack of large lobes on the axial septal extremities and strong septal
dilation in the cardinal quadrants until quite late in ontogeny, although it is
possible those early stages have not been preserved in the Angullong specimens.
Other differences include a less steep-walled calice, different shape of the tabulae
(Neuman, 1969, p. 47, fig. 37J) and a fixing groove and rootlets on the convex
side of the corallite.

The only other Llandoverian species described (G. alternata Saleh in Fligel
and Saleh, 1970, from the Niur Formation of Iran) differs from G. parva in having
much larger size (ccrallite diameter 30 mm), proportionately fewer septa (n/De=5
for G. parva and 3-5 for G. alternata), less tightly anastomosing elements in the
axial structure and a lack of lobes on the major septa.

The specimen sectioned at the base of the calice (SUP 45154, Plate I, fig. 10),
while tentatively included in G. parva, shows a greater number of septa (78),
longer minor septa and very pronounced dilated lobes on the axial extremities
of the major septa.

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46 LOWER SILURIAN RUGOSA FROM NEW SOUTH WALES

Grewingkia neumani sp. nov.
Plate 1, figs 11, 12; Plate u, fig. 1; Text-fig. 3

Derivation of name. After Dr. B. Neuman, for his important studies of
streptelasmatid corals.

Material. Holotype SUP 20115. Paratypes SUP 45156, 45157. Coarse
calearenite lens in Brown Mudstone Horizon, Angullong district. Late Lower or
Early Middle Llandoverian.

Diagnosis. Trochoid or ? subeylindrical Grewingkia characterized by very
deep calice, wrinkled epitheca and very broad peripheral stereozone. Corallite
diameter ranges up to at least 35mm and septal number ranges up to 96 in
subeylindrical forms. Minor septa short, major septa typically showing club-like
dilated lobes at axial extremities. Axial structure 0-3-0-5 corallite diameter,
septal lamellae and lobes coarse in trochoid forms, more tightly intertwined
and finer in smaller representatives. Tabulae slightly sagging peripherally,
weakly convex axially.

Description. Corallum trochoid in two specimens while the other shows
subeylindrical growth form. Epitheca smooth with transverse wrinkles evident
in SUP 45157. Calice very deep and in largest specimen (holotype SUF 20115,
with height of at least 30 mm) calice has approximate depth of 15mm. Corallite
diameter varies from about 50 mm at top of calice to about 30 mm near calice
base in largest specimen. Subcylindrical specimen reaches diameter of 14 mm.
Calical boss low and wide.

Text-fig. 3. Grewingkia neumani sp. nov. SUP 45156a, paratype, transverse section, x 4.

3road peripheral stereozone of dilated septa and ? sclerenchyme. In largest
specimen stereozone thins progressively from about 8 mm in width at base of
calice to approximately 2 mm near its top. Maximum of about 60 septa in
subeylindrical form, ranging to 94-96 in large trochoid specimen. Septa dilated
so as to be in contact in stereozone where trabecular structure of septa is shown.
Trabeculae, although poorly preserved, exhibit coarse fibres radiating obliquely
from central axis (Plate u, fig. 1). Beyond stereozone major septa thin markedly
to about 0-3 of their peripheral width and internal structure is obscured by
recrystallization. Small ‘“ nodes” and undulations are evident on septa in this
region in transverse section (Text-fig. 3). Minor septa barely extend beyond
peripheral stereozone, while in subcylindrical form they extend approximately
0-3 of the distance between stereozone and axial structure. Width of axial
structure ranges from 0-3 to 0-5 corallite diameter and it is composed of loosely
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R. A. MCLEAN 47

intertwined septal lamellae and lobes. In subcylindrical form septal elements in
axial structure are finer and more tightly reticulate. Tabulae sagging adjacent
to peripheral stereozone, weakly arched in axial region. Tabular spacing approxi-
mately 1 mm.

Remarks. The only described species of Grewingkia to show any strong
similarities to G. newmani is the Upper Caradocian G. altaica (Tcherepnina,
1960) from the Altai and Salair areas of south-west Siberia. Features in common
inelude similar corallite dimensions and growth form (for the large, trochoid
representatives), coarse, intertwined septal lamellae of the axial structure, and a
broad stereozone consisting mainly of dilated septa. In particular, the transverse
section illustrated by Ivanovskiy (1961, Pl. XV ; 1965a, Pl. X XVI, fig. 3) appears
closely comparable. However, differences may be seen in the rather more
irregular and wavy nature of the major septa in the region between the stereozone
and axial structure of G. altaica although this is not as apparent in the Altai
material of Tcherepnina (1960, Pl. O-X, fig. 3b). Also in G. altaica the tabulae
are more strongly downflexed near the periphery and there is no development
of club-shaped dilated lobes on the axial ends of the major septa such as is found
in G. newman.

The smaller, subcylindrical specimen assigned to G. newmani, while differing
in possessing longer minor septa, rather wavy major septa and a more densely
compacted axial structure, is included in the species because of its similarly
constructed broad stereozone. If further material is found that confirms there
is a Sharp distinction between these two types, then it may be advisable to consider
the latter form perhaps as a new species or aS a subspecies of G. neumani. For
the present, however, they are grouped together.

G. neumani may be distinguished from G. parva sp. nov. primarily by the
broad peripheral stereozone, shorter minor septa and deeper calice of the former.
A feature in common, however, is the development of the dilated axial lobes on
the major septa. This characteristic is apparently unique to the local repre-
sentatives of Grewingkia. However, some Silurian species of the genus Calostylis
show this feature, Smith (1930a, p. 261) recording that ‘“‘ individual strands of
this peripheral and axial tissue. . . dilate so as to form numerous nodes, and the
numerous free endings are nearly always swollen and rounded’’. Possible
affinities between Grewingkia and Calostylis are discussed further below.

The most useful descriptions of the septal microstructure of Grewingkia
have been those of Kato (1963), Ivanovskiy (1967) and Wang (1950). Kato
(1963, Text-fig. 10. 3) illustrated fibres diverging obliquely from a central axis
in a septum of ‘‘ Grewingkia sp... Ivanovskiy (1967, Fig. 3d) showed a similar
structure in G. altaica. The septal microstructure of the New South Wales
representatives of Grewingkia is very poorly preserved and only the holotype of
G. neumant (SUP 20115) shows this feature in any detail. As can be seen from
Plate , fig. 1, a rather coarse system of fibres diverging from a central axis
occurs in the dilated septa of the peripheral stereozone. Beyond the stereozone
recrystallization has obscured the microstructure but a similar structure on a
smaller scale would be expected, according to Kato and Ivanovskiy. The septal
microstructure illustrated by Ivanovskiy (1967, Fig. 3d) for G. altaica is quite
closely comparable to that of G. neumant.

Family CALOSTYLIDAE Roemer, 1883

Diagnosis (modified from Hill, 1956, p. F296). Solitary and colonial corals
with major and minor septa perforate ; axial ends of major septa lobed and
reticulate, forming spongy axial structure. Tabulae generally distally arched in
axial region; dissepiments lacking. Epitheca typically not developed over
entire corallum.

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48 LOWER SILURIAN RUGOSA FROM NEW SOUTH WALES

Discussion. The family Calostylidae is taken here to include the genera
Calostylis Lindstrém, Palaearaea Lindstré6m and Helminthidiwm Lindsti6m, all
characterized by possession of perforate septa. The taxonomic affinities of this
group of corals, however, are open to question, as mentioned by Hill (1956),
although they are generally recognized as being probably closest to the strep-
telasmatids.

Ivanovskiy (1961), in his discussion of the family Calostylidae, deseribed a
new genus, Cyatholasma, from the Upper Caradocian of the Salair regien of
south-west Siberia. Cyatholasma was characterized by complete, lamellar septa
in the peripheral region of the corallite and a reticulate structure of septal lobes
and lamellae at the axis. The synonymy of this form with Grewingkia Dybowski
was recognized by Ivanovskiy (1967, 1970a). However, the genus Calostylis
possesses perforate septa peripherally but an axial structure similar to that of
‘* Cyatholasma’”’. Ivanovskiy (1961) thus inferred that ‘‘ Cyatholasma”’ repre-
sented a link between the Middle-Upper Ordovician Streptelasmatidae which
have complete, lamellar septa throughout, and the Upper Ordovician-Silurian
Calostylidae which have perforate, incomplete septa. He subsequently (1965a,
1968) included Grewingkia in the family Calostylidae, although the genus has
been generally placed in the Streptelasmatidae by most workers (e.g. Dybowski,
1873 ; Hill, 1956; Neuman, 1969). As representatives of the Streptelasmatidae
(e.g. Streptelasma Hall, see above and Neuman, 1969) may often show traces
of septal lobes and lamellae in the axial zone at some stage of their ontogeny,
it seems preferable to include Grewingkia in that family. However, the evident
close relations of Calostylis to Grewingkia would suggest that the calostylids
are most probably an aberrant branch of the Streptelasmatidae. It may in
fact be best to consider Grewingkia and Calostylis, together with related forms, as
representatives of a distinct subfamily of the Streptelasmatidae, characterized
by forms with well-developed axial reticulate structure. Further work on these
and similar forms is required before a definitive statement on the systematic
position of the calostylids is possible. Particularly important in this regard
would be ontogenetic studies of the species of Calostylis and Grewingkia. Hence
for the present the family Calostylidae is retained for Calostylis and other perforate
genera.

Genus CALOSTYLIS Lindstrém, 1868

1865 ? Clisiophyllum Kjerulf, p. 22 (not seen)
1868 Calostylis Lindstrém, p. 421

1878 Calostylis; Nicholson and Etheridge, p. 65
1906 Calostylis ; Foerste, p. 322

1912 Calostylis; Reed. p. 123

1917 Calostylis ; Foerste, p. 200

1930a Calostylis ; Smith, p. 257

1930b Calostylis ; Smith, p. 294

1958 Calostylis; Kaljo and Reiman, p. 27
1961 Calostylis; Ivanovskiy, p. 119

1963 Calostylis ; (? part.) Ivanovskiy, p. 92
1965 ? Calostylis ; Stumm, p. 47

1966 Calostylis ; Sytova and Ulitina, p. 243
1971 Calostylis ; Lavrusevich, p. 69

Type species. C. cribaria Lindstrom, 1868? Clisiophyllum denticulatum
Kjerulf, 1865. Wenlockian, Gotland.

Diagnosis. Corallum solitary or often witb lateral buds ; having perforate
septa which degenerate axially and peripherally into a reticulate structure.
Tabulae complete and incomplete, distally arched; dissepiments absent.
Epitheca may not be developed over entire corallum.

)

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R. A. MCLEAN 49

Discussion. A useful summary of previous work on the genus Calostylis
has been given by Ivanovskiy (1961). He suggested a phylogenetic scheme
whereby Calostylis was seen as a Lower Silurian intermediate between Cyatholasma
(=Grewingkia), of Late Ordovician age, and the genus Helminthidiwm Lindstrom,
a Middle-Upper Silurian form. Helminthidium differs from Calostylis in that
the septa are completely broken down to a retiform condition and individual
radial septa cannot be distinguished. The other member of the family Calosty-
lidae, the early Silurian Palaearaea Lindstrém, is an astreoid form having perforate
septa and “spongy ”’ axial structure. Ivanovskiy (1961) suggested that Pal-
aearaea was derived from “‘ Cyatholasma”’ but it could equally have been derived
from Calostylis.

The species composition of Calostylis is badly in need of revision. C. tomesi
Smith, 1930, from the Wenlock Limestone (Upper Wenlockian) of Shropshire,
C. parvula Foerste, 1917, from the ‘‘ Laurel Limestone ” (Wenlockian) of Ohio
and C. dravidiana Reed, 1912, from the ? Llandoverian of Spiti, Central
Himalayas, require detailed thin-section study before their affinities are known.
The type species, C. denticulata (Kjerulf, 1865), is a very variable form, and there
are several species which are very similar to it, and may in fact be found to be
Synonymous when they are studied in more detail. Among these forms are
C. togata Smith, 1930, from the Purple Shales (Upper Llandoverian) of Shropshire,
and C. spongiosa Foerste, 1906, from the Waco Limestone (Upper Llandoverian)
of Kentucky, both of which were stated to be closely similar to C. denticulata
by Smith (1930a). C. concavifundatus Reiman in Kaljo and Reiman, 1958,
from the Porkuni Horizon (Upper Ashgillian) of Estonia and Llandoverian of the
Siberian Platform is also similar to the type species and was listed as a synonym
of it by Ivanovskiy (1970b).

C. aberrans Smith, 1930, from the Pentamerus Beds and Purple Shales
(Upper Llandoverian) of Shropshire, is closely similar to C. roemeri Smith, 1930,
differing only in its distorted growth form. There are intermediate forms between
it and C. roemeri (Smith, 1930a, 1930b) and it may be better considered as
synonymous with or a subspecies of C. roemert.

The phaceloid Middle Devonian species from Kentucky, C. ? trigemma
(Davis, 1887), as described by Stumm (1965) differs from the typical Ordovician-
Silurian representatives of the genus in its growth habit and very weakly developed
axial structure. Its systematic position is uncertain.

Range. Upper Ashgillian of Estonia; Lower-Middle Llandoverian of
Estonia, Siberian Platform, Tadzhikistan, New South Wales; Upper
Llandoverian of England, Scotland, Norway, Gotland, Estonia, Urals, Tadz-
hikistan, ? Central Himalayas, Kentucky ; Wenlockian of England, Gotland,
Tadzhikistan, Ohio; ? Ludlovian of Kazakhstan.

Calostylis panuarensis sp. Nov.
Plate U, figs 2-10; Text-fig. 4a, D

Derivation of name. After the major stream in the area, Panuara Rivulet,
an alternative name for Four Mile Creek.

Material. Holotype SUP 69220. Paratypes SUP 45158-45166, 63275,
69221. Brown Mudstone Horizon, Angullong district. Late Lower or Early
Middle Llandoverian.

Diagnosis. Calostylis either solitary or with rare lateral buds, cylindrical,
with average corallite diameter of 12mm. Broad axial and narrow peripheral
reticulate zone; 66-76 strongly perforate septa. Tabulae sagging, with axial
convex zone. Abundant sclerenchyme filling of corallite interior.

PROCEEDINGS OF THE LINNEAN Socrety oF NEw SoutH WALES, Vou. 99, Part 1

50 LOWER SILURIAN RUGOSA FROM NEW SOUTH WALES

Description. Corallum solitary, in three cases (SUP 45163, 45165, 63275)
showing budding. Increase is lateral and non-parricidal. Corallites cylindrical,
slightly tapering proximally with sharp increase in diameter soon after budding
takes place (Plate w, fig. 2). Epitheca not preserved in any specimens. Corallite
diameter varies from 10-5 to 15-5 mm in material available, with an average of
12-13 mm. Weak calicular boss present but complete calice not preserved in
any specimen. Disposition of layering of sclerenchyme in corallites suggests
rather shallow calice.

Septa entirely perforate, 66-76 in number, breaking down axially and to a
lesser extent peripherally, into anastomosing, reticulate structure. Minor septa
long, extending 0-5-0-7 of length of major septa, as best illustrated by holotype
SUP 69220 (Plate u, figs 8, 9; Text-fig. 4a). Individual septa not traceable
axially beyond margin of central reticulate structure, which varies from about
4-5 to 5mm in diameter, i.e. slightly more than 0-3 of corallite diameter.
Peripheral reticulate zone, where it can be clearly differentiated, reaches diameter
of about 1-5mm, averaging 0-1 of diameter of corallite.

Tabulae generally not clearly defined, strongly sagging near margin of axial
structure and prominently convex in the axial zone, although difficult to trace
actual plates in this area. Tabulae may be defined by abundant sclerenchyme
deposits and layers of sediment, suggestive of periodic pauses in growth (e.g.
SUP 45166). Average spacing of tabulae of about 0-7-1 mm.

Text-fig. 4. Calostylis panuarensis sp. nov. a. SUP 69220b, holotype, transverse section, x 4.
b. SUP 45158b, paratype, longitudinal section, x 4.

Remarks. Of the described species of Calostylis, four show some similarities
to C. panuarensis. Of these, the type species, C. denticulata (Kjerulf, 1865)
from the Llandoverian-Wenlockian of Scandinavia and Central Asia has perhaps
the closest affinities. From the descriptions by Smith (1930a) of material from
Gotland and Norway, similarities may be seen in the cylindrical, solitary growth
form together with lateral, non-parricidal increase in some specimens, long minor
septa, almost extending to the axial zone, the size and nature of axial reticulate
structure and the shape and disposition of the tabulae. The transverse section
illustrated by Smith (1930a, Pl. XI, fig. 5) in particular appears very similar
to the local form. However, the general size of corallites (common diameter
20 mm, reaching 35 mm in some cases, although some may be as small as 13 mm),
the usually more clearly defined and wider peripheral reticulate zone, together

PROCEEDINGS OF THE LINNEAN Socrety of New SourH Watss, Vou. 99, Part 1

R. A. MCLEAN 51

with a greater number of septa (average about 90, compared to 66-76 in C.
panuarensis) serve to distinguish it from the Angullong species. According to
Lindstr6m (1868), C. denticulata may even have up to 140 septa.

C. denticulata has also been described from Horizon D (Lower Llandoverian)
to Horizon K (Lower Wenlockian) of the Zeravshan-Gissar region of Tadzhikistan
(Lavrusevich, 1971). The form from this area differs slightly in showing a greater
tendency of budding, larger size with proportionally greater number of septa,
generally thinner septal elements and a wide peripheral reticulate zone. It is
evident that the two species are very closely related.

Several of the inadequately studied species of Calostylis show similarities
to C. panuarensis also. OC. togata Smith (1930a), from the Purple Shales of
Shropshire, is comparable to C. panuarensis in the features of growth form, septal
size and axial structure and is also of more comparable dimensions (average
corallite diameter 13mm). However, it too differs in having a larger number
of septa (approximately 90) and a considerably broader peripheral reticulate zone.
As Smith (1930a, 1930b) pointed out and was discussed above, it is quite likely that
C. togata may represent a smaller variant of C. denticulata.

The two described North American species also show similarities to C.
panuarensis. C. spongiosa Foerste, 1906, from the Waco Limestone (Upper
Llandoverian) of Kentucky is comparable in growth form, size and axial structure
but has a greater number of septa (100), weakly developed minor septa and a
wide, thickened peripheral zone.

C. parvula Foerste, from the Laurel Limestone (Wenlockian) of Ohio is of
comparable size, septal number and axial structure, but is more trochoid in growth
form, has more clearly differentiated minor septa and the peripheral zone is not
as clearly developed. However, this species has apparently not been studied
in thin section and hence the internal structures are imperfectly known.

Therefore, while the local form is assigned here to a new species, further
study of some of the European and North American representatives of Calostylis
may lead to a revision of its taxonomic position. Particularly important in this
respect would be a restudy of the smaller variants of C. denticulata.

ACKNOWLEDGEMENTS

The author is grateful to Dr. B. D. Webby for critical comments on the
manuscript. The work was partly supported by a Commonwealth Post-graduate
Fellowship at the University of Sydney.

References

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Duncan, H., 1956.—Ordovician and Silurian coral faunas of western United States. Prof. Pap
U.S. geol. Surv., 1021-F: 209-236.
Fiucet, H. W., and Sateu, H., 1970.—Die palaozoischen Korallenfaunen Ost-Irans. I. Rugose
Korallen der Niur-Formation (Silur). Jb. geol. Bundesanst., 113: 267-302.
Forrste, A. F., 1888.—Notes on Palaeozoic fossils. Bull. Sci. Lab. Denison Univ., 3: 117-137.
—., 1906.—The Silurian, Devonian and Irvine Formations of East-Central Kentucky etc.
Bull. Ky. geol. Surv., 7: 1-369.
, 1917.—Notes on Silurian fossils from Ohio and other central states. Ohio. J. Sci.,
17: 187-204, 233-266.
Hit, D., 1940.—The Silurian rugosa of the Yass-Bowning District, N.S.W. Proc. Linn. Soc.
N.S.W., 65: 388-420.
, 1956.—Rugosa. In Moore, R. C. (Ed.), Treatise on Invertebrate Paleontology. Part F.
Coelenterata. Lawrence, Kansas: Univ. Kansas Press, pp. F233—F324.
Ivanovskiy, A. B., 1961.—Nekotorie dannie o rugozakh semeystva Calostylidae. Paleont.
Zh., 1961, 2: 119-121.

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEW SoutTH WALES, VOL. 99, Part 1

52 LOWER SILURIAN RUGOSA FROM NEW SOUTH WALES

, 1962.—Dva novikh roda siluriyskikh rugoz. Trudy sib. nauchno-issled. Inst. Geol.
Geofiz. miner. Syr., 23: 126-133.

, 1963.—Rugozy ordovika i silura Sibirskoy Platformi. Moscow: Nauka (Akad. Nauk
S.S.S.R., Sib. Otd. Inst. Geol. Geofiz.), 160 pp.

, 1965a.—Drevneyshie rugozy. Moscow: Nauka (Akad. Nauk 8.8.S.R., Sib. Otd. Inst.
Geol. Geofiz.), 152 pp

, 19656.—Siratigraficheskiy %i paleobiogeograficheskiy obzor rugoz ordovika 7 silura.
Moscow: Nauka (Akad. Nauk 8.8.S8.R., Sib. Otd. Inst. Geol. Geofiz.), 120 pp.

, 1967. Morfogenez septalnogo apparata ordovikskikh 1 siluriyskikh rugoz. In
Ivanovskiy, A. B., and Sokolov, B.S8., (Eds.), Novie dannie po biostratigrafit nizhnego paleozoya
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, 1968.—The evolution of the Ordovician and Silurian rugosa. XXIII Int. geol. Cong.,
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, 1970a.—O sistematicheskom polozhenii nekotori rugoz ordovika i silura. Geol. and
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, 1970b.—-Stratigraficheskie i paleogeograficheskie compleksi rugoz na _ Sibirskoy
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Katso, D. L., 1956—O streptelasmidnykh rugozakh Pribaltyskogo ordovika. Hesti NSV
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. 1958.—Nekotorie novie i maloizvestnie rugozy Pribaltiki. Hesti NSV Tead. Akad.
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, 1961.—Dopolneniya k izuchentya streptelasmid ordovika Estonii. Hestt NSV Tead.
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, and Krnaaman, E., 1965.—The fauna of the Portrane Limestone. III. The corals.
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, and Remay, V. M., 1958.—Dva novikh vida roda Calostylis iz nizhnego silura Estonii.
Eesti NSV Tead. Akad. geol. Inst. Uurim., 2: 27-31.

Kato, M., 1963.—Fine skeletal structures in Rugosa. J. Fac. Sci. Hokkaido Univ., (4), 11:
571-630.

LamBE, L. M., 1901.—A revision of the Canadian Palaeozoic corals; the Madreporaria aporosa
and the Madreporaria rugosa. Contr. Can. Palaeont., 4, (2): 97-197.

Lane, W. D., Smirn, 8., and THomas, H. D., 1940.—Index of Palaeozoic Coral Genera. London :
Brit. Mus. Nat. Hist., 231 pp.

Lavrusevicn, A. J., 1971.—Rugozy rannego silura Zeravshano-Gissarskoy gornoy oblasti. In
Lavrusevich, A. I. (Ed.), Paleontologiya i stratigrafiya. 3. Trudy Uprav. geol. sov. Minist.
Tadzhik. S.S.S.R., pp. 38-136.

, and Menaxova, G. N., 1971.—Stratigrafiya nizhnego silura zapadnoy chasti Zerashano-
Gissarskoy gornoy oblasti. Jn Lavrusevich, A. I. (Ed.), Paleontologiya i stratigrafiya. 3.
Trudy Uprav. geol. sov. Minist. Tadzhik. S.S.S.R., pp. 3-37.

Lrnpstr6m, G., 1868.—Om tvenne nya 6fversiluriska koraller fran Gotland. Ofvers. K. Vetensk.
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Link, A. G., 1970.—Age and correlations of the Siluro-Devonian strata in the Yass Basin, New
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, and Druce, E. C., 1972.—Ludlovian and Gedinnian conodont stratigraphy of the Yass
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McLean, R. A., 1974.—Cystiphyllidae and Goniophyllidae (Rugosa) from the Lower Silurian of
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Mryato, M., 1961.—Ontogenetic study of some Silurian corals of Gotland. Stockh. Contr. Geol.,
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, 1969.—Upper Ordovician Streptelasmatid corals from Scandinavia. Bull. geol.
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PROCEEDINGS OF THE LINNEAN Society or New Sourn WaAtzs, Vou. 99, Part 1

PLATE I

Part 1

Vol. 99,

Soc. N.S.W.,

N.

Proc. LIN

Proc. Linn. Soc. N.S.W., Vol. 99, Part 1 PLATE 1

R. A. MCLEAN 53

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Unders. Afh. (Ca), 19: 1-94.

EXPLANATION OF PLATES
PLATE I

Figs 1-6. Streptelasma recisum sp. nov., Bridge Creek Limestone, Bridge Creek, x 4.
1. SUP 45167 (at left of figure), holotype, transverse section, ephebic stage; SUP 45168 (at
right), paratype, transverse section, ephebic stage. 2. SUP 45175, paratype, longitudinal section.
3. SUP 45171, paratype, transverse section, neanic stage. 4. SUP 45174, paratype, longi-
tudinal section. 5. SUP 45169, transverse section, ephebic stage. 6. SUP 45172, paratype,
transverse section, neanic stage.

Figs 7-10. Grewingkia parva sp. nov., Brown Mudstone Horizon, Angullong, x 4.
7. SUP 4515la, holotype, transverse section. 8. SUP 45153, paratype, transverse section.
9. SUP 45152, paratype, longitudinal section. 10. SUP 45154, paratype, transverse section.

Figs 11-12. Grewingkia neumani sp. nov., coarse calcarenite lens, Brown Mudstone Horizon,
Angullong. 11. SUP 20115a, holotype, transverse section, x2. 12. SUP 20115c, holotype,
longitudinal section, x 4.

PLATE II

Fig. 1. Grewingkia neumani sp. nov., coarse calearenite lens, Brown Mudstone Horizon,
Angullong. SUP 20115b, holotype, transverse section of peripheral region of corallite in calice,
howing poorly preserved trabeculae, x 8.

Figs 2-10. Calostylis panuarensis sp. nov., Brown Mudstone Horizon, Angullong.
2. SUP 45165, paratype, longitudinal section showing lateral bud, x3. 3. SUP 45158b, paratype,
longitudinal section, x4. 4. SUP 45162a, paratype, transverse section, x4. 5. SUP 4516la,
paratype, transverse section, x4. 6. SUP 45163a, paratype, transverse section showing lateral
bud at left, x4. 7. SUP 45162b, paratype, longitudinal section, x 4. 8. SUP 69220a, holotype,
transverse section at level of calice, x4. 9. SUP 69220c, holotype, transverse section, x 4.
10. SUP 45158a, paratype, transveise section, x 4.

PROCEEDINGS OF THE LINNEAN SOcIETY oF NEW SoutTH WALEs, Vot. 99, Part 1

A LABYRINTHODONT TRACKWAY FROM THE MID-TRIASSIC
NEAR SYDNEY, NEW SOUTH WALES

JULIAN PEPPERELL* AND GORDON GRIGG*
[Accepted for publicaticn 24th October, 1973]

Synopsis
A recently discovered trackway of a large Mid-Triassic Labyrinthodont is described. Notable
features of the trackway are the length (22 sequential imprints) and that the digits are laterally
directed, suggesting a primitive stance.

INTRODUCTION

In July 1970, workmen digging a sewerage tunnel at Macquarie Fields,
New South Wales, discovered what appeared to be footprints on the tunnel roof.
The authors visited the site and ascertained 22 definite imprints constituting
an excellent trackway made by a large Middle Triassic quadruped, probably a
labyrinthodont amphibian. Because of its length and some unusual features,
the trackway is of particular interest and warrants description.

METHOD OF STUDY

As the roof of the tunnel was soon to be concreted over, an accurate photo-
graphic record was made. The roof was divided into sections with chalk-marks
and each section numbered. A series of photographs was then taken with cblique
illumination to emphasize the relief of the individual footprints. A scale was
included in each photograph. A photo-montage was then built up to represent
the whole trackway and each imprint given a number. From this montage, a
drawing was produced (Fig. 1). Scale measurements were made from a tracing
of the montage and these dimensions are displayed in Table 1.

TRAVEL ==
=] a =]
e-) = = =
x = aa eeee = 3 3 B x =
= e.] mm m aR Ne o a —
nN Bcd « x) t ay = E-] Ss
= a & _@ x Wh / JERS os S a >
ee ee ee
= FR i ieee S a7 <7 nm Gz Seria v - u
ee ee
i Ne
sions 7 7
EDGE OF TUNNEL ~ (cae Sa Ey
100cm.

Fig.l. Scale diagram of the Labyrinthodont trackway.

DESCRIPTION

The trackway was in the Ashfield shales at the base of the Wianamatta
Group, of Mid-Triassic age. It consisted of a series of casts of prints made in the
underlying shale which had been removed during excavation of the tunnel.
The trackway emerged from the unexcavated section of the roof and was exposed
for three full strides before disappearing again (Fig. 1). The visible trackway
consisted of casts as follows: eight right pes, eight right manus (three indistinct),

* School of Biological Sciences, University of Sydney, New South Wales, 2006.

PROCEEDINGS OF THE LINNEAN Society or New Sourn Wangs, Vou. 99, Part 1

OL
[yi

JULIAN PEPPERELL AND GORDON GRIGG

(0)
Fig. 2. (a) Right manus (RM 2). (6) Left pes (LP 2).

PROCEEDINGS OF THE LINNEAN Soctrety or NEw Soutu Watss, Vou. 99, Part 1

56 A LABYRINTHODONT TRACKWAY FROM THE MID-TRIASSIC

TABLE 1
Dimensions of Trackway*
(Length measurements in centimetres)

Mean Range
Right manus a 5% ge 117 113-122
Stride y Left manus fe one a 111 106-114
Right pes .. as ys ! 115 111-119
Left pes... Me 5 Ae 112 109-114
(Mean stride i ae ae 114)
Trackway width .. a sie se 80
Manus pace oie ay we A 94 84-101
Pes pace aie oe 50 a es 74 63-81
Manus pace angulation .. Ss as 77° 73°—85°
Pes pace angulation te i ae 995 94°-107°
Gleno-acetabular length .. ae He 106

* The reference point for measurements of length was taken as the base of
the second outermost digit of both manus and pes.

four left pes, and five left manus. There were no signs of tail or body drag marks.
Of particular interest is the lateral direction of the digits of both pes and manus
from the axis of travel.

Dimensions that could be ascertained are givenin Table1l. Gleno-acetabular
length was determined by geometry to be 106 centimetres.

Prints RM2 and LP2 were the best preserved of manus and pes respectively.
These are illustrated in Fig. 2. The manus appears to have three digits, the pes
four. Most impressions have a ridge extending anteriorly which prebably
indicates a drag mark made during protraction of the limb. Unfortunately,
These marks could have obscured an extra digit if it were present. The drag
marks are more pronounced in those imprints made by right pes and right manus,
probably because the animal was making a slow left turn. Webbing between
digits is not indicated in either pes or manus, except for a slight suggestion of a
membrane between the third and fourth digits in LP2, LP3, LP4, RP6, RP7
and RP8.

DISCUSSION

Because of the pace angulation and the angle of the digits, and because the
only tetrapods known from this formation are amphibians, the trackmaker was
probably a large Labyrinthodont. If so, then the gleno-acetabular length of
106 centimetres is consistent with a total length of about three metres. The
laterally directed digits of pes and manus indicate a primitive stance close to that
proposed by Morton (1926) as the likely stance for primitive tetrapods. Only
one other trackway has been described with this primitive feature, an upper
Devonian record from Victoria, Australia (Warren and Wakefield, 1972). Absence
of marks indicating body or tail drag suggests either that the animal may have
been wading through a shallow billabong or marsh with body weight partly
supported by water or that the animal was capable of walking with its body clear
of the ground.

ACKNOWLEDGEMENTS

We are grateful to personnel from the Metropolitan Water, Sewerage and
Drainage Board for drawing our attention to their discovery. Advice and help
from Dr. Anne Warren is gratefully acknowledged. Mr. David Stanley prepared
the diagrams.

References

Morton, D. J., 1926.—Notes on the footprint of Thinopus antiquus. Am. J. Sci., 12: 409-414.
Warren, J. W., and Waxeriecp, N. A., 1972.—Trackways of tetrapod vertebrates from the
Upper Devonian of Victoria, Australia. Nature, 238: 469-470.

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A NEW TROGLOBITIC QUADRANNULATE LAND-LEECH FROM
PAPUA (HIRUDINOIDEA : HAEMADIPSIDAE s.1.)

LAURENCE R. RICHARDSON*
[Accepted for publication 24th October, 1973]

Synopsis

A troglobitic quadrannulate land-leech, Lezobdella jawarerensis gen. et sp. nov., is associated
with insectivorous bats in the aphotic zone of acave. its general somital annulation corresponds
with that of Philaemon, but differs in having the recurrent limb of the female median region about
half the length of the procurrent limb, the lambertian organs posterior, elongate cylindrical, the
organ much longer than its duct, and the first pair of nephropores lateral on viii a,.

A 4-annulate land-leech from the aphotic zone of a cave near Jawarere,
Papua, lacks general cutaneous pigment. The retinal cells of the eyes are
pigmented ; otherwise the preserved animal is opaque white. The animal is
the same in life. Although many species of land-leeches are found in the vestibule
of many caves, this is the first record of a troglobitic land-leech.

It is also the first record of a troglobitic leech among the euthylaematous
leeches, which are characterized by a pharynx whose internal muscular ridges
extend along its full length as dorsomedian and ventrolateral ridges, and by the
fact both that somite v is the first complete somite and that it forms the lateral
and ventral margins of the anterior sucker.

Euthylaematous leeches form a very large and diverse assembly. It contains
the majority of leeches with a pharynx and includes not only freshwater,
amphibious, terrestrial and terricolous jawed and jawless macrophagous leeches
which ingest smaller animals entire, but also freshwater, terrestrial and terricolous
jawed sanguivorous leeches.

The only previous record of a troglobitic leech is the aquatic, eyeless, opaque
white Dina absoloni Johansson, 1913, found in some caves in Europe; but there
is now knowledge of a similar leech in a cave in Japan. JD. absoloni belongs to
the smaller compact group having a strepsilaematous pharynx: the internal
muscular ridges as a dorsomedian and ventrolaterals anterior to somite vii,
transposing posterior to this to be dorsolaterals and a ventromedian for the
length of the pharynx ; ; with iv as the first complete somite forming the lateral
and ventral margins of the pharynx. All leeches in this group are freshwater
and macrophagous.

Considering the great diversity of habit and habitat of the euthylaematous
leeches, it is of interest that the first record of a troglobitic species is not that of
a macrophagous leech with its capacity to feed on the lesser fauna within a cave
but of a terrestrial jawed sanguivore so restricted in habit as to be dependent
on a vertebrate.

The 4-annulate land-leeches of the Australian Region are known to me in
specimens from Tasmania, southern Victoria, eastern New South Wales, eastern
Queensland, the Northern Territory, Papua and Lord Howe Island. They are
a complex group which I have not yet been able to bring into satisfactory
zoological order as a whole. Some few are distinctive in external meristic
morphology, in the form of the auricle and in pattern.

* 4 Bacon St., Grafton, New South Wales, 2460.

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58 A NEW TROGLOBITIC QUADRANNULATE LAND-LEECH

The majority are monotonous in external meristic morphology, in the
morphology of the alimentary tract and in the morphology of the reproductive
systems, but they separate by pattern into groups, in the same manner as our
aquatic jawed sanguivores. When the latter are grouped by pattern, genera can
be separated within each group by distinctive differences in the morphology of
the median regions of the reproductive systems. This is not possible in the
4-annulates ‘since the median regions have been monotonous in form in those
previously studied. This seemed to leave only pattern as a primary guide to
genera.

In the absence of pattern, I was initially reluctant to base a new genus on the
Jawarere leech. It seemed preferable to describe it as a new species and to assign
the species to the genus Philaemon, based on P. pungens of Victoria (the traditional
genus to which 4-annulate leeches are assigned), leaving the nature of the new
genus until such time as a patterned species could be described as the type
species. However, I now realize that both the form of the female median region
in the Jawarere leech and the form of the lambertian organs are different from
those of all other 4-annulates known to me. The combination of these with
other features described below fully warrants the provision of a new genus.

A particular feature is the location of the first pair of nephropores in the
marginal field, lateral on viii a, in the Jawarere leech (and also in a second species
of 4-annulate from Papua still to be detailed and named).

Earlier (1969) I drew attention to fig. 8 in Blanchard (1917) which shows
in a diagrammatic manner the dorsal aspect of the anterior somites of Philaemon
minutus Blanchard 1917 of Samoa. In this figure, a pair of nephropores, labelled
the second pair, is shown on viii a,. I suggested there had been a simple error,
or alternatively that these were in fact the first pair of nephropores. The latter
now seems to be the case.

Lambert (1898) fully establishes the first pair of nephropores as labial in
position in P. pungens, i.e. sitiated on the margin of the sucker at the level of
vi a, (posterior to the position in my experience). They are labial and at iii/iv
in P. grandis Ingram 1957 of Tasmania, and at about this level in other Australian
4-annulates I have studied, including Neoterrabdella australis Richardson, 1969.
The first pair of nephropores is not recorded as other than labial in two-jawed
and three-jawed 4-, 5- and 6-annulates elsewhere, and known to me only as
labial in the 4-, 5- and 6-annulates in eastern Australia which are all two-jawed,
with one exception.

The exception was the two-jawed 5-annulate [diobdella seychellensis Harding,
1913, in the Seychelle Islands, a leech with other morphological characteristics
which cause this genus to be separated from terrestrial jawed sanguivores
elsewhere.

To date I have dissected specimens of 10 or 11 species of land-leeches in the
Australian Region. I have published (1969) an account of Neoterrabdella
australis, and there is more recently (Richardson, 1974) an account of Dom-
anibdella tristriata (Goddard, 1909). To this knowledge can be added Lambert’s
description (1898) of Philaemon pungens and Ingram’s descriptions (1957) of
P. pungens and P. grandis. This is not a full fauna, but it is a usable sampling.

In all these, the ovaries are posterior in xii, or in the contiguous annuli
of xii and xiii; the oviducts join at xii/xiii or anterior in xiii ; the initial recurrent
limb of the female median region is essentially equal in length to the length of the
terminal procurrent limb, the latter slightly longer, about the length of an annulus
or a little longer in those in which the female pore is median or slightly anterior
mM Xili.

In the Jawarere leech, the ovaries and the junction of the oviducts are
essentially at the same levels as in the others, but the initial recurrent limb of

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LAURENCE R. RICHARDSON 59

the median region is short, about half or slightly less than half of the length of
the terminal procurrent limb. The difference is the same in a contracted as in
an extended specimen.

Of the 4- and 5-annulate Australian land-leeches which I have dissected,
only Neoterrabdella australis lacks lambertian organs.

In all the 5-annulates known to me, including Chtonobdella limbata Grube,
1866 (described by Lambert in 1899 under the name of “‘ Geobdella australiensis ’’)
of central eastern New South Wales, ‘‘ Geobdella”’ whitmant Lambert, 1899, of
southern eastern Queensland, and Domanibdella tristiata of Papua, the lambertian
organs are either posterior in position, situated.in the contiguous halves of xxii
and xxiii, the organ ovoid about the length of a somite, and the duct and organ
nearly equalin length ; or, asin D. tristriata, the organ longer, elongate cylindrical,
from xxi/xxii to the’ middle of xxiii, the duct very short, its length no more than
the width of the organ.

In the 4-annulates, the lambertian organs are anterior; in xx, the organ
obtusely ovoid, short, and the duct long; or in the contiguous parts of xx and
xxi, the organ longer, the duct shortened ; but in both cases, the organ is much
shorter than the duct, the length of the organ half or less than half of the length
of the duct. The organs are in xx in Philaemon pungens (Lambert, 1899, pl. x,
fig. 2).

In the Jawarere leech, the lambertian organs in xxi and xxii are elongate
cylindrical, folded, the length of two somites, and the organ much longer than
the duct, about twice its length, i.e. the lambertian organs and ducts differ from
the form in the 4-annulates and resemble the form of these structures in the
5-annulates.

Only the general external meristic aneraalnalone is known for the great
majority of land-leeches.

Phytobdella catenifera Moore, 1938, is a two-jawed 6-annulate land-leech
from Johore, India. In his description of this species, Moore shows (1938,
pl. v, figs 10, 11) the initial recurrent limb of the female median region as short,
about half the length of the terminal procurrent limb, and the lambertian organ
aS posterior in position, elongate, and more than twice the length of the short
lambertian duct. Both therefore are the same as the Jawarere leech.

Keegan et al. (1969, fig. 39A) show the female median region in a two-jawed
6-annulate from Laos as having the two limbs equal in length.

Moore (1944) describes the lambertian organs as posterior, extending from
xxi to the level of the end of the postcaecum, the duct short, the gland seven
to nine times the length of the duct, in Phytobdella lineata Moore, 1944, and
Phytobdella maculosa Moore, 1944, both of Papua. He does not adequately
describe the female reproductive system. These are both two-jawed 6-annulates.

A two-jawed 6-annulate collected by Dr. Ewers at Mount Suckling, differs
from the above species in details of the somital annulation. I find in this
specimen that the two limbs of the female median region are essentially equal
in length and that the lambertian organ is elongate cylindrical, folded on itself,
extending anteriorly to xxi, the duct short, the gland six to seven times the length
of the duct. Mount Suckling lies in Papua.

I review briefly elsewhere (Richardson, 1974) the manner in which land-
leeches have been separated systematically. Initially, all 4-annulates were
placed in the genus Philaemon, with the species separated on the location of the
genital pores and the annulation of vii. Later, genera were separated on the
number of jaws and on the presence or absence of lambertian organs.

To this I added (1969) the annulation of xxiv and xxv, the form of the
auricle, and, recently, the annulation of vi, vii, etc., making a further step in
the separation of genera.

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60 A NEW TROGLOBITIC QUADRANNULATE LAND-LEECH

The 4-annulate genera previously recognized in the Haemadipsidae (Soos,
1967) were Philaemon, a name proposed by Blanchard in 1897 for 4-annulates,
but without detail until Lambert (1898); Tritetrabdella Moore, 1938; Neso-
philaemon Nybelin, 1943. To these I added Neoterrabdella Richardson, 1969,
and, from the nature of the reproductive systems, I removed Nesophilaemon
from the Haemadipsidae.

Tritetrabdella of Penang, Malaya, has three jaws, lacks lambertian organs ;
xxiv, 2-annulate; genital pores, xi a,/b; and xii a, (a,/b;).

Neoterrabdella of the Northern Territory, Australia, lacks lambertian organs ;
xxiv, 3-annulate; xxv, 2-annulate; the auricles large, formed by the lateral
ends of xxiv-xxvii; genital pores, xi a, (a,/b;, b;) and xii bs. Philaemon, of
Victoria and Tasmania, Australia, as in Lambert (1898) and Ingram (1957),
has lambertian organs; xxiv, uniannulate; auricles formed by the lateral
ends of xxv—xxvii; genital pores, xi b; (? b;/b,) and xii b, (? b;/b,). Both genera
have two jaws.

With genera separated as above, the Jawarere leech would be assigned to
Philaemon as a species distinct in having the genital pores in xi b, and xii a,/b;.

To the previous criteria for the separation of genera we can add the form
both of the lambertian organ and of the female median region, and also the
location of the first pair of nephropores. This combination excludes the Jawarere
leech from the previously established genera.

If this is adopted, some generic separations can be anticipated in the
6-annulates, all currently in the genus Phytobdella.

Genus LEIOBDELLA gen. nov.
Deriwwation. Leto, smooth +bdella, leech.

Description. Somites viii-xxii, complete 4-annulate (total, 15); xxiii,
incomplete 4-annulate ; xxiv—xxvii, incomplete uniannulate ; six annuli posterior
to xxiii a,; auricles lateral to xxv, xxvi, xxvii, margin lobate, incised ; first
nephropores lateral on viii a, ; duognathous ; jaws housed in an annular groove ;
teeth, minute ; salivary glands, sparse, no obvious columns of aggregated ducts ;
extrinsic radial musculature in vii-ix ; pharynx thin-walled, internal muscular
ridges represented by an undivided dorsomedian and a pair of ventrolateral
ridges; pharynx terminates at viii/ix; crop compartmented, caecate, the
compartments each with a pair of primary caeca at the median level and reduced
secondary anterior and posterior caeca, excepting xix with secondary anterior
caeca and postcaeca originating at the median level, extending in the paramedian
chamber to xxiii/xiv and each continuing as a lambertian duct shorter than the
elongate cylindrical lambertian organs in xxi and xxii; genital pores, xi (b;/b,) be
and xii a,/b;; male median region, amyomeric, micromorphic ; female median
region, myomeric, mesomorphic, haemadipsoid; testes, saccular; anterior
region of the male paired duct reflecting in the median splanchnic chamber at
xi/xii or in xii as a primary posteriorly directed loop, an epididymis on the initial
portion of the recurrent limb, a sperm duct completing the recurrent limb and |
extending along the length of the procurrent limb ; ejaculatory bulbs, present ;
ejaculatory ducts enter the anterior aspect of the basal portion of the minute
atrium ; ovaries, saccular, at xii/xiii or in xiii; oviducts join at xii/xiii to median
in xiii; female median region, a posteriorly directed loop reflecting in xiii—xiii/xiv,
the initial recurrent limb short, half or less of the length of the terminal procurrent
limb; oviducal glandular sac expanded on the posterior aspect of the elbow of
the loop.

Terrestrial. Sanguivorous. Australian Region, Papua.
Pattern. % The type species, cavernicolous and lacking pattern.
Type species. Leiobdella jawarerensis sp. nov.

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LAURENCE R. RICHARDSON 61
an.gr

Fig. 1. Letobdella jawarerensis gen. et sp. nov. 1. Annulation of somites i—ix, left lateral

view, sensillae indicated in v—vii. 2. Annulation of somites xx—xxvii; the auricle; posterior
sucker ; left lateral view. 3. Ventral view of somites xi and xii, showing the location of the
genital pores. 4. Annular groove, jaws, entrance to pharynx, and the internal muscular ridges,
as exposed by a median ventral incision ; arrow marks midpoint in length of pharynx. 5. Caeca-
tion of crop, somites xviii and xix, postcaeca, lambertian organs, intestine and rectum.

1.1-1.5 drawn from the type.

Somites and somital ganglia indicated by roman numerals; somital limits, broken lines ;
annuli, ‘‘a,”’, ete. Scales equal 1-0 mm.

Abbreviations: an.gr., annular groove; dm.r., dorsomedian muscular ridge; f.p., female
pore ; int., intermediate somital sense organs; 1.d., lambertian duct; 1.o., lambertian organ ;

m.p., male pore; nepr., nephropore ; p.c., postcaecum ; pm., paramedian somital sense organ ;
re., rectum ; vl.r., ventrolateral muscular ridge.

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62 A NEW TROGLOBITIC QUADRANNULATE LAND-LEECH

Leiobdella jawarerensis sp. nov.
(Figs 1, 2, 3)

Type. One specimen, 19:5 mm long. Jawarere Cave, Papua. 14/7/68.
Collector W. H. Ewers. Deposited Australian Museum, Sydney, Collection
No. W. 4299. Dissected.

Paratypes. Two specimens, 18-0 and 42:0mm long. Same locality, date
and collector as type. Deposited Australian Museum, Sydney, Collection
No. W. 4303. Dissected.

Representative material from the same collection has been placed in the
Papua and New Guinea Museum and Art Gallery, Port Moresby.

Excepting the reproductive systems, which are detailed from the paratypes,

the following account is based on the type.
General form. In a preserved state, moderately contracted ; the body generally
low convex across the dorsum, the margins obtusely rounded, the venter flat ;
widest along the posterior half of the body, reducing gradually in width anteriorly
to form a narrow short region, a base for the wider anterior sucker which is about
two-thirds of the maximum width of the body ; posteriorly, narrowing abruptly
along the postnephric somites to form the base for the posterior sucker, which is
slightly wider than the body. Richly papillate.

Total length, 19-5 mm ; the anterior sucker, 3-6 mm wide at v/vi; the body,
2°8mm wide and 2-0 mm deep at viii/ix, 3-0 mm from the tip of the velum,
5-2 mm wide and 3-5 mm deep at 10-0 mm from the tip of the velum, 6-0 mm
wide and 4-0 mm deep at 14:0 mm, and of these dimensions to xxii a, ; narrowing
then to the base of the sucker, 2-0 mm wide at xxvii, 18-0 mm from the velum ;
the posterior sucker, 5-8mm wide and 6-0 mm long.

Colour. Preserved in alcohol. Immaculate, opaque china-white on all
aspects without dark pigment other than in the retinal cells of the eyes and a few
spaced small erratic patches lateral to the ocular arch.

Pattern. There is no indication of any pattern.

Annulation (Figs 1.1, 1.2, 1.3). Interannular and intersomital furrows
equivalent and somital limits not directly recognizable.

Richly ornamented with low obtusely-rounded papillae, most being marked
with a white sensory patch. Papillae arranged as a transverse row on each
annulus, large and small on the dorsal and lateral aspects, only small on the
venter ; the large papillae appearing as though arranged in sinuous longitudinal
lines which are not recognizably related to the lines and fields of the somital sense
organs. In somite xvi, each annulus carries about 25 to 27 papillae.

The somital sense organs of the dorsal and lateral aspects are most difficult
to recognize as longitudinal series, and cannot be recognized with any confidence
on the venter. Confusion comes from the fact that b,; is by far the longest
annulus in most somites, and it is only with the identification of a, by the presence
of the minute and obscure nephropores that the somital annulation can be
analysed. The nephropores are central in a, in anterior somites and close to the
posterior edge of the anulus in posterior somites.

There is a soft narrow rounded rim on an undivided thin semitransparent
membrane which forms the lateral and ventrolateral wall of the chamber lateral
to somites i-iv and anterior to v, which in turn forms the ventral portion of the
margin of the sucker, and which, with vi, completes the wall of the chamber of
the sucker. There is a transverse row of some eight large white sense organs on
the dorsal portion of the rim of the sucker.

A weak but distinct short furrow extending across the median field separates
somite i from ii; ii, uniannulate, the first pair of eyes, as also the eyes in ili and
iv, located each in a large areola, and some areolation also in i, ii, iii, with
indications of areolation in iv and v in the median and paramedian fields ; 11,
uniannulate, with the second of eyes ; iv, 2-annulate between the ocular areolae,
the third pair of eyes in a,a,—a,; Vv, incomplete 2-annulate, the fourth pair

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- LAURENCE R. RICHARDSON , 63

of eyes in a,a >a 3, the furrow a,a,/a, terminating at the ocular areolae, v uni-
annulate below this and forming only the ventral margin of the sucker; vi,
incomplete 3-annulate, a,—a, slightly >a,; the fifth eyes in a,, the furrow
a,/a. extending into the marginal area, and vi 2-annulate across the venter ;
vii, 3-annulate, a,—a, slightly >as.

Somites viii-xxii complete 4-annulate (total, 15); the midnephric series
with a, <a,—b;> bg or a, <a. <b; > bg.

The first nephropores lateral on viii ay.

Somite viii, a, =a, slightly <b; >b,; ix, a; <a ,.<b;>b,; x, not assessable ;
Xi, a, <a,—D; > Dg, as also Xil; Xili, a, <a,—b,? b,; Xiv, a, <a, slightly <b, >be,
aS also xv; xvi to xxii, with a,<or slightly <a,<b,>b, (and b,>than the
following a,); Xxiii, incomplete 4-annulate, a, slightly<a,>b,; slightly >b,,
the last pair of lateral nephropores close to the posterior edge of a,, and xxiii a»
tha last annulus fully formed across the venter, b; and possibly b, cross the venter
as very thin ridges.

Somites xxiv, XXV, XXVi, XxXvii, incomplete uniannulate. Anus at the
posterior border of xxvii.

Auricles, relatively small, restricted to the lateral ends of xxv, xxvi, xxvii ;
Xxv expanding into a poorly-formed anterior pointed lobe; xxvi, shorter,
pointed, and bridging between the anterior lobe and the larger rounded posterior
lobe which is formed by xxvii. The auricles roof over a small but distinct
cavern.

