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PROCEEDINGS
OF THE
LINNEAN SOCIETY
NEW SOUTH WALES
VOLUME
106
(Nos 465-468; for 1981-82)
Sydney
The Linnean Society of New South Wales
1983
© Linnean Society of New South Wales
Contents of Proceedings
Volume 106
NUMBER 1 (No. 465)
(Issued 27th October, 1982)
JONSELL, B. Linnaeus and his two circumnavigating apostles ..............
FOSTER, B. A. Two new intertidal balanoid barnacles from eastern Australia...
MCALPINE, D. K., and KENT, D. S. Systematics of Tapezgaster (Diptera:
Heleomyzidae) with notes on biology and larval morphology ...........
SCHNEIDER, M. A. A comparative morphological study of the reproductive
systems of some species of Tapeigaster Macquart (Diptera: Heleomyzidae) . .
SHAW, D. E., CANTRELL, B. K., and HOUSTON, K. J. Neurochaeta inversa
McAlpine (Diptera: Neurochaetidae) and seed set in Alocasta macrorrhiza
(L.) G. Don (Araceae) in southeast Queensland.....................
HUTCHINGS, P. A., and RECHER, H. F. The fauna of Australian mangroves. . .
Notes and Discussion
FORD, R. J. A. W. H. Humphrey, His Majesty’s mineralogist in New South
Wales (803-12 —— Avcomment. 24. 25.2 vie eae ee oe ee se wits
NUMBER 2 (No. 466)
(Issued 27th October, 1982)
POWELL, C.McA., NEEF, G., CRANE, D., JELL, P. A., and PERCIVAL, I. G.
Significance of late Cambrian (Idamean) fossils in the Cupala Creek
Formation, northwestern New South Wales...................20056-
ANDERSON, D. T. Origins and relationships among the animal phyla ........
JENKINS, C. J. Late Pridolian graptolites from the Elmside Formation near Yass,
Ne WAS GUILE WialeSeum tian ante psec Har eee aos Gi ore elect tous ay wake Gees
JENKINS, C. J. Darriwilian (middle Ordovician) graptolites from the Monaro
Trough sequence east of Braidwood, New South Wales ...............
HUTCHINGS, P. A. The fauna of Australian seagrass beds.................
Annexure to Numbers 1 & 2. The Linnean Society of New South Wales. Record of
the Annual General Meeting 1981. Reports and balance sheets .............
123
NUMBER 3 (No. 467)
(Issued 10th May, 1983)
LARKUM, A. W. D., and WEST, R. J. Stability, depletion and restoration of
seagrass! beds oii. ned as Mates ds) ie age ins ed) aie Wc sane a ro
WEST, R. J., and LARKUM, A. W. D. Seagrass primary production — a review.
Kuo, J. Notes on the biology of Australian seagrasses ................-...
GRAY, M. R. The taxonomy of the semi-communal spiders commonly referred
to the species Ixeuticus candidus (L. Koch) with notes on the genera
Phryganoporus, Ixeuticus and Badumna (Araneae, Amaurobioidea) .........
NUMBER 4 (No. 468)
(Issued 10th May, 1983)
WEBB, L. J. Sir William Macleay Memorial Lecture 1982. Ecological values of
the tropical naimtonestasesOUnCe nen .s uate. cues aeons ira tet eer
GRAY, M. R. A new genus of spiders of the subfamily Metaltellinae (Araneae,
Amaurobioidea) from southeastern Australia.......................
CARR, P. F. A reappraisal of the stratigraphy of the upper Shoalhaven Group
and lower Illawarra Coal Measures, southern Sydney Basin, New South
Whales ak ak GSP. cee ot ie ti attra) atid at ANcs oe PSR ON eR OO GE ne Ree ae
HOLMES, W. B. K., HOLMES, F. M., and MARTIN, H. A. Fossil Eucalyptus
remains from the middle Miocene Chalk Mountain Formation,
Warrumbungle Mountains, New South Wales......................
WATSON, I. A., and SOUSA, C. N. A. de. Long distance transport of spores of
Puccinia graminis tritici in the southern hemisphere................-...-
SHAW, D. E., and CANTRELL, B. K. A study of pollination of Alocasia macrorrhiza
(L.) G. Don (Araceae) in southeast Queensland.....................
SELKIRK, P. M., COSTIN, A. B., SEPPELT, R. D., and SCOTT, J. J. Rabbits,
vegetation and erosion on Macquarie Island........................
TINE gilt 21-505, Sto Oe Ro eet Scag oop toate ie Be
263
ZU
287
299
311
323
-
=
Annexure to Proceedings Vol. 106
THE LINNEAN SOCIETY OF NEW SOUTH WALES
RECORD OF THE ANNUAL GENERAL MEETING, 1981
The one hundred and sixth Annual General Meeting was held in the Activities Room, Australian
Museum, 6-8 College Street, Sydney on Wednesday, 25th March 198] at 7.30 p.m.
The President, Dr F. W. E. Rowe, occupied the Chair. He read the minutes of the one hundred and
fifth Annual General Meeting and they were adopted by the meeting.
REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR 1980-81
Publications
The Society’s Proceedings were published on the following dates :
Vol. 104, Parts 1 & 2, July 1980
Vol. 104, Numbers 3 & 4, January 1981
We record our gratitude to Professor T. G. Vallance for his continuing service as Honorary Editor.
Membership
During the year, 9 new members were elected to full membership of the Society and 18 joined as
associate members. There were 4 resignations and 3 deaths; 5 names of unfinancial members were removed
from our membership list and 2 ordinary members became life members after 40 years membership of the
Society, making a total of 300 on the 20th March, 1981.
Council noted with regret the death of life member, Dr I. M. Mackerras who joined the Society in 1922
and was Linnean Macleay Fellow from 1925 until 1927. We also note with regret the death of Mr E. T.
Smith who had been a member since 1953 and of Mr S. L. W. Allman, also a life member, who joined in
1940.
DrJ. M. Beattie and Mr K. L. Taylor became life members this year and we record our appreciation of
their long service.
The Council is pleased to note the growth in the new category of associate membership.
Meetings
Activities during the year included 4 ordinary general meetings, 1 field excursion and 1 full day
symposium.
The first meeting was held on 11th June_at the School of Botany, University of New South Wales. Dr J.
Bowler of the Research School of Pacific Studies, A.N.U. addressed us on the topic ‘Reconstructing climates
of the past 50,000 years: evidence from salt lakes in Northern and South Australia’.
On 9th July, in the Australian Museum, Dr D. A. Adamson of Macquarie University showed slides and
discussed his research in the Antarctic during the summer of 1979-80.
On 24th September, in the Australian Museum, Dr P. Selkirk of Macquarie University spoke to us
about ‘Summer on Macquarie Island through a Biologist’s Eye’.
The biennial Macleay Memorial Lecture was delivered on 10th September by Professor H. B.
Whittington of the Sedgwick Museum in Cambridge, at the Science Centre Auditorium. His subject was
‘Evolutionary problems presented by the appearance of multi-celled animals in the Cambrian’.
On Sunday 28th September, members were escorted on a tour of the Royal Botanic Gardens by the
Director, Dr Johnson and staff members, Mr Rodd and Mr Dangerfield.
A review symposium on ‘The Sydney Basin’ attracted an audience of close to one hundred people to the
Australian Museum on Saturday 25th October to hear a panel of speakers deal with the area from various
scientific and environmental aspects.
Our last meeting for the year was held on 19th November at the Australian Museum when Dr D. A.
Adamson addressed us on ‘The biology and geomorphology of a Blue Mountain valley, the Grand Canyon,
Blackheath’.
Newsletter
The LINN Soc News, edited by Dr Helene Martin, continues to be printed quarterly. It consists of details
of our programme, reports of resolutions of the Council and other items of interest to members.
Library
The Council has for some time been concerned about the Society's problems in meeting the increasing
cost of maintaining the library.
Detailed discussions have been held with officers of the State Library, the National Herbarium, the
Australian Museum and the three universities in Sydney. These discussions established that none of these
institutions has the capacity or the finance te house and service the Society’s library as a whole, but all are
interested in receiving selected parts of it.
After extensive discussion of available options, Council decided in December to inform the State
Government, the Australian Academy of Science and the interested institutions that unless the costs of
running the library could be entirely and permanently undertaken by an outside body within six months,
the Council proposed to dispose of the collection in the following manner:
1. As public institutions, the Australian Museum and the National Herbarium should be given first
refusal of parts of the library in which they are interested.
2. Consideration should be given to donating to the Macleay Museum any Macleayana, including
bound pamphlets.
3. Remaining parts should be offered to the universities of Sydney, New South Wales and
Macquarie.
4. The remainder to be dispersed by a decision of Council.
Linnean Macleay Fellowship
Barbara Porter has completed her second and final year as Linnean Macleay Fellow at the University of
New South Wales. Her studies on the structure and function of macropodid salivary glands have disclosed
the remarkable secretory capacity of these glands, and the importance of salivary secretions in foregut
homeostasis and in electrotype balance in the kangaroos. In many respects, the histology of the glands and
the volume and composition of their secretions, resembles ruminant counterparts; the study therefore
provides further evidence for the evolutionary convergence of macropods and ruminants with respect to
digestive physiology.
Miss Porter submitted her Ph. D. thesis in March 1981.
Linnean Macleay Lectureship in Microbiology
The work this year was concerned mainly with the physiology and cultivation of a new type of oyster
mushroom, Pleuwrotus sajor-caju. The mushroom has many advantages over Agaricus bisporus, the only
mushroom presently grown in Australia. Apart from the high yield, the oyster mushroom can utilize a wide
range of agricultural wastes and grow at a wide range of temperatures.
The work is supported by a grant awarded by the Rural Credit Development. Dr K. Y. Cho continues
to be the Linnean Macleay Lecturer.
Sczence Centre Report
At the end of 1980, Mr E. J. Selby resigned as one of the Society’s representatives on the board of
Science House Pty Ltd. Being experienced in business affairs, Mr Selby has given invaluable service and the
Society is indebted to him. His place has been filled by Mr Harry Wallace, a director of F. T. Wimble with a
background of successful business experience. The Society is currently represented on the board of Science
House by Mr Harry Wallace, Dr Lyn Moffat, Dr Alex Ritchie and Dr Helene Martin.
The Science Centre is now fully let so income from rents has reached a plateau. Hirings of the rooms
are about 42% of capacity. The secretarial service operates well and is near the limit for the present staff.
The activities in conference organizing have shown a spectacular increase — conferences are organized for
the Science Centre and elsewhere, e.g. Wollongong, Broken Hill and overseas.
As a trading operation, all of these avenues work well and the income over expenditure is quite
satisfactory. However, it in no way services the mortgage of $2.6 million and there is no hope of Science
House Pty Ltd ever trading itself out of this debt.
A foundation is being created for tax-deductible fund-raising purposes with Mr Harry Wallace as
chairman of the fund-raising committee.
FINANCIAL REPORT
Report of the Honorary Treasurer (Dr A. Ritchie), presented to the Annual General Meeting with the
audited balance sheets.
Despite increased costs for goods and services, the Society closes the year 1980 with a small surplus. The
amount ($2678) now transferred to accumulated funds is less than the surplus for 1979 ($3648) but
compares favourably with the $2224 in 1978.
Invested funds in the General Account stand at $91434 (down $1782 on 1979, but $2000 redeemed
from the MSW&DB awaits reinvestment) and yielded interest of $10952 in 1980, $634 more than in 1979.
The remainder of our investments (totalling almost $420,000) is tied in loans to Science House Pty Ltd
- which are unlikely to yield a return in the foreseeable future.
Total expenditure for the year 1980 was $30,497, substantially more than in 1979 ($26,392). Among
items contributing to the increase were: higher audit fees, a grant to the Linnean Macleay Fellow for
assistance with research equipment, higher postal costs and greater outlay on Library (a number of
subscriptions had been held over or delayed from 1979). Again, following a necessary review by Council, the
emoluments of the Secretary and Librarian were increased in acknowledgement of present inflation. The
cost of printing the Proceedzngs has also risen but members surely will appreciate the improved quality of
the journal and be aware that all editorial work is done by Prof. Vallance at no cost to the Society. Economy
in the use of the secretarial services of Science House Pty Ltd resulted in our paying only $7903 in 1980, as
against $8097 in the previous year.
The total income of $33,175 for 1980 derived chiefly from subscriptions, interest on General Account
investments, sales of back issues of the Proceedings and, importantly, from the transferred surplus from the
Fellowships Account (see below).
In the Fellowships Account, total investments of $128,196 yielded interest of $13,414 in 1980 ($11,620
in 1979).
The maximum amount ($3200) permitted by the Equity Court was paid to our Fellow during
1980 and the surplus ($10214) transferred to the General Account.
The Bacteriology Account, with investments totalling $36,921 earned interest of $3639 in 1980. The
sum of $4000 paid during the year to the University of Sydney as contribution to the salary of the Linnean
Macleay Lecturer in Microbiology left a deficit of $361 but this is covered by a previous surplus.
Total investments in the Scientific Research Fund (to be known henceforth in honour of its principal
benefactor, the Joyce Vickery Scientific Research Fund) by the end of 1980 stood at $49,890, a considerable
increase on the 1979 figure and due largely to further releases from the late Dr Vickery’s estate. Interest
received during 1980 amounted to $4216; increased income in future is anticipated as the investments start
to yield over a full year. Accordingly, Council has resolved that from now on a certain proportion of the
annual income will be made available as grants for appropriate scientific research. It plans soon to
announce conditions of eligibility for the grants and to invite applications for them during 1981 by a date to
be fixed, and regularly thereafter. Council, however, will reserve the right and capacity to receive and grant
applications for emergency assistance at any time, One such grant (of $300) from the Joyce Vickery
Scientific Research Fund has already been made — to enable Dr S. K. Roy of India to complete a research
project at the Royal Botanic Gardens in Sydney. The Council hopes others will emulate Dr Vickery’s
generosity by donating to the Fund; by so doing they will enhance the capacity of the Linnean Society of
New South Wales to foster the cause of science. Donations to the Fund are tax-deductible.
Following discussion of the Honorary Treasurer’s report, a motion that the audited balance sheets for
1980 and the financial report be adopted was carried unanimously by the members present.
PRESIDENTIAL ADDRESS
Dr Rowe delivered the Presidential Address entitled ‘A Brief Review of Australian Echinoderms since
H. L. Clark (1946)’. This address will be published in full in the Proceedings.
DECLARATION OF ELECTIONS
As there were five nominations to fill six vacancies on Council and no other nominations for President
or Councillors were received, no election was necessary. The President declared the following members
elected: Dr L. A. S. Johnson, Dr Lynette A. Moffat, Mr G. R. Phipps, Dr F. W. E. Rowe, Professor T. G.
Vallance.
Our Auditors continue to be W. Sinclair & Co.
Dr Rowe introduced Dr Helene A. Martin as the President for 1981-82 and invited her to take the
Chair.
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NEW SOUTH WALES
VOLUME 106
NUMBERS 1 &2
NATURAL HISTORY IN ALL ITS BRANCHES
THE LINNEAN SOCIETY OF
NEW SOUTH WALES
Founded 1874. Incorporated 1884.
The Society exists to promote ‘the Cultivation and Study
of the Science of Natural History in all its Branches’. It
holds meetings and field excursions, offers annually a
Linnean Macleay Fellowship for research, contributes to
the stipend of the Linnean Macleay Lecturer in Micro-
biology at the University of Sydney, and publishes the
Proceedings. Meetings include that for the Sir William
Macleay Memorial Lecture, delivered biennially by a
person eminent in some branch of Natural Science. The
Society’s extensive library is housed at the Science Centre
in Sydney.
Membership enquiries should be addressed in the first
instance to the Secretary. Candidates for election to the
Society must be recommended by two members. The
present annual subscription is $20.00.
The current rate of subscription to the Proceedings for non-members is set at $35.00 per volume.
Back issues of all but a few volumes and parts of the Proceedings are available for purchase. A price list will
be supplied on application to the Secretary.
OFFICERS AND COUNCIL 1981-82
President: HELENE A. MARTIN
Vice-Presidents: LYNETTE A. MOFFAT", A. RITCHIE, F. W. E. ROWE,
J. T. WATERHOUSE
Honorary Treasurer: A. RITCHIE
Secretary: BARBARA STODDARD .
Council: D. A. ADAMSON, M. ARCHER, L. W. C. FILEWOOD, L. A. S.
JOHNSON, HELENE A. MARTIN, P. M. MARTIN, LYNETTE A.
MOFFAT", P. MYERSCOUGH, G. PHIPPS, A. RITCHIE, A. N. RODD, F. W.
E. ROWE, C. N. SMITHERS, T. G. VALLANCE, J. T. WATERHOUSE, B.
D. WEBBY, A.J. T. WRIGHT
Honorary Editor: T. G. VALLANCE — Department of Geology & Geophysics,
University of Sydney, Australia, 2006.
Librarian: PAULINE G. MILLS
Linnean Macleay Fellow:
Linnean Macleay Lecturer in Microbiology: K.-Y. CHO
Auditors: W. SINCLAIR & Co.
eh
The office of the Society is in the Science Centre, 35-43 Clarence Street, Sydney,
N.S.W., Australia, 2000. Telephone (02) 290 1612.
©Linnean Society of New South Wales .
' resigned 20 May 1981
Cover motif: Forewing of Permorapisma bisertalis (Neuroptera:
Permithonidae), from the Permian of Belmont, f
New South Wales 4
Adapted by Len Hay from Proc. Linn. Soc. N.S.W.
51, 1926, p. 278 (Fig. 15)
PROCEEDINGS
of the
LINNEAN
SOCIETY
NEW SOUTH WALES
VOLUME 106
NUMBER 1
}
’
ea
bite
Linnaeus and his two
Circumnavigating Apostles
BENGT JONSELL
Uppsala Universitet
Institutionen for Systematisk Botantk
Box 541 75121 UPPSALA Sweden
A Lecture
delivered under the joint auspices of The Linnean Society of London, The Linnean
Society of New South Wales and the Svenska Linnésallskapet in the Macleay Lecture
Theatre, University of Sydney, on 26 August 1981, during the 13th International
Botanical Congress
If you visit Linnaeus’s house in Uppsala and ascend to the floor where his study
remains with inost of the furniture intact, you will see on a high glass cupboard a globe
of the world made in Akerman’s famous workshop in Uppsala in the middle of the
18th century (Fig. 1). The globe shows with surprising accuracy the continents in
broad outline but we can imagine that Linnaeus looked with curiosity and fascination
at the arbitrary, dotted contour that runs along the eastern side of Nova Hollandia up
to Nova Guinea (Fig. 2), across the strait said to have been kept secret by the
Spaniards. Long before Linnaeus knew of the plans for Captain Cook’s first voyage to
the Pacific he may well have wondered about this area on his globe and what such a
remote region might yield in the way of living forms. As it turned out, two of his pupils
— Daniel Solander and Anders Sparrman — were to sail with Cook and by so doing
helped unravel some of nature’s secrets on the other side of the world.
At the time of Cook’s voyages, as we approach Linnaeus to consider his relations
with and expectations of these widest-ranging of his pupils the great teacher is in his
fifties, already prematurely aged (Fig. 3). His great botanical contributions — those
beginning with the explosion of fundamental works during the years 1735-38 in
Holland and culminating in the Speczes Plantarum of-1753 — are finished. But
although his principal new discoveries had been published Linnaeus still maintained
industrious botanical work. He received innumerable additions to his garden and
herbarium and wrote new and essentially augmented editions of Systema Naturae,
Genera Plantarum and Species Plantarum, undertakings that later in his life were
taken over by compilers abroad.
Linnaeus’s garden in Uppsala (Fig. 4) continued to flourish in the early 1760s,
twenty years after its energetic restoration following his appointment in 1741 as one of
the two professors of medicine at Uppsala University. After its first glory in the days of
Olof Rudbeck the elder, the grand old man of Swedish 17th century science, the
garden had long been neglected when Linnaeus took it in hand.
Soon after 1760, however, Linnaeus turned his attention increasingly to his new
estate Hammarby in the countryside southeast of Uppsala. The collections were
housed there in a special small museum building and gardening was more successful
than in the flat, moist plots in the town. His family by this time was fairly grown up.
The four daughters were still unmarried and the only surviving son Carl was now in
charge as demonstrator in the Uppsala garden, an arrangement that had drawn much
criticism. Linnaeus himself did not leave Uppsala any more, except for meetings of the
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
2 LINNAEUS AND HIS CIRCUMNAVIGATING APOSTLES
Fig. 1. The globe in Linnaeus’s study (from a colour photo by O. Lindman).
Fig. 2. Detail of the globe (Fig. 1) showing Nova Hollandia and surroundings (from a colour photo by O.
Lindman).
Academy of Science in Stockholm and visits to the Royal family and castles near the
capital. That, very briefly, is the pattern of those years.
By their journeys his pupils, the apostles, have begun to cover the world and
gather its ‘spoils’ (Fig. 5). Linnaeus’s knowledge of the world and of its botany in
particular increased substantially as a result of the records and specimens, dried and
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
B. JONSELL 3
©
Fig. 3. Linnaeus at the age of 40. From a pencil drawing by J. E. Rehn, probably made 1747.
HoR TI! UPSALIENSIS PROSPECTUS
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Fig. 4. The Botanical Garden in Uppsala in the 1740s. From the dissertation Hortus upsaliensis (1745).
Proc. Linn. Soc. N.S.W. 106 (1), (1981) 1982
4 LINNAEUS AND HIS CIRCUMNAVIGATING APOSTLES
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Fig. 5. The voyages of Linnaeus’s apostles. Map compiled by R. E. Fries (Fries, 1951).
living, brought back by the travellers. Yet along with the new-found facts an
enthusiastic credulity persisted. Linnaeus could still accept that the tropical and
southern regions of the world were populated by strange ‘human cousins’. There was,
for instance, the Homo caudatus — the tail man — from the Nicobar islands in the
Gulf of Bengal, and the Homo troglodytes the white nocturnal species, blind in
daylight, wild and thievish, reported from ‘Ethiopia’ (i.e. tropical Africa), Java and
other places (Fig. 6) ; both species appear in Systema Naturae of 1758.
Linnaeus’s view of the southern lands was based largely on knowledge of
southernmost Africa, the rest south of the tropic of Capricorn being virtually unknown
to him. In the thesis Flora Capenszs of 1759 details on the flora are mixed with remarks
on a strange assemblage of beings between ape and man. ‘Semper aliquid novi ex
Africa was a much used proverb among the old Romans, and this is still true in our
days’ he says in the introduction to the thesis and goes on to list troglodytes, sathyres,
sylvanes, sphinxes, dianae, hamadryades and cynomolges as well as the hottentot —
the most awkward of humans and the most remote of all to be included in the species
Homo sapiens. ‘Africa monstrifera’ he calls this place in Plantae Rarzores Africanae
(Linnaeus, 1760), the end of the world where all miracles of nature seem to have been
concentrated. One might even discern here his disappointment at not having seen this
world. Sparrman tells us there was nothing Linnaeus regretted more in his life than
declining the voyage to South Africa offered him during his years in Holland.
What I have said makes it easy to understand Linnaeus’s great expectations when
he heard from London of the plans for a scientific voyage to the South Seas, to those
very areas still empty on the globe in his study. The great news arrived in a letter of 19
August 1768 from John Ellis, the London merchant and outstanding student of
corallines and other fields of natural history and Linnaeus’s most faithful and intimate
correspondent in Britain. ‘I must inform you’, Ellis begins, ‘that Joseph Banks Esquire,
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
B. JONSELL 5
AN THROPOMORPH
Fig. 6. The ‘human cousins’, as illustrated in the dissertation Anthropomorpha (1760).
a gentleman of £6000 per annum estate, has prevailed on your pupil Dr Solander, to
accompany him in the ship that carries the English astronomers to the newly
discovered country in the South sea, Lat. about 20° South, and Long. between 130°
and 150° West from London. .. . They are to proceed . . . on further discoveries of
the great Southern continent. . . . No people ever went to sea better fitted out for the
purpose of Natural History, nor more elegantly. . . . In short, Solander assured me
this expedition would cost Mr Banks 10 000 pounds’. And as an act of devotion to
Linnaeus Ellis added: ‘All this thing is owing to you and your writings’. Ellis had taken
leave of Banks and Solander three days before writing the letter.
Linnaeus, always sensible, became enthusiastic on receiving the news. The
participation of Solander, one of his most intimate pupils, seemed a guarantee that
Linnaeus would have a share of the collections. Solander wrote from Rio de Janeiro,
where the expedition had landed three months after leaving Britain, explaining that if
‘Mr Banks will be in the same spirit as now to complete studies in Natural History, we
will together make the voyage to Sweden to ask prof. Linnaeus to order our recruits’.
The last was an expression borrowed from Linnaeus himself. This makes it
appropriate to consider how relations between Linnaeus and Solander had evolved so
far.
Solander (Figs 7, 8), whose origin lay in the north of Sweden, arrived in Uppsala
in 1750. Like many before him, he went with quite other academic intentions but
became fascinated by Linnaeus’s teaching in medicine and natural history. Within a
few years he was a favourite of Linnaeus. “The wittiest pupil I ever had’, Linnaeus
wrote many years later and, on another occasion, ‘I have housed him under my roof,
just like a son’. Indeed, Solander in those days was practically one of the family. He
accompanied Linnaeus to the Royal castles and collections and joined the family on
visits to Dalecarlia, where Linnaeus’s father-in-law lived. He was even considered
seriously as husband for Linnaeus’s eldest daughter. Through his close association with
Linnaeus Solander’s botanical expertise became impressive. He also gained experience
in the field travelling, as the young Linnaeus had done more than twenty years earlier,
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
6 LINNAEUS AND HIS CIRCUMNAVIGATING APOSTLES
wee Hie C. Hite
Fig. 7. Daniel Solander (1733-1782). After the painting by J. Zoffany, now in the possession of the Linnean
Society of London.
in the mountains of Lapland, probably during two summers. Several of Solander’s
collections from those journeys are preserved in Stockholm (Fig. 9).
Linnaean pupils began to be sought after in the learned world. The British
wanted a man to teach them Linnaean method and so when Linnaeus thought
Solander mature enough for a trip abroad to enlarge his education — the sort of thing
almost obligatory for learned young Swedes in those days — he proposed for him a
year or two in Britain. This was surely a sign of great trust in the 26-year-old Solander,
whom Linnaeus now apparently hoped to groom as his own successor. Solander was
urged to return after not too long a sojourn.
As we know Solander never returned to Sweden. He was immediately and
cordially received by Ellis and by Peter Collinson, another merchant and student of
natural history, and other Londoners. Solander made friends everywhere — such was
his amiable nature — and soon became a well-known figure in society and scientific
circles (Fig. 8). “Throw him where you will — he swims’ were Boswell’s words about
Solander. His combination of modesty and wit, his skill in conversation made a
general impression, as so many London reports of those days acknowledge.
Linnaeus long persisted with his own plans for Solander, counting on his return.
When he transmitted to Solander an offer of the chair in natural history at the
Academy of Science in St Petersburg Linnaeus may well have regarded some years’
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
B. JONSELL 7
Zhe SIMPLING MACARONI.
Lhe Slant 4 ‘cove oom 2 froxen LoVe. J Bea :
On thellou- Parkes grows Lf SO ia generat
4
Pal accor lo BL by Marly Strered aby (35 772
Fug. 8. Caricature of Solander, published in London after the return of the Endeavour. After the etching
dated 1772.
service in the Russian capital as a way of securing Solander’s succession to the chair at
Uppsala. But Solander’s English friends were appalled at the prospect. Collinson
warned that tumults and riots, perhaps even a revolution, might occur in Russia,
expressing also more rational arguments about the scientific and commercial isolation
of a place where Solander would be buried in obscurity. ‘No doubt you . . . know
persons less eminent but every way qualified in botanic science to teach Russian bears’
Collinson protested to Linnaeus in November 1762. The English had no wish to lose
the young man who had won their favour.
Eventually, but only after repeated pleas from Linnaeus, Solander replied that he
would have accepted the offer with uplifted hands, but on certain conditions.
Thereafter his direct correspondence with Linnaeus seems to have ceased until the
messages from the Endeavour. In 1763 Solander obtained a post at the British
Museum. The Swedish scholar Arvid Uggla has concluded that a breach occurred
between master and pupil, and for two reasons: first, following a particular grace of
the Swedish parliament Linnaeus had his own son nominated as his successor, and
second, his eldest daughter had married an officer, Captain Bergencrantz — soon to
be a most unhappy marriage. Solander could no longer feel like a son in Linnaeus’s
household and was more than content to remain in England.
Solander probably made his first acquaintance with finds from the South Seas in
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
LINNAEUS AND HIS CIRCUMNAVIGATING APOSTLES
ete IV Wit Ait
Rees
ey
nes
Gentianella aurea (1) J. M. Gullerr
JM. Giller
Ge #} Le cl? 2. of
f ee ee, ae
NOBEL.
ee ay hee at ye
LOR Ca. Foto parte
Fig. 9. Gentiana aurea L., collected by Solander at the coast of northern Norway in 1753. Specimen
preserved at the Museum of Natural History, Stockholm.
1765, in a collection of ‘artifacts’ given to the British Museum by Commodore Byron
after his circumnavigation on the Dolphin. About a year earlier Solander had met
Joseph Banks and they had already planned.to make a study trip to Linnaeus in
Uppsala when Banks’s decision to join the Endeavour expedition altered everything.
Banks made Solander enthusiastic for the voyage with a result that the latter soon
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
B. JONSELL 9
asked: “Would you like a fellow-traveller?’. ‘Someone like you would give me untold
treasures and rewards’ was Banks’s reply, and so the business was settled.
I shall not here describe the course of this voyage, many times told and doubtless
well-known in this part of the world. Let us only recall the days 28 April-5 May 1770
when, as William Stearn has said, rarely indeed can so many new and remarkable
plants have been collected in so short a time — enthusiastic collecting that made Cook
find Botany Bay the most appropriate name for the place. May I also remind you of
Solander’s curious comparison of the termite mounds of the Endeavour River area
with the rune stones of the Uppsala plain, perhaps an understandably nostalgic
reaction for a person who for ten years had not seen his native country and now found
himself at the antipodes.
Let us now turn back to Linnaeus in Uppsala and his eagerness for news from the
expedition. Immediately after the return of the Endeavour the faithful Ellis informed
Linnaeus that it came laden with the greatest treasures of natural history ever brought
into any country at one time by two people. Linnaeus replied within the hour that he
had never received a more welcome letter, adding: ‘If I were not bound here by 64
years of age, and a worn out body, I would this very day set out for London to see this
great hero in botany’. He drew a characteristically bold parallel: ‘Moses was not
permitted into Palestine, but only to view it from a distance: so I conceive an idea in
my mind of the acquisitions and treasures of those who have visited every part of the
globe’.
But Ellis also wrote in the same letter to Linnaeus, four days after Endeavour’s
return: ‘as to their Natural History I fear I shall not live to see it. They have sufficient
for one thousand folio plates’. Unfortunately these were prophetic words.
Besides Ellis, Linnaeus also obtained information about the expedition and of
Solander’s activities from two of his pupils, Anders Berlin and Henric Gahn, then in
London, but heard nothing from Solander himself. Berlin, soon to meet his destiny in
Guinea, estimated that 1200 new plant species and 100 new genera had been taken
home. Gahn, later a well-known physician in Stockholm (Gahnza (Cyperaceae) was
named by Forster in his honour), arrived in London just in time to see the collections
and pass on to Linnaeus glimpses, for instance, on the bread fruit tree and about
rubber that could be used to erase pencil writing. He presumed Linnaeus would get
part of the results but already feared for delays in publication.
Ten months after his return Banks signed a letter, drafted by Solander, for
Linnaeus, very polite with many excuses and explanations why the specimens reserved
for him had not been despatched. But already Linnaeus had written in alarm to Ellis
after learning from Berlin that a new voyage to the South Seas was being planned with
Banks and Solander of the company. The report almost deprived Linnaeus of sleep.
He foresaw the collections of the first expedition being put aside untouched, thrust in
some corner to become perhaps the prey of insects and risk destruction. His attitude to
life, so often dark in those years, is revealed in the words: ‘I shall be only more and
more confirmed in my opinion that the Fates are ever adverse to the greatest
undertakings of mankind’. He mentions bitterly that Solander in his letter from Rio de
Janeiro had promised to visit Uppsala and it is touching to read of his disappointed
hopes for such an occasion: ‘If he had brought some specimens with him I could at
once have told him what were new; and we might have turned over books together,
and he might have been informed or satisfied upon subjects, which after my death will
not be so easily explained’.
As we know Banks and Solander never joined Cook’s second expedition — they
and their equipment could not be accommodated as Banks wished. The ships left
Britain with Johann Reinhold and Georg Forster, father and son, as naturalists but at
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
10 LINNAEUS AND HIS CIRCUMNAVIGATING APOSTLES
Fig. 10. Anders Sparrman (1749-1820). Engraving after a drawing by M. Mollard.
Cape Town they were joined by Anders Sparrman (Fig. 10). This pupil of Linnaeus,
16 years younger than Solander but already a far-ranging, observant traveller, is not as
generally known as for instance Solander or Thunberg. Sparrman was brave and full
of confidence. When only 17-years-old he had travelled to Canton as physician on a
Swedish East India ship on which his services, luckily, are said not to have been much
needed. His connection with the famous Swedish East India Company captain Carl
Gustaf Ekeberg, who owned an estate in Sparrman’s home parish, helped in this and
the following voyage. Five years later he got the opportunity to go to South Africa as
tutor to a Dutch family at False Bay in the Cape Province. ‘There after seven months
he met the Forsters and within a few days he was on board the Resolutzon for a 28-
month trip in the South Seas (Fig. 11)
Sparrman was a true Linnaean in his curiosity and devotion to his master whose
words were law for him and eagerly defended. But he was an unusual Linnaean in his
disregard for systematic method and description, at least in published work. His
journeys in virgin South Africa, before and after his voyage with Cook, are vividly
related with a highly-developed sense for nature and people, in marked contrast to
Thunberg’s contemporary, exact and thorough narrative. Sparrman must have been
an amiable and sociable person. He expresses delight in plant hunting with a fellow
Linnaean (Thunberg) on Table Mountain, a venture about which Thunberg has not
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
B. JONSELL 11
XRTA she SODR 6
med de emyaste
er * Vi ceetras:
Utgitven at ef
|
htt
Fig. 11. Map of the South Pacific, from Sparrman’s Voyage, part 2.
a word to tell. Sparrman has only good to say of the Forsters who often came into
conflict with other people, not least with Banks and Solander after Cook’s return to
Britain. There is a touch of credulity and naivety in Sparrman, revealed, for instance,
in his diary by a certain parading of his cleverness. An amusing example comes from
the turning of Resolutzon at its southernmost position among the icebergs (Fig. 12).
On that occasion Sparrman hurried to the stern to claim he had been further south
than anyone else in the world. That was at Lat. 71° 10’, a record not broken until 1823
when James Weddell reached a more southerly point.
Sparrman’s diary is of outstanding value, both readable and informative, but the
section from the South Seas is generally regarded as somewhat inferior to that relating
his years in South Africa. The South Seas part was completed only decades after
Sparrman’s return to Sweden when impressions must have faded and Georg Forster’s
Voyage round the World had aptly told the story. As a scientist Sparrman was
principally a zoologist. His observations in Africa show that but in the South Seas he is
first of all an ethnographer, describing with accuracy and ingenuity the inhabitants of
one island after another. He collected quivers and war clubs, pillows and diadems,
altogether about 60 specimens that were presented to the Academy of Science in
Stockholm where they can now be seen at the Ethnographical Museum. He was also a
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
12 LINNAEUS AND HIS CIRCUMNAVIGATING APOSTLES
Sie Pee annbadl CE eR
=
Sy jf 2
C Mita Sy: Berg, Avarvid ba apitain ¢ oe co Skgpd J os ef: Matin » ined frrfhe vatten .
JSrd Luakted OF ok Ofler om Oe. -dallopps< Uda #28 ;
Fig. 12. The icebergs as illustrated by W. Hodges in Sparrman’s Voyage.
competent, interested botanist who collected quite a lot, now mostly in Stockholm. We
note only one example of interest: a gentian from Dusky Bay (Fig. 13), the first place
reached by Sparrman in New Zealand, where the virgin temperate forest covered the
mountains from top to sea — gentian from the antipodes of that gathered by Solander
on the coast of Norway.
The botanical achievements of Cook’s second voyage have generally been
attributed to the Forsters, to such an extent that in an authoritative work like Merrill’s
The Botany of Cook’s Voyages Sparrman’s contribution passes unnoticed. His role
may never be completely assessed but from diary notes it seems clear the three
naturalists formed a team. Johann Reinhold Forster, the father, bears witness to
Sparrman’s contribution in the preface to his Characteres Generum Plantarum:
‘Sparrman described the plants and my son depicted them. I devoted my whole time to
zoological descriptions. But when Sparrman had more carefully examined the plants,
my son and I were often summoned for advice and we discussed together. After that
my son compiled the described plants in another volume. I revised it all before the
descriptions were again transferred to another volume according to the Linnaean
system. And while Sparrman with my son were so occupied, I gathered again new
plants and other wealths of Nature so that we should not leave any place with empty
hands’.
It is significant that Sparrman stands in the background, he never achieved a
central position like that gained by both Solander and Thunberg. Eventually at home
he was for many years in charge of the collections of the Academy of Science, where his
own ethnographica seem to have been the most spectacular items; the naturalza are
said to have been in more or less disorder. Towards the end of the century Sparrman
was forced to leave this post and lived his last twenty years as a physician among the
lower classes in Stockholm. The appearance, only two years before his death in 1820,
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
B. JONSELL 13
(OOK One OL
Fig. 13. Gentiana saxosa Forst., collected by Sparrman at Dusky Bay, southern New Zealand, in 1773.
Specimen preserved at the Museum of Natural History, Stockholm.
of the final part of Sparrman’s narrative from the South Seas marks the last time a
voice from the Linnaean era was heard.
With the return to Britain of the Resolution in 1775 a new important load of
plants from the South Seas was added to that brought back earlier by Banks and
Solander. In London Solander now worked as a ‘registrator of the world’s botany’, in a
way like Linnaeus himself had done 30 years before. But still he made no
communication with his old teacher who repeatedly complained that ‘Solander, who
may fill hundreds of letters with novelties’ kept silent. “The ungrateful Solander does
not send one herb or insect of all that he collected in Insulis australibus novis’
Linnaeus wrote in one of his five autobiographies. Solander certainly was hard at work
and that ‘sedentary and luxurious life’ which has been ascribed to him was more a
misinterpretation of his seemingly-unhurried manner. He prepared in manuscript
many island floras — for Madeira, Tierra de Fuego, Tahiti and New Zealand — with
numerous descriptions arranged for the printers (Fig. 14). Others, among them the
Australian flora, did not reach that stage. A substantial number of engravings were
completed from sketches and paintings (Fig. 15). In a letter to the younger Linnaeus
in 1778, the father having died in January of that year, Banks told that 550 plates were
then engraved but it would not be possible to include them in a work to be published
in the course of the year.
Having received nothing from Banks and Solander, Linnaeus the elder promptly
got in touch with the Forsters as soon as he learned of their participation in the second
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
14 LINNAEUS AND HIS CIRCUMNAVIGATING APOSTLES
InsuleOcPacifi
onect (AYERS Perea, ES Awe FUE LE Cay rer on UTS
ole. LY, Laws cathe Bes Se ela ys
Poca wla: Pee hi wer ca Cf PFE A epee. cerrak'~
eee Parr caepticcer fathece coma hirer) a ea
CEPA S fee Valeamunien ee bane nl ee Pen as
Sepxtceins wd p aaa,
operatic Led. We 1 Linn Cp. ph NAL2Z A
Vy art eke Finn
vcs HEXANDRIA
pontap hy le Op ee ann Se pL NACD 1. Prrvcr » 462
oe Kala Ce Chesed . Frrnl. 63 7.2
CLheecrre queen gee fob ; NO oe
Che - blr eden i beed
unt ts 3
Pals. bor gr Lawler e ‘ rr, cmerrre ad -
Bubb. aecMnes 2 ‘
Lice 2 ner tgarren ah vebherrmapher dhe vefosler
foie prec Leore co a
bok. Mia, ovata, comcard, an havin att vd;
PA
FED Pes van 7 fra allerrra alicces oa wees pla cute
: : . : =
bv ere Caparica, pela Lan Qrtherce rena por-
Fog. 14. A page of the manuscript of Solander’s unpublished Flora of Tahiti, in the British Museum
(Natural History), London.
Cook voyage. Indeed, Linnaeus was to encourage them to publish their results before
Banks and Solander. And, as Linnaeus probably calculated, some collections from the
Forsters reached him through the agency of his most intimate friend, the Stockholm
physician Abraham Back. These were the only plants from the South Seas to come
under Linnaeus’s inspection. As a result the Supplementum Plantarum, the
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
B. JONSELL 15
%
Fig. 15. Banksia integrifolia L.f., collected at Botany Bay, painted by Sydney Parkinson on the Endeavour
and engraved by Miller in the 1770s. From Britten (1905).
manuscript of which was partly elaborated by the elder Linnaeus and edited and
extended by his son, includes a number of new Pacific plant genera and species. The
work provoked considerable tension between the Banks-Solander circle and Linnaeus
filtus, exacerbated by disagreement over the botanical application of Banksza, a name
the older Linnaeus in his first enthusiasm had proposed for the whole continent, Terra
Australis.
Solander, who knew he might have been in the younger Linnaeus’s chair in
Uppsala, may have had cause for resentment. But his generous nature took over when
Linnaeus ftlzus visited London; Solander cordially introduced him into the Banksian
circle. During this time, in May 1782, Solander suffered a sudden stroke at breakfast
at Banks’s house, a hard blow to Linnaeus as well as to the absent host. He died a few
days later and what had long been feared now became all too clear. Without Solander
the professional botanist, publication of the botany from the voyage could hardly be
imagined, the more so since Banks was now deeply involved with official duties.
Solander is unique among Linnaeus’s pupils in his influence on botanical science.
Yet his published works are remarkably few. Besides being a gifted systematist, as is
evident from his flora manuscripts, he was a keen observer of morphology. One
example of the latter skill is his observation of the Eucalyptus bud with its debated,
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
16 LINNAEUS AND HIS CIRCUMNAVIGATING APOSTLES
o
Fig. 16. Sparrmannia africana L.f. From Curtis's Botanzcal Magazine, plate 516 (1801).
M1874.
WA Al 3
ee
S\N
Fig. 17. Solandra grandiflora Sw. From Curtis’s Botanical Magazine, plate 1874 (1817).
deciduous cap, for which he suggested an interpretation. In particular he was
influential in spreading Linnaean thought and method in Britain, amply realizing the
hopes of those who had once begged Linnaeus to send them a pupil.
To this day only a fraction of Solander’s work has been published, the main parts
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
B. JONSELL 17
printed being the more than 700 copper engravings of Australian plants with
descriptions edited by Britten at the beginning of this century (Fig. 15) and now
followed by the magnificent editions of Captain Cook’s and Banks’s Florilegza. But the
work was never shut away. The generous access granted by Banks to Solander’s
material after his death enabled many scientists to profit by and even incorporate
many of Solander’s results in their own works. Some, like the Forsters, did so without
giving fair credit to the source while Solander was still alive, action that led to their
being ostracized by the London scientific community. Most, however, paid full credit
to the fundamental importance Solander’s work had for their own, as Gaertner in his
De Fructibus et Seminibus Plantarum or Robert Brown in Prodromus Florae Novae
Hollandiae and other works that marked the break with Linnaean classification. So
the diversity and peculiarity of the material collected and described by the influential
Linnaean Solander contributed essentially to the adoption of a non-Linnaean system.
Solander’s influence can thus be traced at various levels which ought to be further
analysed.
I will stop here with only a short retrospective epilogue. Linnaeus, Solander and
Sparrman were all brought up in provincial Swedish rectories — Linnaeus in Smaland
in the south, Solander in northern Vestrobothnia and Sparrman in the central
province of Uppland. Their fathers were clergymen, their grand- or great
grandfathers, on one side or the other, had been peasants. Their links with rural life
and knowledge were vital, and particularly obvious for Linnaeus. These three
Fig. 18. Linnaea borealis L. From a copper plate engraved for J. W. Palmstruch: Svensk Botanzk (1802-
1843) and used in Lindman: Bilder ur Nordens Flora (Stockholm, 1922).
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
18 LINNAEUS AND HIS CIRCUMNAVIGATING APOSTLES
distinguished men possessed a social and educational background typical of many of
those men of learning who made the 18th century in Sweden a brilliant period for
science.
Their names live by their achievements and also through the well-known plants
named in their honour: Sparrmannia L.f. (Fig. 16), the South African tiliaceous
genus of many greenhouses, Solandra Sw. (Fig. 17), the spectacular member of the
Solanaceae from Central and South America, now widely cultivated, and Linnaea
Gron. (Fig. 18), which Gronovius in Holland dedicated to Linnaeus and to which the
latter claimed to find a likeness in himself. Linnaeus’s personality may not seem to us
to have much in common with this shy, secluded plant of the northern coniferous
forests but it has nevertheless become a sort of symbol for Linnaeus and his era.
ACKNOWLEDGEMENTS
My thanks are due to staff members of the University Library, the Institutes of
Art Science and of Systematic Botany at Uppsala University, and of the Department of
Botany, Museum of Natural History, Stockholm, for multifarious assistance and
photographic work. The Hon. Editor of the Proceedings, Prof. Vallance, is thanked
for revising the English text.
References
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Public Library of N.S.W.
BLunT, W., and STEARN, W. T., 1973. — Captain Cook’s Florilegium. London: Lion & Unicorn Press.
BRITTEN, J., 1901-05. — Illustratzons of Australzan Plants collected in 1770 during Captain Cook’s voyage.
London: British Museum.
BroserG, G., 1975. — Homo sapiens L. Studzer ¢ Carl von Linnés naturuppfattning och manniskolara.
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author ].
ERIKSSON, G., 1969. — Botanzkens historia 7 Sverige intill adr 1800. Stockholm: Almgqvist & Wiksell.
Forster, G., 1777. — A Voyage round the World in His Britannic Majesty's sloop Resolution. 2 vols.
London: B. White, J. Robson, P. Elmsly and G. Robinson.
ForSTER, J. R., and Forster, G., 1776. — Characteres generum plantarum. London: B. White, T. Cadell
and P. Elmsly. 5
Fries, R., 1940. — Daniel Solander. K. Svenska Vet. Akad. Arsb. 38: 279-301.
———, 1951. — De linneanska apostlarnas resor. Kommentarer till en karta. Svenska Linnésallsk. Arsskr.
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GAERTNER, J., 1788-91. — De fructibus et seminibus plantarum. Stuttgart/Tubingen: Typis Academiae
Carolinae/G. H. Schramm.
Granit, R., 1978. — Banks och Solander — tva vanner i 1700-talets London: Jn Granit, R., ed., Utur
stubbotan rot (pp. 41-59). Stockholm: Norstedts. ‘
JuEL, O., 1924. — Notes on the-herbarium of Abraham Back. Svenska Linnésallsk. Arsskr. 7: 68-82.
LinDMAN, C., 1907, 1909. — A Linnean herbarium in the Natural History Museum in Stockholm, I-II.
Arkiv f. Botantk 7(3) ; 9(6).
LINDROTH, S., 1967. — Kungl. Svenska Vetenskapsakademiens historia 1739-1818. Vols 1, 2. Stockholm:
Almgvist & Wiksell.
LINNAEUS, C., 1758-59. — Systema naturae. Ed. 10. Holmiae: Impensis L. Salvii.
———, 1759. — Flora capensis (resp. C. H. Wannman). Diss. Upsaliae.
———, 1760. — Plantae africanae rariores (resp. J. Printz). Diss. Upsaliae.
———, 1760. — Anthropomorpha (resp. C. E. Hoppius) . Diss. Upsaliae.
——, 1909-12. — Bref och skrifvelser af och till Carl von Linné (Frirs, T. M., ed.), I, part 3-6.
Stockholm: Ljus.
———, 1957. — Vita Caroli Linnaez (MALMESTROM, E., and UcGLa, A., eds). Stockholm: Almgvist &
Wiksell.
Linn—E, C. von, fil., 1781. Supplementum plantarum. Braunschweig (Brunsvigae): Impensis
Orphanotrophei.
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B. JONSELL 19
MERRILL, E. D., 1954. — The botany of Cook’s voyages. Chronica Botanica 14 (5/6).
RAUSCHENBERG, R. A., 1968. — Daniel Carl Solander, naturalist on the ‘Endeavour’. Trans. Amer. phil.
Soc. n.s. 58 (8). .
SELLING, O., 1962. — Daniel Solanders naturaliekabinett och dess 6den. Svenska Linnesallsk. Arsskr. 45:
128-137.
SMITH, B., 1960. — European Vision and the South Pacific. Oxford: Oxford Univ. Press.
SmiTH, J. E., 1821. — A Selection of the Correspondence of Linnaeus and other Naturalists. 2 vols.
London: Longman, Hurst, Rees.
SGDERSTROM, J., 1939. — A. Sparrman’s ethnographical collection from James Cook’s 2nd expedition
(1772-1775). Ethnogr. Mus. Sweden, Stockholm, n.s. Publ. 6.
SPARRMAN, A., 1783, 1802, 1818. — Resa till Goda Hopps-Udden, Sodra Pol-Kretsen och Omkring
Jordklotet samt till Hottentott- och Caffer-landen Gren 1772-76. 2 vols. Stockholm: A. J.
Nordstr6ém/C. Delén.
STEARN, W. T., 1969. — A Royal Society appointment with Venus in 1769: The voyage of Cook and Banks
in the Endeavour in 1768-1771] and its botanical results. Notes Rec. Roy. Soc. London 24: 64-90.
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Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
Co (a Re eer :
SOF tale
Bae ‘ang Py
Re et eee) ae
Two new intertidal
balanoid Barnacles
from eastern Australia
BRIAN A. FOSTER
(Communicated by D. T. ANDERSON)
Foster, B. A. Two new intertidal balanoid barnacles from eastern Australia. Proc.
Linn. Soc. N.S.W. 106 (1), (1981) 1982: 21-32.
Two species of balanomorph barnacles are described from harbours and
estuaries of New South Wales where they are locally very abundant intertidally.
Elminius covertus sp. nov., is closely related to E. modestus from New Zealand, and
less so to Elminzus kingw from South America, and has long been mistaken for
Elmanius modestus which is now known reliably only in Australia from some ports.
Hexaminius popeiana gen. nov. and sp. nov. is superficially similar to Elmznzus spp.
but has 6 parietal plates. Both species are assigned to the Elmininae, a new subfamily
in the Archaeobalanidae.
Brian A. Foster, Department of Zoology, University of Auckland, Private Bag,
Auckland, New Zealand; manuscript received 21 October 1980, accepted for
publication in revised form 19 August 1981.
INTRODUCTION
Elminius modestus Darwin has .become a well-known barnacle since its
introduction and spread in Europe (see full literature citation in Newman and Ross,
1976). It has been assumed that the species was carried to Britain on ships in the 1930-
40 period from Australia and New Zealand (Bishop, 1947).
In New Zealand E. modestus is very common in harbours and estuaries where it
occurs on a wide variety of substrata in the midlittoral and shallow sublittoral zones
(Moore, 1944; Morton and Miller, 1968; Foster, 1978). It is the commonest fouling
barnacle in New Zealand harbours.
For Australia, the literature on E. modestus is perplexing to one used to the
species on New Zealand (and European) shores. The early records of Darwin (1854),
and the later account of Pope (1945) for Sydney shores, seem to be the basis for the
Australian presence and identity of the species. For Port Jackson, Pope (1945)
reported that E. modestus occurred ‘only towards the upper limit of the tidal range
which prevents it from becoming much of a nuisance on ships and small craft’. This
anomaly in the intertidal range, plus the lack of data on the distribution of the species
away from Sydney, prompted a closer look at the sheltered-shore barnacles of New
South Wales. It was concluded (Foster, 1980) that the account of Pope (1945) refers
to a different species of Elmznzus which occurs along with E. modestus and an
undescribed 6-plated balanid on harbour shores in the vicinity of Sydney. This present
paper describes both new species and considers their relationships within the
Balanomorpha as recently revised by Newman and Ross (1976).
The barnacles described in this paper are abundant where they occur. Samples
collected and studied contain at least 10, up to 50, specimens, at various stages of
development and erosion. The two species of Elminzus are difficult to identify when
the shell is extremely eroded, and close examination is required to distinguish all 3
species when they are mud encrusted.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
22 NEW BALANOID BARNACLES
SYSTEMATICS
Suborder BALANOMORPHA Pilsbry, 1916
Superfamily BALANOIDEA Leach, 1817
Family ARCHAEOBALANIDAE Newman and Ross, 1976
Balanomorph barnacles with labrum thin with pronounced medial cleft; shell wall
either solid or with pores of the non-balanine type, inter-laminate figures lacking or
simple.
Remarks: The Archaeobalanidae contain 10 extant and 4 extinct genera. The solid-
walled genera fall into two groups, those that associate with various sponge or
coelenterate habitats (Armatobalanus, Membranobalanus, Acasta and Conopea),
and those that are free-living (Chzrona, Solidobalanus, Elminius and Notobalanus) .
The former have hooks and teeth on cirri III and IV, the latter do not; the latter are
considered to be the more primitive. The various forms of tubiferous parietes as seen
in Semzbalanus and the extinct Archaeobalanus and Actinobalanus are considered to
be independent developments from the much more prevalent parietal pore condition
of the Balanidae (Newman and Ross, 1976).
I have argued before (Foster, 1978, p. 130), that the calcareous base of
balanomorph barnacles has been variously evolved in different groups, and that the
primitive condition is a membranous base. Three of the archaeobalanid genera have
membranous bases: Semzbalanus, Membranobalanus and Elminius. Elminius is the
only 4-plated genus in the archaeobalanids, but its simple cirri, its membranous base,
and the non-interlocking radii indicate a basic origin for Elmznzus. The new 6-plated
form described below also has simple cirri and a membranous base, and this
combination of characters denies its inclusion in any of the aforementioned genera. It
also has non-interlocking radii and seems closely allied to Elmznzus. Rather than
amend the diagnoses of Elmznzus or of Chzrona (which has a calcareous base) a new
genus is proposed for this undescribed species. Furthermore, this species, and those of
Elminius, seem to constitute a southern hemisphere subfamily as well defined as the
A — 1mm B —_1mm
Fig. 1. A: Elminius modestus Darwin, drawn from a New Zealand specimen; B: Elminius covertus n. sp.,
drawn from a specimen from Swansea, Lake Macquarie.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
B. A. FOSTER 23
northern hemisphere Semibalaninae. I propose the name Elmininae for this
subfamily.
Genus Elminius Leach, 1825
Balanoid barnacles with a medially cleft labrum; 4 thin, solid parietes, radii simple,
base membranous; cirri III and IV without hooks or teeth; basi-dorsal point of penis
absent.
Type species Elmanzus kingaz Gray, 1831 (subsequent designation, Pilsbry 1916).
Remarks: Elminius includes 3 extant species: E. kengz from southern South America,
E. modestus from New Zealand, Australia and Europe, and the new species described
below. The identity of E. crzstallinus Gruvel from the Azores needs confirmation; it
might be a case of introduced E. modestus (Newman and Ross, 1976) or the young 4-
plated stage of some other species. The subspecies E. modestus molluscorum Kolosvary
from Auckland, and E. modestus laevis Nilsson-Cantell from Melbourne are just two
of the forms assumed by this species when growing on various intertidal substrata
(Moore, 1944).
Elminius modestus Darwin
Figs 1A, 2
Synonymy: Elmznzus modestus Darwin, 1854:350; Guiler, 1952:20; Pope,
1966:181; Foster, 1978:95 (see for New Zealand references) ; Newman and Ross,
1976:52 (in part, viz. the European and New Zealand references) ; Foster, 1980: figs
2, 3. Not Pope, 1945:368; Underwood, 1977:23, 217.
Elminius modestus molluscorum Kolosvary, 1942 :147
Elminius modestus laevts Nilsson-Cantell, 1925 :49
Australian material examined. Personal collection: Port Jackson (The Spit), 1978;
Melbourne (coll. J. S. Buckeridge, 1976) ; Adelaide (coll. A. Gackle, 1979). Could
not be relocated in Port Jackson in 1980. Australian Museum: P21284 (Newport
Power House, Victoria, 1950), P21287 (Derwent River, Tasmania, 1950), P21288
(Sandy Bay, Hobart, Tasmania, 1951), P21291 (Port Lincoln, South Australia,
1951), P23569 (Port Phillip, Melbourne, boat fouling 1950), P23571 (Port Jackson,
Sydney, ship fouling, 1954). There are numerous lots of specimens that have been
classified under E. modestus but are referable to E. covertus (below). Description as
given by Moore (1944) and Foster (1978).
Designation of lectotype: British Museum (Natural History), one of several specimens
attached to an intertidal limpet Cellana ornata (Dillwyn), Registered No. B.M.
47.1.15.39, as arrowed in Fig. 2 and now held separately, Reg. No. B.M. 1981.274.
The limpet shell is one of four shells on a wooden slab bearing on reverse a label in
Darwin’s handwriting. These shells are endemic to New Zealand, and despite the lack
of collecting data it is assumed that the Elmznius specimens are part of Darwin’s
syntype material and originated from New Zealand. Specimens of the barnacle
Chamaesipho columna (Spengler) are included on two of the limpet shells. This
association of barnacles and limpets is common on New Zealand harbour shores.
Remarks: Darwin (1854, p. 348) stated ‘at Sydney I found E. modestus adhering to
oysters in a muddy lagoon, almost separated from the sea, and apparently very
unfavourable for cirripedes’. The question arises whether Darwin found specimens of
the other species of Elminzus described below, and whether he confounded the two
species in his description of E. modestus. Specimens now in the British Museum, some
with labels identifying them as E. modestus in Darwin’s handwriting, are indeed E.
modestus as known in New Zealand and as has been introduced into Europe. It is
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
24 NEW BALANOID BARNACLES
A B
C.
—2
d Lt tr. fue ee, ff Bt oe o hac oe
Lk.
e
Fig. 2. Elminius modestus Darwin. A: lectotype arrowed, on shell of Cellana ornata (Dillwyn). B: lectotype
and syntypes on New Zealand gastropod shells — Cellana radians (Gmelin), left, Turbo smaragda Gmelin,
bottom, and Cellana ornata (Dillwyn), top and right. C: Darwin’s own label as applied to reverse of slab
pictured in B. Depository: British Museum (Natural History) , London.
suspected not all Darwin’s material is now in the British Museum, and there is doubt
(Foster, 1980) about the labelling and the localities of some specimens he had to
study. Nor are his Sydney specimens identifiable, if indeed he collected them. Darwin
did not designate types and it seems purely academic to seek out all remnants of
Darwin’s barnacles in the possibility that some may not be E. modestus.
Insofar as it goes, Darwin’s description of E. modestus does describe E. modestus
as it is known in New Zealand and Europe. The barnacles Darwin ‘found’ at Sydney
were on oysters, and on Sydney shores oysters are ‘zoned’ below the confounding
species. Therefore, it is possible that Darwin did collect E. modestus at Sydney in
1836. More discerning collecting and identification of Elmznzus is required to
ascertain the details of the geographic distribution of both species in Australia. If it is
found to be restricted to major shipping ports, and absent from the intervening
estuaries on the coast, then it is possible that ship-aided dispersal to Australia has
occurred, some at least pre-1836. ;
Without examination of specimens, it is not possible to determine the true
identity of the barnacles listed by Guiler (1952) and Pope (1966), and figured by
Underwood (1977).
Elminius covertus n.sp.
Figs 1B, 3, 4A-C
Synonymy: Elminius modestus Pope, 1945:368; Underwood, 1977:23, ?17;
Guiler, 1952:20; Pope, 1966:181
Elminius sp. Foster, 1980:614 (figs 1, 3, 4)
Material examined. Personal collection: in 1975, Pittwater and Port Jackson; in 1978,
Coffs Harbour, Nambucca River (Macksville), Hastings River (Settlement Point,
Port Macquarie), Camden Haven, Wallis Lake (Tuncurry), Port Stephens (Tea
Gardens, Nelson Bay), Port Hunter (Stockton), Lake Macquarie (Swansea), Broken
Bay (Gosford, Bobbin Head, Pittwater), Port Jackson (Vaucluse, The Spit, Bantry
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
B. A. FOSTER
G H J
Fig. 3. Elminius covertus. A: inner view of shell; B: scutum; C: tergum; D: labrum; E: mandible; F:
maxilulle; G-I: cirri I-III; J: seta from 5th segment, posterior ramus, cirrus III; K: middle segment,
posterior ramus, cirrus VI. All drawn from a 9 mm rostrocarinal diameter specimen.
Bay, Roseville Bridge), Botany Bay (Captain Cook Bridge), Port Hacking
(Cronulla) ; in 1979, Port Jackson (Rose Bay, coll. M. F. Barker) ; in 1980, Pittwater
and Bennelong Point.
Australian Museum: Queensland localities, P21306, P21313 (both Stradbrooke
Island, 1961); New South Wales localities, P21296 (Hawkesbury, 1967), P21314
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
26 NEW BALANOID BARNACLES
(Pittwater, 1962), P21301-(Port Jackson, 1953), P21303 (Port Hacking, 1952) ;
Victorian localities, P11643 (Kangaroo Island, Bass Strait, 1945), P21283 (Fisher
Island, Bass Strait, 1930), P21297 (Mallacoota Inlet, 1957), P21309 (Westernport
Bay, 1962); Tasmanian localities, P21286 (Recherche Bay, 1953), P21288 (Sandy
Bay, Hobart (195) 221289 s(Port Anchunme 195i) 220 Ons Gear are oy)
P21295 (Port Arthur, 1951); South Australian localities, P21294 (Port Adelaide,
1950) ; Western Australian localities, P21311 (Bunbury Jetty, 1959), P21316 (Oyster
Bay, Albany, 1959). ;
Deposition of types: Type specimens from Lake Macquarie (Swansea), coll. 8 March
1978. Holotype; Australian Museum Cat. No. P30976. Paratypes; Australian
Museum, Sydney (Cat. No. P30977), National Museum, Wellington (Cat. No.
Cr2238), British Museum (Natural History), London (Cat. No. 1980. 305-314), U.S.
National Museum, Washington (Cat. No. 181718).
Description: Shell (Figs 1B, 3A) : flat, with 4 parietal plates, up to 15 mm across and
4 mm high. Parietes thin, solid; basal margin sinuous from within. In uneroded
specimens, parietes have pale narrow ribs alternating with pale wine-red coloured
spaces between: 4 or 5 ribs per plate in young specimens but up to 9 in larger
specimens. In eroded specimens, the shell is grey and granulate apically, but usually
with some evidence peripherally of dark reddish-purple laminae and ribbing.
Radii narrow, not completely covering underlying alae in uneroded specimens,
dark wine-red in colour, not interlocking with adjacent parietes. On erosion, the
sutures between parietes remain obvious but the radii and alae are indistinguishable.
Base membranous. Orifice pentagonal in outline, elongate in the rostrocarinal axis
with a very short rostral side.
Opercula (Figs 3B, C) : Scutum about as long or longer than high, the articular ridge
occupying half articular margin and curving into the articular furrow. Internally
there is no adductor ridge, nor crests for depressor muscles, but a faint adductor
muscle scar. Tergum elongated with a very wide articular furrow; articular ridge
standing at right angles to the pitted innerface of the plate, curving directly to a spur
that merges with the basiscutal angle; basal margin concave, with 4 or 5 crests
depending at the carinal end.
Externally in live, uneroded specimens, the scuta are dark red with a conspicuous
widening white band along the tergal margin of each scutum; opercular membranes
are white with 6 pairs of dark spots as shown in Fig. 1B.
Mouthparts (Figs 3D-F) : Labrum with 3 teeth on each side of central cleft. Mandible
with 5 teeth, the 2nd about % along the cutting edge from the upper tooth, the 5th
merging with a pectinate lower angle which bears a prominent curved spine at the
bottom point. Maxillule with a wide but shallow notch below the upper pair of spines,
the main cutting edge not protuberant bearing 3 or 4 major spines and a clump of
short ones at the lower angle.
Cirri (Figs 3G-K) : Number of segments in the rami of 5 specimens as follows, anterior
ramus first :
Shell length (mm) I II Ill IV V VI
8.5 EG 7,6 10,9 16,17 16,18 20,19
5.2 (eroded) 12,6 @) 7 11,10 15,17 23,21 25,23
12.0 14,6 9,7 11,10 17,20 24,19 21,19
12.6 16,6 9.8 U2, 18,19 22) 24 25,26
9.2 (eroded) 16,7 9.8 12,10 D3.) 26,27 BI 27
Cirrus I with anterior ramus 4% as long again as posterior ramus, distal segments
elongate with long setae; posterior ramus with segments slightly protuberant
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
B. A. FOSTER 27
Fig. 4. A: Elminius covertus, group of specimens from Pittwater, photographed alive, X 0.9; B: E.
covertus left and centre, Hexaminius popeiana right, from Swansea, Lake Macquarie, photographed alive,
2.5: C-F: medial face of middle segments of posterior ramus of cirrus III of E. covertus (X 220) C, H.
popetana (X 125) D, Elminzus modestus (x 60) E, Elminius king@i (X 220) F.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
28 NEW BALANOID BARNACLES
anteriorly. Cirrus II with anterior ramus slightly longer than posterior ramus,
segments of both rami setose and slightly protuberant anteriorly. Cirrus III with
subequal rami, all but basal 1 or 2 segments on medial face of posterior ramus with
pectinate setae (Fig. 4C). These pectinate setae are much stouter than the finely
feathered setae of the posterior rami. Cirri IV to VI with square segments with 4,
rarely 5, pairs of setae on the anterior edge. Penis 1 x length of cirrus VI, no basal
dorsal point or setae.
Etymology: with reference to the species being so long unnoticed on harbour shores.
Remarks: As both E. modestus and E. covertus may occur together it is desirable to
emphasize the distinguishing characters. In young uneroded shells the reddish to buff
colour of E. covertus is in marked contrast to the vivid white shell of E. modestus. Also,
the narrow contrastingly-coloured ribs of E. covertus differ from the uniformly white
and broadly-folded parietes of E. modestus.
Eroded shells are harder to identify; close inspection may reveal a stellate basal
margin where ‘the continuation of the ribs stick out round the margin in a series of
points’ (Pope, 1945:369).
As with all intertidal barnacles, there is some variation in the dimensions of the
scutum, ranging from almost isosceles (with articular and basal margins equal) to the
basal margin being 114 times the length of the articular margin, perhaps a feature of
flat conic species growing on different substratum contours.
The terga also vary in shape mostly due to the degree of erosion of the outer face.
The terga of E. covertus are more elongate than those of E. modestus (‘hatchet-
shaped’ of Pope, 1945), with the articular ridge overhanging the spur with which it
merges (forming the ‘handle of the hatchet’) and extending as a thickened rim around
the apico-carinal region. The triangular region between the depressor muscle crests
and the spur is notably thinner, often pitted. In E. modestus the inner surface slopes to
the apex and articular ridge, and is roughened in larger specimens.
The major point of difference in the appendages is the presence of pectinate setae
on the posterior ramus of cirrus III in E. covertus (Fig. 4C), absent in E. modestus
(Fig. 4E).
In the gaping animals, the tergoscutal flaps reveal the most marked
distinguishing character. In E. covertus (see Fig. 4A, B; Fig. 1B) they are white with
6 pairs of discrete black spots, one at the groove, 3 to the carinal side and 2 to the
rostral side.
On the basis of the opercula, Elmznzus covertus and E. modestus are more closely
related to each other than to E. kengz of South America; in E. kengz the tergal spur is
separate from the basiscutal angle and aligned with the articular margin as it is in
Solidobalanus spp. However, the posterior ramus of cirrus III of E. kengz bears
pectinate setae (Fig. 4F) like those in E. covertus (Fig. 4D), lacking in E. modestus
(Fig. 4E).
Genus Hexaminius nov.
Balanoid barnacles with a medially cleft labrum; 6 thin, solid parietes; rostrum not
elongated; radii simple; base membranous; cirri III and IV without hooks or teeth;
basi-dorsal point of penis vestigial.
Remarks: This is a monospecific genus proposed to accommodate the species
described below, hereby designated the type species.
Hexaminius popezana n.sp.