The dorsum of the posterior sucker carries some six concentric rows of
papillae. The ventral surface has a small central papillate disc, about one-quarter
of the diameter of the sucker, and radiating from the disc are muscular ridges
which subdivide to end as about 100 ridges on the margin of the sucker. The
clamp, small, acutely triangular.

Genital pores, male anterior in xi bg; female, xii a,/b;.

Alimentary tract (Figs 1.4, 1.5). Central in the dise of the anterior sucker
is a small transversely oval aperture, 1-0 mm wide. This is at the level of
vi/vii, immediately anterior to the ventral portion of the anterior ganglionic
mass which is compact, almost globular, and separated from the first independent
ganglion (vii) of the nerve cord by a space approximately equal to the length
of the ganglion.

The rim of the oval aperture is rounded, and the rim continuous with a thin
membrane.

A median ventral longitudinal incision shows a very well-developed deep
wide annular groove internal to the aperture, the anterior wall of the groove
being formed by the thin membrane and the posterior wall by the entrance to the
pharynx and the jaws.

There is no jaw or muscular pad in the dorsomedian position. The dorso-
median and dorsolateral portions of the entrance to the pharynx have the
appearance of a narrow transverse flat uniformly muscular band, continued
posteriorly as an initially narrow dorsomedian primary internal muscular ridge,
wider in the posterior half of the pharynx and without indications of secondary
subdivision.

The two ventrolateral jaws are narrowly separated ventrally, housed in the
annular groove without any indication of crypts or pockets ; each jaw, minute,
the height of the medial aspect about 0-2mm, the width of the base about
0-3 mm, and this aspect obtusely rounded in profile; the dental margin about
0-3 mm long, armed with a single row of minute sturdy teeth which appear to be
spaced, taller at the median end, and the height diminishing along the row.
(Because of the small size of the jaw, this could not be removed for a count of
the teeth. No teeth were seen on jaws dissected from other specimens.)

Each jaw continues posteriorly as a primary internal muscular ridge, initially
wide, then narrowing in the posterior half of the pharynx.

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64 A NEW TROGLOBITIC QUADRANNULATE LAND-LEECH

There are no dorsolateral or ventromedian internal muscular ridges.

The entrance to the pharynx appears wide in the absence of a dorsomedian
jaw. The lumen is tubular, tapering.

Salivary glands are sparse, located in vii-ix, without any indication of
dorsal columns of aggregated ducts.

The pharynx terminates at viii/ix followed by the compartmented crop ;
the compartment in ix, simple ; in x, xi, each with a small caecum at the median
level on each side ; xii,? ; xiii and posteriorly including the postcaeca, containing
coagulated blood, the casts indicating a pair of primary caeca at the median
level extending into and lobed in the paramedian chambers, an anterior pair
of small simple caeca retained in the median splanchnic chamber, and also
possibly a posterior pair of small simple caeca.

Fig. 2. Leiobdella jawarerensis Anterior region of male paired duct ; male median region
and female reproductive system of paratypes. 6. Strongly contracted 18:0 mm specimen, male
atrium turned posteriorly to show the termination of the ejaculatory ducts; arrows indicate
dorsal aspect of sperm duct and ejaculatory bulb, and black squares indicate the original position
in the median chamber of the posterior end of the primary loop on the anterior regions of the
male paired ducts. 7. Fully extended 42-0 mm specimen.

Abbreviations: at., atrium; ej.b., ejaculatory bulb; ej.d., ejaculatory duct; epid.,
epididymis ; ov., ovary; ovd., oviduct ; ovd.s., oviducal glandular sac ; pre.l., procurrent limb,
female primary loop; rec.|., recurrent limb, female primary loop; sp.d., sperm duct ; te., testis ;
v.d., vas deferens; v.ef., vas efferens.

The compartment in xix with postcaeca originating at the median level and
extending into the paramedian chambers, each postcaecum reducing in the
posterior half of xxiii to form a narrow duct which passes through the paramedian
palisade of dorsoventral muscles into the median splanchnic chamber, then
anteriorly to the level of xxii/xxiii, where it expands into the elongate cylindrical
lambertian organ which terminates bluntly at about xx/xxi. The lambertian
organs are parallel, ventral to the intestine.

The lambertian duct is relatively short; the lambertian organ is about
twice the length of the duct.

The compartment in xix reduces to be narrowly tubular, connecting
terminally at xix/xx to the much wider intestine which commences with a bilobed

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LAURENCE R. RICHARDSON 65

chamber in xx, continues posteriorly as a tapering tube connecting at xxiii/xxiv
to the short tapering rectum.

Reproductive systems (Figs 2.6, 2.7). Detailed and figured from the paratypes.

Haemadipsoid: the anterior region of the paired male ducts reflected in
the median splanchnic chamber on a posteriorly directed loop; ejaculatory
bulbs, present ; male median region, a weakly muscular atrium ; female median
region formed on a posteriorly directed loop with a glandular asymmetrical
oviducal sac.

Genital pores, xi b;/b,(b,), and xii a,/b;.

The type, male, mature, the organs strongly compacted, and in general
unsuitable for detailed description. Dissected sufficiently to establish that the
morphology and relationships of the reproductive systems agree with the following
description based on the dissection of an 18:0 mm contracted specimen and a
42-0 mm extended specimen.

Two facts should be noted. Both the anterior region of the male paired
ducts and the median regions are located in the median splanchnic chamber
ventral to the crop ; the anterior region of the male paired duct and the female
median region are anchored anatomically only at the anterior ends and are
subject to displacement posteriorly on contraction.

The distinct paired small sacs on the vas deferens in xii in the 18-0 mm
contracted specimen are new to me. They are not present either in the type or
in the 42-0 mm extended specimen. They are located on the duct in the para-
median splanchnic chamber at the point where the duct passes through the
paramedian palisade of dorsoventral muscles. They might possibly be a
temporary artefact, a consequence from a strangulation of the duct in contraction.

Testes saccular, 10 pairs in the median chamber, the most anterior pair
in xiv; each testis connecting laterally by a short vas efferens to the tortuous
white vas deferens in the paramedian chamber; the vas deferens, reducing in
diameter in xiii, continuing as a very narrow (diameter less than 0:1 mm)
thin-walled semitransparent straight duct which passes through the paramedian
palisade in xii and extends in the median chamber as a posteriorly directed loop
reflecting at xii/xiii or in Xiii.

The initial half of the recurrent limb of the loop is a thin-walled narrow duct,
slightly tortuous as though a poorly defined epididymis, continuous with the
tortuous much-folded wider thin-walled sperm duct which completes the initial
limb, increasing in diameter as it passes around the elbow of the loop and becomes
loosely folded along the greater length of the procurrent terminal limb to about
xi/xii, where it narrows abruptly before joining the ejaculatory bulb.

The right and left loops of the male paired ducts are parallel, not tandem,
in arrangement, and the posterior portion of each loop forms a large globular
mass dorsal to the female system.

The ejaculatory bulbs are stoutly fusiform, muscular, opalescent, terminating
obtusely ; the duct continues as a narrow opalescent ejaculatory duct which
connects to the anterior aspect of the basal portion of the male atrium. The
right and left ducts enter independently.

The male atrium is small, extending above the ventral nerve cord, and the
wall is thinly muscular.

The female reproductive system is contained wholly within the median
splanchnic chamber.

The ovaries are pyriform, situated in the contiguous annuli of xii and xiii,
each connected by a short oviduct to the recurrent limb of the median region
which is formed on a posteriorly directed primary loop reflecting in the posterior
annuli of xiii or at xiii/xiv.

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66: A NEW TROGLOBITIC QUADRANNULATE LAND-LEECH

The recurrent limb is thin-walled, narrower than the strongly muscular
procurrent limb which terminates at the genital pore and is twice or more the
length of the recurrent limb.

The oviducal glandular sac expands from the posterior aspect of the elbow
of the primary loop and terminates obtusely in xiv.

No distinct, i.e. compact, albumin or prostate glands were recognizable.

Additional material. Australian Museum, Sydney. Collection No. W. 5421.
Musgrave River area, Central District, Papua. Inside cave. 28.11.1960. Col-
lector J. Barnaby. Ex Collection No. c/6, Department of Agriculture, Port
Moresby. One specimen, 32:0mm long. Leiobdella jawarerensis. Determined
L. R. Richardson, 1973.

Discussion. Leiobdella jawarerensis is one of the large 4-annulate leeches.
Fully extended specimens measure up to 55:0 mm. _ It is variable in appearance.
Strongly to moderately extended specimens are distinctly papillate, each papilla
tipped with a white sense organ. Fully extended specimens are smooth ; somital
sense organs and sensillae showing as small white patches, of much the same size ;
interannular furrows, weak and difficult to identify ; some but not all of the testes
showing through the ventral body wall.

Cavernicola have been the basis for much speculation on adaptation and
isolation as factors inducing directive or non-directive evolution.

The troglobites, animals committed to persistent darkness, constant
temperature and humidity, and a diet restricted because it is not mediated by
photosynthetic plants, include animals from many phyla. Among these, the
common reduction of visual organs, the loss of cutaneous pigment and other
associated changes are taken as major evidence of convergent regressive evolution.

As a sanguivorous land-leech, L. jawarerensis is an interesting addition to
this area of speculation. It inhabits the aphotic zone. This fact and the fact of
the absence of general cutaneous pigment, qualify it for classification as a
troglobite. I am unable to point to any other feature which supports this
classification.

The eyes are not obviously reduced and the retinal cells are pigmented,
both as in some other troglobites. In the study of the land-leech in the bush
I reached the conclusion (1968) that vision has no part in bringing the leech to
the host. JL. jawarerensis provides further evidence in support of this conclusion.

Dr. Ewers informs me that the portion of the cave inhabited by this leech is
tunnel-like, reducing to a height of four feet at one place where there are some six
leeches to the square foot. Swiftlets, fructivorous bats and insectivorous bats
enter the cave. No vertebrate other than the insectivorous bat enters the region
occupied by the leech. The leech pays no attention to human beings.

Photographs sent to me by Dr. Ewers show leeches taking a blood meal
from bats. In these photographs the posterior sucker of the leech is attached
to the wall of the cave ; the body, extended ; the anterior sucker attached to the
arm, or anterior aspect of the fore-arm, or to the base of the web of the hand.

In reply to my enquiry, Dr. Ewers informs me that they have never seen a
leech entirely attached on a bat. When the bat flies, the leech remains on the
wall of the cave.

L. jawarerensis is accordingly a troglobite obtaining its nutriment from a
host which finds its food beyond the limits of the cave. Blood is a well-balanced
tissue, its composition held within the close limits necessary for the well-being
of the animal. Other than a monotony of diet, in the sense that DL. jawarerensis
feeds from only one host species, the nutrition does not differ from that of the
land-leech in the bush.

[ have shown (1968) that the land-leech in the bush has a closely defined
habitat preference. Activity is dependent on soil moisture and humidity, the
leech becoming inactive when these are low and anhydrobiotic in drought.

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LAURENCE R. RICHARDSON 67

In many collections from the field, the majority of the crops are empty.
When engorged land-leeches are held in captivity, the crop contains blood for
six to nine months after the meal. The indications are that blood meals are few
and widely-spaced for the land-leech in the bush, a circumstance arising both
from the task of finding a host and from its varying periods of inactivity.

Dr. Ewers informs me that most of the Jawarere leeches contain blood.
This is the case for the specimens he has sent to me. From this it seems that the
stable environment of the cave permits long periods of activity, even possibly
continuous activity with a host which is readily available.

Fig. 3. Leiobdella jawarerensis feeding from an insectivorous bat in the Jawarere Cave.
Photo: A. Johnstone.

The presence of this land-leech in the aphotic zone raises the usual queries
as to the manner in which the troglobite becomes committed to this zone. In
this case, the survival of the leech is wholly dependent on the presence of the
insectivorous bat.

I have provided (1968) the basis for showing both that the land-leech is
a poor traveller, and that a land-leech population is restricted to the limited area
of a suitable habitat. From this it becomes difficult to regard the land-leech as
an animal capable of active dispersal. This would require travel through
environments unsuitable for activity on the part of the leech.

Land-leeches are known in the vestibule of caves. In the case of L.
jawarerensis, the invasion of the aphotic zone can be assessed as having high
probability since the uniformity of the cave environment does not provide

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68 A NEW TROGLOBITIC QUADRANNULATE LAND-LEECH

barriers to the active dispersal of the leech. It required only that the leech
come into an area occupied by a suitable host animal for a community to be
established.

The alternative is that the leech was brought to this part of the cave by
passive transport on the host animal. The probability for this can be assessed
as very low, unless the bat behaves in a manner differing from other insectivorous
bats, e.g. crawls over the ground or climbs through low vegetation in leech-
infested areas in addition to feeding on the wing. If there is such behaviour,
the probability continues to be low because of the manner in which the fed leech
readily departs from the host.

In either case, it seems that the population of L. jawarerensis in this cave is
isolated.

The land-leech is a convenient laboratory animal. IL. jawarerensis is a
troglobite suitable for experimental studies. It would be valuable to use leeches
from this cave to study the consequences both of a varied diet and of exposure
to light, in order to see if these lead to the establishment of general cutaneous
pigmentation.

ACKNOWLEDGEMENTS

I am grateful to the Australian Research Grants Committee, whose award
for studies of the zoology of the Australian freshwater and terrestrial leeches
assisted this research.

I thank Dr. W. H. Ewers, University of Papua and New Guinea, for the
specimens on which this account is based and for information on Leiobdella
jawarerensis. I also thank Mr. A. Johnstone, Biology Department of the same
University, for the photographs of the leech and the bat taken in the cave.

References

Iveram, D. M., 1957.—Some Tasmanian Hirudinea. Proc. R. Soc. Tas., 91: 191-232.
KEEGAN, H. L., TosHioxa, 8., and Susux1, H., 1969.—Blood-sucking Asian Leeches of Families
Hirudidae and Haemadipsidae. U.S. Army Medical Command, Japan. 406th Med. Lab.
Spec. Rept., pp. i-vi, 1—130.
Lampert, A. M., 1898.—The structure of an Australian land leech. Proc. R. Soc. Vict., 10:
211-235.
, 1899.—Descriptions of two new species of Australian land leeches, with notes on their
anatomy. Proc. R. Soc. Vict., 11: 156-163.
Moore, J. P., 1938.—Leeches (Hirudinea) principally from the Malay Peninsula, with descriptions
of new species. Bull. Raffles Mus., 14: 64-80.
, 1944.—Leeches in the British Museum, mostly Haemadipsidae from the South Pacific,
with descriptions of new species. Ann. Mag. nat. Hist., Ser. ii, 11: 389-409.
Ricwarpson, L. R., 1968.—Observations on the Australian land-leech Chtonobdella limbata
(Grube, 1866) (Hirudinea: Haemadipsidae). Aust. Zool., 14 (3): 294-305.
———., 1969.—On a distinctive new subequatorial Australian quadrannulate land-leech, and
related matters. Aust. Zool., 15 (2): 201-213.
, 1974. Domanibdella gen. nov. a duognathous 5-annulate land-leech genus in New
Guinea (Hirudinoidea : Haemadipsidae s.l.), with a discussion on general somital annulation
Mem. Nat. Mus. Vict., 35: 97-109.
Soos, A., 1967.— Identification key to the leech (Hirudinoidea) genera of the world with a catalogue
of the species. IV. Family: Haemadipsidae. Acta zool. hung., 13 (3/4): 417-432.

PROCEEDINGS OF THE LINNEAN Soctery of New SourH Watss, VoL. 99, Part 1

SIR WILLIAM MACLEAY MEMORIAL LECTURE, 1974*

‘A SOCIETY OF NATURAL HISTORY. I HOPE
THEY MAY SUCCEED.”

The First Hundred Years
RUTHERFORD N. ROBERTSON +

The title I chose for my Macleay Memorial Lecture on the occasion of the
centenary year of the Society is a quotation from Sir William Macleay (1874)
himself : ‘‘ A Society of Natural History. I hope they may succeed ’’ and my
sub-title is ‘‘ the first hundred years’. Perhaps I should have added ‘‘ and the
future’’. I hope to develop the idea of continuity of achievement in the past
and promise for the future which, I believe, will be even more exciting.

One hundred years ago yesterday, 23rd April 1874, William Macleay was at
the North Shore Steam Ferry wharf, Circular Quay, at eight o’clock in the
morning to meet his guests for a fishing party. His diary tells us that in a short
time all were mustered and having got their provisions, etc., on board they made
a good start (Fletcher, 1920). This fishing expedition was one of a series of events
in the year 1874 which was to have profound effects not only for this Society but
also for science in Australia and, indeed, for science generally. The fishing
picnic was in honour of the scientific visitors of HMS Challenger which had arrived
in Sydney on 6th April. The party consisted of Professor Wyville Thomson, the
scientific leader on board the Challenger, and Mr. Murray, his Assistant, a
physicist ; the local guests were Professor Badham (Classics), Mr. Stephens (later
Professor), Captain Onslow, Mr. Mackay, Mr. Masters, Dr. Cox, Mr. Forster (a
grazier) and William Macleay. Of that party, Stephens, Onslow, Cox, Mackay
and Macleay were to become members of the first Council of the Linnean Society
of New South Wales when it was founded later in the same year. The coat of
arms of HMS Challenger now belongs to the Australian Academy of Science.

On the picnic day they called at HMS Dido for Captain Chapman and then
at Point Piper for Ned Hill who brought with him a blackfellow and a lot of
dynamite-torpedoes for fish—it is not recorded whether the blackfellow or the
dynamite-torpedoes had more success. Two nights later, one hundred years ago
tomorrow, the Captain and the scientific leader on board the Challenger returned
the hospitality with a dinner at 6.30 p.m. for Macleay, Onslow, Badham, Hill,
Cox, Morris, Captain Chapman and Captain Macleary. Macleay’s diary records :
‘“‘ we had a good dinner, spent a most delightful evening and had a look at some
of the current things collected. It was 11 0’clock before we left’. The Challenger
was sent out by the British Government on a circumnavigating scientific ex-
ploration of the sea in the years 1872 to 1874. A steam corvette of 2,036 tons,
she was furnished with every scientific device for examining the sea from surface
to bottom—natural history work room, chemical laboratory, aquarium, ete. It
is not difficult to imagine the immense stimulus which this visit gave to those
interested in natural history in Sydney.

* Delivered on 24th April 1974, at the Uaerey of Sydney, on the occasion of the celebration
of the Society’s centenary.
+ Research School of Biological Sciences, Tertaian National University, Canberra, A. (OS Us

PROCEEDINGS OF THE LINNEAN Society or New Soutu WAtEs, VOL. 99, Part 1

70 THE FIRST HUNDRED YEARS

It is interesting, now, to think about the Sydney in which these picnics and
this dinner on board the Challenger were taking place. Its population was about
140,000—not very different from the size of Canberra today but so different in
design. There was no need for an artificial lake to enhance its beauty but the
unplanned city whose streets and roads grew along the old bullock-dray tracks
stirred Richard Twopenny (1883) to say here a little later : ‘‘ One feels quite angry
with the town for being so unworthy of its site’’. The size and handsomeness
of the individual buildings were lost in the narrow Sydney streets and the people,
said Twopenny, ‘ succeed in looking pre-eminently dowdy’. To move from
one part of Sydney to another, you would have ridden on steam trams which
had been introduced in 1869. The Sydney Ferries Company was in its palmy
days and the main means of transport was by sea. In 1876 Sydney had 571
registered vessels of which 176 were steamers (Birch and Macmillan, 1962). You
would perhaps have been able to afford a ride in a hansom cab, though most of us
would have belonged to the classes for which such fares would have been a
luxury, for it was a time of great contrast between wealth and poverty.

Macleay, who had had a good education, had succeeded in his own activities
with his wool-growing in the country but had moved to Sydney when he was
elected to Parliament in 1855, first as Member for the Lachlan and Lower Darling,
but later as Member for the Murrumbidgee. He would have been very familiar
with the river at Wagga. He was a member of the select committee which
prepared a report in 1866 “‘ on the distress at present existing among the working
classes’. David Macmillan (1957) records that at that time conditions in the
older parts of Sydney, especially around the docks of Darling Harbour, were
horrific. The water supply was filthy, germ-infested and inadequate. Under a
crust of hopeful respectability was a world of vice, crime and hopeless poverty
which makes the worst in contemporary social conditions seem mild.

In the 1870’s the city was lit by gas introduced some thirty years before ;
only in 1874 was this lighting being extended to Redfern. In the 1860’s the
railways had gradually pushed out from Sydney and the southern railway, on
which William Macleay would have travelled towards his station on the lower
Murrumbidgee, had gone as far as Goulburn. In 1870 he had served on another
select committee on the ‘‘ best mode of facilitating the inland traffic and extension
of the railways generally’. Victoria had already pushed its railways to the
border at Albury but it was to be some time before the New South Wales railways
extended to that area.

The capital cities of the colonies had been in touch with each other by tele-
graph since 1858 and in 1874, with the completion of the transcontinental tele-
graph line from Adelaide to Darwin, telegraphic communication with Europe
was established. Other developments too were taking place. Marcus Clark was
publishing For the Term of his Natural Life. At Sydney Hospital some of the
medical staff resented the authority being assumed by nurses and the Chairman
of the Hospital Council, William Windeyer, had to write to Florence Nightingale
about the place of the nurses, some of whom she had trained (Birch and Mac-
millan, 1962). In her reply she said ‘‘a man can never govern women ”’ and
advised that the matron’s power should be increased ; advice which was followed.
Elsewhere, Australia was still very much in the process of development. John
Forrest had completed his long walk from Perth to Adelaide in 1870. As Macleay
held his fishing picnic, Warburton was just back in Adelaide after more than a
year crossing from Alice Springs to the west coast under conditions of almost
incredible hardship. In Adelaide, the University was in its first year.

On the world scene, science was just emerging into that strong development
which characterised the last part of the nineteenth century. It was only fifteen
years after the publication of The Origin of Species, the greatest biological event
of all time. Those interested in natural history in Sydney were avidly reading

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RUTHERFORD N. ROBERTSON (ale

the thrust and counter-thrust of the sometimes violent debate which took place
on Darwin’s revolutionary views. They did not know then that nine years earlier
that quiet monk who was also a scientist, Gregor Mendel, had published a most
important paper in the Proceedings of the Natural History Society of Briin,
which not to be discovered by the scientific world until the year 1900.

Sir Joseph Dalton Hooker, who had visited Australia with Ross on HMS
Hrebus on the way to the Antarctic, was now President of the Royal Society of
London and Director of Kew Gardens. With his interests in the varied flora
which he had seen, particularly in Tasmania, he was largely responsible for the
work undertaken by Bentham who was second only to Hooker amongst nineteenth
century British systematists. The collaboration between Bentham and von
Mueller resulted in Flora Australiensis. The seventh and concluding volume of
this great work which was some sixteen years in progress was not published until
1878. In first part issued in 1863, Mr. Bentham expressed an intention of forming
a supplementary volume to contain an account of new species added to existing
knowledge during the progress of the work. In 1878 we read that owing to
increasing age and infirmity, Mr. Bentham found himself unable to undertake the
amount of literary scientific labour involved in such a task and hoped that Baron
von Mueller would complete their joint enterprise.

The Reverend W. B. Clarke was nearing the end of his long and productive
life devoted to the geology of Australia. When he died in his eighty-first year in
1878, he was putting the last touches on his work which recapitulated his views
and the controversies on the geology of the sedimentary deposits in New South
Wales. Despite both the relationships that had been observed in the dis-
tributions of some of the flora and fauna of different parts of the world and the
developments in palaeontology taking place at a rapid rate at the end of the
nineteenth century, there was still no idea that the explanations might lie in the
movements of the continents. It was another thirty-seven years before Wegener
made his suggestion.

It was only eight years since Pasteur in France had done his famous work in
isolating the two disease bacilli from the silkworm; now he was studying the
fermentation process in beer. Koch in Germany had developed the techniques
for the isolation and study of bacteria generally and two years later was to
isolate the bacillus of anthrax. The development of bacteriology at this time
had special significance for the Linnean Society because it was undoubtedly an
important influence in bringing about the later establishment of the Macleay
Bacteriologist.

This then was the intellectual atmosphere of that time, almost exactly one
hundred years ago, when William Macleay was dining with the scientists of HMS
Challenger. He was to have a busy year. He had just been back to his station
at Kerarbury, ninety miles west of Wagga near Hay. Both there and in the
country through which he journeyed he had been collecting with the curator
whom he had recently appointed, George Masters—Mr. Masters, as he always
called him—who was to build up the William Macleay part of the collection that
had been initiated by Macleay’s cousin and uncle. Macleay was thoroughly
familiar with the low, level skyline of the Hay plain, broken only by the river red
gums and the trees which the squatters had planted near the banks of the slow
moving river of muddy water and sluggish creeks. Macleay was now fifty-four
years old. He had arrived in Sydney in March 1839. Soon after his arrival he
had gone to the station on the lower Murrumbidgee as a squatter with his cousin
and there they had had several difficult years. It was not until 1844 with the
recovery in the price of wool—as happens today—that they began to have
financial success. When he went into politics he decided that he could manage
with occasional visits to his property and a permanent residence in the city of
Sydney. There the young man of thirty-seven met the much younger woman

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72 THE FIRST HUNDRED YEARS

of eighteen, the daughter of Sir Edward Deas Thomson, and they were married
in 1857. A difference of some nineteen years in their ages and William Macleay’s
preoccupation with science probably account for the way in which they apparently
agreed that she would go to social functions and he would engage himself in
science. About a month before the fishing picnic Macleay had been taken,
somewhat unwillingly, to the University of Sydney for the ceremony in which
his father-in-law, as Chancellor, accepted on the University’s behalf the Macleay
Collection, which had been started by his uncle, Alexander Macleay, and cousin,
William S. Macleay (Fletcher, 1920). His own real interests are shown by the
extract from his diary: “ Susan attended a ball at Government House. [I]
worked alone on the collection till well after midnight. The octopus which
Masters brought in this morning is interesting—a new variety. Unfortunately
must go out to dinner tomorrow ”’.

When his cousin, William 8S. Macleay, died in 1865 it was possible for William
and Susan to move into the Macleay residence—Elizabeth Bay House—a gracious,
beautiful piece of architecture standing in some sixty acres of land. It had been
built for Uncle Alexander in the early 1830’s to the designs of John Verge. It
was a gentleman’s residence with an unrivalled situation, before the city it once
neighboured engulfed it. There he lived under quite luxurious circumstances—
he took his butler on his first fishing picnic in 1874. However, he had by no
means lost the tough resilience which had helped him make a success of the
outback life, as is shown by an incident in December 1864. Accompanied only
by the boy who was driving the buggy, he came upon the bushrangers Gilbert,
Hall and Dunn (one of whom had recently killed a policeman). They were
holding up the Goulburn coach, several teams and a number of travellers. Rifle
in hand Macleay raised the siege and routed the bushrangers. Ten years later
he was awarded a gold medal for ‘‘gallant and faithful services’ during the
bushranging period (Fletcher, 1893).

By the 1870’s William Macleay was beginning to enjoy his preferred style of
life, embracing the thrill of the discovery of a new species and the discussion of
matters of scientific interest with his friends. His idea of social iife was the
Sunday whisky party where intense and eager discussion was part of the gathering.
Captain Arthur Onslow, Dr. Cox and others who were to become the founders of
the Linnean Society of New South Wales were among those who regularly attended
these weekly discussions.

On the 13th October Macleay wrote in his diary : ‘‘ Dr. Alleyne and Captain
Stackhouse are trying to get up a Society of Natural History. I hope they may
succeed. Such a Society, embracing all branches of Natural History, and
issuing a Monthly Magazine, ought to be both useful and successful’? (Walkom,
1925). Thus we find the earliest mention of the Linnean Society. This was
followed by a notice of a preliminary meeting to be held in the boardroom of the
Public Library on 29th October 1874. There, Macleay was elected as President,
Sir William Macarthur as Vice-President, Captain Stackhouse as Honorary Sec-
retary and H. H. Burton Bradley Esq. as Honorary Treasurer of the provisional
committee.

The President-elect steered the business of the meeting away from the
suggestion that it should be called the Macleayan Society, and the name of the
great Linnaeus was commemorated. The President-elect, Mr. W. J. Stephens
and Dr. Alleyne were chosen to draft the rules. Some eleven years later Macleay
was to say : “‘ Our rules state that the Society is for the promotion of the study of
Natural History in all its branches. Few people, I suspect, are aware of the
wide significance of that sentence ”’ (quoted by Fletcher, 1893). The first annual
meeting of this Society was held in Lloyd’s Chambers in 362 George Street on
13th January 1875.

When we look at this first hundred years of our Society, how far can we say
with confidence that Macleay’s hopes for its success have been fulfilled? I

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RUTHERFORD N. ROBERTSON 73

shall speak for a little while on the kinds of things for which the Society has been
responsible and let you judge whether you would regard it as a successful venture.
I shall finish my lecture by speculating a little on what might be done in the next
hundred years.

There is no doubt whatever that the Linnean Society immediately became a
focal centre for scientific activities in the areas of biology and geology in this
country. It had significance that went wider than the city of Sydney and the
colony of New South Wales, for before long it was truly a national body. This is
illustrated splendidly by reference to The Macleay Memorial Volume (Fletcher,
1893), published in 1893 to record by scientific publications (which he would
have appreciated) the great regard in which the biologists and geologists of
Australia then held Sir William who had died in 1891. This volume contains
thirteen different scientific contributions ; half the contributors were or were to
become Fellows of the Royal Society of London, in those days the greatest
distinction that could be conferred upon a British scientist. Those writing these
papers were the leaders in biology and geology in Australia and New Zealand :
the Professor of Biology in the University of Melbourne, Baldwin Spencer, the
Challis Professor of Biology in the University of Sydney, William Haswell, the
Professor of Geology in Canterbury College, Christchurch, F. W. Hutton and
two great men of science, Professor J. T. Wilson, who was then Professor of
Anatomy in the University of Sydney and was to become Professor of Anatomy
in the University of Cambridge, and Mr. C. J. Martin (later Sir Charles), one of
the great physiologists of the British Empire. The famous Baron von Mueller
contributed Notes on an Undescribed Acacia from New South Wales and Description
of a New Hakea from Eastern New South Wales. The important and interesting
contribution of the Palaeontologist to the Australian Museum and Geological
Survey of New South Wales, Mr. RK. Etheridge, described the weapons and
implements of the Alligator Tribe, Port Essington, North Australia. You will
recall that Port Essington failed as a settlement and was later succeeded by
Port Darwin. A section in Mr. Etheridge’s work describes the ‘‘ trumpets ”’ of
these tribes, made from hollow bamboo—what we call the didgeridoo. Inciden-
tally the bamboo was not indigenous but had been planted by the Port Essington
settlers and used by the tribes.

There is no doubt that the success of the Society is reflected in the greatness of
the individuals who were associated with it in its early years. The importance
of its ability to publish the papers that described the Australian natural history
can hardly be overestimated. The first volume of the Proceedings started in
the two years 1875/76. Thereafter a volume was published every year so that
in the first fifty years of the Society’s existence, forty-nine volumes of the Pro-
ceedings were published. In those days the publication of these descriptions and
of the speculations of our scientific colleagues were, relatively speaking, much
more important than they would be today. First, opportunities for publication
were more restricted than those now available in the wide range of current journals.
Second, the exchange of information between scientific colleagues was much more
difficult. Bear in mind that travel was slow and arduous, opportunities for
conferences infrequent, that the radio did not exist and that consequently the
printed word was the pre-eminent form of communication. Jn addition to the
Proceedings, the Abstracts of Proceedings played an important part in the days
when there were so many new forms of naturally occurring objects to be described.

The Society quickly built up a library, some of the early parts of it being due
to the generosity of Sir William Macleay. Some of it was unfortunately lost in the
disastrous fire of 1882 when the Garden Palace was destroyed. The Garden
Palace, which had been built for the International Exhibition of 1879, had been
retained as a building both for government offices and society premises. The loss
of records, library and microscopes to the Linnean Society as a result of that
fire was nothing short of crippling.

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74 THE FIRST HUNDRED YEARS

It is not my intention, in a short talk, to attempt anything like another
historical account of the Linnean Society ; it has been written before (Walkom,
1925). I would, however, like to refer to its success by some examples of the
types of men and women who have been associated with it. My selection will
show you where my admiration lies.

In 1882 a young Welshman, Tannatt William Edgeworth David, arrived in
Australia to take up an appointment as Assistant Geologist-Surveyor in govern-
ment employ. It was not long before he was a member of the Linnean Society.
In 1883 he surveyed the tin fields of the New England district and in 1886, the
coal-bearing strata of the lower Hunter Valley in which he discovered the hitherto
unsuspected existence of the important coal seams which became the great Mait-
land Coalfield. In 1891 he was appointed Professor of Geology and Physical
Geography in the University of Sydney and assumed his place as a leader in the
scientific world of Australia. He was President of our Society from 1893 to
1895 and continued his active geological work first with the expeditions to the
coral reefs to ascertain the actual structure of coral, incidentally confirming the
theory of Charles Darwin about reef formation, and later with trips to the moun-
tains to consider the effects of glaciation on the Australian continent. Towards
the end of 1907 he joined the Shackleton expedition to the Antarctic and not
only led the climbing party on the first ascent of Mt. Erebus, no mean feat since
it was 13,000 feet in those regions, but also led the small party of himself, Mawson
and Mackay to the exact site of the south magnetic pole, which they reached on
16th January 1909. There, following the instructions of Shackleton (1911),
they hoisted a Union Jack on a small flagpole and uttered the words, as David’s
diary says: ‘‘ I hereby take possession of this area, now containing the magnetic
pole, for the British Empire”’. At the same time he fired the trigger of the
camera, and the string can be seen vibrating in the photograph of the three men
at the south magnetic pole. What is not seen is that the flag which they had
taken with them was one which had to be made from pieces of cloth from various
garments since there was no spare flag on the Nimrod. That flag we now proudly
possess among the archives in the Academy of Science. They started that long
trek on 25th September and were picked up again by the ship Nimrod on 4th
February. I have quite a personal interest in this distinguished member and
former President of our Society who for me was the greatest character of our
many great people. When I was a student he used to lecture occasionally,
telling the story of the Antarctic adventures to the students’ Science Society, and
I still remember two things quite clearly. The first was that he said that people
who were concerned about his welfare used to say to his wife: ‘‘ Why ever did
you let that poor thin Professor David go to the Antarctic? . He will feel the
cold so terribly!” ‘‘ But ’’, said David, ‘‘ I didn’t feel the cold because it’s not
so important to be fat as to have a good circulation, and I have a good circu-
lation!’ The other thing is his amusing description of how, when they were in
their tent and were in great concern about whether they would be picked up by
the ship, they heard the ship’s gun. The tent door was not made to fit three
large men in arctic dress emerging together and they collided and trampled over
each other in so doing. After picking themselves up they had not run very far
outside the tent when Mawson disappeared twenty feet down a crevasse ; for-
tunately he was not hurt. It is no wonder that David was welcomed home by a
crowded function in the Great Hall of the University of Sydney.

Finally, as though David had not done enough already in his life, he became
Chief Geologist to the British Army supervising the practical application of
geological knowledge to tunnelling, water-supply and locating deep dugouts
throughout the British sphere of operations, winning the D.S.O. in 1918.

One of the characteristics of natural history in the nineteenth century was
that the work—including wo1k of great distinction—was often carried out by

PROCEEDINGS OF THE LINNEAN Society or New SourH Watszs, Vou. 99, Part 1

RUTHERFORD N. ROBERTSON 75

people to whom it was more of a hobby than a profession. I have already
mentioned Macleay and the clergyman-geologist W. B. Clarke. In this century
too there were people of similar devotion to natural history—the Reverend
H. M. R. Rupp working on the orchids, Mr. G. A. Waterhouse ‘ the butterfly
man ’’, Mr. 8. J. Copland the herpetologist who used to be the night crime reporter
for the Sydney Morning Herald. I want to mention one name both because of a
personal reminiscence and to make an important point. Mr. A. H. 8S. Lucas was
Headmaster of Sydney Grammar School, a member of our Council for many
years and President from 1907 to 1909 ; he was the only person in Australia then
to undertake a systematic investigation of our marine algae. He was still on the
Council when I, a brash young recent graduate on a Science Research Scholarship
of $120 a year, became an applicant for a Linnean Macleay Fellowship which paid
the princely sum of $400 a year. Some of my friends at Sydney University were
speculating on my chances when I said that I was not very hopeful since I was
interested in experimental science and the Linnean Council was composed of a
number of old fogies mainly interested in ‘‘ stamp-collecting ’’ biology. I quite
forgot that the grandson of A.H.S. Lucas was among those present. Not long
after I was awarded a Fellowship I had a message via the grandson, in the form
of a question : ‘‘ Did I think the old fogies sometimes did the right thing?’ A
splendid rebuff for my youthful arrogance and intolerance. I am glad that I
have lived long enough to realise that we need people like Clarke, Waterhouse,
Lucas, Copland and others. The reason for the paucity of knowledge of biology
and geology in district details in this country, compared with such knowledge in
Britain, is that such people are so thin on the ground. The ordinary member
here, with his notes and publications, has an important role to play in the estab-
lishment of this kind of science.

I could not attempt to enumerate the professional scientists whose out-
standing work has been associated with our Society. Among those I personally
admire greatly are Professor W. L. Waterhouse of the Faculty of Agriculture,
University of Sydney, and Dr. W. R. Browne, who in addition to his distinguished
research did so much for the Society.

The Linnean Macleay Fellowships, of course, have made an enormous
contribution to the scientific development of this country ; at one time they
provided opportunities for research which were without parallel anywhere else in
Australia. The first Macleay Fellow, H. I. Jensen, appointed in 1905, was to
write The Soils of New South Wales. The second, J. M. Petrie, held his appoint-
ment for eigateen years during which time he raised a family, one member of
which was A. H. K. Petrie, the first plant physiologist at the Waite Agricultural
Institute and one of our three pioneers of plant physiology. Father Petrie was
really a plant chemist with biochemical leanings ; he wrote on nitrogen met-
abolism in plants, a subject which his son was to develop strongly. The third
and fourth Fellows later became professors and the fifth, Dr. A. B. Walkom, in
addition to being a palaeontologist, became an organiser for science who left not
only this Society but also ANZAAS, the former Australian National Research
Council and various other bodies permanently indebted to him. Among the
other distinguished Fellows were R. J. Tillyard, the entomologist who became a
Fellow of the Royal Society and first Chief of the CSIRO Division of Entomology,
the late Professor P. D. F. Murray and Dr. I. M. Mackerras, now one of our most
revered senior scientists in the Australian Academy. Others are Dr. Lilian
Fraser, Professor N. A. Burges, now a Vice-Chancellor, Dr. Ilma Pidgeon, Dr.
Germaine Joplin, Dr. June Lascelles, and so on. Present Council members,
Dr. Mary Hindmarsh, Dr. Elizabeth Pope and Professor T. G. Vallance have
been Fellows.

Incidentally, you will notice a high proportion of female names among those
I have mentioned. I am pleased to say that although the Linnean Society

PROCEEDINGS OF THE LINNEAN SociETy OF NEw SoutH WALES, Vou. 99, Part 1

76 THE FIRST HUNDRED YEARS

began in the tradition of being a male preserve one hundred years ago, it was
reasonably early in breaking through that awful and now almost incomprehensible
barrier, the exclusion of women. Women were admitted as associate members in
1885 and as ordinary members in 1909 and the first woman Linnean Macleay
Fellow, Miss Irwin Smith, was appointed in 1919.

The Macleay Bacteriologist was the far-sighted scheme of an imaginative
man—William Macleay—who endowed the position as a long term appointment.
Only four people have held that position ; Mr. Greig Smith for 29 years, Dr. H. L.
Jensen for 19 years, Dr. Y. T. Tchan (including the period since the arrangement
was made with the University of Sydney) for 14 years and now Dr. K. Y. Cho
since 1969. This endowment showed the importance that William Macleay
placed upon the coming development of bacteriology as a science. When you
think now of all in health, genetics and biology that depends upon our knowledge
of the bacterial kingdom—one might almost call it the Third World of biology—
you can appreciate the imagination of this man who was associated with our
Society in the beginning. If Macleay were to return I think he would be par-
ticularly pleased both with his intelligent guess about the importance of bac-
teriology and with the success of the Macleay Bacteriologists.

I am equally sure that there are some areas where he would be bitterly
disappointed in the stewardship of those who have succeeded him. I shall
mention only two: one a scientific disappointment, the other a social failure.
You will recall that Mr. Bentham completed his work on Flora Australiensis in
1878. He specifically stated that the writing of a flora is an ever-changing,
never-ending activity and expressed the intention of forming a supplementary
volume to contain an account of the new species added to existing knowledge.
Now one hundred years later, despite the excellent work in the regional floras
and local handbooks, we are still talking about the need for a revised Flora Aust-
raliensis. True, we have made a modest beginning with some work started by
the Academy but we hope it will be stepped up to more vigorous activity by the
newly formed Biological Resources Survey sponsored by the Federal Government
and under the chairmanship of Dr. D. F. Waterhouse one of our members and
the bearer of a well-known Linnean Society name.

I find the social failure quite frightening. In 1866 William Macleay was a
member of a Select Committee of Parliament in New South Wales to report on
‘“‘ the distress at present existing in the working classes’. In 1972 the Federal
Government appointed Professor Ronald Henderson to report on ‘‘ poverty in
Australia’ and that survey is still going on. How can we as a community fail
to be ashamed that amidst all our affluence we should still be talking about
surveying poverty instead of having made some social arrangement to eliminate
it? JI would not like to have to make excuse for this to William Macleay.

What shall we make of the next hundred years? I cannot be a prophet
except of one thing—the next hundred years will be different from the last. The
only constant and predictable thing in our world is change but the nature of any
change is unpredictable and often startlingly unexpected. Perhaps we can
define how our starting point for a hundred years from now differs most from
the starting point in 1874. If I asked you to define the most important difference
I would get a variety of answers. I am not going to enumerate the obvious
changes like those in travel and communication. JI want to draw your attention
to one vital change. Sir William Macleay said that he thought few people
realised the significance of the study of natural history. Natural history was of
course the generic phrase used in those days for the biological and earth sciences.
Now one hundred years later that statement is no longer true.

A most important difference is that the awareness of science in the whole
community is changing and changing rapidly. 1874 saw an age in which only a
favoured few escaped illiteracy and scientific ignorance. The ordinary person

PROCEEDINGS OF THE LINNEAN Society or New Sourn Wass, Vou. 99, Part 1

RUTHERFORD N. ROBERTSON (7

today realises how large a part science and its daughter, technology, fathered by
Science and mothered by industry, plays in his life. A recent survey by the
Sydney Morning Herald recorded that 49° of those surveyed have great faith in
Science, 39% had some faith and only 2% had no faith. Today the student in
our high schools has courses in science aimed not merely at expanding his know-
ledge of scientific facts and phenomena but also at giving him the experience of
thinking for himself as a scientist thinks. We therefore have some hope that
problems will be solved by rational thinking. Some of the unrest among the
young is undoubtedly associated with reaction against the older irrational thought
and superstitions. There is always some backlash in such circumstances too,
Since it is more comfortable for some people to cling to superstition than to
face realistic reason. There is increasing realisation that science is an all-per-
vading influence in society. It must be considered not a thing apart but rather
as an intrinsic part of our way of thinking—a topic I have dealt with at some
length in my Oscar Mendelsohn Lecture (Robertson, 1974). We have therefore
an almost explosive increase in the awareness of the methods of science and
scientific thought. On the international scene we are witness to the greatest
revolution of all time, in China, a country which I visited earlier this year. There,
children at school from the age of 6 to 17 are receiving a scientific education
specifically directed towards the elimination of the older superstitions with which
that country was so rife. It will be interesting to see how this great attempt at
egalitarian socialism based on modern science and technology develops. What a
contrast to China as it was one hundred years ago ; it has emerged from feudalism
only in the last twenty-five years.

What will these new starting points in our attitudes to science mean to the
Linnean Society? Farlier I referred to the importance of the non-professional
Scientist in contributing to our knowledge of the flora, fauna and geology of this
country. I hope that with our increasing leisure and increasing scientific back-
ground there will be a corresponding increase in the number of people associating
themselves with bodies like the Linnean Society, better to describe and explain
our Australian surroundings in scientific terms. In this way we shall fill the
many gaps in our knowledge of this vast continent in which we have nothing
like the detailed scientific exploration seen in Britain and other countries of
Europe, where every district is so well-known and described scientifically. In
this area the non-professional scientist, equipped with the improved background
in scientific training now provided by the high schools, will be able to make a
first-rate contribution. I hope that the Linnean Society will continue to act
both as a stimulus for such workers and as a publisher of their results. I think
there is great room for encouragement of these kinds of activities. I have, for
example, just read a quite brilliant little paper summarising personal research
on plant chemistry done by a fifteen-year-old high school student.

On the international scene all of us will have an important role to play. We
Shall be developing science in this country parallel with the urgent need for the
development of scientific knowledge and, more particularly, scientific attitudes in
our neighbouring countries. I gather that the Council of the Linnean Society,
which together with the Royal Society of New South Wales has acquired the new
building that will become a Science Centre in Sydney, has given consideration to
its possible international significance. Many national scientific bodies are
expected to have their headquarters in this Science Centre and there will be
opportunities to provide for international scientific visitors and to make contact
with the science in the developing countries—an extremely important activity
for Australian scientists.

I would have another hope and expectation for the next hundred years:
that some colourful and devoted personalities such as those I have described will

PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WALES, VoL. 99, Part 1

73 THE FIRST HUNDRED YEARS

emerge, like Macleay, David, Walkom, W. L. Waterhouse, Browne and others,
who are just as interesting though with activities suited to their own times,
whose work and personalities will be talked about a hundred years hence.

I have been much involved in recent discussions of national science policy
which means among other things the best use of science and technology for the
welfare of the community. Inevitably the question is: For what purpose shall
we use our science and technology ? What are the national goals to which we
would apply our science? Politicians and others are extremely coy in defining
national goals. If you came from outside and looked at our behaviour with
detached objectivity, you might well judge our goals to be those of selfish sectional
interest or of personal self interest, each man for himself or his group. I wonder
if it is too much to expect that some time in the next century we as a nation shall
evolve socially to the point of having a united major goal—the provision of
equality in welfare and opportunity for each member of our community. If so,
science and technology could properly be used to that end. If we were to attempt
the fulfilment of this ideal and particularly if we succeeded we should be in a
much stronger position to supply help in science and technology to developing
countries. Until something of this kind is done we shall never be free of the
tensions between the underprivileged and those living in luxury, whether we are
talking about individuals or nations, and science and technology will not be used
for the advantage of all.

I have tried to show how our Society, starting one hundred years ago, was a
creation of its time and of the social conditions in which it was born. I should
think that the members of the Linnean Society of the next hundred years who
will keep changing with changing thought will be increasingly aware of the fact
that science and technology have not been properly used. Even if the Society
as such is not involved, certainly as individuals, members will become increasingly
involved and will take leading roles in seeing to the proper use of science. For
if the world does not increasingly aspire to the ideals of the use of rational thought
and scientific attitudes, what can save it ?

I owe the Linnean Society a great deal. It was partly to express my grati-
tude to the Society that I accepted the honour of being Macleay Lecturer for a
second time. In the next hundred years I hope that there will be many who
will gain as much from the Society as I have done in my forty years of association.
If I were here when the Society celebrates its bicentenary, I would accept an
invitation to be Macleay Lecturer for a third time so that I could wish the Society
as much good fortune for its third hundred years as I now sincerely wish it tor the
second.

References

Brreuw, A., and Macminnan, D. S., 1962.—The Sydney Scene 1788-1960. Melbourne : Melbourne
University Press.

FiLetcuer, J. J. (ed.), 1893.—The Macleay Memorial Volume. Sydney : Linnean Society of New
South Wales.

Frercuer, J. J., 1920.—The Society’s Heritage from the Macleays. (Part 1) Proc. Linn. Soo.
N.S.W., 45 : 567-635.

Fietcuer, J. J., 1929.—The Society’s Heritage from the Macleays. (Part 2) Proc. Linn. Soc.
N.S.W., 54: 185-272.

Macieay, W.—Diary for 1874. MS, Mitchell Library, Sydney.

Macminian, D.S., 1957.—A Squatter went to Sea. Sydney : Currawong Publishing Company.

Rogertrson, R. N., 1974.—Survival—Science or Superstition? (Third Oscar Mendelsohn
Lecture) Search 5 (5): 191-197.

SHACKLETON, E., 1911.—The Heart of the Antarctic. London: Heinemann.

Tworenny, R. E. N., 1883.—Town Life in Australia. London: E. Stock.

Watxom, A. B., 1925.—The Linnean Society of New South Wales. Historical Notes of Its Furst
Fifty Years. (Jubilee Publication) Sydney: Australasian Medical Publishing Company.

PROCEEDINGS OF THE LINNEAN Sociery or New SourH WALEs, VoL. 99, Part 1

A NEW SPECIES OF TYPHLODROMUS SCHEUTEN (ACARINA :
PHYTOSEIIDAE) FROM APPLE IN AUSTRALIA

E. ScuicHa* AND G. DOSSET

[Communicated by M. Casimrr]

[Accepted for publication 12th December, 1973]

Synopsis

Typhlodromus (Typhlodromus) helenae n. sp. (Phytoseiidae) is described from apple in New
South Wales, Australia. It is closely related to 7. (Z'.) pyri Scheuten (=. (Z'.) taliae Oudemans)
and to 7. (f.) setubali Dosse.

INTRODUCTION

Phytoseiid mites from various plants in Australia have been studied and
described by Womersley (1954), but none of these has been found on apple.
The species described below was collected from apple trees at Bathurst, New
South Wales, and could be mistaken for Typhlodromus (Typhlodromus) pyri
Scheuten, a predaceous mite found on apple trees in Europe, Canada and New
Zealand. A key reference is Dosse (1961). Measurements listed are the
mean +standard deviation of 20 specimens in microns, if not otherwise indicated.

Genus TYPHLODROMUS Scheuten
Typhlodromus Scheuten, 1857, Arch. Naturgesch., 23: 104-112.

Typhlodromus (Lyphlodromus) helenae, uv. sp.
(Figs 1-7)

Diagnosis. This species is closely related to Typhlodromus pyri Scheuten
(=T. (f.) tiliae Oudemans) and to Typhlodromus setubali Dosse. According to
Nesbitt (1951) and Chant (1958, 1959) it keys out to Typhlodromus pyri. However,
it is distinguished from 7. pyri by having one tooth (instead of two teeth) on
the digitus mobilis and from 7. setubali by having three pores, instead of four,
on the dorsal shield. It is distinguished from 7. pyri and T. setubali by having
setae M, smooth. The spermeatheca differs from that of T. pyri by having
chitinized lips, a broader macroduct and a cervix without a thin neck ; it is
similar to that of 7. setubali but the macroduct is thicker.

The males of the three species differ in length and width and the male of the
new species can also be distinguished from that of 7. pyri by having four pre-anal
setae instead of three. The spermatodactyl is rodlike but does not terminate
with an ‘‘ inverted boot ” as in T. pyri (Nesbitt, 1951). However, its posterior
part ends with a suctorial dise similar to that of 7..setubalt.

* N.S.W. Department of Agriculture, Agricultural Research Station, Bathurst, New South
Wales, 2795.

{ Institute for Plant Protection, University of Stuttgart-Hohenheim, West Germany.

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80

PROCEEDINGS

A NEW PHYTOSEHD MITE

Fig. 1. Dorsal shield of female.

oF THE LINNEAN Socirry or NEw Sourn Wass, Vou. 99, Part 1

EK. SCHICHA AND G. DOSSE 81

Fig. 2. Venter of female. -

PROCEEDINGS OF THE LINNEAN SocreTy oF NEw Soutu Wates, Von. 99, Part 1

82 A NEW PHYTOSEDD MITE

Female

Dorsum: Length, 333-+-7, width at L, 179+38. Dorsal shield (Fig. 1) is
imbricated and bears 17 pairs of setae distributed as follows: six dorsal, two
median (one anterior and one posterior), six prolateral and three postlateral.
These setae measure

D,=21+1, D,=15+1, D,;=14+1, D,=15+1, D;=16+1, D,=5+1,

L,=22+1, L,=15+1, Lg=18+2, L,=19+2, L;=21+42, L,=23+41,

L,=26+2, Le=29+1, Lo=47 +3,

M,=15-+40, M,=29 +2.

The setae are of moderate length, except D,. Setae L, are equal to the
distance between their bases and the bases of the setae L,; all other setae are
shorter than the distances between their bases and the bases of the setae following

Figs 3, 4. 3. Chelicera of female. 4. Spermatheca (of female).

next in the series. Setae M, are longer than the distance between their bases and
the bases of setae Ls. All setae are smooth except setae Ly, which are slightly
serrated. There are three pairs of large pores present. One pair is near Ly,
one is postlateral to L, and one pair is between M, and Ly. In addition there are
10 pairs of small pores present.

Sublateral setae S,=20+2 and S,=21-+1 in length, situated on the dorsal
interscutal membrane.

Peritremes extend slightly beyond setae L,.

Venter (Fig. 2): Sternal shield, length 71-+-1, width 71-+1, is smooth and
bears two pairs of setae. Sternal setae III are situated on small irregularly
shaped shields. Setae IV are on irregularly shaped metasternal shields. Genital
shield, length 70 +-1, width 68 +0, is normal with a pair of genital setae. Ventri-
anal shield, length 101-+3, width 102-+-4 (10 measurements), is shield-shaped
with four pairs of preanal setae, and the posterior part is reticulated.

PROCEEDINGS OF THE LINNEAN Society or New Sourn Waxes, Vou. 99 Part 1

E. SCHICHA AND G. DOSSE 83

Chelicera (Fig. 3): Fixed digit, length 38+2 (11 measurements), of the
chelicera bears four teeth and a pilus dentilis. The movable digit, length 40 +1,
bears one tooth on its inner margin.

Spermatheca (Fig. 4): Atrium is knee-like and bears strongly chitinized
lips. The funnel-like cervix has a diameter of 17 where it joins the vesicle.
Cervix plus vesicle are pear-shaped.

Legs: Only basitarsus of leg IV (Fig. 5) bears a macroseta, 46-+2 in length.

Ny)

Yh} | | k

!/ NU
Hy We

Figs 5, 6. 5. Leg IV of female. 6. Ventri-anal shield of male.

Male

Dorsum: Imbricated, length 253+9, width at L, 165+4; the chaetotaxy
and the shape of the setae resemble those of the female. The setae are, however,
relatively shorter.

D,=16+1, D,=12+1, D,=11+1, D,=12+1, D,;=13+1, D,=4+0,

L,=18+1, L,=11-+1, Lzs=138+1, L4=15+1, L,=18+1, L,=19+1,

L,=21+1, Lg=21 +2, L,=37 +1,

M,=11+1, M,—25 +1,

S,=15+1, S,=15+41.

Venter: Sternal shield with five pairs of setae and three pairs of pores.
Ventri-anal shield (Fig. 6), length 100+2, width 142+4 (10 measurements),
bears four pairs of preanal setae and three pairs of small pores.

Spermatodactyl (Fig. 7): Rod-like and terminating in a small suctorial disc.

Type. Female holotype from Bathurst Agricultural Research Station,
New South Wales, Australia (W. G. Thwaite) on leaves of Granny Smith apple
trees, 1970. In South Australian Museum, Adelaide, Australia.

PROCEEDINGS OF THE LINNEAN Society oF New SoutH WALES, Vou. 99, Part I

84 A NEW PHYTOSEDD MITE

Paratypes. Hight females and eight males taken from the same locality
as the holotype. Four females plus four males at Biological and Chemical
Research Institute, Rydalmere; two females plus two males at Institute for
Plant Protection, University of Stuttgart-Hohenheim, West Germany; two
females plus two males at South Australian Museum, Adelaide.

Fig. 7. 7. Spermatodactyl (of male).

References

Cuant, D. A., 1958.—Immature and adult stages of some British Phytoseiidae Berl., 1916

(Acarina). J. Linn. Soc., Zool., 43 : 599-643.
. 1959.—Phytoseiid mites (Acarina: Phytosetidae). Part II. A taxonomic review of

the family Phytoseiidae, with descriptions of 38 new species. Can. Ent. 91, Suppl. 12: 45-166.

Doss, G., 1961.—Zur Klaerung der Artenfrage von T'yphlodromus (Typhlodromus) pyri Scheuten
1857 (=T. tiliae Oud. 1929) und TLyphlodromus (Typhlodromus) setubalt n. sp. (Acar.,
Phytoseiidae). Z. angew. Ent., 48: 313-323.

Nessitt, H. H. J., 1951.—A taxonomic study of the Phytosetidae (Family Laelaptidae) predaceous
upon Tetranychidae of economic importance. Zool. Verh., Leiden, 12: 1-64.

WoMERSLEY, H., 1954.—Species of the subfamily Phytoseiidae (Acarina : Laelaptidae) from
Australia. Aust. J. Zool., 2: 169-191.

PROCEEDINGS OF THE LINNEAN Society oF New Sours Wares, Vou. 99, Part 1

The Proceedings, of the Linnean Society of New South Wales contain original papers
relative to “the Science of Natural History in all its branches’’, and reports of business
transacted at the meetings of the Society.

NOTES AND INSTRUCTIONS FOR AUTHORS

Original papers may be submitted for consideration for publication in the Society’s
Proceedings. Submission of a manuscript must imply that the contents are original and
have not been published previously in the same, similar, or abbreviated form, and that no
similar paper is being or will be submitted for publication elsewhere until it has been
either (a) not accepted for publication by the Society or (b) withdrawn by the author.

Papers submitted should embody the results of a significant piece of work. Interim
reports, serialized studies and student theses should not be submitted.

Papers are accepted for publication (or otherwise) by the Council of the Society
after taking into consideration advice received from at least one referee. The decision
of Council is final.

Once accepted for publication a paper becomes the property of the Society, and
copyright is vested in the Society. Reproduction of the author’s synopsis in abstracting
journals is authorized.

A paper may be withdrawn by the author at any time prior to acceptance.

Papers may be submitted by members or non-members of the Society but a paper
submitted by a non-member must be communicated by a member of the Society, and
will not ordinarily be considered for acceptance before July in each calendar year.

SUBMISSION OF MANUSCRIPTS

Manuscripts should be forwarded to the Secretary, The Linnean Society of New
South Wales, 157 Gloucester Street, Sydney, N.S.W. 2000.

Text and illustrations must be submitted in duplicate. The illustrations may be
duplicated by any acceptable clear method and the duplicate set need not necessarily
be identical in size with the originals. The author should retain a third copy for
checking proofs. If a paper is not accepted for publication, the original copy of the
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Proceedings, Volume 99, Part 1, 1974

CONTENTS

Annual General Meeting :
Report on the Affairs of the Society for the Year
Elections

Balance Sheets
Domrow, R. Miscellaneous mites from Australian vertebrates. 1-48

McLzeAN, R. A. The rugose coral genera Streptelasma Hall, Grewingkia

Dybowski and Calostylis Lindstrém from the Lower Silurian of New .

South Wales

PEPPERELL, J., and Griee, G. A Labyrinthodont trackway from the Mid-
Triassic near Sydney, New South Wales

RICHARDSON, L. R. A new troglobitic quadrannulate land-leech from
Papua (Hirudinoidea : Haemadipsidae s.1.)

ROBERTSON, R. N. Sir William Macleay Memorial Lecture, 1974. ‘A
Society of Natural History. I hope they may succeed.” The first
hundred years

ScuicHA, H., and Dosse, G. A new species of Typhlodromus Scheuten
(Acarina : Phytoseiidae) from apple in Australia

Notes and instructions for authors

36

54

57

69

79

85

Proceedings of the
Linnean Society .__..
of New South Wales

Issued 12th February, 1975

VOLUME 99
PART 2
No. 438

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, 1974-75

President
T. G. Vallance, B.Se., Ph.D.

Vice-Presidents

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Secretary
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Elizabeth C. Pope, M.Sc., C.M.Z.S.*

* Resigned 28/8/74. + Elected 20/11/74.
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The Society’s Headquarters are in Science House, 157 Gloucester Street, Sydney,
N.S.W. 2000, Australia

FOUR NEW DAMSELFISHES (POMACENTRIDAE) FROM THE
SOUTHWEST PACIFIC

GERALD R. ALLEN*

(Plates III and IV)

[Communicated by Gilbert P. Whitley]

[Accepted for publication 20th February 1974]

Synopsis

Four new species of damselfishes which were collected at Indonesia, Melanesia, Lord Howe
Island and New South Wales are described. Glyphidodontops niger n.sp. was taken at Cape Nelson,
New Guinea. It is closely allied to G. biocellatus, G. glaucus, G. leucopomus, and G. unimaculatus,
all of which are widely distributed Indo-Pacific species. G. notialis n.sp. was collected at New
Caledonia, Lord Howe Island and New South Wales. It bears a close resemblance to G. rapanut
from Easter Island. G. talbott n.sp. was taken at Indonesia, Solomon Islands,’ New Hebrides,
Fiji Islands and the Great Barrier Reef. It is related to G. traceyi from the Marshall and Caroline
Islands. Pomacentrus albimaculus n.sp. was collected at Madang, New Guinea. It is allied to
P. amboinensis from the western Pacific.

INTRODUCTION

The damselfishes (family Pomacentridae) are one of the largest families of
tropical reef fishes, both in number of species and number of individuals. It is
estimated that there are at least 250 species, approximately 150 of which inhabit
the seas of Australia and Melanesia. Most of our knowledge of the group in
this region is based on the insufficient works of Bleeker (1877) and Fowler and
Bean (1928). In 1973 the author made extensive collections of damselfishes at
Fiji Islands, New Caledonia, New Hebrides, Solomon Islands, New Britain, New
Guinea, the Great Barrier Reef of Australia, and Lord Howe Island. These
collections contain 132 species, a substantial increase compared with 85 species
reported by de Beaufort (1940) for the region which includes Malaysia, Indonesia,
Melanesia and Australia. At least 12 of the species are new, including four
species, Glyphidodontops niger, G. notialis, G. talboti and Pomacentrus albimaculus,
which are described herein.

The Pomacentridae represent an extremely diverse lineage. The majority
of Indo-West Pacific members are usually assigned to five major genera : Amphi-
prion Bloch and Schneider, Dascyllus Cuvier, Chromis, Cuvier, Abudefduf Forskal
and Pomacentrus Lacépede. However, recent studies by the author (unpublished)
revealed the necessity for a comprehensive revision of certain genera, in particular
Abudefduf and Pomacentrus. In the South Pacific region 93 of 143 species have
been assigned to these two ‘ genera.’? Under this scheme those individuals
with a serrate preopercle margin are assigned to Pomacentrus and those with a
smooth margin to Abudefduf. This arrangement leads to a multiplicity of
problems as there are certain groups of species which display intermediate
criteria, being characterised by a crenulate or weakly serrate preopercle. A

* Western Australian Museum, Francis Street, Perth, W.A., 6000.

PROCEEDINGS OF THE LINNEAN SociETY oF NEw SoutH WaAtEs, Vou. 99, Part 2

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PROCEEDINGS oF THE LINNEAN Society or- New SourH Wauus, Vou. 99, Part :2

GERALD R. ALLEN 89

revision of the Pomacentridae of the western Pacific, (Allen, in press a), reveals
there is justification for splitting these groups into at least 13 genera. Surpris-
ingly, most of these divisions were recognised nearly a century ago by Bleeker
(1877). Abudefduf, as presently recognised, is particularly diverse and obviously
constitutes a polyphyletic assemblage. This group is separable into the following
genera: Abudefduf Forskal, Amblyglyphidodon Bleeker, Glyphidodontops Bleeker,
Hemiglyphidodon Bleeker, Paraglyphidodon Bleeker, Plectroglyphidodon Fowler
and Ball and two additional genera which will be described by the author. The
genus Pomacentrus, as currently recognised, is divisable into Amblypomacentrus
Bleeker, Dischistodus Bleeker, Hupomacentrus Bleeker and Pomacentrus Lacépede.
Most of the species pertaining to this group will remain in the latter genus.

METHODS OF COUNTING AND MEASURING

The methods of counting and measuring are the same as those described by
Allen (1972), except the length of the dorsal and anal spines are measured proxi-
mally from the base of the spine rather than from the point at which the spine
emerges from the scaly sheath. Measurements were made with needle-point dial
calipers to the nearest one-tenth millimetre. Standard length is abbreviated as
SL. The fraction 4 which appears in the dorsal and anal fin ray formulae refers
to a bifureate condition of the last ray.

The counts and proportions which appear in parentheses under the descrip-
tion section for each species apply to the paratypes when differing from the:
holotype. A summary of counts for the dorsal, anal and pectoral fin rays, tubed.
lateral line scales and gill rakers on the first gill arch is presented in Tables 1 and 2.

Type specimens have been deposited at the following institutions : Australian.
Museum, Sydney (AMS) ; Bernice P. Bishop Museum, Honolulu (BPBM) ; British
Museum (Natural History), London (BMNH); Muséum National d’Histoire
Naturelle, Paris (MNHN) ; United States National Museum of Natural History,
Washington, D.C. (USNM).

DESCRIPTIONS
Glyphidodontops niger, ni. sp.
(Plate ITI, Fig. a)

Holotype. AMS 1.16708-004, 41-8 mm SL, coll. G. Allen, 26/V/1972, near
mouth of Tufi Inlet, Cape Nelson, New Guinea (9°05’S, 149°19’E), 1-2 m, quin-
aldine and dipnets.

Paratypes. AMS 1.16708—005, 5 specimens, 17-2-30-6 mm Sh, collected
with the holotype.

Diagnosis. A species of Glyphidodontops with the following combination of
characters : body depth 2-0 to 2-1 in SL; dorsal spines 13 ; horizontal scale rows
between middle of lateral line and dorsal fin base 14; predorsal scales reaching
level of nostrils ; preorbital, suborbital and inferior limb of preopercle naked ;
teeth biserial; membranes between dorsal spines slightly incised; colour in
alcohol uniformly dark brown with intense black spot covering pectoral base,

Description. Proportional measurements for the holotype and two selected
paratypes are expressed as percentage of the standard length in Table 3.

Dorsal rays XTII, 124; anal rays II, 13 ; pectoral rays 17 ; pelvic rays I,.5;
branched caudal rays 13 ; gill rakers on the first arch 21 (20) ; tubed lateral line
scales 17 ; vertical scale rows from upper edge of gill opening to base of caudal

PROCEEDINGS OF THE LINNEAN Society oF New SoutH WALES, VoL. 99, Part 2

90 DAMSELFISHES FROM THE SOUTHWEST PACIFIC

TABLE 3
Measurements of the holotype and two paratypes of Glyphidodontops niger
(expressed in thousandths of the standard length)

NT NP eo ss

Holotype Paratype Paratype
Characters AMS AMS AMS
1.16708-004 1.16708-005 1.16708-005

Senda bse an A 41-8 29-3 25-5
Greatest body depth .. ae 498 478 502
Greatest body width .. Ee 203 184 192
Head length .. oy ee 316 338 353
Snout length .. ae wy 81 82 82
Eye diameter .. Mis oh 107 116 126
Interorbital width a 96 92 90
Least depth of caudal peduncle 156 154 149
Length of caudal peduncle .. 103 116 102
Snout to origin of dorsal fin .. 431 420 439
Snout to origin of anal fin 694 683 675
Snout to origin of pelvic fin .. 407 399 416
Length of dorsal fin base Ay 615 517 620
Length of anal fin base a 254 266 235
Length of pectoral fin ile 311 273 255
Length of pelvic fin. .. ate 323 341 306
Length of pelvic spine 184 198 204
Length of lst dorsal spine .. 52* 68 82
Length of 7th dorsal spine .. 163 143 153
Length of last dorsal spine .. 172 160 165
Length of longest soft dorsal

ray . 220 205 216
Length of Ist anal spine Ne 86 79 82
Length of 2nd anal spine as 165 147 157
Length of longest soft anal ray 239 205 216
Length of middle caudal rays 251 215 235

* damaged.

fin 27 ; horizontal scale rows from base of dorsal fin to terminal lateral line scale
(exclusive of dorsal base sheath scales) 1 ; from lateral line to anal fin origin 8 ;
predorsal scales about 20, extending to level of nostrils ; teeth biserial, those of
outer row slender, close-set with flattened tips numbering about 36 in each jaw,
inner row of buttress teeth about 4 width of outer row teeth.

Body ovate, laterally compressed, the greatest depth 2-0 (2-0 to 2-1) in the
standard length. Head profile conical, the head length contained 3-2 (2-8 to
3-2) times in the standard length ; snout 3-9 (4-1 to 4-3), eye diameter 2-9 (2-8
to 2-9); interorbital width 3-3 (3-7 to 3-9), least depth of caudal peduncle 2-0
(2-2 to 2-4), length of caudal peduncle 3-1 (2-9 to 3-6), of pectoral fin 1-0 (1-2 to

4), of pelvic fin 1-0 (1-0 to 1-2), of middle caudal rays 1-3 (1-5 to 1-6), all in
the head length.

Single nasal opening on each side of snout; mouth oblique, terminally
located ; lateral line gently arched beneath dorsal fin, terminating one scale row
below base of 13th dorsal spine ; preorbital, suborbital, snout tip, lips, chin, and
isthmus naked ; remainder of head and body scaled; scales finely ctenoid ;
preopercle scale rows 2, inferior and posterior limb broadly naked ; small sheath
scales covering basal 1/2 to 2/3 of membraneous portions of dorsal, anal, and
caudal fins ; margin of preorbital, suborbital, preopercle and opercle entire.

Origin of dorsal fin at level of third tubed lateral line scale ; spines of dorsal
fin gradually increasing in length to last spine, length of first dorsal spine 3-9
(4-3 to 4-5), of seventh dorsal spine 4-3 to 4-5 (paratypes only, holotype dam-
age d), of last dorsal spine 1-8 (2-1), of longest soft dorsal ray 1-4 (1-6 to 1-7), of

PROCEEDINGS OF THE LINNEAN Sociery or New SoutrH WAuzEs, Vou. 99, Part 2

GERALD R. ALLEN 91

first anal spine 3-7 (4-3), of second anal spine 1-9 (2-3), of longest soft anal ray
1-3 (1:6 to 1-7), all in the head length ; caudal fin emarginate, lobes rounded ;
pectoral fins slightly rounded.

Colour of holotype in alcohol: Head and body dark brown, breast paler,
median fins dark brown except posterior portion of soft dorsal and caudal fins
pale ; pelvic fins brown ; pectoral fins pale with black spot covering base and axil.

Colour of holotype in life: Uniformly bluish-black with intense black spot
covering pectoral base and axil ; pectoral fin translucent.

Individuals under about 30 mm SL are uniformly bluish-black with a series
of neon blue stripes on the sides, one per horizontal scale row.

Remarks. G. niger belongs to a species complex which includes G. biocellatus
(Quoy and Gaimard), G. glaucus (Cuvier), G. leucopomus (Cuvier), and G. unim-
aculatus (Cuvier). These species are sympatric inhabitants of tropical Indo-
West Pacific reefs, usually frequenting depths of less than 2-3 metres. They are
similar with regards to counts and general body shape, but the adults at least are
easily distinguished, both alive and in preservative, on the basis of colour. G.
biocellatus is characterised by a brown ground colour and single whitish bar,
several scales wide, just behind the pectoral fin. The juveniles of this species,
as well as those of G. leucopomus and G. wnimaculatus are similar to adult G.
leucopomus which are predominately yellow with a blue stripe extending from the
eye to the soft dorsal fin and with one or two occelli on the posterior part of the
dorsal. The “ amabilis ”’ variety of G. leucopomus, an ecological variant discussed
by Allen (in press, 6), is characterised by two or three whitish bars on a dark
ground colour. G. glaucus is uniformly pale. Adults of G. unimaculatus are
predominantly brownish with a small black spot at the base of the hindmost
dorsal rays. The head and anterior portion of the body is abruptly pale in some
individuals (‘‘ hemimelas”’ variety). G. niger is the only uniformly blackish
member of the complex and also the only species which possess a black pectoral
base and axil.

G. niger was encountered at Cape Nelson, New Guinea and Goodenough
Island, D’Entrecasteaux Islands. It was relatively common at both localities in
areas of moderate surge between 0-5 and 2 metres. The largest individuals
encountered were approximately 50 mm SL. The stomach contents of two
paratypes indicate an algal diet.

The specific name refers to the characteristic black colouration of the body.

Glyphidodontops notialis, n. sp.
(Plate III, Fig. 5)

Holotype. AMS 1.17402-008, 49-2 mm SL, coll. J. Randall and B. Goldman,
17/11/1973, off Phillip Point, Lord Howe Island (31°32’S, 159°04’E), 15 m,
rotenone.

Paratypes. UORD HOWE ISLAND: AMS 1.17357-016, 50-8 mm SL,
same data as holotype except coll. G. Allen, B. Goldman and W. Starck, 5/1/1978 ;
AMS 1.17358-012, 6 specimens, 44-7-61-3 mm SUL, coll. G. Allen and party,
5/11/1973, off west side of Mt. Lidgbird, 20-25 m, rotenone ; AMS I.17374-007, 2
specimens, 49-3 and 52-0 mm SUI, coll. F. Talbot and party, 18/11/1973, on outer
reef between Rabbit Island and Erscott’s Passage, 16 m, explosives; AMS
1.17377-014, 3 specimens, 28-2-51-5 mm SUL, coll. G. Allen and J. Randall,
24/11/1973, North Islet, 25 m, rotenone; BMNH 1973.10.17.4-9, 6 specimens,
51-0-64-5 mm SLI, coll. F. Talbot and party, 25/11/1973, on outer reef between
Rabbit Island and Erscott’s Passage, 12 m; BPBM 15935, 18 specimens, 45 -4—
64:2 mm SL, same data as preceding paratypes except coll. 26/11/1973 ; BPBM
15936, 16 specimens, 43 -5-62-8 mm SL, same as preceding except coll. 19/IT/1973 ;
MNHN 1973-59, 3 specimens, 45-0-52-4 mm SL, same data as holotype except
coll. G. Allen, 15/17/1973, quinaldine ; USNM 211292, 4 specimens, 24-8—-51-9 mm

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutH WALES, VoL. 99, Part 2

92 DAMSELFISHES FROM THE SOUTHWEST PACIFIC

SL, same data as holotype except coll. J. Randall and party, 26/11/1973 ; NEW
CALEDONIA: AMS I.,17466-001 59-0 mm SL, coll. G. Allen, 15/V1/1973,
Puetege Reef, off southeast tip of New Caledonia, 20 m, multi-prong spear.

Diagnosis. A species of Glyphidodontops with the following combination of
characters : body depth 2-3 to 2-5 in SL; dorsal spines 13 ; horizontal scale rows
between middle of lateral line and dorsal fin base 14; predorsal scales reaching
level of nostrils ; suborbital and inferior limb of preopercle scaly ; teeth uniserial,
dorsal outline uniform without incisions between spines ; colour mostly bluish-
black (dark brown to blackish in preservative).

Description. Proportional measurements for the holotype and two selected
paratypes are expressed as percentage of the standard length in Table 4.

TABLE 4

Measurements of the holotype and two paratypes of Glyphidodontops notialis
(expressed in thousandths of the standard length)

Holotype Paratype Paratype

Characters AMS AMS USNM

1.17402-008 1.17466-001 211292

Standard length (mm) ae 49-2 59-0 39-0
Greatest body depth .. BG 403 436 400
Greatest body width .. aH 180 186 180
Head length .. Bie a 298 285 308
Snout length .. Ao ee 63 71 72
Eye diameter .. a Bes 103 92 121
Interorbital width : 78 75 80
Least depth of caudal peduncle 151 144 146
Length of caudal peduncle .. 103 109 115
Snout to origin of dorsal fin .. 308 336 346
Snout to origin of anal fin .. 608 612 669
Snout to origin of pelvic fin .. 384 398 426
Length of dorsal fin base a 660 644 577
Length of anal fin base 5 298 312 277
Length of pectoral fin ae 333 310 308
Length of pelvic fin .. of 346 305 318
Length of pelvic spine se 166 159 167
Length of lst dorsal spine .. 55 54 56
Length of 7th dorsal spine. . 151 153 144
Length of last dorsal spine .. 157 166 151

Length of jonpeet soft dorsal

ray : 359 322 235
Length of lst ‘anal spine a 84 80 87
Length of 2nd anal spine .. 141 125 128
Length of longest soft anal ray 340 322 244
Length of middle caudal rays 241 224 231

Dorsal rays XIII,14% (XIJI,134 to 154); anal rays 11,154 (11,154 to 164) ;
pectoral rays 17 (16 to 18) ; pelvic rays 1,5 ; branched caudal rays 13 ; gill rakers
on the first arch 20 (20 to 23); tubed lateral line scales 17 (16 to 18) ; vertical
scale rows from upper edge of gill opening to base of caudal fin 28 ; horizontal
scale rows from base of dorsal fin to terminal lateral line scale (exclusive of dorsal
base sheath scales) 13; from lateral line to anal fin origin 9; predorsal scales
about 18 to 20, extending to level of nostrils ; teeth uniserial, incisiform, about
40 to 44 in each jaw.

30dy elongate, laterally compressed, the greatest depth 2-5 (2-3 te 2-5) in
the standard length. Head profile conical, the head length ereninedl 3°4 (3°3
to 3-5) times in the standard length ; snout 4-7 (4:0 to 4-6), eye diameter 2-9
(2-6 to 3-1) ; interorbital width 3-8 (3-7 to 3-9), lez By depth of caudal peduncle
2-0 (2-0 to 2-1), length of caudal peduncle 2-9 (2-5 to 2-7), of pectoral fin 1-0
(0-9 to 1-0), of pelvic fin 0-9 (0-9 to 1-0), of middle ceitidal rays 1-2 (1-2 to 1-4),
all in the head length.

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GERALD R. ALLEN 93

Single nasal opening on each side of snout; mouth oblique, terminally
located ; lateral line gently arched beneath dorsal fin, terminating 14 scale rows
below base of second to third soft dorsal ray ; preorbital, snout tip, lips, chin and
isthmus naked ; remainder of head (including suborbitals) and body sealed ;
head scales cycloid, remainder finely ctenoid ; preopercle scale rows 2 with
additional row of scales on inferior limb ; small sheath scales covering basal 1/3
to 2/3 of membraneous portions of dorsal, anal and caudal fins ; margin of pre-
orbital, suborbital, preopercle and opercle entire.

Origin of dorsal fin at level of second tubed lateral line scale; spines of
dorsal fin gradually increasing in length to last spine ; length of first dorsal spine
5:5 (5-3 to 6:3), of seventh dorsal spine 2-0 (1:9 to 2-3), of last dorsal spine
1-9 (1:7 to 2-1), of longest soft dorsal ray 0-8 (0-9 to 1-3), of first anal spine
3:6 (3:2 to 4-1), of second anal spine 2-1 (2-0 to 2-4), of longest soft anal ray
0-9 (0:9 to 1-3), all in the head length ; caudal fin forked, the lobes filamentous ;
pectoral fins pointed.

Colour of holotype in alcohol : Ground colour of head and body dark brown,
scale centres slightly lighter ; breast and abdomen tannish ; median fins dark
brown to blackish ; pelvic fins dusky with blackish tips ; pectoral fins mainly
translucent with black wedged-shaped spot covering upper portion of base, axil
of fin pale.

Colour of holotype in life: Head and anterior portion of body blue-grey
grading to blackish behind pectoral fins; dorsal, anal, and basal portion of
caudal fin bluish-black ; dorsal and anal fins with bright blue margin ; distal half
of caudal fin blue ; pelvic fins dusky with bluish anterior edge, tips black ; pectoral
fins light blue with blackish spot superiorly at base.

Remarks. G. notialis most closely resembles G. rapanwi (Greenfield and
Hensley, 1970) which is endemic to Easter Island. Both species are elongate
(depth 2-3-3-0 in SL), possess elongate posterior dorsal and anal rays and are
predominately blackish in colouration with blue fin margins. G. rapanui differs
by having biserial dentition, a slightly deeper body (average depth 2-7 in SL),
and a colour pattern which includes blue spots on the head and anterior portion
of the body, a pale caudal fin and peduncle, and a black spot covering the base
of the pectoral fin. The counts for the fin rays, tubed lateral line scales and gill
rakers are virtually identical in these two species.

G. notialis is one of the most common damselfishes outside the lagoon at
Lord Howe Island. The habitat, which is essentially subtropical, consists largely
of rocky canyons, ledges and boulders covered with algae. The species was
observed at depths ranging from 7 to 45 metres. G. rapanui was observed by the
author to inhabit a similar environment at Easter Island. The stomach contents
of several paratypes of G. notialis contained algae and crustacean remains (largely
copepods).

This species is named notialis (Gr. ‘* southern ”’) in reference to its geographical
distribution. It is the most southerly occurring species of Glyphidodontops and
a common inhabitant of Lord Howe Island, the world’s southernmost coral reef.

Glyphidodontops talboti, n. sp.
(Plate III, Fig. c)

Holotype. AMS 1.15684—-004, 39-8 mm SL, coll. F. Talbot and party,
1/XII/1969, One Tree Island (23°30'S, 152°05’E), Capricorn Group, Great Barrier
Reef, 30 m, explosives.

Paratypes. GREAT BARRIER REEF: AMS 1.15486—038, 34:2 mm SL
coll. H. Choat, 31/1/1966, Heron Island, Capricorn Group, 9 m, rotenone ; AMS,
1.16479—-001, 39-7 mm SL, coll. W. Starck, 1/VII/1972, Pixie Reef, off Cairns,
Queensland, 12 m, multi-prong spear; AMS I1.15647-001, 36-5 mm Sh, same
data as holotype except coll. 9/X/1968, 13-5 m; AMS 1.15637-009, 31-8 mm

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94 DAMSELFISHES FROM THE SOUTHWEST PACIFIC

SL, same data as holotype except coll. 5/X/1967, 23 m; INDONESIA : USNM
209745, 2 specimens, 22-7 and 34:7 mm SL, coll. V. Springer and M. Gomon,
11/1/1973, off Tandjung Suli, Ambon Island, Moluccas, 2-3 m, rotenone ; USNM
210026, 2 specimens, 32-3 and 36:5 mm SL, coll. V. Springer and M. Gomon,
19/1/1973, offshore west of Tandjung Namatatuni, Ceram, Moluccas, 14-16 m,
rotenone ; SOLOMON ISLANDS : AMS 1.17479-002, 38-8 mm SH, coll. G. Allen,
5/V11/1973, off Tassafaronga Point, about 7 nautical miles west of Honiara,
Guadaleanal, 20 m, quinaldine; BMNH 1973.10.17.10-12, 3 specimens, 32-5-—
42-0 mm SL, coll. J. Randall and B. Goldman, 18/VII/1973, off southwest corner
of Savo Island, 10 m, rotenone ; MNHN 1973-60, 30-8 mm SH, coll. G. Allen and
J. Randall, 30/VI1/1973, off Tanavulu Point, northern end of Florida Island,
20-30 m, rotenone; NEW HEBRIDES: AMS I1.17472-001, 3 specimens, 28 -5—
38:2 mm SU, coll. G. Allen, 22/V1/1973, off Malapoa Point, near Vila, Efate,
10 m, rotenone; AMS I[.17473-001, 25:7 mm SU, coll. G. Allen and D. Popper,
23/V1/1973, about 34 nautical miles south of Vila, Efate, 13 m, rotenone ; AMS
T.17475-001, 39-8 mm SUL, coll. G. Allen, D. Popper and W. Starck, 25/V1/1973,
Pula Iwa Reef (17°02’S, 168°17’E) off Emae Island, 25 m, rotenone; FIJI
ISLANDS : AMS 1.17464- 001, 36-8 mm SH, coll. G. Allen, 8/V1I/1973, outer reef
at mouth of Nukulau Pass, near Suva, Viti Levu, 30 m, multi-prong spear ;
BPBM 14564, 37-0 mm SUL, coll. J. Randall, 11/II1/1972, outside Mbengga
barrier reef, off Yanutha Island, 10 m, rotenone.

TABLE 5

Measurements of the holotype and two paratypes of Glyphidodontops talboti
(expressed in thousandths of the standard length)

Holotype Paratype Paratype
Characters AMS AMS AMS
1.15684-004 1.15647-001 1.15637-009

Standard length (mm) Rs 39-8 36-5 31-8
Greatest body depth .. ae 445 460 459
Greatest body width .. bs 190 209 203
Head length .. ae ss 299 315 315
Snout length .. ae oe 75 66 66
Eye diameter .. as Bs 108 115 132
Interorbital width Be ae 73 71 76
Least depth of caudal peduncle 151 145 148
Length of caudal peduncle .. 111 134 129
Snout to origin of dorsal fin .. 392 419 393
Snout to origin of analfin .. 673 669 670
Snout to origin of pelvic fin 387 411 409
Length of dorsal fin base a5 575 597 576
Length of anal fin base Ke 211 211 226
Length of pectoral fin as 299 329 280
Length of pelvic fin... ot 324 321 305
Length of pelvic spine te 176 197 204
Length of Ist dorsal spine... 58 74 60
Length of 8th dorsal spine... 138 153 154
Length of last dorsal spine .. 91 93 91
Length of longest soft dorsal

ray .. Be Me ae 216 214 186
Length of Ist anal spine 37 50 58 66
Length of 2nd anal spine 2 151 181 182
Length of longest soft anal ray 219 216 208
Length of middle caudal rays 302 329 320

———

Diagnosis. A species of Glyphidodontops with the following combination of
characters : dorsal spines 13 ; horizontal scale rows between middle of lateral line
and dorsal fin base 14 ; predorsal scales reaching level of nostrils ; suborbital and
inferior limb of preopercle scaly ; teeth biserial; membranes between dorsal

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GERALD R. ALLEN 95

spines deeply to moderately incised ; colour largely pale purple (red-brown in
preservative) with prominent black spot near base of hindmost dorsal spines.

Description. Proportional measurements for the holotype and two selected
paratypes are expressed as percentage of the standard length in Table 5.

Dorsal rays XIII,114 (XIIT,11 to 114) ; anal rays IT,12 (11,114 to 13) pectoral
rays 16 (15 to 16) ; pelvic rays I,5 ; branched caudal rays 13 ; gill rakers on the
first arch 19 (18 to 20) ; tubed lateral line scales 16 (14 to 16) ; vertical scale rows
from upper edge of gill opening to base of caudal fin 27 ; horizontal scale rows
from base of dorsal fin to terminal lateral line scale (exclusive of dorsal base
Sheath scales) 14; from lateral line to anal fin origin 9 ; predorsal scales about
17 (17 to 20), extending to level of nostrils ; teeth of jaws biserial, at least ant-
eriorly, those in outer row somewhat spatulate with rounded tips, about 34 in
lower jaw and 38 in upper jaw ; teeth of inner row slender, about 4 width or less
of outer row teeth.

Body ovate, laterally compressed, the greatest depth 2-2 (2-2 to 2-3) in the
standard length. Head profile conical, the head length contained 3-3 (3-2 to
3:3) times in the standard length ; snout 4-0 (4-3 to 5-5), eye diameter 2-8 (2-4
to 3-0), interorbital width 4-1 (4-0 to 4-6), least depth of caudal peduncle 2-0
(1-8 to 2-2), length of caudal peduncle 2-7 (2-2 to 2-5), of pectoral fin 1-0 (1-0 to
1-2), of pelvic fin 0-9 (1:0 to 1-1), of middle caudal rays 1-0 (1-0 to 1-1), all in
the head length.

Single nasal opening on each side of snout; mouth oblique, terminally
located ; lateral line gently arched beneath dorsal fin, terminating 1} scale rows
below base of last dorsal spine ; suborbital scaled ; snout tip, lips, chin, isthmus,
and most of preorbital naked ; remainder of head and body scaled ; scales finely
ctenoid ; preopercle scale rows 2 with additional row of scales on inferior limb ;
small sheath scales covering about basal 1/2 to 1/3 of membraneous portion of
dorsal, anal, and caudal fins; margin of preorbital, suborbital, and bones of
opercle series entire (edge of preopercle weakly crenulate in some specimens).

Origin of dorsal fin at level of fourth tubed lateral line scale ; spines of dorsal
fin gradually increasing in length to about eighth spine, remaining spines gradually
decreasing in length ; length of first dorsal spine 5-2 (4-3 to 5-8), of eighth dorsal
Spine 2-2 (1-9 to 2-2), of last dorsal spine 3-3 (2-8 to 3-4), of longest soft dorsal
ray 1-4 (1-5 to 1-8), of first anal spine 6-0 (3-8 to 5-5), of second anal spine 2-0
(1-7 to 1-9), of longest soft anal ray 1-4 (1-4 to 1-7), all in the head length ;
caudal fin emarginate ; pectoral fins pointed.

Colour of holotype in alcohol: Head and body generally reddish-brown ;
predorsal region, side of head and ventral surface of body suffused with yellow
to tan; prominent black spot about size of eye near base of hindmost dorsal
spines (9th to 13th), extending about halfway out on fin; anal papilla black ;
fins pale, slightly yellowish.

The 31-8 mm paratype from One Tree Island is faded due to the action of
the preservative and is basically pale tan with the black dorsal spot and black
anal papilla. The other paratypes from Melanesia and Queensland are similar
to the holotype in colouration.

Colour in life : Most of body faded purple ; predorsal region and head yellow ;
prominent black spot about 1-14 size of eye at base of hindmost dorsal spines ;
anal papilla black; dorsal and anal fins usually translucent, yellow basally ;
pectoral and caudal fins translucent to pale yellow ; pelvic fins yellow.

Remarks. G. talboti is closely related to G. traceyi from the Marshall and
Caroline Islands. Morphologically the two species are nearly identical, but they
differ significantly in colouration. The colour of several G. traceyi observed at
Palau was as follows : head and body mostly purple grading to yellow at level of
soft dorsal fin origin ; anterior half of anal fin and spinous dorsal fin purple with
black spot about size of eye at base of hindmost dorsal spines ; soft dorsal fin,

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96 DAMSELFISHES FROM THE SOUTHWEST PACIFIC

caudal fin, and posterior half of anal fin yellow ; pectoral fins translucent, pelvic
fins dark purplish-brown. ‘Thus, the colour pattern of G. traceyi is nearly the
opposite of that exhibited by G. talboti, which is primarily yellow anteriorly and
purplish posteriorly. Two specimens of G. traceyi which I collected at Palau
also differ in having 104 soft dorsal rays, one less than the usual number for
G. talbott. However, Woods and Schultz (1960) gave a range of 10 to 12 rays
for 12 specimens from the Marshall Islands. They included traceyi in the genus
Pomacentrus on the basis of the finely serrate preopercle. However, they stated
that the serrations are so small they may be seen only with difficulty unless
magnified and some specimens have the serrations only near the angle of the
preopercle. Likewise, some specimens of G. talbott exhibit a weakly crenulate
preopercle margin, usually confined to that portion near the angle. Examination
of the general morphology of these species indicates a relationship to the genus
Glyphidodontops rather than Pomacentrus. G. rollandi from the Indo-Australian
Archipelago is a closely related form. G. talboti and G. traceyi appear to have
allopatric distributions. The latter species is known from the Marshall and
Caroline Islands and probably occurs at the Philippines. G. talboti is apparently
confined to Indonesia, Melanesia and Queensland. In addition to the localities
listed for the types, it was observed at Madang, New Guinea, at Rabaul, New
Britain, at Egum Atoll (Solomon Sea), at the D’Entrecasteaux Islands and at
Osprey Reef (Coral Sea).

Abudefduf bonang (non Bleeker) of Fowler and Bean (1928) is probably
synonymous with G. traceyi judging from their description of specimens from the
Philippines. The counts, measurements, size range (35-53 mm SL) and general
colour pattern are in agreement. Bleeker’s (1853 ; 1877) description and figure
of Paraglyphidodon bonang indicate that it is a distinct species, differing from
G. traceyi and G. talbott in several respects. The counts given by Bleeker which
include 15 to 16 dorsal rays, 13 to 14 anal rays and 19 to 20 pectoral rays, represent
significant differences. In addition, P. bonang grows to a much larger size. The
largest specimen recorded by Bleeker was 150 mm TUL, nearly three times the
length of the author’s largest G. talbott. The author has observed hundreds of
individuals of G. traceyt and G. talbott in the sea, none of which exceeded about
60mm Th. The 36-5 mm SL paratype of G. talboti is a mature female with eggs,
also indicative of the small size of the species. Specimens of P. bonang were
reported by Bleeker (1877) and de Beaufort (1940) as having a white-edged
dark brown ocellus near the middle of the dorsal fin. According to Bleeker, this
ocellus may disappear in large specimens. G. traceyi and G. talboti, by contrast,
do not have a pale ring around the dark dorsal spot and the spot persists in
individuals of all sizes.

The status of Glyphidodon mutabilis Cartier (1874), relegated to a synonym
of Abudefduf bonang (non Bleeker) by Fowler and Bean (1928) is uncertain. The
soft dorsal and anal fin ray counts are too high for either G. traceyi or G. talbott.

G. talboti was generally observed on outer reef slopes in depths ranging from
about 8 to 30 metres. It is a solitary dweller which hovers a short distance
above the bottom while feeding. The stomach contents of two paratypes
indicate a diet consisting largely of pelagic tunicates, copepods, isopods and a
small amount of algae.

Named in honour of Dr. Frank H. Talbot, Director of the Australian Museum,
the collector of the holotype.

Pomacentrus albimaculus, 0. sp.
(Plate IV, Figs. a and Db)

Holotype. AMS 1.16691—-006, 61-7 mm SL, coll. G. Allen, 7/IV/1972, near
main wharf at Madang, New Guinea (5°14’S, 145° 45’), 16 m, multi-prong spear.

Paratypes. AMS 1.16691-007, 3 specimens, 45-5-56-5 mm SL, collected
with the holotype.

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GERALD R. ALLEN 97

Diagnosis. A species of Pomacentrus with the following combination of
characters : body depth 1-9 to 2-0 in SL; dorsal spines 13 ; preorbital and sub-
orbital naked ; prominent notch between preorbital and suborbital ; inferior edge
of suborbital denticulate ; teeth biserial anteriorly ; colour grey (brown in preser-
vative) with darker scale edges giving overall reticulated appearance, prominent
white spot on upper half of caudal peduncle when alive.

Description. Proportional measurements for the holotype and two selected
paratypes are expressed as percentage of the standard length in Table 6.

Dorsal rays XIII,144; anal rays 11,154 (11,144 to 154) ; pectoral rays 17;
pelvic rays 1,5 ; branched caudal rays 13 ; gill rakers on the first arch 21 (20 to
21) ; tubed lateral line scales 17 (16) ; vertical scale rows from upper edge of gill
opening to base of caudal fin 28 ; horizontal scale rows from base of dorsal fin to
terminal lateral line scale (exclusive of dorsal base sheath seales) 14 ; from lateral
line to anal fin origin 9 ; predorsal scales about 18, extending to level of nostrils ;
teeth conical with rounded tips, biserial at front of jaw, about 34 primary teeth
in the lower jaw and 40 in the upper.

TABLE 6

Measurements of the holotype and two paratypes of Pomacentrus albimaculus
(expressed in thousandths of the standard length)

Holotype Paratype Paratype
Characters AMS AMS AMS
I.16691-006 1.16691-007 1.16691—007

Standard length (mm) sis 61-7 56:5 45-5
Greatest body depth .. Si 519 492 517
Greatest body width .. ote 195 188 198
Head length .. is ae 300 301 306
Snout length .. BE Sh 75 81 75
Eye diameter .. me oe 97 96 106
Interorbital width oe a 73 78 84
Least depth of caudal peduncle 146 149 143
Length of caudal peduncle .. 97 94 88
Snout to origin of dorsal fin .. 381 372 374
Snout to origin of anal fin... 622 639 666
Snout to origin of pelvic fin .. 376 372 391
Length of dorsal fin base Xe 674 646 633
Length of anal fin base ae 327 303 286
Length of pectoral fin Be 292 294. 297
Length of pelvic fin .. Ne 324 324 328
Length of pelvic spine as 175 184 199
Length of Ist dorsal spine. . 86 71 88
Length of 7th dorsal spine .. 154 154 171
Length of last dorsal spine .. 173 168 185
Length of longest soft dorsal

ray .. ae AS, ae 259 258 286
Length of Ist anal spine oe 73 73 73
Length of 2nd anal spine he 165 161 174
Length of longest soft anal ray 256 264 259
Length of middle caudal rays 248 265 2M

Body ovate, laterally compressed, the greatest depth 1-9 (1-9 to 2-0) in the
standard length. Head profile conical, the head length contained 3-3 times in
the standard length ; snout 4-9 (3-7 to 4:1), eye diameter 3-1 (2-9 to 3-1);
interorbital width 4-1 (3-7 to 3-9), least depth of caudal peduncle 2-1 (2-0 to
2:1), length of caudal peduncle 3-1 (3-2 to 3-5), of pectoral fin 1-0, of pelvic fin
0-9, of middle caudal rays 1-2 (1-1), all in the head length.

Single nasa] opening on each side of snout; mouth oblique, terminally
located ; lateral line gently arched beneath dorsal fin, terminating 1} scale rows
below base of second soft dorsal ray ; preorbital, suborbital, snout tip, lips, chin,
and isthmus naked ; remainder of head and body scaled ; scales finely ctenoid ;

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98 DAMSELFISHES FROM THE SOUTHWEST PACIFIC

preopercle scale rows 3; small sheath scales covering basal 1/2 to 2/3 of mem,
braneous portions of dorsal, anal, and caudal fins ; margin of preorbital, suborbital-
and preopercle denticulate ; margin of opercle entire.

Origin of dorsal fin at level of third tubed lateral line scale ; spines of dorsal
fin gradually increasing in length to last spine ; length of first dorsal spine 3-5
(3-5 to 4-3); of seventh dorsal spine 1-9 (1-8 to 2-0); of last dorsal spine 1-7
(1:7 to 1-8); of longest soft dorsal ray 1-2 (1-1 to 1-2); of first anal spine 4-1
(4-1 to 4-2), of second anal spine 1-8 (1:8 to 1-9), of longest soft anal ray 1-1 (1-1
to 1-2), all in the head length ; caudal fin slightly emarginate ; pectoral fins
pointed.

Colour of holotype in alcohol: Head and body brown, scale margins and top
of head darker ; upper portion of caudal peduncle pale tan ; dorsal and anal fins
brownish with one or two pale submarginal bands, distal tips of membranes
between dorsal spines black and small dark spot in middle of soft dorsal between
9th and 11th ray ; caudal fin brown ; pectoral and pelvic fins pale tan ; small
blackish spot on opercle near lateral line origin and diffuse dark spot on upper
pectoral base, axil of pectoral pale.

Colour of holotype in life: Head and body grey with dusky scale edges
giving overall reticulated appearance ; median fins pale grey with one or two
pale blue submarginal bands on dorsal and anal fins ; pelvic fins pale blue-grey ;
pectoral fin translucent with blackish spot at upper portion of fin base ; small
bluish spot on opercle near origin of lateral line ; prominent white spot on upper
half of caudal peduncle.

The small dark spot between the 9th and 11th soft dorsal rays is a remnant
of the juvenile ocellus which disappears with growth. The ocellus is generally
present on specimens under about 40 mm SL.

Remarks. P. albimaculus belongs to the subgenus Pseudopomacentrus
Bleeker and is closely allied to the following species from the western Pacific :
P. amboinensis Bleeker, P. grammorhynchus Fowler, P. melanochir Bleeker, P.
melanopterus Bleeker, P. moluccensis Bleeker, P. nigromanus Weber, P. nigro-
marginatus Allen, P. philippinus Evermann and Seale, and P. tripunctatus Cuvier
and Valenciennes. The relationships between most of the members of this
group have been discussed by Allen (1973). P. albimaculus appears to be most
closely related to P. amboinensis, but this latter species differs by having a slightly
higher gill raker count (22 to 24), and is primarily yellowish in colouration. P.
nigromargimatus has a similar reticulated pattern and grey ground colour, but
differs by having a black margin on the soft dorsal and caudal fins, an intense
black spot covering the entire pectoral fin base and axil, and usually 14 or 15
tubed lateral line scales.

P. albimaculus was only encountered at Madang during a cruise which
included several stops along the northeastern coast of New Guinea from Madang
to Samarai. It was relatively common in the harbour and lagoon in turbid areas
of limited visibility at depths between 10 and 20 metres. It shares this habitat
with P. amboinensis. The stomach contents of the holotype consisted mainly
of algae.

The specific name albimaculus refers to the characteristic white spot on the
upper caudal peduncle.

ACKNOWLEDGEMENTS
The author is indebted to Dr. Walter A. Starck II, Research Associate of
the B. P. Bishop Museum, Honolulu, and the Australian Museum, Sydney, for
providing facilities aboard his research vessel Hl Torito. Thanks are also due to
Dr. Frank H. Talbot, Director, and to the Board of Trustees of the Australian
Museum, who provided funds for the completion of this study. Dr. John E.

PROCEEDINGS OF THE LINNEAN Society or New SourH WALES, Vou. 99, Part 2

Proc. LINN. Soc. N.S.W., Vol. 99, Part 2 PLATE I

Fig. a. Glyphidodontops niger, holotype, 41-8 mm SL, Cape Nelson, New Guinea.

Fig. b. Glyphidodontops notialis, holotype, 49:2 mm SL, Lord Howe Island.

Fig. c. Glyphidodontops talboti, paratype, 37:0 mm SL, Fiji Islands,

Proc. LINN. Soc. N.S.W., Vol. 99, Part 2 PLATE IV

Fig. a. Pomacentrus albimaculus, holotype, 61:7 mm SL, Madang, New Guinea.

Fig. b. Pomacentrus albimaculus, approximately 70 mm SL, underwater flash photograph
taken at Madang, New Guinea, near the type locality.

GERALD R. ALLEN 99

Randall provided the initial stimulus for my studies of Indo-Pacific damselfishes
aided by National Science Foundation Grant GB-8732. Financial assistance for
trips to Kaster and Lord Howe Islands was provided by the National Geographic
Society, Washington, D.C. Finally I thank my wife, Connie, for her care in the
preparation of the typescript.

References

ALLEN, G. R., 1972.—The anemonefjishes, their classification and biology. Neptune, New Jersey ‘
T.F.H. Publications Inc.,

————, 1973.—Three new species of deep-dwelling damselfishes (Pomacentridae) from the
south-west Pacific Ocean. Australian Zool., 18 : 31-42.

, In press a.—Abudefduf amabilis (De Vis) an ecological colour variety of Abudefduf
leucopomus (Cuvier). Undersea Biology.

, In press b.—Damselfishes of the South Seas. Neptune, New Jersey : T.F.H. Publications
Inc.

Beavrort, L. F. de, 1940.—The fishes of the Indo-Australian Archipelago. Vol. 8. Percomorphi
(continued), Cirrhitoidea, Labriformes, Pomacentriformes. Leiden: E. J. Brill : 508.

BLEEKER, P., 1853.—Nieuwe tientallen diagnostische beschrijvingen van nieuwe of weinig bekends
vischsoorten van Sumatra. Nat. Tijdschr. Ned. Ind., 5 : 495-534.

, 1877.—Memoire sur les Chromides marins ou Pomacentroides de |’Inde archipélagique.
Nat. Verh. Holl. Maatsch. Haarlem., Ser. 2, 3(6) : 1-166.

CaRTIER, O., 1874.—Beschreibungen neuer Pharyngognathen ; ein Beitrag zur Kenntiss der
Fische des philippinischen Archipels. Verh. Phys. Med. Wurzburg, 5 : 96-106.

Fow ter, H. W., 1946.—A collection of fishes obtained in the Riu Kiu Islands by Captain Ernest
R. Tinkham. Proc. Acad. Nat. Sci. Philad., 98 : 173-218.

, and Bran, B. A., 1928.—Contributions to the biology of the Philippine Archipelago
and adjacent regions. The fishes of the families Pomacentridae, Labridae, and Callyodontidae
collected by the United States Bureau of Fisheries Steamer “‘ Albatross ”’ chiefly in Philippine
Seas and adjacent waters. Bull. U.S. Nat. Mus., 100(7) : 1-525.

GREENFIELD, D. W., and HENstEy, D. A., 1970.—Damselfishes (Pomacentridae) of Easter Island,
with descriptions of two new species. Copeza, 4: 689-695.

Woops, L. P., and ScHutrz, L. P., 1960.—Family Pomacentridae ; damselfishes. Jn Schultz,
L. P., Chapman, W. M., Lachner, E. A., and Woods, L. P., ‘‘ Fishes of the Marshall and
Marianas Islands.’ Bull. U.S. Nat. Mus., 202.(2) : 47-120.