Figs 4B, 4D, 5, 6
Synonymy: Solzdobalanus sp. Foster, 1980: p, 614, fig. 3
Proc. LINN. Soc. N.S.W., 106 (1), (1981) 1982
B. A. FOSTER 29
Material examined. Personal collection: New South Wales localities: In 1975, Careel
Bay; in 1978, Hastings River (Settlement Point, Port Macquarie), Wallis Lake
(Tuncurry), Port Stephens (Tea Gardens, Karuah, Nelson Bay) ; Lake Macquarie
(Swansea) ; Broken Bay (Gosford, Careel Bay, Bobbin Head); Port Jackson (The
Spit, Roseville Bridge, Vaucluse) ; Botany Bay (Captain Cook Bridge) ; Port Hacking
(Cronulla). Further specimens from Rose Bay (Port Jackson) in 1979, (coll. M. F.
Barker) ; in 1980, Pittwater (Broken Bay) and Bennelong Point, Port Jackson.
Deposition of types: Holotype and paratypes — Australian Museum, Sydney, Cat. No.
P30974 (holotype) Cat. No. P30975 (paratypes) from Lake Macquarie, 8 March
1978. Paratypes — National Museum, Wellington (Cat. No. Cr2239), from Cowan
Creek, Hawkesbury, coll. 9 March 1978; British Museum (Natural History) , London
(Cat. No. 1980.315-320) , from Lake Macquarie, coll. 8 March 1978; U.S. National
Museum (Cat. No. 181719), Cowan Creek, Hawkesbury, coll. 9 March 1978.
Description: Shell (Figs 5, 6A): conic, up to 10 mm across and 4 mm high. Rostrum
wide; carinolatera about 1/3 width of latera. Compartments solid throughout, thin,
sometimes broadly sinuous, without ribs internally, with basal edge truncated. Radii
narrow, with simple edges, not interlocking with adjacent parietes, exposing much of
the underlying ala. In young and uneroded specimens, shell compartments with broad
pale bands alternating with bands of purplish to wine red colour, the latter with white
flecks which underly short projecting spines that line the growth ridges. In eroded
specimens, colour generally white with narrow darker stripes confined to the
depressions between the broad ‘ribs’. Base membranous. Orifice pentagonal in
outline, with a very short rostral side.
Opercula (Figs 6B, C): Scutum longer than high, articular ridge occupying 2/3
articular margin, with deep articular furrow; internally with very faint muscle
attachment scars, and a weak adductor ridge. Tergum waisted by having a concave
basal margin, but straight articular margin; deep articular furrow; spur close to but
separate from the basiscutal angle, forming an acute angle with the short part of the
—_ Imm
Fig. 5. Hexaminius popecana n.gen. and n. sp., drawn from a specimen from Rose Bay, Port Jackson.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
30 NEW BALANOID BARNACLES
Fig. 6. Hexaminius popezana. A: inner view of shell; B: scutum; C: tergum; D: labrum; E: mandible;
F: maxilulle; G-I: cirri I-III; J: seta from 7th segment, posterior ramus, cirrus III; K: middle segment,
posterior ramus, cirrus VI. All drawn from a 9 mm rostrocarinal diameter specimen.
basal margin on that side, spur curves gradually to the concave basal margin, which
on the carinal side is prominent with 4 or 5 crests for the carinal depressor muscles.
Externally in live specimens, the scuta are dark purple red with an inconspicuous pale
flange along the tergal articular edge of each scutum; opercular membranes off-white
with paired markings as shown in Fig. 6C.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
B. A. FOSTER 31
Mouthparts (Figs 6D-F): Labrum cleft, with 3 teeth on either side. Mandible with 5
teeth, the uppermost separated from the 2nd by half the length of the cutting edge;
the 3rd, 4th and 5th merging into a molariform lower angle which may bear a few
short spines. Maxillule with a slight notch below the upper pair of spines, the edge
below not protuberant, with 5 or 6 spines, the lowest 2 of which equal in length the 2
above the notch.
Cirri (Figs 6G-K) : The numbers of segments in the rami of cirri of 3 specimens as
follows, anterior ramus first :
Shell length I II Ill IV V VI
4.8mm 9.6 8,7 9.9 16,17 20,20 21,20
9.6mm 14,7 8,8 10,10 20,21 24,24 25,25
9.6mm 7 8,9 11,10 21,20 2424 26,26
Cirrus I with anterior ramus only slightly longer than posterior ramus. Cirrus IT with
subequal rami. Cirrus III with subequal rami, setae on distal 3 or 4 segments of both
rami serrated more on one side than the other (Fig. 4D and Fig. 6J). Basal segment of
cirri III very broad. Cirri IV to VI with segments slightly longer than wide, each with 5
pairs of setae on the anterior edge. Penis longer than cirrus VI, with a small basal
dorsal point bearing 2 setae, and with a prominent pedicle.
Etymology: named in honour of Miss Elizabeth Pope, student of Australian barnacles,
whose collections in the Australian Museum contain numerous lots of this species. It is
a species she has clearly pondered on.
Remarks: The poreless shell structure and unmodified 3rd and 4th cirri indicate an
affinity with Notobalanus and Chirona; the position of the tergal spur, set aligned to
the articular margin and very near to the basiscutal angle, resembles Solzdobalanus
and Elminius kingz. The absence of a calcareous base precludes its inclusion in any of
these genera. Nor can it be included in the membranous-based genera Semzbalanus
and Membranobalanus, because of the poreless shell and simple cirri respectively.
The similarity of H. popezana with Elmznzus is striking, particularly with respect
to the primitive features of non-toothed cirri, basiscutal position of tergal spur, the
membranous base, and the non-interlocking radii. Indeed, Hexamznzus satisfies the
requirements of an Elmznzus ancestor as discussed by Foster (1978, p. 97).
All species of Elmznzus and Hexaminius are characteristic of shallow sea habitats
and estuaries. Their weakly-constructed shells are inappropriate to surf habitats.
These species, the Elmininae, may represent the survivors of an early stage in balanid
radiation before the development of stronger radially-interlocked shells like those of
Solidobalanus. They perhaps indicate a southern hemisphere development to parallel
the Semibalaninae (e.g. Semzbalanus balanozdes) of the northern hemisphere. The
fouling proclivity of E. modestus has enabled it to overcome oceanic barriers and
become sympatric with related species in Australia and Europe.
ACKNOWLEDGEMENTS
I am grateful to the Director of the Australian Museum for allowing me to study
specimens therein, and acknowledge the work of Miss Elizabeth Pope whose personal
collecting significantly contributed to the barnacles collection in the Museum. For
other specimens I am grateful to Miss A. Gackle, Dr John Buckeridge and Dr Michael
Barker. I thank Prof. D. T. Anderson for reading the manuscript, Mr Charles
McKenzie and Mr Gary Batt for help with the illustrations, and the Auckland
University Research Committee for funding travel to Australia.
References
BisHop, M. W. H., 1947. — Establishment of an immigrant barnacle in British coastal waters. Nature
159:501.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
32 NEW BALANOID BARNACLES
Darwin, C., 1854. — A monograph of the subclass Cirripedia with figures of all the species. The Balanidae,
the Verrucidae etc. London: Ray Society.
Foster, B. A., 1978. — The marine fauna of New Zealand: Barnacles (Cirripedia Thoracica) . Mem. N.Z.
Oceanogr. Inst. 69:1-160.
——, 1980. — Biogeographic implications of re-examination of some common shore barnacles of Australia
and New Zealand. Procs. Int. Symp. Mar. Biogeography & Evolution in the Southern Hemisphere,
613-623. N.Z. DSIR Information Ser. 137.
GuILER, E. R., 1952. — A list of the Crustacea of Tasmania. Rec. Queen Victoria Mus. 3:15-44.
Ko.osvary, G., 1942. — Studien un Cirripedien. Zool. Anz. 137:138-150.
Moore, L. B., 1944. — Some intertidal sessile barnacles of New Zealand. Trans. Proc. R. Soc. New
Zealand 73 :315-334.
Morton, J. E., and MILER, M. C., 1968. — The New Zealand Sea Shore. London, Auckland: Collins.
Newman, W. A., and Ross, A., 1976. — Revision of the balanomorph barnacles; including a catalog of the
species. Mem. San Diego Soc. Nat. Hist. 9:1-108.
NILSsON-CANTELL, C. A. 1925. — Neue und wenig bekannte Cirripeden aus den Museen zu Stockholm und
zu Upsala. Ark. Zool. 18A:1-46.
Pope, E. C., 1945. — A simplified key to the sessile barnacles found on the rocks, boats, wharf piles and
other installations in Port Jackson and adjacent waters. Rec. Australian Museum 21:351-372.
——, 1966. — Sessile barnacles (Thoracica, Cirripedia). Port Phillip Survey, 1957-1963. Mem. Natl Mus.
Melbourne 27: 179-182.
SOUTHWARD, A. J., and Crisp, D. J., 1963. — Barnacles of European waters. In: Catalogue of main marine
fouling organisms, Vol. 1, Barnacles, pp. 1-46. Paris: Organisation for Economic Cooperation and
Development.
UNDERWOOD, A. J., 1977. — Barnacles. Science Field Guides. Sydney: Reed Education.
Proc. LINN. Soc. N.S.W., 106 (1), (1981) 1982
Systematics of Tapezgaster
(Diptera: Heleomyzidae)
with notes on biology
and larval morphology
DAVID K. McALPINE and DEBORAH S. KENT
McALpineE, D. K., & KENT, D. S. Systematics of Tapezgaster (Diptera: Heleomyzidae)
with notes on biology and larval morphology. Proc. Linn. Soc. N.S.W. 106 (1),
(1981) 1982: 33-58.
The genus Tapezgaster, which inhabits temperate Australia, is reviewed
systematically and placed in the monogeneric tribe Tapeigastrini of the family
Heleomyzidae. Four species are described as new. New synonymy is recorded for five
species. Lectotypes are designated for four nominal species. A neotype is designated
for Dryomyza cingulipes Walker. The third instar larva and puparium are described
for two species. Brief notes on the biology of the insects are given, with a list of
recorded host fungi.
David K. McAlpine and Deborah S. Kent, The Australian Museum, Box A285,
Sydney South, Australia 2000; manuscript received 22 April 1981, accepted for
publication 19 August 1981.
INTRODUCTION
Although the genus Tapezgaster was reviewed taxonomically by Paramonov
(1955) this is considered an appropriate time to present a revision of the genus for
several reasons. Type material of all nominal species has been examined (except for
one which appears no longer to exist) as well as a considerable amount of new
material. As a result it becomes necessary to change the names in use for several
species, and to describe four new species. As no additional new species has come to the
attention of the senior author in more than twenty years it appears that the register of
species is complete or almost so.
In listing material the names of the following collectors are abbreviated to the
initials: D. H. Colless, I. F. Common, G. Daniels, B. J. Day, G. A. Holloway, Z. R.
Liepa, D. K. McAlpine, K. R. Norris, M. S. Upton. Institutions housing material are
abbreviated thus: AM, Australian Museum, Sydney; ANIC, Division of Entomology,
Commonwealth Scientific and Industrial Research Organization, Canberra; BCRI,
Biological and Chemical Research Institute, (N.S.W. Department of Agriculture) ,
Rydalmere, Sydney; BM, British Museum (Natural History), London; CNC,
Canadian National Insect Collection, Agriculture Canada, Ottawa; PM, Muséum
National d’Histoire Naturelle, Paris; SPHTM, Commonwealth Institute of Health
(formerly School of Public Health and Tropical Medicine), University of Sydney;
UQ, Department of Entomology, University of Queensland; USNM, National
Museum of Natural History, Smithsonian Institution, Washington, D.C.; WAM,
Western Australian Museum, Perth; WM, Naturhistorisches Museum, Vienna.
Tribe TAPEIGASTRINI
The tribal characters are those of the only genus, but the main distinguishing
characters may be summarized as follows:
Fronto-orbital plates short and parallel with eye margin; face concave, uniformly
Proc. Linn. Soc. N.S.W.,106 (1), (1981) 1982
34 SYSTEMATICS OF TAPEIGASTER
sclerotized; prelabrum reduced. Antenna decumbent, with arista inserted sub-basally
on segment 3. Thorax with one or 2 dorsocentral bristles and no mesopleural bristle.
All femora with ventral spines. Costal spines absent; subcosta complete, diverging
from vein 1 distally; vein 6 long, sometimes attaining wing margin.
We place the genus Tapezgaster alone in the tribe because it is only distantly
related to any other heleomyzid genus, and its nearest ally is unknown. This tribe is
considered to have similar status to other groups in the Heleomyzidae, which are
probably best regarded as tribes e.g. Suilliini, Allophylopsini, Cnemospathini,
Rhinotorini, Trixoscelidini. The species of Tapezgaster are apparently restricted to
temperate parts of Australia. Probably, then, the group has evolved in isolation on the
Australian continent for much of the Tertiary; or, alternatively, Tapezgaster may be a
relict group, whose nearest allies in other regions are extinct.
Griffiths (1972) has associated Tapezgaster with the Rhinotorini, to which he
gives family rank. We find the resemblances between Tapezgaster and the other
genera referred by Griffiths to the Rhinotoridae unconvincing as indicators of close
phylogenetic affinity, because most of them are widely distributed among heleomyzoid
groups or seem likely to be readily duplicated by convergence. On the other hand,
Tapeigaster shows some fundamental differences from the rhinotorine genera. In
Tapeigaster the antenna is decumbent, with the arista inserted near the base of
segment 3; the prelabrum is reduced; the subcosta forms a distinct tubular vein right
to its junction with costa, and diverges from vein 1 distally; vein 6 is gradually reduced
distally and generally long (most reduced in T. pulverea) ; vein 7 is distinct; the anal
crossvein is transverse and forms with vein 6 a definite posterodistal angle to anal cell.
In these characters Tapezgaster contrasts with all true rhinotorine genera (including
Apophoneura and Anastomyza). The larva of Tapezgaster is very different from that
of the only known rhinotorine larva (Cazrnsimyza robusta (Walker), see McAlpine,
1968). The pupa of Cazrnsimyza lacks the prothoracic respiratory horn which
perforates the puparium in Tapezgaster. Griffiths gives as a ground plan condition of
Rhinotoridae: ‘Cerci of males small, fused below anus’. In Tapezgaster the cerci are
neither significantly reduced in size nor fused.
McAlpine (1968) gives reasons for considering the Rhinotorini to be an integral
part of the family Heleomyzidae. As Griffiths’s fragmentation of the Heleomyzidae
depends largely on the erroneous assumption that characters of the male postabdomen
are appropriate for higher classification within the Schizophora, we continue to regard
Rhinotorini and equivalent groups as tribes. Because structural divergence between
closely related species is usually greater in male postabdominal organs than in any
other part of the insect, these characters must be regarded as the least stable above the
species level and of low reliability as indicators of broad relationships.
Genus TAPEIGASTER
Tapeigaster Macquart, 1847: 86-87 (in reprint). Type species T. annulzpes
Macquart.
Scromyzoptera Hendel, 1917: 46. Type species S. annulata Hendel.
A genus of medium-sized to rather large flies of medium to robust build, having
most of the characters of the family Heleomyzidae s.l. as defined by Colless and
McAlpine (1970).
Head rounded, with convex occiput; fronto-orbital plate not sharply
differentiated, short, parallel with eye margin, bearing one or 2 fronto-orbital bristles
which are sometimes very fine and hair like; postvertical bristles strong, convergent or
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
D.K. McALPINE AND D.S. KENT 35
crossed; uppermost postocular bristle well developed; face uniformly sclerotized,
concave in profile, without distinct median carina but often with median line slightly
raised, with epistomal margin more or less prominent. Antenna deflexed from
articulation between segments 1 and 2; segment 1 short, setulose at least on dorsal
margin; segment 2 setulose, with one longer dorsal bristle; segment 3 rounded oval,
compressed ; arista inserted before middle of dorsal margin of segment 3; segments 4
and 5 small, not notably swollen; segment 6 long, not much swollen basally, with
numerous minute hairs throughout. Prelabrum not prominent, only slightly
projecting from general level of peribuccal area; proboscis somewhat elongate, with
short labella.
Thorax with the following bristles: humeral, 1 + 1 notopleurals, presutural
(sometimes absent), supra-alar, postalar, intra-alar, one or 2 dorsocentrals,
prescutellar acrostichal (sometimes absent), 2 pairs of long scutellars, inserted
apically and dorsolaterally respectively, one sternopleural; propleural and
mesopleural bristles absent; scutellum somewhat rounded, with short hairs and
pubescence-pruinescence. All femora with strong ventral spines, mostly aligned in the
anteroventral and posteroventral series; mid femur with one or more posterior
preapical bristles, often in a short, oblique series; tarsi with distal segments depressed
(least so in T. swbglabra) ; in males fore coxa, ventral surface of femora and tibiae
and ventral part of sternopleuron often with long, fine, dense hairs. Wing without
dark markings; costa much weakened and notched but not deeply incised at end of
subcosta, not broken or weakened just beyond humeral crossvein, with usually 2 to 4
bristles on section bordering first costal cell, otherwise without differentiated bristles,
with a series of short stout spinules dorsally bordering subcostal and marginal cells;
subcosta well developed throughout, diverging from vein | distally; anal crossvein
with sigmoid curvature, almost transverse; vein 6 long, usually gradually reduced
distally to an unpigmented fold which reaches wing margin or is discontinued a short
distance from it, sometimes pigmented to margin (T. subglabra) ; vein 7 represented
by a usually well defined, dorsally convex crease in membrane which is often
pigmented.
Abdomen with preabdominal spiracles in pleural membrane; d postabdomen
with abbreviated tergite 6; sternites 6 and 7 well developed, displaced to left side,
connected to the dorsal sternite 8; cerci distinct, separate. 9 post-abdomen with no
particularly elongate segments; segments 6 and 7 with separate tergites and sternites;
cerci free.
Key to Species of Tapezgaster
1. Prosternum without long hairs, with minute pubescence only ........ 2
Prostscnwiaen wun Momer eu > $3 Ss sc ae oe assess asus Se aes ea 3
2. Vein 6 not reaching wing-margin; presutural bristle present; vibrissa
arising from brownish spot, subtended by a second fairly strong
bristles (uilbrissaenotiso: in amy Other Species) vr. 2 oe a ee pulverea
Vein 6 reaching margin though weakened apically; presutural bristle
absent (present in all other species) ; vibrissa not arising from a
pigmented spot, subtended only by fine hairs .............. subglabra
3. Propleuron with numerous long, fine hairs near centre; mid and
liitaderenitoraanotsmotablysswolllemters a) ses Mele oeesolun ye Meee ne so 64 - 4
Propleuron not haired near centre; other characters variable ....... 6
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
36 SYSTEMATICS OF TA PEIGASTER
4. Mesoscutum entirely pruinescent, not shining, with three broad, well-
defined, longitudinal grey bands; fore and mid tibiae with only the
apical black band distinct; surstylus unequally bilobed, the lobes
strongly: procurved seahiich. aii AS cen SH eae ee tener area paramonovt
Mesoscutum without longitudinal bands, shining and sparsely _
pruinescent; all tibiae with a black band at apex and another above
middle esunstylasio titer wal Sesh © tern Cle epee een eee ee 5
5. Fore femur browned on no more than distal quarter; thoracic pleura
in part sparsely pruinescent, parts of mesopleuron and sternopleuron
somewhat shining; male terminalia with surstylus inserted close to
cercus, undivided, broad basally, with apical part narrow ..... annulata
Fore femur with brown anterodorsal stripe from well before middle
to apex; thoracic pleura entirely densely pruinescent, not shining;
male terminalia with surstylus inserted a short distance in front of
cercus, forked, the posterior lobe broadly obtuse ....... Se Aeriiah digitata
6. Prescutellar acrostichal bristle absent or reduced; mesoscutum tawny
with three regular, grey-pruinescent, longitudinal bands; abdominal
tergite 9 of male with pair of incurved horn-like processes .... annulipes
Prescutellar acrostichal bristle well developed; mesoscutum not
marked as above; abdominal tergite 9 of male with at most a
sibbositysonveaehside i) fs gil icslekpie dt aaron. aaah des ae ele ike aac a as 7
7. Vein 6 discontinued before reaching wing margin; wing membrane
with narrow bare zone in front of vein 7; mid femur stouter than
hindtone (viewedifirom above)) #4441) a ieee ama Lee PemTies
Vein 6 reaching wing margin though faint distally; wing membrane
entirely microtrichose near vein 7; hind femur usually stouter than
TMMUGIVONME 4 ee Ena Mya Her coe 2eobl sarc ais: cole Re ys ek tO eh eA ee aces ee Eee 8
8. Mid tibia darkened only at apex; hind tibia with grey pruinescence
on most of surface; posterior surface of fore femur with neither
depression nor transverse grey-pruinescent band; wing not yellowish
basally; postfrons with complete silvery-pruinescent orbital stripe,
but without silvery orbital spots; abdominal tergite 9 (epandrium) of
maleswith eb bOsibyaonkeachysic cy sie sa lenient an aeeee nigricornis
Mid tibia darkened sub-basally as well as apically; hind tibia shining,
without obvious pruinescence; posterior surface of fore femur with
either depression or transverse grey-pruinescent band just beyond
middle; wing yellowish basally; postfrons with silvery-pruinescent
orbital spots; abdominal tergite 9 of male without paired
PUD DOSTEVES H1.0 75. Stele joviolke Pane se eRe eee sac oe meat cies eRe ee ea 9
9. Mesoscutum pale brownish with a large round blackish spot before
and behind transverse suture on each side; fore femur with
whitish-pruinescent transverse stripe on posterior surface beyond
FagvKO (CG | (eas Wiens Neat MR MeeRecind Sear cael Sea Ue ge STROM. Rare ee cinctipes
Mesoscutum differently coloured; fore femur without such stripe .... 10
10. Mesoscutum tawny-orange, with distinct white-pruinescent markings
on margins only; fore coxa entirely fulvous-orange, largely pruinescent
ON OUCER/SUTPACE 2 Silo Daa se a eee ee argyrospila
Mesoscutum deep brown with whitish-pruinescent markings on central
part; fore coxa with glossy brown zone on outer surface ... brunnezfrons
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
D. K. McALPINE AND D. S. KENT 37
Tapergaster nigricornis (Macquart) n. comb.
(Fig. 3)
Sctomyza nigricornis Macquart, 1851: 277-278, pl. 25, figs. lla, 11b.
Tapergaster marginifrons Bezzi, 1923: 74-75; McKeown, 1942: 228, fig.;
Paramonov, 1955: 459-461. N. syn.
Bezzi has given a detailed description which is quoted by Paramonov. The species
is not very closely related to any other species, and is easily recognized from the
characters given in the key.
Distribution: Queensland — south-east districts as far north as Bundaberg; New
South Wales — very widely distributed; Victoria; Tasmania; South Australia —
mainly southern districts; Western Australia — southern districts north to Perth and
east to southern Nullarbor.
Type material of S. nigricornis: “Tasmanie’ (lectotype d, here designated, PM no.
4/8/47); ‘Akarea, Nouvelle-Zélande’ (probably Akaroa, Banks Peninsula),
(paralectotype 9, PM no. 4/46). The paralectotype is not conspecific with the
lectotype but is a lauxaniid of the genus Sapromyza in the sense in which that genus is
employed in New Zealand. It may well be referable to one of the species included by
Harrison (1959) in his revision of this group. After careful consideration and
discussion we have decided to designate the Tapezgaster specimen as lectotype.
Although this means altering the established name Tapezgaster margznifrons, it
removes a potential problem for lauxaniid workers.
Type material of T. marginifrons: Blue Mountains, New South Wales, 22.iv.1922
(lectotype d, here designated, right wing missing, AM), anon.; Sydney, New South
Wales, 2.vi.1922 (paralectotype 2, much damaged, head glued to label, AM), anon.
Other material examined (localities only given). Qld.: Woowonga Range, SW of
Bundaberg (ANIC) ; Tibrogaren Creek, via Beerburrum (UQ) ; Dayboro (ANIC) ;
Camp Mountain (UQ); Capalaba, near Brisbane (UQ); Rosewood (ANIC) ;
Lamington National Park (UQ); Inglewood (UQ); Braeside (UQ); Stanthorpe
(UQ). N.S.W.: Boonoo Boonoo, near Tenterfield (ANIC) ; Deepwater, near Glen
Innes (AM); Graman (BCRI); Mount Gibraltar (AM, ANIC); Bourke (BCRI) ;
near Rylstone (ANIC); Kandos (AM); Mount Coricudgy (AM) ; Derriwong, near
Condobolin (BCRI) ; Parkes (ANIC) ; Winburndale Nature Reserve, near Bathurst
(ANIC) ; 37 km (23 miles) N of St Albans (BCRI) ; Mount Wilson, Mount Tomah,
and Mount York, Blue Mountains (AM); Kurrajong (AM); Ooma Creek, NW of
Grenfell (ANIC); Katoomba (AM); Blackheath and Leura (BCRI); Wentworth
Falls (AM); Maroota, Gordon, Lane Cove, Bronte, Como and Sutherland, Sydney
district (AM) ; Manly Reservoir, near Sydney (ANIC) ; Port Hacking (AM) ; Royal
National Park (AM); Jenolan Caves (AM); Kanangra Plateau (AM); Mittagong
(BCRI); Moss Vale (BCRI); Wombeyan Caves (ANIC); Hilltop (UQ);
Abercrombie River, Bummatoo Forest (AM) ; Marulan (ANIC); Robertson (AM) ;
Gerroa (AM); Yanco (AM); Narrandera (BCRI); Leeton (BCRI); Bungendore
(AM); Royalla (ANIC) ; Billabong Creek, Wanganella (ANIC) ; Billabong Creek,
Conargo (ANIC); near Deniliquin (ANIC); Pilot Hill, Bago Forest, near Batlow
(ANIC); Talbingo (ANIC); Clyde Mountain (ANIC); Monga, near Clyde
Mountain (ANIC); Broulee (ANIC); Gerogery (ANIC); near Adaminaby
(ANIC) ; Nimmitabel (ANIC) ; Brown Mountain, near Nimmitabel (AM, ANIC) ;
Pambula (BCRI); Wilson’s Valley, Snowy Mountains (AM); The Creel, Snowy
Mountains (ANIC) ; Alpine Creek, Snowy Mountains Highway (ANIC). Vzc: Mount
Buffalo (ANIC); Seaford (ANIC); Mill Park, Melbourne (AM); near Woodend
(AM); Lorne (ANIC); Mount Buangor, near Beaufort (ANIC); Wedderburn
(ANIC) ; Mount William, Grampians (AM); Wannon River, Grampians (AM) ;
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
38 SYSTEMATICS OF TA PEIGASTER
Figs 1-2. Tapezgaster lutetpennis. 1-2. db genitalia. 1. lateral view. 2. posterior view.
Warrnambool (UQ); Dimboola (ANIC). Tas.: Derwent Bridge (UQ); Tyenna
River, near Mount Field National Park (AM); Mount Wellington (ANIC). S.4.:
Moorlands (ANIC); Mount Lofty Range (ANIC); Prospect (UQ); Second Valley
Road, Cape Jervis (ANIC) ; Seal Bay, and Bale’s Bay, Kangaroo Island (AM) ; Cape
Borda, Kangaroo Island (ANIC) ; Sleaford Bay (ANIC) ; 8 km SSE of Mount Hope
(ANIC); N of Nullarbor Homestead (AM). W.A.: Cocklebiddy (ANIC) ; Junana
Rock (ANIC) ; Mount Ragged (ANIC) ; NW of Mount Arid (ANIC) ; Thomas River
estuary, Esperance district (ANIC); Gibson (ANIC); E of Ravensthorpe (ANIC) ;
Mount Magog, Stirling Range (AM) ; Porongurup National Park (ANIC) ; E of Jewel
Cave, Augusta (ANIC) ; Darlington (AM).
Tapeigaster cinctipes (Walker) n. comb.
(Figs 4-5)
Heteromyza cinctipes Walker, 1853: 404.
Tapergaster bella Paramonov, 1955: 454-455, N. syn.
The description of Paramonov, together with the characters indicated in our key,
is adequate for identification.
The species is polymorphic in abdominal coloration. A majority of specimens
from eastern states, including the holotypes of Walker and Paramonov, have the
abdominal tergites fulvous with much of tergite 2 and all of tergites 3 and 4 blackish.
Specimens from Bendora (allotype 9 of T. bella), Mount Ainslie (paratype d of T.
bella), and Monga (1 G) have the abdominal tergites entirely fulvous. The three
specimens from Western Australia (both sexes) are intermediate between the two
eastern forms, having the dark coloration reduced to a brown patch on each side of
tergites 2 to 4, smallest on tergite 2. No difference in the male genitalia can be
discerned between the two eastern forms and the difference in shape of the surstylus in
the one western male examined is so slight that a significant difference between the
populations may not be indicated.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
D.K. McALPINE AND D. S. KENT 39
SO postabdomen. Surstylus broadly ovate, broadly gibbous on posterior margin from a
little beyond base, most strongly haired on anterior margin, with shallow, oblique
apical cleft producing a blunt tooth on inner surface; sides of hypandrium without
series of hairs.
Distribution: southern New South Wales; Australian Capital Territory; Victoria;
Tasmania; southern Western Australia.
Holotype 9 of H. cinctipes: “Van Diemen’s Land’ (Tasmania) (BM), anon.
Type materzal of T. bella: Blundell’s clearing, near Canberra, 6.iv.1947 (holotype d,
paratype 0, ANIC), S. J. Paramonov; Bendoora, A.C.T., 15.iv.1953 (allotype 2,
ANIC), S. J. Paramonov; Mount Ainslie, near Canberra, i1i.1950 (paratype 6,
ANIC), A. Floyd; National Park (i.e. Mount Field National Park), Tasmania,
xii.1922 (paratype 0d, ANIC), A. L. Tonnoir; Beverley, W.A., 1913 (paratypes 1d,
12, ANIC), ‘D.B.’
Other material examined (localities only given). N.S.W.: Murrumbateman
(ANIC); Monga, near Braidwood (ANIC); Broulee (ANIC, AM). A.C.T.: Black
Mountain, Canberra (ANIC). Vic.: Mount Macedon (AM). W.A.: Nedlands
(ANIC).
Tapeigaster argyrospila Bezzi
(Fig. 6)
Tapeigaster argyrospila Bezzi, 1923: 77-78; Paramonov, 1955: 458-459.
The species has been described in some detail by Bezzi and is correctly interpreted
by Paramonov. The surstylus is, as usual, quite distinctive.
Abdomen of G. Sternite 5 not divided into 2 plates, but rather weakly sclerotized
medially; surstylus three-lobed; central lobe rather elongate, tapering; anterior lobe
broadly rounded, with short hairs towards margin, not deeply divided off from central
lobe; posterior lobe more slender and deeply set off than anterior lobe, much shorter
than central lobe.
Distribution: Queensland — southern districts only; New South Wales — coast to
western plains; Victoria; South Australia.
Holotype 3 : Blue Mountains, 11.iii.1922 (AM, ex Health Dept. collection) , anon.
Other material examined (localities only given). Qld.: Mount Mowbullan, Bunya
Mountains (AM) ; Emuvale (UQ) ; Stanthorpe (UQ). N.S.W.: 11 km (7 miles) E of
Mendooran (AM); Brummagem Creek, near Dubbo (AM); Bogan River (AM) ;
Comark, near Oberon (BCRI); Leura Falls, Blue Mountains (ANIC) ; Thornleigh,
near Sydney (BCRI) ; Upper Middle Harbour, Sydney (AM); Mittagong (BCRI) ;
Leeton (AM) ; Billabong Creek, near Wanganella (ANIC). Vzc. : Upper Main Creek,
Mornington Peninsula (AM); Hobson’s Bay, Melbourne (AM); near Woodend
(AM) ; Cobrum (ANIC). S.A.: Sleaford Bay, Port Lincoln (ANIC).
Tapeigaster brunnezfrons Malloch
(Fig. 7)
Tapeigaster brunnezfrons Malloch, 1927: 16.
Tapeigaster vernalis Paramonov, 1955: 457. N. syn.
Descriptions have been given by Malloch and Paramonov. These are brief, but as
the species resembles T. argyrospila very closely, except in male genitalia, a detailed
redescription is not necessary.
Although the type of T. brunnezfrons was undoubtedly darker-coloured when
Malloch described it the year after the collection date, its coloration is now quite in
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
40 SYSTEMATICS OF T4 PEIGASTER
Figs 3-10. 3-8. 3 surstyli. 3. Tapecgaster nigricornis. 4. T. cinctipes (Mt. Ainslie, A.C.T.).5. T. cinctipes
(Beverley, W.A.). 6. T. argyrospila. 7. T. brunnezfrons. 8. T. annulipes. 9-10. T. paramonow. 9. d
genitalia. 10. db 5th sternite.
agreement with material of the species subsequently named vernalis by Paramonov.
Though, from the descriptions, the small size of the fronto-orbital bristles and the
single pair of dorsocentral bristles in T. brunnezfrons might seem to separate it from
T. vernalis, these characters are variable in the available series and asymmetrical
combinations of the bristle characters occur.
Proc. Linn. Soc. N.S.W.,106 (1), (1981) 1982
D. K. McALPINE AND D. S. KENT 4]
Abdomen of 6. Tergite 5 undivided, but with median part of posterior margin weakly
sclerotized ; surstylus not lobed as in T. argyrospila, subtriangular, with subacute apex
very slightly inclined forwards and short posterobasal projection, on inner surface a
strongly developed blade running from posterobasal projection towards, but not
reaching, apex.
Distribution: New South Wales — coast, tablelands and south-western plains;
Australian Capital Territory; Victoria; Tasmania.
Holotype 2 of T. brunnetfrons: Newcastle, N.S.W. 7.iv.1926 (USNM), anon.
Type material of T. vernalis. Lectotype d, here designated: Wahroonga, near
Sydney, 31.x.1926 (SPHTM), anon.; same data (paralectotype 9, SPHTM). These
specimens were marked “Typus 6’ and ‘Typus 9’ by Paramonov, and there is no
indication of a holotype in his description.
Other material examined (localities only given). N.S.W.: near Spring Creek, Ebor-
Point Lookout Road (AM); Mooney Mooney Creek, near Gosford (AM); Mount
Wilson, Blue Mountains (AM); Mount Tomah, Blue Mountains (ANIC);
Katoomba (AM); Wentworth Falls (AM) ; Glenorie (BCRI) ; Gordon, near Sydney
(AM); Royal National Park (AM); Manar, near Braidwood (AM); Monga, near
Braidwood (ANIC); Batehaven, near Bateman’s Bay (ANIC); Billabong Creek,
near Wanganella (ANIC). A.C.T.: Black Mountain, Canberra (ANIC). Tas.:
Hellyer Gorge (ANIC) ; Huon-Picton Junction (ANIC).