PROCEEDINGS OF THE LINNEAN Society or New SoutH WAtzESs, Vou. 99, Part 2

POWDERY MILDEW ON CUCURBITACEAE: IDENTITY,
DISTRIBUTION, HOST RANGE AND SOURCES OF RESISTANCE

BARBARA BALLANTYNE*
[Accepted for publication 20th February 1974]

Synopsis

The species of powdery mildew fungi recorded on cucurbits are reviewed and the value of
several characteristics of the imperfect stage in distinguishing these species in the absence of the
perithecia is considered. One hundred and fifty collections were made in New South Wales from
a wide area and climatic range and on numerous commonly grown species, cultivars and plant
introductions of Citrullus, Cucumis and Cucurbita. All field collections resembled the imperfect
stage of Sphaerotheca fuliginea in having conidia which are borne in chains, have well-developed
fibrosin bodies and which produce germ tubes, some of which are forked. These characters have
occurred consistently in mildews identified as S. fuliginea on the basis of perithecial characteristics
by several workers in several countries.

Before 1958 Erysiphe cichoracearum had generally been assumed to be the most common and
widespread powdery mildew species on Cucurbitaceae, but recent reports and results of this
investigation now indicate that a mildew resembling the imperfect stage of S. fuliginea is generally
the more important species.

Cucurbita lundelliana and numerous cultivars and plant introductions of Cucumis melo and
C. sativus, which have previously been reported to have resistance to a mildew referred to as
Erysiphe cichoracearum in the U.S.A. and in other countries, were found to have resistance to the
powdery mildew in New South Wales.

A list has been made of cultivars and plant introductions of Cucumis melo, C. sativus and
Cucurbita spp. which have shown resistance and which are suitable for commercial production or
use in breeding programmes.

INTRODUCTION

Six species of powdery mildew fungi are recorded on Cucurbitaceae in various
parts of the world : Hrysiphe cichoracearum DC. ex Mérat, EL. communis (Wallr.)
Link., H. polygoni (DC.) St.—Am., H. polyphaga Hammarlund, Leveiilula taurica
(Lev.) Arnaud and Sphaerotheca fuliginea (Schlecht. ex. Fr) Poll. There are
also records of conidial powdery mildew fungi as Oidiwm sp. More than one
species may occur in the same locality (Teterevnikova-Babayan and Pe
1956) and on the same plant (Deckenbach and Koreneff, 1927).

The two species most commonly recorded are H. cichoracearum and S.
fuliginea but, since they rarely produce perithecia on cucurbits and their imperfect
forms have many similar features, the validity of most published records based
on the conidial stage requires confirmation.

In the U.S.A. resistance of cultivars and plant introductions was specifically
claimed to be to H. cichoracearum. No mention has been made of resistance to
S. fuliginea. However, Bohn and Whitaker (1961) stated that ‘ it seems desirable
that studies designed to determine the true identity or identities of Oidiwm stages
of Erysiphaceae on cucurbits should be made.’ They drew attention to the
conflicting host range reports in the literature, the demonstration of strains with
different temperature requirements which ‘ suggested that the relationships need
clarification.’

* silo gice al ae Ghanueal Research etnies Private Mail Bag No. 10, Rydalmere, New
South Wales, 2116.

PROCEEDINGS, OF THE LINNEAN Society or New SoutH WatzEs, Vou. 99, Part:2

BARBARA. BALLANTYNE 101

In New South Wales powdery mildew is common on rockmelon (muskmelon)
(Cucumis melo L.), cucumber (C. sativus L.) pumpkin (Cucurbita maxima Duch.),
marrow (C. pepo L.) and other related plants (Noble et al., 1934). The fungus
was referred to as H. cichoracearwm in earlier publications, but more recently has
been termed Oidiwm sp. (Anon., 1959), as only the imperfect stage has been
collected. !

In the late 1950’s the most widely grown rockmelon cultivar, Powdery
Mildew Resistant No. 45 (PMR 45), became severely affected by powdery mildew
in the south western areas of the State (Anon., 1959). The cultivars PMR Nos.
5 and 6 were resistant in 1959 but were severely affected the next season in the
same districts (Anon., 1960). The conidia of all collections of the fungus on
PMR Nos. 5, 6 and 45 contained well-developed fibrosin bodies (Zopf, 1887 ;
Blumer, 1933 ; Homma, 1937 ; Clare, 1958), which may be used as a characteristic
for distinguishing Hrysiphe and Sphaerotheca. Their presence suggested that
the fungus was the imperfect stage of S. fuliginea and not EL. cichoracearum.

A survey of powdery mildews on Cucurbitaceae in New South Wales was
therefore carried out from 1963 to 1965 to determine the species present and to
compare the disease reaction of cultivars and breeding lines grown both here and
in other countries.

: THE FUNGI—TAXONOMY
Perithecia

Six species of powdery mildews have been recorded on cucurbits in the
perithecial stage. Most records are of EH. cichoracearum and S. fuliginea and
their distributions are given in Tables 1 and 2. Erysiphe polygoni was recorded
on Cucurbita pepo in Poland by Schroeter (1893) and in Japan on three uncommon
‘species by Homma (1937).

There are several records of perithecia of powdery mildews on cucurbits under
the names H. communis and H. polyphaga (Berlese and Peglion, 1892 ; Hammar-
lund, 1945 ; Blumer, 1952). Both the taxonomy and nomenclature of these two
fungi need further study (Junell, 1965 ; 1967). The records of H. cichoracearum
in Table 1 include those of #. polyphaga.

Leveillula taurica was listed, together with H. cichoracearum and S. fuliginea
on cucurbits in the U.S.S.R. by Gordeeva (1961) and by Tarr (1955) with S.
fuliginea in the Sudan. Golovin (1956) refers to Leveillula on Cucurbitaceae in
her detailed survey of the genus, in which she split LZ. tawrica into species for each
host family. She described L. cucurbitarum on cucumber, marrow and Cucurbita
sp. in the U.S.S.R. but the description is invalid as no Latin diagnosis is given.

Imperfect Stage : Taxonomic Value of Characters of the Imperfect Stage in' distin-
guishing Powdery Mildew Species

As the identification of genera of powdery mildew is based primarily on
characters of the perithecia which are not always developed, the definition of
characteristics which could be used for precise identification in the absence of
- ‘the perfect stage would be of great value. Length and structure of conidiophores,
presence or absence of well-developed fibrosin bodies in conidia, and the mor-
phology of the germ tubes and appressoria, all appear to be suitable charac-
teristics (Ballantyne, 1963 ; Clare, 1964). The presence or absence of mycelial
appressoria and hyphal swellings also provide useful characters for Gustine mshite
Species.

Leveillula taurica is easily distinguished from other powdery mildews on
cucurbits in having internal mycelium and Oidiopsis-type conidiophores. E.
polygont is the only mildew with a single matured spore at the end of the conidio-

PROCEEDINGS OF THE LINNEAN SOCIETY oF NEw SoutH WALES, Vou. 99, Part 2

CUCURBITACEAE

POWDERY MILDEW ON

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PROCEEDINGS OF THE LINNEAN Society or New Souru WALEs, YOu. 99, Part 2

103

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104 POWDERY MILDEW ON CUCURBITACEAE

phore. The only confusion likely is between H. cichoracearum and S. fuliginea
which both have external mycelium and Oidiwm-type conidiophores with long
chains of conidia.

Fibrosin bodies were first reported by Zopf (1887) in the conidia and conidio-
phores of Podosphaera oxycanthae. Foex (1912, 1925) and Bouwens (1927)
considered that they were of value in distinguishing certain species and described
them in more detail. Blumer (1933) considered that presence or absence of
fibrosin bodies was probably of some value in distinguishing the oidial stages of
E. cichoracearum and 8S. fuliginea on cucurbits. This was confirmed by Homma
(1937) who described two forms, a granular form characteristic of Hrysiphe and
a form characteristic of Sphaerotheca which is either cylindrical or disc-, cone-,
or truncated-cone shaped.

Clare (1958, 1964) recognised the significance of well-developed fibrosin
bodies and used their presence or absence in distinguishing mildews on cucurbits
and other hosts in south eastern Queensland. Several workers, including Uozumi
and Yoshii (1952), Hashioka (1937), Sawada (1959), Dingley (1959) and personal
communication and Nagy (1970), either describe or figure well-developed fibrosin
bodies in cucurbit powdery mildews identified on perithecial characters as S.
fuliginea from Japan, Taiwan, New Zealand, The Netherlands and Hungary -
respectively. Ellert (1966) reported that these structures were present in powdery
mildews identified as S. fuliginea from perithecia on non-cucurbitaceous hosts.

Klika (1922) reported fibrosin bodies in numerous species including #.
cichoracearum, E. polygoni and S. humuli but without specifying the type.

The shape of the germ tube was claimed to be characteristic of the species
of powdery mildew by Hirata (1942, 1955) and later by Zaracovitis (1965). The
germ tubes of H. cichoracearum were simple with inconspicuous appressoria, those
of EH. polygont formed complex appressoria, and some but not all of the germ
tubes of S. fuliginea were forked. Hashioka (1937), Boerema and Van Kesteren
(1964) and Nagy (1970) reported forking of the germ tubes of S. fuliginea on
eucurbits in Taiwan, The Netherlands and Hungary. Homma (1937) and Salmon
(1900) figure the many-lobed appressoria of H. polygoni on other hosts.

Conidia of H. cichoracearum, EH. polygoni and S. fuliginea have generally
similar shape and size. However, some workers (Bouwens, 1924; Yarwood,
1957 and Nagy, 1970) consider that measurements of length and breadth are of
value in distinguishing between these species. Details of conidial measurements
recorded by other workers on cucurbits are given by Ballantyne (1971).

A species of Oidium resembling the imperfect stage of S. fuliginea in having
long chains of conidia, well-developed fibrosin bodies and a proportion of forked
germ tubes, has been recorded throughout Australia and in several overseas
countries (Table 3). Only two records of an imperfect stage lacking well-
developed fibrosin bodies and resembling H. cichoracearum in other ways have
been reported. These were on Momordica charantia L. and Sechiwm edule Sw.
in Hawaii (Raabe, 1966) and on several cucurbits in Hungary, where it occurred
with S. fuliginea which was identified on perithecial characteristics (Nagy, 1970)

THE Hosts

Powdery mildew is a serious disease of susceptible rockmelon cultivars in
many countries, particularly in arid areas where large scale commercial production
often takes place. It is usually less severe on pumpkins, marrows, squashes and
cucumbers, although it can be a serious problem in glasshouse cucumbers in
Europe, mainly because of continuous culture and very favourable conditions
for development of the disease. Watermelon (Citrullus lanatus (Thunb.) Mans-
feld var. caffer Mansfeld is not often affected, but occasional severe outbreaks

PROCEEDINGS OF THE LINNEAN Socrery oF New Sourn WaAtzEs, Vou. 99, Part 2

BARBARA BALLANTYNE 105

occur. In New South Wales two common cucurbitaceous weeds, the prickly
paddy melon (Cucumis myriocarpus Naud.) and the wild watermelon or camel
melon (Citrullus lanatus var. lanatus) may also be affected.

Rockmelon

The primary gene centre of Cucwmis melo is probably in tropical Africa, and
well-developed secondary gene centres of cultivated melons are in India, Iran,
southern U.S.S.R. and China.

Many powdery mildew resistant lines have been collected in Asia and Africa
and extensive breeding for resistance has been carried out. Details are in the
Appendix.

TABLE 3
Records of Oidium sp. resembling the imperfect stage of Sphaerotheca fuliginia

Africa
South Africa ae ate ae i .. Gorter, 1966
Asia
India .. Jhooty, 1967
Israel . Rudich eé al., 1969
Australasia

New South Wales, Australia
Northern Territory, Australia
Queensland, Australia ..
South Australia . .

Western Australia

Hurope
England ..

Greece. ae
The Netherlands

North America
California, U.S.A.
New York State, U.S.A.
Ohio, U.S.A.

South America
No recerds

.. Ballantyne, 1963 ; Clare, 1964
.. Ballantyne, unpublished data
. Clare, 1958, 1964
.. Harrison, personal communication
.. MeNish, 1967

. Zaracovitis, 1965
. Zaracovitis, personal communication
. Boerema and Van Kesteren, 1964

Kooistra, 1968

. Yarwood and Gardiner, 1964

Paulus eé al., 1968
Bohn, personal communication

. Kable and Ballantyne, 1963

Schroeder and Provvidenti, 1968

. Hllert, 1966

The most widely grown cultivar is Powdery Mildew Resistant No. 45 (PMR
45) which was released in California in 1936 (Jagger and Scott, 1937), and is still
widely grown though it was affected by a new race (race 2) in 1938 (Jagger et al.,
1938a). It is reported to be resistant to at least some of the races of powdery
mildew present in some of the Eastern States of the U.S.A. (Markarian and
Harwood, 1967).

Five genes for powdery mildew resistance have been designated, Pm}!->
(Jagger et al., 1938b ; Whitaker and Pryor, 1942 ; Bohn, 1961 ; Bohn and Whitaker,
1964 ; Harwood and Markarian 1968a and J).

Cucumber

India is considered to have been the centre of origin of the cucumber (Leppik,
1966b). Many powdery mildew resistant collections have been made in India and
Africa and some breeding for resistance has been carried out. Details are in the
Appendix.

PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WALES, Vout. 99, Part 2

106 POWDERY MILDEW ON CUCURBITACEAE

Inheritance of resistance has been shown to be complex (Smith, 1948;
Kooistra, 1968 ; Shanmugasundarum et al., 1971).

Cucurbita species

America, possibly Central America and southern Mexico is the centre of
origin of the genus Cucurbita (Whitaker, 1956).

Powdery mildew resistance has been reported in Cucurbita lundelliana
L. H. Bailey where it is controlled by a single dominant gene (Rhodes, 1959 ;
1964) and in C. martinezii L. H. Bailey. Further details are in the Appendix.

Watermelon
The watermelon probably originated in tropical Africa (Whitaker and Davis,
1962). There are no published reports of varietal resistance to powdery mildew.
Many rockmelon and cucumber cultivars bred for resistance in one country
also have resistance in other countries. Details are given in Table 4.

MATERIALS AND METHODS

One hundred and fifty collections of powdery mildew were made from a
wide range of locations and climates and on numerous cucurbit species and
cultivars.

The fungus was mounted in 3% aqueous potassium hydroxide for examin-
ation of fibrosin bodies and in tap water for examination of the conidiophore and
measurement of 20 conidia.

Germ tubes were obtained by germinating conidia on strips of onion bulb
epidermis according to the method of Hirata (personal communication). Epid-
ermis was stripped from the adaxial surface of the swollen leaf base, immersed
for three to five minutes in 80°, ethanol and washed in running water for two
hours. The strips were placed on a microscope slide with the cuticular surface
uppermost and blotted to remove excess moisture. Conidia were dusted onto
the strip and tap water was added with a dropper so that the strip floated. Early
collections were checked only for presence or absence of forking, but as investi-
gation proceeded the need for more precise data was recognised and the per-
centage of germ tubes showing forking in at least 500 germinating conidia was
determined for later collections.

The specimens are filed in the Herbarium of the Biology Branch, Biological
and Chemical Research Institute, Rydalmere (DAR). Permanent mounts of
the germinated and fresh spores were not made as no method of preserving these
in a satisfactory condition was known. Herbarium specimens of conidial powdery
mildews deteriorate with age and whilst structures resembling fibrosin bodies
were detected in herbarium specimens ten years old, they were faint and in-
frequent.

THE Hosts

As considerable variation had previously been observed in the reaction to
powdery mildew of commercial lines of powdery mildew-resistant rockmelon
cultivars, seed of such cultivars was obtained from the original breeder wherever
possible. Where commercial seed was used, lines of each resistant cultivar were
obtained from two different seedsmen. Details of seed source are given by
Ballantyne (1971).

During 1963, 53 lines of Cucumis melo were grown at Rydalmere near Sydney,
New South Wales, in hills with eight plants of each line per hill. Where seed of
some of the U.S. Plant Introductions was limited, no fewer than four plants of
each line were grown.

PROCEEDINGS OF THE LINNEAN SocrerTy or New SourH Watss, Vou. 99, Part 2

107

BARBARA BALLANTYNE

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PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WALES, VoL. 99, Part 2

108 POWDERY MILDEW ON CUCURBITACEAE

During 1964, 36 lines of C. melo, and ten lines of Cucurbita species were grown.
These lines included cultivars of C. maxima, C. moschata and C. pepo, the three
Cucurbita species cultivated in New South Wales, and C. lundelliana.

In the 1963 season, the results were recorded as resistant or susceptible
(except for one line) as the plants were either free from disease or severely affected.

In the 1964 season, disease ratings were recorded on the following scale and
the time of fruit maturity noted. The results (Table 5) were recorded when the
fruit first ripened.

R—Fully resistant ; no mildew seen.
R-—Resistant ; mildew on less than 5% of leaf surface.
MR—Moderately resistant ; mildew on 5-30% of the leaf surface.

S-—Susceptible ; mildew on more than 30% of leaf surface; ripe fruit
produced.

S—Fully susceptible ; mildew on more than 30% of leaf surface ; no ripe

fruit produced.

Observations were also made on the powdery mildew reactions of various
lines grown in small scale replicated and unreplicated trials at Griffith and Yanco
in the Riverina district in the south west of New South Wales during 1964 and
1965 and at Rydalmere in 1966.

Limited cross-inoculation experiments were carried out by transferring
mildew spores with a scalpel from french bean (Phaseolus vulgaris L.), cucumber,
and noogoora burr (Xanthium chinense Mill.) on to rockmelon (cv. Bender’s Sur-
prise) plants raised under bell jars.

RESULTS
The fungi
The 150 collections on naturally-infected cucurbits from New South Wales
showed conidial characteristics of S. fuliginea. The percentage of forking in 80
specimens varied between 5 and 60°, with most specimens in the range 3 to 5%.
Seventy specimens were checked only for presence or absence of forking. The
conidial measurements were (24) 27-40 x 16-24 (27) um.

The host plants on which mildew collected included : Citrullus lanatus var.
lanatus, wild watermelon ; C. lanatus var. caffer, watermelon, one cv. ; Cucumis
melo subspecies conomon (Thunb.) Greb., oriental pickling melon, eight accessions ;
C. melo subspecies melo, cultivated rockmelon, 17 cvs; C. melo subspecies and
cultivar unknown, three accessions; C. myriocarpus, prickly paddy melon ;
C. sativus, cucumber, 11 evs; Cucurbita ficifolia Bouché, fig leaf gourd; C.
lundelliana, the peten gourd ; C. martinezti ; C. maxima, pumpkin and hubbard
squash, 7 cvs; CU. moschata Duch. ex Poir, gramma, pumpkin and trombone;
2 evs; C. palmata Wats. ; C. pepo, marrow and squash, 5 evs ; C. radicans Naud. ,
C. texrana A Gray ; Cucurbita spp. unknown and a gourd, genus and species
unknown.

Most of the specimens were collected in February (78), March (24), January
(22) and April (16), with less in May (5), June (3), July (1) and October (1).
Ninety-four of the specimens were collected in the Sydney Metropolitan Area,
29 in the Riverina, 7 in the Central Tablelands, 4 in the North West Slopes, 5 in
the Northern Tablelands, 3 each in the Manning and Central Coast area, 2 each
in the Australian Capital Territory and on the North Coast and one in the North
West Plains. One collection from the Northern Territory was examined. This
specimen resembled those in the New South Wales area in having long chains of
conidia, a similar size, well-developed fibrosin bodies and a proportion of forked
germ tubes.

PROCEEDINGS OF THE LINNEAN Society of New Sours WALES, Vou. 99, Part 2

BARBARA BALLANTYNE 109

Both a collection from cucumber and a collection from french bean which
resembled the cucurbit powdery mildew fungus, readily infected rockmelon plants
in inoculation tests. A collection of powdery mildew on noogoora burr which
differed from the cucurbit powdery mildews also infected rockmelon plants in an
inoculation test. The collection showed the same characteristics on both the
noogoora burr and the rockmelon : the conidia were borne in chains, lacked well-
developed fibrosin bodies and produced simple unforked germ tubes. However,
it grew sparsely on the rockmelon and soon died. Many of the spores of this
fungus produced germ tubes from the end of the conidium whereas in other
collections from cucurbits the germ tube usually grew from the side of the
conidium.

Details of some representative collections are given below. Full details are
given by Ballantyne (1971).

Conidial collections

Citrullus lanatus var. caffer, watermelon cv. Blacklee, Rydalmere glasshouse,
March, 1963, DAR 7954, B. Ballantyne ;

Cucumis melo subspecies conomon, oriental pickling melon, C* 46, (U.S.P.1.
157070, Lt 90128), Rydalmere, February, 1963, DAR 7914, B. Ballantyne ;

C. melo subspecies melo, rockmelon (muskmelon) cv. PMR 45, Yoogali,
February, 1963, DAR 7984, B. Ballantyne.

C. sativus, cucumber cy. Polaris, Duranbah, October 1963, DAR 12226,
F. Autry Hall ;

Cucurbita moschata, pumpkin ev. Butternut, C153, Eastwood, February
1964, DAR 12801, B. Ballantyne.

Perithecial collections
Erysiphe cichoracearum. On Cucumis sativus, 1951, Dominion Laboratory
of Plant Pathology, KP 1798c, KP 1798d, Nova Scotia, Canada, D. Creelman.
Sphaerotheca fuliginea. On Cucurbita pepo, 1925, Herbarium mycologicum
Romanicum Exsicatti Fase. 1, No. 19, Herastrau, Romania People’s Republic,
Tr. Savalescu (ex CUP).

THE Hosts

Disease reactions as reported in the U.S.A. and observed in New South
Wales are given in Table 5.

Rockmelon. In 1963 at Rydalmere, the cultivars Delta Gold, Edisto,
PMR Nos. 6, 45 and 88, Rio Gold, Seminole and U.S.P.I. Nos. 124111, 124112
and subline L90209 of 183310 showed no signs of mildew. Other cultivars and
U.S.P.I. Nos. were severely affected.

In 1964 at Rydalmere, PMR Nos. 6 and 88, Seminole, LJ 430, breeding lines
151, 157 and 180 of M. B. Hughes, the P. lines (2-9, 10) of G. W. Bohn and
U.S.P.I. 234607 were fully resistant. Delta Gold, Edisto, PMR 45, Rio Gold
and Wescan were resistant. United States P.I. Nos. 164756, 165525 and 183307
gave mixed reactions with some resistant, some moderately resistant and others
susceptible. Florida No. 1 was moderately resistant, Florisun, Floridew and
U.S.P.I. 134200 were susceptible and other cultivars were fully susceptible.

In 1964 at Griffith, PMR Nos. 6 and 88, Seminole, 151, 157, 180, LJ 430 and
the P lines were fully resistant and PMR 45 and Edisto were fully susceptible.
In 1965 at Griffith the same results were obtained except that the P lines were
not included. In 1966 at Rydalmere, Campo and Jacumba were fully resistant.

* indicates the accession number of the authors collection.
+ indicates the numbers given by the U.S. Horticultural Field Station, La Jolla, which
provided seed of such lines.

PROCEEDINGS OF THE LINNEAN SocreTy oF New SoutH WaAtzzEs, Vou. 99, Part 2

POWDERY MILDEW ON CUCURBITACEAE

110

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PROCEEDINGS OF THE LINNEAN Society or New Sourn WALES, VOL. 99, Part 2

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PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WALES, VoL. 99, Part 2

POWDERY MILDEW ON CUCURBITACEAE

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PROCEEDINGS OF THE LINNEAN Society or New SourH Waxzs, Vou. 99 Part 2,

BARBARA BALLANTYNE 113

Cucumber. In 1964 at Rydalmere, Pixie and Polaris were resistant, Ashley,
Palomar and Stono were moderately resistant, U.S.P.I. 197087 gave a mixed
reaction with two plants moderately resistant and the other two susceptible.
This plant introduction was probably not a pure line (Barnes, personal com-
munication). Other cultivars and U.S.P.I. Nos. were susceptible including
179260 and 181910 (probably not pure lines [Bohn, personal communication ])
on which Randall and Menzies (1956) recorded perithecia of HL. cichoracearum in
the U.S.A. No perithecia of any mildew fungus were seen on these introductions
in this trial or on any other cucurbit host in New South Wales.

Cucurbita species. In 1964 at Rydalmere two plants of C. lundelliana
which produced mature fruit were moderately resistant, one which did not produce
mature fruit was resistant and the fourth was fully resistant. Other cultivars
of C. maxima, C. moschata and C. pepo were susceptible.

DISCUSSION

Considerable confusion has arisen in the literature because the name of the
perfect stage of a powdery mildew fungus has so commonly been given to the
imperfect stage without adequate identification. There is no doubt that both
EH. cichoracearum and S. fuliginea can occur on cucurbits in several countries,
e.g. India and the U.S.S.R., because perithecia of these two fungi have been
recorded on several occasions.

Before 1958 H. cichoracearum had generally been assumed to be the most
common and widespread powdery mildew species reported on Cucurbitaceae.
However, this investigation and recent reports from many countries indicate
that a mildew having major features of the imperfect stage of S. fuliginea is the
predominant mildew in some countries, and probably the only species in others.
Only two instances of an imperfect stage resembling LH. cichoracearum have been
reported on cucurbits. One was in Hawaii on Momordica charantia and Sechium
edule. The other was in Hungary where two powdery mildews commonly occur
on cucurbits ; one identified as S. fuliginea from perithecia, and the other iden-
tified as EL. cichoracearum on conidial characteristics (Nagy, 1970).

All naturally infected collections of cucurbit powdery mildew from many
areas and a wide range of climates in New South Wales resembled the imperfect
stage of S. fuliginea. The poor vigour of the mildew resembling HE. cichoracearum
from noogoora burr when artificially inoculated on to rockmelon suggests that
this fungus is not important on cucurbits in the field in New South Wales.

Observations by many investigators support the contention that the type of
conidiophore, the presence or absence of well-developed fibrosin bodies and the
mode of germination are useful criteria for distinguishing between the powdery
mildew species recorded on cucurbits. The very consistent data obtained in
the present study also support this. The reliability of these characters could be
further checked by culturing these species from ascopores under controlled
conditions and examining the colonies. In addition, characters which would
permit mixtures of species to be detected would be valuable if these could be
found. The conidial dimensions of HL. cichoracearum, E. polygoni and S. fuliginea
have generally been considered to be too similar to be of value in distinguishing
these species. Nagy (1970) compared length: width ratios of two cucurbit
powdery mildew fungi in Hungary ; one was identified as S. fuliginea on the
basis of perithecial characteristics and the other as EH. cichoracearum from the
morphology of the conidia. He found that the length : width ratio was signifi-
cantly different for the two species. However, he only quoted mean measure-
ments for length, width and the ratio between. The range of measurements of
50 conidia was not given.

PROCEEDINGS OF THE LINNEAN Socrety oF NEw SoutH WA tgs, Von. 99, Part 2

114 POWDERY MILDEW ON CUCURBITACEAE

It is considered that the mildew present on naturally infected cucurbits in
New South Wales is the imperfect stage of S. fuliginea.

Many cultivars, breeding lines and plant introductions of several species of
cucurbits with reported resistance to H. cichoracearwm were shown to have
resistance to a fungus resembling S. fuliginea in this investigation.

Tables 4 and 5 show that there is generally a close similarity in powdery
mildew reaction in different countries of various species and cultivars with
different genes and sources of resistance. An exception is in the reports of
Kooistra (1968), who found only slight resistance in many of the cucumber lines
reported to have a higher degree of resistance in other countries. However, his
testing was carried out under glasshouse conditions which are very favourable
for powdery mildew development and in which reactions cannot be regarded as
typical of field behaviour. Leppik (1966a) reported that some lines with good
resistance in the field showed less resistance under glasshouse conditions.

Some cultivars have a long history of resistance whereas others have resist-
ance when first grown and are later affected by another race of the fungus. For
example, the rockmelon cultivar PMR 45, whose resistance is due to the
single dominant gene Pm1+, was resistant when first grown in California, New South
Wales and Israel but later was affected by a new race in these three areas. There
have been no reports of the disease affecting cultivars such as Campo and Jac-
umba, whose resistance may be derived from several genes including one from
U.S.P.I. 124111. Where breeding programmes are being initiated or new
sources of resistance are being introduced into existing programmes, preference
should be given to lines having a history of resistance in several areas, including
some where races capable of severely affecting a range of cultivars are present.
The race 2 which occurs in California and Texas appears to be similar to the
race which occurs in south western New South Wales, except that the rockmelon
cultivar Seminole, which has shown no sign of mildew here, is only moderately
resistant in California. It is likely that the races designated 1 and 2 are both
complexes of races.

The severe outbreak of powdery mildew on the cultivar PMR 6 in the 1960
season and its resistance in the 1959, 1963, 1964 and 1965 seasons could be due to
the disappearance of the PMR 6—attacking form at the end of the 1960 season.
The conidia are short-lived and winter conditions in inland and southern coastal
New South Wales are too cold for survival of cucurbits. Perithecia have not
been found despite thorough search and overwintering on some other host is
therefore considered to be the likely means of survival. Alcorn (1967, 1969)
found seven non-cucurbit genera were alternative hosts of the cucurbit powdery
mildew fungus in Queensland, but these are sub-tropical or tropical species and
do not survive winter conditions in southern New South Wales.

Perithecia of powdery mildew fungi occur commonly in some countries and
rarely or not at all in others. Perithecia of numerous powdery mildew species
occur frequently in the Northern Hemisphere (Salmon, 1900; Blumer, 1933 ;
Viennot-Bourgin, 1956; Junell, 1967; Saville, 1968 and Solheim, Mycolflora
Saximont. Exsice. various Nos. in Herb. DAR), but perithecia of very few species
have been recorded in Australia.

Heterothallism is a possible explanation of perithecia being common in
some areas and rare or lacking in others. They may be more common in the
centre of origin of the species and rare or lacking in other areas where only one
mating type of fungus was introduced.

Perithecial formation is more frequent on some host species and cultivars
than others (Yarwood, 1957 ; Khan and Khan, 1970; Price, 1970). Most records
of S. fuliginea and FB. cichoracearwm have occurred on Cucumis sativus and Cucur-
bita pepo (Tables 1 and 2). Alcorn (1969) distinguished at least four patho-

PROCEEDINGS OF THE LINNEAN Society of New Souru WALES, Vou. 99, Part 2

BARBARA BALLANTYNE 115

genically distinct races of the cucurbit powdery mildew resembling 8S. fuliginea
in Queensland, all of which infected Cucumis sativus, Cucurbita maxima and C.
pepo, but not all infected the two cultivars of Cucumis melo and the single cultivar
of Citrullus lanatus var. caffer onto which they were inoculated. Perithecia may
occur on particular hosts such as Cucumis sativus and Cucurbita pepo more often
because these are susceptible to a wider range of isolates. The probability of
opposite mating types occurring together is therefore greater. More widespread
culture of these two species could also be the reason.

Homma (1937) reported that heterothallic species seemed to be more common
than homothallic ones in Japan. She showed that an isolate which she identified
as S. fuliginea on Taraxacum ceratophorum DC. was homothallic. However this
isolate has smaller than typical conidial and perithecial stages and may be a
different species.

Smith (1970) reported heterothallism in four mildew species in England and
suggested that any lateness or irregularity in perithecial formation in the field is
due to absence of the necessary mating types rather than to an unfavourable
environment or the nutritive condition of the host.

One might expect to find resistance in areas where both the pathogen and
host have been evolving side by side. Most of the powdery mildew resistant
collections of Cucumis have been made in the primary, secondary and tertiary
gene centres of this genus in Africa, India and nearby Asian areas, where the
imperfect stage of the mildew resembles S. fuliginea and perithecia of this fungus
have been recorded on several occasions. Perithecia of H. cichoracearum, which
is heterothallic (Morrison, 1961), have also been recorded on cucurbits in some
of these countries.

Many powdery mildew-resistant lines of Cucumis have been recorded but
little resistance has been reported in Cucurbita. This could be explained by the
presence or absence of powdery mildew in the gene centres in the early stages of
evolution of these genera. It is likely that S. fuliginea has been present in
Africa, India and nearby Asian areas for very long periods of time, as suggested
by the occurrence of perithecia. This would have favoured selection of mildew
resistance in Cucumis. No perithecia of S. fuliginea have been recorded on
cucurbits in the Americas, although they have been found on other hosts. A
mildew resembling the imperfect stage of S. fuliginea is common and widespread
on cucurbits in North America.

ACKNOWLEDGEMENTS

Grateful acknowledgement is made to Mr. J. Walker for guidance, to the
Directors of Herb. CUP, and the Canada Department of Agriculture Research
Station, Kentville, Canada, who lent perithecial specimens, to persons who
supplied seed and to Mr. A. Searle for photographs.

References

Atcorn, J. L., 1967.—Cucurbit powdery mildew on pawpaw. Qld. J. Agric. and An. Sci., 25:
161-164.
, 1969.—Infection experiments with cucurbit powdery mildew. Aust. J. Sci., 31: 296-
297.
Awnon., 1959.—Plant Disease Survey for twelve months ending 30th June, 1959. Sydney : New
South Wales Department of Agriculture. 54 pp.
, 1960.—Plant Disease Survey for twelve months ending 30th June, 1960. Sydney :
New South Wales Department of Agriculture. 42 pp.
, 1963.—Wescan—a cantaloupe adapted to Texas conditions. Leafl. Texas agric. Exp.
Stn., L-588 : 1-3.
, 1964.—State news. Texas. Valley develops ‘ Pearl’ of a cantaloupe. Am. Veg.
Grower, 12 (2): 38.
, 1965.—Institute of Horticultural Plant Breeding, Ann. Rep. for 1964. Wageningen.
Medeling 240. 60 pp.
, 1969a.—Dulce cantaloupe. Seed World, 104 (2) : 22.

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutH WALES, VOL. 99, Part 2

116 POWDERY MILDEW ON CUCURBITACEAE

, 19696.—Report of the Department of Agriculture for the year ended 30th June, 1968.
Sydney : New South Wales Department of Agriculture. 176 pp.

, 1971.—Tam dew honeydew melon. Seed World, 109 (2) : 22.

BALLANTYNE, Barbara, 1963.—A preliminary note on the identity of cucurbit powdery mildews.
Aust. J. Sci., 25 : 360-361.

, 1971.—Powdery mildew fungi on Cucurbitaceae in relation to breeding disease-resistant
cultivars. M.Sc. Agr. Thesis, University of Sydney. 73 pp.

BanGa, O., 1956.—International conference on the improvement of vegetable varieties at Wagenin-
gen, Netherlands on August 26th and 27th, 1955. Huphytica, 5 : 18-32.

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BARBARA BALLANTYNE 119

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APPENDIX
POWDERY MILDEW RESISTANT COLLECTIONS AND CULTIVARS

Rockmelon (Cucumis melo L.)

Many plant introductions have been collected from Asia, Hurope and Africa.
Several workers (Pryor et al., 1946 ; Mortensen, 1961, 1962 and personal com-
munication ; Leppik, 1966a ; Corley, 1966 and Oda, 1969) have reported on the
reaction of some of these to powdery mildew. Of 202 introductions from Asia,
Europe and Africa for which disease resistance was given in Corley’s compilation,
56 were resistant to powdery mildew in the U.S.A. Most of these were from
India (49) with others from Turkey (2), Africa (1), Iran (1), Peru (1), Saudi

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120 POWDERY MILDEW ON CUCURBITACEAE

Arabia (1) and the origin of one was not listed. Leppik (1966a) reported an
additional five resistant introductions from India. Oda (1969) reported four
introductions from India and one from Burma had a very high level of resistance
to powdery mildew, six from India and one from Nepal had intermediate resistance
and three others from India had a low level of resistance.

Resistant Cultivars

Cultivars bred in California and Texas for resistance to race 2 are PMR Nos.
5, 6 and 7 (Pryor et al., 1946), PMR 88 (Bohn, 1958), Wescan (Anon., 1963),
Perlita (Anon., 1964), Campo, Jacumba (Bohn e¢ al., 1965), Dulce (Anon, 1969a)
and Tam-dew (Anon., 1971). Those bred elsewhere in the U.S.A. for powdery
mildew-resistance include Georgia 47 (Minges, 1972), Delta Gold (Brown et al.,
1960), Seminole (Whitner, 1960), Floridew, Florida No. 1, Florisun (Jamison et al.,
1962 ; 1963), Golden Perfection (Brasher, 1965), Gulfstream (Minges, 1972),
Southland (Norton, 1970) and Gulfcoast (Norton, 1971).

Cultivars bred or selected for resistance in other countries are: in Israel,
Yokniam 54 and 56 (Ilan, 1963) and Ananas PMR, Pearl of En Dor, Yellow
Honeydew E1313 and Green Honeydew E3412 bred with PMR 45, Seminole and
Davis X as sources of resistance (Rudich et al., 1969) ; in Japan, Iyo 1 bred from
PMR 5 and Earl’s Favourite (Tamai et al., 1962); and in New South Wales,
Yanco Treat and Yanco Delight bred from subline 36739 of U.S.P.I. 124111
(Anon., 1969D).

Cucumber (Cucumis sativus L.)

Powdery mildew resistant collections have been made from Burma, U.S.P.I.
Nos. 200815 and 200818 (Wilson ef al., 1956) ; from Japan, U.S.P.I. 279465 and
from Ethiopia, U.S.P.I. 233646 (Leppik, 1966a), from India, U.S.P.I. 197087
(Barnes, 1961) and from China, several varieties including Vladivostoksky 155,
Di-huan-guas and Ty-hy-cy (Mescherov, 1961) and Puerto Rico Nos. 37 and 40
(Roque and Adsuar, 1939 ; Smith, 1949). Leppik (1966b) reported that several
wild cucumbers from India were immune and several wild species from Africa
were resistant to powdery mildew but they could not be crossed easily with
cultivated cucumber.

Resistant Cultivars

Resistance to powdery mildew has been reported in the following cucumbers
bred in the U.S.A.: Ashley, Stono and Palomar, three cultivars whose resistance
appears to be derived from Puerto Rico 40 (Barnes and Epps, 1956) ; Polaris,
Pixie, Pointsett and Cherokee, four cultivars whose resistance appears to be
derived from U.S.P.I. 197087 (Barnes, 1961; Minges, 1972) and Tablegreen,
whose source of resistance is unknown (Minges, 1972). The cultivar Natsufus-
hinari is reported to be resistant to powdery mildew in Japan (Hujieda and Akija,
1962) and Favor is listed as resistant to the disease in Sweden (Banga, 1956).

Cucurbita species

Cucurbita lundelliana lL. H. Bailey, the peten gourd, which is cross compatible
with the cultivated species of Cucurbita (C. maxima Duch., C. moschata Duch. ex
Poir. C. mizta Pang. and C. pepo L.) has resistance to powdery mildew in the
U.S.A. (Whitaker, 1956). Rhodes (1959) developed a gene pool among these
species and recovered hybrid plants tolerant to powdery mildew. Whitaker
(1965) developed relatively stable lines from crosses involving C. moschata and
C. pepo with C. lundelliana. These lines have a higher degree of resistance than
the susceptible commercial cultivars. Bemis (personal communication) reported
that C. martinewi L. H. Bailey has powdery mildew resistance similar to C.
lundelliana.

PROCEEDINGS OF THE LINNEAN Society oF New SourH WALES, VOL. 99, Part 2

THE GLYCASPIS SPP. (HOMOPTERA: PSYLLIDAE) ASSOCIATED
WITH HUCALYPTUS CAMALDULENSIS

K. M. Moorr*
[Accepted for publication 20th February 1974]

Synopsis

The distributions of three Glycaspis spp. populations associated with Eucalyptus camaldulensis
are illustrated and the distinctive lerp typical of two of the species is figured.

Additional information on the previously recorded distributions and hosts of other Glycaspis
spp. is given.

Previous studies on the Glycaspis spp. associated with HL. camaldulensis suggested that the
three psyllid species which utilise this plant species as host might indicate that the species H.
camaldulensis consists of more than a single taxon.

The present intensive study of the relevant Glycaspis spp., their distributions and host plant
associations, indicates that the effects of temperature apparently exert limitations on the dis-
tribution of the three species, although there is some broad correspondence between the
distributions of these psyllid species and the races of their host eucalypt.

INTRODUCTION

The species Hucalyptus camaldulensis Dehnh. has been considered as con-
sisting of a single species (Blake, 1953), of a single species and five varieties
(Blakely, 1955), and currently of a single species consisting of two subspecies
(Pryor and Johnson, 1971).

These differing interpretations of the species, and the general interest shown
by workers in various disciplines (Banks and Hillis, 1969 ; Pryor and Byrne, 1969)
suggested that a more comprehensive evaluation of the associated psyllids,
Glycaspis blakei Moore, G. brimblecomber Moore and G. eremica Moore, throughout
the range of distribution of their host plant, might provide information of value
in any taxonomic reassessment of the host.

Banks and Hillis (1969) established an intermingling of their ‘‘ northern ”
and ‘‘ southern ” chemotaxa of H. camaldulensis in four widely separated localities.
An intermingling of two of the three Glycaspis spp. associated with this species
as host, over an extensive area, had also been determined (Moore, 1970b, 1972).
It was therefore decided to examine more extensively the distributions of the
three Glycaspis spp. during this project, and attempt to determine whether
nymphs of each of the species surviving to the adult stage, utilised different
trees of H. camaldulensis, or completed their life cycle on the same tree.

These investigations included that portion of the L. camaldulensis distribution
encompassed by the Central Australia-Lake Eyre river drainage systems,
together with the three psyllid species previously found to be associated with that
plant as host.

METHODS

Lerps were collected, and nymphs reared to the adult stage, from a number
of localities, particularly in the area where the distributions of G. blakei and
G. brimblecombet were known to overlap. Lerps on portions of leaves from selected
trees were held in containers for up to 12 days and were examined daily. Adults
bred from these nymphs and lerps were preserved for examination and deter-
mination of the species.

Net collections of Glycaspis spp. were made at 65 collection sites (Fig. 1 and
Table 1), and details of the lerp shapes observed at each site were also recorded.

* Statue Bay, M/S 76, Rockhampton, Queensland, 4700.

PROCEEDINGS OF THE LINNEAN Society or New SoutH WALES, Vou. 99, Part 2
cc

122 GLYCASPIS SSP. ON EUCALYPTUS CAMALDULENSIS

TABLE 1

Collection localities for Glycaspis spp. Sites 1-64: collections from Eucalyptus camaldulensis.
Site 65 : collections from Eucalyptus tereticornis.

Number

Site Lerps of

No. Location River or Creek Present Species Specimens
1 16:7 mls. N. Wentworth Darling River OR _ brimblecombei 1
amnicola 15
2 31-5 mls. N. Pooncarie Darling River OR __ brimblecomber 3
amnicola 5
3 42 mls. E. Broken Hill Stephen Creek C eremica 21
4 16mls. W. Broken Hill Umberumberka Creek C eremica 13
5 Menindee Darling River OR _ brimblecomber 15
amnicola 14
6 S8mls. S. Wilcannia — Darling River OR __brimblecombet 14
amnicola NG
7 12 mils. N. Wilcannia Darling River 20 R__ brimblecombet 13
amnicola 2
8 Tilpa Darling River COR brimblecombet 22
amnicola 1
9 Louth Darling River C R_ brimblecombet 18
blake 5
10 14 mls. 8. Bourke Darling River C?0 R_— brimblecomber 18
blaket 1
11 12mls. E. Milparinka Warratta Creek C _ eremica 4
12 9mls. W. Milparinka Depot Glen Cc eremica 29
d brimblecombet 5
blaket 1
13. Wompah Gate Yalpunga Creek COR_ eremica 5
brimblecombet 13
blakez 1
14 7:3 mls. W. Warri Gate Stoney Creek 2 eremica 6
brimblecomber 1
blaket 1

15 9mls. NE. Innamincka Cullyamurra Waterhole

Cooper’s Creek OR brimblecomber 6
blake 1
16 9mls. N. Innamincka 2 COR eremica 2
brimblecomber 2
17 40 mls. N. Innamincka Patchewara Creek 2 brimblecombet 7
blake 2
18 23 mls. 8. Cordillo Dns. 2 C R._ brimblecomber 8
blake 3
19 44 mls. N. Cordillo Dns. 2 OR brimblecombei 2
blake 1
eremica 1
20 Copley Leigh Creek C eremica 23
21 33 mls. N. Leigh Creek ? C i 28
22 22mls. NW. William Creek Anna Creek C hi ll
23. Edwards Creek Edwards Creek CO e 38
24 11 mls. 8. Oodnadatta Allandale Homestead CO 33 11
25 89 mls. W. Oodnadatta ?Evelyn Creek C .5 19
26 61lmls. N. Welbourne Hill ?Alberga River C A 14
27 75mls. N. Welbourne Hill Tarcoonyinna Creek C is 5
28 5mls. N. DeRose Hill The Marryatt River Cc Me 43
29 Mt. Olga (Valley of Winds) Bubia Creek CO iN 24
30 Kings Canyon Kings Creek C 0 36
3 31 mls. E. Wallarah Palmer River C 2 50
32 Henbury Finke River C As 13
33 57 mls. S. Alice Springs Hugh River C 9p 19
34 Alice Springs Todd River C 0 36
35 47 mis. E. Alice Springs Trephina Creek C 4 10
36 Ormiston Gorge ? C y 13
37 18 mls. N. Alice Springs 16-mile Creek (@) ue 33
38. +36 mls. N. Alice Springs Burt Creek C oF 29
39 44 mls. on Harts Ra. Rd. Gillen Creek (Sandover) C eremica 22

PROCEEDINGS OF THE LINNEAN Society or New Soursn Watszs, Vou. 99, Part 2

k. M. MOORE

123

TABLE 1—continued
Number
Site Lerps of
No. Location River or Creek Present Species Specimens
40 84mls. EK. Harts Ra. Police
Station Marshall River C R- eremica 5
blaket 4
brimblecombei 2
41 81 EK. Harts Ra. Police
Station Plenty River C eremica 14
brimblecombei Dy
blaket 1
42 14mls. 8. Barrow Creek ? C eremica 3
43 25 mls. N. Barrow Creek Taylor Creek C eremica 38
44 Wauchope ? C eremica 18
45 Devil’s Marbles — C R_ eremica 15
46 23 mls. N. Wauchope McLaren Creek C R-_ eremica 7
brimblecombez 2
47 Daly Waters ?Katherine River C?0 blakea 9
48 16mls.S. Renner Sprmgs Tomlinson Creek C blaket 23
49 Attack Creek Attack Creek C R blakei 18
50 6mls. N. Tennant Creek Tennant Creek (Oe) blakez 6
eremica 2
51 8mls. E. Camooweal ? COR. blaket 5
52 62 mls. E. Camooweal Buckley River CoO blakew 2
53 3mls. N. Mt. Isa Leichhardt River C blaket 7
j brimblecombei 1
54 2 mls. EH. Mt. Isa Breakaway Creek COR. blakei 19
55 Urandangi Georgina River CO blaket 34
brimblecomber 1
56 66 mls. 8S. Dajarra ~ ? C R__ blakei 17
brimblecombet 2
57 Boulia Burke River C R blaket 6
: brimblecombei 4
58 23 mls. SE. Springvale Diamantina River C R_ blakei 1
59 104 mls. W. Windorah Farrar’s Creek C RR blaket 1
60 7mls. NE. Windorah Cooper’s Creek C?0 blaker 18
brimblecombei 2
61 © Isisford Barcoo River COR. blake 9
62 10mls. NW. Longreach Dingo Crk. (Thompson R.) C R __ brimblecombei 2
63 Alice Alice River COR. blakei 5
brimblecomber i
64 E. of Drummond Range Medway Creek COR blakew 4
brimblecomber 1
From E. tereticornis
65 Meteor Creek Meteor Creek COR. brimblecombei 13
blaket 3

Total specimens examined : 1082

Lerp shape : C = clover leaf; O = oval; R = round

All insect material has been placed in The Australian National Insect Collec-
tion, C.S.I.R.O., Canberra, A.C.T. Methods of storage and treatment of
specimens for examination and identification were essentially as those previously
recorded (Moore, 1961, 1964, 1970a).

A sample of buds, seed capsules and mature leaves, when each or all were
available, was obtained from a selected tree of H. camaldulensis on which Glycaspis

lerps occurred, at each collection site.

Corresponding site numbers on aluminium

labels were attached to the samples which have been lodged, together with
relevant details of the sites, with the New South Wales National Herbarium,
Royal Botanic Gardens, Sydney, N.S.W.

PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WALES, Vou. 99, Part 2

124 GLYCASPIS SSP. ON EUCALYPTUS CAMALDULENSIS

RESULTS
A. Glycaspis spp. associated with EH. camaldulensis.

An extensive area where the three Glycaspis spp. blaket, brimblecombei and
eremica intermingle, was determined. This approximate area is delimited by
the heavier outline in Fig. 1.

From the rearing of nymphs to the adult stage, it was found that two or
more of these Glycaspis spp. were able to coexist and survive on the same tree.
The non-selectivity between trees of H. camaldulensis by these three species in
areas where they intermingle therefore lessens the likelihood that they indicate

blakei e

brimblecombei @

as

Lc
an}
\

=e:
=

\

uci

a!

Fig. 1. Distributions of three Glycaspis spp. on Hucalyptus camaldulensis, and of G. confinis
on E. rudis and EH. cornuta.

separate taxa of their host plant, at least in that area of intermingling. Never-
theless, their known distributions and overlap may indicate, in a broad sense,
the approximate areas of distribution for more than a single taxon of H. cam-
aldulensis.

The typical shape of lerps constructed by G. blaket and G. eremica, atypical
of those constructed by other species of Glycaspis, are illustrated in Fig. 2, and
the general shape suggests the name “‘ cloverleaf ”’ lerps. Each lerp is constructed
by a different nymph, and the figure illustrates the superimposed lerps of three
nymphal instars. Lerps are usually white, or rarely yellow, and their composition
appears similar to that of lerps of other species in the subgenus Glycaspis. The
close phylogenetic relationship of these two species, aS previously indicated
(Moore, 1970a), is thus confirmed by the lerp shapes. <A few lerps were different
in shape, being more or less like a Maltese Cross (i.e. with two crossbars at right

PROCEEDINGS OF THE LINNEAN Society or New SoutH WALES, Vou. 99, Part 2

K. M. MOORE 125

angles) or single-bar-shaped. These aberrant lerp shapes were not investigated,
but it is suggested that such atypical forms might result from parasitism of the
associated nymph.

The distribution of G. blaket was not found to extend southwards beyond its
previously determined southern limit at Wileannia, New South Wales. It
occurred in large numbers on H. tereticornis Sm. at Site 65 (Meteor Creek, Queens-
land), adult specimens of this species and of G. brimblecombei being reared from
the sample tree. Although H. tereticornis is plentiful from that site eastward to
the coast, an investigation of this area revealed no further cloverleaf lerps. The
occurrence of G. blakei on EL. tereticornis is a new host record for this species.

ga

Fig. 2. Cloverleaf lerp typical of G. blake: and G. eremica.

Known only from west of the Darling River, G. eremica intermingles with
G. blakei and G. brimblecombei at a few sites in the drier inland areas of the Nor-
thern Territory, Queensland, South Australia and New South Wales, although it
occurs alone, throughout the greater portion of its range (Fig. 1).

The species G. gradata Moore, previously recorded as possibly occurring on
EH. camaldulensis and E. largiflorens F. Muell., was not obtained during this project.
Its absence from the intensively sampled former host thus might be interpreted
as indicating that its correct host is H. largiflorens, and from its distribution
records, other closely related ‘‘ box” species also. It is possible that adverse
seasonal factors may have prohibited its occurrence on H. camaldulensis during
these investigations, but this seems unlikely.

At Sites 1 to 10, round lerps were utilised for rearing Glycaspis spp. nymphs
to the adult stage for determination of the species. Table 1 indicates that lerps
of G. amnicola Moore intermingle with those of G. brimblecombei on the same
tree. G. amnicola was previously recorded as constructing oval lerps, but it is
now evident that many of the lerps of this species are round, as shown by emer-
gences from the lerps used in the rearing of nymphs, and they are apparently
indistinguishable from the round lerps of G. brimblecombei. It is maintained that
some lerps of G. amnicola are oval, thus indicating a possible transitional evolution-

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutH WatsEs, Vou. 99, Part 2

126 GLYCASPIS SSP. ON EUCALYPTUS CAMALDULENSIS

ary Stage from the formation of round to oval lerps by nymphs of G. amnicola.
Northwards from Site 8 (Tilpa) G. amnicola was replaced by G. blakei, and the
former species now is known to coexist with G. brimblecombei from Euston on the
Murray River, along the Darling River to Tilpa.

B. Glycaspis spp. associated with other hosts. -

Additional collections for Glycaspis specimens were made from a number of
other hosts. Results, arranged according to the subgeneric classification of
Eucalyptus and the system of coded species of Pryor and Johnson (1971), are
now given.

(i) Subgenus Blakella

Species BAA :A Hucalyptus tessellaris F. Muell. was sampled in several areas,
but no Glycaspis specimens were obtained or lerps observed. It is now considered
that this species is most unlikely to be a host of Glycaspis spp., even though it
was recorded as a possible host of G. brunneincincta Moore (1970a). This con-
clusion is consistent with previously recorded results of collections from other
eucalypt species in this subgenus (Moore, 1970D).

(ii) Subgenus Hudesmia

Species EAADC HE. odontocarpa F. Muell. was sampled in three widely
separated areas. No Glycaspis specimens or their lerps were obtained or observed,
these results being consistent with those previously recorded (Moore, 1970D).

EAADE HE. gamophylla F. Muell. was sampled in several areas with the same
results. This project presented an opportunity to become well acquainted with
this eucalypt in the field, and it is now almost certain that the plant previously
collected from, at Joffre Falls, Western Australia (Moore, 1970a), was not H.
gamophylla but probably SNABG EL. brevifolia F. Muell. At that time, HE. gamo-
phylla in the field was not familiar to the writer.

EFAAA E. similis Maiden was intensively and extensively sampled in one
locality, but no Glycaspis specimens were obtained, or their lerps observed.

Intensive sampling of EFABA E. baileyana F. Muell. in one locality produced
the same result. ,

Within this subgenus, 11 of the 15 species have now been sampled for Gly-
caspis spp. which were obtained only from EAC:A E. tetrodonta F. Muell., EFC:A
E. miniata A. Cunn. ex Schau. and EFC:B H. phoenicea F. Muell. (Moore, 1970a,
1970b).

(iii) Subgenus Gaubaea

GAA:A EH. curtisii Blakely and White was intensively sampled in one area,
but no Glycaspis specimens or their lerps were obtained or observed.

GAA:C EH. tenuipes (Maiden and Blakely) Blakely and White was intensively
sampled in one area, with the same results.

(iv) Subgenus Monocalyptus

Glycaspis specimens from MATEL EH. radiata Sieber ex DC. collected by
A. Yen of La Trobe University, at Healesville, Victoria, were identified as G.
endasa Moore. This species was originally collected and named from LH. ‘‘ robert-
soni”’ Blakely at Towamba, South Coast, New South Wales, by the writer ;
however, the trees in that area are H. radiata ssp. radiata.

(v) Subgenus Symphyomyrtus
Species SBA:C FL. raveretiana ¥. Muell. Trees on Moore’s Creek, and in the

Botanic Gardens, Rockhampton, Queensland, were sampled for Glycaspis and
their lerps, but no specimens were obtained or observed.

PROCEEDINGS OF THE LINNEAN Society oF NEw SourH WALES, Vou. 99, Part 2

K. M. MOORE 127

SIVEO HF. pachyphylla F. Muell. was previously sampled (Moore, 1970b),
when no. Glycaspis specimens were obtained or lerps observed. During this
project it was sampled in two widely separated localities where it was established
that Glycaspis specimens of the occidentalis group were utilising it as host. The
species obtained, which constructs round lerps, requires more study before it can
be assigned a correct position within the occidentalis group of species.

SNABG EL. brevifolia F. Muell. was sampled at Skull Creek, near Central
Mount Stuart, Northern Territory. Adults collected were identified as G. onychis
Moore, and this becomes the most southern locality where G. onychis has been
obtained. The previous most southern locality was Joffre Falls, Western Aust-
ralia.

SNEEB LE. tereticornis Sm. At Site 65, intermingling round and cloverleat
lerps were collected from the sample tree, and nymphs reared to adults. The
species were determined as G. brimblecombei and G. blakei respectively.

SNEEP #. camaldulensis. Additional to the species previously mentioned

in this paper, the following Glycaspis spp. were obtained during net collections
from this plant species :

G. cnecosia Moore from Sites 15, 18, 19, 31, 59, 60.
G. sudicola Moore from Sites 6, 23, 49.

G. froggatti Moore from Sites 32, 49, 54.

G. buxalis Moore from Site 25.

G. retrusa Moore from Site 51.

The sweeping of these species from the foliage of H. camaldulensis does not
necessarily mean that the eucalypt is utilised as host by the Glycaspis species
obtained ; e.g. it has been found that species accidental to the host may often be
collected during periods of windy weather. The positions of trees in relation to
each other and the prevailing wind direction are also contributing factors ;
hence the large number of queried hosts recorded in Moore (1970a). It seems
certain that none of the species G. buxalis, G. retrusa or G. froggatti utilise EL.
camaldulensis aS host, when their previously recorded hosts are considered.
EH. microtheca F. Muell., which also occurred at each of these sites, is the probable
host of these species.

SNEEX E. exserta F. Muell. Previously recorded from 34 miles north of
Clermont, Queensland, adults of G. exsertae Moore were obtained from this host
four miles west of Alice, Queensland, and it was verified that their lerps were
round in shape. The original description of this species was based on one male
Specimen only (Moore, 1970a).

SUADE E. microneura Maiden and Blakely. G. froggatti was previously
recorded from this host and eight other eucalypt species, from Nannine in Western
Australia to Clermont, Queensland (Moore, 1970a). It has now been collected
from this host at two miles east of Mt. Isa, Queensland, where it was verified that
the shape of its lerp was round to oval, as previously recorded.

SUADJ EH. cyanoclada Blakely. This eucalypt species previously had not
been sampled for Glycaspis specimens. At 56 miles east of Frewena, Northern
Territory, adults of G. retrusa were obtained during this project, but no lerps were
found, although intensively sought.

SUG:A H. cambageana Maiden. G. cnecosia was collected from this, its
originally recorded host, at 36 miles north of Biloela, Queensland, where its
numerous round lerps occurred mainly on the twigs and blossoms, thus indicating
its possible affinities with the occidentalis group of species.

SUP:Y EH. pruinosa Schau. was again sampled for Glycaspis spp. but, as
previously, no adults or lerps were obtained or observed.

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutH WALES, Vou. 99, Part 2

128 GLYCASPIS SSP. ON EUCALYPTUS CAMALDULENSIS

(vi) Melaleuca argentea W. V. Fitzg.

Specimens of G. devexa Moore were obtained from this host at Moonal Creek,
28 miles south-east of Urandangie, Queensland, this record constituting a new
host for G. devera. The range of its previously recorded distribution is thus
considerably extended.

CONCLUSION

The area where the three Glycaspis spp. on EH. camaldulensis intermingle
shows some correlation with the 31°C. summer isotherm. This isotherm extends
from the approximate centre of the Northern Territory to the south-eastern
corner of that State, the north-eastern corner of South Australia almost to the
north-western corner of New South Wales, to south-west and central Queensland,
passing from latitude c. 18° S. to ¢. 28° S., through the four areas of differing
summer isohyets of 0 mm to 125 mm, 125 mm to 250 mm, 250 mm to 500 mm and
500mm to 1000 mm. A temperature effect on the distribution limits of the three
Glycaspis spp. is thus indicated as the most probable limiting factor.

Collections of the three Glycaspis spp. and the sampling of EL. camaldulensis
during this project were necessarily confined to the planned route. Presence or
absence of the host on a discrete river or creek contacted at more than one locality,
varied considerably ; e.g. although the host species was present at Sites 58 and
59, it was absent from the lower reaches of the same watercourse at Birdsville.
Differing soil types appeared to be associated with this variability.

The atypical ‘‘ cloverleaf”’ lerps of G. blakei and G. eremica suggest phylo-
genetic divergence of these two species from the remainder of the species which
construct round, oval or rectangular lerps within the subgenus Glycaspis.

ACKNOWLEDGEMENTS

The Committee of the Science and Industry Endowment Fund, C.S.I.R.O.,
to whom the writer is most grateful, supported this project with a substantial
monetary grant.

Thanks are also expressed to Dr. L. A. S. Johnson and Mr. D. Blaxell, of the
National Herbarium, Royal Botanic Gardens, Sydney, for their interest in, and
care of, the Hucalyptus spp. samples, and to Dr. Barbara Briggs, of the same
institution, for her identification of the Melaleuca species.

References

Banks, J. C. G., and Hiruis, W. E., 1969.—The characterization of populations of Hucalyptus
camaldulensis by chemical features. Aust. J. Bot., 17: 133-146.

Brake, 8. T., 1953.—Studies on Northern Australian species of Hucalyptus. Aust. J. Bot.,
1: 186—352.

BuaKkety, W. F., 1955.—A key to the Hucalypts. 2nd ed. Canberra: Forestry and Timber
Bureau.

Moors, K. M., 1961.—The significance of the Glycaspis spp. associations with their Hucalyptus spp.
hosts : erection of a new subgenus and descriptions of thirty-eight new species of Glycaspis.
Proc. Linn. Soc. N.S.W., 86: 128-167.

————,, 1964.—-Additional information on the genus Glycaspis : erection of a new subgenus and
descriptions of six new species. Proc. Linn. Soc. N.S.W., 89 : 221-234.

~ —, 1970a.—A revision of the genus Glycaspis (Homoptera : Psyllidae) with descriptions of
seventy-three new species. Aust. Zoologist, 15 : 248-342.

, 1970b.—Results from a study of the genus Glycaspis. Aust. Zoologist, 15: 343-376.
. 1972.—The Glycaspis spp.—Eucalyptus camaldulensis associations. J. ent. Soc. Aust.
(N.S.W.), 7: 3-7.

Pryor, L. D., and Byrne, O. B., 1969.—Variation and taxonomy in Hucalyptus camaldulensis.
Silwae Genetica, 18 7 64-71.

Pryor, L. D., and Jonnson, L. A. 8., 1971.—A classification of the Hucalypts. Canberra: A.N.U.

Press.

PROCEEDINGS OF THE LINNEAN Soctety or NEw SoutH WAxzEs, VOL. 99, Part 2

BIOGENIC STRUCTURES IN THE ORDOVICIAN BOWAN PARK GROUP,
NEW SOUTH WALES

V. SEMENIUK*

{Communicated by B. D. Webby]

(Plates v and v1)

[Accepted for publication 20th February 1974]

Synopsis

Chondrites, large horizontal meandering burrows, small pellet-filled burrows and burrow-
mottled limestones are the main biogenic structures in the Daylesford and Quondong Limestones
of the Ordovician Bowan Park Group. Synsedimentary diagenetic phenomena associated with
biogenic structures include obliterated primary fabrics, fragmented thin-shelled skeletons and
vertically mixed sequences of sediments. The occurrence of burrows in the limestones has in-
fluenced the distribution of later alteration such as dolomitisation, grain growth, stylolitisation
and silicification. The various burrows occupied consistent and unique facies during deposition
of the Daylesford and Quondong Limestones : Chondrites occurred in dark grey lime mudstones,
wackestones and packstones in intermediate to offshore environments ; large horizontal burrows
and small pellet-filled burrows occurred in dark grey wackestones and packstones in intermediate
environments. These burrows are comparatively rare in grainstones and grey skeletal wacke-
stones/packstones of nearshore environments.

INTRODUCTION

Biogenic structures are common in many sedimentary rocks and there is a
considerable literature dealing with their description, environmental significance
and role in diagenesis (Simpson, 1957 ; Seilacher, 1964, 1967). Studies by Seil-
acher (1962), Farrow (1966), Goldring (1962) and others have shown their im-
portance in unravelling ancient environments. Biogenic structures in modern
sediments have been studied using plastic impregnated cores, plastic casts of
burrow systems, epoxy relief peels and X-ray radiographs (Frey and Howard,
1969).

Burrows and other evidence of biogenic activity are conspicuous and
important in many horizons on the Bowan Park Group (Semeniuk, 1970, 1973b).
These biogenic structures are significant because of the associated synsedimentary
diagenetic effects (particularly the mixing of facies) and because of their consistent
occurrence within recurring facies in the sequence. This paper is an account of
biogenic structures in the lower two formations of the group, the Daylesford and
Quondong Limestones. It deals with burrow description, the role of burrowing
organisms in synsedimentary diagenesis and the influence of burrows on later
alteration. The occurrences of biogenic structures are placed in an inferred
environmental framework of the Daylesford and Quondong Limestones.

This study is based on some 600 samples collected from five section localities
(Semeniuk, 1973a). Samples were studied as thin sections, polished slabs and
acid-etched hand specimens. Numbers prefixed by U.S.G.D. and 8.U.P. refer to
material catalogued in the petrology and palaeontology sections respectively of
the Department of Geology, University of Sydney. Limestone classification of
Dunham (1962) is used in this paper. Muddy limestone is used as a collective
term for rocks with abundant lime mud, i.e. packstone, wackestone and lime
mudstone.

* Department of Geology, University of Western Australia, Nedlands, W.A., 6009.

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutH WALES, Vou. 99, Part 2

130 BIOGENIC STRUCTURES IN THE ORDOVICIAN BOWAN PARK GROUP

Outline of stratigraphy

The Ordovician Bowan Park Group (Semeniuk, 1970; 1973a) is located in
the Central Fold Belt (Packham, 1969) of New South Wales, and crops out in the
main limestone belt between Eurimbula and Cowra (Fig. 1). The group discon-
formably overlies the Cargo Andesite and is disconformably overlain by the
Malachi’s Hill Beds (Semeniuk, 1970 ; 1973a).

The Bowan Park Group consists mainly of limestone and contains three
formations (Semeniuk, 1970 ; 1973a) :

Ballingoole Limestone (top), 280m thick; mainly massive, generally
unfossiliferous limestone ; conformably overlies Quondong Limestone, 34m
thick ; fossiliferous, thinly bedded limestone and marl; disconformably
overlies—Daylesford Limestone, 250m thick ; terrigenous sediment and marl
in basal part, thinly bedded to massive limestone in middle part, and mainly
massive limestone in upper part ; disconformably overlies Cargo Andesite.

»Eurimbula

voonad

\

Now South
Wales
a

BOWAN PARK

Fig. 1. Location map, distribution of Ordovician limestones between Eurimbula and Cowra
and location of the Bowan Park Group.

At the type section the Daylesford Limestone contains six members, which
in ascending order are : Ranch Member, Bourimbla Limestone Member, Manooka
Limestone Member, Gerybong Limestone Member, Glenrae Limestone Member
and Davys Plains Limestone Member. East of the type section, the Oakley
Limestone Member is laterally equivalent to the Manooka and Gerybong Lime-
stone Members. Thicknesses and diagnostic lithologies of members are sum-
marised in Table 1.

The Quondong Limestone consists of thinly bedded grainstone, pellet
packstone, skeletal wackestone and packstone and lime mudstone. Though not

PROCEEDINGS OF THE LINNEAN Socrery of New Sours WaAxEs, Vou. 99, Part 2

V. SEMENIUK Sill

formally subdivided into members, the formation contains a lower part mainly
of thinly bedded grainstone and pellet packstone, and an upper part mainly of
marly lime mudstone and pellet packstone.

Tithofacies

The Daylesford and Quondong Limestones contain four main limestone
types, and minor intercalations of terrigenous sediment. limestones are:
(1) grainstones, (2) grey skeletal wackestones and packstones, (3) dark grey
burrowed wackestones and packstones and (4) dark grey burrowed lime mudstones.

Grainstones are mostly light-coloured and commonly exhibit lamination,
cross-lamination and graded bedding. Gravel-sized fossils and pisolites occur
in bands and platy grains are arranged parallel to lamination. Most grainstones
contain well-sorted, subangular to rounded, medium to coarse sand-sized skeletal

TABLE |
Daylesford Limestone stratigraphy

Unit Thickness Diagnostic Lithologies
(metres)
Davys Plains Lime- 95 Skeletal lithoclast grainstone, skeletal wackestone and pack-
stone Member stone, pellet packstone.
Glenrae Limestone 25 Light grey, vuggy and weathered limestone in upper part ;
Member intercalated and burrow-mottled grainstone, skeletal wacke-

stone, lime mudstone in lower part.

Oakley Limestone 90 Skeletal grainstone, skeletal lithoclast grainstone.
Member
Gerybong Limestone 64 Lime mudstone, skeletal wackestone.
Member
Manooka Limestone 16 Skeletal grainstone, skeletal lithoclast grainstone, skeletal
Member packstone and wackestone, lime mudstone.
Bourimbla Limestone 16 Skeletal wackestone and packstone, lime mudstone.
Member
Ranch Member 34 Unit 3: Skeletal wackestone, lime mudstone, terrigenous mud-
stone.

Unit 2: Skeletal grainstone, packstone and wackestone,
skeletal pellet grainstone, lime mudstone.

Unit 1: Lithic sandstone, terrigenous mudstone, lime mud-
stone.

and lithoclast grains. Grey skeletal wackestones and packstones are mostly light
grey to medium grey, or, less commonly, dark grey. Burrows are uncommon.
Whole gravel-sized fossils are in situ, in layers, or randomly oriented. Frag-
mented, thin-shelled skeletons are angular, poorly-sorted to well-sorted, fine sand-
to gravel-sized, and comprise a minor part of the sediment.

Dark grey wackestones and packstones are commonly burrowed, containing
gravel-sized and sand-sized whole and fragmented fossils in a dark grey lime
mud matrix (skeletal wackestones and packstones); some sediments contain
abundant pellets (pellet packstones). Large fossils are oriented in layers, or,
more commonly, randomly oriented and disrupted by burrows. Sediments
contain abundant poorly-sorted, angular, fine to coarse sand-sized skeletal
fragments which are randomly oriented or concentrated in burrows. Skeletal
fragments include thin-shelled brachiopods, sponge spicules, dasycladacean algae,
ostracods, trilobites, bryozoans, small gastropods and thin-walled coral tubes.

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutH WALES, VOL. 99, Part 2

132 BIOGENIC STRUCTURES IN THE ORDOVICIAN BOWAN PARK GROUP

Dark grey lime mudstones are commonly burrowed and consist of clay-sized
calcite (micrite of Folk, 1965) and patches of recrystallised micrite (microspar of
Folk, 1965) ; some contain a high proportion of silt- to fine sand-sized skeletal
debris. Sand-sized angular fragments of dasycladacean algae, small gastropods,
small thin-shelled brachiopods, ostracods, trilobites and sponge spicules are
minor components of the sediment.

Terrigenous sediments include lithic sandstone, and brown and green mud-
stone. lLithic sandstones are well-laminated, medium to coarse sediments.
Brown mudstones are structureless ; green mudstones are burrowed, and grade
into burrowed, marly lime mudstone.

Stratigraphic relationships of lithofacies

The Bowan Park Group trends approximately east-west for 9 km, and this
has permitted analyses of facies changes in a direction probably perpendicular to
the palaeo-shoreline (Semeniuk, 1973b). Grainstones and grey skeletal wacke-
stones and packstones are more common in eastern sections of the Daylesford
Limestone (as evidenced by the Oakley, Bourimbla and Ranch Members) and are
laterally equivalent to dark grey lime mudstones, wackestones and packstones
that dominate western sections (Semeniuk, 1973b). Eastern sections also contain
abundant lithoclasts in grainstones ; these lithoclasts are absent from the thick
muddy sections to the west, except in thin horizons immediately above discon-
formities.

Lithofacies also tend to occur in sequential arrangement immediately above
disconformities. The fully developed, idealised sequences are :

Type 1. Grainstone grading up into dark grey, burrowed, wackestone (or
packstone) which grades into dark grey, burrowed lime mudstone ;
total average thickness 1-5 m.

Type 2. Grey skeletal wackestone and packstone (or, in marly sections,
brown terrigenous mudstone) grading up into dark grey, burrowed
skeletal wackestone and packstone (or green terrigenous mudstone)
which grades into dark grey, burrowed lime mudstone; total
average thickness 0-5 m.

These sequences are well developed above disconformities in horizons of the
Quondong Limestone and in the Ranch, Manooka and Davys Plains Limestone
Members of the Daylesford Limestone. Elsewhere, the sequences are only
partly developed with either the basal or top lithology absent. Reversals of the
sequences may occur beneath disconformities.

DESCRIPTION OF BIOGENIC STRUCTURES

Five types of biogenic structures are represented in the Bowan Park Group :
(1) Chondrites, (2) large, meandering horizontal burrows, (3) small, pellet-filled
burrows, (4) burrow-mottled limestones and (5) borings.

Chondrites

The most common burrow system in the sequence is the ichnogenus Chon-
drites. Burrows are cylindrical, smooth-walled tunnels which branch laterally
and regularly (Pl. V, Figs. a, b, d); several orders of branching are developed
with tunnels radially arranged (Pl. V, Fig. a) and horizontal to gently inclined
(Simpson, 1957). Burrow systems are spread on bed surfaces over areas up to
100 sq em, representing a working area approximately 6 em radius. Maximum
burrow penetration, inferred from vertical slabs, is 6-5 em; more commonly,
burrow penetration is 3-0 to 4-0 cm (PI. V, Fig. c). Tunnel diameter ranges
from 1-5 to 3 mm but is constant in any one burrow system ; rare tunnels reach

PROCEEDINGS OF THE LINNEAN Society or NEw SourH Wass, Vou. 99 Part 2,

V. SEMENIUK 135

4 mm diameter. Tunnel walls are generally smooth and some are lined with a
thin layer of light-coloured carbonate mud (Pl. VI, Fig. f), a feature which
presumably represents a mucus/mud lining made to maintain burrow walls.
Burrows are lined mostly when they occur in sandy host sediments such as
packstones and grainstones.

Burrows are filled with sediment, usually lighter in colour than the host
sediment. The fill commonly exhibits a concentric lamination in transverse
section (Pl. VI, Figs. a, c) and downward draping of laminae in longitudinal
sections. There is no increase in pellet content within burrows as compared
with outside, but some burrows contain a high proportion of fragmented skeletal
material (Pl. VI, Fig. e). Burrows may penetrate lithologically distinct laminae
and are filled by sediment similar to the upper layer (Pl. VI, Fig. ¢). Some tunnels
are only partly filled with sediment and the remaining void contains sparry calcite
cement. Burrows totally filled by sparry calcite are rare. Most of the Bowan
Park Chondrites appear to have been filled early since filled tunnels are reworked
by as many as four later burrows (Pl. VI, Fig. b). Although most contem-
poraneous burrows do not intersect because of some chemical control which
prevented the animal from working sediment that already had been exploited,
later burrows may intersect earlier ones (Simpson, 1957).

Chondrites occurs in dark grey lime mudstone and dark grey wackestone and
packstone that are present in most units of the Daylesford and Quondong Lim e-
stones, and in green terrigenous mudstone of Unit 1 of the Ranch Member.

Horizontal meandering burrows

Large meandering burrows that occasionally bifurcate (Pl. V, Fig. h), are
second in abundance in the sequence. As many as 30 burrows have been found
on a square metre of bedding. Buirows are circular in cross section (1 em
diameter) and tend to follow bedding planes, with some extending over distances
of Im. Numerous vertical pipes (1 em diameter) connect the horizontal tunnels
GRIEVE ohio 7g):

Tunnels may be partially to completely filled by sandy or silty mud, with
sparry calcite filling any remaining pore space (Pl. V, Fig. f) ; some tunnels are
lined with carbonate mud forming a smooth wall. Most burrow-fills, however,
are partly to completely replaced by a coarse dolomite mosaic (Pl. V, Fig. g) ;
the burrow/host sediment contact may also be obliterated. In some, dolomite
has advanced beyond the original burrow resulting in vague, cylindrical, inter-
connected patches of dolomite mosaic.

Large meandering burrows occur in dark grey skeletal wackestone of Bourim-
bla, Manooka and Gerybong Limestone Members, in burrowed pellet packstone
and skeletal wackestone that occur in the transitional facies between Gerybong
Limestone Member and Oakley Limestone Member and also in burrowed pellet
packstone at the contact of Quondong and Ballingoole Limestones.

Pellet-filled burrows

Small pellet-filled burrows are circular in cross section (1 mm diameter) and
consist of meandering and branching tubes up to 3 cm long. Some are lined
with carbonate mud. Burrows are filled partly or completely with faecal pellets
(Pl. VI, Fig. a); sparry calcite fills any void space. These burrows are best
developed in dark grey wackestones of the Manooka Limestone Member where
they are abundant between the pillars of the receptaculitid [schadites (Semeniuk
and Byrnes, 1971). Here the burrows consist of randomly oriented tubes.
Small pellet-filled burrows also occur in dark grey skeletal wackestones in Unit 2
of the Ranch Member.

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134 BIOGENIC STRUCTURES IN THE ORDOVICIAN BOWAN PARK GROUP

Burrow-mottled limestones

Burrow-mottled limestones are those sediments so thoroughly reworked by
burrowers that both the original fabric and the burrow form have been largely
destroyed. The limestones vary from homogeneous wackestones and packstones
to sediments with patches of grainstone, packstone, wackestone and lime mud-
stone. Cross sections of small to large burrows are evident in vertical slabs and
thin sections but, apart from Chondrites, the geometry of most burrows cannot
be determined since they have been obliterated by later biogenic activity.

Burrow-mottled limestones occur in some horizons of Manooka Limestone
Member, in the transitional facies between Gerybong Limestone Member and
Oakley Limestone Member (Fig. 2) and in the Davys Plains Limestone Member.

Borings
Borings are cylindrical holes (generally 2 to 4 mm in diameter) that truncate
rock fabrics and sparry calcite mosaics (Pl. VI, Fig. g) and indicate tunnelling
into cemented sediment. They are commonly filled by contemporary marine
sand or later sparry calcite. Borings also occur in brachiopods (Pl. VI, Fig. h)
and corals ; these holes are generally small (0-4 to 0-6 mm diameter) but may
reach 5 mm diameter, particularly in corals.

Borings have been found only in grainstones of Manooka and Davys Plains
Limestone Members of the Daylesford Limestone, and also in limestones of the
Quondong Limestone ; where they penetrate rock fabrics they illustrate hard
ground conditions were present. Borings, however, are rare in the sequence
and will not be discussed further here.

ROLE OF BURROWING ORGANISMS IN SYNSEDIMENTARY DIAGENESIS

Burrowing organisms have affected sediments of the Bowan Park sequence
in the following ways : (1) by vertically mixing sequences of facies, (2) by obliter-
ating original fabric and lamination and disorienting large oriented grains within
facies and (3) by fragmenting skeletal material.

Vertical mixing of facies

The ability of burrowers to mix a vertical sequence of sediments has important
geological implications as the resulting sediment may be a mixture of two or
more facies (Logan et al., 1969). Where grain types are facies-restricted and
where end-member sediment-types have been found elsewhere within the sequence
in a partly mixed or unmixed section, it is possible to ascertain how many litho-
facies have been mixed. The problem is easily resolved where one sediment
occurs aS patches in another.

Burrowing organisms in the Bowan Park sequence have been responsible for
mixing of sediments. Detailed studies of some sections have provided com-
parisons of unmixed, partially mixed, and thoroughly mixed sequences. Sections
in Unit 2 of the Ranch Member have illustrated partial mixing. Biogenic re-
working has not been intense but infiltration of overlying sediment into burrows
has produced a patchy distribution of sediment-types within the beds (Pl. VI,
Fig. c). In the Manooka Limestone Member, burrowing has been more intense
and burrowing organisms have mixed sections which originally consisted of
interbedded grainstone, packstone, wackestone and lime mudstone. At the
west end of one horizon near the base of the Manooka Limestone Member, for
example, grainstone/muddy limestone intercalations were mixed by burrowing
organisms to produce a 3 m-thick relatively homogenous burrow-mottled grain-
stone/packstone bed within a predominantly lime mudstone/skeletal wackestone
sequence. Burrowing is less intense 4 km to the east and patches of individual

PROCEEDINGS OF THE LINNEAN Society oF NEw SourH WAtgss, VOL. 99, Part 2

A

LOCALITY 1
(type section)

GLENRAE LIMESTONE
MEMBER

GERYBONG LIMESTONE
MEMBER

(muddy limestones)

MANOOKA LIMESTONE
MEMBER

BOURIMBLA LIMESTONE Por
MEMBER

SEMENIUK 135

LOCALITY 3
(east of type
section)
2km
GLENRAE LIMESTONE

Locality 2 MEMBER

(for details j
see below)

OAKLEY LIMESTONE
MEMBER
(grainstones)

BOURIMBLA LIMESTONE
MEMBER

DETAILED SECTION AT LOCALITY 2

LEGEND FOR DETAILED SECTION

(all limestones bedded
in 0.3-1.0m units)

burrow-mottled skeletal
pellet wackestone/packstone;

4 minor lime mudstone
Ss with grainstone/packstone/
wackestone patches
sree BFainstone; minor muddy
| limestone patches and

a3 burrow mottling

burrow-mottled limestone

Fig. 2.
Gerybong Limestone Members.

INTERPRETATION

tongue of grainstone;
minor biogenic mixing

tongue of grainstone
thoroughly mixed with
muddy limestones

thoroughly mixed
grainstone/muddy limestone
intercalations

tongue of grainstone;
minor biogenic mixing

A. Generalised diagram showing facies relationships between Oakley Limestone and
B. Detail at locality 2 of transitional sequence between Oakley

Limestone and Gerybong Limestone Members ; section illustrates biogenic mixing of interbedded

grainstones and muddy limestones.

136 BIOGENIC STRUCTURES IN THE ORDOVICIAN BOWAN PARK GROUP

sediments types are recognisable : the horizon consists of muddy limestone-filled
burrows and grainstone-filled burrows within a mottled grainstone to packstone
host-sediment. Finally, at the east end of outcrops, only massive grainstone
occurs and no burrows are evident.

Similar mixing occurs at localities where burrowed muddy sediments of the
Gerybong Limestone Member interfinger with and pass laterally into grainstones
of the Oakley Limestone Member (Fig. 2). The transitional section consists
mainly of thickly bedded, intercalated grainstone, skeletal pellet wackestone and
packstone, pallet packstone, lime mudstone and burrow-mottled mixtures of
these sediments. Vertical contacts between sediment types are commonly
burrowed.

The results outlined above have been used in the interpretation of environ-
ments of the Davys Plains Limestone Member (Semeniuk, 1973b). Sediments
in the member include grainstone and dark grey muddy limestones. Although
unburrowed sediments such as laminated and cross laminated grainstone are
recognisable in many horizons, portions of the member consisting of interbedded
grainstone/muddy limestone appear to have been mixed by burrowing organisms
that kept pace with sedimentation. Burrowing organisms have produced, locally,
a burrow-mottled to relatively homogeneous packstone or wackestone where
pisohtes and rounded sand grains (which are both diagnostic of grainstone
lithofacies) occur in a muddy matrix (PI. VI, Fig. d) ; more commonly, burrowing
organisms have produced a mottled limestone in which patches of grainstone
and muddy limestone are still discernible.

Obliteration of original fabric

Biogenic activity disrupts primary laminations and other sedimentary
structures and ultimately destroys all original fabric (Ginsburg, 1957). In
muddy sediments of the Bowan Park sequence, patches of original sediment are
observed where burrowing has not been intense. There are two primary fabrics :
(1) massive, unlaminated lime mudstone with scattered fossils or (2) skeletal
wackestone and packstone with laminae of silt-, sand-, or gravel-sized skeletal
debris. Intense burrowing obliterates original fabrics and produces a mottled
fabric in which circular to cylindrical outlines of filled tunnels are dominant
GEL Vieiiowze):

Burrows also disrupt oriented larger grains from original bedding positions
to form fabrics in which most grains are randomly oriented. For instance, the
large brachiopod Hodinobolus occurs beak down in growth position or as dis-
articulated valves oriented convex-up or convex-down in some unburrowed
horizons of the Bourimbla Limestone Member. Where burrows occur, articulated
and disarticulated valves are found in random orientations. Reorientation of
grains is striking where encrusting coralline organisms are present. Coralline
organisms would have encrusted only upper portions of shell gravel layers but
disruption of the fabric by burrowers has resulted in random orientations of both
shells and encrustations.

3urrows generally are uncommon in grainstones, which are bedded with
lamination, cross-lamination and graded-bedding. Burrows in these sediments
truncate sedimentary structures to varying degrees depending on the intensity of
biogenic activity.

Pragmentation of skeletal material
surrowing organisms can modify grain size (Dapples, 1938 ; Rhoads, 1967).
Feeding, ingestion or the mechanical act of excavating tunnels can comminute
skeletal material, particularly if skeletons are fragile. Intense burrowing can
produce a sediment composed of angular, highly fragmented shells, explaining
how skeletons are highly fragmented in quiet-water environments (Matthews,

PROCEEDINGS OF THE LINNEAN Society oF New Sourn WatusEs, Vou. 99, Part 2

V. SEMENIUK lo

1966). In the Bowan Park sequence, weakly burrowed horizons may contain
articulated and disarticulated complete valves of thin-shelled brachiopods,
together with recognisable sponge spicules, dasycladacean algae, trilobites,
ostracods and coral tubes. As burrows become more numerous, thin-shelled
skeletons become fragmented and form angular, skeletal sand and silt, which is
common in many dark grey burrowed lime mudstones, wackestones and pack-
stones (Pl. VI, Figs. b, e).

INFLUENCE OF BURROWS ON LATER ALTERATION

Biogenic structures have influenced distribution of some late (diagenetic or
low-grade burial metamorphic) alteration phenomena such as dolomitisation,
grain growth, silicification and stylolitisation. In addition to undergoing normal
diagenesis, limestones of the Bowan Park Group have been subjected to low-
temperature burial metamorphism of the prehnite-pumpellyite-metagreywacke
facies of Coombs (1960), as evidenced by alteration in the overlying rocks of the
Malachi’s Hill Beds (Semeniuk, 1970). It has been difficult, however, to differ-
entiate late diagenetic alteration from that produced by burial metamorphism.

Apart from some dolomite associated with faults, stylolites and chert nodules,
most dolomite in the sequence is associated with large burrows (Pl. VI, Fig. g)
and, to a less extent, Chondrites burrows. This dolomite commonly connects
with thin sheet-like dolomite mosaics that are localised along cracks and stylolites.
Dolomite appears to have replaced predominantly burrow-fill sediment, forming
a coarse interlocking mosaic with calcite patches in inter-rhomb areas. Dolo-
mitised burrows grade into undolomitised and partly dolomitised burrows ;
within these latter burrows, burrow-fills commonly are sandier than the enclosing
host sediment, suggesting selective dolomitisation of burrow-fills may have been
related to the higher porosity of material in burrows relative to the surrounding
host sediment.

Many burrow-fill sediments also exhibit grain growth mosaics, especially if
there is a contrast in character between burrow-fill material and host sediment.
Grain growth is most pronounced where burrows in lime mudstone are filled
with grainstone. Sparry calcite cement of the grainstone is recrystallised to the
extent that it is poikiloblastic. Wackestone and mudstone filled burrows in a
lime mudstone host are recrystallised to a mosaic of microspar (Folk, 1965) while
the surrounding host undergoes comparably little or no recrystallisation. This
results in burrow-fills which are granular, and lighter-coloured in thin section.
Where lime mudstone, wackestone or packstone fills a burrow in grainstone,
similar grain growth phenomena occur but mosaics are ragged at the burrow
periphery.

Less important alteration phenomena associated with burrows include
silicification, where portions of dolomitised burrows are silicified, and stylolitis-
ation, where the stylolites follow the physical discontinuity at the burrow/host
rock contact.

OCCURRENCE OF BURROWS WITHIN INFERRED DEPOSITIONAL ENVIRONMENTS

In the Daylesford Limestone, the convergence of disconformities to the east,
the dominance of grainstones and grey skeletal wackestones and packstones in
eastern sections and the occurrence of lithoclasts in the thick grainstone sequences
all suggest a positive area to the east of the depositional basin (Semeniuk, 1973)).
Grainstones and grey skeletal wackestones and packstones probably bordered
the positive area as fringing deposits. Dark grey burrowed muddy limestones
occurred in the subsiding area to the west. The convergence of disconformities
to the east and similar suites of lithologies within the Quondong Limestone
suggest a similar palaeogeographic and environmental setting for this formation.

PROCEEDINGS OF THE LINNEAN Soctety or NEw SoutH WALES, VoL. 99, Part 2

138 BIOGENIC STRUCTURES IN THE ORDOVICIAN BOWAN PARK GROUP

The lateral facies changes also are reflected (immediately above discon-
formities) in the vertical sequences which probably were deposited under trans-
eressive conditions and relate to deepening water. The sequences generally
indicate increased effects of burrowing, increased accumulation of mud relative
to grains, decreasing energy, and relatively increased reducing conditions.

Grainstones commonly form the base of Type 1 transgressive sequences
(p. 132). Grey skeletal wackestones and packstones (or brown terrigenous mud-
stones) form the base of Type 2 transgressive sequences (p. 132). This
stratigraphic position and relationship to the positive area suggests the grainstones
and grey skeletal wackestones and packstones are nearshore, shallow water
sediments. In marly sequences, brown terrigenous mudstones substituted for
grey skeletal wackestones and packstones.

Dark grey burrowed lime mudstones occur at the top of most transgressive
successions and predominate in western localities. Abundance of lime mud and
lack of shallow water indicators suggest the sediments formed under low energy
conditions, probably beneath wave base. These sediments are interpreted as
offshore deposits.

Stratigraphically the dark grey burrowed wackestones and packstones
generally occur above grainstones (or grey skeletal packstones and wackestones)
and below dark grey lime mudstone. The sediments are interpreted as having
formed in intermediate depths of water. In marly sequences, green terrigenous
mudstones substituted for dark grey wackestones and packstones.

The various burrows occurred in consistent facies throughout deposition of
the Daylesford and Quondong Limestones. For instance, Chondrites burrows
always occur in abundance within dark grey lime mudstones, wackestones and
packstones that are interpreted as offshore to intermediate facies, and which are
present in all phases of sedimentation of these formations. Bathymetry, rather
than sediment type, appears to have controlled their distribution : Chondrites
burrows are uncommon in all shallow water sediments, e.g. in grey skeletal
wackestones and packstones of the Bourimbla Limestone Member and the
Quondong Limestone and in grainstones of the Quondong Limestone and Man-
ooka, Oakley and Davys Plains Limestone Members of the Daylesford Limestone.
Chondrites distribution within reconstructed depositional environments of Ranch,
Bourimbla, Manooka, Gerybong and Oakley Limestone Members is illustrated
in Fig. 3.

Horizontal meandering burrows occur predominantly in dark grey skeletal
wackestones and packstones, sediments interpreted to have formed in inter-
mediate depths of water. The burrows overlap the distribution of Chondrites.
Their position relative to Chondrites is consistent during four depositional phases
of the Daylesford Limestone (Fig. 3). Small pellet-filled burrows appear in two
units of the Daylesford Limestone ; both appearances occur in dark grey skeletal
wackestone interpreted as sediments formed at intermediate depths (Fig. 3).

The occurrences of biogenic structures in these unique environmental
positions aid environmental interpretation of some vertically or laterally in-
complete sections. Where they occur, burrows can be used to infer relative
bathymetry. For instance, Chondrites-burrowed lime mudstones may be placed
as the deepest facies and other sediment types treated as shallow water
equivalents. Biogenic structures are used to complement information given by
other criteria and there is good correlation between environments suggested by
burrows and that suggested by petrographic and palaeontologic data.

ACKNOWLEDGEMENTS

The author wishes to thank B. D. Webby and D. P. Johnson for critically
reading the manuscript. C. Hughes assisted with photography. The work,

PROCEEDINGS OF THE LINNEAN Society or New Sours Wass, Vou. 99, Part 2

V. SEMENIUK 139

COMMON RARE

Chondrites
Large horizontal burrows

Small pellet-filled burrows

OFFSHORE

E
Upper Gerybong/

x
Oakley Limestone
SS ET LE LD LET LIT ED LE LETT SITE SE Members
x x

——X —— XX — x — X — x — KX — x — 7K —
Lower Gerybong/

Oakley Limestone
Members

x ye
—~ Oe Se Limestone
eee ee Le, Member :

Bourimbla
Limestone
Member

Unit 3,
Ranch Member

Unit 2,
(e oe Ranch Member
ee

Unit 1,
Ranch Member

LEGEND

Terrigenous mudstones with gastropods, bivalves
i ith gastropods,Alleynodictyon,
Grainstones [AA\" Fatradiun

Grey skeletal wackestones/ Dark grey eal 4
aaeketoree (with Tetradium, skeletal with Vermiporella
Eodinobolus, gastropods) wackestones/

packstones

Dark grey burrowed with Ischadites, sponges,
lime mudstones Vermiporella

Dark grey burrowed lime a - ;

mudstones (with Maclurites) BE ue rauiliniea sauorads

Fig. 3. Distribution of burrow types within reconstructed facies during various phases of
sedimentation of lower to middle Daylesford Limestone. (Alleynodictyon=cylindrical strom-
atoporoid ; Hodinobolus=inarticulate brachiopod ; Ischadites=receptaculitid ; Vermiporella=—
dasycladacean alga).

PROCEEDINGS OF THE LINNEAN Society or New SoutH WALES, VoL. 99, Part 2

140 BIOGENIC STRUCTURES IN THE ORDOVICIAN BOWAN PARK GROUP

carried out in the Department of Geology, University of Sydney, was supported
by a Commonwealth Postgraduate Scholarship. Costs of manuscript preparation
were met by a University of Western Australia Research Grant.

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in South Florida. Jn Le Blanc, R. J., and Breeding, J. G., (eds), “‘ Regional aspects of
carbonate deposition.”’ Soc. Econ. Palaeontologists and Mineralogists Spec. Pub., 5: 80-99.

GoupRine, R., 1962.—The trace fossils of the Baggy Beds (Upper Devonian) of North Devon,
England. Palaeontol. Z., 36 : 232-257.

Loacan, B. W., Harpine, J. L., AnR, W. M., Wittiams, J. D., and SNEAD, R. G., 1969.—Carbonate
sediments and reefs, Yucatan Shelf, Mexico. Am. Assoc. Petroleum Geologists Mem., 11 : 1-196.

MartHews, R. K., 1966.—Genesis of Recent lime mud in Southern British Honduras. J. Sediment.
Petrol., 36 : 428-454.

PackuHam, G. H., (ed.) 1969.—The geology of New South Wales. J. Geol. Soc. Aust., 16: 1-654.

Ruoaps, D. C., 1967.—Biogenic reworking of intertidal and subtidal sediments in Barnstable
Harbor and Buzzards Bay, Massachusetts. J. Geol., 75 : 461-476.

SEILACHER, A., 1962.—Palaeontological studies on turbidite sedimentation and erosion. J.
Geol., 70 : 227-234.

, 1964.—Biogenic sedimentary structures. In Imbrie, J., and Newell, N., (eds)
Approaches to palaeoecology. New York: John Wiley : 296-316.
, 1967.—Bathymetry of trace fossils. Marine Geol., 5 : 413-428.

SEMENIUK, V., 1970.—The Lower-Middle Palaeozoic stratigraphy of the Bowan Park area, central
western New South Wales. J. Proc. Roy. Soc. N.S.W., 103 : 15-30.

, 1973a.—The stratigraphy of the Bowan Park Group, New South Wales. J. Proc. Roy.
Soc. N.S.W., 105 : 77-85.

, 1973b.—Nearshore to offshore facies, and depositional history of the Ordovician Dayles-
ford Limestone, New South Wales. J. Geol. Soc. Aust., 20 : 449-463.

SEMENIUK, V., and Byrnes, J. G., 1971.—Occurrence and significance of Ischadites Murchison in
Ordovician limestones at Bowan Park, New South Wales. J. Geol. Soc. Aust., 18 : 235-241.

Smvpson, S., 1957.—On the trace fossil Chondrites. J. Geol. Soc. Lond., 112 : 475-500.

EXPLANATION OF PLATES

PLATE V

Fig. a. Chrondites burrows showing radial arrangement of tunnels on bedding plane surface
of limestone. Upper Bourimbla Limestone Member of Daylesford Limestone, 8.U.P. 23859.

Fig. b. Chondrites burrows showing branching pattern of tunnels on bedding plane surface
of limestone. Quondong Limestone, 8.U.P. 23860.

Fig.c. Vertically oriented slab showing depth of penetration (arrowed) of Chondrites burrows.
Bourimbla Limestone Member of Daylesford Limestone, U.8.G.D. 42195.

Fig. d. Chondrites burrows and large horizontal burrows on bedding plane surface of lime-
stone of upper Gerybong Limestone Member near “‘ Quondong”’ property. Lens cap diameter
approximately 5 cm.

Fig. e. Burrow-mottled fabric produced by intense bioturbation. Ranch Member of
Daylesford Limestone, U.S.G.D. 46676, polished slab.

Fig. f. Vertical slab of (undolomitised) large horizontal burrow (left of photo) showing
sediment partly filling tunnel and sparry calcite filling remaining void space ; surrounding limestone
is burrow-mottled. Glenrae Limestone Member of Daylesford Limestone, U.8.G.D. 46737.

PROCEEDINGS OF THE LINNEAN Soctety or New SourH Waxzs, Vou. 99, Part 2

Proc. LINN. Soc. N.S.

V., Vol. 99, Part 2 PLATE V

PLATE VI

Proc. LINN. Soc. N.S.W., Vol. 99, Part 2

V. SEMENIUK 141

Fig. g. Short vertical pipes that connect large horizontal burrows ; burrow-fills have been
dolomitised. Gerybong Limestone Member of Dayvlesford Limestone, U.S.G.D. 46732, acid-etched
slab.

Fig. h. Dolomitised large horizontal meandering and branching burrows on bedding plane
surface of limestone of lower Gerybong Limestone Member, 1-5 miles east of ‘‘ Quondong ”’
homestead.

PLATE VI

Fig. a. Small pellet-filled burrows (two are arrowed), and section through Chondrites at
lower portion of photomicrograph. Ranch Member of Daylesford Limestone, U.S.G.D. 47038,
thin section, plane polarised light.

Fig. b. Chondrites burrows (light grey) transecting earlier burrows (dark grey) ; note abund-
ance of highly fragmented angular skeletal grains. Ranch Member of Daylesford Limestone,
U.S.G.D. 46681, thin section, plane polarised light.

Fig. c. Chondrites tunnel filled with skeletal wackestone penetrating skeletal pellet grain-
stone; note concentric lamination exhibited by burrow-fill. Ranch Member of Daylesford
Limestone, U.S.G.D. 46671, thin section, plane polarised light.

Fig. d. Pisolitic burrow-mottled wackestone/packstone formed by burrowing organisms
mixing a pisolitic grainstone and muddy limestone. Davys Plains Limestone Member of Dayles-
ford Limestone, U.S.G.D. 47049, thin section, plane polarised light.

Fig. e. Angular skeletal fragments occurring predominantly in Chondrites burrows. Gery-
bong Limestone Member of Daylesford Limestone, U.S.G.D. 41799, thin section, plane polarised
light.

Fig. f. Carbonate silt (arrowed) lining wall of Chondrites burrow. Manooka Limestone
Member of Daylesford Limestone, U.S.G.D. 46704, thin section, plane polarised light.

Fig. g. Boring in sparry calcite-cemented grainstone ; note truncated grain at right margin of
bore (arrowed) ; the hole has been partly filled with sand. Davys Plains Limestone Member of
Daylesford Limestone, U.S.G.D. 41814, thin section, plane polarised light.

Fig. h. Smooth walled, sparry calcite-filled boring (arrowed) in brachiopod ; the hole truncates
internal shell structure and is parallel to the shell surface. Quondong Limestone, U.S.G.D. 47043,
thin section, plane polarised light.

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Proceedings, Volume 99, Part 2, 1975

CONTENTS

ALLEN, G. R. Four new damselfishes (Pomacentridae) from the south-
west Pacific...

BALLANTYNE, B. Powdery mildew on cucurbitaceae: identity, dis-
tribution, host range and sources of resistance .

Moore, K. M. The Glycaspis spp. ee ee +? ae associated
with Hucalyptus camaldulensis a : ai AG

SEMENIUK, V. Biogenic structures in the Ordovician Bowan Park Group,
New South Wales ..

Page

87

100

121

129

Proceedings of the
Linnean Society

of New South ee:

y Ma vine Biologica

oe Ae

ened) March Sth, 1975 LIB -
f= MAY 15

VOLUME 99

PART 3

No. 439

Registered for posting as a periodical—Category A.

The Linnean Society of New South Wales

Founded 1874. Incorporated 1834

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

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N.S.W. 2000, Australia

A NEW SPECIES OF 4AMBLYSEIUS BERLESE
(ACARINA : PHYTOSEIIDAE) FROM APPLE IN AUSTRALIA

H. A. DENMARK* AND E. SCHICHAT

[Accepted for publication 17th April 1974]

Synopsis
A new species of mite (Acarina : Phytoseiidae) is described from apple in New South Wales,

Australia : Amblyseius lentiginosus Denmark and Schicha n. sp. is closely related to Amblyseius
rhabdus Denmark.

INTRODUCTION
The species of phytoseiid mite described in this paper was collected in
commercial orchards of Granny Smith apple trees at Bathurst, New South Wales.
This species was also found in an orchard of young, unbearing apple trees and on
single neglected ‘“‘ backyard trees ”’ during the years 1971 to 1973.

Measurements listed are the mean of three specimens in microns.

Genus AMBLYSEIUS Berlese

Amblyseius Berlese, 1914, Redia, 10: 143. Type of the genus:
Zercon obtusus Koch, 1839, by indication of Berlese (1914).
Amblyseius (Amblyseius) lentiginosus, 0. sp.

(Figs 1-8)

Diagnosis. Amblyseius lentiginosus n. sp. is closely related to A. rhabdus
Denmark (1965), but the latter has no teeth on the movable digit of the chelicerae
and the spermathecal atrium is more rod-like. A. lentiginosus has lunate areas
and numerous pores on the dorsal shield, three teeth on the movable digit of the
chelicerae, and the spermathecal atrium is about twice as wide as in A. rhabdus.

FEMALE

Dorsum: The dorsal shield (Fig. 1), length 352-360, width at L, 199-202,
is well sclerotised and smooth, with 11 pairs of pores and 17 pairs of setae, six of
which are dorsal, two median (one anterior and one posterior), four prolateral
and five postlateral. The setae measure in length :
Di —50-30. 5-6. Dao Di D —( D(_ os ti 0008), nO
ie — 6-7 = lu, —85-90): 7 > and? lh, —7_-9 5 ‘—203-210; M, —3—-4;
M, =123-130. Setae D,, L, and Ly, are longer than the distances between their
bases and the bases of the setae following next in line. Setae 8,, 14-16 and 8,,
9-10 long, are located on the dorsal interscutal membrane.

The peritremes extend forward beyond the bases of setae D,.

Venter: The sternal shield (Fig. 2) measures 116-130 in length and 101-109
in width. It bears three setae and two pairs of pores near the first and the third
pair of setae. The fourth pair of setae is placed on metasternal shields which
each bear a caudomedial pore. The genital shield (Fig. 2), width 80-100, is
normal with a pair of setae and a straight posterior margin. The smooth,
pentagonal, ventrianal shield (Fig. 2) is 116-130 long and 101-109 wide. It is

* Division of Plant Industry, Florida Department of Agriculture and Consumer Services,
Gainesville, Florida, 32601, U.S.A. (Contribution No. 287).

7 N.S.W. Department of Agriculture, Agricultural Research Station, Bathurst, New South
Wales, 2795.

PROCEEDINGS OF THE LINNEAN SocretTy oF NEw SoutH WALES, Vou. 99, Part 3

146 A NEW PHYTOSELUD MITE

Fig. 1. Dorsal shield of female.

PROCEEDINGS OF THE LINNEAN Society or New SouruH WALES, Vou. 99, Part 3

H. A. DENMARK AND E. SCHICHA 147

provided with three pairs of preanal setae and a pair of pores caudomedial to the
third pair of setae.

Spermatheca : (Fig. 3) The major duct is nearly as broad as the tube-like
cervix ; an atrium is not distinct.

f §
% 1

Fig. 2. Venter (sternal shield, genital shield, ventrianal shield) of female.