Comparative notes. T. brunnezfrons is very closely related to T. argyrospila, there
being few external characters separating them, though the former species is darker.
The surstylus of the male shows a quite remarkable degree of difference as described
above.
Tapeigaster lutecpennis Bezzi
(Figs 1-2)
Tapeigaster lutetpennis Bezzi, 1923: 76-77.
Tapeigaster taylort Malloch, 1935: 94-95. N. syn.
A detailed description has been given by Bezzi, but the male postabdominal
characters have not been recorded.
Abdomen of G : tergite 9 produced posteriorly into two short rounded protuberances ;
cerci concealed in a roughly oval membranous cavity beneath and between the
protuberances; surstylus very complex (Figs 1-2), with tubercle on basal region and
two divergent lobes; hypandrium with numerous rather long hairs medially.
Distributzon: Queensland — south-east; New South Wales — coast and warmer
eastern margins of tablelands. This species shows a greater degree of restriction to
warmer areas than any other in the genus. The record from Wentworth Falls and
probably that from Tallong indicate localities in nearby valleys at much lower
elevations than the towns named.
Holotype 3 of T. luteipennis: Eccleston, near Dungog, N.S.W., 1.i1i.1921 (not 1922
as given by Bezzi) (AM, ex Health Dept. collection) , anon.
Holotype 6 of T. taylort: Tallong, near Marulan, N.S.W., no date (SPHTM), F. H.
Taylor.
Other material examined. Qlid.: Bundaberg (UQ); Montville (UQ), Bunya
Mountains (UQ); Brisbane (UQ, ANIC); Tamborine Mountain (AM) ;
Cunningham’s Gap, near Maryvale (UQ); Lamington National Park (UQ).
N.S.W.: Legume (UQ); Tooloom (UQ); Ulong, East Dorrigo district (AM) ;
Dorrigo (ANIC) ; Wentworth Falls, Blue Mountains (AM) ; Heathcote (AM) ; Royal
National Park (AM) ; Otford (AM).
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
42 SYSTEMATICS OF TAPEIGASTER
Notes. It is clear from examination of types and additional material that T.
lutecpennis and T. taylor? are conspecific. There is a little variation in the amount of
white pruinescence on the mesoscutum and in the brown markings on the abdomen.
The only other species with paired gibbosities or tubercles on tergite 9 of the male
are T. annulipes and T. nigricornis. In T. annulzpes the tubercles are more elongate
and horn-like. In T. nzgricornzs the paired gibbosities are less prominent than in T.
lutecpennis and the cerci are much more prominent.
Tapezgaster annulipes Macquart
(Fig. 8)
Tapezgaster annulipes Macquart, 1847: 87, pl. 6, figs. 1, la-le; Bezzi, 1923: 75-76;
Colless and McAlpine, 1970, fig. 34.30A.
Dryomyza cingulipes Walker, 1857: 220, synonymized by Bezzi, 1923: 75.
A useful description is given by Bezzi. It may be added that the propleuron is not
haired just below the humeral callus, and vein 6 does not quite reach the wing-margin
though longer than in T. dzgztata and allied species.
Distribution: Queensland — southern border districts; New South Wales — mainly
coast and tableland districts; Victoria; Tasmania; South Australia — Lofty Ranges
and Kangaroo Island. The species is very common and widely distributed, being found
in forested areas and sometimes inner urban areas. It is apparently absent from some
of the warmer, drier districts inhabited by T. nzgricornzs and T. argyrospila.
Type material of T. annulipes: no locality on label but Macquart gives ‘Nouvelle-
Hollande’, ‘Tapeigaster annulipes. d @. n. g. n. sp. Macq’ (in Macquart’s
handwriting) and on separate disc “TYP’ (lectotype G here designated, OX).
Macquart probably had 2 specimens in his type series as he gives a single measurement
for each sex. The lectotype is the only specimen which we can identify with certainty as
belonging to the type series, and its designation as such is essential to preserve current
usage of the name T. annulzpes. Also above the cabinet label T. annulzpes in the Bigot
collection at Oxford, there are two other flies at least one of which should be a
paralectotype of T. annulipes though neither is conspecific with the lectotype. The
first is a female pyrgotid with a “IYP’ disc on the pin which could indicate type
material if its authenticity could be established. The second specimen is a male
Tapeigaster, probably of T. argyrospila but very mouldy. Some credence is given to its
possible type status, because Macquart’s pl. 6, fig. 1 (whole insect) , looks more like T.
argyrospila than T. annulipes in some respects. On the other hand Macquart’s
illustrations are so exceedingly inaccurate that they scarcely provide grounds for safe
conclusions.
Neotype of D. cingulipes: We designate the lectotype of Tapezgaster annulipes
Macquart as the neotype of Dryomyza cingulipes Walker. In so doing we stabilize the
otherwise doubtful synonymy first given by Bezzi and repeated by Paramonov (1955).
Fixing the identity of D. cingulipes in this way will prevent the use of the epithet for
other species of Tapezgaster here referred to by names of later date. In accordance
with Article 75(C) of International Code of Zoological Nomenclature we make the
following comments. (1) The species is as described by Bezzi (1923) and is
distinguished as given above in our key to species of Tapezgaster. (2) The label data
are those on the lectotype of T. annulipes given above. (3) Walker’s types of
Australian Diptera are, so far as we know, housed in the British Museum (Natural
History) ; the Hope Department of Entomology, University Museum, Oxford; and the
National Museum of Victoria, Melbourne. McAlpine has been unable to find type
material of D. congulipes at the British Museum or Oxford and B. H. Cogan confirms
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
D. K. McALPINE AND D.S. KENT 43
its apparent absence from BM. A. Neboiss informs us that there is no type material of
D. cingulipes at NMV. We therefore regard this original type material as probably
lost. (4) The original description of D. cingulipes provides the only available
information on the type material of that species. It almost certainly refers to a species
of Tapezgaster and agrees fairly well with T. annulzpes though it also resembles other
species, particularly the much less common T. pulverea. (5) The type locality for T.
annulzpes is given as ‘Nouvelle Hollande’, that for D. cingulipes as ‘New South Wales’.
As the term New South Wales at that time applied to most of the eastern half of
mainland Australia (or New Holland), there is no evidence that the two type localities
were remote from one another. (6) The neotype belongs to the University Museum,
Oxford, England.
Other material examined (localities only given). Qld.: Lamington National Park
(UQ) ; Stanthorpe (UQ). N.S.W.: 32 km (20 miles) , Glen Innes to Grafton highway
(ANIC) ; Mt. Kaputar, near Narrabri (AM, ANIC); New England National Park
(ANIC) ; near Spring Creek, Ebor— Pt. Lookout road (AM); Point Lookout, near
Ebor (AM) ; Wright’s Lookout, New England National Park (AM) ; near Mt Dagola,
Warrumbungles (AM); Nundle State Forest (ANIC); Tomalla (AM) ; Coachwood
Gully, 21.7 km SE Threeways (AM); Mt Irvine (AM) ; Mt Wilson, Blue Mts (AM) ;
Mt Tomah, Blue Mts (AM, ANIC) ; 8 km NW of Kurrajong-Bell road (ANIC) ; near
Mt Banks, Blue Mts (AM); Mt Boyce, Blue Mts (AM); Leura Falls (ANIC) ;
Govett’s Leap, Blue Mts (AM); Katoomba (AM); Wentworth Falls, Blue Mts
(AM) ; Sassafras Gully, Springwood (AM); Jenolan (AM); 6.4 km (4 miles) N of
Jenolan Caves (AM); Mt Queen Pin, Kanangra road, Boyd Plateau (AM);
Kanangra (AM); Boyd River Crossing, Kanangra road (AM); Budthingaroo Ck,
Kanangra-Boyd National Park (AM) ; Cowan (AM); Milson Island (AM) ; Sydney
(AM, UQ); Gordon (AM); Mosman (ANIC); Bexley (AM); Northwood (AM) ;
Royal National Park (AM, ANIC); Otford (AM); Cataract Ck, Bulli (ANIC); Mt
Keira, near Wollongong (AM) ; Colo Vale (ANIC) ; Mt Gibraltar, Bowral (ANIC) ;
6 km SE of Robertson (AM) ; Fitzroy Falls (AM); Beaumont, 6 km SE of Kangaroo
Valley (AM); Seven Mile Beach State Park, Gerroa (AM); Jervis Bay (ANIC) ;
Clyde Mountain, near Braidwood (ANIC); Mongarlowe River, Clyde Mountain
(ANIC) ; 9 km SE of Bateman’s Bay (ANIC) ; Broulee (ANIC) ; Rutherford Creek,
Brown Mtn (AM, ANIC); Brown Mtn, Bega district (ANIC) ; 6.4 km (4 miles) E of
Nimmitabel (ANIC); Tumut (ANIC); Pilot Hill, Bago Forest, Batlow (ANIC) ;
Kiandra, Alpine Ck (ANIC); Eucumbene-Lookout, Snowy Mts (ANIC); 19.2 km
(12 miles) NW of Adaminaby (ANIC); The Creel, Kosciusko (AM, ANIC) ;
Wilson’s Valley, Snowy Mts (AM) ; Sawpit Creek, Snowy Mts (AM, ANIC); 3 km S
Wragge’s Creek, Kosciusko National Park (ANIC). A.C.T.: Black Mountain
(ANIC); Mt Coree (ANIC); Bull’s Head (AM); Uriarra State Forest (ANIC) ;
Paddy’s River (ANIC); Lee’s Creek, Brindabella Range (AM); Condor Creek,
Brindabella (AM); Brindabella Range (AM). Vzc.: Dynamite Creek, Bonang
Highway (ANIC); 21 km (13 miles) W of Matlock (AM); Cement Creek, near
Warburton (AM); Warburton (ANIC); Sherbrooke Forest, near Kallista (AM) ;
Belgrave (AM); Hobson’s Bay, Melbourne (AM); Mt Macedon (AM); Daylesford
(ANIC); Mt Buangor, NE of Beaufort (ANIC). Tas.: 5 km (3 miles) S of Oonah,
Waratah Highway (AM); Hellyer Gorge (AM, ANIC); 19.2 km (12 miles) S of
Wilmot (AM); Mt Barrow, via Launceston (UQ); 5 km (3 miles) E of Waratah
(ANIC); Needles, near Deloraine (AM); Marakoopa Caves, near Mole Creek
(AM); 25.6 km (16 miles) NE of Cradle Mt (AM); Breona (UQ); Lake St Clair
(AM); Derwent Bridge (ANIC); Franklin River Crossing, Lyell Highway (AM) ;
National Park (UQ); Hobart (UQ); Mount Wellington (ANIC). S.4.: Mt Burr
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
44 SYSTEMATICS OF TAPEIGASTER
(ANIC); Glen Osmond (AM); Waterfall Gully, Burnside (AM); Engelbrook
Reserve, near Bridgewater (AM); National Park, Mt Lofty Range (ANIC) ; Upper
Ravine des Casoars, Kangaroo Island (AM).
Tapeigaster paramonovwi n. sp.
(Figs 9-10)
d Q. Closely related to T. dégztata and agreeing with description given for that
species except as indicated below.
Coloration. Postfrons orange-tawny, often deeper tawny-brown laterally, sometimes
darkened anteriorly, with whitish pruinescence along orbital margins and to lesser
extent on ocellar triangle. Mesoscutum tawny-brown with pale grey, yellowish-edged
median stripe from anterior extremity almost to scutellar suture where it is narrowed
to a point, and with broad yellowish grey lateral margins; scutellum tawny-brown on
central and anterior part of dorsal surface, broadly pale tawny-buff on lateral and
posterior margins; pleura with deeper tawny ground colour than in T. digztata, and
with even thicker covering of whitish pruinescence. Fore femur with brown
anterodorsal longitudinal stripe as in T. dzgitata but broader and more diffuse; fore
tibia with distal brown zone occupying nearly half length of tibia, without sub-basal
dark band; mid tibia browned apically, with sub-basal dark band at most indistinct;
hind tibia coloured approximately as in T. dzgztata.
Head with one or 2 fronto-orbital bristles, in the latter case anterior bristle very short.
Thorax. Femora of 6 with very numerous, rather short, fine ventral hairs.
Abdomen. Sternite 5 of db with each of paired lobes very rounded posteromedially. d
postabdomen: surstylus with 2 divergent lobes, both procurved and obtuse, the
anterior one larger, also with anterior basal gibbosity which bears much longer hairs
than those on lobes; hypandrium with a series of long hairs on each side, the longest
ones about as long as surstylus; cercus with anterior subapical tubercle.
Dimensions: total length, d 5.2-6.8 mm, 2 4.9-6.5 mm; length of thorax, 5 2.5-3.3
mm, 2 2.7-3.1 mm; length of wing, d 5.6-6.9 mm, 9 6.2-7.5 mm.
Distributzon: New South Wales (tablelands) ; Victoria; South Australia; Western
Australia (south-west) .
Holotype 3: Leather Barrel Creek, Snowy Mountains, 11.11.1979 (AM), B.J.D. and
D.K.M.
Paratypes. N.S.W.: Mount Kaputar, near Narrabri, xi.1964 (4d, AM), D.K.M.;
Mount Wilson, Blue Mountains, xii.1956 (1 9, AM), D.K.M.; 8 km (5 miles) S of
Mount Wilson, iv.1971 (1d, AM), D.K.M.; Mount Boyce, Blue Mountains, iii.1963
(1 d, AM), D.K.M.; Mount York, Blue Mountains, x.1960 (1 9, AM), D.K.M.;
Falls Creek, near Nowra, x1i.1926 (1 Q, AM), B. Bertram; Clyde Mountain (west
slope), near Braidwood, v.1965 (1 9, ANIC), D.H.C.; The Creel, near Mount
Kosciusko, xi.1961 (5 d, 29, ANIC), D.H.C.; Wilson’s Valley, Snowy Mountains,
11.1979 (1 d, AM), B.J.D. and D.K.M.; 19.2 km (12 miles) NW of Adaminaby,
xi.1961 (1d, ANIC), D.H.C.; Geehi River, xi.1961 (1d, ANIC), D.H.C. 4.C.T.:
Black Mountain, Canberra, viii.1968 (1 0, ANIC), I.F.C.; Mount Coree, iv.1968 (2
d, ANIC, 1d, 1 9, CNC), D.H.C., J. W. Boyes; Bendora, ii.1950 (1 2, ANIC),
K.R.N.; Cotter River, x.1956 (1 d, ANIC), Z.R.L.; Lee’s Springs, iv.1958 (1 9,
ANIC), Z.R.L.; Condor Creek, Brindabella Range, iv.1972 (1 9, AM), D.K.M.
Other material examined. Vic.: Frenchman’s Gap, near Wood's Point, iv.1963 (2 ©),
AM), D.K.M.; Daylesford, viii.1968 (1 9, ANIC), N. Dobrotworsky; Wannon
River, near Jimmy’s Creek, Grampians, xii.1977 (1 9, AM), M. A. Schneider and
D.K.M. Tas.: Cradle Mountain, i.1923 (19, ANIC), A. L. Tonnoir. S.A. : Second
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
D. K. McALPINE AND D. S. KENT 45
Valley Road, Cape Jervis, x.1975 (1d, ANIC), Z.R.L. W.A.: Darlington, ix-xii.1964
(1d, AM), G. L. Bush; 17.6 km (11 miles) SW of Collie, x.1964 (2d, 292, AM), G.
L. Bush; 24 km (15 miles) S of Mumballup, x.1964 (1 d, AM), G. L. Bush; Mount
Toolbrunup, Stirling Ranges (10, AM, 1d, WAM), G.A.H.; Warren River, 9.6 km
(6 miles) SE of Pemberton, 1.1971, (16, AM, 1d6, WAM), G.A.H. and H. Hughes;
Pemberton, xii.1936 (1d, ANIC), K.R.N.; Channybearup, near Pemberton, x.1970
(12, ANIC), D.H.C.; Pimelia, near Pemberton, x.1970 (12, ANIC), D.H.C.; 14.4
km (9 miles) W of Pemberton, x.1970 (1 Q, ANIC), D.H.C.; Porongurup National
Park, x.1970 (20, ANIC), D.H.C.; Mount Chudalup, S of Northcliffe, x.1970 (19,
ANIC), D.H.C.; Rest Point, Walpole, x.1970 (1 9, ANIC), D.H.C.; Nornalup
National Park, x.1970 (2d, 19, ANIC), D.H.C.
Comparative notes. T. paramonovwt is related to T. annulata and especially T. digztata
but differs from these in having a median grey stripe on the mesoscutum much as in T.
annulipes. It differs from T. annulzpes in the presence of hairs on the propleuron and
absence of paired horn-like tubercles on the epandrium. The shape of the surstylus is
distinct from all related species.
Paramonov (1955) referred specimens of this species doubtfully to T. fulva.
Tapeigaster digitata n. sp.
(Figs 13-17)
6 2. Coloration. Head dull fulvous; cheek and face paler; ocellar spot black.
Antenna black to blackish brown. Palpus pale fulvous; labella brown. Mesoscutum
brownish fulvous to tawny brown, entirely thinly greyish- to brownish-pruinescent,
somewhat shining, with pair of ill-defined whitish-pruinescent marks on anterior
margin; pleura pale fulvous to light brown, entirely densely whitish-pruinescent and
nowhere shining. Legs fulvous; femora blackish apically; fore femur with rather
broad brown to black longitudinal anterodorsal stripe from about basal quarter to
apex; tibiae each with brown to blackish ring at apex and before middle; tarsi
variably browned distally. Wing hyaline, tinged with greyish brown; haltere dull
yellowish. Abdomen brownish, often paler at base and apex, but variable, more
densely pruinescent in d than in 9, largely shining in latter sex.
Head with usually one distinct fronto-orbital ; vibrissa not duplicated.
Thorax. Propleuron with fine rather long hairs below humeral callus; presutural and
2 pairs of dorsocentral bristles present; prescutellar acrostichal bristle absent. Fore
coxae usually with pale hairs only, sometimes with one or 2 incipient bristles; fore
femur moderately stout, mid and hind femora relatively slender for genus and of
almost equal thickness; ventral spines of femora restricted to anteroventral and
posteroventral series but usually only the former present on hind femur; femora and
tibiae without long, dense ventral hairs in either sex. Wing with veins 3 and 4 apically
only very slightly convergent; vein 6 discontinued at a short distance from margin.
Abdomen. Sternite 5 of d divided medially into 2 plates, each of which is rather
rounded posteriorly with only slight indication of posteromedian angle. d
postabdomen with left spiracle 6 situated within lateral extremity of tergite; left
spiracle 7 situated higher up behind tergite 6; right spiracles 6 and 7 close together
and horizontally aligned in membrane; sternite 6 situated on lower part of left side,
with a narrow strip extending round ventral surface, connected to sternite 7 by a short,
narrow isthmus; sternite 7 more dorsally placed on left side, broadly continuous with
the large, dorsal sternite 8 at its dorsal extremity; epandrium without tubercles or
gibbosities, with well-developed free cerci and one pair of basally articulated surstyli a
short distance in front of them, surstylus forked into 2 subequal lobes, anterior lobe
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
46 SYSTEMATICS OF TAPEIGASTER
12
Figs 11-16. 11-12. Tapecgaster annulata. 11. 3 genitalia. 12. d 5th sternite.
13-16. T. digitata. 13. db genitalia. 14. d 5th sternite. 15. hypandrium. 16. wing of holotype.
obtuse, straight, posterior lobe more broadly obtuse slightly posteriorly curved;
hypandrium broad, with a series of very short hairs on each side, with pair of slender
anterior extensions which join together on median line to enclose a subcircular
membranous area, with a broad horizontal plate posteriorly on each side of aedeagus,
without distinct gonites; a pair of sclerites in the form of well-developed plates
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
D.K. McALPINE AND D.S. KENT 47
between bases of surstyli and posterior to hypandrium, embracing bases of cerci and
connected to each side of the rather small median transverse posthypandrial plate ;
aedeagus short, stout, curved forwards, with posterior surface membranous, with a
sclerotized skeletal strip on each side, and complex apical part.
Dimensions: total length, 5 4.8-6.4 mm, 2 4.9-6.1 mm; length of thorax, d 2.4-3.1
mm; length of wing, d 6.0-8.0 mm, 2 7.0-7.8 mm.
Distribution: New South Wales (principally tablelands); Australian Capital
Territory; Victoria; Tasmania; South Australia (Lofty Ranges and Kangaroo
Island).
Holotype 6 : below Govett’s Leap, Blue Mountains, 7.x.1956 (AM), D.K.M.
Paratypes. N.S.W.: Mount Kaputar, near Narrabri, xi.1964 (1 d, AM), D.K.M.;
Wright’s Lookout, New England National Park, iii-iv.1961 (6¢,592, AM, 3d, 39,
BM), D.K.M.; Point Lookout, New England National Park, x.1962 (2 3, ANIC),
D.H.C.; Barrington Tops via Salisbury, xii.1965 (3 d, 1 9, UQ), B. Cantrell;
Tubrabucca, near Barrington Tops, x.1956 (1 3,192, AM),D.K.M.; Mount Wilson,
Blue Mountains, 1.iii.iv.vi.x.xi.xii.1956-1977 (12d, 149, AM, 2d, 39, CNC,24, 3
Q, USNM), G.D., M. A. Schneider, D.K.M.; below Govett’s Leap, ix.1957 (2 6,
AM), D.K.M.; Katoomba, xii.1956 (1 d, AM), D.K.M.; Leura Falls, Blue
Mountains, 1.1973 (1 6, ANIC), D.H.C.; Wentworth Falls, Blue Mountains,
x1.xii.1956-59 (4 d, AM), D.K.M.; Woodford, Blue Mountains, xi.1925 (1 4,
ANIC), I. M. Mackerras; Royal National Park, x.1956-65 (20, AM), D.K.M.; Bola
Creek, Royal National Park, ix.1961 (1 9, ANIC), D.H.C.; Otford, Mlawarra
District, x.1957 (1d, 19, AM). D.K.M.; Mount Gibraltar, near Bowral 111.1975 (3
Q, ANIC), Z.R.L.; Lake George, xii.1950 (2 0, ANIC), K.R.N.; Clyde Mountain,
near Braidwood, ii.v.x.1960-65 (1d, AM, 5 d, ANIC), S. J. Paramonov, D.H.C.,
D.K.M.; 5-8 km (3-5 miles) S of Monga, near Braidwood, v.1968 (3d, 32, ANIC),
D.H.C. and Z.R.L.; Mount Jagungal, Snowy Mountains, 11.1951 (1 3d, ANIC), L.
Pryor; The Creel, Snowy Mountains, xi.1961 (2d, 22, ANIC), D.H.C.; Wilson’s
Valley, Snowy Mountains, 11.1963-1979 (1 d, 1 2, AM), D.K.M. and Bye:
Charlotte Pass, Snowy Mountains, 11.1963 (19, AM), D.K.M.; Leather Barrel Creek,
Snowy Mountains, ii.1979 (8d, 82, AM), B.J.D. and D.K.M.; Rutherford Creek,
Brown Mountain, near Nimmitabel, xi.1974 (6d, 39, AM), G.D. A.C.T.: Black
Mountain, Canberra, iv.x.1979-80 (1 d, 1 9, ANIC), D.H.C.; Blundell’s, near
Canberra, x.1930 (1d, ANIC), A. L. Tonnoir; Uriarra State Forest, iv.xi.1960-61 (1
6, 29, ANIC), D.H.C.; Bendora, 11.1950 (1 d, ANIC), K.R.N.; Condor Creek,
Brindabella Range, iv.1972 (1 d, AM), D.K.M.; ‘Piccadilly Circus’, Brindabella
Range, iv.1972 (69, AM), D.K.M.; Bull’s Head, Brindabella Range, iv.xi.1968 (3
3,12, ANIC), D.H.C.
Other material examined. Vic.: Mount Beauty, x.1961 (3d, 12, ANIC), D.H.C.;
Dynamite Creek, Bonang Highway, x.1961 (3d, ANIC), D.H.C.; Bonang Highway,
Bendoc Road Junction, xi.1976 (1 6, ANIC), D.H.C. and Z.R.L.; Nowa Nowa,
x.1961 (1 d, ANIC), D.H.C.; Noojee, xi.1964 (1 2, ANIC), D. E. Havenstein;
Mount Baw Baw, near Tanjilbren, 111.1964 (1 @, AM), G. L. Bush; Frenchman’s Gap
near Woods Point, iv.1963 (32, AM), D.K.M.; 20.8 km (13 miles) W of Matlock,
iv.1963 (1 2, AM), D.K.M.; Mount Dom Dom (Black Spur), near Healesville,
x.1961 (3d, 39%, ANIC), D.H.C.; 8 km SW of Narbethong, xii.1979 (1 3, ANIC),
K.R.N.; Cement Creek, near Warburton, iv.1963 (1 29, AM), D.K.M.; Warburton,
iv.1963 (2 3, 22, AM), D.K.M.; Sherbrook Forest, i.1966 (2d, AM), D.K.M.;
Belgrave, 1.1966 (1 9, AM), D.K.M.; near Melbourne, 11.1931 (1 d, AM), A.
Musgrave; Hobson’s Bay, Melbourne, xii.1922 (1 6, AM), A.F.B. Hull; Mount
Macedon, near Woodend, xii.1931 (5 d, 3 2, AM), A. Musgrave; Beech Forest,
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
48 SYSTEMATICS OF TAPEIGASTER
i.1967 (12, ANIC), Z.R.L.; Otway Range, 11 km W of Apollo Bay, xii.1978 (1d,
AM), G.D.; Mount William, Grampians, x.1977 (20,39, AM), D.K.M. and M. A.
Schneider. Tas.: Mount Barrow, near Launceston, 1.1960 (3 d, AM), D.K.M.;
Western Tiers, Lake Highway, 1.1960 (26, AM), D.K.M.; Marakoopa Caves, near
Mole Creek, 1.1960 (30,39, AM), D.K.M.; 19.2 km (12 miles) S of Wilmot, i.1960
(3d, AM), D.K.M.; 3.2km (2 miles) S of Oonah, Waratah Highway, 1.1960 (3d, 7
Q, AM), D.K.M.; Hellyer Gorge, Waratah Highway, i.1960 (4¢0,49, AM, 24,192,
ANIC), D.K.M., E. F. Riek; Meredith River, 19.2 km (12 miles) from Corinna,
1.1954 (12, ANIC), T. G. Campbell; 11.2 km (7 miles) W of Rosebery, ii.1963 (1
3d, 22, ANIC), I.F.C. and M.S.U.; Lake Saint Clair, 1.1960 (4 d, 3 2, AM),
D.K.M.; Franklin River crossing, Lyell Highway, 1.1960 (13d, 5 Q, AM), D.K.M.;
near Russell Falls, Mount Field National Park, 1.11.1960-63 (1 2, AM, 2d, 1 @,
ANIC), D.K.M., D.H.C.; Mount Wellington, ii.1963 (2 5d, 6 9, ANIC), D.H.C.;
Cambridge, x.1965 (10, ANIC), K. L. Taylor; Myrtle Gully, iii.1935 (12, ANIC),
W. Rafferty. S.4.: Engelbrook Reserve, near Bridgewater, iv.1967 (1d, 192, AM),
D.K.M.; Ravine des Casoars, Kangaroo Island, xi.xii.1977 (5 d, AM), M. A.
Schneider and D.K.M.
Comparative notes. T. digttata forms with T. annulata and T. paramonow a group of
three closely related species characterized by the haired propleuron, absence of strong
thickening of the mid and hind femora, and incomplete vein 6. T. digitata differs
from T. annulata in its more densely pruinescent thorax, which generally has a less
definite orange hue, in having the dark distal zone of the fore femur extended basad as
an anterodorsal stripe, in having the lobes of sternite 5 much blunter posteriorly, the
lateral hairs of the hypandrium much shorter, and the surstylus not distinctly forked.
For comparison with T. paramonow see under that species.
Tapergaster annulata (Hendel), n. comb.
(Figs 11-12)
Scromyzoptera annulata Hendel, 1917: 47.
Tapeigaster fulva Malloch, 1926: 553. N. syn.
To Malloch’s description we add the following:
Coloration generally of a brighter orange tone than in T. dzgztata. Thoracic pleura a
little less thickly whitish-pruinescent than in that species with mesopleuron somewhat
shining. Fore femur fulvous, dark brown to black only on apical quarter or less.
Thorax. Propleuron with fine hairs below humeral callus; presutural bristle present ;
prescutellar acrostichal bristle absent.
S postabdomen as described for T. digztata with the following notable differences:
lobes of sternite 5 more narrowly produced and subacute; surstylus inserted very close
to cercus, not forked, very broad basally, contracted into a narrow, straight,
posteriorly directed distal part with attenuated apex; hypandrium with a series of
conspicuously long hairs, almost as long as surstylus, on each side.
Distribution: Queensland — extreme south-east; New South Wales — coast and
wetter parts of tablelands districts; recorded in error by Paramonov (1955) from
Australian Capital Territory, Victoria, and Tasmania.
Type material examined : ‘N. Holl. 878° (i.e. Australia) no other data (holotype d of
Sciomyzoptera annulata Hendel, WM); Botany Bay, N.S.W., no date (holotype 6,
paratype G, latter damaged, of T. fulva Malloch, USNM), H. Petersen.
Other material examined (localities only given). Q/d.: Binna Burra, Lamington
National Park (AM). N.S.W.: Mount Gibraltar National Park, 102.4 km (64 miles)
W of Grafton (AM); Ulong, East Dorrigo district (AM); Dorrigo (ANIC); New
Proc. LINN. Soc. N.S.W., 106 (1), (1981) 1982
D.K. McALPINE AND D.S. KENT 49
Figs 17-21. 17. Tapeigaster digitata, head of holotype. 18-21. T. swbglabra. 18. head of holotype. 19. wing
of holotype. 20. db genitalia. 21. hypandrium.
England National Park (AM, ANIC, UQ); Upper Allyn, near Gresford (ANIC) ;
Wallaroo State Forest, near Karuah (ANIC); 46.7 km (29 miles) NW of Putty
(AM); Mount Wilson, Wentworth Falls, and Springwood, Blue Mountains (AM) ;
Mooney Mooney Creek, near Gosford (AM); Middle Creek, near Narrabeen
(ANIC); Bronte, near Sydney (AM); Royal National Park (AM, ANIC); Otford,
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
50 SYSTEMATICS OF TAPEIGASTER
Illawarra district (AM, ANIC) ; Cataract Creek, Bulli (ANIC) ; Minnamurra Falls,
near Kiama (AM); Kangaroo Valley (ANIC); Clyde Mountain, near Braidwood
(ANIC).
Paramonov (1955) misidentified T. fulva, most of his specimens being referable
to T. digitata and one to T. subglabra. Bezzi placed S. annulata as a synonym of T.
annulipes but examination of the holotype shows this to be incorrect.. T. annulata is
most closely related to T. dzgztata, q.v. for comparison.
Tapergaster pulverea n. sp.
(Figs 22-23)
3 2. Coloration. Head fulvous to tawny; cheek and face much paler; ocelli
surrounded by a rather large dark brown spot; orbital margins of postfrons with
creamy-white pruinescence; vertex with V-shaped creamy-white pruinescent mark
pointing posteriorly visible from some angles, its arms passing to each side of
postvertical pair of bristles; upper occiput with brownish patch of varying intensity on
each side; extremity of vibrissal angle brown. Antenna dark brown; base of segment 3
reddish brown. Palpus pale fulvous; labella brown to tawny. Thorax with tawny
ground colour; mesoscutum with pair of whitish-pruinescent paramedian
longitudinal stripes which are very marked anteriorly but gradually become narrower
and indistinct posteriorly; whitish-pruinescent spots also present sublaterally, one
before and one behind transverse suture, one between supra-alar and intra-alar
bristles, one on scutellar bridge; a whitish suffusion on humeral callus and on
notopleuron; scutellum with whitish spot anteriorly on each side; pleura with brown
spot on upper posterior part of mesopleuron and sometimes with additional brownish
suffusion, rather extensively whitish-pruinescent. Fore coxa pale yellowish; other
coxae yellowish to tawny; femora fulvous, brown apically and less intensely so near
middle; tibiae fulvous, each browned at apex, near middle, and, less extensively so, at
base; tarsi fulvous, becoming brownish apically. Wing membrane with faint, almost
uniform yellowish brown tinge. Haltere pale yellowish. Abdomen tawny, sometimes
with variable brown suffusion.
Head with usually one strong upper fronto-orbital bristle, occasionally another weaker
one in front of it; one or 2 setulae of peristomial series just below vibrissa stronger than
others and usually at least half as long as vibrissae.
Thorax. Propleuron pruinescent, without hairs; presutural and 2 pairs of
dorsocentral bristles present; prescutellar acrostichal bristle absent. Fore coxa with
bristles all rather weak and hair-hke, but quite long in 6: fore femur much stouter
than other femora, with ventral spines numerous and not entirely seriate; other
femora with ventral spines weaker and less numerous; in 6 femora and tibiae with
numerous long, fine ventral hairs. Wing with veins 3 and 4 very slightly converging
apically; vein 6 shorter than in other species, discontinued at about 0.8 of distance
from anal crossvein to margin (measured in direction of vein 6); anal crossvein
sloping a little more towards base at posterior end than in other species, posterodistal
angle of anal cell thus a little more obtuse.
Abdomen. Sternite 5 of d not divided medially. d postabdomen somewhat resembling
that of T. digztata; right spiracle 7 not found, possibly absent; surstylus slender
beyond the thickened base, falcate, tapering to the narrow obtuse apex, with rather
long hairs anteriorly near base; hypandrium with hairs of moderate length in a series
on each side; posthypandrium plate much larger than in T. dzgztata and related
species, touching hypandrium anteriorly on each side and connected to basal plates of
surstyli posteriorly.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
D. K. McALPINE AND D.S. KENT
j mh
ls
Figs 22-26. 22-23. Tapeigaster pulverea. 22. head of holotype. 23. d genitalia.
24. T. nigricornis, larval cephalopharyngeal skeleton. dc = dorsal cornu. es = epistomal sclerite. hs =
hypostomal sclerite. ls = ligulate sclerite. mh = mouth-hook. pb = parastomal bar. ps = pharyngeal
sclerite. ve =
ventral cornu. 25-26. T. annulipes. 25. puparium. 26. pupal respiratory horn. is = inner
spiracles. lc = larval cuticle (puparium wall). pe = pupal cuticle.