Chelicera : (Fig. 4) The fixed digit is 40-41 long and bears seven teeth and
a pilus dentilis. The movable digit is 50-51 long and is provided with three teeth.

Legs: (Fig. 5) Sge IV, 90-101 ; Sti IV, 91-94 ; St IV, 75-87.
Peritrematal Shield : (Fig. 6).

PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WAtgEsS, Vou. 99, Part 3

148 A NEW PHYTOSEIID MITE

Fig. 3. Spermatheca of female.

Fig. 4. Chelicera of female.

PROCEEDINGS OF THE LINNEAN Society or New SourH Wares, Vou. 99, Part 3

H. A. DENMARK AND E. SCHICHA 149

(Fig. 5. Leg IV of female.

Fig. 6. Peritrematal shield of female.
Fig. 7. Ventrianal shield of male.
Fig. 8. Spermatodactyl of male.

PROCEEDINGS OF THE LINNEAN Society or NEw SoutH WALES, VoL. 99, Part 3

150 A NEW PHYTOSEHD MITE

MALE

Dorsum: The dorsal shield measures 283 in length and 199-202 in width
at Ly. The chaetotaxy of the shield resembles that of the female but the setae
are shorter. D,=22-23; D,=—5-7; D,=3-5; D,=3-6; D;=—3-6; D,—6-7 ;
bb, =38-43 ;) b,—3-7 ; L—3—7 3), —712-715); Li, —7 5 7 ; lu, —5—-7 = 6-7
Lig =152-154 ; M, =4-5 ; M,=87.

Venter: The ventrianal shield (Fig. 7), length 109 and width 159, bears three
pairs of preanal setae and four pairs of pores in addition to a fifth pair which is
caudo-medial to the third pair of setae.

Spermatodactyl : (Fig. 8) The lateral process of the foot (length 7-9) is
rounded ; the toe is a rounded knob.

Holotype. Female from Bathurst, N.S.W., Australia, 28.1.1971, E. Schicha,
on apple trees. Deposited in Biological and Chemical Research Institute,
Rydalmere, N.S.W., Australia.

Paratypes. Six females and six males taken from the same locality as the
holotype. Deposited as follows : two females and two males at Biological and
Chemical Research Institute, Rydalmere ; two females and two males at Division
of Plant Industry, Florida Department of Agriculture and Consumer Services,
Gainesville, Florida, U.S.A. ; two females and two males at the South Australian
Museum, Adelaide, 8.A., Australia.

Reference

DENMARK, H. A., 1965.—Four new Phytoseiidae (Acari : Mesostigmata) from Florida. Fla Ent.
48 : 89-95.

PROCEEDINGS OF THE LINNEAN SocieTy or New Souru WaxsEs, Vou. 99, Part 3

A NEW SPECIES OF TILEFISH (FAMILY BRANCHIOSTEGIDAE)
FROM EASTERN AUSTRALIA

JAMES K. DOOLEY* AND JOHN R. PAXTON}
[Accepted for publication 20th March 1974]

Synopsis

Until recently the genus Branchiostegus was represented in Australia only by B. wardi, thought
to be restricted to waters off southern Queensland and New South Wales. A specimen of this
species has been taken recently from New Caledonia and a new species of Branchiostegus trawled
off New South Wales. The new form is characterised by a colour pattern of 18-19 black body
bars, two broad yellow bands on the caudal fin, yellow spots on the dorsal fin and 67—72 pored
lateral line scales. Both Australian species are benthic carnivores. 6. wardi appears to spawn
a number of times per year.

INTRODUCTION

The genus Branchiostegus is represented by eleven nominal species found
mainly in the tropical Indo-Pacific ; exceptions are B. semifasciatus from West
Africa and B. sawakinensis which ranges into the Red Sea from the Indian Ocean.
The only species previously known from Australia, B. wardi, is not restricted to
the tropics but occurs off southern Queensland and New South Wales (Marshall,
1965).

A revision of the family Branchiostegidae has recently been completed
(Dooley, 1974). During this study a second Australian species became apparent
which is strikingly different from B. wardi, particularly in colour pattern.

MATERIALS AND METHODS

Measurements were made with dial calipers to the nearest 0-5 mm and
conform to those defined by Hubbs and Lagler (1958). Measurements are
expressed as per cent standard length (SL) or per cent head length (HL). Head
length was taken from the tip of the snout to the tip of the opercular spine.
Cheek depth was measured vertically from the lower rim of the orbit to the
lower margin of the preoperculum. Opercular length is the horizontal distance
from preoperculum to opercular spine ; opercular scale counts were also made
along this line. Vertebral counts were made from radiographs.

The holotype has been deposited at the Australian Museum, Sydney (AMS)
and the paratypes at the Australian Museum, at the Queensland Museum,
Brisbane (QM), at the California Academy of Sciences, San Francisco (CAS), at
the United States National Natural History Museum, Washington, D.C. (USNM),
at the British Museum of Natural History, London (BMNH) and at the Museum
National d’Histoire Naturelle, Paris (MNHN). AIl specimens were obtained by
bottom trawl.

Branchiostegus serratus n. sp.

Holotype. AMS I. 16207-004, 285 mm Sh, Neweastle, New South Wales,
Australia, 8 May 1971, Sydney Fish Markets, J. Paxton.

Paratypes. New South Wales: AMS I. 16207-002, USNM 209532, USNM
209533, (3) 245-290 mm SL, data as holotype ; AMS I. 15916—-003, —004, —005,

* Institute of Marine Sciences, University of North Carolina, Moorehead City, North Carolina,
28557, U.S.A. Present address: Department of Marine Biology, Institute of Marine Sciences,
Adelphi University, Garden City, Long Island, New York, 11530, U.S.A.

+ Department of Ichthyology, The Australian Museum, College Street, Sydney, N.S.W., 2000.

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutH WALES, VOL. 99, Part 3

152 A NEW TILEFISH FROM EASTERN AUSTRALIA

-—006, (4) 263-275 mm SL, Newcastle, 2 Feb. 1971, N.S.W. State Fisheries ;
AMS IB. 5074, 240 mm SL, off Lake Illawarra, Jan. 1961, J. Woore; AMS I.
15885-006, 228 mm SL, Newcastle (33° 13’-32’ S ; 151° 50’-52’ EB), 27 Oct. 1970,
110-150 m depth, T. Gorman and T. Johnson ; AMS IB. 2908, 405 mm SL, off
Coff’s Harbour, 21 Jan. 1953, J. Woore ; AMS I. 17312—-001, —002, -003, —004,
(4) 237-302 mm SL, BMNH 1973.7.17.3, 232 mm SL, MNHN 1973-36, 235 mm
SL, CAS 28355, 229 mm SL, Newcastle, Sept. 1971, Sydney Fish Markets,
J. Paxton. Queensland : QM I-8707, 268 mm SL, N.E. Cape Moreton, 162 m
depth ; QM I-8968, 213 mm SL, off Cape Moreton.

Diagnosis. The new species is distinguished from all congeners by the
following combination of characters: the presence of a series of 18 or 19 dark
vertical bars on the dorsal portion of the body, 67-72 pored lateral line scales,
usually seven dorsal spines and the absence of dark pigmented areas on the
operculum, dorsal head ridge and pectoral axillae.

DESCRIPTION

The following counts are of the holotype, with those showing variation in the
19 paratypes in parentheses : dorsal fin rays VII, 15 (VI (2), VII (17)); anal II,
12 (12 or 138); pectoral 17 (17 or 18); pelvic I, 5; principal caudal rays 17;
cheek scales from preopercular angle to orbital rim 10 (10-13) ; opercular scale

TABLE 1
Frequency distribution of pored lateral line scales in' Branchiostegus wardi and B. serratus

A8:: 49.50. 51 52-662 .67. 68) 69 (70,7 seo

B. wardi Pas os ats Ae Deals: 4 1 4
B. serratus os Ae MA 55 2 4 3 5 2 4 6

rows 7 (5-7); scales above lateral line 8 (7-10); scales below lateral line 22
(20-25) ; pored lateral line scales to hypural crease 70 +2 on tail (67-72, Table 1) ;
gill rakers 7+11 (18-20, Table 2); branchiostegal rays 6; vertebrae 10+14.

Measurements are given as per cent SL in Table 3 ; the following measurements
are additionally presented as per cent HL: head depth 98° (98-108) ; snout
length 39% (36-45); upper jaw 41% (37-43) ; lower jaw 48% (45-51); cheek
depth 44%, (39-47); opercular length 27% (24-29); snout to upper margin of
preoperculum 77°, (74-79); orbit diameter 26% (25-33); suborbital depth
23%, (19-25).

TABLE 2
Frequency distribution of the number of first arch gill rakers in Branchiostegus wardi and B. serratus

Gill Rakers (upper and lower limbs)

18 19 20 ahi 22 23 x
B.wardi .. a ALS of ne Se 8 a 3 1 19-9
B. serratus 7 ae ies A as 10 9 1 18°6

Head blunt, snout steep, body elongate. Mouth oblique, upper jaw
moderately protrusile; maxillary anteriorly reaching to below first nostril,
posterior extension to below centre of pupil. Upper lip set in groove along
entire edge; lower lip not grooved medially. Teeth on upper jaw in double

bands, inner row of very fine canine teeth posteriorly, widening into four to five
rows anteriorly ; outer row of larger ineurved canines with enlarged teeth

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JAMES K. DOOLEY AND JOHN R. PAXTON 153

anteriorly and single large canine at extreme posterior margin of upper jaw.
Lower jaw with fine canine teeth in patch on either side of symphysis ; larger
incurved canines in single row along outer jaw margin. No teeth on palatines,
vomer or tongue ; well-developed pharyngeal teeth.

Lateral line pores easily visible in low curve, passing slightly dorsal to
middle of body bars and extending just past hypurals. Cephalic lateral line
with five lower jaw pores from symphysis to preopercular margin, four-five pores
in preorbital series, ten-twelve pores circumorbital, two pores below anterior
nostril and one pore above posterior nostril ; other cephalic pores present but
not easily visible.

TABLE 3
Measurements of B. wardi and B. serratus in per cent standard length

B. wardi B. serratus
Holotype Specimen sats Holotype Paratype hive
AMS IA. 5130 Ranges X AMS 1.16207— Ranges x
(n= 18) 004 (n=19)

Standard length (mm) ee 327 215-336 263 285 213-405 267

Body depth .. ure = 26-0 22-6-26-1 24-57 26-1 24-6-27-7 26-77
Body width .. a ee 14-1 11-8-13-3 12-55 13-2 11-8-13-7 12-82
Dorsal fin length ah us 54-1 56-1-62-1 58-87 60-7 55:5-63-:2 60-88
Anal fin length ae a 29-4 27-4-31-:6 29-71 33:2 30-3-32:8 31-52
Pectoral fin length .. a. 22-6 22-7-25:1 23-87 23°7 22-8-25-5 24-29
Pelvic fin length ae ae 14-1 13-0-15:3 14-07 11-2 11-4-14-1 12-48
Peduncle length Ae ie 16-2 15-2-17-4 15-88 16-4 15:6-18-4 16-76
Peduncle depth AG ve 11-6 10:0-14-:2 10-88 11-4 10-3-11:9 | 11-01
Head length .. is a 27°5 25:-1-27-6 26-50 25-4 23°8-26:3 25:62
Head depth .. ae 8 24-5 21-2-26-4 23-73 24-8 25:1-27-:9 26:32
Snout length .. BS ae 11-2 8-6-11-1 9-86 10-0 8-9-12:0 10-57
Predorsal length Be 5 31-8 28:9-35-7 31-04 30-9 28-6-32°3 30-99
Length of upper jaw .. a6 11-3 10-0-12-3 11-21 10-3 9-5-11-4 10-61
Length of lower jaw .. ie 10-1 11-7-13-2 12-51 12-3 11-3-13:0 12-30
Opercular length a oe eu 6:9- 8-0 7-26 6-8 5:6— 7:3 6-51
Diameter of orbit So ie 5-7 5:8— 7:8 6-82 6-7 6-2— 8-5 6-97
Cheek depth .. at ae 11-3 9-1-11-:0 10-13 11-1 9-4-11-8 10-86
Suborbital depth A Me 5:8 4-4—- 5-7 5:07 5:8 4-6— 6-4 5-78

Gill membranes free from isthmus and notched medially. Raised medial
ridge on top of head from above posterior margin of orbit almost to dorsal origin.
Anterior nostril within a thin cutaneous tube, posterior nostril open and oval
Shaped. Preopercular angle approximately 100-115°, with fine serrae on upper
limb, less distinet serrae on lower limb. Operculum ending in single blunt spine
or tab. Pseudobranch well developed ; gill rakers blunt, generally three or more
times longer than wide, although reduced toward distal ends of limbs.

Seales large, embedded in pockets, ctenoid over most of body, cycloid in
head region ; scales on cheek, operculum and extending on top of head to posterior
margin. of pupil. Scales on base of pectoral ; caudal fin finely scaled, other fins
naked ; body with mostly regenerated scales.

Dorsal fin continuous, origin above pectoral bases ; dorsal spines slender,
first two close together, united at their bases and slightly shorter than remaining
five spines ; latter nearly equal in length and about ? length of anterior seven
rays. Rays 8-12 about + longer than anterior rays ; ray 13 elongate, reaching
just past hypurals and + longer than next longest dorsal ray ; rays 14 and 15
progressively shorter.

Anal fin continuous, origin below fourth dorsal ray ; two anal spines slender
and close together ; first spine about 4 length of second, second spine $ or less

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154 A NEW TILEFISH FROM EASTERN AUSTRALIA

length of anal rays. First three rays progressively longer, fourth ray about 4
length of all but last two rays ; penultimate anal ray elongate, reaching base of
hypurals.

Pectoral fins pointed, with scaled bases ; tip of fin reaching to level of anus ;
rays branched except stout first ray ; length of first ray 4 or less than longest
pectoral ray. Pelvic fins rounded or slightly pointed, inserted below posterior
margin of pectoral base and reaching to middle of pectorals. Caudal fin truncate,
tips extended, dorsal tip longer than ventral tip ; 17 principal and 15 branched
rays ; 11 dorsal and 10 ventral procurrent caudal rays.

Colour of holotype silver-white with some overlying yellow-orange and 18
(18 or 19 in paratypes) dark blue-violet wedge-shaped bars ; bars shorter anteriorly
and posteriorly, each tapering to a point about in line with lower margin of orbit.
Iris golden, black area on dorsal margin of orbit ; cheeks silver-white with an
oblique dark band reaching from preoperculum nearly to orbit ; snout yellow-
orange with black area on medial portion of upper lip; dorsal portion of head
coloured as snout with overlying greyish hue, a raised yellow ridge on top of

Fig. 1. Branchiostegus serratus.

head from vertical of posterior margin of orbit to near dorsal origin. Dorsal fin
membrane dusky, spinous portion dusky with narrow yellow margin; some
diffuse yellow in front of first ray and elliptical yellow spots along dorsal margin
between each ray, decreasing between soft rays 8 and 9 and disappearing between
rays 12 and 13; last ray dusky. Anal fin membrane translucent. Pectoral fin
opaque with black edging on first and second rays. Pelvic fins translucent.
Caudal fin base orange ; central portion yellow with medial black area not reaching
posterior margin ; medial posterior margin edged in black, dorsal and anal lobes
black (Fig. 1).

Distinguishing Features. Three species of Branchiostegus have a series of
vertical dark body bars, B. serratus, B. semifasciatus from West Africa and B.
doliatus from East Africa and Mauritius. B. semifasciatus has six dorsal spines
(usually seven in B. serratus), 47-50 pored lateral line scales (67-72 in B. serratus),
a dark head ridge and large dark area above the pectoral fin axillae and behind the
dorsal margin of the operculum (both light in B. serratus). B. doliatus has six
dorsal spines, 16-18 vertical body bars (18-19 in B. serratus), and a dark blotch
on the operculum (none in B. serratus). The two Australian species can be
distinguished by coloration, with B. wardi lacking vertical bars, pored lateral
line scales (Table 1), and snout profile, with B. serratus having a steeper snout.

Etymology. The specific name is in reference to the series of dark body bars.

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JAMES K. DOOLEY AND JOHN R. PAXTON 155

DISTRIBUTION AND BIOLOGY

Branchiostegus serratus is known only from the coast of New South Wales,
where it is trawled from Coff’s Harbour (30° 20’ 8) to Lake Ilawarra (34° 30’ 8).
The only recorded depth range is 110-150 m, from a State Fisheries trawl off
Newcastle (33° S). The other Australian species, B. wardi, is known from Noosa
Heads, southern Queensland (26° 25’ S) to off Broken Bay, New South Wales
(33° 30’ 8). A 410 mm specimen of this species was recently taken off the edge
of a reef at 250 m off Noumea, New Caledonia (P. Fourmanoir, pers. comm.).
Apparently both species are trawled in similar depths off New South Wales.

Fluctuations in the abundance of the North American tilefish Lopholatilus
chamaeleonticeps have been well documented (Collins, 1884). The limited
available evidence indicates that the Australian species may also undergo
population fluctuations. B. wardi was first collected off Queensland in 1928
(Marshall, 1928) and a second specimen was trawled in 90-110 m off Port Stephens,
New South Wales, in 1931 (Whitley, 1932). In 1972-73 from 50 to 300 kg of
B. wardi per day were sent to the Sydney wholesale markets from the Newcastle-
Port Stephens area. Fewer B. serratus arrived at the markets, from none to
about 25 kg per day. Neither the Thetis Expedition of 1898, which trawled
off Port Stephens—Newcastle in 60-100 m (Waite, 1899 : 20), nor the Hndeavour
Collections of 1909-1910, which were taken from 30-110 m off Port Stephens-
Neweastle (McCulloch, 1911: 3, 23), took any specimens of Branchiostegus. It
is possible that both species are more abundant now than they were 60 or 70
years ago. However, this is impossible to verify since the amount of fish
marketed in past years cannot be broken down by species ; moreover, methods
of trawling have changed over the years, further making any comparison of
numbers unreliable.

Stomach contents of 12 specimens of B. wardi and 7 specimens of B. serratus
were examined. Both species are benthic carnivores, with remains of fish
(Apogonops anomalous), molluses (bivalves and gastropods), crustaceans (crabs,
amphipods and stomatopods) and polychaetes in the stomachs and intestines
of both species.

All of the type material of B. serratus proved to be mature. Selected market
specimens of B. wardi (totalling 31) were dissected to determine the sex and
reproductive state. A deviation from the expected 50/50 sex ratio was found
both for B. wardi and for B. serratus. B. wardi specimens under 300 mm were
predominantly females ; above 300 mm, only males were identified. Twenty-
nine specimens of B. serratus were examined. Below 280 mm there were
approximately equal numbers of males and females, but, as in B. wardi, only
males were identified in the larger sizes. It can be hypothesised either that only
males reach a larger size or that there is protogynous sex reversal. There is as
yet no strong evidence for the latter hypothesis. Ovaries from specimens of
B. wardi taken in June, July and September 1972 and in January 1973 were
removed, preserved in 10°, formal-saline and later dissected. The ovaries
measured 35-50 mm in length and 10-20 mm in diameter ; ova of various sizes
ranging from 0:2—0-8 mm in diameter were distributed evenly in the ovaries.
No seasonal differences in gonad condition were noted. The range of ova size
and the presence of large ova in all ovaries examined are indicative of multiple
Spawning during the year. The only ovaries of B. serratus examined were
collected in June ; they were also filled with ova from 0-2—0-9 mm in diameter.
The larvae, although unknown, would probably resemble those of B. japonicus
(Okiyama, 1964). Larvae in four of the five tilefish genera (larvae of Lopholatilus
are unknown) have been found to have similar arrangements of spinules and head
serrations.

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156 A NEW TILEFISH FROM EASTERN AUSTRALIA

ACKNOWLEDGEMENTS

W. Brinsley, Fisheries Inspector at the Sydney Fish Markets, extended
every courtesy in making available both specimens and information. T. Gorman
and T. Johnson of N.S.W. State Fisheries provided specimens from FRV Kapala.
P. Fourmanoir provided information on and a photograph of the Noumea
specimen. B. Campbell (QM) and V. Springer (USNM) loaned specimens under
their care. M. Peoples drew Fig. 1. J. Randall and B. Russell made suggestions
on the manuscript. To all goes our appreciation.

References

Cottins, J. W., 1884.—History of the tilefish. Report U.S. Fish Commission, 10: 237-294.

Dootey, J. K., 1974.—Systematic revision and comparative biology of the tilefishes (Perciformes :
Branchiostegidae, Malacanthidae). Ph.D. dissertation, University of North Carolina.

Husss, C. L., and Laaumr, K. F., 1958.—F shes of the Great Lakes region. rev. ed. Bloomfield Hills,
Michigan : Cranbrook Institute of Science.

MarsuHatt, T. C., 1928.—Ichthyological notes, No. 3. Mem. Qld. Mus., 9 : 189-193.

, 1965.—Fishes of the Great Barrier Reef. Sydney : Angus and Robertson.

McCuutoc#, A. R., 1911.—Report on the fishes obtained by the F.I.S. “‘ Endeavour ’”’, on the
coasts of New South Wales, South Australia and Tasmania. PartI. Jn Zoological results of
the fishing experiments carried on by the F.1.8. ‘‘ Endeavour ”’, 1909-1914. Vol.1. Sydney :
Commonwealth Department of Fisheries : 1-87.

Oxtyama, M., 1964.—Early life history of the Japanese blanquillos, Branchiostegus japonicus.
Bull. Jap. Sea Regional Fish. Research Lab., 13 : 1-14.

Waits, E. R., 1899.—Scientific results of the trawling expedition of H.M.C.S. “‘ Thetis ’’, off the
coast of New South Wales. Part I: Introduction. Mem. Aust. Mus., 4: 1-23.

Wuittey, G. P., 1932.—Studies in ichthyology, No. 6. Rec. Aust. Mus., 8 : 321-348.

PROCEEDINGS OF THE LINNEAN Socitgery oF New Souru WALES, Vou. 99, Part 3

THE SYSTEMATIC AND PHYLOGENETIC POSITION OF
PALAEOSPONGILLA CHUBUTENSIS (PORIFERA : SPONGILLIDAE)

A. A. RACEK* AND F. W. HARRISONT
[Accepted for publication 19th June 1974]

Synopsis
Palaeospongilla chubutensis Ott and Volkheimer (1972), the first fossil spongillid ever recorded,
was detected during an examination of stromatolithic crusts of blue-green algae from lacustrine
sediments of the Cretaceous of Patagonia. It is represented by a fully intact skeletal arrangement
of megascleres and microscleres, both of two types, and also contains numerous gemmules with
gemmoscleres in situ.

Based on a thorough reexamination of parts of the type material, which demonstrates the
early perfection of gemmule-producing freshwater Demospongiae, the present study reveals
additional criteria which are used to interpret more precisely the relationship of the Mesozoic
fossil to extant genera and species. P. chubutensis displays gemmoscleral characteristics from
which both acerate and birotulate spicular types could have evolved by divergence, and it is
significant to find it the almost direct ancestral form of the extant genus Radiospongilla which has
retained identical gemmular and gemmoscleral criteria and which today represents such a central
stock. Furthermore, the fossil spongillid is obviously closely related to two additional extant
genera, 1.e. Spongilla and Pectispongilla, which also share some of its structural characteristics.

Recent advances in the fields of palaeolimnology, plate tectonics, and continental drift
provide a feasible background for speculation on spongillid evolution, dispersal routes and
distribution since the occurrence of the Patagonian fossil, some 100 million years ago.

INTRODUCTION

The discovery of extensive fully fossilised crusts of the spongillid
Palaeospongilla chubutensis in lacustrine sediments of the Cretaceous in the
Chubut River Valley of Patagonia, by Ott and Volkheimer (1972), can only be
described as exciting. Although spicular remains of freshwater Porifera, mostly
comparable with those of extant genera and species, have been recorded from
a number of lentic sediments of Tertiary age (Traxler, 1895, 1896 ; Reul, 1954 ;
Racek, 1966, 1969, 1970), no palaeontological evidence could hitherto be
obtained for Mesozoic occurrences of gemmule-producing freshwater Demos-
pongiae, nor was it considered likely that a well-preserved spongillid fossil could
ever be found. Factors preventing successful fossilisation of freshwater sponges
would seem to include both their comparatively flimsy skeleton, and, in particular,
the established rapid deterioration of their symplasm after death. The
exceptionally perfect preservation of P. chubutensis, over a period of more than
100 million years, can thus only be explained by the fact that its entire surface
was rapidly covered by a thick layer of calcareous algae (Cyanophyceae) which
smothered the sponge itself and simultaneously prevented it from being crushed
during sedimentation.

Realising the uniqueness of their unexpected discovery, made during a
routine examination of stromatolithic crusts of blue-green algae, Ott and
Volkheimer (1972) proceeded with the description of P. chubutensis, which they
rightly considered generically and specifically different from any extant spongillid
hitherto recorded. However, in spite of the clarity and morphometric accuracy
of the original description, the present authors considered it imperative to
reexamine parts of the type material in order to attempt a reliable assessment
of the fossil’s systematic and phylogenetic position in line with the recent
comprehensive revision of Penney and Racek (1968), which was unavailable to
the describing palaeobotanists. This reexamination, made possible by courtesy

* School of Biological Sciences, University of Sydney, New South Wales, 2006.

{ Present address : Department of Anatomy, Albany Medical College of the Union University,
Albany, New York 12208, U.S.A.

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158 POSITION OF PALAEOSPONGILLA CHUBUTENSIS

of Dr. E. Ott, revealed additional information of multidisciplinary significance,
which enabled the present authors to solve a number of problems previously
encountered.

MATERIAL AND METHODS

The fossilised remains of P. chubutensis, used for this study, consisted of
three slides containing thinly ground sections of a topotype, i.e. material adjoining
that selected as a holotype by Ott and Volkheimer. Although these sections
permitted most useful general observations, and were at least as informative as
the slides used for the original description of the sponge, it became necessary to
separate the various spicules from the rocky matrix so that their detailed
structures could be thoroughly examined. This was first achieved by removing
the mounting medium of the slides with xylene and acetone, after which the
matrix was treated with boiling nitric acid in order to dissolve all non-siliceous
matter and to reveal the structural characteristics of the spicules. The
illustrations for this paper were prepared from photomicrographs and camera
lucida drawings, using a Zeiss-Winkel microscope and its accessories.

REVISED MORPHOLOGICAL EVALUATIONS
Palaeospongilla chubutensis Ott and Volkheimer, 1972, pp. 49-63.

Holotype. Three sections and one polished rest-cut of the branch holding
the sponge incrustation, in the collections of the Museo Argentino de Ciencias
Naturales ‘‘ B. Rivadavia ’’, Buenos Aires.

Topotype (selected by the present authors). Three sections from another
part of the fossil, of which one was fully dissolved in nitrie acid, resulting in
several spicule slides; material deposited in part in the collections of the
Australian Museum, Sydney, and in part in those of the Smithsonian Institution,
U.S. National Museum, Washington, D.C.

Type locality. Cretaceous lacustrine sediments of the Chubut Group,
Valley of the Rio Chubut, 16 km NNW of Cerro Condor, Patagonia, Argentina.

Distribution. Hitherto known only from the type locality.

Redescription. Sponge encrusting the fossilised remains of what appears to
be a gymnosperm stem or root, in cushions from 2 to 6 mm in thickness, and
surrounded by a rather thick layer of covering calcareous algae (Cyanophyceae).
Skeleton forming several clearly defined tiers of megascleres, displaying a well-
arranged and rather regular meshwork of vertical and horizontal spicule fibres,
indicating the original presence of a considerable amount of binding spongin.
Spicule fibres consisting of joined bundles of up to 40 megascleres, forming
meshes of about 1000 um in diameter. Original surface of sponge difficult to
assess since the surrounding thick crust of calcareous algae adjoins very closely.

Megascleres of two distinct types, both feebly curved or almost straight.
The much more common type A represented by rather stout and almost cylindrical
amphioxea, as a rule completely smooth, occasionally bearing one or two irregular
smaller spines without characteristic arrangement or position (Fig. la). Length
range 420-540 um, width range 10-22 um. Megascleres of type B less prevalent
in the pre parations made, smaller and more slender , typically fusiform with rather
sharp apices, armed with inconspicuous spines except at their tips (Fig. 1b).
Length range 180-308 um, width range 4-14 ym. Megascleres of type A forming
the main skeleton of basal and central parts of the sponge, those of type B
associated more with peripheral spicule fibres.

Free microscleres (Fleischnadeln), like the megascleres, of two distinct
types, both almost straight or only very slightly curved amphioxea with distinctly
sharp apices. The much more common type A represented by rather long and

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A. A. RACEK AND F. W. HARRISON 159

slender scleres, incipiently to conspicuously spined along almost their entire
length (Fig. 1c). Length range 179-190 wm (partly assessed from fragments),
width range 3-4 um, spines rarely exceeding 1um in height. Microscleres of
type B somewhat smaller and entirely smooth amphioxea (Fig. 1d). Length
range 77-147 um (partly assessed from fragments), width range 3-5 um. Both
types seem not to be indicative of a certain position in the sponge, although they
appear more abundant above the layer of gemmules.

00pm

—=

Fig. 1. Palaeospongilla chubutensis. a. Range of megascleres of type A. b. Rangef of
megascleres of type B. c. Range of microscleres of type A. d. Range of microscleres of type B.

Gemmoscleres moderately long and slender amphistrongyla, often possessing
a Spine in the prolongation of their axis so that they appear amphioxeous. They
are armed with numerous acute spines along their entire shaft, while in the
vicinity of the extremities of the sclere these spines are distinctly aggregated,
often forming sceptre-like distal arrangements of perceptibly recurved teeth
(Fig. 2a). Pseudorotules could not be observed. Length range of gemmoscleres,
which are only slightly curved, 77-168 um, width range 3-5-6 um.

Gemmules quite abundantly produced, not confined to base of sponge but
instead freely scattered through skeletal meshwork. Although they are generally
spherical, ranging in diameter 490-570 um, their thick and well-developed
pneumatic coat is of irregular height, varying from 63 to 95 um (Fig. 2b).
Gemmoscleres embedded in this coat more or less radially but mostly crossing
each other at various angles, arranged in one layer only, their distal extremities
not appreciably penetrating the outer gemmular membrane. None of the
sections available displayed a cut through the region of the micropyle; thus
foraminal structures remain unknown.

Discussion. Ott and Volkheimer (1972), the discoverers of this first
spongillid known from the Mesozoicum, provided a very useful general description
and carefully assessed measurements of the variety of spicular components.
However, some of their morphological evaluations must now be amended. Since

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160 POSITION OF PALAEOSPONGILLA CHUBUTENSIS

they had no access to recent taxonomic revisions (Penney and Racek, 1968 ;
Racek, 1969, 1970), which include data from subfossil spicular remains, and
based their description on the study of thin sections only, it is understandable
that they faced difficulties in expressing the status of some scleral components.
While the megascleres of type A, including those which possess a small number
of irregular spines, were correctly observed, the spined type B were not recognised
as megascleres, but were instead included in the range of occurring free
microscleres, thus indicating a greater average length of the latter. The
assessment of all scleres from sections, the matrix of which grossly distorts the

Ss ead senile an NS er tenance ES UNS es EAS Ta

100m

Fig. 2. Palaeospongilla chubutensis. a. Range of gemmoscleres. b. Section through
fossilised gemmule, with gemmoscleres in situ.

characteristics of the spicules, seems to have resulted in somewhat misleading
line drawings, which show little resemblance to those scleres illustrated in the
excellent photomicrographs of the original description. Following the isolation,
by the present authors, of spicular components from the matrix, it can now be
established that there are two types of both megascleres and microscleres, a
condition which has not been retained in any extant genus of the now redundant
subfamily Spongillinae Vejdovsky, i.e. those spongillids possessing acerate (non-
birotulate) gemmoscleres. The possibility of a spicular mixture of two distinct
species, both fossilised simultaneously, can be readily dismissed. Although such
a condition can occasionally be observed in living spongillids (usually a result of
competition for a restricted substrate), there is always evidence of growth overlap,
i.e. two distinctly separated skeletal floors, each displaying the spicular
characteristics of one of the adjoining or overgrowing spongillids involved. The
careful examination of the skeletal arrangement of P. chubutensis did not reveal
such a condition and demonstrated beyond doubt that the spicular components
of the fossil belong to a single species. Both types of megascleres take part in

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A, A. RACEK AND F. W. HARRISON 161

the construction of a rather uniform skeleton, uninterrupted by membranes,
tiers or floors. The free microscleres of both types occur side by side throughout
the quite insignificant height of the sponge crust, without any indicative position.
The typical gemmules are not restricted to either the upper or lower portion of
the flat sponge growth but are scattered throughout the skeletal meshwork.
No other types of gemmules nor even single gemmoscleres of any other species
could be observed in the material available.

Considering all the morphological characteristics of P. chubutensis, as
reexamined by the present authors, and in spite of the close affinities of both
gemmules and gemmoscleres of the fossil sponge with those occurring in a number
of species of the recent genus Radiospongilla Penney and Racek (Fig. 3a-g), the
choice of a new genus for this Mesozoic spongillid by the describing authors is
fully supported by the present work. In the following section, the fossil sponge’s
systematic and phylogenetic position will be discussed in the light of recent
taxonomic, ecological and geological studies.

100ym

> 7 ‘ < cS
4m aS

Fig. 3. Morphological features of extant Radiospongilla spp. gemmoscleres. a. R. cerebellata.
b. R. philippinensis. c. R. hemephydatia. d. R. sceptroides. e. R. crateriformis. f. R. canton-
ensis. g. Optical section through gemmule of R. cerebellata (left half of gemmoscleres omitted).

SYSTEMATIC, PHYLOGENETIC AND ECOLOGICAL EVALUATIONS

Penney and Racek (1968), by introducing a new genus, Radiospongilla, as a
natural morphological link between the spongillids with acerate and those with
birotulate gemmoscleres, demonstrated the fallacy of the previously widely
applied division of the family Spongillidae into the meaningless subfamilies
Spongillinae and Meyeninae. Although lacking fossil evidence, Penney and
Racek suggested that the two different types of gemmoscleres present in these
artificial subfamilies had resulted from divergence from shapes originally present
in an hypothetical stock, from which the extant genus Radiospongilla could also

PROCEEDINGS OF THE LINNEAN Society or NEw SourH WatzgEsS, Vou. 99, Part 3

162 POSITION OF PALAEOSPONGILLA CHUBUTENSIS

have evolved. While most species of Radiospongilla possess distinctly acerate
gemmoscleres, whether of amphioxeous or amphistrongylous shape, they are
characteristic in having terminal aggregations of gemmoscleral spines, often
forming pseudorotules. One of these species, i.e. R. cantonensis (Gee) (Fig. 3f),
produces almost perfect birotulate gemmoscleres which are straight and therefore
strictly radially arranged, thus documenting the likelihood of a gradual
evolutionary divergence of both acerate and birotulate spicules from scleres
present in an ancestor, from which both these types could have been derived.
The antiquity of the extant genus Radiospongilla has now been established by
palaeolimnological examinations of Pliocene and Pleistocene sediments (Racek,
1969) and there is no doubt that this genus occupies a central phylogenetic
position. However, what could have been more gratifying than to obtain
palaeontological evidence of the existence of an even earlier spongillid, which
clearly displays ancient traits from which both acerate and _ birotulate
gemmoscleres could have arisen ?

As can be expected from a fossil of about 100 million years of age,
P. chubutensis differs in several ways from any presently living genus or species.
It thus deserves a taxonomic status of its own, at generic as well as specific level,
although lying close to Radiospongilla. At the same time, the Mesozoic fossil
displays spicular and constructional criteria which are now found separately in
species of three extant genera, i.e. Radiospongilla, Spongilla, and Pectispongilla.
With Radiospongilla the fossil shares almost unchanged gemmular and
gemmoscleral characteristics and at least one type of megascleres, i.e. those
which are incipiently spined, but differs by the presence of free microscleres,
which are typically absent in Radiospongilla. Spongilla, too, seems to have
inherited some of the characteristics of P. chubutensis. These are in particular
the flesh spicules, of which only the spined type is retained in Spongilla, and also
the cylindrical shape of the smooth megascleres. Even though Spongilla must
be considered more distantly related to the fossil than Radiospongilla, especially
with respect to gemmular and gemmoscleral structure and shape, its evolution
from this ancestral type is most likely. Pectispongilla, an extant genus closely
related to Radiospongilla, usually possesses two types of flesh spicules, and thus
shares an apparently important morphological criterion with P. chubutensis.
Pectispongilla, in addition, displays a similar skeletal construction and rather
insignificant depths of encrustations. However, although the gemmoscleres are
inserted in a manner similar to that both in P. chubutensis, and in almost all
Radiospongilla spp., the gemmules of Pectispongilla are much smaller, lack the
thick pneumatic coat, and their gemmoscleres display a unique and specialised
arrangement of mostly unilateral spines.

As with all fossil evidence, the phylogenetic relationship of the Cretaceous
spongillid to living forms can only be assumed until earlier ancestors are found.
However, the above comparison of combined characteristics displayed by P.
chubutensis with those found separately in the three extant genera makes it
more feasible to speculate about the possible pathway of spongillid evolution
during the past 100 million years, both chronologically and spatially. The fossil
clearly demonstrates the early existence of perfected gemmule-producing sponges
in inland waters, long before the isolation of non-spongillid Porifera of rather
marine facies in the so-called ** ancient freshwater lakes ”’. Recent comprehensive
studies by the present authors of a great number of gemmule-less Porifera from a
range of thalassoid environments (Racek; Racek and Harrison, unpublished
data) show conclusively that they must have arisen not only from different
ancestors but also in much more recent geological times. The polyphyletic
origin of the ‘‘ freshwater sponges ’’, already assumed by Marshall (1883),
commented on by Penney and Racek (1968) and Racek (1969), and discussed by
Brien (19664, 1966b, 1969, 1973), has thus become well documented.

PROCEEDINGS OF THE LINNEAN Socrety or New SoutH WAtzEs, Vou. 99, Part 3

A. A. RACEK AND F. W. HARRISON 163

Until recently, morphological comparisons between fossilised presumed
ancestral stocks and extant related organisms were the only means of assessing
evolutionary pathways. However, recent continental drift reeonstructions from
plate tectonics (Oxburgh, 1971 ; Smith, 1971; Vine, 1971) can now be used to
demonstrate feasible pathways of dispersal and speciation leading from the
Mesozoic fossil of Patagonia to related extant groups of spongillids. Since
P. chubutensis can be recognised as the almost direct ancestral form of the genus
Radiospongilla in general, and of the &. cerebellata group in particular, the present
distribution of these essentially southern sponges suggests an evolutionary
pathway following a west-east course along the Mesozoic continuity of the
combined landmass of parts of South America, Antarctica and Australasia as
one route, and along southern Africa and the then connected Indo-Pakistani
subeontinent as another. That this is not an isolated case can best be
demonstrated by the present range of two different extant spongillids, i.e.
Spongilla alba and the Hphydatia ramsayi complex (Penney and Racek, 1968).
While both these extant spongillids display a markedly uniform morphology
over their entire range from South America through Australia to at least south-
eastern Asia, S. alba shows drastic speciation trends from South America
northward (S. cenota in central America, and S. wagneri in the southeastern
United States), whereas some distant relatives of the H. ramsayi complex (i.e. E.
robusta and H. subdivisa) are not represented south of the United States. Since
it has been established (Fooden, 1972) that Australia must have separated from
the Antarctic plate some 45-50 million years ago, it is obvious that both S. alba
and the H. ramsayi complex must have existed long before the final dismember-
ment of ‘“‘Gondwanaland’”’. On the other hand, clear speciation trends, and
even discontinuity, seem to have occurred along the distributional pathway from
South America northward, indicating the earlier existence of major distributional
barriers, and ensuing isolation of sponge populations. Another indication of
such a development is the absence of the R&R. cerebellata group of sponges, one of
the closest descendants of P. chubutensis, north of Brasil. In spite of recent
detailed studies by specialists in South America, none of the species of the above
group have hitherto been recorded from that area, although some more remotely
related species are known to range discontinuously from central Argentina
Bonetto and Ezcurra, 1964) through Mexico to the United States (Penney and
Racek, 1968).

While the evolutionary pathway leading from the Mesozoic fossil to those
few most closely related species of Radiospongilla now seems well documented,
the rise of the extant genus Spongilla is more difficult to assume. It consists
today of two groups of species, both markedly different in morphological
characteristics, ecological preferences and, last but not least, zoogeographic
distribution. The S. alba complex, already discussed above, must be considered
another close relative of P. chubutensis, and has a distinct southern distribution
and a pronounced preference for alkaline habitats. The geologically more
recent S. lacustris group appears to be another evolutionary line. It is presently
restricted to the Northern Hemisphere, and occurs in acidic to only slightly
alkaline lentic and lotic habitats. Although the general morphological criteria
within Spongilla, as restricted by Penney and Racek (1968), are fairly constant,
and certainly generically characteristic, the possibility that these two groups
arose separately and at different geological periods cannot be dismissed. Until
further fossil spongillids are found, particularly in northern landmasses of the
past, any definite conclusions as to the ancestry of the extant genus Spongilla
S.s. must remain strictly speculative.

In dealing with presumed pathways of evolution and speciation, one naturally
has to consider possible dispersal routes. In the family Spongillidae, these do

PROCEEDINGS OF THE LINNEAN SocieTY OF New SoutH WALEs, Vou. 99, Part 3

164 POSITION OF PALAEOSPONGILLA CHUBUTENSIS

not necessarily require fully connected landmasses, since their gemmules are
well known to be distributed by aquatic birds. This often occurs over
considerable distances, including extensive stretches of oceans, with occasional
dispersal by wind from dried water beds. While such an indirect dispersal could
possibly have accounted for the advance of certain spongillid groups from
Australia to nearby Asia, or vice versa, the unlikely presence either of true birds
or of any other animals capable of intercontinental flight at the time of the
occurrence of P. chubutensis would seem to rule out such a possibility. Con-
sidering all the factors discussed, therefore, the evolutionary advance from the
Patagonian fossil to at least those species of Radiospongilla which have retained
identical gemmular and gemmoscleral structures and characteristics, and which
today abound in Australia, Africa and S8.E. Asia, can only be suggested to have
taken a west-east course prior to the dismemberment of ‘‘ Gondwanaland ”’,
which would have permanently disrupted such an advance. In the light of this
new evidence, the previously held belief that some of the Australian spongillid
fauna is a mere southward extension of that of Asia (Racek, 1969), seems severely
affected. It is now most feasible to believe that such an advance to both these
continents occurred independently, and that any later exchange of certain
species from one to the other has been a mutual one.

With respect to ecological evaluations of the probable physical, chemical
and climatic conditions present during the occurrence of P. chubutensis in the
Cretaceous, those characteristic for related Radiospongilla spp. may be used for
comparison. Both the R. cerebellata group of species, and to a certain extent
also R. hemephydatia, are essentially tropical species. Their conspicuously thick
pheumatic coat with its gemmoscleral armature, identical with that found in
P. chubutensis, is indicative of hot or even arid climates. The two Radiospongilla
spp. compared above prefer lentic habitats or still backwaters of the lotic series.
They furthermore appear to prefer distinctly alkaline conditions in their optimal
habitat and show little tolerance of sharp falls in hydrogen-ion concentrations.
The thickness of the gemmular pneumatic coat and its dense gemmoscleral
armature point to a warm to very warm, and perhaps even arid, climate during
the life of the Patagonian fossil in the Cretaceous. Its habitat undoubtedly
belonged to the lentic series, as already suggested by Ott and Volkheimer (1972).
And the fact that its natural environment must have been strongly alkaline is
demonstrated by the abundant, and apparently rapid, growth of the surrounding
calcareous algae, as well as by the actual mode of fossilisation.

ZUSAMMENFASSUNG

Der Fund des ersten fossilen Spongilliden, Palaeospongilla chubutensis Ott
and Volkheimer (1972), aus der Kreide Patagoniens offenbarte interessante
morphologische Strukturen, die griindliche systematische und phylogenetische
Studien erforderten. Dieses guterhaltene Fossil aus lakustrinen Sedimenten,
etwa 100 Millionen Jahre alt, besteht aus zusammenhangenden Skelettfasern von
Megaskleren und freien Fleischnadeln, beide vertreten mit zwei verschiedenen
Typen. Es enthalt ausserdem eine grosse Anzahl von Gemmulae mit deren
Belagsnadeln, in situ. Die vorliegende Neuuntersuchung dieses mesozoischen
Spongilliden warf neues Licht auf die Annahme einer frihen Abzweigung der
Gemmulae-erzeugenden Siisswasserschwaémme von deren ‘‘ marinen ’’ Vorfahren.
Diese Arbeit beweist, dass P. chubutensis als Ahnform dreier rezenten Gattungen,
d.i. Radiospongilla Penney and Racek, Spongilla Lamarck, und Pectispongilla
Annandale anzusehen ist, da sie kombinierte Nadelformen besitzt, die in den
oben genannten Gattungen separat vorkommen, obwohl das Fossil der Gattung
Radiospongilla am nachsten steht. Die Ergebnisse dieser Untersuchung sind

von besonderer phylogenetischer Bedeutung, da sie auf die Abzweigung der
friiher anerkannten Unterfamilien Spongillinae Vejdovsky und Meyeninae

PROCEEDINGS OF THE LINNEAN SooreTY oF NEw SourH WALES, Vou. 99, Part 3

A. A. RACEK AND F. W. HARRISON 165

Vejdovsky von einem zentralen Ahnenstamm hinweisen, der heute durch die
spezialisierten Arten der Gattung Radiospongilla reprasentiert wird.

Gegenwartige Fortschritte in den Gebieten der Palaeolimnologie,
Plattentektonik, und Kontinentverschiebung werden dazu verwertet, um genauere
Unterlagen fiir die Annahme phylogenetischer Verwandschaftsverhaltnisse zu
erhalten, und auch die heutige zoogeographische Verbreitung rezenter Arten zu
erklaren. Es wird ausserdem versucht, die physikalischen, chemischen, und
klimatischen Verhaltnisse wahrend des Vorkommens des mesozoischen Fossils
zu erklaren.

ACKNOWLEDGEMENTS

We are greatly indebted to Dr. Ernst Ott, Institut fiir Geologie, Technische
Universitat, Munchen, for his kind cooperation in this project, and for his
donation of type material of P. chubutensis.

We also wish to thank Professor D. T. Anderson and Associate Professor
J. R. Simons, School of Biological Sciences, University of Sydney, for valuable
criticism, and Mrs. T’. Manalang for her care in the preparation of the typescript.

Financial support from the University of Sydney’s Research Committee
(in provision of a Research Grant), and from the Presbyterian College, Clinton,
South Carolina (to F.W.H., while on sabbatical leave in Australia) is gratefully
acknowledged.
References
Bonetto, A. A., and Ezcurra, I. D., 1964.—Nuevas esponjas de agua dulce de la Republica
Argentina. Physis, 24 : 329-336.
Brien, P., 1966a.—Le polyphylétisme des éponges d’eau douce. L’embryogénése et la larve
chez Potamolepis stendelli (Jaffé). C.R. Acad. Sc. Paris 263 ; sér. D : 649-652.

, 1966b.—Le polyphylétisme des éponges d’eau douce. Formation de statoblastes chez
Potamolepis stendelli (Jaffé). C.R. Acad. Sc. Paris, 263 ; sér. D : 725-728.

, 1969.—Nouvelles éponges du Lac Moero. Res. Sci. Hxpl. Hydrob. Bassin Lac Bangweolo
et Luapula, 11 (2): 1-39.

, 1973.—Malawispongia echinoides Brien, études complémentaires, histologie, sexualité,
embryologie, affinités systematiques. Rev. Zool. Bot. Afr., 87: 50—76.

Foopen, J., 1972.—Breakup of Pangaea and isolation of relict mammals in Australia, South
America and Madagascar. Science, 175 : 894-898.

MarsHaty, W., 1883.—On some new siliceous sponges collected by Mr. Pechiiel-Loesche om the
Congo. Ann. Mag. Nat. Hist., 12 : 391-412.

Ort, E., and VoLKHEIMER, W., 1972 2.—Palaeospongilla chubutensis n.g. et n. sp.—ein Sie
erschwamm aus der Kreide Patagoniens. N. Jb. Geol. Paldont. ADbh., 140 : 49-63.

OxspureH, E. R., 1971.—Plate Tectonics. In Gass, I. G., Smith, P. as and Wilson, R. C. L.,
Understanding the earth. Horsham, Sussex : Artemis Press : 263-285.

Penney, J. T., and Racrex, A. A., 1968.—Comprehensive revision of a worldwide collection of
freshwater sponges (Porifera : Spongillidae). Bull. Smiths. Inst. U.S. Nat. Mus., 272 : 1-184.

Racexk, A. A., 1966.—Spicular remains of freshwater sponges. Mem. Connecticut Acad. Arts Scv.,
17 : 78-83.

, 1969.—The freshwater sponges of Australia (Porifera : Spongillidae). Austr. J. Mar.
Freshw. Res., 20 : 267—310.

, 1970.—The Porifera. In Hutchinson G. E. et al. ‘‘ Ianula : An account of the history
and development of the Lago di Monterosi, Latium, Italy”. Trans. Amer. Phil. Soc.
Philadelphia, n.s., 60 (4) : 148-149.

Revut, K., 1954.—Spongilliden-Erde im Miocén des Fichtelgebirges. Ein Vergleich mit der
miocanen Kieselgur von Beuern. Geol. Bl. NO-Bayern, 4: 14-20.

Smiru, A. G., 1971.—Continental Drift. In Gass, I. G., Smith, P. J., and Wilson, R. C. L.,
Understanding the earth. Horsham, Sussex : Artemis Press : 213-231.

TraxterR, L., 1895.—Spikule von Siisswasserschwammen aus Brasilien. J dldtani Kdzlony, 25:
238-242.

, 1896.—Subfossile Siisswasserschwamme aus Australien. Féldtani Kdézlény, 26 : 95-97.

Vine, F. J., 1971.—Sea floor spreading. In Gass, I. G., Smith, P. J., and Wilson, R. C. L.,

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Reprints

Please address all requests for reprints to: Dr. F. W. Harrison, Department of Anatomy,
Albany Medical College, Albany, N.Y. 12208, U.S.A.

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutTH WALES, VoL. 99, Part 3

MUTAGENIC EFFECTS OF ETHYL METHANESULPHONATE ON THE
OAT STEM RUST PATHOGEN (PUCCINIA GRAMINIS F. SP. AVEN AE)

C. TEo* AND EK. P. BAKER*

(Plate vm)
[Accepted for publication 17th April 1974]

Synopsis

Ethyl methanesulphonate (EMS) induced a high degree of stable variability in the dikaryotic
uredial stage of the oat stem rust fungus (Puccinia graminis f. sp. avenae). Traits affected were
uredospore colour, interval between uredial stage and subsequent telial development, virulence
and pathogenicity. Mutations for virulence were induced on the oat cultivar Saia. Increases
in mutagen concentration, duration of treatment and temperature during treatment significantly
increased mutation rates. Recurrent mutagenic treatment increased mutation frequency and in
addition produced a wider spectrum of mutants. Potential uses of mutants induced by EMS
are indicated.

INTRODUCTION

Ethyl methanesulphonate (EMS) causes mutations in the dikaryotie uredial
stage of the oat stem rust fungus, Puccinia graminis f. sp. avenae Eriks. and
Henn. (Baker and Teo, 1966). In their studies aberrant uredospore colour and
rapid telial-forming variants were described. The present studies provide
estimates of mutation rates for these traits, and describe the effects of mutagen
concentration, duration of treatment and temperature during treatment on
mutation rates. In addition, alterations in virulence and changes in certain
pathogenic traits were demonstrated.

MATERIALS AND METHODS

The strains of oat rust used were 2-0, 7-H and 8-H, the cultures of which
are designated as BC1, BC2 and BC3, respectively, in the University of Sydney
Rust Accession Register. Infection types produced on the oat differential
varieties when inoculated with these cultures are shown in Table 1.