Dimensions: total length, d 4.2-5.9 mm, 2 4.7-5.0 mm; length of thorax, d 2.2-3.0
mm; 2 2.5-2.9 mm; length of wing, d 5.1-6.5 mm, 9 5.5-6.3 mm.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
5]
52 SYSTEMATICS OF TA PEIGASTER
Distribution: Queensland — southern border districts; New South Wales — coast and
tablelands; Victoria.
Holotype 3 : Royal National Park, near Sydney, 14.x.1956 (AM), D.K.M.
Paratypes (all from N.S.W.) : same data as holotype (1 Q, AM); Mount Wilson, Blue
Mountains, i.i1i.1958-1976 (1d, ANIC, 2d, AM), E. F. Riek, J. J. T. Evans, M. A.
Schneider and D.K.M.; Wentworth Falls, Blue Mountains, xi.xil.1956-1958 (2 d,
AM), D.K.M.; Sassafras Gully, Springwood, x1.1956 (1 Q, AM), D.K.M.; Seven
Mile Beach State Park, near Gerroa,, v.1976 (29, AM) G.A.H.; 24 km (15 miles)
SSE of Braidwood, ix.1956 (1d, ANIC), I.F.C.
Other material examined. Qld. : Mitchell Gully, 3.2 km (2 miles) E of Cunningham’s
Gap, vi.1966 (1 3, ANIC), Z.R.L.; Lamington National Park, x.1934 (1 Q, UQ), F.
A. Perkins; Binna Burra, Lamington National Park, v.1964 (1 9, UQ), B. Genn.
Vic. : Powelltown, 60 km E of Melbourne, x.1961 (12, ANIC), D.H.C.
Comparative notes. T. pulverea is not very close to any other species of the genus,
differing from all in having a rather strong bristle just below the vibrissa. In the
hairless prosternum it resembles T. subglabra, but it differs from that species in the
thicker fore femur, shortened vein 6, more extensively haired mesoscutum, and
presence of the presutural bristle.
Tapezgaster subglabra n.sp.
(Figs 18-21)
3 2 Coloration. Head tawny; cheek and face paler; postfrons often with two
longitudinal brown stripes inside orbital margins (varying in intensity); ocelli
surrounded by a dark brown spot; anterior region of orbital margins of postfrons with
white pruinescence. Antenna dark brown. Palpus and labella tawny to fulvous.
Thorax tawny; mesoscutum with faint median longitudinal brown stripe, fading
posteriorly to suture; lateral margins of mesoscutum dark brown, extending to wing
bases (seen in lateral view as a stripe tapering posteriorly) ; scutellum tawny; pleura
tawny, with a faint band of brown tapering anteriorly, running above coxae;
prosternum pale fulvous. Fore coxa pale tawny, other coxae tawny; femora tawny,
browned apically; fore femur also with longitudinal dorsolateral brown stripe, fading
basally, on inner surface; tibiae tawny, each browned at apex and slightly at basal
joint; tarsi tawny, becoming brownish apically. Wing membrane with uniform faint
yellowish brown tint. Haltere with brown capitellum, otherwise fulvous. Abdomen
tawny to brown.
Head with two strong upper fronto-orbital bristles, without strong bristle behind
vibrissa.
Thorax. Presutural bristle absent; two pairs of dorsocentral bristles present;
prescutellar acrostichal bristles absent. Propleuron without hairs below humeral
callus; prosternum hairless; fine hairs on mesoscutum reduced to pair of acrostichal
series, pair of dorsocentral series, and some irregularly placed lateral ones. Fore coxae
with a row of bristles along the inner basal edge; mid and hind coxae with a row of
bristles along the anterior basal edge; femora only slightly thickened; fore and mid
femora slightly stouter than hind femur; ventral spines of femora restricted to
anteroventral and posteroventral series, with a reduction in spines from fore to hind
femur; femora and tibiae with numerous fine hairs. Wing with vein 6 pigmented all
the way to the wing margin; veins 3 and 4 not diverging distally, almost parallel.
Abdomen. Sternite 5 of 5 not divided medially. 6 postabdomen: cercus short and
small; surstylus large in comparison, broadly L-shaped, its basal section with
numerous long hairs along ventral margin, apical section obtusely rounded.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
D. K. McALPINE AND D. S. KENT 53
Hypandrium broad anteriorly, with row of long incurved hairs along inner anterior
margin, with a broad plate posteriorly on each side of aedeagus; anterior margin of
plate extended into a finger-like projection bearing two incurved hairs.
Dimensions: total length, 3 4.6-6.5 mm, 2 5.0-7.0 mm; length of thorax, dS 2.2-3.0
mm, 9 2.2-2.8mm; length of wing, d 5.7-7.3 mm, 2 6.0-7.4mm.
Distribution: New South Wales — principally tableland districts; Australian Capital
Territory; Victoria.
Holotype 3 : Mount Wilson, Blue Mountains, 5.x.1957 (AM), D.K.M.
Paratypes. N.S.W.: Wright’s Lookout, New England National Park, iv.1961 (1 6,
AM), D.K.M.; Coachwood Gully, 21.7 km SE of Three Ways, xii.1977 (1 Q, AM),
G.D.; Tubrabucca, near Barrington Tops, x.1956 (1 d, AM), D.K.M.; Mount
Wilson, Blue Mountains, i.iii-iv.v.vi.viii.ix.x.xi.1957-1980 (27d, 289 AM, 24,192,
AINE, 2G, BQ, TIME, er, We ING, Bos 4. USINIMD), IDI (Ce, GzlDg, Gosia, Se 2.
Simba) he Vien View ACs Schncidens AnG i. lees Springs. ive l958) (a 3, ANIC)
Z.R.L.; Mount Coree, x.1961 (1 d, ANIC), D.H.C.; Bull’s Head, Brindabella
Range, iv.1958 (2d, ANIC), D.H.C.
Other Material Examined. Vic.: Bonang Highway, Bendoc Rd. Junction, 10 km S of
Bonang, xi.1976 (20,29, ANIC), D.H.C. and Z.R.L.; Mount Beauty, x.1961 (19,
ANIC), D.H.C.; Mount Donna Buang, iv.1963 (1 Q, AM), D.K.M.; Warburton,
iv.1963 (30,392, AM), D.K.M.; Belgrave, 1.1966 (1d, AM), D.K.M.
Comparative notes. T. subglabra is not very close to any other species of the genus,
differing from all in the absence of presutural bristles and reduction of hairs on the
mesoscutum. In the hairless prosternum it resembles T. pulverea, but differs from that
species in other characters, as indicated in the description of T. pulverea.
MORPHOLOGY OF EARLY STAGES
The morphology of the early stages of Heleomyzidae s.1. is too poorly known to
enable any broad taxonomic inferences to be drawn from the new data. Two points
may be made here.
The larva of Tapeigaster differs greatly from that of Cazrnsizmyza (McAlpine,
1968) although the two genera have been considered to be related (Griffiths, 1972).
We are unable to find any points of resemblance which are not shared with larvae of
numerous other schizophoran families. The differences in the facial organs and form
of the terminal spiracle-bearing segment are particularly marked.
The pupa has a prothoracic respiratory horn which penetrates the puparium and
is thus visible externally. Such a process is frequently present in muscoid (calyptrate)
pupae, but is apparently known only in certain north-temperate heleomyzids among
the acalyptrates. According to Hennigian reasoning the presence of such a
plesiomorphic character cannot be employed as evidence for close phylogenetic
relationship between the tribes Heleomyzini and Tapeigastrini. Nevertheless it may be
statistically significant that the only known retention of this kind of respiratory horn
among non-muscoid Schizophora is in groups which, from other characters, seem
referable to the family Heleomyzidae.
Tapeigaster annultipes
Larval material of T. annulpes was obtained from an adult female collected at
the Australian Museum, 24 April 1980. The fly laid eggs on a cultivated mushroom
(Agaricus sp.). Of the subsequent larvae raised, some were preserved in alcohol for
later dissection and the rest left to pupate. Adults failed to emerge so the remaining
puparia were either dry mounted or placed in alcohol. It was from this latter material
that the detail of the pupal respiratory horn was obtained.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
54 SYSTEMATICS OF TA PEIGASTER
Figs 27-32. Tapeigaster annulipes, larva. 27. entire larva in lateral view. 28. facial region. 29. anterior
spiracle. 30. posterior view. 31. posterior spiracle. 32. cephalopharyngeal skeleton, with ventral view of
epistomal sclerite and dorsal view of hypostomal sclerite and ligulate sclerites. dec = dorsal cornu. ds =
dentary sclerite. es = epistomal sclerite. hs = hypostomal sclerite. Is = ligulate sclerite. mh = mouth-
hook. pb = parastomal bar. ps = pharyngeal sclerite. ve = ventral cornu.
Last instar larva (Figs 27-32) creamy-white, somewhat elongate, circular in cross-
section; anterior end tapered; posterior three quarters almost uniform in width;
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
D.K. McALPINE AND D.S. KENT 55
posterior end truncate and sloping (Fig. 27). Segment 1 (Fig. 28) divided by a
median groove; each lobe with a short, pale yellowish-brown, 2-segmented antennal
papilla dorsoapically and a circular sensory plate ventroapically; a smaller third pair
of papillae located on each side of atrial opening. Facial area with numerous parallel
and branching, minutely serrated, sclerotized ridges posteriorly, and with numerous
cuticular teeth anteriorly and on each side of atrial opening, the latter of 4 different
types: the foremost ones (type 1) consisting of numerous small subglobular serrated
plates occupying a narrow transverse band from each side of anterior extremity of
atrial opening to lateral extremity of atrial opening to lateral extremity of facial
region; teeth of type 2 lying across anterior extremity of atrial opening and extending
into it, more elongate than those of type 1 with fewer, longer cusps, the inner ones
simply conical or unicuspid ; teeth of type 3 large and few, lying behind third pair of
papillae, each with 6-10 cusps; a triangular sheath-like structure located at posterior
margin of the atrial opening projecting into the opening. Segment 2 bearing paired,
light yellowish-brown, fan-shaped anterior spiracles on short processes postero-
laterally, each bearing 10-14 short apical papillae; a yellowish-brown sclerotized ring
encircling base of each process (Fig. 29). Segments 3-12 bearing narrow encircling
bands of unpigmented spicules anteriorly. Segments 5-12 bearing ventral creeping
pads on anterior margins, each pad bearing unpigmented spicules arranged in short
parallel rows, directed both anteriorly and posteriorly. Segment 12 (Fig. 30) bearing a
pair of spiracular tubercles and five pairs of papillae. Posterior spiracles set in a
concave depression, with 4 pairs of papillae encircling depression on outer rim, and a
fifth and largest pair ventrally directed and associated with the anal pore. Posterior
spiracular plates (Fig. 31) yellowish brown, located on separate, short tubercles; each
plate with 3 narrowly oval apertures at about 65° to each other, containing lamellae
just within, and with 4 colourless, much branched interspiracular hydrophobe hairs; a
yellowish brown sclerotized subcuticular ring below _ spiracular plate.
Cephalopharyngeal skeleton (Fig. 32) black to dark brown. Mouth-hooks well
developed, paired, separate, without accessory teeth below the curved apical section,
with small window in posteroventral corner and distinctive hook-like nodules on the
posterodorsal margin of the basal sections. Dentary sclerites paired, separate, near
posteroventral margin of mouth-hooks. Epistomal sclerite not fused to parastomal
bars, largely dark-pigmented posteriorly, lighter on anterior median process and
margin, with pair of narrow rod-like sclerites extending posteriorly to fuse with inner
posterodorsal margin of hypostomal sclerite. Parastomal bars narrow, dilated
anteriorly, darkly pigmented, separate from upper margin of hypostomal sclerite and
fused posteriorly with pharyngeal sclerites. Hypostomal sclerite H-shaped, not fused
posteriorly to pharyngeal sclerites; anterior rami shorter and broader than posterior
rami; hypostomal bridge darkly pigmented on all but narrow posterior band. Ligulate
sclerites curved, paired and separate; situated anterior to hypostomal sclerite and
between posterior margins of mouth-hooks. Pharyngeal sclerites with anterodorsal
bridge joining anterior ends of dorsal cornua, bridge with several windows medially;
with antero-ventral projections lying below posterior rami of hypostomal sclerite;
dorsal cornua narrower and slightly longer than heavily pigmented part of ventral
cornua, which are lightly pigmented posteroventrally. Pharyngeal ridges between
ventral cornua well developed but lacking pigmentation.
Length 10-11 mm.
Puparium (Fig. 25) light reddish brown, darker at each end, elongate-ovoid, slightly
more curved dorsally than ventrally; posterior end rounded, with posterior spiracular
tubercles heavily pigmented and surrounded by concentric ridges; anal plate
surrounded by a narrow band of ridges; anterior end tapered in lateral view; anterior
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
56 SYSTEMATICS OF TAPEIGASTER
spiracles heavily pigmented and compressed, but otherwise as in larva; thoracic
segments 1 and 2 with prominent encircling ridges; thoracic segment 3 with slightly
less distinct ridges restricted to anterior margin; abdominal segment 1 (segment 4)
bearing on its dorsal surface the paired, brown outer pupal respiratory horns, which
project through the puparium wall (as in Musca). Outer respiratory horn (Fig. 26)
with heavily sclerotized wall and obliquely aligned pores, arising from a short
outgrowth of the dorso-lateral edge of the precursor prothorax of the pupa; this
outgrowth also bearing inner bilobed spiracles on posterior surface; spiracles slightly
domed, with oval pores, divided medially by a bulbous process, opening into cavity
between puparium wall and pupal cuticle, each subtended by a crescent-shaped
sclerite.
TABLE 1.
Host Records of Tapezgaster
Species Host Records
Adults Reared From: Adults Collected On:
T. nagricornis ‘Pleurotus lampas’ = Pleurotus nidiformis Berk. ‘Pleurotus lampas’
Amanita grisea Mass. & Rodw.
Amanita ochrophylla (Cke. & Mass.) Clel.
Boletus luridus Schaeff. ex Fr.
‘Boletus granulatus’ = Suzllus granulatus
(L. ex. Fr.) O. Kuntze
‘Boletus portentosus’ = Phaeogyroporus
portentosus (Berk. & Br.) McNabb.
Cortinarius brunneus (Pers. ex. Fr.) Fr.
Agaricus sp.
Boletus sp.
“Puff ball fungus’
T. annulipes ‘Psalliota campestris’ = Agaricus campestris L. ‘Pleurotus lampas’
ex Fr.
T. digitata ‘fungi’
T. cinctipes ‘Pleurotus lampas’
T. paramonow ‘mushrooms’
T. argyrospila ‘Puff ball fungus’
T. brunneifrons ‘fungi’
Tapergaster nigricornis
A similarly detailed study of T. négr¢cornis larvae was not possible as the available
material consisted of only two larvae (possibly last instar), collected ex fungus from
Broulee, N.S.W., 6 May 1980 by Z.R.L. (ANIC). Only one specimen was dissected,
the other was left entire.
Larvae of T. nigricornis differ little from those of T. annulipes. The minor
differences evident are as follows: (a) in facial area, the patterns of cuticular teeth
differed slightly in number and arrangement, i.e. teeth of type 2 are all unicuspid and
extend into the atrial opening in only a single row; (b) the anterior spiracle, is more
broadly fan-shaped, bearing only 10 papillae; each papilla has a single pore opening
into a roughly circular cavity lined by tear-shaped protuberances. The major
difference between the larvae is in the cephalopharyngeal skeleton; the mouth-hook
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
D. K. McALPINE AND D. S. KENT 57
has an elongate, acute, inwardly directed process on the posterodorsal surface; the
mouth-hook has a large window in the basal section and a lightly pigmented anterior
region. The parastomal bar is parallel-sided along its entire length with slight upward
curve anteriorly. The dentary sclerite is absent in both specimens. The ligulate sclerite
is large, with incurving dorsal bulge; this sclerite is visible in lateral as well as ventral
view (Fig. 24).
A comparison of puparia and pupae showed that those of T. negricornis differed
little from those of T. annulzpes.
NOTES ON BIOLOGY
Adults of Tapezgaster annulitpes are often observed on fruiting bodies of agaric
and other fungi, and other species of the genus have either been observed by the
authors on fungi or recorded on labels as inhabiting fungi. These species are T.
nigricornis, T. brunnetfrons, T. cinctipes, T. annulipes, T. digitata, T. paramonovi
and T. argyrospila. In the case of T. annulipes males frequently take a position on top
of the pileus and defend the position from rival males. Females approaching and
flying beneath the pileus (perhaps for oviposition) have been seen to be followed by
the male and mounted, sometimes after a struggle, but no time was given to courtship
in such instances.
Adults of Tapezgaster species are often swept from ferns and low foliage during
insect-collecting activities. They are rarely found at night, perhaps because they are
mainly diurnal, and they do not habitually settle on tree trunks.
Larvae of at least five species of Tapezgaster live in fruiting bodies of fungi. T.
annulipes and perhaps all or most other species can be reared in the laboratory on
commercial mushrooms, but they are not considered to be pests of this crop. The large
number of mushroom spores in facial grooves of preserved T. annulipes larvae suggest
that they had been feeding particularly on the gills. Presumably the fungi on which
the adults are found are normally those that the larvae infest. Table 1 gives the host
records available to us.
ACKNOWLEDGEMENTS
We are indebted to Dr D. H. Colless, C.S.I.R.O. Division of Entomology,
Canberra, and Miss M. A. Schneider, Department of Entomology, University of
Queensland, St Lucia, for loans of material. Mr B. H. Cogan, British Museum
(Natural History), London, Dr R. Contreras-Lichtenberg, Natural History Museum,
Vienna, Dr W. Mathis, National Museum of Natural History, Washington, D.C., and
Dr A. Neboiss, National Museum of Victoria, Melbourne, supplied information on
types.
References
Bezzi, M., 1923. — Note on the Australian genus Tapezgaster Macq. (Diptera) with descriptions of new
species. Aust. Zool. 3: 72-78.
Co..ess, D. H., and McAtpine, D. K., 1970. — Chapter 34. Diptera. In The znsects of Australia: 656-740.
Melbourne: Melbourne University Press.
GrirFiTHs, G. C. D., 1972. — The phylogenetic classification of Diptera Cyclorrhapha with special
reference to the structure of the male postabdomen. The Hague: W. Junk.
HENDEL, F., 1917. — Beitrage zur Kenntnis der acalypraten Musciden. Dé. ent. Zettschr., 1917: 33-47.
HEeNNic, W., 1958. — Die Familien der Diptera Schizophora und ihre phylogenetischen
Verwandtschaftsbeziehungen. Beztr. Ent. 8: 505-688.
Macquarr, P. J. M., 1847. — Diptéres exotiques, supplément 2: 104 pp. 6 pl.
——, 1851. — Diptéres exotiques, supplément 4 (pt. 2) : 161-364, pl. 15-28.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
58 SYSTEMATICS OF TA PEIGASTER
MALLocH, J. R., 1926. — Notes on Australian Diptera. No. ix. Proc. Linn. Soc. N.S.W. 51: 545-554.
——, 1927. — Notes on Australian Diptera. No. x. Proc. Linn. Soc. N.S.W. 52: 1-16.
, 1935. — Notes on and descriptions of new species of Australian Diptera. Aust. Zool. 8: 87-95.
McALPINnE, D. K., 1968. — The genus Cazrnsemyza Malloch (Diptera, Heleomyzidae, Rhinotorini). Rec.
Aust. Mus. 27: 263-283.
PARAMONOY, S. J., 1955. — Notes on Australian Diptera (xx). A Review of the genus Tapezgaster Macq.
(Neottiophilidae, Acalyptrata) , Diptera. Ann. Mag. nat. Hist. (12) 8: 453-464.
WALKER, F., 1853. — Insecta Saundersiana. Diptera 4: 253-414.
——, 1858. — Characters of undescribed Diptera in the collection of W. W. Saunders, Esq., F.R.S. & c
(Continued). Trans. ent. Soc. Lond. (2) 4: 190-235.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
A Comparative Morphological Study of the
Reproductive Systems of some Species of
Tapeigaster Macquart (Diptera: Heleomyzidae)
MARGARET A. SCHNEIDER
(Communicated by D. K. McALPINE)
SCHNEIDER, M. A. A comparative morphological study of the reproductive systems of
some species of Tapeigaster Macquart (Diptera: Heleomyzidae). Proc. Linn.
Soc. N.S.W. 106 (1), (1981) 1982: 59-65.
The female reproductive systems of Tapezgaster annulipes Macquart, T.
annulata (Hendel), T. swbglabra McAlpine and Kent, T. dzgztata McAlpine and
Kent, T. pulverea McAlpine and Kent, and T. lutezpennzs Bezzi, and the male re-
productive systems of T. annulata, T. lutecpennis, T. pulverea, T. annulipes and T.
subglabra are discussed and illustrated either fully or in part to show specific
differences.
Margaret A. Schneider, Department of Entomology, University of Queensland, St
Lucia, Australia 4067; manuscript received 22 July 1981, accepted for publication 23
September 1981.
INTRODUCTION
Alcohol-preserved specimens of six species of Tapeigaster Macquart
(Heleomyzidae) were dissected to investigate whether the male and female
reproductive systems would show differences at the species level and similarities which
could be considered generic characters. There has been little published on either the
male or female reproductive systems of other genera of Heleomyzidae for comparison.
As in many families of the Schizophora, Heleomyzidae appear to have a groundplan
number of three spermathecae, with the occasional increase to four or decrease to two
being apomorphic (Hennig, 1958). In his investigations Sturtevant (1926) found
three spermathecae in all the genera except one, in which there were four. The
females of all six species of Tapezgaster which were dissected in the present study have
only two spermathecae and Hennig found only two in T. margznzfrons Bezzi (= T,
nigricornis (Macquart) ). Further comparative studies are needed to determine how
often this reduction in the number of spermathecae occurs in Australian genera of
Heleomyzidae. In two other endemic genera so far examined, only two spermathecae
were found in each of the two species of Austrolerxa McAlpine dissected while three
were found in each of the three species of the closely related genus, Diplogeomyza
Hendel. It is interesting to note, however, that in Diplogeomyza the two spermathecae
on the same side share a common duct throughout its length and are very closely
associated. The shape and overall appearance of the spermathecae of Diplogeomyza
spp. and Austroleria spp. are somewhat similar and very different from the simple,
spherical form on a long duct as found in Tapezgaster spp.
The females of Tapezgaster show only slight specific differences such as the
relative positions of the spermathecal and accessory gland duct openings into the
vagina, the relative size of the structures concerned and the presence or absence of a
sclerite in the wall of the vagina in the region of the spermathecal and accessory gland
duct openings. Differences between species are more obviously shown in the male
reproductive systems, with shape of testes and length and form of accessory glands and
the ejaculatory duct being useful characters. Male specimens of T. dzgztata McAlpine
and Kent were not available for dissection.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
60 REPRODUCTIVE SYSTEMS OF SOME TA PEIGASTER
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
M. A. SCHNEIDER 61
FEMALE REPRODUCTIVE SYSTEM
Tapergaster annulipes Macquart
(Fig. 1)
The ovaries, which occupy the greater part of the abdominal cavity in a gravid
female, end in short lateral oviducts which meet to form a muscular, median oviduct.
There are two spherical, brown-black spermathecae, one on each side; each
spermatheca is surrounded by a white fat mass which, in the undisturbed state, is
embedded in the tissue surrounding the ovary on the same side. The spermathecal
ducts are long, thin, relatively stout-walled and darkly sclerotized at the distal ends;
they enter the vagina at the midline. Each of the two accessory glands is an elongate,
pale, thin-walled, semi-transparent structure embedded in tissue adjacent to the
spermatheca on the same side. The distal end of each accessory gland duct, adjacent
to the gland, is somewhat expanded and muscular, presumably with a sphincter
function; the accessory gland ducts enter the vagina at the middle immediately
posterior to the spermathecal ducts. The vagina, or posterior part of the median
oviduct, is slightly expanded, with thick muscular walls in the region of these ducts,
and a small dark sclerite lies in the ventral wall. No morula gland or ventral receptacle
is present. Strong muscle bands originating from the posterior end of the vagina and
inserted in the wall of the vagina just anterior to the spermathecal ducts had to be cut
to expand the vagina to its full length.
Tapergaster lutetpennis Bezzi
(Fig. 2)
Differs from T. annulipes as follows. Median oviduct shorter, about three-fifths
length of that in T. annulzpes. Spermathecal ducts pale yellow along most of length,
colour slightly darker at proximal ends where they enter the vagina a little lateral to
midline on each side. In this region, vagina bulbous with greatly thickened muscular
walls and internal yellow, thickened supporting bands associated with openings of the
ducts. Accessory glands long, as in T. annulzpes, but distinctly bulbous distally (Fig.
2). Accessory glands ducts more expanded distally, entering vagina immediately
posterior to spermathecal duct on each side.
Tapezgaster annulata (Hendel)
Fig. 3
Differs from T. annulipes as follows. Median oviduct short, about half length of
that in T. annulipes. Accessory glands and muscular distal ends of accessory gland
ducts much shorter but otherwise accessory gland ducts a little longer than in T.
annulipes. Spermathecal and accessory gland ducts meeting vagina lateral to midline
as shown in Fig. 3. Nosclerite in ventral vaginal wall.
Tapergaster pulverea McAlpine and Kent
(Fig. 4)
Resembles most closely T. annulata but differs as follows. Vagina in region of
spermathecal duct openings more bulbous with thicker muscular walls. Accessory
Figs 1-5. Female Tapezgaster spp.: (1) T. annulipes, reproductive system; (2) T. lutezpennis, accessory
gland; (3) T. annulata, reproductive system, ovaries not shown; (4, 5,) proximal ends of spermathecal and
accessory gland ducts: (4) T. pulverea; (5) T. digitata. ac = accessory gland, acd = accessory gland duct, c
= cercus, ft = fat mass, lo = lateral oviduct, m = cut muscle band, mo = median oviduct, ov = ovary, sp =
spermatheca, spd = spermathecal duct, T9 — tergite 9, v — vagina.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
62 REPRODUCTIVE SYSTEMS OF SOME TA PEIGASTER
~aC
vd—
05mm
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
M. A. SCHNEIDER 63
glands and ducts smaller and shorter than in 7. annulata, entering vagina distinctly
lateral to spermathecal ducts (Fig. 4).
Tapergaster subglabra McAlpine and Kent
Only specimen of this species available for dissection proved to be very immature
with ovaries not developed and accessory glands small and very thin-walled. Sclerite in
ventral wall of vagina not present. Sclerotization of distal ends of spermathecal ducts
absent, these ducts meeting vagina as in T. annulipes but accessory gland duct
appearing to meet vagina lateral to midline.
Tapeigaster digitata McAlpine and Kent
(Fig. 5)
Although generally agreeing with description given for T. annultpes, differs as
follows. Spermathecae mid-brown and spermathecal ducts meeting vagina lateral to
midline (Fig. 5). Accessory glands about three-fourths length of those of T.
annulipes, pale yellowish rather than white and not as thin-walled.
MALE REPRODUCTIVE SYSTEM
Tapezgaster annulata
(Fig. 6)
The testes are golden brown, apically tapered, elongate sacs. The stout, golden-
brown vasa deferentia narrow proximally where they enter the apex of the pale-
coloured ejaculatory duct. This duct is at first relatively stout and thick-walled but
abruptly narrows about half way along its length. Posteriorly it passes into the
ejaculatory bulb which has articulated with it, the muscle-covered ejaculatory
apodeme. A narrow, strong duct connects this with the aedeagus to which is
articulated the large, flat, muscle-covered aedeagal apodeme. The accessory glands
which enter the apex of the ejaculatory duct ventral to the vasa deferentia are long,
pale yellow and only slightly expanded apically.
Tapeigaster annultpes
(Fig. 7)
Generally similar to T. annulata but differs as follows. Testes (Fig.7) shorter and
more expanded basally. Vasa deferentia a little longer. Accessory glands about half as
long as those of T. annulata and almost as thick as distal portion of ejaculatory duct,
which is wider and longer than in T. annulata.
Tapeigaster luterpennis
Specimens of this species available for dissection were immature so it 1s difficult to
make accurate comparisons with other species. Testes small but of similar shape to
those of T. annulipes except not so markedly expanded basally and more rounded
apically. Ejaculatory duct wide distally as in T. annulipes but accessory glands in
specimens examined long and narrow, about half width of ejaculatory duct, as in T.
annulata.
Fig. 6-7. Male Tapezgaster spp.: (6) T. annulata, reproductive system; (7) T. annulipes, testis and vas
deferens. aa = muscle-covered aedeagal apodeme, ac = accessory gland, ae = aedeagus, ea = muscle
covered ejaculatory apodeme, ej = ejaculatory duct, t = testis, vd = vas deferens.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
64 REPRODUCTIVE SYSTEMS OF SOME TA PEIGASTER
Tapeigaster pulverea
Appears most similar to T. annulata although once again available specimens
seemed immature judging from small size of testes, especially in one specimen in which
there was also very little pigmentation of testes and vasa deferentia. Testes relatively
long and narrow but not as apically tapered as in T. annulata. Accessory glands
narrow but about half length of those in T. annulata.
Tapeigaster subglabra
(Fig. 8)
General appearance of male reproductive system of this species quite different
from that of any other species dissected as shown in Fig. 8. Testes golden-brown with
8
> €a
05mm
Fig. 8. Tapeigaster subglabra, male reproductive system. ac = accessory gland, ae = aedeagus, ea = muscle
covered ejaculatory apodeme, ej = ejaculatory duct, t = testis, vd = vas deferens.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
M. A. SCHNEIDER 65
rounded lobe distally. Pale brown vasa deferentia short, entering median ejaculatory
duct at its slightly expanded apex. Accessory glands much shorter than those of T.
annulata, distinctly expanded at distal ends and entering distal end of ejaculatory
duct posterolateral to vasa deferentia. Aedeagal apodeme present although not shown
in Fig. 8.
DISCUSSION
All female specimens dissected had two spherical, dark spermathecae and males
had large, golden-brown testes. These could perhaps be regarded as generic
characters but further comparative studies need to be made to confirm this. The
dissection also showed that even in the females there are specific differences in the
morphology of the reproductive systems but it would be difficult to make accurate
interpretations concerning closeness of relationships between species from the
differences observed. However, in conjunction with the information gained from the
dissections of males it would seem that T. pulverea and T. annulata are quite closely
related. T. lutezpennis is perhaps closest of T. annulipes but with some similarities to
T. annulata. T. subglabra is not very close to any of the other species dissected.
ACKNOWLEDGEMENTS
I wish to thank Dr D. K. McAlpine of the Australian Museum, Sydney, for his
assistance and for supplying specimens for dissection.
References
Hennic, W., 1958 — Die Familien der Diptera Schizophora und ihre phylo-genetischen Verwandt-
schaftsbeziehungen. Beztr. Ent. 8: 505-688.
STURTEVANT, A. H., 1926 — The seminal receptacles and accessory glands of the Diptera, with special
reference to the Acalypterae. J. New York ent. Soc. 34: 1-21, pls 1-3.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
Neurochaeta inversa McAlpine
(Diptera: Neurochaetidae)
and Seed Set in
Alocasta macrorrhiza
(L.) G. Don (Araceae) in
southeast Queensland
DOROTHY E. SHAW, B. K. CANTRELL, and K. J. HOUSTON
SHAw. D. E., CANnTRELL, B. K., & Houston, K. J. Neurochaeta inversa McAlpine
(Diptera: Neurochaetidae) and seed set in Alocasta macrorrhiza (L.) G. Don
(Araceae) in southeast Queensland. Proc. Linn. Soc. N.S.W. 106 (1), (1981)
1982: 67-82.
Forty one intact infructescences of Alocasta macrorrhiza (L.) G. Don were
examined for presence or absence of Neurochaeta inversa McAlpine and for numbers
of seed set. N. znversa was recorded in all the spathal chambers of 20 infructescences in
two rainforest localities, in only two chambers of nine infructescences from two
natural stands in cleared areas, but not at all in chambers of 12 infructescences from
cultivated garden plants. Numbers of seed were not correlated with the numbers of N.
mmversa in the chambers, and seed was still set at sites where no N. inversa was
recorded. Pollination, therefore, is probably independent of ovipositing females of N.
mversa.
Dorothy E. Shaw, Plant Pathology, and B. K. Cantrell and K. J. Houston,
Entomology, Department of Primary Industries, Meters Road, Indooroopilly,
Australia 4068; manuscript received 2 June 1981, accepted for publication 23
September 1981.
INTRODUCTION
Alocasta macrorrhiza (L.) G. Don isa large aroid commonly known as ‘cunjevoi’
in Australia. Its inflorescence consists of a spathe and spadix (Fig. 1, A). The spathe is
divided into a terminal expanded portion (the spathal blade) and a convoluted
persistent base (the spathal tube) which forms the spathal chamber. The spadix
consists of four sections (Fig. 1, B) which, from the tip downwards, are: the sterile
terminal appendage; the staminate portion with synandria bearing the pollen; the
median sterile (m.s.) florets of the constricted portion and the pistillate flowers or
ovaries (with ovules) at the base. The stigmas are receptive prior to anthesis. After
fertilization, the tip of the spathal tube tightly clasps the constricted portion of the
spadix, forming the ‘sealed’ spathal chamber. The unfertilized ovaries in the sealed
chamber persist on the maturing infructescence and remain about the same size as at
anthesis, but fertilized ovaries (berries) enlarge to accommodate the developing
fertilized ovules (seed). The berries, green at first, tuyn yellow and then red at
maturity. The spathal tube also enlarges to accommodate the developing seeds. At
maturity, the walls of the spathal tube turn from green to yellow, and split
longitudinally into four or five segments which unfurl by rolling outwards revealing
the ripe berries:
McAlpine (1978) described Neurochaeta inversa, noting that it is only known to
occur in association with A. macrorrhiza in its original habitat in or at the edge of
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
NEUROCHAETA INVERSA McALPINE AND SEED SET
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Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
69
D. E. SHAW, B. K. CANTRELL AND K. J. HOUSTON
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Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
70 NEUROCHAETA INVERSA McALPINE AND SEED SET
rainforest and not with cultivated examples. Adults live on both sides of the leaf blade,
on the outer surface of the petiole, within the petiolar cleft and in the spathe of the
inflorescence at anthesis. Eggs (which were not observed) are apparently laid on the
female part of the inflorescence and larvae live between the developing fruitlets where
they are in contact with a ‘watery liquid’ found within the spathe which forms a sealed
chamber during the whole of the larval development. Pupariation, and often
emergence of adults, takes place while the fruitlets are still unripe and the spathe
sealed. The adults, therefore, cannot escape until the spathe splits at maturity. He
stated that ovipositing females of N. znversa may act as pollinators of 4. macrorrhiza,
but observed that, as many insects visit the flowers, they are probably not the sole
pollinating agents.