TABLE 1

Infection types exhibited on seedlings of various oat cultivars (resistance gene(s) in brackets) by three
cultures of oat stem rust at 17 +.2°C

Rust culture
Cultivar —_———
BCl BC2 BC3
(a) Canadian differentials
Richland, *08(A) .. is a se ae Bt 1+ 1 4
Rodney, 0654(B) .. 2 ae Se we ui 2— J— 1
White Tartar, 05 (D) 22 Hi a dif 2 2= 2— 4 2—
Jostrain, 0617(#) .. eis A ate ap a 3+ 4 4
Eagle? x C.1.4023, 0658 (F) oe ae ae .. 2= 2— 2 2—
C.1.5844-1, 0661 (H) See mK; ci EN .. 2= 2—3e 3 3
(b) Supplementary differentials
Minnesota Ag.331, 0615 (A D) we He AN .. I+ 1+ 2—
Saia, 0589 (Sa) us Age An ae he oh eh ein Bll Roll
Strain designation ac si aus i oh .. 2f-OF 7-H 8-H.
Virulence formula (Green, 1963) A ce se .. ABDFH/E§ ABF/DEH BDF/AEH

* ( numbers refer to Sydney University Oat Accession Register.

+ Race designation according to Newton and Johnson (1944).

t The symbol 0 indicates avirulence on genes A, B, D, F and H. The symbol H. indicates
virulence on gene H.
* Department of Agricultural Botany, University of Sydney, N.S.W., 2006.

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEW SourH WALES, VOL. 99, Part 3

C. TEO AND E. P. BAKER 167

Seedling infection types, recorded 14-16 days after inoculation, were
designated in accordance with the scale suggested by Stakman et al. (1923).

The oat cultivars employed were Bond and Saia. Bond is a susceptible
hexaploid cultivar. Saia (an accession of Avena strigosa Schreb., a diploid
species) is resistant to most oat stem rust strains in Australia. However, certain
field isolates collected in eastern Australia (Luig and Baker, 1973) have proved
virulent on this cultivar.

The current experiments were conducted to study, in part, the effect of
mutagen concentration, duration of treatment, and temperature during treatment,
on mutation rate for certain rust characters. In this regard culture BC2 was
treated as follows :

1. With 9-8x10-7, 14-7x10- and 19-6x10-#M aqueous solutions of
EMS for 2-0 hrs at 25°C. Distilled water was used in the preparation
of the solutions. (M.W. of EMS is 124-16 and its density at 20°C was
determined to be 1-2175).

2. For 2-0, 3-5 and 5-4 hrs with a 9-8 x10-2M solution at 20°C in separate
experiments.

3. At 20, 25 and 30°C for 2-0 hrs with a 9-8 x 10-2M solution.

Cultures BC1 and BC3 were also treated with EMS, each with a 14-7 x 10-?M
solution for 2-0 hrs at 20°C. Finally, the effect of treating an EMS-induced
grey-brown uredospore colour mutant of culture BC2 with 12-5x10-?M EMS
solution for 2-0 hrs at 22°C was studied.

In recurrent mutagenic treatments cultures BC1 and BC2 were used. The
former was treated for four and the latter for five successive uredial generations
with 14-7 x10-*M EMS for 2-0 hrs at room temperature. Cultures BC1 and
BC2 were also treated for two and three consecutive uredial generations, respec-
tively, with 15-7 x10-?M EMS for 2-0 hrs at room temperatures in attempts to
induce mutations for virulence on the cultivar Saia.

Uredospores were treated with solutions of EMS (100 mls per 0-2g of spores)
in a stoppered flask shaken with a Griffin flask shaker. After treatment, the
Spores were rapidly filtered in a Buchner filtering apparatus. After washing,
filtering was continued to remove as much water as possible from the spore mass.
A sample was retained for assessment of spore viability. The remainder was
dispersed in ‘‘ Mobil Sol 100” (Rowell and Olien, 1957) and the suspension
immediately sprayed with a ‘‘ Desaga Spray Can ”’ atomiser onto primary leaves
of 1,500—2,000 seedlings of Bond. However, in experiments designed to induce
mutations for virulence, 1,500—2,000 seedlings of Saia and 300 seedlings of Bond
(to provide inoculum for the next recurrent mutagenic treatment) were inoculated
simultaneously. A control experiment set up in each case comprised 500-600
seedlings of the appropriate cultivar inoculated with untreated spore samples.
Inoculated plants were placed in misting chambers for 12—24 hrs and then trans-
ferred to well-lit glasshouse benches.

Assessment of Spore Germination

Samples of mutagen-treated and untreated spores were spread lightly and
evenly on 2% water-agar in three petri dishes and placed in an incubator at
18—20°C for 24 hours. After this period, germination of at least 500 spores per
dish was examined. The percentage spore germination was determined after
totalling the results for each dish.

Spores were classified as inviable if no germination was shown or if initial
germination occurred but germ tubes failed to continue growth. In the latter
the germ tube barely appeared through a pore in the wall after 24 hours whereas

germ tubes from viable spores showed extensive hyphal development after the
same period.

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutH WALES, VOL. 99, Part 3

168 ETHYL METHANESULPHONATE ON OAT STEM RUST FUNGUS

Assessment of Mutation Rate

All leaves were examined and the number of mutants recorded. The total
number of pustules appearing on all leaves was then counted. In certain
experiments, where indicated, the total number of pustules was assessed by
sampling. In these instances, the number of pustules on seedlings in ten randomly
selected pots (approximately 35 seedlings per pot) was counted and the
approximate total number of pustules in the experiment estimated on the basis
of such sampling. This method of estimation was less reliable as the rate of
infection varied somewhat from pot to pot. However, slight inaccuracies in the
estimation of the total number of pustules by sampling were shown to have no
significant effect on statistical assessment of mutation rate.

EXPERIMENTAL RESULTS
Induction of Mutants

During the course of various experiments spore viability following EMS
treatments under similar conditions was found to vary markedly depending on
the source and batch of the chemical. For example, spore viability ranged
from 0-96°, of controls (which showed 85—92°, viability) after treatment with
9-8 <10-2M solutions for 2-0 hrs at 20°C in different experiments. Apparently
EMS may contain impurities, such as mercapto ethanol, which are fungicidal, thus
presumably accounting for reduced spore viability, the variation in which may
have been due to relative amounts of impurities.

It was not possible to obtain EMS free from impurities on all occasions.
Consequently, the use of EMS from sources which did not result in undue lethality
offered the only practical solution to this problem. In experiments where strict
comparisons were necessary, as in studies concerned with the effect of treatment
variables, EMS relatively free from impurities and from the same bottle was
used in all treatments.

Within the range of reduction in spore viability tolerated there was no
evidence that impurities affected either the mutation rate or the type of mutants.

The influence of treatment conditions on spore viability and mutation rate
in culture BC2 is recorded in Table 2. Spore viability decreased with increases
in the mutagen concentration, duration of treatment and temperature during
treatment. Conversely, the total mutation rate (which was based on both
aberrant colour mutants and rapid telial-forming variants) significantly increased
with increases in mutagen concentration, duration of treatment and temperature
during treatment. A similar result was obtained when the rate of colour mutants
alone was considered. In the case of rapid telial-forming variants increases in
rate were significant only in instances where the 95% limits of expectation
(calculated by the method of Stevens (1972)) did not overlap.

Two alternatives for determining mutation rates can be considered. The
mutation rate can be estimated in terms of the total number of spores treated or
in relation to the total number of spores which germinated and produced pustules.
For practical considerations the latter estimate was used in these studies.

As with culture BC2, yellow and orange colour mutants and rapid telial-
forming variants were readily induced in cultures BC1 and BC3 (Table 3).
Mutation rates for these traits in the three strains did not differ significantly. No
mutants were observed in the control experiments.

The results of recurrent mutagenic treatments of cultures BC1 and BC2 are
shown in Table 4. With each successive cycle statistically significant increases
in total mutation rate occurred. Moreover, a wider range of aberrant uredospore
colour types was obtained. Grey-brown colour mutants, which were not obtained
following single mutagenic treatment except on one occasion at the highest

PROCEEDINGS OF THE LINNEAN Society or New Sourn Waxes, Vou. 99, Part 3

169

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PROCEEDINGS OF THE LINNEAN Socrety oF New SoutH Watzs, Vou. 99, Part 3

170 ETHYL METHANESULPHONATE ON OAT STEM RUST FUNGUS

Effect of recurrent mutagenic treatments on mutath

Rust Recurrent Spore Total

culture Treatment cycle viability number Yellow

number (% control) pustules and
orange

no.

20+1°C Initial 50-2 12,163 38

14-7x10-?M, 22+1°C 1 38-2 (11,353 90

BC1 2-0 hrs 22-+1°C 2 45:8 8,917 157

19+1°C 3 57-1 7,688 244

20+1°C Initial 47-5 15,050 63

BC2 14-7x10--M, 21+1°C 1 43-4 11,439 118

2-0 hrs 19+1°C 2 59-6 7,302 109

22+1°C 3 37-7 10,725 329

19+1°C 4 53-4 8,120 416

* O= Observed ; L=Lower limit ; U= Upper limit.

EMS concentration (Table 2), were readily induced following recurrent mutagenic
treatments. Mutants possessing a markedly darker spore colour than normal
were also found. Certain variants categorised in Table 4 as showing rapid telial
formation were also characterised by aberrant uredospore colour.

In all of the above, single or recurrent mutagenic treatments mutations for
pathogenicity were observed. In the current context, usage of the terms
‘‘ pathogenicity ’> and ‘‘ virulence ”’ follows that adopted by Watson (1970).
Pathogenicity is a general term and is used in connection with such characters
observed on the host as pustule size, volume of spores produced and the rate of
growth of the pathogen. If a strain is virulent on a host, the gene or genes for
resistance in the host are matched in the fungus by corresponding genes for
virulence.

Pathogenic mutants exhibited either small uredopustule size or delayed
pustule growth and development. Pathogenic mutants exhibiting subepidermal
uredial formation were also found following recurrent mutagenic treatments.
Certain aberrant colour mutants were characterised by one or more such patho-
genic traits.

No mutants virulent on Saia were induced following initial EMS treatments.
However, virulent mutants were induced in cultures BC] and BC2 after one and
two recurrent mutagenic treatments, respectively (Table 5).

TABLE 5
Effect of recurrent EMS treatments on mutation rates for virulence on Saia BC1 and BC2 of oat
stem rust
Pertinent Recurrent Total Virulence mutants (Vga)
Rust virulence cycle number
culture gene* Treatment number pustules Number Rate ( x 1074)
15-7x 10-2M, Initial 20,000F —
BCl Vsa 2 hrs, 21 ae 1°C
2 hrs, 21+1°C 1 25,600F 4 1-56
a 15-7 10-M, Initial 22,7004 =
2 hrs, 21+1°C
BC2 Vsa 2 hrs, 21-+1°C 1 23,400t —-
2 brs, 21+1°C 2 24,850 2 0-80

Vsa and vg, indicate avirulence and virulence on Saia, respectively.
* A single dominant gene was considered to govern resistance (see Discussion).
+ Estimated by sampling procedure.

PROCEEDINGS OF THE LINNEAN Society or New SourH WALsES, Vou. 99, Part 3

Uredospore col¥

Grey-
brown

Tota

TAr

C. TEO AND E. P. BAKER IRAE

m cultures BCL and BC2 of oat stem rust
ae EEE EE RIE ANU ANNE OE nee a LUA CREE EI, | PCL Aa ed SA) LS a STOEL = BAC

Rapid telial-forming variants

ants with uredial colour Total mutants
Normal Yellow Grey- Mutation rate and 95% limits
Tutation and brown Total Mutation No. (x 10-8)
rate orange rate
P< 1055) no. no. no. no. (x 10-4) O* L U
3°01 9 9 7:13 47 3-73 2-74 4-95
7:93 12 1 13 11-45 103 9-07 7:41 10-99
17-94 25 25 28-04 185 20-75 17-78 23:93
32-26 19 1 1 21 27°31 269 34-99 31-01 39-36
4-19 8 8 5:31 71 4-72 3-69 5-95
10-49 14 14 12-24 134 11-71 9-83 13-86
15-61 16 1 17 23-28 131 17-94 15-03 21:26
30-96 43 3 46 42-89 378 35-24 31-84 38-92
52-09 38 4 42 51-72 465 57-70 52-33 62-56

ES OE

Subjecting an EMS-induced grey-brown colour mutant to mutagenic
treatment yielded new variants. Of a total of approximately 20,000 pustules
(estimated by sampling procedures) examined, eight white uredospore mutants
were found.

The mutants, excepting those characterised by induced virulence on Saia,
will be described in a subsequent paper.

Mutants Virulent on Cultivar Saia

The infection types produced by two virulent mutants of culture BC2
compared with the original culture are shown in Plate vu.

The four mutants virulent on Saia induced from culture BC1 exhibited
identical infection types when tested on differential sets. The two mutants
virulent on Saia induced from culture BC2 also produced identical infection
types on differential sets. Characteristic infection types produced on differential
Sets by one representative mutant virulent on Saia induced from each of cultures
BC1 and BC2 are shown in Table 6.

TABLE 6

Infection types exhibited on seedlings of various oat cultivars (resistance gene(s) in brackets) at 17 +2°C
by mutants of oat stem rust with induced virulence

Mutant rust culture
Cultivar

BCl-a BC2-a

(a) Canadian differentials

—

Richland, 08(A) .. a oe Me He be as .. 11 I— 1+
Rodney, 0654(B) .. oh ae oe ee as ise sigma k ; 1—
White Tartar, 05 (D) Me ee BY fate =e ae i lf 2= 33+
Jostrain, 0617(H) .. te a she aie 23 ae ae), Be 33+
HKagle? x C.1.4023, 0658 (F) ue sls und ae ne .. 2= 2— 1+ 2
C.1.5844-1, 0661 (A) te if is i ee ah -. 1+23e¢ 33+
(6) Supplementary differentials
Minnesota Ag. 331,0615(A D) .. S16 i By oe sje: sieg, Lt l1i+
Saia, 0589 (Sa) we a ai Hs Aig Ae 3c 33+
Strain designation ui er “fs ag a ad = .. 2-0-1* 7-H-1

* Indicates virulence on Saia.

PROCEEDINGS OF THE LINNEAN Society or NEw SoutH WALES, VoL, 99, Part 3

172 ETHYL METHANESULPHONATE ON OAT STEM RUST FUNGUS

Except for virulence on Saia, mutants produced no changes in infection
types when tested on genotypes constituting the differential set. However,
mutants of culture BC1 gave “‘ 3c” infection types on the universally susceptible
cultivars Algerian, Bond, Fulghum, Victoria and Belar compared with ‘ 4”
infection types produced on these cultivars by the original culture. Likewise,
on these susceptible cultivars mutants of culture BC2 produced infection types
(‘°3 3+” with some chlorosis) similar to those exhibited on Saia. These were
slightly but distinctly lower than the ‘‘ 4” infection types produced on these
cultivars by the original cultures. Further, mutants showed 2-3 days’ longer
incubation and developmental periods and a general reduction in sporulation
vigour compared with the respective original strains. These observations
suggest that associated changes in pathogenicity were implicated in the production
of these mutants.

DISCUSSION

Ethyl methanesulphonate proved highly mutagenic on Puccinia graminis
avenae. Variability induced in the dikaryotic uredial stage involved traits
differing as widely as uredospore colour, interval between uredial stage and
subsequent telial development, virulence and pathogenicity.

Genetic rather than temporary phenotypic changes were involved in the
variability induced. Mutants remained stable over several generations of
subculturing. Contingent upon proof from inheritance studies, the changes
induced are believed to be due to recessive chromosomal gene mutations.

Mutants virulent on Saia differed from the original strains only by high
infection types produced on this cultivar. The number of genes conditioning
resistance in Saia is not definitely known. Murphy ef al. (1958) concluded that
the same two independent dominant genes in Avena strigosa accession C.D.3820
(which they considered agronomically identical with Saia) conditioned resistance
to each of four races. On the other hand, Dyck (1966) found that the same
single dominant gene in several accessions of A. strigosa governed resistance to
each of a wide range of races. If Dyck’s conclusions are accepted on the basis
of the gene-for-gene hypothesis (Flor, 1956) mutations for virulence on Saia
would involve a single locus and be recessive since avirulence is usually dominant.

Except for mutants virulent on Saia, no alterations in virulence were shown
by the variants examined. Although certain aberrant colour mutants exhibited
rapid telial formation or one or more pathogenic changes, absence of invariable
association between the various traits indicated that independent mutations were
involved. However, changes in incubation and developmental periods associated
with a change in spore colour may have been due to pleiotropic gene effects.

The variants induced were, therefore, intrinsically similar to those reported
to arise from spontaneous mutation or genetic recombination, although the rate
of, and range in variability induced was greater than that ascribed to spontaneous
mutation. This does not imply that such types would not occur naturally at a
low frequency. However, most abnormal variants would obviously have low
survival ability under natural conditions and would not be expected to persist.

Mutations for virulence were less readily induced than variation in other
traits. Mutations for uredospore colour, particularly those affecting loss of
spore wall pigment formation resulting in yellow or orange uredosori, occurred
at the highest frequency. Increases in mutagen concentration, duration of
treatment and temperature during treatment increased significantly mutation
rate for colour mutants and rapid telial-forming variants. Recurrent mutagenic
treatment also increased mutation frequency and, in addition, produced a wider
spectrum of mutants.

PROCEEDINGS OF THE LINNEAN SocieTY of New Sourn Waxzs, Vou. 99, Part 3

PLATE VII

Proc. LINN. Soc. N.S.W., Vol. 99, Part 3

i?)

Cc. THEO AND E. P. BAKER L733

Variation induced by EMS obviously has many applications, some of which
are listed as follows :

1. Elucidation of the extent, causes, and nature of spontaneous variability.

2. As cultures for assessing the role of somatic hybridisation in creating
variability in the oat stem rust fungus.

3. The rust reaction of segregating populations can be tested simultaneously
with two or more rust strains differing in virulence and uredospore colour.

4. Difficulty is often experienced in producing teleutospores from field
strains under glasshouse conditions. Teleutospores produced on mature
plants in the field usually originate from uredospores of unknown geno-
types. Rapid telial-forming variants induced from identified field strains
are obviously useful for determination of their genotypes and for studies
on the inheritance of virulence and other traits by infection of the barberry.
Inheritance studies can be greatly expedited since rapid telial-forming
variants provide abundant teleutospores after an extremely short
transitory uredial stage.

5. Analysis of uredospore pigments in differently coloured mutants may be
useful in elucidating the biosynthetic pathway of carotenogenesis in rust
fungi and in fungi in general.

References

Baker, E. P., and Txo, C., 1966.—Mutants of Puccinia graminis avenae induced by ethyl methane
sulphonate. Nature, 209 : 632-633.

Dyck, P. L., 1966.—Inheritance of stem rust resistance and other characteristics in diploid oats,
Avena strigosa. Can. J. Genet. Cytol., 8 : 444-450. :

Ftror, H. H., 1956.—The complementary genic systems in flax and flax rust. Advan. Genet., 8:
29-54.

GREEN, G. J., 1963.—Stem rust of oats in Canada in 1963. Can. Plant Disease Survey, 43 : 173-176.

Luie, N. H., and Baxer, E. P., 1973.—Variability in oat stem rust in eastern Australia. Proc.
Linn. Soc. N.S.W., 98 : 53-61.

Murpuy, H. C., Zituinsky, F. J., Srmons, M. D., and GRINDELAND, R., 1958.—Inheritance of
seed colour and resistance to races of stem and crown rust in Avena strigosa. Agron. J., 50:
539-541.

Newton, M., and Jounson, T., 1944.—Physiologic specialization of oat stem rust in Canada.
Can. J. Res., C, 22: 201-216.

Rowe tt, J. B., and Oren, C. R., 1957.—Controlled inoculation of wheat seedlings with uredospores
of Puccinia graminis tritici (Eriks. and Henn.) Guyot. Phytopathology, 47 : 650-655.

STAKMAN, E. C., Levine, M. N., and Battry, D. L., 1923.—Biologic forms of Puccinia graminis
on varieties of Avena spp. J. agr. Res., 24: 1013-1018.

STEVENS, W. L., 1942.—Accuracy of mutation rates. J. Genetics, 43 : 301-307.

Watson, I. A., 1970.—Changes in virulence and population shifts in plant pathogens. Ann. Rev.
Phytopathology, 8 : 209-230.

EXPLANATION OF PLATE VII

Infection types exhibited on Saia by original and two virulent mutants of strain 7-H of oat
stem rust (x3-5). a. Original strain 7—H : “‘; 1-” infection types. b-c. Mutant strains : ““33-+”’
infection types.

PROCEEDINGS OF THE LINNEAN SOCIETY oF NEw SoutH WALES, VoL. 99, Part 3

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PROCEEDINGS OF THE LINNEAN Society or New Sours WALES,
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Proceedings, Volume 99, Part 3, 1975

CONTENTS

DENMARK, H. A., and ScHIcHA, E. A new species of Amblyseius Berlese
(Acarina : Phytoseiidae) from apple in Australia

Doo.ey, J. K., and PAxton, J. R. A new species of tilefish ae
Taenioniocconitae) from eastern Australia oy, a :

RAcEK, A. A., and HARRISON, F. W. ‘The systematic and phylogenetic
position of Palaeospongilla chubutensis (Porifera : Spongillidae)

TEO, C, and BAKER, E. P. Mutagenic effects of ethyl methanesulphonate
on the oat stem rust pathogen (Puccinia graminis f. sp. avenae)

Notes and instructions for authors

Page

145

151

157

166

175

Proceedings of the
Linnean Society
of New South Wales i

Issued 23rd April, 1975

VOLUME 99
PART 4
No. 440

Registered for posting as a periodical—Category A.

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, 1974-75

President
T. G. Vallance, B.Sc., Ph.D.

Vice-Presidents

H. G. Cogger, M.Se., Ph.D.; L. A. S. Johnson, D.Sc. ;
P. J. Stanbury, Ph.D.; N. G. Stephenson, M.Sec., Ph.D.

Honorary Treasurer
Joyce W. Vickery, M.B.E., D.Sc., F.L.S.

Secretary
Mrs. Ruth J. Inall

Council
D. J. Anderson, Ph.D. A. Ritchie, B.Se., Ph.D.
Barbara G. Briggs, Ph.D. P. J. Stanbury, Ph.D.
H. G. Cogger, M.Se., Ph.D. N. G. Stephenson, M.Se., Ph.D.
D. W. Edwards, B.Sc.Agr. P. G. Valder, B.Sc.Agr., Ph.D.
L. A. 8. Johnson, D.Se. T. G. Vallance, B.Se., Ph.D.
D. McAlpine, M.Sc., Ph.D. Joyce W. Vickery, M.B.E., D.Sc., F.L.8.
Helene A. Martin, Ph.D. B. D. Webby, M.Se., Ph.D.
Lynette A. Moffat, B.Sc., Ph.D. G. P. Whitley, F.R.Z.S.
P. Myerscough, M.A., D.Phil. A.J. T. Wright, B.Se., Ph.D.

Councillors E’meriti

W. BR. Browne, D.Sc., F.A.A. A. B. Walkom, D.Sc.

Auditors

W. Sinclair and Company, Chartered Accountants

Iinnean Macleay Lecturer in Microbiology, University of Sydney
Ky Chosrh.D:

Librarian
Constance B. McKay, B.A., A.L.A.A.

Associate Editor
Janet Donald, B.A., Dip.Lib.

The Society’s Headquarters are in Science House, 157 Gloucester Street, Sydney,
N.S.W. 2000, Australia

A NEW SPECIES OF PHYTOSEIUS RIBAGA
(ACARINA: PHYTOSEIIDAE) FROM APPLE IN AUSTRALIA

H. A. DENMARK* AND HE. SCHICHAT

[Accepted for publication 19th June 1974]

Synopsis

Phytoseius fotheringhamiae n. sp. (Acarina: Phytoseiidae) is described from apple in New
South Wales, Australia. It is closely related to Phytoseius woodburyi DeLeon.

INTRODUCTION

The species of phytoseiid mite described in this paper was collected from
apple trees at Bathurst, New South Wales. It was found mainly on the under-
sides of leaves in orchards of young unbearing apple trees, in commercial orchards
of Granny Smith apple trees and on single neglected apple trees (‘ backyard
trees ’’) during the years 1971 to 1973.

Genus PHYTOSEIUS Ribaga
Phytoseius Ribaga, 1904, Riv. Patol. Veg., 10: 177

Phytoseius (Phytoseius) fotheringhamiae, n.sp.
(Figs 1-8)

Diagnosis. Phytoseius fotheringhamiae n. sp. is closely related to Phytoseius
woodburyt DeLeon from which it is distinguished by the following characteristics.
The lateral and the anterior sublateral setae are longer in the new species than
in P. woodburyi. The clunals are serrate in the new species but smooth in P.
woodburyi. A pair of large pores caudomedial to M, in the new species is missing
in P. woodburyi. The cervix of the spermatheca in the new species is much
longer than in P. woodburyt.

Measurements listed are the mean and standard deviation of 20 specimens
in microns, if not otherwise indicated.

* Division of Plant Industry, Florida Department of Agriculture and Consumer Services,
Gainesville, Florida, 32601, U.S.A. (Contribution No. 275.)

+ N.S.W. Department of Agriculture, Agricultural Research Station, Bathurst, New South
Wales, 2795.

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutH WALES, Vout. 99, Part 4

178 A NEW PHYSTOSEIID MITE

FEMALE

Dorsum: (Fig. 1) The dorsal shield, length 293-++6, width at L, 132-+3, is
rugose and bears 15 pairs of setae (inclusive S,), eight lateral, one median, three
dorsal, one pair each of clunals and verticals, and one pair of anterior sublateral
setae S, (on the dorsal shield). The setae measure in length: verticals 35 +2 ;
D;, D,, DD, and M,—4-5--0:5; clunals’ 38=-1; %,—52--4; 6-2
Lb, =30-2 5 Gb, 12-62 3 be tb 7s i, = 83-45 lb —82--5 ; 1 86--4
anterior sublaterals (S,) 50-2. Setae L,, L;, L., L,, Lz, S, and the verticals

Fig. 1. Dorsal shield of female. Fig. 2. Venter of female.

are serrate, L, and the clunals are slightly serrate. All other setae are smooth.
Setae 8,, L,, L;, L,, L, and L, are unusually thick. There is one pair of large
pores posterolaterally, one pair of large pores caudomedial to the first median
setae (M,), and one pore between the vertical and L,. Peritremes extend to the
vertical setae.

Venter: (Fig. 2) (Measurements of holotype only) Sternal shield, length 63,
width 91, smooth with three pairs of setae. Setae IV are on oval shaped meta-
sternal shields. Genital shield, length 94, width 101, normal with a pair of
genital setae. Ventrianal shield, length 119, width 64, elongate vase shaped
with three pairs of preanal setae.

Chelicerae: (Fig. 3) The fixed finger measures 36-+1 (ten measurements)
in length and is provided with three teeth and a pilus dentilis. The movable
finger, length 40+-1, has a single tooth.

Peritrematal shield: (Fig. 4).

PROCEEDINGS OF THE LINNEAN SocrETY oF New SourH WAtES, VoL. 99, Part 4

DENMARK AND SCHICHA 179

Length of macrosetae on leg IV as follows: (Fig. 5) Sge IV 17+2; Sti IV
a 2 2
4943; St IV, 2543; St IV, 20-2; Genn IL2, ~~,1; Genu ILI, a i
Spermatheca: (Fig. 6) (Two measur Coren) cane cup or bowl shaped cervix.
The major duct is 35-40 long. The atrium is 5 broad and 4-5 long. The cervix

has a diameter of 12-14 where it joins the vesicle.

MALE

Dorsum: (Measurements of three specimens) Length 260-274, width 120-
130, smaller than female; the chaetotaxy and the shape of the setae resemble
those of the female. The setae are, however, relatively shorter. Verticals =

0

Figs 3-8. 3. Chelicerae of female. 4. Peritrematal shield of female. 5. Leg IV of female,
6. Spermatheca of female. 7. Ventrianal shield of male. 8. Spermatodactyl of male.

29-33, D,, D, and D, all approximately 4; M,—5-6, clunals=6-7, L, 38-46,
fe 1-17 ls 95 90) L,=12-13, T7080) Tie 48"69) L,=51- _55, L,=43-49,
anterior sublaterals (S,)=35—43.

Venter: Ventrianal shield (Fig. 7) smooth with three pairs of preanal setae.

Spermatodactyl: (Fig. 8) Spermodactyl has a terminal heel and its lateral
process is distinct (no crest present on shank).

PROCEEDINGS OF THE LINNEAN SOCIETY oF NEw SoutH WALES, Vou. 99, Part 4

180 A NEW PHYTOSEIID MITE

Type. Female holotype from Bathurst, N.S.W., Australia, January, 1971,
E. Schicha, on leaves of sprayed Granny Smith apple trees. Deposited in South
Australian Museum, Adelaide, 8.A., Australia.

Paratypes. Eight females and four males taken with the holotype from the
same locality. Deposited as follows: four females and two males at Biological
and Chemical Research Institute, Rydalmere, N.S.W., Australia; two females
and one male at Division of Plant Industry, Florida Department of Agriculture
and Consumer Services, Gainesville, Florida, U.S.A.; two females and one
male at South Australian Museum, Adelaide.

Reference

DeLzon, D., 1965.—Phytoseiid mites from Puerto Rico with descriptions of new species (Acarina :
Mesostigmata). Fla Hnt., 48: 121-131.

PROCEEDINGS OF THE LINNEAN Society or New Souru WALzs, VOL. 99, Part 4

SILURIAN RUGOSE CORALS FROM THE MUMBIL AREA,
CENTRAL NEW SOUTH WALES

R. A. McLEAN*

(Plates vuI—x)

[Accepted for publication 19th June 1974]

Synopsis

Six species of rugose corals (three new) are described from strata of probable late Wenlock
to early Ludlow age in the Mumbil area, near Wellington in central New South Wales. They
comprise representatives of the genera Holmophyllum Wedekind, Holmophyllia Sytova, Coronoruga
Strusz, Pilophyllum Wedekind and Palaeophyllum Billings. Several of the forms described from
this area by Strusz (1961) are revised. The holotype of ‘‘ Baeophyllum’”’ colligatum Hill, 1940
is refigured and its affinities discussed.

INTRODUCTION

The first detailed study of the Palaeozoic strata of the Mumbil-Neurea-
Dripstone area, some 24 km south-east of Wellington, was that of Strusz (1960).
He defined a sequence consisting in the lower part of the Oakdale Formation of
Upper Ordovician age, overlain by the Mumbil Formation, regarded as being
Upper Llandovery to Ludlow in age. The Mumbil Formation was subdivided
into a lower, Narragal Limestone Member, and upper, Barnby Hills Shales
Member. Aspects of the coral fauna of this sequence were examined by Strusz
(1961). Subsequent study of this region by Vandyke (1970, unpubl.), particularly
in the region of the Oakdale Anticline (Text-fig. 1), has led to considerable
reappraisal of the stratigraphic sequence and the age of the constituent formations
(Vandyke and Byrnes, 1974). The Oakdale Formation is now elevated to group
status and comprises a lower, Cypress Hill Voleanics and an upper, Mona Vale
Siltstone (Vandyke and Byrnes, 1974). Unconformably overlying this is the
Dripstone Group, consisting of the Wylinga Formation, Warderie Volcanics and
Catombal Park Formation, in ascending stratigraphic order. The Dripstone
Group is equivalent to the upper part of the former Oakdale Formation of Strusz
(1960). Finally, the Mumbil Formation was also raised to group status and
includes the Narragal Limestone and overlying Barnby Hills Shale.

The rugose corals described herein are represented in three horizons of the
Dripstone and Mumbil Groups. From the Catombal Park Formation of the
Oakdale Anticline Palaeophyllum oakdalensis sp. nov., Holmophyllum struszt
sp. nov. and Holmophyllia maculosa sp. nov. are described. P. oakdalensis is
particularly abundant in the calcareous beds at the base of this formation while
the other two species are rare. The Narragal Limestone was divided in some
places into an upper and lower member separated by a thin horizon of volcanic
sandstone (Vandyke, 1970; WVandyke and Byrnes, 1974). From the upper

* Department of Geology and Geophysics, University of Sydney, New South Wales, 2006.
Present address : Institute of Sedimentary and Petroleum Geology, Geological Survey of Canada.
3303—-33rd St. N.W., Calgary. Alberta, Canada, T2L 2A7.

PROCEEDINGS OF THE LINNEAN SOCIETY oF NEw SoutH WALES, Vot. 99, Part 4

182 SILURIAN RUGOSA FROM CENTRAL NEW SOUTH WALES

member of this formation Holmophyllia cf. simplex (Lewis) is described, and
Coronoruga dripstonense Strusz probably also comes from this upper member.
Pilophyllum multiseptatum (Strusz) occurs in a limestone lens of the Barnby Hills
Shale south of the Bell River (probably the lens marked as Loc. 7 in Text-fig. 1).
This lens was regarded as equivalent to the Narragal Limestone by Strusz (1961)
but was considered to overlie that formation by Vandyke (1970).

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3. Collecting Locality

Fig. 1. Geological sketch map of the Mumbil area, showing collecting localities. Geology
based on Vandyke (1970, unpubl.).

An unconformity between the Oakdale Group and Wylinga Formation was
recognised by Vandyke (1970). Inthe Oakdale Anticline, shales with Retiograptus
occur below the unconformity and, to the west, definite Ordovician graptolites
and trilobites are found in shales of the Oakdale Group (Strusz, 1960, 1961 ;
Vandyke, 1970; Webby, 1973). In the limestone lenses of the Wylinga
Formation in the Oakdale Anticline, the rugosan Rhizophyllum is represented,

PROCEEDINGS OF THE LINNEAN SocreTy oF New SoutH WALES, Vou. 99, Part 4

R. A. McLEAN 183

comprising forms showing similarities to those of the ?late Wenlock to Ludlow
of the Yass District (see also Vandyke, 1970). This would suggest that the
Llandovery and probably much of the Wenlock is missing in the sequence. The
species of Rhizophyllum show no close similarities to the large from represented. in
the Upper Llandovery (Rosyth and Quarry Creek Limestones) of central New
South Wales (McLean, 1974bd).

From horizons low in the Barnby Hills Shale overlying the Narragal Lime-
stone, abundant Pristiograptus bohemicus (Barrande) is represented (Strusz,
1960), together with rare Neodiversograptus nilssoni (Lapworth) (Vandyke, 1970 ;
C. J. Jenkins, pers. comm., 1974). Hence the Narragal Limestone would not
appear to range in age higher than the nilssoni zone (basal Ludlow). Therefore
the Dripstone and Mumbil Groups as defined by Vandyke and Byrnes (1974)
and the faunas described in this paper would appear to range most probably
from late Wenlock to early Ludlow in age.

Specimen numbers in the University of Sydney Palaeontological Collections
bear the prefix SUP and where more than one section has been prepared from
the one specimen, they have the suffix a, b, ete. Numbers of specimens in the
palaeontological collections of the Australian Museum, Sydney, have the prefix
AM.F and thin sections in these collections bear the prefix AM.

SYSTEMATIC PALAEONTOLOGY
Family CYSTIPHYLLIDAE Edwards and Haime, 1850
Genus HOLMOPHYLLUM Wedekind, 1927

1927 Holmophyllum Wedekind, p. 31

1940 Nipponophyllum Sugiyama, p. 115

1940 Baeophyllum Hill, p. 403

1961 Nipponophyllum (part.); Strusz, p. 346

1963 Nipponophyllum (part.); Ivanovskiy, p. 112
non 1934 Holmophyllum; Lewis, p. 95

For further synonymy see McLean, 1974).
Type species. H. holmi Wedekind, 1927. Eke Beds (Ludlow), Gotland.

Diagnosis. Solitary or fasciculate corallum. Septa consist of discrete
rhabdacanthine trabeculae piercing several dissepiments or tabulae. Tabularium

generally clearly differentiated from peripheral zone of axially inclined
dissepiments.

Discussion. The affinities and species composition of the genus
Holmophyllum were reviewed by McLean (1974b). It was suggested in that paper
that the branching form Nipponophyllum Sugiyama, 1940, was closely allied to
Holmophyllum, but was tentatively separated from that genus on the basis of
having more contiguous septal spines, particularly in the Tadzhikistan rep-
resentatives of the genus. There is insufficient figured material of the type
species of Nipponophyllum, N. giganteum Sugiyama, 1940, for detailed comparison
but from what is available the septal spines would appear to be mainly discrete
(Sugiyama, 1940, Pl. XXX, figs 3, 4; Text-fig. 3). The Australian genus
Baeophyllum Hill, 1940 (regarded as a synonym of Nipponophyllum by most
subsequent workers, e.g. Hill, 1956; Strusz, 1961; Ivanovskiy, 1963 ;
Lavrusevich, 1967) was described as having both complete septa and discrete
septal spines (Hill, 1940, p. 404). However, re-examination of the type material
of B. colligatum Hill has shown no definite evidence of complete septa, although
material is extensively recrystallised. Discrete septal spines may be sparsely
seen and the holotype of the species (AM.F 9148) is re-illustrated herein for

PROCEEDINGS OF THE: LINNEAN SocieTY oF New SoutH WALES, Vou..99, Part 4

184 SILURIAN RUGOSA FROM CENTRAL NEW SOUTH WALES

comparison with other forms (Text-fig. 2; Pl. VIII, figs 1-5). Lavrusevich
(1967) regarded Nipponophyllum as a branching form of Holmophyllum but it is
felt that such a difference is probably not of generic importance. Hence both
Nipponophyllum and Baeophyllum are here regarded as synonyms of
Holmophyllum.

Fig. 2. Holmophyllum colligatum (Hill), Bowspring Limestone, Yass. a. AM704, holotype,
transverse section, x6. b. AM 704, holotype, longitudinal section, x 6.

The Tadzhikistan species of ‘“‘ Nipponophyllum” described by Lavrusevich
(1967) are referred to Holmophyllia Sytova, 1970 (see below).

Species of Holmophyllum bearing a branching corallum (Group III of McLean,
1974b) are now taken to include H. obscurum Smelovskaya, H. subtenuis
Shurygina, ?H. primaevum Wang, H. gigantewm (Sugiyama), H. colligatum (Hill)
and H. struszi sp. nov.

Holmophyllum struszi sp. nov.

Pl. VIII, figs 6-9, Pl. IX, figs 1-3; Text-fig. 3

1961 Nipponophyllum sp. cf. giganteum Sugiyama, 1940; Strusz, p. 347,
Pl. 43, figs 3, 4; Pl. 45, fig. 12.

Derwation of name. After Dr. D. L. Strusz, who first worked on the coral
faunas of the Mumbil area.

Material. Holotype SUP 74220, Paratypes SUP 10289, 40279 Catombal
Park Formation, Oakdale Anticline, Mumbil area (Loc. 4). ?Late Wenlock.

Diagnosis. Corallum fasciculate or phaceloid. Septal spines stout, sparsely
developed, of moderate length. Dissepiments large, strongly elongate ; tabulae
complete and incomplete, widely spaced, in strongly sagging series.

Description. Corallum fasciculate or phaceloid, showing outgrowths of the
dissepimentarium for support of adjacent corallites. Dimensions of corallum
reach at least 27 em high, with diameter of 25 em. Increase is apparently
calicinal. Corallites are closely spaced with diameter ranging from 9-12 mm in
mature specimens.

Septal spines are rather stout, in 30-36 radial rows which are usually clearly
defined. Spines are confined mainly to surface of dissepiments and are only
rarely developed on tabulae. They generally pierce no more than 2-3 dissepi-

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R. A. MCLEAN 185

ments. Trabecular microstructure is obscured by recrystallisation (Pl. VIII,
fie) 4).

Dissepiments are mainly strongly elongate, of very variable size although
generally large, particularly at corallite periphery. They occur in 2-5 rows and
are very steeply inclined towards axis of corallite. Tabulae are both complete
and incomplete, very widely spaced and form strongly sagging series.
Tabularium occupies approximately half of corallite diameter.

Fig. 3. Holmophyllum struszi sp. nov., Catombal Park Formation, Mumbil area. a. SUP
74220e, holotype, transverse section, x4. b. SUP 74220h, holotype, longitudinal section, x 4.

Remarks. Strusz (1961) closely compared this form with H. gigantewm
(Sugiyama, 1940) from the Lower Ludlow of the Kitakami Mountains, Japan,
the latter being type species of ‘‘ Nipponophyllum”’. However, H. gigantewm
may be clearly distinguished by its larger number of septal spines and flatter,
closer, more complete tabulae. It is felt therefore that the New South Wales
form warrants erection of a new species.

H. colligatum (Hill, 1940) from the Bowspring Limestone (early or middle
Ludlow) of Yass, New South Wales, is closely comparable to H. struszi in growth
form but differs in having generally sparser, more irregularly developed septal
spines, together with flatter more complete tabulae.

Genus HOLMOPHYLLIA Sytova, 1970
1934 Holmophyllum; Lewis, p. 95 (non Wedekind, 1927)
21952 Holmophyllum (part.); Bulvanker, p. 12
1967 Nipponophyllum; WUavrusevich, p. 18 (non Sugiyama, 1940)
21968 Gukoviphyllum Sytova, p. 54
1970 Holmophylliia Sytova, p. 68
21973 Gukoviphyllum ; Fedorowski and Gorianov, p. 32

Type species. Holmophyllia boreale Sytova, 1970. Vaygach Horizon (Lower
Devonian), Vaygach Island.

Diagnosis. Solitary or fasciculate corallum. Septa comprise
rhabdacanthine trabeculae partly fused vertically near corallite periphery,

PROCEEDINGS OF THE LINNEAN SOCIETY oF NEw SoutH WALES, VoL. 99, Part 4

186 SILURIAN RUGOSA FROM CENTRAL NEW SOUTH WALES

becoming discrete at their axial ends and in tabularium. Tabulae complete and
incomplete, dissepiments very variable in shape and abundance.

Discussion. Holmophyllia is distinguished from Holmophyllum Wedekind
on the basis of having its rhabdacanthine trabeculae partly fused vertically to
give the impression of complete septa in the peripheral region of the corallite.
This small difference could perhaps be regarded as of subgeneric importance only
but it does appear to be a basic difference and so for the present full generic status
for Holmophyllia is retained.

The similarities between Gukoviphyllum Sytova, 1968, and Holmophyllum
and ‘*‘ Nipponophyllum ” were discussed by McLean (19746). In its long, partly
fused septal spines Gukoviphyllum is also closely allied to Holmophyllia. However,
according to Sytova (1968, p. 54; 1970, p. 68) the former lacks rhabdacanths,
the trabeculae instead having a fan-shaped appearance, although this may be a
feature of preservation. In all other respects, especially the fused spines,
Gukoviphyllum appears identical to Holmophyllia (see G. septatum, Bulvanker,
1952, Pl. IV, figs 2-4; Sytova, 1968, Pl. I, fig. 5b; Fedorowski and Gorianov,
1973, Pl. VI, fig. 7b) and most probably Holmophyllia will prove to be a junior
synonym of Gukoviphyllum. However, until the septal microstructure of
Gukoviphyllum is illustrated and its synonymy with Holmophyllia confirmed or
denied, it is felt that the latter genus should be retained for forms bearing definite
rhabdacanths.

Holmophyllum simplex Lewis, 1934, from ? Wenlock pebbles in the Stack
Series of the Isle of Man, has rather short rhabdacanthine trabeculae but these
are clearly fused peripherally (Lewis, 1934, Pl. VIII, fig. a). The species is here
regarded as a representative of Holmophyllia. Sytova (1970, p. 67) included it
in Gukoviphyllum, but its definite rhabdacanths (Wang, 1947) would seem to
preclude it from that genus.

The three species of ‘“‘ Nipponophyllum” (N. turkestanicum, N. minimum
and N. reimani) described by Lavrusevich (1967) from the Upper Silurian of
Tadzhikistan all exhibit partly fused, long rhabdacanthine trabeculae and hence
are included here in Holmophyllia.

The following species are here regarded as representative of Holmophyllia :
H. boreale Sytova (type species), H. simplex (Lewis), H. turkestanicum
(Lavrusevich), H. minimum (Lavrusevich), H. retmani (Lavrusevich) and H.
maculosa sp. Nov.

Range. % Wenlock of Isle of Man; ? Late Wenlock of New South Wales ;
Upper Silurian of Tadzhikistan; Lower Devonian of Vaygach Island.

“ Gukoviphyllum”’ is represented in the Pridoli of Podolia and late Ludlow-early
Pridoli of Estonia.

Holmophylia maculosa sp. nov.
Pl. IX, figs 4-7; Text-fig. 4

Derivation of name. Latin, maculosus =spotted, referring to appearance of
tabularium due to septal spines in transverse section.

Material. Holotype SUP 40261. Catombal Park Formation, Oakdale
Anticline, Mumbil area (Loc. 5). ? Late Wenlock. Collected by A. Byrnes,
1970.

Diagnosis. Corallum fasciculate. Septal spines partly fused vertically in
peripheral regions of corallite, becoming discrete axially. Entirely discrete
septal spines abundantly developed in tabularium. Dissepiments in narrow
zone, elongate ; tabulae closely spaced, forming strongly sagging series.

Description. Only one small fragment of a corallum was available for
study. Growth form is fasciculate, the corallites quite widely spaced (average
5 mm). Corallite diameter ranges from 9-13 mm in mature stage.

PROCEEDINGS OF THE LINNEAN Society or New SourH WALES, Vou. 99, Part 4

R. A. McLEAN 187

Septa comprise long rhabdacanths, partly fused vertically in peripheral
regions of corallite, becoming discrete at their axial ends. In inner parts of
dissepimentarium, septal spines are entirely discrete, composed of broad
rhabdacanths. In tabularium, spines are thinner and abundantly developed on
tabularial crests. Spines pierce up to 8 dissepiments or tabulae. In transverse
section well-defined radial rows of spines are evident with differentiation into
major and minor septa. Minor septa consist mainly of one spine in a transverse
section, extending 0-2—0-3 of length of major septa which are composed of several
spines. Major septa, which range in number from 31-35, cannot usually be clearly
traced far into tabularium, where they are replaced by an irregular cluster of
spines (Text-fig. 4a; Pl. IX, fig. 7). Minor septa are not always clearly
apparent owing to extensive recrystallisation around most corallite margins.

Fig. 4. Holmophyllia maculosa sp. nov., Catombal Park Formation, Mumbil area. a. SUP
40261la, holotype, transverse section, x5. b. SUP 40261c, holotype, longitudinal section, x 5.

Dissepiments are weakly to strongly elongate, of moderate size and very
steeply inclined to corallite axis. They occur generally in 2-3 rows. Tabulae
are both complete and incomplete, occurring in strongly sagging series. Peripher-
ally tabulae are widely spaced and steeply inclined to axis, becoming flat or
weakly arched in axial region where they are very closely spaced.

Remarks. In its strong development of discrete septal spines in the
tabularium and long major septa this species is comparable to the branching
form H. reimani (Lavrusevich, 1967) from the Upper Silurian of the Turkestan
Range of Tadzhikistan. However, the latter species may be distinguished by
having fewer septa, longer minor septa and a very narrow tabularium.

Holmophyllia cf. simplex (Lewis, 1934)
PIS Xe hie Sic Pl ex fies — 7). Next-tig.-

Material. SUP 39184, representing several corallites embedded in a
stromatoporoid. Upper member of Narragal Limestone, Oakdale Anticline,
Mumbil area (Loc. 6). Collected by A. Byrnes, 1970. SUP 74228, same horizon,
western area of outcrop (Loc. 1). Late Wenlock or early Ludlow.

Diagnosis. Corallum solitary. Septal spines of short to moderate length,
in partial lateral and vertical contact peripherally, forming distinct major and
minor septa in late growth stages. Septal spines mainly discrete where based
on dissepiments, but not developed in tabularium. Dissepiments small ;
tabulae widely spaced, strongly sagging.

= T 4
sa Es by o > J
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188 SILURIAN RUGOSA FROM CENTRAL NEW SOUTH WALES

Description. Corallum appears to be solitary, growing commensally with
a stromatoporoid. Corallites are subcylindrical with height at least 30 mm and
diameter reaching 9 mm.

Septal spines are of short to moderate length, rhabdacanthine, dilated at
corallite wall to be in lateral contact, forming a very narrow septal stereozone in
some places. Spines are also partly fused vertically at corallite periphery
(Pl. X, figs 2, 4) but where spines are based on dissepiments, which is uncommon,
they are typically discrete.

It is rare for spines to extend through dissepimentarium to tabularium.
Septa range in number from 46-50 in mature corallites, minor septa being
0-2-0-5 of length of major septa, the latter extending up to 0-6 of corallite
radius. Minor septa do not appear to be developed until late in ontogeny.

Fig. 5. Holmophyllia cf. simplex (Lewis), Narragal Limestone, Mumbil area. a. SUP 39184a,
transverse section, x7. b. SUP 39184c, longitudinal section, x 5.

Dissepiments are small, globose to weakly elongate, in 3-4 rows steeply
inclined to corallite axis. Tabulae form very strongly sagging series, complete
and incomplete, widely spaced.

Remarks. This form appears very closely similar to the ? Wenlock species
H. simplex (Lewis, 1934) from the Stack Series, Isle of Man. The only difference
would seem to be greater development of small, globose dissepiments at earlier
growth stages in the New South Wales material. However, as only one figured
longitudinal section of the type material is available (Lewis, 1934, Pl. VIII, fig. a),
the New South Wales form cannot be more closely compared.

Genus CORONORUGA Strusz, 1961
1961 Coronoruga Strusz, p. 347

Type species. CU. dripstonense Strusz, 1961. Narragal Limestone, Mumbil
area. Late Wenlock or early Ludlow.

Diagnosis. (Based on Strusz, 1961, p. 347.) Large solitary corallites with
a wide dissepimentarium separated from a rather narrow tabularium by a narrow
stereozone in which are embedded short, discrete trabeculae.

Discussion. Ivanovskiy (1965a, b) suggested Coronoruga may be a synonym
of Gyalophyllum Wedekind, 1927. However, the latter has coarse, very closely
spaced septal spines almost completely occupying the space of the dissepiment-
arium whereas in Coronoruga the spines are largely confined to the stereozone
separating the dissepimentarium and tabularium. The septal structure of
Coronoruga with trabeculae, wrapped in lamellar sclerenchyme, occurring in
layers on dissepiments and tabulae is comparable to that of Angullophyllum

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R. A. McLEAN 189

McLean, 1974), found in the Upper Llandovery of Angullong, central New South
Wales. The latter differs in its smaller skeletal elements and compound growth
form. Coronoruga appears to be endemic to Australia.

Coronoruga dripstonense Strusz, 196i
Pl. X, figs 8-10, Pl. XI, figs 1-3
1961 Coronoruga dripstonense Strusz, p. 348, Pl. 42, figs 9, 17

Material. Holotype SUP 11104, Paratypes SUP 11101, 11212-11215,
11195, 11198. Narragal Limestone, Mumbil area (Loc. 2). Late Wenlock or
early Ludlow.

Diagnosis. See Strusz, 1961.

Description. See Strusz, 1961.

Remarks. From further sections made of the paratypes it is evident that
where layers of sclerenchyme occur on dissepiments and tabulae, short septal
Spines are sparsely present although recrystallisation generally obscures this
feature. The trabecular microstructure is largely also obscured by recrystallisa-
tion but it appears most probably to be rhabdacanthine, the individual trabeculae
being wrapped in lamellar sclerenchyme.

Family ENDOPHYLLIDAE Torley, 1933
Genus PILOPHYLLUM Wedekind, 1927

1927 Pilophyllum Wedekind, p. 39
21937 Sinospongophyllum Yoh, p. 56
71942 Sinospongophyllum ; Hill, p. 20
1944 Pilophyllum; Wang, p 23
1952 Pilophyllum; Bulvanker, p. 20
1956 ? Pilophyllum; Hill, p. F301
1961 Nipponophyllum (part.); Strusz, p. 346 (non Sugiyama, 1940)
1966 Pilophyllum (? part.) ; Sytova and Ulitina, p. 227
1968 Pilophyllum (part.); Lavrusevich, p. 110
non 1958 Pilophyllum; Kaljo, p. 115
non 1960 Pilophyllum; Zheltonogova, p. 77
non 1963 Pilophyllum ; Ivanovskiy, p. 60
non 1969 Pilophyllum ; Hill et al., p. 5, 12.

Type species. P. keyserlingi Wedekind, 1927. Hemse Beds (Ludlow),
Gotland.

Diagnosis. Solitary or fasciculate corallum. Septa thin, with well-defined
stereozone on corallite wall and dissepimental surfaces. Dissepiments typically
large, lonsdaleoid, sparsely developed. Tabulae incomplete, in arched series.