In studies carried out in southeast Queensland in 1980 and 1981, numbers and
percentages of fertilized ovaries (berries), numbers of fertilized ovules (seed) and
insects found in the sealed chambers of infructescences were recorded in order to
determine whether any correlation exists between the numbers of seed set and
numbers and types of insects recorded in the chambers.
SITES AND SAMPLES
Samples were from plants in three general habitats, viz., rainforest, intermediate
zones of remnant natural stands in mainly cleared areas, and in gardens. The localities
and brief descriptions of the sites are as follows :
1. Rainforest Samples were from clumps in small private holdings, adjacent
to large or relatively large reserved areas of rainforest, as
below:
Lamington Site O’R, on creek in mixed rainforest /sclerophyll area.
Plateau:
Mt Tamborine: Site BC, on creek in open area near rainforest and sclerophyll
forest.
Mt Glorious: Sites F, R(L), R(R), H, M, in rainforest, some near dirt road,
some near small clearings or thinned rainforest.
2. Intermediate zones
Upper Brookfield: Site Sm, isolated natural stand on dry creek in cleared area of
secondary growth and weeds; general area under crop and
artificial grassland; nearest rainforest remnant about 0.3 km
distant.
Beechmont: Site K, isolated natural stand on dry creek surrounded by
artificial grassland and weeds; nearest rainforest remnants
about 0.5 km distant.
3. Garden
Chapel Hill: Site S, one cultivated plant in suburban garden; area with
housing and gardens, and remnant sclerophyll and rainforest.
Mt Coot-tha: Sites AR, ER and MC, scattered planted clumps, some near
creek, in suburban Mt Coot-tha Botanic Gardens; area
originally sclerophyll forest, planted to simulated rainforest
during the last three years.
St Lucia: Site NR, large planted clump near creek in small simulated
rainforest surrounded by parklands in the University of
Queensland.
Each site was sampled once except Mt Coot-tha (twice) and Mt Glorious (three
times) .
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
D. E. SHAW, B. K. CANTRELL AND K. J. HOUSTON 71
The samples consisted of 54 infructescences or heads which were examined for
insects. Forty-four heads had intact and sealed spathal chambers; of these, 41 were
examined for seed set and three especially for physiological maturity. Ten other heads
were too young for seed or had damaged spathal tubes. The samples were at various
stages of development from anthesis to near-maturity.
Counts for insects and other organisms were made by removing each spathal tube
in four or five longitudinal strips and examining the inside of each strip immediately
under the stereomicroscope for larvae, pupae, etc., any detected being removed to
vials for later study. Any liquid in the spathal chamber was also examined. The outer
surfaces of the ovaries were examined microscopically in situ and any organisms
removed to vials. Each ovary was then removed with forceps while still under
microscopic observation, this being especially necessary in order to detect colourless
larvae around the ovary bases and on the white spadix core.
SEED AND INSECT RECORDS
Numbers of fertelized ovaries (berries) per head
The number and percentage of fertilized ovaries (berries) per head for 41 heads
with intact, sealed chambers, are given in Table 1. There was considerable variation
in the number of berries per head (six to 191), and in the percentage of berries per
head, which ranged from 3.5% to 98.1%. The three heads from Beechmont (Site K)
all had a low number of fertilized ovaries, viz., 3.5%, 10.1% and 17.0%. The
numbers were also low for the three heads from Site R(L), Mt Glorious, with 18.3%,
5.8% and 4.2%.
Numbers of seed per head
The numbers of seed per head are given in Table 1, ranging from six seeds in six
berries (an average of 1.0 seeds per berry) to 354 seeds in 191 berries (an average of
1.9 seeds per berry).
The lowest averages of seed per head per site were for Site R(L), Mt Glorious,
and Site K, Beechmont, with 10.7 and 27.7 seeds per head per site respectively, while
the highest were at Site M, Mt Glorious, and Site ER, Mt Coot-tha, with 222.0 and
222.5 seeds per head per site respectively. Statistical analysis showed that differences
between some sites are significant at either the 5% or 1% level (Table 2).
TABLE 2
Statistical analysis of average numbers of seeds
per head per site
HABITAT, Site* Seeds Sites (numbered as in third column)
locality Design- Num- per head significantly less than designated site
ation ber# per site
Av. No. 5% level 1% level
RAINFOREST
Lamington OR 144.7 Sites 3, 8
Plateau 1
Mt Glorious F 2 183.0 Sites 8, 11 Site 3
R(L) 3 10.7
R(R) 4 180.5 Site 8 Site 3
M 5 220.0 Sites 9, 12 Sitesi3i16) 7,8) 1
H 6 89.6
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
72 NEUROCHAETA INVERSA McALPINE AND SEED SET
TABLE 2—continued
HABITAT, Site* Seeds Sites (numbered as in third column)
locality Design- Num- per head significantly less than designated-site
ation ber# per site
Av. No. 5% level 1% level
INTERMEDIATE
ZONES
U. Brookfield Sm 7 102.3 Site 3
Beechmont K 8 Dl
GARDEN
Mt Coot-tha AR 9 75.5
ER 10 222.5 Sites 6, 7, 9, 12 Sites 3, 8, 11
MC 11 63.3
St Lucia NR 12 109.3 Site 3
Garden site 1 at Chapel Hill omitted because only one sample.
# Sites numbered consecutively for statistical analysis.
Less variation was shown in the average numbers of seed per head per locality,
which ranged from 27.7 for Beechmont to 144.7 for Lamington and even less variation
in the average numbers of seed per head per habitat, being 77.4 for the intermediate
zones, 112.3 for garden areas and 137.1 for rainforest (Table 1).
Insects
The only insects found within undamaged, still-sealed heads were 1) N. znversa,
2) Nitidulidae and 3) a few miscellaneous organisms, as below:
1. N. znversa
The eggs were not located. The numbers of larvae, puparia (with pupae or
empty) and adults recorded in the spathal chambers of the intact heads with seed are
shown in Table 3, and the totals are also inserted in Table 1. The numbers of adults
are given in brackets in Table 3, and are not included in the totals, as those individuals
are presumably already included in the numbers of empty puparia.
TABLE 3
Occurrence of N. inversa and Nitidulidae in 41 heads of
A. macrorrhiza
HABITAT, Head N. inversa Nitidulidae
locality, design- Larvae Puparia Adults Total Larvae Adults
site ation with empty
pupa
No. No. No. No. No. No. No.
RAINFOREST
Lamington Plateau
O’R 1 31 0 0 0 31 3 0
2 2 18 0 0 20 0
3 0 6 11 0* 17 1 0
Mt Glorious
F 2 0 0 46 0* 46 n.s.* n.s.*
3 0 0 42 0* 42 n.s.* n.s.*
R(L) 2 0 16 10 0* 26 2 0
3 4 25 5 0* 34 2 0
4 8 19 0 0 27 1 0
Proc. Linn. Soc. N.S.W.,106 (1), (1981) 1982
D. E. SHAW, B. K. CANTRELL AND K. J. HOUSTON
TABLE 3—continued
Occurrence of N. inversa and Nitidulidae in 41 heads of
A. macrorrhiza
HaBIiTAT, Head N. inversa
locality, design- Larvae Puparia
site ation with empty
pupa
No No. No.
R(R) 6 2 29 18
7 21 5 ]
M 3 35 0 0
4 32 44 1
5 64 11 1
6 0 1 3
7 0 44 1
H 2 ] 2 0
3 16 2 0
4 0 0 2
5 0 4 2
6 0 6 32
INTERMEDIATE ZONES
Upper Brookfield
Sm ] 0 0 0
2 0 0 0
3 0 3 10
4 0 0 0
6 0 0 5
Beechmont
B 1 0 0 0
D) 0 0 0
3 0 0 0
GARDEN
Chapel Hill
S 1 0 0 0
Mt Coot-tha
AR 4 0 0 0
5 0 0 0
ER 6 0 0 0
7 0 0 0
MC 3 0 0 0
4 0 0 0
5 0 0 0
St Lucia
NR 1 0 0 0
2 0 0 0
3 0 0 0
4 0 0 0
Adults
Total
Soe qocose
ooo °&
Nitidulidae
Larvae Adults
No. No.
2 0
2 2
3 0
0 0
1 0
2 0
0 0
1 0
9 0
5 0
0 0
] 0
11 0
0 0
2 0
0 1
0 0
7 0
2 0
0 0
n.s.* n.s.*
0 0
2 0
0 0
0 0
0 0
0 0
0 0
2 0
0 0
0 0
0 0
* Some adults apparently escaped without detection during the opening of the chamber
n.s. = not sought
73
# Numbers of adults given in brackets not included in the total, because the individuals already
counted in the numbers of empty puparia.
All heads sampled from rainforest sites had one or more stages of N. znversa,
while in heads from intermediate zones, only two out of nine heads sampled had any
stage of the fly, and then only in low numbers. No N. znversa were recorded in heads
from garden areas.
Because of the number of zero values in the intermediate and garden sites, the
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
74 NEUROCHAETA INVERSA McALPINE AND SEED SET
data were transformed (x = \/x + 4) and the transformed means show that both the
intermediate and garden sites differ significantly at the 5% or 1% levels from the
rainforest sites in numbers of N. znversa present (Table 4).
The numbers of N. znversa individuals per head, together with average numbers
per site, locality and habitat, are shown in Table 1. N. znversa was recorded in all
heads at all the rainforest sites, as reported previously, and seed set per head for these
ranged from 4.2% to 96.0%. In the intermediate zones, only two heads at the Upper
Brookfield site had a small number of N. znversa with 15.7% and 80.3% seed set
respectively, while the four heads without N. znversa at this site had 27.5% to 76.6%
seed set. None of the three heads from Beechmont had N. znversa, but seed set was
3.5%, 10.1% and 17.0% respectively. In the garden sites, where N. inversa was
absent, seed set ranged from 5.5% to 98.1%. This latter figure, in fact, was the
highest percentage of seed recorded in this study.
TABLE 4
Statzstzcal analysis of average numbers of N. inversa
per head per site
HABITAT, Site* N. inversa Sites (numbered as in third column
locality Design- Num- per head significantly less than designated site
ation. ber# per site
Av. No.* 5% level 1% level
RAINFOREST
Lamington O’R 1 22.3 Sites 7, 8, 9, 10, 11, 12
Plateau
Mt Glorious F 2 44.0 Sites 6, 7, 8, 9, 10, 11, 12
R(L) 3 28.9 Site 6 Sites 7, 8, 9, 10, 11, 12
R(R) 4 37.2 Site 6 Sites 7, 8, 9, 10, 11, 12
M D) 36.3 Sites, 6, 7, 8, 9, 10, 11, 12
H 6 10.4 Sites 7, 8, 9, 10, 11 Site 12
INTERMEDIATE
ZONES
U. Brookfield Sm 7 1.7
Beechmont K 8 0.0
GARDEN
Mt Coot-tha AR 9 0.0
ER 10 0.0
MC 11 0.0
St Lucia NR 12 0.0
x Garden Site 1 at Chapel Hill omitted because only one sample.
Sites numbered consecutively for statistical analysis.
Transformed means.
A co-variance analysis was carried out on seed number per site using the numbers
of N. znversa as a co-variate, and the regression is not significant.
2. Nitedulidae
Larvae of Nitidulidae (Coleoptera) and more rarely, adults, were found in low
numbers in some heads at most sites (Table 3), particularly at rainforest and
Beechmont sites. The larvae mainly occurred in the ms. florets undergoing
deliquescence, on the inner face of the spathal tube and only rarely on the berries.
3. Other insects etc.
Other insects and organisms found in undamaged, sealed spathal chambers are
recorded in Table 5. They were mainly larvae of unknown Diptera, Ceratopogonidae
(biting midges) , oligochaete worms and crustaceans (Syncarida: ?Bathynellacea) .
Proc. Linn. Soc. N.S.W.,106 (1), (1981) 1982
D. E. SHAW, B. K. CANTRELL AND K. J. HOUSTON 75
TABLE 5
Occurrence of other insects etc. and damaged berries in 41
heads of A. macrorrhiza
HABITAT, Head Insects etc. Damaged berries with
locality, design - Prick Hole with Wet rot# into
site ation lesion* some rot spadix core
No. Ge, No. %
RAINFOREST
Mt Glorious
R(L) 2 = 18 19.4
4 7 28 19.7
R(R) 6 He ] 0.6
7 Few Oligochaeta (worms) at 6 4.9"
M 3 ie D 1.8
4 a 37 N78
5 3 2 1.0
6 oe I 0.
7 z 2 1.6
H 5 y
6 6 Ceratopogonidae larvae z 4 3.9 29 22
3 Unknown Diptera larvae
2 Diptera puparia
1 dead flying ant
GARDEN
Mt Coot-tha
MC 3 3 dead adult Thripidae >
St Lucia
NR 1 16 Ceratopogonidae larvae i 4 3.5
1 Aphididae (1 apterous,
1 immature)
1 ? Hymenoptera (prepupa)
2 3 Oligochaeta (worms) =
3 4 Oligochaeta (worms) z 2 le
4 10 (approx). ? Bathynellacea 5 4
(Syncarida) Crustacea
Fusarium sp. (fungus) (sp.1)
(fine web and spores over the
berries)
* * = ‘prick’ lesions present; ** = abundant; ~ = absent.
# Impossible to determine whether rotted structures were berries or unfertilized ovaries.
Rot extending into spadix core.
INSECTS IN YOUNG OR DAMAGED HEADS
Insects and other organisms in ten other heads which were either too young to
have seed, or where the spathal tube was damaged or gaping, are reported in Table 6.
N. inversa larvae, puparia and adults were recorded inside the spathal chambers of
most of these rainforest heads. Nitidulid larvae and a few adults were found either
outside or inside the spathal chambers, in the latter case mainly near or in the
decaying m.s. florets at the tip of the chamber. Eggs of Chloropidae (Diptera) were
found mainly inside the chambers of heads picked while the spathal limb was still
extended and unwilted, while the larvae were found in decaying tissue outside and
inside the spathal chambers. Two species (Cadrema sp. and Hippelates sp.) were
reared from larvae in the laboratory, and one specimen of each species was parasitized
by a cynipid wasp (Hymenoptera: Cynipidae: Eucoilinae). The large (0.6 x 0.24
mm) opaque white eggs on Mt Glorious heads have not been identified. Only one
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
76 NEUROCHAETA INVERSA McALPINE AND SEED SET
ceratopogonoid larva was recorded on a Mt Glorious head. Brachypterous female
Phoridae (Diptera) were found in the decaying tissue of two damaged heads from Mt
Coot-tha.
TABLE 6
Insects recorded in ten young or damaged heads of A. macrorrhiza, not reported in previous tables
HABITAT, Insects, etc.
locality Head maturity and position of N. inversa Nitid- Chlor- Miscell-
site, head insects ulidae opidae aneous
designation No. * No. * No.* No. *
RAINFOREST
Mt Glorious
R(L)
Head 5 Very young, spathal limb still
present; no difference in ovary
sizes
In plastic bag after transport 1L 1 Diptera L
4A 1 Staphy-
lididae L
In debris at tip on outside of
spathal tube 11L 20L
Inside spathal chamber 33L 1L 98#E 1 Large white
€ss
Few flatworms?
Head 8 Prior to anthesis, held in lab. 18
days; no enlargement of ovaries
but constriction closed
Inside chamber near tip 1L 20L 28#E 1 Ceratopo-
gonidae L
3P 15L
4P
2A
R(R)
Head 9 Very young, spathal limb still
present; held in lab. 4 days; no
perceptible difference in ovary sizes
In decaying florets outside
spathal chamber 1L 3L
Inside chamber 2L aan E 1 large white
ess
7L
Head 10 Very young, spathal limb still
present; held in lab. 5 days;
hardly perceptible difference in
Ovary sizes
Inside chamber 40L 97#E 8 large white
eggs
24L
M
Head 1 Head net bagged before anthesis; 1
ovary slightly enlarged F
Inside spathal flap 4E
Inside spathal chamber 47L (Some ‘prick’
lesions present)
5P
Head 8 Head ripe with partially opened
spathal tube
Inside chamber 8P
H
Head 7 Head with slight rot of spathal tube tip
Proc. Linn. Soc. N.S.W.,106 (1), (1981) 1982
D. E. SHAW, B. K. CANTRELL AND K. J. HOUSTON 77
TABLE 6 — continued
Insects recorded in ten young or damaged heads of A. macrorrhiza, not reported in previous tables
HABITAT, Insects, etc.
locality Head maturity and position of N. inversa Nitid- Chlor- Miscell-
site, head insects ulidae opidae aneous
designation No. * No. * No. t No. *
Inside chamber 2P 4L
2A
Head 8 Head with slight rot of spathal
tube tip
Inside chamber 15P 2P
GARDEN
Mt Coot-tha
MC
Head 1 Staminate portion deliberately
severed before anthesis; later rest
of spadix rotting
In rotting florets 3 Phoridae A
Head 2 Small hole with some rot into
spathal chamber
In rotting spadix 19L 15 Phoridae A
1A
*E = egg; L = larva; P = puparium; A = adult.
# Probably more eggs present but uncounted.
+ Eggs uncounted.
BIOLOGY OF NEUROCHAETA INVERSA
The development of N. znversa from larva to adult within the sealed spathal tube
of A. macrorrhiza seems fairly precisely adjusted to the maturation of the
infructescence. Details of the maturation of the head will be given separately by
D.E.S., but the part of the process relevant to the spathal chamber and the chamber
liquid may be summarized as follows: The portion of the spadix (consisting of the
terminal sterile appendage, the staminate section and the upper part of the m:s.
florets) distal to the constriction withers and falls off. The spathal blade also withers
and falls off, usually in its entirety. The tip of the spathal tube constricts to form the
sealed spathal chamber (Fig. 1, C) which encloses the lower part of the ms. florets
and the pistillate flowers or ovaries. The enclosed m.s. florets then undergo
deliquescence, leaving the spadix core bare and contributing to a broth of liquid and
flower-part remains. This broth bathes the surface and sides of the ovaries in the
sealed chamber and the broth debris often accumulates near or at the base of the
chamber. In some young heads, the enclosed liquid, which may not all have been
derived from the dissolution of the m.s. florets, measured up to 8 cc.
Larvae of N. znversa were usually found on the inner face of the spathal tube, in
the broth from the spathal chamber (Fig. 1, D) or on the outer faces of the exposed
berries and unfertilized ovaries. These latter larvae, however, moved rapidly between
the ovaries when the spathal tube was removed, and were usually only retrieved by the
removal of the berries and unfertilized ovaries. The development of the larvae
coincides with the deliquescence of the m.s. florets, as described above, so that when
the broth resulting from the dissolution is abundant, the larvae are present. Not all the
larvae in any one head pupariate at exactly the same time, so that in some cases both
larvae and puparia occur together in the same chamber (Table 3), often with the
remains of the m.s. florets still discernible. However, by the time all the larvae have
pupariated the remains of the m.s. florets have virtually disappeared, together with
most of the liquid. There is some latitude in this sequence, however, regarding
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
78 NEUROCHAETA INVERSA McALPINE AND SEED SET
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
D. E. SHAW, B. K. CANTRELL AND K. J. HOUSTON 79
TABLE 7
Infructescence maturity, amount of liquid in spathal
chamber and stages of N. inversain heads from
Mt Tamborine
Site, head Maturity of head Amount of Stages of N. znversa
designation (All heads with spathal liquid in Larvae Puparia Adults
tubes still green) spathal with empty
chamber pupae
No. No. No. No.
BC
Head 2 Berries enlarged, green trace 0 2 11 Da
Head 4 Berries larger, green about 0 16 30 295
0.5 cc
Head 3 Berries largest, turning about 1 4 7 i
yellow at base (therefore lec
ripest)
* Some adults apparently escaped without detection during the opening of the chamber.
maturity of the head, amount of liquid remaining in the chamber, and stage of
development of N. znversa. This is shown, for example, in three heads from Mt
Tamborine (Table 7), not included in the other tables. Head 3 from Mt Tamborine
was the only one, out of over 50 heads studied, where all three stages (larva, puparia
and adults) occurred at the same time. Although its berries were more mature than
those on Heads 2 and 4, its chamber contained more liquid than the other two
chambers, and this may have contributed to the presence of the larva.
The puparia were oriented with the posterior end downwards between the
berries, and with the flattened or slightly depressed facet at the anterior end usually
parallel to the surface of the spathal tube (Fig. 1, E). In a few cases puparia were
found between the base of the ovaries, probably dislodged during transport of the
head from the field.
McAlpine (1978) reported that the adults which emerge from the puparia cannot
escape from the spathal chamber until the fruits are ripe and the spathal tube splits.
In this study, adults were present in Heads 5 and 6, Site H, and in Head 7, Site M, at
Mt Glorious (Table 3) and also in all three heads from Mt Tamborine (Table 7), and
in all cases the spathal tubes were still green, without any yellowing which indicates
approaching maturity and which precedes splitting of the tube. The adults were quick
to escape from any artificial opening made in the tube, and perhaps escape naturally
as soon as the slighest relaxation occurs in the tube, even possibly prior to splitting.
Information is still required as to whether the adults feed after emergence while
still within the chamber, and whether they remain quiescent between the berry
shoulders, or make their way to the tip of the chamber and congregate around the
spadix core bared by the dissolution of the m.s. florets, waiting for the first
opportunity to escape.
N. inversa larvae were never noted causing damage to the ovaries or to the spadix
core. No parasites were recorded in any of the specimens.
Fig. 1. A, Spathe (left, detached) with spathal blade (sb) and spathal tube (st) and spadix (right) of
Alocasia macrorrhiza. B, Later stage of the spadix showing sterile terminal appendage (sta) ; staminate
flowers (s); median sterile flowers (msf) and pistillate flowers (p). C, ‘Sealed’ spathal tube enclosing
berries. D, Diptera larvae and ‘broth’ from spathal chamber. E, Puparia of Neurochaeta inversa and
berries of A. macrorrhiza. F, Tip of spathal tube with rot and sporulating fungus (Fusarzum sp.).
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
80 NEUROCHAETA INVERSA McALPINE AND SEED SET
DAMAGE TO INFRUCTESCENCE PARTS
Damage to berries in sealed, intact heads was of three types, viz., 1) ‘prick’
lesions, 2) hole in the berry wall, and 3) wet rot of the berry. Rot of the spathal tube
tip (point 4. below), resulting in gaping, occurred in three other heads.
1. ‘Prick’ lesions
In some heads (Table 5) at sites at Mt Glorious, minute red ‘prick’ lesions were
noted on some berry walls. In a few cases a narrow border of slightly more translucent
tissue surrounded the ‘prick’, but even then the whole diameter of the lesion was still
only about 0.5 mm. In no case did any rot of the berry wall or the underlying seeds
appear to be associated with these ‘prick’ lesions. The lesions may have been caused by
stylet penetration of the pistillate flowers before anthesis by an unknown insect.
2. Hole in berry
A hole in the berry wall, sometimes with damage to the enclosed seed, occurred in
14 berries from Mt Glorious and 10 from St Lucia (Table 5). The hole was usually
near the base of the berry, and such damage tended to cause a premature change in
the colour of the berry wall from green to yellow and even to red. In some cases the
damage extended into the spadix core, causing a reddish wet rot at the site of berry
attachment. Although larvae of Ceratopogonidae were occasionally found feeding
within the holes in the berries, this may merely indicate opportunism and not
necessarily a causal relationship.
3. Wet rot of ovaries
Wet rot of 116 ovaries (or berries, as it was impossible to determine whether
fertilization had taken place or not) was noted in six heads from Mt Glorious (Table
5), one case involving 22.7% of the ovaries on the head. If adjacent ovaries were
affected, counts were made on stigmas rather than on the ovaries themselves, as it was
difficult to distinguish the individual rotting bodies. In each case the rot extended into
the spadix core, again with a wet reddish discolouration of the tissue. It is not known
whether the wet rot is an extension of the ‘hole’ condition described above, or whether
it is quite distinct. The latter may be the case, as no wet rot was recorded in the St
Lucia heads, whereas holes did occur in 10 berries on three heads from this site.
4. Rot of tip of spathal tube
Three heads, including Heads 7 and 8 from Site H, Mt Glorious (Table 6), and
an unlisted pathology specimen head from Site R(R) at Mt Glorious, had a rot of the
spathal tube beginning at the tip, and proceeding downwards, with white sporulating
fungal clumps on the rotted tissue (Fig. 1, F). The fungus, which was obtained in
axenic culture, was Fusarium sp. 2 (see Addendum), different from Fusarzum sp. 1
recorded as a fine sporulating web on Head 4, St. Lucia (Table 5). Further
investigation 1s required to determine whether Fusarzum sp. is a primary pathogen, or,
as would seem more likely, a secondary invader following damage by an unknown
agent.
GENERAL DISCUSSION
McAlpine (1978) reported that larvae of Cadrema sp. (Diptera: Chloropidae)
were found in the moist decaying upper part of the spadix of A. macrorrhiza, and has
confirmed it (pers. comm.) to be the same species as that found in the present study.
We also found larvae of Hippelates sp. (Chloropidae) and Nitidulidae occupying that
niche provided by the dissolution of the m.s. florets at the tip of the spathal chamber
and in the decaying spadix parts outside the spathal chamber. It is not known if they
also occur on the decaying distal portion of the spadix which usually withers and falls
off the infructescence. If so, the life cycle of these two groups may be completed in the
decaying tissue on the ground.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
D. E. SHAW, B. K. CANTRELL AND K. J. HOUSTON 81
Some of the miscellaneous organisms found in the sealed chambers, such as the
Ceratopogonidae (biting midges), the oligochaete worms and the ?Bathynellacea
(Crustacea: Syncarida) may be chance visitors caught within the chambers before
sealing. Oligochaete worms and midges were only recorded in sealed intact heads from
St Lucia and Mt Glorious. The worms, however, are difficult to see, and low numbers
may have been missed in heads from other sites. The presence of the worms seemed to
be associated with thin rings of debris on the inner surface of the spathal tube,
although it was not possible to confirm the connection.
One of the interesting aspects revealed by this study was the considerable
variation in seed set on different heads at the one site (Table 1). Whether this was the
result of differences in numbers of the pollinating agent(s), or in the amount of
available pollen, or in microclimatic factors at the time of stigma receptivity of each
head, is not known.
McAlpine (1978) stated that ovipositing N. znversa females may well be polli-
nators of A. macrorrhiza, but observed that, as many insects visit the flowers, they are
probably not the sole pollinating agents. As shown in Table 1, the numbers of seed set
were not related to the numbers of N. znversa recorded in the chambers at the sites
where the fly was found. McAlpine (1978) further observed that N. znversa had not
been found on cultivated examples of ‘cunjevoi’ and this was confirmed in the present
study, as no N. znversa were recorded at these sites, although seed was still set as shown
in Table 1. We conclude, therefore, that pollination of A. macrorrhiza is probably
independent of ovipositing females of N. znversa in the inflorescence.
Carson and Okada (1980) previously noted that there appeared to be no
difference in seed set of flowers of Colocasza esculenta (Araceae) from which adult
Drosophiella pzstilicola were excluded by bagging, and unbagged controls. They
concluded, therefore, that the presence of. this fly was not necessary for full pollination
of the flowers. Their work, like that of the present authors, was concerned specifically
with an insect associated with the sealed spathal chambers, and not with the
elucidation of the roles which may be played by many other insects known to visit the
exposed flower parts.
ACKNOWLEDGEMENTS
The owners of the sites sampled and other individuals who assisted in obtaining
the collections, including Mr and Mrs E. J. Frazer, Ms J. Grimshaw, Mrs J. Henry, Mr
A. Hiller, Mrs J. Hope, Ms K. Howdesell, Mrs B. Kennedy, Mr and Mrs L. J. Manning,
Mr M. Olsen, Mr P. O'Reilly, Mr and Mrs D. Sands, Mr and Mrs W. H. Smith, and
Mr H. Caulfield and Mr J. Donnelly (Mt Coot-tha Botanic Gardens) and Dr A. B.
Cribb, Dr D. Priest and Mr A. R. Steginga (University of Queensland) are thanked
for their co-operation in this study.
We are also grateful to Dr P. J. F. Davie, Mr J. F. Donaldson, Dr I. D. Galloway,
Mrs M. M. Harris and Dr D. K. McAlpine for identifications, to Ms J. Alder for the
statistical analyses, and to the Director, Plant Pathology Branch, Department of
Primary Industries, Indooroopilly, for facilities to D. E. S. during the study.
References
Carson, H. L., and Oxapa, T., 1980. — Drosophilidae associated with flowers in Papua New Guinea. 1.
Colocasia esculenta. Kontyu, Tokyo 47: 15-29.
McALPINE, D. K., 1978. — Description and biology of a new genus of flies related to Anthoclusea and
representing a new family (Diptera, Schizophora, Neurochaetidae). Ann. Natal. Mus. 23: 273-295.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
82 NEUROCHAETA INVERSA McALPINE AND SEED SET
ADDENDUM
Identifications of Fusarzum spp. 1 and 2, mentioned in the section ‘Damage to Infructescence Parts’, now
received from the Commonwealth Mycological Institute, United Kingdom, are Fusartwm solani (Mart.)
Sacc. and F. oxysporum Schlecht., respectively.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
1/06 C/a) ¢
The Fauna of
Australian Mangroves
P. A. HUTCHINGS and H. F. RECHER
Hurcuincs, P. A., & RECHER, H. F. The fauna of Australian mangroves. Proc. Linn.
Soc. N.S.W. \@@€®, (1981) 1982: 83-121.
Mangrove forests are among: the world’s most productive ecosystems. In
Australia, mangroves have an extensive distribution and are probably important for
the maintenance of estuarine fisheries. Despite these values, there has been relatively
little research on the ecology of Australian mangrove communities. Botanical studies
have been most extensive, work on mangrove fauna has been largely restricted to a few
commercially-important or pest species. Many of the data available are non-
quantitative, anecdotal or unpublished. This lack of data on mangrove fauna prevents
the development of detailed plans of management and therefore poses long-term
conservation problems. In order to identify areas where research on Australian
mangrove fauna is needed and, it is hoped, to stimulate such work we summarize
information on the Australian mangrove ecosystem and discuss the structure and
evolution of these communities.
P. A. Hutchings, Department of Marine Invertebrates, and H. F. Recher,
Department of Terrestrial Ecology, Australian Museum, P.O. Box A285, Sydney
South, Australia 2000; manuscript received 19 May 1981, accepted in revised form 21
October 1981.
INTRODUCTION
Mangroves are among the most productive of the world’s forests (Westlake, 1963;
Lugo and Snedaker, 1974) and contribute importantly to the productivity of tropical
and sub-tropical estuaries. Working in southern Florida, Odum and Heald (1975)
demonstrated that the organic matter produced by mangroves formed the base of a
detritus food chain that culminated in the rich fisheries of Florida Bay. Mangroves
occur around the coast of Australia (Lear and Turner, 1977) (Fig. 1). In the
estuaries of northern New South Wales, Queensland, the Northern Territory and
parts of Western Australia, mangroves form extensive forests which, with seagrasses
and salt marshes produce the organic matter that is the base of the fisheries of
northern Australia. Many of Australia’s commercial fish species feed on mangrove
fauna or use the mangroves as a nursery (Newell and Barber, 1975; Pollard, 1976,
1981). Crab, prawn and oyster are other fisheries intimately associated with
mangroves (Ruello, 1973; Staples, 1980 a,b).
Despite their ecological and economic importance, research on Australian
mangroves has been largely botanical, describing vegetation (MacNae, 1966, 1967;
Chapman, 1975; Saenger et al., 1977; Bunt and Williams, 1980, 1981; Williams and
Bunt, 1980), floral biology (Clarke and Hannon, 1970, 1971; Duke and Bunt, 1979;
Tomlinson et al., 1978, 1979), geomorphology in relation to distribution of
mangroves (Thom e¢ al., 1975) and mangrove physiology (Clough and Andrews,
1982). Faunal studies have mostly been restricted to surveys of species present in
mangroves (e.g. Hutchings and Recher, 1974; Saenger et al., 1977). Studies on
ecosystem dynamics have been initiated (Bunt e¢ al., 1979; Clough and Attiwill,
1982, a, b; Goulter and Allaway, 1979), but there is little information on mangrove
productivity or the pathways by which energy flows from mangroves through
Australian estuaries. The information that is available tends to support the findings of
Odum and Heald (1975).
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
84 FAUNA OF AUSTRALIAN MANGROVES
12
7]
3
NO. OF MANGROVE ee, _TROPIC OF CAPRICORN _
SPECIES (ene alae SS ae
38 eee 24
ONE SPECIES 1
20-29
30
~A
10- 20
SE IONS oe ee ae oe
: ONE SPECIES 36
ONE SPECIES
Fig. 1. The number of mangrove species increases from temperate areas to the tropics. Only one species,
Avicennia marina, occurs around the coast from South Australia to Cape York. With increasing species
number, there is a concomitant increase in community complexity.
Odum and Heald (1975) examined the food webs in a simple estuarine mangrove
community, Rhzzophora mangle, in southern Florida. The community was
characterized by a large vascular plant production with little input from plankton or
algae. Most of the leaves and stems produced by the mangroves (Duke et al., 1982;
Williams et al., 1982) were not eaten by herbivores, but became part of the decaying
matter (‘litter’) on the forest floor. Mangrove detritus formed the basic component in
the diet of primary marine consumers (‘detritivores’). Among the primary consumers
studied by Odum and Heald, detritus of vascular plant origin formed 20 per cent of
the diet. The authors concluded that mangrove leaf detritus was the most important
element in the food web. However, Redfield (1982) has presented a modified version
of their model with increased emphasis on the consumption of mangrove leaves by
insects (see Onuf et al., 1977) and mud dwelling herbivores (see Malley, 1978) and
decreased emphasis on the importance of bacteria and fungi in the breakdown of
mangrove detritus.
As in all communities, animals are an integral part of the mangrove ecosystem.
Herbivores and detritivores living in the mangrove forest have been shown to be
important in the transfer of energy from the mangroves to the estuary (Goulter and
Allaway, 1979; Odum and Heald, 1972; Malley,1978). In the absence of detailed
information about the fauna of Australian mangroves, models of local mangrove
ecosystems must be considered incomplete. In turn, the lack of data on the mangrove
ecosystem, its fauna and the relation of these to the estuary seriously hampers the
conservation of mangrove forests and the management of estuarine fisheries. As we
have previously pointed out (Hutchings and Recher, 1977; Recher and Hutchings,
1980), mangroves are a threatened habitat throughout eastern and southern
Australia. The development of a national conservation strategy for mangroves and
Proc. LINN. Soc. N.S.W., 106 (1), (1981) 1982
P. A. HUTCHINGS AND H. F. RECHER 85
their management is of critical importance for the sustained production of estuarine
and near-shore fisheries (Hutchings and Recher, 1977).
In this paper we review what is known of the mangrove fauna in Australia. Data
are often sketchy or anecdotal and much of the information presented here is as yet
unpublished. We do not apologize for these deficiencies; our intention is to focus
future research work on areas of greatest need.
MANGROVE FAUNA
The mangrove fauna is composed of animals from terrestrial, marine and
freshwater environments. It is a diverse fauna with an abundance of vertebrate and
invertebrate forms. Some are restricted to mangroves; others occur more widely. For
many species the full range of habitats used is unknown and species currently classed
as Mangrove endemics may occur elsewhere. Some animals are only temporary
residents. Birds may use mangroves for roosting, fish move into and out of the
mangroves with the tides, others come in seasonally like flying foxes (Pteropus spp.) to
feed on nectar. In this review we have ignored records of animals which do not occur
regularly in mangroves.
VERTEBRATES
Few vertebrates are restricted to mangroves. The majority are casual visitors or
accidental occurrences. Saenger et al. (1977), for example, record over 200 species of
birds from Australian mangrove habitats, but fewer than half occur regularly
(Schodde et al., 1982).
Mammals
No mammal is endemic to mangroves in Australia, nor is there any species for
which mangroves are the principal habitat. The rat, Xeromys myozdes, has been
recorded living among mangroves on the Tomkinson River, Arnhem Land, and on
Andranang Creek, Melville Island in the Northern Territory (Magnusson et al.,
1976). It has also been recorded as frequenting mangroves near Mackay in central
Queensland and on North Stradbroke Island in southern Queensland (Van Dyck et
al., 1979). In these situations it builds nests of leaves and mud among the mangrove
roots of Bruguzera; the nests are flooded on the highest tides (Magnusson et al.,
1976). Xeromys feeds on crabs and is at least partially arboreal, but also occurs in a
wide range of habitats other than mangroves (R. Strahan, pers. comm.) .
Many other terrestrial animals enter mangroves and may obtain part of their food
there. Frith (1973) records a number of rodents (Rattus colletti, Mus musculus,
Melomys spp., Mesembriomys spp. and Conilurus spp.), bandicoots (Perameles,
Isoodon) and the northern brush-tailed possum Tr¢chosurus arnhemensis as occurring
in mangroves. The water rat Hydomys chrysogaster forages regularly in mangroves
(Frith, 1973; Recher, pers. obs.) . Two species of flying foxes, Pteropus poliocephalus
and P. alecto commonly come into the mangroves to feed on nectar or to camp. P.
poliocephalus occurs along the entire east Australian coast whereas P. alecto is
restricted to tropical mangroves. Another species, P. conspicillatus, sometimes camps
in mangroves. Some species of tree-roosting bats such as Tadarzda planiceps, the flat-
headed mastiff-bat and T. loriae, the little northern mastiff-bat, are occasionally
found in mangroves. The northern blossom bat, Macroglossus lagochilus occurs in
stands of Sonneratza alba in Western Australia.
Mangroves do not appear to be used by marine mammals. Heinsohn and Wake
(1976) stress the importance of sheltered waters for dugong Dugong dugon and the
association of dugong with seagrass beds but make no mention of dugong using
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
86 FAUNA OF AUSTRALIAN MANGROVES
mangroves. Recently dugongs and dolphins have been seen in the mangrove channels
on Hinchinbrook Island (Bunt, pers. comm.).
Birds
Birds are a conspicuous component of all mangrove forests, although they are not
abundant. Schodde et al. (1982) suggest that the comparative uniformity of structure
within the canopy of mangrove forests, which provides little variety of foraging
surfaces, accounts for the small number of individuals. In many regions, mangroves
have a fragmented distribution which Schodde et al. suggest may also be important in
limiting numbers. Feeding time for ground foragers is restricted by tidal flooding.
Nest sites may be limited. These reasons may also explain why many birds occurring in
mangroves, also occur in other habitats. Data are not available on whether these
species occupy similar niches in all habitats or if they broaden their niche in the
mangrove forest.
Over 200 bird species have been recorded from Australian mangroves (Saenger et
al., 1977). Of these, 14 species are virtually restricted to mangroves; 12 species use
mangroves as a primary habitat in part of their range, and 60 species use mangroves
regularly throughout the year or in particular seasons (Schodde et al., 1982). This is a
rich mangrove bird fauna compared with other parts of the world (MacNae, 1968).
The number of species of birds using mangroves and the number of mangrove
specialists increases from south to north with increasing floristic richness (Fig. 2).
Thus near Sydney (34°S) only one species, the mangrove heron Butorides striatus, is
restricted to mangroves. The other birds in temperate mangroves are widely
distributed in a variety of aquatic or forest habitats (e.g. rufous whistler Pachycephala
rufiventris and white-face heron Ardea novaehollandiae). In contrast, on the Cape
York Peninsula (19°S) nineteen species of birds are considered mangrove specialists
and seven of these are endemic to mangrove habitats (Schodde et al., 1982). Birds
which are wholly or partially dependent on mangroves include two herons, a kite, a
rail, a pigeon, a cuckoo, three kingfishers, five pachycephaline flycatchers, four
myiagrine flycatchers, three acanthizine warblers, three honeyeaters, a silvereye and a
butcher-bird. None shows major morphological adaptations to the mangrove
environment. However there is a tendency for bills to be longer than in non-mangrove
congeners (Schodde et al., 1982) . The white-breasted whistler Pachycephala laniozdes
has a comparatively heavy and hooked bill for handling marine invertebrates.
Birds which regularly use mangroves, but which cannot be considered mangrove
specialists, include aquatic species and a wide range of passerines. Aquatic and
wading birds (e.g. herons, ibis, sandpipers) forage among mangroves and many use
mangroves as high tide roosts. Some species (e.g. pied cormorant Phalacrocorax
varius, straw-necked ibis Threskzornis aethiopicus) nest in mangroves (White, 1917;
Seton, 1971; Schodde et al., 1982). Among the passerines, honeyeaters commonly
visit mangroves for nectar when blossom is abundant. The sulphur crested cockatoo
Cacatua galerita has been seen feeding on the fruits of Lumnitzera littorea in north
Queensland by Bunt and his co-workers (pers. comm.). Other birds (e.g. rufous
fantail Rhipidura rufifrons and grey fantail R. fuliginosa) frequent mangroves during
migration.
Reptiles
Although they may be important feeding grounds, for the majority of terrestrial
reptiles using mangroves, mangroves are peripheral habitats. In some areas,
mangroves are the only habitable forest and may, therefore, be important corridors
for the movement of individuals.
Reptiles are common in tropical mangroves, but are rarely seen in temperate
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
P. A. HUTCHINGS AND H. F. RECHER 87
35 (10 )
tae
45 (8)
Faunal areas based on terrestrial birds ( after Kikkawa & Pearse, 1969 )
No. of bird species ( no. of species confined to or most abundant in mangroves )
Total for Australia is ~ 58 (14) species of terrestrial birds
Fig. 2. Kikkawa and Pearse (1969) divided Australia into regions based on the similarity of the bird fauna.
These regions are convenient to illustrate the number of species of birds which occur regularly in
mangroves. The number of bird species endemic to mangroves in each region are shown in parentheses.
Species number and the number of endemics increase from south to north. This latitudinal gradient in bird
species diversity is correlated with an increasing area of mangroves and the richer floristic and structural
complexity of tropical mangrove forests.
ones. The salt water crocodile Crocodylus porosus is probably the best known
mangrove resident; a notoriety derived from its occasional consumption of humans.
Salt water crocodiles occur in river systems from Broome in Western Australia, around
the Northern Territory and south to Maryborough in Queensland (Cogger, 1979).
The animal is most abundant in the tropics and comes into mangroves to feed.
Juvenile crocodiles take crabs (especially sesarmids) , prawns, mud skippers, and small
fish. Larger animals take mud crabs Scylla, birds, and mammals including Xeromys
and flying foxes (Taylor, 1979). Crocodiles do not nest in mangroves (Webb et al.,
1977).
The fresh water crocodile C. johnstond occurs rarely in mangroves and is probably
accidental at the landward margins of the mangroves and in less saline areas. Cogger
(1979) has recorded the pitted-shelled turtle Carettochelys insculpta in the Daly,
Victoria and Alligator River systems in the Northern Territory where it may enter
mangrove areas.
A small number of snakes and lizards occur regularly in mangroves. Mangroves
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
88 FAUNA OF AUSTRALIAN MANGROVES
are the primary habitat for the mangrove monitor Varanus ¢ndicus which occurs in the
Northern Territory, Torres Strait and Cape York (Cogger, 1979). It feeds on crabs
and mud skippers. Four other goannas, V. semiremex, V. prasinus, V. ttmorensis and
V. tristts are common in tropical mangroves, but range widely in other habitats
(Cogger, 1979). The littoral skink Emoza atrocostata is common in mangrove areas of
the Torres Strait and northern Cape York Peninsula (Cogger, 1979). The Northern
Water Dragon (Lophognathus temporalis) feeds on insects in the mangroves of
northwest Australia and the Northern Territory (Swanson, 1976).
Several species of pythons (Lzaszs fuscus, L. oltvaceus) occasionally use
mangroves and are attracted by large camps of flying foxes on which they feed. The
carpet snake (Morelia spilotes) and Liaszs amethystinus and two common colubrid
snakes, the green tree snake Dendrelaphis punctulatus and brown tree snake Bozga
rregularts also occur in mangrove habitats. The file snake Acrochordus granulatus is
virtually restricted to mangroves in northeastern and northern Australia. It lives on
the mud flats in front of the mangroves where it feeds on crabs and fish (Cogger,
1979). Mangroves are also an important habitat for the bockadam Cerberus
rhynchops, the white-bellied mangrove snake Fordonza leucobalia, and the mangrove
snake Myron richardsoni which occur in northern Australia (Cogger, 1979.). Some
sea snakes are estuarine and occur commonly in mangroves (Cogger, 1979.).
Ephalophis mertoni, E. greyt and Hydrelaps darwiniensis are found in mangroves in
the Northern Territory and Hydrophis elegans and Aipysurus eydouxw are found in
central Queensland.
Amphibians
Two frogs have been reported from mangrove habitats in the Northern Territory.
The northern dwarf tree frog Lztorza bicolor has been recorded on the Wildman
River, in an area greatly affected by freshwater, and the marbled frog Lymnodynastes
convexiusculus occurs in the Cerzops tagal zone of the South Alligator River (Hegerl et
Glo, UDI)
Fishes
Fishes are a conspicuous component of the mangrove ecosystem with large
numbers invading the mangrove forest at high tide and retreating to deeper waters as
the tide falls. Either as juveniles or adults, virtually all the common fishes of the
estuaries can be found in mangroves (Ellway and Hegerl, 1972; Shine et al., 1973;
Saenger et al., 1977; Beumer, 1978; Blaber, 1980). Few species are restricted to
mangroves.
The major group of fishes adapted to living in mangroves belong to the gobiid
sub-family Oxcidercinae. This includes mud skippers which are represented in
Australia by five species in two genera, Perrophthalmus and Perzophthalmodon. Mud
skippers are common throughout the tropics but are absent from temperate
mangroves. They occur only as far south as Hervey Bay in Queensland (25°S). Other
representatives of this family are the genera Boleophthalmus and Scartelaos which
burrow in the mud and occur in the Northern Territory and Queensland.
Other gobies are common in mangroves, but are also found elsewhere in the
estuary. Included are the genera Mugilogobius, Taenzoides and Arenigobzus which
occur throughout Australia. The oyster blenny Omobranchus occurs in frontal
mangroves where residual water is trapped. All these fish can stand some exposure to
air. Most live in burrows where some water is present even during low tide.
A diverse fish fauna occurs in creeks or lagoons in the mangroves where
permanent water occurs. These will be estuarine species, characteristic of the
geographical region (Saenger et al., 1977; Ellway and Hegerl, 1972 and Shine et al.,
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
P. A. HUTCHINGS AND H. F. RECHER 89
1973). Among the more common estuarine species which frequent mangroves are
toados Torquiger hamiltoni, mullet Myxus elongatus, and fortesques Centropogon
australis.
Blaber (1980) has designated feeding categories of the fish of Trinity Inlet,
Cairns. Beumer (1978) has studied the feeding ecology of black bream
(Acanthopagrus berda), bony bream (Anodontostoma chacunda), hair fin goby
(Ctenogobius crineger) and milk-spotted toad fish (Chelonodon patoca) in a
mangrove creek in the Townsville region.
INVERTEBRATES
There are two components to the invertebrate fauna of mangrove communities,
spiders and insects associated with the forest canopy and aquatic animals inhabiting
the intertidal. This fauna is richer than the vertebrate community. Our approach in
reviewing the information available on mangrove invertebrates has been to use a
classification based on taxonomic and ecological attributes. We first consider insects
and spiders, but then review the aquatic fauna according to where they occur in the
mangroves.
Insects
Although neither the insects nor spiders of mangroves have been extensively
studied in Australia there is a great variety of species. These show a variable degree of
dependence on mangroves. Whether the diversity in mangroves is similar to that in
other forest habitats at the same latitude or whether mangrove forests have an array of
mangrove specialists is unknown. C. Smithers (pers. comm.) suggests that each
species of mangrove probably will have an associated unique suite of insect species.
In general, insects in mangroves appear to have the same range of adaptations
that they show in other forest environments. For many species, mangroves are an
important habitat and insects may be favoured by the often large, almost mono-
specific stands of trees within the forest.
A comprehensive list of insects from a temperate mangrove system is provided in
the Australian Littoral Society Report of Towra Point, Botany Bay (1977). The only
detailed study of the insect and spider communities of mangroves has been carried out
by Simberloff and Wilson (1970) and Wilson and Simberloff (1969a, b) in the
Florida Keys and more recently in north Queensland*. Simberloff and Wilson
fumigated small mangrove islands and observed the recolonization of the islands with
time.
Most work in Australia on mangrove insects has been on species of potential
commercial importance (termites) or which may be involved in the transmission of
disease (mosquitoes, biting midges) .
Termites are an important component of tropical mangroves. Three species of
Kalotermitidae occur in the Darwin region: Neotermes insularis, Cryptotermes
secundus and C. domesticus. C. domesticus is widely distributed in South-East Asia
and the Western Pacific and occurs in both natural and man-made habitats. Whether
the mangrove populations in Australia are indigenous, or were introduced is not
known. Cryptotermes secundus has also been recorded from Rhzzophora at
Kalumburu in Western Australia, and from Avecennza at Groote Eylandt. At Cooloola
in southern Queensland Cryptotermes primus occurs in species of Avecennza and
*The extensive insect collections made by Simberloff on Hinchinbrook Island have been deposited with Dr
R. Taylor, C.S.I.R.O.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
90 FAUNA OF AUSTRALIAN MANGROVES
Rhizophora. At Weipa, mangroves support large colonies of Inczsttermes barrettz.
Another species, Nasutztermes graveolus, is common on the landward fringe. This
termite builds arboreal nests externally on the trunk or branches. Some species may
range below high tide level. Mastotermes darwiniensts has been recorded in this
situation on Cerzops in the Darwin region, but this is unusual (Miller and Watson,
pers. comm.). The location of this termite at high tide is unknown. It may nest in the
forest canopy. The same species has been recorded from Corio Bay, Queensland, at
the southern limit of distribution of Mastotermes on the Queensland coast (Ellway,
1974).
The grasshopper Valanga irregularis has been seen feeding in the mangrove
canopy at Corio Bay (Ellway, 1974). The pygmy grasshopper Coptotettzx mastricatus
has been recorded on the mud of mangroves. In the New World mangroves, many
species of crickets and a number of tettigoniids occur (Rentz, pers. comm.).
The beetle fauna of mangroves is diverse and largely undescribed. It probably
includes species which feed on dead wood as larvae, others whose larvae feed on
mangrove seed pods, while the adults feed on foliage. Some weevils have been
recorded from mangrove seed capsules (Zimmerman, pers. comm.). MacNae (1968)
records coccids covering the leaves and young twigs of Rhizophoracea (species not
given) in north Queensland.
Diptera occur in mangroves as both free living and parasitic forms and are a
diverse component of the mangrove fauna. Fourteen species were collected by
Hutchings and Recher (1974) at Careel Bay, near Sydney. Some Diptera, such as
Melanagromyza avicenniae which is restricted to mangroves, appear to feed on new
shoots. Diptera are also associated with rotting wood. The adults and larvae of
Copidita nigronotata are secondary timber borers. Other dipterans specialize on the
fruits and propagules of mangroves (see Hutchings and Recher, 1974, and Spencer,
WIT) «
According to Common and Waterhouse (1972) several species of butterflies are
commonly associated with mangroves. All belong to the Lycaenidae (blue
butterflies). Hypochrysops epicurus which occurs from Port Macquarie to Brisbane,
appears to be restricted to stands of Avicennza marina. Hypochrysops apelles and H.
narcissus are characteristic elements of the mangrove fauna, although their larvae also
feed on non-mangrove species. Hypochrysops apelles which occurs along the east
Australian coast from north of Yeppoon, feeds on Rhizophora stylosa, Bruguzera
gymnorhiza, Ceriops tagal and Avicennia marina. Ogyris amaryllis hewttsonz which
occurs from Maryborough to Cairns, is common around mangroves, as its larvae feed
on the mistletoe A myema mackayense which grows on mangroves.
A less common butterfly Nacaduba kurava occurs from the Richmond River
(New South Wales) to Cape York. It feeds on Aegiceras corniculatum but also uses
other plants. The larvae and pupae of Acrodtpsas illidgez have been found in the nests
of the ant Crematogaster laeviceps near Brisbane. The ant nests in hollow branches of
mangroves. Three to four larvae or pupae of Acrodipsas were present in each colony;
it seems likely that the larvae of A. zlledgez feed on the immature stages of the ant
(Common, pers. comm.). Similarly, Hypochrysops appollo is associated with ants in
mangroves, as it breeds in ant house plants (Saenger et al., 1977).
The larvae of the tortricid moth Procalyptis parooptera feed on the leaves of
Ceriops tagal at Yeppoon, firmly joining adjacent leaves with silk to form a shelter in
which the larvae live. Pupation occurs in this shelter. This empty shelter is then used
by the larvae of the butterfly Hypochrysops apollo as a daytime retreat where it is
attended by ants and from which they emerge at night to feed on the surrounding
foliage (Common and Waterhouse, 1972). The moth Macrocyttara expressa
Proc. LINN. Soc. N.S.W., 106 (1), (1981) 1982
P. A. HUTCHINGS AND H. F. RECHER 9)
(Cossidae) is restricted to mangroves. The larvae of this species tunnel gregariously in
the trunks of Excoecaria agallocha. The larvae of Cenoloba_ obliteralis
(Oxychirotidae) develop in the cotyledons of fallen seeds of Avicennia marina
(Common, 1970).
All the Lepidoptera mentioned occur in eastern Australia but other species
closely associated with mangroves probably occur in other regions. Two unidentified
species have been recorded from South Australian mangroves (Butler et al., 1977).
The ant fauna of Australian mangroves has not been studied in detail but many
species occur. No species has been found which is totally restricted to mangroves. For
ants, colonization of mangroves means coping with the tidal cycle. Thus most
mangrove ants are arboreal and several species nest in hollow twigs. Dead mangroves
provide shelter for three species of ants (not identified) in Corio Bay, Queensland
(Ellway, 1974). Some species nest in the canopy while other ants nest on the ground
where they are inundated by the tide (Taylor, pers. comm.). Overseas, the ant fauna
in the Florida Keys has been well studied by Wilson (1964).
MacNae (1968) records the weaver ant Oecophylla smaragdina in Bruguiera,
Cerzops and Sonneratia forests in north Queensland. This species has a similar life
history to species of Oecophylla occurring in coconut palm forests in Zanzibar which
have been described by Vanderplank (1960). MacNae also suggests that species of
ants which occur below high tide level may be able to trap air in their burrows with
plugs of mud. Whether ants living in inland areas where freshwater flooding regularly
occurs, adopt a similar strategy is unknown. If they do not, then the ants found in
mangrove muds have developed a unique adaptation enabling them to invade the
intertidal environment. An unidentified species occurs at low tide in the Hinchinbrook
region (Hutchings, pers. obs.) and presumably adopts this strategy. Another
unidentified species at Hinchinbrook has been seen living in tunnels made by Teredo
in the prop roots of Rhizophora.
Two species of ant house plants Myrmecodia antoinim and Hydnophytum
formicarum, occur in northeastern Queensland. Their tuberous stems are hollow and
often used by ants of several species. The ant Phezdole myrmecodizae is restricted to
this habitat, where it obtains protection from desiccation and predation. The
advantages to the plants are less clear, but the ants may protect them from herbivores.
In contrast to other groups of insects, there is considerable information about the
mosquito (Lee et al., 1980; Griffiths, in ms.) and biting midge fauna of mangroves.
Mosquitoes are typically found in the ephemeral pools at the back of the mangroves,
rather than within the mangroves where there is regular tidal exchange. In disturbed
mangroves, stagnant pools may be created as a result of dredge and fill operations.
These pools are ideal mosquito habitat. The three most common mosquito species
occurring in Australian mangroves are Aedes vigilax, A. alternans and Culex sttiens.
The distribution of A. wgzlax is largely coastal, with a special relationship to the low-
lying land of mangroves and mangrove zones (Sinclair, 1976) . It also occurs inland.
Typically, A. wgilax breeds in temporary waters and is intermittent in
occurrence. The water must be stagnant and exposed to the sun. Sunlight is necessary
for the growth of the algal plankton on which the larvae feed. Reproduction occurs
largely during the summer wet season. Aedes can complete larval development within
10 days. Low temperatures inhibit breeding and greatly prolong the duration of the
larval stages (Iyengar, 1965; Griffiths, in ms. and pers. comm.).
Aedes alternans occurs in all mainland States, and is particularly associated with
estuarine areas. It also occurs inland. A. alternans breeds in temporary stagnant
pools, both fresh and brackish, exposed to the sun. The eggs are laid singly (probably
on mud at the edge of drying pools) and can withstand drying. The eggs hatch when
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
92 FAUNA OF AUSTRALIAN MANGROVES
the depression fills with water (Marks, 1967, 1971). The eggs take seven days to hatch
under favourable conditions but below 21°C hatching is delayed. In coastal situations
the larvae of A. alternans are often found with those of A. wgzlax. The larvae of A.
alternans are predacious from the second instar onwards on other mosquito larvae.
The adults are strong fliers, and are known to migrate with A. wgzlax, but only in the
summer (Hamlyn-Harris, 1933).
Culex sittens occurs from New South Wales northwards, and westwards to
Western Australia. In the Northern Territory, Hill (1917) found C. sztzens abundant
near the coast and inland as far as 55 km south of Darwin, breeding in pools, hollow
stumps, tins, crab holes, mangrove swamps, weedy lagoons, inland waterholes and
shallow wet-season accumulations of water on grassland. Marks (1953) also found it in
a variety of habitats, in sunlit, muddy, brackish pools and in deep shade in mangrove
swamps, both amongst roots and in more open water, and in footprints on mud flats.
Three species of Anopheles also occur in mangroves. Anopheles amictus hilli
breeds most abundantly in brackish water, and occurs in north Australia from
Western Australia to the Queensland/New South Wales border. A. annulipes
occasionally breeds in brackish water and occurs throughout Australia. A. farautz is
found in the north of Australia from Western Australia to the east coast of
Queensland. It sometimes breeds in brackish water. Perry (1946) reports it breeding
in extensive brackish water pools in the Solomon Islands. These were high in organic
debris and subject to tidal fluctuations. Similar habitats in Australian mangroves are
likely to be suitable.
Biting midges belong to the family Ceratopogonidae and are erroneously called
sandflies. More than twenty species are associated with mangroves; many are still
undescribed. For most, knowledge is restricted to the adult form. This information is
largely based on emergence trapping which may or may not indicate larval habitat.
A considerable amount of work has been carried out on the biology of coastal
biting midges, with a view to developing control techniques. Debenham (1979, and
pers. comm.) has reviewed the biology of coastal species of biting midges. Of the
twenty-odd species, seven are common in mangroves. Culzcozdes henry occurs in
coastal southern Queensland, coastal and subcoastal New South Wales; it appears to
prefer muddy sand as a larval habitat, possibly with tree cover. The pupae float on the
water surface with their long axes parallel to the surface, and are unable to submerge.
The major source of blood is unknown, but people are attacked. Reye (1972a) ranks it
as the seventh most important pest midge of coastal Queensland, while noting that it
reaches pest proportions only in restricted localities. C. hzstr¢o occurs in Australia from
coastal northeastern Australia south to the Sydney region. Immature stages have been
collected from a Juncus pool at Careel Bay (New South Wales). At Townsville, C.
historto occurs on mangrove flats well inside the mouth of the Ross River. Populations
are maximal during the summer and negligible in winter (Kay and Fanning, 1974).
It has been only recorded feeding on birds. C. magneszanus occurs on the coast and
offshore islands of the Northern Territory and Queensland. It has been collected
biting humans (Lee and Reye, 1955), but it is probably an avian feeder. Activity is
nocturnal; peak activity in the Townsville area occurs between 2100 and 9300 hr (Lee
and Reye, 1961). C. marmoratus occurs in Australia from north Queensland to
southern New South Wales. Larvae occur in low-lying estuarine zones, often in
association with Salicornia, Sporobolus wirginicus or Sueda maritima. The eggs cannot
withstand desiccation and breeding occurs after spring tides which flood the salt
marsh. This species is known to occur outside its breeding area. It has been reported to
bite humans and can be a pest in coastal regions. Lee et al. (1963) suggest it is an
opportunistic feeder, with wallabies its primary native host. Biting activity is primarily
Proc. LINN. Soc. N.S.W., 106 (1), (1981) 1982
P. A. HUTCHINGS AND H. F. RECHER 93
crepuscular. C. molestus occurs throughout coastal eastern Australia. Reye (1972a)
has characterized the larval habitat as clean sand in the open or among trees disturbed
by slight to moderate wave or current action. A wide range of salinities is tolerated.
The larvae occur with the top 7.5 cm of sand, and there is evidence that the pupae will
drift in on a rising tide, be stranded and emerge there. This species has invaded the
sandy banks of canal (housing) estates in southern Queensland (Reye, 1971). C.
molestus has been recorded attacking people, but except in canal estates, it is not
considered a problem (Kettle et al., 1975). C. subtmmaculatus occurs throughout
coastal eastern Australia. Immature stages are found on sandy estuarine foreshores,
and in the Salzcornza zone. They have also been found in the vicinity of crab holes
(often associated with Avwcennza), and Reye (1969a,b) suggests that the presence of
soldier crabs Mictyrzs platycheles is essential for the breeding of this species. Kettle
(1977) has recorded the larvae feeding on polychaete worms and desmids. Mass
emergences appear to be correlated with neap tides. The adults are opportunistic
feeders. Activity is largely crepuscular but can be diurnal if humidity is high and there
is little wind.
The last common species is C. ornatus which occurs in the Torres Strait islands
and coastal northern Australia. It does not occur south of Tin Can Bay in Queensland
but this species has been often confused taxonomically giving it an erroneous
distribution. The larval habitat is the mean neap tide zone of estuarine areas, mud
substrate, completely sheltered from wave action, and with a dense tree cover, often
Rhizophora stylosa. Reye (1972a) has suggested that there is an obligatory association
with Aegiceras corniculatum. Adults are abundant in mangrove swamps, and Reye
(1972a) regards this species as the most important Queensland pest species. It also
feeds on flying foxes and birds.
Spiders
The spider fauna of Australian mangroves is undescribed but from casual
observations appears to be rich in species and individuals. Hutchings and Recher
(1974) recorded 18 species in temperate mangroves at Careel Bay, New South Wales.
At Towra Point, Botany Bay (New South Wales), 35 species were found (Australian
Littoral Society, 1977). Forty two species were collected from Trinity Bay, Cairns
(Hegerl and Davie, 1977), and 56 species in Corio Bay, near Rockhampton (Ellway,
1974). Far fewer species (6) have been recorded from South Australia (Butler et al.,
1977). These few observations do not allow for many generalizations.
The diversity of spiders associated with mangroves increases rapidly from
temperate to tropical zones, but all the spiders recorded from mangroves also occur in
other terrestrial habitats. McCormick (1978) suggests that spiders in mangroves may
be highly seasonal in occurrence and in areas subjected to regular flooding the fauna
may be severely restricted. From the information available, it appears that the spider
fauna of mangroves may be recruited from adjacent habitats.
Of the spiders found in mangroves the orb web building spiders are the most
evident. This group includes both the slant orb web weavers of the genus Tetragnatha
(the large jawed spiders) which elsewhere are associated with stream or lake-side
habitats and the vertical orb web weavers of the genus Eriophora. Both are adapted
for catching flying prey. Foliage dwelling spiders seem to be rather uncommon.
Other groups of spiders occur on the ground or on the lowest vegetation. Most
evident in these situations are the wolf spiders (Geolycosa sp,) and allied hunting
spiders of the family Pisauridae (Dolomedes sp.). Presumably these are opportunistic
hunters, foraging on the exposed mud at low tide. Dolomedes, however, is often
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
94 FAUNA OF AUSTRALIAN MANGROVES
associated with aquatic habitats and has been observed to move underwater to avoid
predators or take prey (Gray, pers. comm.).
Scorpions and mites have been reported from Hinchinbrook Island (Bunt, pers.
comm.).
Aquatic Invertebrates
This review is restricted to macro-invertebrates. Although microfauna such as
nematodes and protozoa is abundant in mangroves, its presence has been poorly
documented in Australia. Only a single study has been carried out on the nematode
fauna. Dacraemes and Coomans (1978) collected 25 species of nematodes from a
single sample in the mangroves at Lizard Island, Great Barrier Reef, suggesting that a
rich fauna is awaiting study. Overseas, more attention has been directed to the
microfauna (see Newell, 1974, for review).
The aquatic invertebrates of mangroves are primarily marine in origin with some
fresh water animals occurring at the freshwater, marine interface in the upper reaches
of the estuaries. In addition there is a group of marine pulmonate molluscs which is
probably related to terrestrial and freshwater pulmonate gastropods. However they
have had a long history of independent evolution (Zilch, 1959).
Within the intertidal zone, the mangrove fauna is dominated by polychaetes,
crustaceans and molluscs (Appendices 1,2,3). In addition to the intertidal fauna, tide
pools and channels with permanent water have a characteristic estuarine subtidal
fauna. Again this is dominated by polychaetes, crustaceans and molluscs, but
echinoderms, ascidians, coelenterates and sponges may be present (Saenger et al.,
1980) . Sponges and ascidians are also associated with sea grasses. This subtidal fauna
is arguably part of the mangrove system, but for the purposes of this review, we
consider only the intertidal fauna.
According to where they live, there are five distinct groupings of marine
invertebrates within the intertidal of mangrove forests: encrusting epifauna,
mangrove epifauna, substrate epifauna, substrate infauna and wood-boring infauna.
The composition of the fauna within each of these categories is determined by the
physical environment and there are distinct patterns of zonation (Hutchings and
Recher, 1974). Zones are determined by sediment structure, tidal and salinity regimes
and period of inundation. The distribution of mobile animals will also vary according
to the time of day and night. Thus the intertidal fauna changes through time and is
patterned on both a horizontal and vertical scale in space. Little of this complexity has
been quantified and the summary which follows must be accepted as a simplification
of an exceedingly complex situation. Studies which provide information on relative
abundances of marine invertebrates within mangrove habitats are limited to Butler et
al. (1974, 1975) (South Australia), Hutchings and Recher (1974), Hutchings et al.
(1977), Weate (1975), McCormick (1978) (New South Wales), Shine et al. (1973)
(Queensland) and Hegerl et al. (1979) (Northern Territory) .
Epifauna
The epifauna consists of sessile and mobile animals which live on the surface of
the forest floor, on mangrove trunks and on mangrove roots and pneumatophores.
Encrusting fauna
Throughout most of Australia, the encrusting fauna is dominated by the oyster
Saccostrea commerczalts and barnacles. The dense mass of these animals provides a
sheltered environment for a rich and mobile fauna of errant polychaetes, crustaceans
and gastropods. Encrusting animals occur throughout the mangroves, but require
inundation on each tide and are best developed at the edges of the mangrove forest
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
P. A. HUTCHINGS AND H. F. RECHER 95
where the flow of water is high and prolonged inundation occurs. As in other
intertidal environments (e.g. rocky foreshores), barnacles tend to dominate in the
upper reaches and oysters or mussels at lower tide levels.
In New South Wales, encrusting fauna has a similar pattern of zonation
irrespective of locality (McCormick, 1978). The greatest concentration of animals
occurs in the frontal areas. McCormick (1978) suggests that frequency of tidal
inundation and shade are the most critical factors affecting the abundance of this
fauna. In areas of low light intensity, epiphytic algae (Bostrychia sp., Caloglossa sp.,
Catenella sp.) grow over the pneumatophores of Avicennia and around the base of
tree trunks. These algae provide a microhabitat for small gastropods and amphipods.
The algae, together with oysters may in some places retain enough moisture for
spionid and nereid polychaete worms to occur. Broadly similar patterns and
communities of encrusting organisms have been reported from South Australia,
Victoria and Queensland.
The barnacle Elmznzus modestus and the serpulid Galeolaria caespitosa occur on
the pneumatophores and lower trunks of Avicennia marina in the upper part of
Spencer and St Vincent Gulf and at Ceduna in South Australia (Womersley and
Edmonds, 1958). In Victoria, the fauna on Avcennza is characterized by the snail
Bembiccum melanostomum, the mussel Mytzlus edulis planulatus, and the barnacle
Chamaesipho columna (Smith et al., 1975). The Sydney rock oyster Saccostrea
commercialis and the barnacle Balanus amphitrite dominate the encrusting epifauna
on Avicennza in New South Wales (Dakin et al., 1952; Hutchings and Recher, 1974;
Hutchings et al., 1977). The snail Lzttortna scabra and the isopod Ligza australienszs
are also typical of this community.