Discussion. The genus Sinospongophyllum Yoh, 1937, may well be
synonymous with Pilophyllum. Hill (1942) considered the type species, S.
planotabulatum Yoh, 1937 (Givetian, Kwangsi) differed from Pilophyllum in
lacking greatly thickened septa and having more complete tabulae. However,
from the illustrations of Wedekind (1927, Pl. 8, figs 1-6), the extent to which the
septa of Pilophyllum are thickened in the stereozone is quite variable, that of
P. progressum Wedekind being quite comparable to that of S. planotabulatum
(Wedekind, 1927, Pl. 8, figs 5,6; Yoh, 1937, Pl. VI, figs 2-5). The tabulae of
Sinospongophyllum are certainly more complete than is evident in the Gotland
specimens of Pilophyllum illustrated by Wedekind (1927) but this is probably not
a significant generic difference. The material described below as P. multiseptatum
(Strusz) has tabulae roughly intermediate in character between P. keyserlingi
Wedekind (type species) and S. planotabulatum. Consequently Sinospongophyllum
is herein tentatively regarded as a synonym of Pilophyllum. ‘“ Sinospongo-

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190 SILURIAN RUGOSA FROM CENTRAL NEW SOUTH WALES

phylum” is also represented in the Lower Devonian of Queensland (‘‘ 8.”
abrogatum Hill, 1942, p. 20). Affinities of ‘‘ Sinospongophyllum”’ with other
Devonian forms have been reviewed by Hill and Jell (1970, p. 64).

Of Silurian genera, Pilophyllum shows perhaps closest similarities to
Kyphophyllum Wedekind, 1927 (probably a synonym of Strombodes Schweigger,
1819—see McLean, 1974a). As pointed out by Merriam (1972, p. 37), Pilophyllum
differs from ‘“* Kyphophyllum ” in having a broader septal stereozone and sparser
development of dissepiments, the latter tending to be larger than those of
‘“ Kyphophyllum”’. The type species of Strombodes, S. stellaris (Linnaeus) with
its narrower dissepimentarium shows close similarities to Prlophyllum in
longitudinal section (Smith, 1945, Pl. 29, fig. 2), but its lack of stereozone serves
to distinguish it from Prlophyllum.

The genus Pseudopilophyllum was proposed by Lavrusevich (1971) for forms
similar to Pilophyllum but differing in having numerous small dissepiments as
opposed to the large, sparse dissepiments of Pilophyllum. It may be distinguished
from Pilophyllum also by possessing thickened septa in the tabularium.
** Pilophyllum ” moyeroense Ivanovskiy, 1963, from the Upper Wenlock of the
Siberian Platform was designated as type species of Pseudopilophyllum and the
species is also represented in the Lower Wenlock (Horizon K) of Tadzhikistan
(Lavrusevich, 1971, p. 69).

The three species of Pilophyllum described by Kaljo (1958) from the late
Ordovician-early Silurian of Estonia (P. porosum, P. zonatum and P. massivum)
all possess an axial structure and thus would not appear to be representative of
that genus. This has been recognised also by Sytova and Ulitina (1966, p. 227)
and Lavrusevich (1968, p. 110). Ivanovskiy (1965b) has listed these three species

as possible representatives of Paliphyllum Soshkina.

Zheltonogova (1960) described two species of Pilophyllum (P. insolitum and
P. angustum) from the Tom-Chumysh Beds (Lower Devonian) of the Salair,
south-west Siberia. Both these forms require further study but do not appear
to be representative of Pilophyllum as suggested by Sytova and Ulitina (1966,
p. 228) and Hill (1967, p. 668). ‘‘ P.” insolitum possesses numerous dissepiments
and ‘*‘ P.” angustum, while appearing rather similar to Pilophyllum in longitudinal
section (Zheltonogova, 1960, Pl. S—18, fig. 1), does not appear to have clearly
developed lonsdaleoid septa and has been referred to the new genus Salairophyllum
by Besprozvannykh (1968).

P. bimurum Sytova im Sytova and Ulitina, 1966, from the Lower Devonian
Nadaynasuy Horizon of Kazakhstan differs from typical Pilephyllum in having
an ‘‘ inner wall”’ of thickened septa. However, in all other respects it appears
closely similar to that genus.

The form illustrated by Hill et al. (1969, Pl. SVI, fig. 9) as Pilophyllum sp.
from the upper part of the Jack Limestone Member of the Graveyard Creek
Formation, Broken River, North Queensland (?. M.-U. Silurian, Hill et al., 1969,
p-s 12) differs from Pilophyllum in lacking a septal stereozone and having a very
Saas axial vortex. It may be congeneric with a form described as
Tabulophyllum ? lowry Hill and Jell, 1970, from the Upper Devonian Napier
Formation of the Canning Basin, Western Australia. As mentioned by Hill and
Jell (1970, p. 66) this latter form probably represents a new genus.

The species composition of Pilophyllum is here taken to be: P. keyserlingt
Wedekind, P. munther Wedekind, P. sayuhoense Wang and P. multiseptatum
(Strusz). Doubtfully included in the genus are P. bimurum Sytova,
“ Sinospongophyllum ” planotabulatum Yoh and “ 8.” abrogatum Hill.

Range. Wenlock of China; Ludlow of Gotland, Podolia and New South
Wales; Upper Silurian of Tadzhikistan and Kazakhstan ; Lower Devonian of
? Kazakhstan and ? Queensland ; Middle Devonian of ? China.

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R. A. MCLEAN 191

Pilophyllum multiseptatum (Strusz, 1961)
Pl. XI, figs 4-6; Pl. XII, figs 1-3; Text-fig. 6
1961 Nipponophyllum multiseptatum Strusz, p. 346, Pl. 44, fig. 12; Pl. 45,
He9:
Material. Holotype SUP 11099, limestone lens in Barnby Hills Shale, south
of Bell River (Loc. 7?). Early Ludlow.

Diagnosis. Fasciculate Pilophyllum with well developed peripheral
Stereozone, dissepiments weakly developed and only late in ontogeny and tabulae
differentiated into axial and periaxial series.

Description. Corallum fasciculate with lateral increase. Corallites sub-
cylindrical with deep, steep-sided calice. Diameter of corallites ranges commonly
from 12-14 mm.

Septal number ranges up to 64 in mature corallites with all septa set in a
stereozone of lamellar sclerenchyme up to 1-5 mm wide. Sclerenchyme occurs
thickly on corallite wall and to a lesser extent on the dissepimental surfaces
(Pl. XI, fig. 5). Major septa are thin, extending 0-6—0-7 of corallite radius,
interrupted peripherally late in ontogeny by lonsdaleoid dissepiments. Minor
Septa are up to 0-5 of length of major septa, generally not extending far beyond
Stereozone on corallite wall or dissepimental surfaces. They may show weak
Separation of trabeculae at their inner margins.

Fig. 6. Pilophyllum multiseptatum (Strusz), limestone lens in Barnby Hills Shale, Mumbil
area. SUP 11099, holotype, transverse section, x 4.

Dissepiments are lacking in early growth stages, septa extending uninter-
rupted from corallite wall (Pl. XI, fig. 4; Pl. XII, fig. 3). In late ontogeny,
dissepiments developed in 1—2 rows, large, strongly elongate and steeply inclined
with marked sclerenchymal coating. Tabularium width is approximately 0-7
of corallite diameter. Tabulae occur in two series: an outer, distally inclined
Series comprising closely spaced, incomplete tabulae and an axial series of widely
spaced, mainly flat, complete tabulae.

Remarks. This form was originally included in the genus Nipponophyllum
Sugiyama, 1940, by Strusz (1961) but differs from that genus in lacking discrete
septal spines. Further thin sections prepared from the holotype material have
confirmed the lamellar character of the septa. Unfortunately the longitudinal
section of the holotype figured by Strusz (1961, Pl. 44, fig. 12) has been partially
destroyed but an additional section figured herein (Pl. XII, fig. 2), although not
as well oriented, confirms the character of the tabulae and dissepiments. The
material listed as a paratype of this species (SUP 10299, Strusz, 1961, p. 346) is
of a poorly preserved carinate form and would not appear to be conspecific with
the above described species.

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192 SILURIAN RUGOSA FROM CENTRAL NEW SOUTH WALES

P. multiseptatum shows perhaps closest internal similarities to P. weissermelr
Wedekind, 1927 (?=P. keyserlingt), from the Klinteberg Beds (Upper Wenlock-
Lower Ludlow) of Gotland and Akkan Horizon (Upper Silurian) of Kazakhstan
(Wedekind, 1927; Sytova and Ulitina, 1966). P. multiseptatum may be dis-
tinguished however, by its much smaller size, broader stereozone on the corallite
wall and differentiation of the tabulae into two series.

P. multiseptatum also shows similarities to the type species of ‘* Sinospongo-
phyllum”, “8S.” planotabulatum Yoh, 1937, from the Middle Devonian of China.
Particularly comparable are the degree of peripheral septal thickening and the
complete tabulae in the axial region. However, ‘‘ 8S.” planotabulatum is a much
larger, solitary form and has less development of peripheral, incomplete tabulae,
the axial complete tabulae being more closely spaced.

Family STAURIIDAE Edwards and Haime, 1850
Genus PALAEOPHYLLUM Billings, 1858

1969 Palaeophyllum ; Ivanovskiy, p. 80 (cum syn.)

1972 Palaeophyllum ; Webby, p. 151

1972 Palaeophyllum; Merriam, p. 29

Type species. P. rugosum Billings, 1858. Lake St. John, Little Discharge,
Quebec. Black River or early Trenton.

Diagnosis. (Modified from Hill, 1961, p. 4.) Corallum phacelo-cerioid.
Corallites have narrow peripheral stereozone and lack dissepiments. Major
septa are long and thin and minor septa are generally short. Tabulae are thin,
commonly complete and arched, sometimes with axial depression.

Discussion. Ivanovskiy (1969) proposed that the known species of
Palaeophyllum could be grouped into three broadly defined species: P. thomi
(Hall, 1857); P. fasciculum (Kutorga, 1837) and P. lebediensis (Tcherepnina,
1960). The writer agrees with Webby (1972) that these three “ species’? may
be better considered as species groups (or ? subgenera). P. oakdalensis sp. nov.
described below, with well-developed minor septa and flattened tabulae, is
representative of the P. fasciculum group, as suggested by Webby (1972). nhs

Range. Lower Caradoc of Scotland, North America and New South Wales ;
Upper Caradoc of North America, ? Urals and ? south-west Siberia; Ashgill of
Norway, Estonia, Urals, Siberian Platform, south-west Siberia, North America
and ? China; Lower Llandovery of Estonia, Urals, Siberian Platform and
? North America; Upper Llandovery of Norway, Greenland and ? Siberian
Platform ; ? Late Wenlock of New South Wales ; Late Wenlock-Ludlow of New
York; Ludlow of ? California.

Palacophyllum oakdalensis sp. nov.
Pl. XII, figs 4-8; Text-fig. 7
1961 Palaeophyllum rugosum Billings ; Strusz, p. 341, Pl. 42, figs 7, 8, 15;
Text-fig. 3.

Derivation of name. After the property of ‘‘ Oakdale ”’.

Material. Holotype SUP 74229 Catombal Park Formation, Oakdale
Anticline, Mumbil area (Loc. 3). Paratypes SUP 10240, 10244-10246, 10248—
10250, 74230-74235. Same horizon, Locs 3, 4, 5. ? Late Wenlock.

Diagnosis. Palacophyllum with average corallite diameter 7 mm. Major
septa extend to nearly 0-8 of corallite radius at maturity, minor septa generally
0-4 length of major septa. Peripheral stereozone well developed and major

septa average 20 in number. Tabulae range in shape from flat to strongly
arched, with gently arched types most common; spacing very variable.

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R. A. MCLEAN 193

Description. Corallum phaceloid. Increase is calicinal and parricidal,
with four offsets from the one corallite being most common (Pl. XII, fig. 8).
Possible lateral budding is also represented. Corallite diameter ranges from
6-5-8 mm in mature specimens with an average of 7mm. Septa are of two orders,
set in a narrow but clearly developed peripheral stereozone of lamellar
sclerenchyme. ‘This stereozone averages 0-1 of radius of corallite. Major septa
are thin, tapering slightly towards axis. In early growth stages, major septa
extend almost to corallite axis (Pl. XII, fig. 8), but in maturity they are somewhat
withdrawn extending 0-7—0-8 of corallite radius (Pl. XII, fig. 5) and numbering
20-22. Minor septa reach 0-35-0-5 of length of major septa.

Cc d

Fig. 7. Palaeophyllum oakdalensis sp. nov., Catombal Park Formation, Mumbil area.
a. SUP 74233, paratype, transverse section of immature specimen, x6. b. SUP 74229b, holotype,
transverse section of mature specimen, x6. c. SUP 74231b, paratype, longitudinal section
showing most commonly observed tabular shape, x4. d. SUP 74230b, paratype, longitudinal
section showing rarely developed highly arched tabulae, x 4.

Tabulae are generally flat with weakly downturned edges. Degree of
downturning is very variable, some barely showing this feature while others are
markedly domed (Text-figs 7c, d). Tabulae are only rarely incomplete. Spacing
of tabulae is quite variable. In a few examples they are closely spaced (0-1 mm)
and grouped in series (Pl. XII, fig. 4) but in general they are evenly and widely
spaced, average 0-:8-1-2 mm. Axial depression of tabulae is very weak or more
commonly absent.

Remarks. Strusz (1961) originally described this form as belonging to
P. rugosum Billings, 1858, the Upper Ordovician type species from Quebec.
However, from the descriptions of Hill (1961) it may be clearly distinguished
from P. rugosum by having parricidal increase, longer minor septa and tabulae
lacking a marked axial depression. Webby (1972, p. 152) also recognised the
distinction of Strusz’s form from the type species. }

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194 SILURIAN RUGOSA FROM CENTRAL NEW SOUTH WALES

Species of Palaeophyllum have not been commonly recorded from the Silurian
and the only forms of comparable age to P. oakdalensis are P. multicaule (Hall,
1852) redescribed by Oliver (1963), and Palaeophyllum ? sp. G of Merriam (1972).
P. multicaule is represented in the Lockport Dolomite (late Wenlock-late Ludlow)
of Lockport, New York, and may be distinguished from P. oakdalensis in having
smaller corallite diameter, longer major septa and shorter minor septa.
Palaeophyllum? sp. G occurs in the Gazelle Formation (Ludlow) of the Klamath
Mountains, California. It has flattened tabulae comparable to some specimens
of P. oakdalensis but differs in having shorter septa, particularly in early growth
stages (Merriam, 1972, Pl. 2, figs 9, 10). In its short septa (approximately 0-5
of corallite radius), Palaeophyllum? sp. G would appear to be intermediate in
character between typical Palaeophyllum and Pycnostylus Whiteaves.

None of the previously described Australian representatives of Palacophyllum
(Webby, 1972) bear any close similarities to P. oakdalensis.

ACKNOWLEDGEMENTS

The author is grateful to Associate Professor B. D. Webby for critical review
of the manuscript and to Mrs. A. Byrnes for providing both material for study
and unpublished information about the stratigraphy of the Mumbil area.

The project was. supported by a Macleay Fellowship of the Linnean Society
of New South Wales:

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area. Unpubl. B.Sc. (Hons.) thesis, University of Sydney.

and Byrnss, J. G., 1974.—Review of the Palaeozoic succession beneath the Narragal
Limestone in the Oakdale Anticline near Mumbil, N.S.W. (In manuscript.)

Wana, H. C., 1944.—Silurian rugose corals from Eastern and Northern Yunnan. Bull. geol.
Soc. China, 24: 21-32.

1947.—New material of Silurian rugose corals from Yunnan. Bull. geol. Soc. China,
27: 171-192.

Wessy, B. D., 1972.—The rugose coral Palaeophyllum Billings from the Ordovician of central
New South Wales. Proc. Linn. Soc. N.S.W., 97: 150-157.

, 1973.—Remopleurides and other Upper Ordovician trilobites from New South Wales.
Palaeontology, 16: 445-475.

WEDEKIND, R., 1927.—Die Zoantharia Rugosa von Gotland (bes Nordgotland). Sver. geol.
Unders. Afh., Ser. Ca, 19: 1-94. .

You, S. 8., 1937.—Die Korallenfaunen des Mitteldevons aus der Provinz Kwangsi, Stidchina.
Palaeontographica, Abt. A, 87: 45-76.

ZHELTONOGOVA, V. A., 1960.—Siluriyskaya Sistema. Podklass Tetrakorally. In Khalfin,
L. L. (ed.), ‘° Biostratigrafiya paleozoya Sayano-Altayskoy gornoy oblasti. Tom. 2: Sredniy
Paleozoy’”’. Trudy sib. nauchno-issled. Inst. Geol. Geofiz. miner. Syr., 20: 74-86.

EXPLANATION OF PLATES

PLATE VIII

Figs 1-5. Holmophyllum colligatum (Hill), Bowspring Limestone, Hatton’s Corner, Yass
district, AM 704, holotype, x5. 1-2. transverse sections. 3-5. longitudinal sections.

Figs 6-9. Holmophyllum struszi sp. nov., Catombal Park Formation, Oakdale Anticline,
Mumbil area. 6. SUP 74220c, holotype, transverse section, x4. 7. SUP 10289, paratype,
transverse section showing recrystallised septal spines, x 25. 8. SUP 10289, paratype, transverse
section, x3. 9. SUP 74220g, holotype, longitudinal section, x 3.

PLATE IX
Figs 1-3. Holmophyllum struszi sp. nov., Catombal Park Formation, Oakdale Anticline,
Mumbil area. 1. SUP 74220d, holotype, longitudinal section, x4. 2. SUP 74220k, holotype,
longitudinal section, x4. 3. SUP 74220h, holotype, longitudinal section, x 4.
Figs 4-7. Holmophyllia maculosa sp. nov., Catombal Park Formation, Oakdale Anticline,
Mumbil area, x5. 4. SUP 40261d, holotype, longitudinal section. 5. SUP 4026le, holotype,
longitudinal section. 6. SUP 40261c, holotype, longitudinal section. 7. SUP 4026la, holotype,

transverse section.
Fig. 8. Holmophyllia cf. simplex (Lewis), Narragal Limestone, Mumbil area, x3. SUP

74228, tangential longitudinal section.

PLATE X

Figs 1-7. Holmophyllia cf. simplex (Lewis), Narragal Limestone, Oakdale Anticline, Mumbil
area. 1. SUP 39184a, transverse section showing rhabdacanthine septa, x25. 2. SUP 39184c,

PROCEEDINGS OF THE LINNEAN SocIETY OF NEw SoutH WALES, Vou. 99, Part 4

BB

196 SILURIAN RUGOSA FROM CENTRAL NEW SOUTH WALES

longitudinal section showing partially fused septal spines on corallite wall, x25. 3. SUP 39184c,
longitudinal section, x3. 4. SUP 39184c, longitudinal section showing fused septal spines on
eorallite wall, x25. 5-7. SUP 39184a, transverse sections of different corallites, x 7.

Figs 8-10. Coronoruga dripstonense Strusz, Narragal Limestone, Mumbil area, x3.
8. SUP 111038f, paratype, transverse section of early growth stage. 9. SUP 11103a, paratype,

longitudinal section. 10. SUP 11212, paratype, longitudinal section showing sclerenchymal
coating on tabulae.

PLATE XI

Figs 1-3. Coronoruga dripstonense Strusz, Narragal Limestone, Mumbil area, x3. 1. SUP
11215, paratype, transverse section. 2. SUP 11198a, paratype, transverse section. 3. SUP
11103b, paratype, longitudinal section showing sclerenchymal coating on dissepiments.

Figs 4-6. Pilophyllum multiseptatum (Strusz), limestone lens in Barnby Hills Shale, Mumbil
area. 4. SUP 11099f, holotype, transverse section, x4. 5. SUP 11099c. holotype, longitudinai
section showing lateral increase and sclerenchymal coating of dissepiments of incomplete corallite
at right, x3. 6. SUP 11099h, holotype, transverse section, x 4.

PLATE XII

Figs 1-3. Pilophyllum multiseptatum (Strusz), limestone lens in Barnby Hills Shales, Mumbil
area. 1. SUP 11099g, holotype, longitudinal section, x4. 2.SUP 11099d, holotype, longitudinal
section, <3. 3. SUP 110991, holotype, transverse section of early growth stage, x 4.

Figs 4-8. Palaeophyllum oakdalensis sp. nov., Catombal Park Formation, Oakdale Anticline,
Mumbil area. 4. SUP 74235, paratype, thin section (note variation in tabular spacing), x 3.
5. SUP 74229a, holotype, thin section, x4. 6. SUP 74229e, holotype, longitudinal section, x 4.
7. SUP 74229f, holotype, longitudinal section, x4. 8. SUP 74233, paratype, transverse section
(note quadripartite offsets at right), x 4.

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Proc. Linn. Soc. N.S.W., Vol. 99, Part 4 PLATE XI

Proc. Linn. Soc. N.S:W., Vol. 99, Part 4 PLATE XI

NUCLEIC ACIDS IN CHLOROPLASTS OF A CHLORELLA SP.
MOR. binpAy = Vo I CHAN; AND Kooy") CH0%

(Plates xm and XIV)

[Accepted for publication 24th July 1974]

Synopsis

Nucleic acids in chloroplasts of Chlorella (NM1) are demonstrated by light and electron
microscopy. The chloroplasts are positively stained by Feulgen stain. Electron microscopical
examination shows fine filamentous fibrils in association with lamellar membranes and ribosomes,
The fibrils are linear, single and double looped, and morphologically are not unlike that of DNA.
They are distributed throughout the chloroplast matrix and tend to accumulate around the
pyrenoid and in areas of low density between the chloroplast lamella. Polysomes are detected at
these sites. The fibrils are not observed after treatment with deoxyribonuclease. It is concluded
that these fibrils contain DNA and their significance is discussed.

INTRODUCTION

Biochemical studies indicate protein synthesis in the chloroplast as distinct
from cytoplasmic protein synthesis (Kirk, 1964; Ray and Hanawalt, 1965 ;
Smillie et al., 1967; Selsky, 1967). Such findings agree with the cytological
demonstration of RNA and DNA in chloroplasts of plants and algae (Ris and
Plaut, 1962; Brawerman, 1962, 1963; LHisenstadt and Brawerman, 1964 ;
Kislev et al., 1965 ; Gunning, 1965; Sagen eft al., 1965). In Chlorella however,
there is still no direct demonstration of DNA in the chloroplast although bio-
chemical investigations strongly support its presence (Kirk, 1971). The present
paper reports the location and morphology of DNA and RNA in the chloroplast
of Chlorella (NM1) using light and electron microscopy.

MATERIALS AND METHODS
Organism and Growth Conditions

A species of Chlorella, isolated from a garden soil and designated NM1, was
used.

Cells were grown in Erlenmeyer flasks in a defined mineral solution (Tchan,
1959) at pH 6-8 and aerated with CO,-enriched air. The flasks were placed in
a water shaker-bath at 28°C and illuminated by two (60W) incandescent lamps.
The intensity of incident light on growth flasks was 50 lux. Samples were taken
from two-day old cultures for cytological studies. Preliminary work indicated
that the removal of starch grains from the cells was necessary to facilitate the
detection of fine structure. This was achieved by keeping actively growing
poy old culture in the dark without agitation for 12 hours at room temperature
(22°C).

Light Microscopy

Cells were fixed in alcohol-acetic acid (3:1) for 10 minutes. They were
mounted on glass slides, washed with absolute ethanol and then brought to water

* Mauritius Sugar Industry Research Institute, Mauritius.
+ Department of Microbiology, University of Sydney, New South Wales, 2006.

t Linnean Macleay Lecturer in Microbiology, Department of Microbiology, University of
Sydney, New South Wales, 2006.

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198 NUCLEIC ACIDS FROM CHLOROPLASTS OF A CHLORELLA SP.

through a graded series of alcohol. Some slides were treated for 15 minutes in
N HCl at 60°C, and stained with Feulgen’s reagent for 90 minutes in darkness.
Other slides, after hydrolysis treatment, were stained with a Feulgen-fluorescent
dye for 60 minutes in the dark. Control slides were stained without hydrolysis.
All slides were washed in distilled water, and bleached in a mixture of sodium
bisulphite: N HCl, and distilled water (1: 3:3 w/v/v) for 10 minutes with two
changes. The slides were washed with tap water and mounted in glycerine.

Preparations stained with the fluorescent dye were observed by phase
contrast and fluorescence microscopy using a 100 oil immersion objective.
Fluorescence illumination was provided by an illuminator equipped with a
HB200 high-pressure mercury are lamp and a 3 mm BG12 excitation filter, and
K530 barrier filter. The phase and fluorescence images were recorded on Kodak
Super XX film with an initial magnification of 600 and a final enlargement
to x3,350.

Electron Microscopy

A. Cells were prefixed at room temperature in 2-5% glutaraldehyde in
sodium cacodylate buffer (0-045M, pH 6-2) for two hours, washed in the buffer
for 24 hours (three changes) and followed by :

(i) post-fixation in 1°% OsO, in Michaelis buffer (pH 6-0) overnight at room
temperature (Pease, 1964). The cells were washed three times in a
3-fold dilution of the buffer and then suspended in a small amount of
1:5% agar (50°C); or

(ii) post-fixation in freshly prepared 0-5°, unbuffered KMnO, solution for
10 minutes then washed five times in a 3-fold dilution of Palade’s (1952)
acetate buffer (pH 6-2) and suspended in a small amount of 1-5°% agar
(50°C).

Digestion by Deoxyribonuclease

B. Cells were fixed in 10% formalin in veronal acetate buffer (Pease, 1964)
(pH 6-2) for 10 minutes, washed in distilled water and digested with 0-001%
deoxyribonuclease I (Sigma) in veronal acetate buffer at 40°C for four hours.

The control sample was incubated in buffer only. After treatment cells
were prefixed in glutaraldehyde and post-fixed in potassium permanganate
solution as in A (ii), above. :

The small agar blocks from fixations A and B were stained with 0:5%
uranyl acetate buffered solution for two hours. Dehydration was carried out at
five-minute intervals in 30%, 50%, 70% and 80% ethanol followed by 10 minutes
in 90°% and two changes in 100% ethanol. The blocks were cut into approxim-
ately 2 mm pieces, infiltrated in three changes of araldite for one hour each and
overnight at room temperature. Polymerisation was realised in fresh araldite
at 60°C for 36-48 hours. Sections were cut on a Porter Blum ultra-microtome
MT-2 using glass knives and collected in nitrocellulose coated grids. After
staining with lead citrate (Reynolds, 1963), sections were examined and -photo-
graphed with a Philips EM200 or a Siemens Elmiskop I operating at 60 or 80 kV.

RESULTS
Light Microscopy
In Feulgen stained cells, the chloroplast gives a colour reaction of about
the same intensity as the nucleus (Pl. XIII, Fig. 1). A comparison of the internal
structures of cell shown by Feulgen-fluorescent staining with those observed in
phase contrast microscopy (Pl. XIII, Figs 2 and 3) clearly demonstrated the
presence of DNA positive sites in the chloroplasts.

PROCEEDINGS OF THE LINNEAN Society or New Souru Wats, Vou. 99, Part 4

A. R. PILLAY ET AL. 199

Electron Microscopy

Pl. XIII, Fig. 4 shows the presence of starch around the pyrenoid and in
areas between the lamellae of the chloroplast. Pl. XIII, Fig. 5 was printed for
maximum contrast to demonstrate areas of low electron density. These areas
are not bounded by membranes and contain fibrils (Pl. XIII, Fig. 5). They are
present in the vicinity of the pyrenoid region and randomly distributed within
the chloroplast (Pl. XIII, Fig. 5).

In OsO, fixed cells the low density areas contain conspicuous fine fibrils only
prior to deoxyribonuclease treatment (Pl. XIII, Fig. 7; Pl. XIV, Fig. 1). These
fibrils are removed after treatment by deoxyribonuclease (Pl. XIII, Fig. 6). At
a higher magnification, unattached filaments, and numerous DNA structures
attached to the lamellar membranes are well resolved in linear, single and double
looped configuration.

In sections of chloroplasts a number of DNA-containing areas occur in the
vicinity of the pyrenoid and between the lamellae (Pl. XIII, Figs 5 and 7;
Pl. XIV, Fig. 1). Their distribution is much more conspicuous than that
reported by Ris and Plaut (1962) for Chlamydomonas. Unlike the brown algal
(Gibbs, 1967, 1968; Bisalputra and Burton, 1969; Bisalputra and Bisalputra,
1969) they lack regular patterns.

Ribosomes and polyribosomes are frequently seen in close proximity or
adhering to fibrils near the lamellar membranes. Free ribosomes are in the
vicinity of pyrenoid and in the stroma regions between the lamellae (Pl. XIV,
Figs 1 and 2).

DISCUSSION

Direct fixation with OsO, or KMnO, usually fails to preserve adequately the
fine structures within the chloroplast (Gunning, 1965). KMnO, is reported to
extract RNA but stabilises DNA as a branching network of fibrils (Luft, 1956 ;
Bradbury and Meek, 1960; Nass and Nass, 1963; Nass et al., 1965). OsO,
can also result in leaching of proteins. It is not clear whether DNA fibrils are
restricted to areas of low electron density in the chloroplast or more widely
dispersed and obscured by ribosomes. It is known that direct KMnO, fixation
removes ribosomes (Hayat, 1970). The present study shows that KMnO, could
be used as a suitable post-fixative after the stabilisation of RNA and proteins by
gluteraldehyde.

The absence of fibrils in the low density areas of the ground substance in
the chloroplast after deoxyribonuclease treatment strongly suggests that these
fibrils contain DNA, and this is in accord with the appearance of the chloroplasts
before treatment and the positive Feulgen staining reaction (Pl. XIII, Figs 1,
2 and 3). The disappearance of ribosomes and the preservation of fibrils by
KMn0O, fixation also lends further support.

CONCLUSION

The present cytological investigation clearly demonstrated the presence of
DNA in chloroplast of Chlorella (NM1). It exists as a number of short single
units (Pl. XIV, Fig. 2). They are not restricted to the low electron density
areas but are also found in electron dense areas where their presence may some-
times be obscured by ribosomes. This suggests a wider distribution of the DNA
chloroplast. However, a few areas containing DNA appear to be randomly
distributed through the chloroplast matrix and around the pyrenoid region.
These areas may represent sites of greater activity in the chloroplast. Thus the
early biochemical evidence is now supported by direct cytological observation.

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutH WALES, Vot. 99, Part 4

200 NUCLEIC ACIDS FROM CHLOROPLASTS OF A CHLORELLA SP.

References

Bisatputra, T., and BisaupuTrRA, A. A., 1969. The ultrastructure of chloroplast of a brown
alga Sphacelaria sp. I. Plasted DNA configuration—the chloroplast genophore. J. Ultrastruct.
Res., 29: 151-170.

BisaLputra, T., and Burton, M., 1969.—The ultrastructure of chloroplast of a brown alga
Sphacelaria sp. II. Association between the chloroplast DNA and the photosynthetic lamellae.
J. Ultrastruct. Res., 29: 224-235.

BRADBURY, S., and MEEK, G. A., 1960.—A study of potassium permanganate fixation for electron
microscopy. Quart. J. Micro. Soc., 101: 241-250.

BRAWERMAN, G., 1962.—A specific species of ribosomes associated with the chloroplasts of
Euglena gracilis. Bioch'm. Biophys. Acta., 61: 313-315.

, 1963.—The isolation of a specific species of ribosomes associated with chloroplast
development in Huglena gracilis. Biophys. Acta., 72: 317-331.

Ersenstapt, J. M., and BrRawErMAN, G., 1964.—The protein-synthesizing systems from the
eytoplasm and the chloroplasts of Huglena gracilis. J. Mol. Biol., 10: 392-402.

GisBs, S.,1967.—Synthesis of chloroplast RNA at the site of chloroplast DNA. Biochem. Biophys.
Res. Commun., 28: 653-657.

, 1968.—Autoradiographic evidence for the in situ synthesis of chloroplast and mito-
chondrial RNA. J. Cell Sci., 3: 327-330.

GuNnNING, B. E. 8., 1965.—The fine structure of chloroplast stroma following aldehyde osmium-
tetroxide fixation. J. Cell Biol., 24: 79-93.

Hayat, M. A., 1970.—Principles and techniques of electron microscopy. Vol. 1: Biological applica-
tions. New York: Van Nostrand: pp. 59-65.

Kirk, J. T. O., 1964—DNA-dependent RNA synthesis in chloroplast preparations. Biochem.
Biophys. Res. Commun., 14: 393-397.

, 1971.—Will the real chloroplast DNA please stand up. Jn Boardman, N. K., Linnane,
A. W., and Smillie, R. M., Autonomy and biogenesis of mitochondria and chloroplasts.
Amsterdam: North-Holland: 267-276.

Kistev, N., Swret, H., and Bocorap, L., 1965.—Nucleic acids of chloroplasts and mitochondria
in Swiss chard. J. Cell. Biol., 25: 327-344.

Lurt, J., 1956.—Permanganate—a new fixative for electron microscopy.. J. Biochem. Biophys.
Cytol., 2: 799-800.

Nass, M. M. K., and Nass, S., 1963. Intramitochondrial fibres with DNA characteristics.
I. Fixation and electron staining reactions. J. Cell Biol., 19: 593-611.

; , and Arzetius, B. A., 1965.—The general occurrence of mitochondrial DNA.
Expt. Cell. Res., 37: 516-539.

PawapE, G. E., 1952.—A study of fixation for electron microscopy. J. Hxp. Med., 95: 285-298.

Pease, D. C., 1964.—Histological techniques for electron microscopy. New York: Academic
Press: 42-43.

Ray, D. S., and Hanawatr, P. C., 1965.—Satellite-DNA components in Huglena gracilis cells
lacking chloroplasts. J. Mol. Biol., 11: 760-768.

Reyno tps, E. S., 1963.—Use of lead citrate at high pH as an electron opaque stain in microscopy.
J. Mol. Biol., 17: 208-212.

Ris, H., and Praut, W., 1962.—Ultrastructure of DNA-containing areas in the chloroplasts of
Chlamydomonas. J. Cell Biol., 13: 383-391.

Sacan, L., Ben-SHaun, Y., Epstein, H. T., and Scurrr, J. A., 1965. Studies of chloroplast
development in Huglena. XI. Radioautographic localisation of chloroplast DNA. Plant
Physiol., 40: 1257-1260.

Sexsky, M. I., 1967.—Effects of puromycin aminonucleotide on growth and chloroplast develop-
ment of Huglena gracilis. Hxpt. Cell Res., 47: 237-245.

Suruire, R. M., Granam, D., Dwyer, M. R., Grieve, A., and Tosrn, N. F., 1967. Evidence for
the synthesis in vivo of proteins of the Calvin cycle and the photosynthetic electron transfer
pathway in chloroplast ribosomes. Biochem. Biophys. Res. Commun., 28: 604-610.

Touan, Y. T., 1959.—Study of soil algae. III. Bioassay of soil fertility by algae. Plant and
Soil, 10: 220-232.

EXPLANATION OF PLATES
PLATE XIII
Fig. 1. Feulgen stained Chlorella NM1 cell, showing nucleus (N) and chloroplast (C).
¥ 3,350.
Fig. 2. Feulgen-fluorescent dye stained Chlorella NM1 cells examined by phase contrast
microscopy. Chloroplast (C), nucleus (N). 3,350.

Fig. 3. The same cells as in Fig. 2 examined by fluorescent microscopy. Note fluorescence
in regions identical to those identified as chloroplast (C) and nucleus (N) in Fig. 2. 3,350.

Fig. 4. Chlorella NM\. Note starch (S) in pyrenoid area (Py) and starch granules (S)
scattered in areas between lamellae (La) of chloroplast. Glut, OsO,. % 22,100.

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PLATE XII

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Proc. Linn. Soc. N.S.W., Vol. 99, Part 4 PLATE A xay

A. R. PILLAY ET AL. 201

Fig. 5. Cell printed for maximum contrast. Low electron dense areas (Lo) around the
pyrenoid (Py) and between lamellae of chloroplast. Glut. KMnO,. x 24,750. Inset A shows
DNA fibrils (f) in low density areas. Glut. KMnO,. x 60,000.

Fig. 6. Cell printed for maximum contrast. Note absence of fibrillan material in low
density areas (Lo) after deoxyribonuclease digestion. Glut. KMnO,. x 29,700.

Fig. 7. Portion of chloroplast showing pyrenoid area. Note starch sheath (S) and fibrils
(f) around pyrenoid (Py). Glut. OsO,. x 53,000.

PLATE XIV

Fig. 1. Section of Chlorella NM1 showing chloroplast. Fine fibrils (f) and ribosomes (r)
can be seen in the organelle. White arrow indicates attachment of DNA fibril to lamellae

membrane: Pyrenoid (Py), lamellae (La) and cytoplasmic ribosomes (CR). Glut. OsQ,.
x 53,000.

Fig. 2. Magnified portion of Fig. 1. Linear DNA fibrils (double arrows), single loop (L)

and double loops (Ls), lamellar membrane (La), polyribosomes (Po) are found close to DNA
fibrils. Glut. OsO,. x 105,000.

PROCEEDINGS OF THE LINNEAN SOCIETY OF NEW SoutH WALES, Vot. 99, Part 4

NOTES AND EXHIBITS

Dr. D. McAlpine exhibited specimens of Boea hygroscopia (Family
Gesneriaciae), a rain forest plant from the Atherton Tableland, Queensland,
which shows remarkable drought adaptation. The plant becomes shrivelled
and dessicated in the dry season but is totally reconstituted after a few hours’
submergence in water. The plant is closely related to the familiar African
violet.

Dr. I. V. Newman presented a note on the journey made by George Caley
in February 1805, from Pennant Hills to Narrabeen and back. Caley was
collector for Sir Joseph Banks from April 1800 to May 1810. Dr. Newman
exhibited a copy of Caley’s own map of the journey and a route plan of it, drawn
on a current road map of the area, calculated from Caley’s original description
and map (both of which are housed in the Mitchell Library, Sydney). Caley’s
near passage on this journey to an unnamed creek in Wahroonga has provided
support for a recommendation that the creek be named ‘ Caley’s Creek ”’ and
that a proposed recreational area nearby be named “‘ Caley’s Common ”’’.

Professor T. G. Vallance (President) exhibited two copies of the Society’s
first publication ; both are preserved in the library at Science House. Each
bears on its title-page: The/Linnean Society/of/New South Wales./=/Sydney :/
F. Cunninghame & Co., Steam Machine Printers, 186 Pitt Street./—/1874. Both
copies Measure 195 x 130 mm and have pp. 10 (the last blank) with the sheets
sewn, the last leaf being tipped-in. The issue with printed blue paper wrappers
agrees with that described as 11655c in Ferguson: Bibliography of Australia
(Vol. VI, Sydney, 1965); the other, with printed yellow paper wrappers, is
otherwise identical. The text lists, first, Offlcers and Council (p. 3), then, 107
names of Original Members (pp. 3-6) and, finally, the Rules (pp. 7-9) of the new
Society. Both copies have been extensively annotated and corrected by hand,
perhaps by way of preparation for the second edition, dated 1876, which is to
be found in some bound copies of the first volume of our Proceedings.

As the official records of the Society’s early years were lost in the Garden
Palace fire (22 September 1882) it does not appear possible to discover the actual
date of these first publications. Indeed, despite the 1874 imprint the pamphlets
may not have been issued until early in the following year. We know that the
committee appointed at the inaugural meeting of the Society (29 October 1874)
to prepare a draft set of Rules completed its work promptly as the Rules were
adopted on 5 November 1874. But according to J. J. Fletcher (Macleay memorial
volume, 1893, p. XX XJ) the first Council was not elected until 13 January 1875.
Dr. A. B. Walkom, however, in the Society’s Jubilee Publication (Historical
notes of its first fifty years, Sydney, 1925) does not refer to this meeting and lists
the first councillors as having been elected from 1874. Whether printed late in
1874 or early in 1875, these pamphlets survive as a previous record of the Society’s
membership in its foundation year. That the present enrolment is only about
three times greater than it was one hundred years ago is a sobering fact.

Again referring to our centenary, Professor Vallance reminded members of
the recent gift from the Linnean Society of London of a splendid engraving after
H. W. Pickersgill’s portrait of Robert Brown. The engraving is currently on
display with selected material from our archives in the galleries of the Library of

PROCEEDINGS OF THE LINNEAN Society or New SourH Wass, Vou. 99, Part 4

NOTES AND EXHIBITS 203

New South Wales. By way of celebrating this important donation as well as
the bicentenary of Robert Brown’s birth (21 December 1773), the President
commented briefly on Brown’s great contribution to Australian science begun
while naturalist aboard H.M.S. Investigator under the command of Matthew
Flinders. This fruitful association is commemorated in a set of the two-volume
work of Flinders: A voyage to Terra Australis (London, 1814) owned by the
Linnean Society of New South Wales. The half-title leaf in Volume I of this set
bears the inscription: To/Mr. John Sangster/from His Relation & Friend Robert
Brown/a member of the Voyage. Below, in pencil, is the date 1857, the year
before Robert Brown’s death. These volumes were presented to the Society in
1887 by Mr. Sangster; the donation is acknowledged in our Proceedings (Vol. 2,
1887, p. 454) where, strangely, no mention is made of the distinguished
provenance. The work was transferred recently, on extended loan, to the Rare
Books department of the University of Sydney Library; its brief return for
exhibition at this meeting has been effected with the ready cooperation of the
University Librarian.

Mr. G. P. Whitley exhibited (by permission of the Curator of the Macleay
Museum, University of Sydney) photographs of the Whistle Insect. The Whistle
Insect is the oldest known specimen in the Macleay Museum, being labelled ‘‘ A
curious insect from Barbary, the only one of its kind in England. Geo.
Edwards, 1756’.

A picture of a ‘‘ Whistle Insect”? in the anonymous WNaturalist’s pocket
magazine (Vol. 2, 1799, no pagination, plates unnumbered) was recognised as a
depiction of the ‘‘ curious insect’’ specimen in the Macleay Museum. The
picture is evidently a copy of one of three figures of the Macleay Museum specimen
found in George Edwards’s Gleanings of natural history (Vol. 2, 1758: 160,
pl. 285, figs 3-5), a rare book in the collection of the National Library, Canberra.
Edwards’s figures were names Gryllus spinulosus by Linneaus in 1763, so the
specimen, brought to Sydney by Alexander Macleay in 1825, is obviously the
holotype of that species, which has been referred to the genus Eugaster by
Kirby, 1906 (Orthoptera: Acridoidea).

PROCEEDINGS OF THE LINNEAN SOCIETY oF New SoutH WAtgS, VoL. 99, Part 4

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Letatd STACY face aa a ee a ee a en at
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Ritvias ee okay ni atbaertadye CUAL ED es aA fay pia ae m Bit ee i eae Yi:

Index

Vol. 99
Page Page

Allen, G. R., Four new damselfishes Glyphidodontops niger n.sp. ae SHON SHE
(Pomacentridae) from the southwest Glyphidodontops notialis n.sp. .. ISAT
Peo. : eae oe co eu Glyphidodontops talbott n.sp... esi

Amblyseius Berlese (Acarina : Phyto- Grewingkia neumant sp. nov. .. ws O
selidae), a new species from apple) in j :

PNaetrolia ete : eas Grewingkia parva sp.nov. aus SO
Amblyseius lentiginosus n.sp. 145 Grigg, G., see Pepperell, J... -. 54
Annual General Meeting 1 Guntheria vegrandis n.sp. .. 14
Bak EP. T C 186 Harrison, F. W., see Racek, A. wn yn) lltas7/

aaa DeSa mindue ae Holmophyllia maculosa sp.nov. . . .. 186

Balance Sheets Soho F

Holmophyllum struszi sp.nov. .. .. 184

Ballantyne, B., Oder: fader on : :
Cucurbitaceae : identity, distribu- Labyrinthodont trackway from the Mid-
tion, host range and sources of geste ONSEN BONE New South a
resistance .. 100 EINEIS) 91 :

Biogenic structures in che Ordouicun Leech, land., WENT troglobitic
Bowan Park eae: New South eae - fon aoe ae
Wales .. ee _. 129 irudinoidea : Haemadipsidae) ..

Branchiostegus serratus n.sp. 151 Leiobdella jawarerensis gen.et sp.nov .. 57

; : Linnean Macleay Fellowship .. st 2

Calostylis panuarensis sp.nov. oO TEA TMI AEG UcGabe nner MInees

Cho, K. Y., see Pillay, A. R. S97 a obrs as eo ies

Corals, rugose ti : 36, 181 McLean, R. A., The rugose dorad genera

Cucurbitaceae, powdery mildew on 100 Streptelasma Hall, Grewingkia

Dybowski and Calostylis Lindstr6m
eae eee) four new, Fe BGO ar Grn SulinrianulG@n ING

rom the southwest Hack Cc 87 South wales. er an 36
‘Dioamngas, LL re Sunol 1B (Shelelatay) eV Oy) McLean, R. A., Silurian rugose corals

Se CLES i. Amblysevus Berlese from the Mumbil area, central New

(Acarina : Phytoseiidae) from

South Wales.. ae pe eed
epee ueele ue Uidkay, Sie (Alon, cco Sie Wilko

Denmark, H. A., and E. eoniena A how " Macias. Tass a 69
species of Phytoseius Ribaga is
(Acarina : Phytoseiidae) ae apple Mildew, powdery, on Gucarbitatees en LOO
in Australia .. 177 Mites hs 15, 79, 145, 177

DNA in chloroplasts of a (Chlorella sp. 197 Moore, K. M., The Glycaspis spp.

Domrow, R., Miscellaneous mites foi (Homoptera : Psyllidae) associated
Wusiralian’ vertebrates. 1-48 15 with Hucalyptus camaldulensis .. 121

Dooley, J. K., and J. R. Paxton, A new Mutagenic effects of ethyl methane-
species of tilefish (Family Branchio- sulphonate on the oat stem rust
stegidae) from eastern Australia .. 151 pathogen (Puceinia graminis f.sp.

Dosse, G., see Schicha, E 79 ee) ed a echoes

tad ae Notes and Exhibits as a .. 202

Elections 29 oe 6 Nucleic acids in chloroplasts of a

Ethyl methanesulphonate, mutagenic Chlorella sp. .. 6 so ae NN
effects on the oat stem rust pathogen Oat stem rust pathogen (Puccinia
(Puccinia graminis f.sp. avenae) 166 graminis f.sp. avenae), mutagenic

Eucalyptus camaldulensis, Glycaspis spp. effects of ethyl methanesulphonate 175
Cees Psyllidae) associated Palaeophyllum oakdalensis sp.nov. lO 2,
iy A ri rs) co Ll Palaeospongilla chubutensis (Porifera :

Glycaspis spp. (Homoptera: Psyllidae) Spongillidae), systematic and ee
associated with Hucalyptus camal- genetic position 20 . 157
dulensis 3 a aie 50 Ul Paxton, J. R., see Dooley, J. K. .. 161

206

INDEX

Page

Pepperell, J., and G. Grigg, A Laby-
rinthodont trackway from the Mid-
Triassic near yan New South
Wales ..

Phytoseius Foihoninahariae: n.sp..

Phytoseius Ribaga (Acarina : Phytosei
idae), a new epee: from apple in
Australia

1Eull ENE va Ri NG ALE Tenn Sue Y. Cho,
Nucleic acids in chloroplasts of a
Chlorella sp. ..

Pomacenirus albimaculus u.sp. nis
Powdery mildew, see Mildew, powdery. .
Presidential Address

Racek, A. A., and F. W. Hanon The
systematic and phylogenetic position
of Palaeospongilla chubutensis
(Porifera : Spongillidae)

Reports on the Affairs of the Society

Richardson, L. R., A new troglobitic
quadrannulate land-leech from
Papua eng mele B Haema-
dipsidae s.l.) .

Robertson, R. N., “‘ A Society of Natinal
History. I hope they may succeed.”’
The first hundred years (Sir William
Macleay Memorial Lecture, 1974)..

Rugose corals, see Corals, rugose

157

57

69

Page

Schicha, E., and G. Dosse, A new species
of Typhlodromus Scheuten (Acarina :

Phytoseiidae) from oper in
Australia ; sine
Schicha, E., see Denmark, H. A. 145,

Semeniuk, V., Biogenic structures in the
Ordovician Bowan Park Group,
New South Wales

Sir William Macleay Memorial ‘Lecture,
1974 oe

Stanbury, P. J., The role of scientific
societies—Some suggestions for the
future (Presidential address)

Streptelasma recisum sp.nov.
Tchan, Y. T., see Pillay, A. R.

Teo, C., and E. P. Baker, Mutagenic
effects of ethyl methanesulphonate
on the oat stem rust pathogen
(Puccinia graminis f.sp. avenae)

Tilefish (Family Branchiostegidae), a
new species from eastern Australia

Typhlodromus helenae n.sp.

Typhlodromus Scheuten (Acarina :
Phytoseidae), a new species from
apple in Australia

79
177

129

79

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PROCEEDINGS OF THE LINNEAN SocriETY oF NEw SoutH WALES
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Proceedings, Volume 99, Part 4, 1975

CONTENTS

Page

DENMARK, H. A., and ScuicHa, E. A new species of ee Be
(Acarina : Phytoseiidae) from apple in Australia .. 177

McLzeAN, R. A. Silurian rugose corals from the Mumbil oy central
New South Wales . Rec Be ne ns a ; ool SH

Pray, A. R., TCHAN, Y. T., and A CEOs K.Y. Nucleic acids in Cae
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Notes and Exhibits... o5 50 ate ac tas 3b way) 202

Index Ne a Bite a: BA Ne sis sie ae ne 205

Proceedings of the

Linnean Society
of New South Wales

VOLUME 100
Nos. 441-444

CONTENTS OF PROCEEDINGS, VOLUME 100

PART 1 (No. 441)
(Issued Ist August 1975)

CONTENTS
Page
Annual General Meeting :
Report on the Affairs of the Society for the Year ae ae ae 1
Balance Sheets ve a tye ae Bee oo a ai 7

VALLANCE, T. G. Presidential address. Origins of Australian geology 13

ASHTON, D. H., BonD, H., and MorRIs, G. C. oven! eee on Mount
Towrong, Victoria : : 44

SELKIRK, D. R. Tertiary fossil fungi from Kiandra New South Wales 70

STAFF, I. A. The fruits and seed productivity in Xanthorrhoea .. AAC RD

PART 2 (No. 442)
(Issued 10th November 1975)

CONTENTS
Page
GRIFFIN, D.J.G. Anew giant deep-water isopod of the genus Bathynomus
(Flabellifera : Cirolanidae) from eastern Australia a cf see Oe
KAImoLa, P. J. Notes on some fishes of the families Uranoscopidae,
Scorpaenidae, Ophichthidae and Muraenidae from Torres Strait .. 110
Sanps, V. E. The cytoevolution of the Australian Papilionaceae BA 1 Lie)

SmirHeErRs, C.N.,and THorntTON,I.W.B. The first record of Stenopsocidae
(Psoe optera) from New Guinea with descriptions of new species Sag lllaXts

PART 3 (No. 443)
(Issued 6th February 1976)

CONTENTS

AULD, B. A., and MARTIN, P. M. Morphology and distribution of Bassia
birch (F. Muell.) F. Muell. ifs ae Bk Be ;

McLEAN, R. A. Aspects of the Silurian Aes coral fauna of the Yass
region, New South Wales .. : ;

McLrEop, RK. W. Sources of resistance to Heterodera avenae Woll. in
New South Wales ..

WHITLEY, G. P., and STANBURY, P. J. Type specimens in The Macleay
Museum, University of Sydney. VII. The holotype of Gry ie
spinulosus Johansson (Insecta : Orthoptera : Tettigonioidea)

PART 4 (No. 444)
(Issued 25th March 1976)

CONTENTS

ANDERSON, D. T., WHITE, B. M., and Eean, EB. A. The larval
development and metamorphosis of the ascidians Pyura praeputialis
(Heller) and Pyura ene (Herdman) (Pleurogona, Family
Pyuridae) é us ais Bil te a ne ate

BEDFORD, G. O. Defensive behaviour of the New Guinea stick insect
Hurycantha (Phasmatodea : Phasmatidae : Eurycanthinae)

DANIELS, G. Three new species of ye te Dakin and Fordham
(Diptera: Asilidae) from Australia a

McLEAN, R. A., and WEBBY, B. D. Alen Ordovician ee corals of
central New South Wales . :

Index

Page

167

179

195

202

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