Weate (1975) working on the lower Myall River on the central coast of New
South Wales (32°S) recorded ten species of molluscs and four crabs living on
mangroves. Some, such as the gastropods Salenator solida and Meloszdula zonata,
grazed both on the trunks of Avicennza and the surface of the mud substrate. Other
molluscs, including Patellocda mimula, Xenostrobus securis, Tatea ruftlabris and
Lasaea australis occurred only on the encrusting oyster Saccostrea commerczalis. On
the central coast of New South Wales, Hutchings and Recher (1974) recorded 26
species of molluscs and crabs living on or in the mangroves at Careel Bay (Appendices
2 and 3). The mangrove epifauna at Brooklyn, also on the Hawkesbury River estuary
but upstream from Careel Bay, has a less diverse fauna, although the total number of
individuals appears similar (Hutchings et al., 1977; pers. obs.). Species such as
Balanus amphitrite, Paragrapsus laevis, Austrocochlea constricta and Melostdula
zonata are absent at Brooklyn, but common at Careel Bay. The difference is probably
the result of lower salinities and possibly of higher silt loads at Brooklyn.
One of the most conspicuous animals in the mangrove epifauna is the gastropod
Littorina scabra. L. scabra occurs throughout the mangrove forest and can be
extremely abundant. In Queensland and northern Australia, L. scabra occurs with a
second species of Littorzna, n. sp., (Ponder, pers. comm.). Other molluscs on
mangrove trunks include Cerzthedea obtusa and Nerita lineata. Oncis sp., a
pulmonate slug, is also present.
Several studies in Queensland mangroves have not recorded the precise habitat
where species were collected (e.g. Hegerl and Davie, 1977; Shine et al., 1973).
However, the mangrove epifauna seems to be more diverse with decreasing latitude.
Saenger et al. (1979) studied subtidal fouling organisms in the Calliope River,
Queensland. Several of the species which colonized their settlement plates also occur
on pneumatophores and tree trunks in the frontal zone of mangroves. These include
such species as the barnacle Balanus amphitrite, the serpulid polychaete Fecopomatus
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
96 FAUNA OF AUSTRALIAN MANGROVES
uschakow and the oyster Saccostrea commercizalis. Yhey found that Balanus,
Ficopomatus and the bryozoan Electra sp. characterize a pioneer community which is
replaced by a community dominated by Saccostrea, Xenostrobus, Balanus and
Ficopomatus. A mosaic of pioneer and climax phases is the common condition found
on naturally occurring substrates such as mangrove pneumatophores and tree trunks
in the study area.
Two species of oysters (Saccostrea commerczalis and S. echinata) have been
recorded from mangroves adjacent to the coral reef at Low Isles. The gastropod
Morula was a common associate (Stephenson et al., 1958). A recent survey of the
fauna of mangroves in the Kakadu National Park, Northern Territory, by Hegerl et
al. (1979) does not indicate specific habitats for the molluscs collected, although the
encrusting oysters Saccostrea echinata and S. commerczalis were collected. Barnacles
were common on the trunks of Sonneratia, Camptostemon and Rhizophora, but were
not identified.
Substrate Epifauna
The animals which live on the surface of the forest floor are mainly molluscs.
Crabs are also common, but the majority live in burrows, coming out only to forage
and are discussed under infauna.
The molluscs are dominated by gastropods which are zoned in relation to tidal
inundation. McCormick (1978) found that the frontal zone of the mangroves is the
most diverse, although the diversity varied at the six sites he studied along the New
South Wales coast. He found no trend towards increasing diversity from southern to
northern New South Wales. However the sites varied greatly in their salinity regimes
and mangrove structure which may explain these findings. Except for McCormick’s
(1978) study seasonal fluctuations in substrate epifauna have been almost completely
neglected. He found maximum densities during the winter and spring.
Five families of molluscs dominate the epifauna, the Neritidae, Littorinidae,
Potamididae, Cerithiidae and Ellobiidae (MacNae, 1968). Scattered information
exists on the preferred habitats within the mangroves for some species. The two species
of Terebralia, T. palustris and T. sulcata, prefer muddy substrates whereas species of
Cassidula occur among decaying vegetation. The pulmonate slugs Onchzdium sp. are
cryptic, burying themselves in mud or hiding under debris. Stephenson et al. (1958)
found Onchidium, Quoyia and Bembicium concentrated among shingle under
Bruguzera bushes at Low Isles. Many gastropods occur throughout the mangroves, on
the surface of mud, on pneumatophores and in association with logs.
The number of molluscs in the surface epifauna increases from the temperate
zone to the tropics. Although almost certainly underestimates, 16 species have been
recorded in South Australia, 24 in Victoria, 33 in New South Wales and 57 in
northern Queensland (Appendix 3). There are distinct temperate and tropical
elements to this fauna.
Conuber melanostoma and C. sordida (Naticidae) are restricted to temperate
mangroves and are absent from central Queensland and further north. In contrast
Cassidula augulifera (Ellobiidae) is restricted to tropical mangroves although C.
augulzfera does extend as far south as southern Queensland. In the family Neritidae, 6
species are restricted to tropical or sub-tropical mangroves and only N. atramentosa
occurs in temperate mangroves and southern Queensland. As more information on the
molluscs becomes available the distribution patterns will become clearer and will
probably continue to substantiate the pattern of greater diversity of molluscs in
tropical mangroves than in temperate systems (Fig. 3).
MacNae (1968) hypothesized that the molluscs occurring in temperate
mangroves such as Westernport Bay and the gulfs of South Australia are typical
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
P. A. HUTCHINGS AND H. F. RECHER 97
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Fg. 3. Several groups of marine invertebrates appear to be more abundant and richer in species in the
tropics than in the temperate zones. Data on molluscs and crustaceans are most complete and are used here
to illustrate this point for Australian mangrove forests.
estuarine species and that specialized mangrove species are restricted to tropical areas.
MacNae (1966) also suggested that the fauna associated with Sydney Harbour
mangroves is a depauperate tropical one with all the species derived from Queensland.
However he based this statement on incomplete species lists provided by Dakin e¢ al.
(1952). The mangrove mollusc fauna near Sydney is in fact quite rich and is a mixture
of tropical and temperate forms.
There is relatively little information on the abundance of any of these animals.
Hutchings et al. (1977) recorded densities of 0.4 individuals per m? for Littorzna
scabra and 55.5 individuals per m? for a species of Tatea in the mangroves at Brooklyn
on the Hawkesbury River. Numbers of Tatea can exceed 10,000 individuals per m? in
salt marsh at the edge of mangroves (Ponder, pers. comm.). Butler et al. (1975)
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
98 FAUNA OF AUSTRALIAN MANGROVES
provide a list of macrobenthic animals found in mangrove areas at different locations
in South Australia and rank these by relative abundance.
Except for McCormick’s (1978) work in New South Wales, there are no measures
of the seasonal change of the substrate epifauna. McCormick (1978) found that
maximum densities occur in winter and spring at Towra Point on Botany Bay and
Patonga Creek on Broken Bay, Hawkesbury River (Fig. 4).
Substrate Infauna
This fauna includes animals that live in the sediment for all or part of their lives.
It is restricted to the upper layers of mud and consists of sedentary and mobile
animals. The mobile species such as crabs feed on the surface of the mud. The
majority of the fauna live in semi-permanent or permanent burrows. Burrows
penetrate through the oxygenated layers of mud into the anaerobic layers, where they
are restricted in depth by the fibrous matted root systems of the mangroves. Some of
the terrestrial fauna such as ants may trap air in their burrows during high tide and
marine animals such as crabs trap water during low tide.
The infauna has been neglected in mangrove faunal studies and well-planned,
long term collecting throughout Australian mangroves is needed. The sedentary
animals are represented by sipunculans, echiuroids, polychaetes and nemerteans.
There is a single species of sipunculan, Phascolosoma arcuatum. It occurs in tropical
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Fig. 4. McCormick (1978) studied the horizontal distribution of animals in several areas of mangroves in
New South Wales. These data on species number for the mangrove stands at Patonga on the Hawkesbury
River are presented in this figure. Both epifauna and infauna are richest in species at the lower tide levels
where there is regular inundation by the tides. The terrestrial invertebrate fauna is most abundant in the
middle zones where refuges during high tide occur and absent from frontal areas (area 3) which are covered
by high tides.
Proc. LINN. Soc. N.S.W., 106 (1), (1981) 1982
P. A. HUTCHINGS AND H. F. RECHER 99
mangroves in Australia and extends into South East Asia (Edmonds, 1980).
Phascolosoma occurs throughout the mangroves, constructs extensive galleries in the
substrate, and may occur in large numbers. The echiuroid Ochetostoma australiense
occurs on the mud flats in front of the mangroves from northern New South Wales to
southern Queensland. This species has a commensal crab, bivalve and gastropod
associated with it in Queensland (Ponder, pers. comm.).
Polychaetes probably occur in all mangrove areas in Australia although they have
rarely been collected systematically. The species which have so far been recorded are
listed in Appendix I but many more species probably occur. Polychaetes are restricted
to the wetter seaward margins of the mangroves or near regions of permanent water
(Hutchings, pers. obs.). They occur in the less consolidated sediments where
burrowing is easy. Burrows do not extend into the dense root systems of the
mangroves. Several families are represented but nereids, spionids and capitellids are
the most abundant groups. From the data available it appears that the polychaete
fauna is more diverse in tropical regions. Most of the species occurring in temperate
mangrove systems also occur in other estuarine habitats. Ai) exception is the terebellid
Hadrachaeta aspeta which occurs only in mangroves or on mud flats immediately
adjacent to the mangroves (Hutchings, pers. obs.). Whether more mangrove
specialists occur in the tropics remains to be investigated. Some of the species such as
the nereids and spionids construct semi-permanent mucous-lined tubes. Many of the
species occurring in the substrate also occur in rotten logs or in pockets of water
trapped among the encrusting epifauna on pneumatophores, prop roots and tree
trunks.
There are limited data on the abundance of polychaetes in mangroves. Hutchings
et al. (1977) and McCormick (1978) have some data for New South Wales. In
general densities of polychaetes appear low with less than one individual per m7’.
However, McCormick (1978) found that numbers changed seasonally and more
extensive sampling could reveal higher population sizes. Numbers of polychaetes in
Queensland mangroves appear to be higher than in New South Wales.
Nemerteans occur in frequently-inundated areas of mangroves where sediments
are relatively unconsolidated. A tropical species, Pantenonemertes winsort, occurs
beneath bark and in cavities in rotten wood of Avicennia marina and Ceriops tagal
(Moore and Gibson, 1981). At Careel Bay, Tubulanus polymorphus occurs in sea
grass beds (Zostera sp.) adjacent to the mangroves and probably extends into the
frontal zone of the mangrove forest (Hutchings and Recher, 1974). Turbellarians also
occur in mangroves throughout New South Wales (Hutchings and Recher, pers.
obs.), but have not been identified.
Molluscs are represented in the substrate of mangrove habitats by large numbers
of bivalves. McCormick (1978) reported up to four species from his sites in New South
Wales. Glauconome plankta, Arthritica helmsi sp., Laternula sp., Tellina deltoidalis
and Venerupis crenata were the most abundant species. Species diversity is higher in
tropical mangroves and a number of species may be restricted to mangrove habitats
(Fig. 3).
MacNae (1968) suggested that some mangrove bivalves literally form a ‘cocoon’
among the matted roots of the mangroves. Bivalves are most abundant along the
seaward fringe of the mangrove forest and decrease in abundance and diversity with
decreasing tidal inundation (McCormick, 1978).
Crustaceans are a conspicuous component of the substrate fauna (Appendix 2).
Crabs are the most obvious. The snapping shrimp Alpheus and the burrowing prawn
or nipper Callianassa as well as isopods and amphipods are less visible, but are
probably equally abundant. Certainly one of the most characteristic sounds of
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
100 FAUNA OF AUSTRALIAN MANGROVES
mangrove environments at low tide is the ‘snapping’ of alpheids in their burrows.
Crabs are represented by large numbers of species in several families (Appendix 2)
and are among the best known of the mangrove fauna (Fig. 3).
In tropical mangroves the burrows of the mud lobster Thalasstna anomola and
the mud crab Scylla serrata are conspicuous. Juvenile Scylla are particularly common
in mangroves although they also occur among sea grasses and the retention of
extensive mangrove forests may be critical for the management of this commercially-
important species. Once the juveniles reach a carapace width of 60-80 mm they tend
to move into subtidal areas, but continue to forage among the mangroves on the rising
tide (Hill, 1979, 1980, and pers. comm.). The use of mangroves is accentuated in
places where the density of sub-tidal benthic animals is low, perhaps as a result of
freshwater conditions during the wet season, such as in far north Queensland.
There is a distinct zonation of crab species within the mangrove forest. In
Western Australia, George and Jones (in press) correlated the distribution of fiddler
crabs Uca with sediment grain size. Elsewhere zonation appears to be determined
largely by period of tidal inundation. Yates (1978) studied the ecology of crabs in
mangroves at Patonga Creek on Broken Bay, New South Wales (33°S). Two
ocypodids Heloeczus cordiformis and Australoplax tridentata, are most abundant on
the seaward fringe of the mangroves with numbers decreasing towards land.
Heteropanope serratifrons and Ilyograpsus paludicola occur only on the seaward
margins. By contrast Sesarma erythrodactyla is most abundant in the highest zones
with numbers decreasing towards the seaward edge. Similarly, Helograpsus
haswellianus occurs only on the landward margins. Paragrapsus laevis is restricted to
the middle regions of the mangroves. Helice leachi, which occurs only in small
numbers, is restricted to the higher levels of the shore.
At Patonga there are only two species of mangrove, Avicenna marina and
Aegiceras corniculatum. The distribution and abundance of crab species cannot be
correlated with either species (Yates, 1978). In the tropics some species of crab are
restricted to particular zones and may be dependent on the species of mangrove.
According to MacNae (1968), Uca lactea is associated with Avicennia, especially
when it is growing in sand. If the sand is very fine U. bellator occurs and is also
common in areas shaded by outermost bushes of Cerzops. However, George and Jones
(in press) disagree with MacNae’s identifications and the precise nature of the
distribution of his Uca species in Australian mangroves must now be viewed with
caution. The larger sesarmids, being mainly herbivores, occur in areas of seedlings of
Avicenna, Bruguzera and Ceriops. Whereas Macrophthalmus depressus, an
omnivore, is restricted to sandy substrates in drainage channels (MacNae, 1968).
Yates (1978) recorded seasonal changes in the abundance of crabs. Maximum
densities of most species occur during summer. Paragrapsus laevts is most abundant in
winter. Some species, such as Sesarma erythrodactyla and Paragrapsus laevis, move to
lower zones in the mangroves to spawn. Crabs also show daily activity patterns. During
the day, at low tide, crabs tend to remain in their burrows. Hutchings and Recher
(1974) recorded maximum concentrations of crabs at night on a rising tide. Many of
the crabs were climbing pneumatophores and trees to forage.
Wood Infauna
The fauna associated with dead and living wood is not restricted to mangroves,
but because logs tend to be trapped and accumulate in mangrove habitats, mangroves
have a rich wood-boring fauna. In Australia, the wood-boring fauna is dominated by
the teredinid molluscs of which 31 species in eight genera have been described
(Turner et al., 1972; Turner and McKay, 1979). Holdich and Harrison (1980) have
Proc. Linn. Soc. N.S.W.,106 (1), (1981)1982
P. A. HUTCHINGS AND H. F. RECHER 101
recently described several species of isopods Gnathia from dead wood found in
Queensland mangroves.
Animals such as the ship worms (Teredinidae) and gnathid isopods are
important in the breakdown of wood in marine habitats. However, there is some
evidence that the wood must first be invaded by fungi before it can be colonized by
wood boring animals (Cragg and Swift, 1980). Leightley (1980) has found marine
fungi in the wood of the five species of Australian mangrove so far examined. The
succession of marine fungi in wood has been extensively studied in Florida by Newell
(1974) and similar patterns can be expected in Australia. In Papua New Guinea,
Cragg and Swift (1980) have drawn analogies between terrestrial decomposers such as
termites and beetles and the marine timber borers.
The wood-boring fauna is not restricted to dead wood. In Florida, an isopod
Sphaeroma terebrans attacks the live prop roots of Rhizophora mangle. Simberloff et
al. (1978) suggest that the damage by isopods coupled with insect attack on aerial
roots may stimulate root branching. For every root produced by the tree, 1.4 roots
reach the substrate thereby providing greater support for the plant. There is no
information for Australia on the wood-boring fauna of live mangroves.
Associated with these wood-boring organisms is a range of animals seeking
protection from predation, desiccation, etc., such as barnacles, limpets, crabs,
amphipods, nemerteans and polychaetes. This fauna is most diverse in logs trapped in
the frontal margins of the mangroves which are frequently inundated. Under logs,
small pockets of water are often trapped which provide suitable microhabitats for
many of these animals. In temperate mangroves the crab Sesarma erythrodactyla
commonly occurs under logs (Yates, 1978). Graham et al. (1975) found 34 species of
molluscs associated with logs in Trinity Bay, Cairns. These molluscs often use the
tunnels and cavities created by the wood-boring teredinids. Also, some species such as
Ellobtum aurtsjudea seek refuge in the log during the day, and feed on the muddy
substrates in the Rhzzophora and Bruguzera zones at night or on cloudy, rainy days.
One species of nemertean Pantznonemertes winsori has been recorded from tropical
mangroves (Moore and Gibson, 1981). It occurs beneath bark or in cavities in rotten
fallen timber of Awicennia marina and Ceriops tagal in the upper tide levels.
At Gladstone, central Queensland (23°S), many species of molluscs including
Melampus castaneus, M. striatus, Ellobtum aurisjudea, Onchidium sp.,
Bactranophorus sp., Nerita lineata, Isognomon ephippium have been collected from
logs (Saenger, pers. comm.). Microgastropods (e.g. Assemznae spp. and Iravadza sp.)
are also found on logs in tropical mangroves (Ponder, pers. comm.). Ponder has
evidence that differences in the mollusc fauna can be attributed to differences in the
moisture content of the log and the frequency of inundation.
In the Northern Territory several species of crab utilize rotting logs in the
mangroves of Kakadu National Park (Hegerl et al., 1979). Hegerl et al. (1979) found
that Metopograpsus quadridentatus, Metopograpsus sp., and Clzstocoeloma
merquzensis were most abundant in logs in the upper part of the tidal range.
Nannosesarma batavicum and Sesarma sp. occur in logs in the Rhizophora zone. S.
krausst borneensis is restricted to sites which are frequently inundated and Epzxanthus
dentatus occurs in Rhizophora, Sonneratia/Camptostemon zones. S. darwinensis
commonly occurs under decaying logs. Saenger (pers. comm.) collected the crabs
Clistocoeloma merguiense, Sesarma molluccensis and S. elongata from logs among
mangroves at Gladstone.
PATTERNS
It helps to understand the ecology of mangrove ecosystems if it is realized that
mangroves are essentially forests with a muddy intertidal substrate. The abundance of
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
102 FAUNA OF AUSTRALIAN MANGROVES
animals, their distribution within the mangrove forest, and their adaptations to the
physical and biological environment can then be explained in the same ways that one
would discuss a terrestrial forest or estuarine mudflat. For much of the fauna,
mangroves are no more than an extension of their ‘normal’ habitat. Relatively few ”
animals are restricted to mangroves or show specific adaptations to the mangrove
environment.
Mangrove endemics have probably evolved in response to a limited array of
unique features. Mangroves are flooded regularly by the tide, for varying periods
depending on the tidal cycle, and at varying times during the day. For terrestrial
organisms, this restricts movement and limits the amount of time available for
foraging on the forest floor. McCormick (1978) has suggested that the canopy fauna
in mangroves which is subject to partial or complete flooding of the canopy is
depauperate for this reason. High tannin content and concentrations of salt on or
within the leaves of mangroves cause special problems for herbivores. The resistance of
mangrove leaves to breakdown may also limit the ability of marine detritivores to use
them.
The mangrove community reflects the merging of the marine and terrestrial
systems. In temperate areas, at their most complex, mangroves form fairly open forests
with a simple canopy; in the tropics, mangroves may be open or closed and are often
layered with distinct zonation. The terrestrial fauna is largely restricted to the forest
canopy, and the marine fauna to the forest floor and lower levels of trees. Thus there is
a horizontal partitioning of these two parts of the mangrove community.
Within the two faunal groups, subdivisions occur. For terrestrial animals, these
divisions may be related to the horizontal zonation of the mangroves. For example,
stands of Rhizophora may provide vertical complexity, nesting sites or shelter for a
particular species of bird, whereas stands of Avicennia may not offer these advantages.
As mangroves are often a continuation of nearby forest, the partitioning within these
forests should be compared with that occurring in mangroves. Such comparisons may
explain, for example, why some birds which occur in adjacent habitats are absent
from mangroves.
For the marine fauna, mangroves provide additional habitat to that found on
muddy shores. The dense canopy of the forest provides protection against desiccation
and may offer cover against predators. Mangrove conditions are less rigorous in terms
of salinity and temperature change than those on an exposed mudflat. The less
rigorous environment and the firm substrates provided by roots, stems and logs enable
some animals to survive which might otherwise be absent from the intertidal
environment of estuaries.
As a generalization, most of the marine fauna occurring in temperate mangroves
also occurs on adjacent muddy or rocky shores. Mangrove specialists or endemics are
by and large restricted to the tropics (e.g. Figs 2 and 3). Either the tropical mangrove
forest offers habitats not available in tropical estuaries (e.g. solid substrates) or the
cooler conditions, high humidity and shelter from the sun within’ the mangrove forest
are critical for survival.
For the marine fauna, zonation horizontally across the substrate from low to high
water mark and vertically up roots and trunks and within the substrate, occurs in
response to the periodicity of tidal inundation (Fig. 4). This may be modified by
drainage patterns and tidal creeks within the mangroves. Behavioural, physiological
and morphological adaptations to water loss will largely determine the tide levels at
which particular species can live. The distribution of animals may vary during a tidal
cycle or during their life cycle; some species for example migrate seawards to spawn.
Like the terrestrial fauna, the distribution of the marine fauna may be determined by
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
P. A. HUTCHINGS AND H. F. RECHER 103
the zonation of the mangroves themselves. For example, the prop roots of Rhzzophora
and the buttress roots of Bruguzera may provide different microhabitats for marine
animals in comparison to the pneumatophores of Avicennia. The density of
pneumatophores, the amount of debris on the floor, and fallen logs are important for
cryptic species and wood fauna, in providing refuges and protection from desiccation.
Zonation of sediment within the mangroves is important to infaunal species. Also the
density of seedlings and the canopy cover which provide shade, affect animals living on
the surface of the mud. Salinity gradients along a river will determine how far each
species penetrates upstream.
Superimposed upon all these factors are latitudinal gradients along the coast.
The number of mangrove species increases from south to north with the greatest
diversity occurring in northeastern Queensland and northwestern Western Australia
(Lear and Turner, 1977) (Fig. 1). Coincident with the increasing number of species,
the extent of mangrove forests and their structural complexity also increase.
Although data are lacking for most tropical mangroves and for Western Australia in
particular, there appears to be a precipitous increase in the diversity of the mangrove
fauna which parallels that of the forest (Figs 2 and 3). As in the case of birds
(Schodde et al., 1982), the increased diversity of species is accompanied by an
increased number of mangrove specialists and endemics (Fig. 3). The three major
groups of marine animals, polychaetes, crustaceans and molluscs (Fig. 3), all increase
in species number from south to north (Appendices 1, 2, 3) and similar patterns can
be expected for other groups.
Pianka (1978) and others have reviewed the many theories advanced to explain
the increased diversity of tropical animal communities relative to those in similar
habitats within the temperate zones. It is not possible to separate completely the
different factors which may have led to the greater diversity of tropical systems. In the
case of mangroves, the greater structural complexity of the tropical forests, the
increased number of plant species and larger areas are all related to increased faunal
diversity. The area of mangroves and the distance between stands, in particular, may
explain the much greater numbers of birds which use mangroves as a primary habitat
in the tropics when compared to temperate mangrove forests. This is simply a logical
extension of the theories of island biogeography (see MacArthur and Wilson, 1967).
The increased structural complexity and greater number of plant species would also
permit a greater number of bird species to co-exist (Recher, 1971). Schodde et al.
(1982) provide examples of resource apportionment among closely related birds co-
existing in northern mangrove forests. The white-breasted whistler Pachycephala
laniotdes apparently forages extensively on the muddy forest floor where it takes
various marine invertebrates while the co-occurring mangrove golden whistler P.
melanura forages for insects in the canopy. Two co-habiting kingfishers Alcedo azurea
and A. pusilla differ in size (28 to 35 grams versus 10 to 13 grams) and would be
expected to take different sized prey. Other examples given by Schodde e¢ al. (1982)
include flycatchers Myagra, fantails Rhzpidura, and warblers Gerygone. Similar
patterns are evident among marine invertebrates with various crabs separated within
the mangroves by size of sediment particles (Uca spp.), type of food or capacity to
withstand desiccation.
As data become available, it is likely that significant regional differences in
community composition and diversity will be evident between tropical mangrove
forests in northern Australia. Stands of mangroves in northwestern Australia are
apparently much more uniform in species composition than those in northeastern
Australia. Historical events related to recent glacial periods and the separation of
northeastern and northwestern marine environments by the Torres Strait land bridge
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
104 FAUNA OF AUSTRALIAN MANGROVES
have probably had a significant effect on the evolution of distinctive mangrove
communities in these regions. During the time the land bridge existed there were
enhanced opportunities for plants and animals to move between mangrove forests in
eastern Australia and eastern New Guinea. These were isolated from mangroves
occurring along the western fringes of the continent.
Schodde et al. (1982) discuss the origins of the mangrove avifauna in the context
of the Torres Strait land bridge. They suggest that northwestern Australia was linked
to New Guinea by a mangrove-rich land connection, Arafura Land. This, they argue,
was the centre of evolution for mangrove bird specialists. In addition, the mangroves
in this area are geologically older and have been stable over geological time. Those in
northern Queensland have been affected by fluctuations in sea levels which
periodically eliminated mangrove areas (Coventry et al., 1980). Cape York was
connected to the Gulf of Papua which had relatively few mangrove forests and hence
fewer mangrove birds.
Whether similar patterns exist for other groups of mangrove animals is unknown.
As is evident from the material presented in this review, information on mangrove
fauna is sketchy. There is a particular need for quantitative studies of fauna and for
work in tropical mangrove forests. Until such information is available it will not be
possible to evaluate fairly the theories of Schodde and his colleagues or those of
MacNae (1968) regarding the evolution and dispersion of mangrove animals in
Australia. Nor, we submit, will it be possible to develop sound management and
conservation strategies for what must be regarded as one of Australia’s richest
ecosystems.
ACKNOWLEDGEMENTS
This review would have been impossible without the help of many people, and we
should like to thank especially H. Cogger, I. Common, P. Davie, M. Debenham, R.
George, M. Griffiths, D. Hoese, I. Loch, P. Marks, L. Miller, W. Ponder, D. Rentz,
E. Reye, C. Smithers, J. Taylor, T. Taylor, T. Watson, F. Wells and E. Zimmerman,
who provided unpublished information. We also thank J. Davie, M. Gill and T.
Kingston for reviewing the manuscript and A. Murray for artwork and K. Handley for
literature searching. However, we take full responsibility for any inaccuracies in
reporting the distribution of the fauna.
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Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
P. A. HUTCHINGS AND H. F. RECHER 107
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108 FAUNA OF AUSTRALIAN MANGROVES
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Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
P. A. HUTCHINGS AND H. F. RECHER 109
monitoring techniques. Ecology 50: 267-278.
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Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
110 FAUNA OF AUSTRALIAN MANGROVES
APPENDIX 1
POLYCHAETES RECORDED FROM AUSTRALIAN MANGROVES
NT Qld NSW Vict. SA WA
Nth Cent. Sth
POLYCHAETA
Polynoidae
Lepidonotus sp. xX
Sigalionidae xX
Amphinomidae
Eurythoe complanata (Pallas) ».4
Phyllodocidae
Phyllodoce duplex McIntosh xX
P. malmgreni Gravier Xx x
P. novaehollandiae Kinberg x
Phyllodoce sp. x
Nereididae
Australonerezs ehlers: (Augener)
Ceratonerezs erythraeensis Fauvel* xX
C. mirabilis Kinberg
xx x
es
Namalycastis abiuma Mueller
Namalycastis sp. x
Namanereis quadraticeps (Blanchard) x
Neanthes cricognatha (Ehlers) x
N. vaalu Kinberg x
Nerezs uncinula (Russell) :
Nerets sp. X
Perinerevs vallata (Grube) x
Platynereis cf. dumerili antipoda Hartman x
x x
Pseudonerets rottnestiana (Augener) Xx
Nephtyidae
Nephtys australiensis Fauchald xX Xx x
Glyceridae
Glycera americana Leidy
Glycera sp.
Gonzada sp.
x x
Eunicidae
Eunice antennata (Savigny) xX
Marphysa sanguinea Montagu Xx x
Marphysa sp. x
Lumbrineridae
Lumbrineris sp. x
Onuphidae
Diopatra dentata Savigny x x
Orbiniidae
Leztoscoloplos sp. Xx
Scoloplos (S.) simplex (Hutchings) xX
Scoloplos sp. Xx x
Spionidae
Boccardia sp. Xx
Polydora sp. xX
Prionospio sp. Xx
Scolecolepis indica Fauvel Xx
Spionidae spp. Xx
Magelonidae x
Chaetopteridae
Chaetopterus sp. x DS Xx
Proc. Linn. Soc. N.S.W.,106 (1), (1981) 1982
P. A. HUTCHINGS AND H. F. RECHER 111
APPENDIX 1
POLYCHAETES RECORDED FROM AUSTRALIAN MANGROVES
NT Qld NSW Vict. SA WA
Nth Cent. Sth
Mesochaetopterus minutus Potts x
Cirratulidae
Cirriformia tentaculata Montagu Xx
Opheliidae
Armandtia intermedia Fauvel x
Capitellidae
Barantolla lepte Hutchings x x
Capitella capitata (Fabricius) x
Notomastus torquatus (Hutchings &
Rainer) x x
Notomastus sp. Xx Xx
Maldanidae
Euclymene sp. x
Oweniidae
Owenza fuszformis Delle Chiaje xX X
Terebellidae
Hadrachaeta aspeta Hutchings Xx
Lysilla pacifica Hessle x
Pista sp. x
Rhinothelepus sp. xX
Thelepus setosus (Quatrefages) x
Amphitrite rubra (Risso) Xx
Terebella ehrenbergz Xx
Sabellidae : x
Laonome sp. xX
Serpulidae
Ficopomatus uschakow (Pillai) Xx
Galeolaria caespitosa Savigny x
Salmacina sp. Xx
*This species has recently been split into several species, by Hutchings and Turvey (manuscript) and
Hutchings and Glasby (in press) . Ceratonerezs erythraeensis probably does not occur in Australia.
Sources of references :
Butler et al., 1977; Hutchings and Recher, 1974; Hutchings et al., 1977; McCormick, 1978; Queensland
Museum (unpublished report), 1974; Saenger et al., 1977; also unpublished data of Hutchings.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
112 FAUNA OF AUSTRALIAN MANGROVES
APPENDIX 2
CRUSTACEANS RECORDED FROM AUSTRALIAN MANGROVES
NT Qld NSW Vict. SA WA
Nth Cent. Sth
CIRRIPEDIA
Iblidae
Ibla cumingz Darwin ».«
Balanidae
Balanus amphitrite Darwin X x X xX x xX X
Elminius modestus Darwin xX
Tetraclitidae
Tetraclita coerulescens (Splenger)
T. squamosa (Bruguiére)
T. vitiata Darwin
Chthamalidae
Chamaesipho columna (Spengler) xX
Chthamalus caudatus Pilsbury Xx xX
C. malayenszs Pilsbury xX Xx
x x
Tanaidacea
Paratanaidae
Paratanais sp. Xx
Isopoda
Anthuridae
Cyathura sp. xX
Haliophasma sp. x
Cirolanidae
Cirolana sp. x
Limnoriidae
Limnoria lignorum (Rathke)
Limnorza sp. x
Sphaeromatidae
Amphrovdella sp.
Eubranchiata sp.
Exosphaeroma ali Baker
E. alata Baker xX
Exosphaeroma sp. xX
Pseudosphaeroma sp.
Sphaeroma quoyanum Milne-Edwards Xx Xx Xx
S. terebrans Baker Xx
S. walkerz Stebbing
Sphaeroma sp. Xx x
Chitonopsts sp.
Ligiidae
Ligza australiensts (Dana)
Ligia sp. Xx
Amphipoda
Cheluridae
Chelura terebrans Philippi xX
Corophiidae
Corophium sp. xX xX xX
Ericthonzus sp. x
Gammaridae
Victorzopisa australiensts (Chilton) Xx
Talitridae
Orchestza sp. x
Talorchestza sp. xX
mK KM KK x -K KM *
*
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
P. A. HUTCHINGS AND H. F. RECHER 113
APPENDIX 2
CRUSTACEANS RECORDED FROM AUSTRALIAN MANGROVES
NT Qld NSW Vict. SA WA
Nth Cent. Sth
Hyalellidae
Parhyallela sp. x
Indet sp. x
DECAPODA
Penaeidae
Metapenaeus macleay: (Haswell) xX
Penaeus latisulcatus Kishinouye x
P. plebejus Hess Xx
Alpheidae
Alpheus edwardsi (Andouin) x x
Alpheus sp. xX
Atyidae
Paratya sp.: Xx
Palaemonidae
Leander intermedius Stimpson x
L. littoreus McCulloch x
Macrobrachium intermedzum Stimpson x
Palaemon serenus Heller Xx
Callianassidae
Callianassa australiensis Dana x Xx 4
Laomediidae
Laomediza healyi Yaldwyn & Wear Xx
Grapsidae
Eriocheir spinosa Milne-Edwards x
Xanthidae
Pilumnopaeus serratifrons Kinahan x
Ocypodidae
Scopimerinae
Ilyoplax dentata Ward
Ilyoplax sp. x
Scopimera inflata Milne-Edwards
Scopimera sp.
Tmethypocoelis sp. X
Heloecius cordiformzs (Milne-Edwards) x Xx x
Ocypode ceratophthalma (Pallas) x x Xx ».4
O. convexa Quoy & Gaimard
O. cordimanus Desmarest x ».« x Xx
O. fabricit Milne-Edwards
Uca capricornis Crane
Uca sp. A
Ucasp. B
Uca sp. C
Uca dussumierz (Milne-Edwards) Xx Xx
U. australiae Crane
U. coarctata Milne-Edwards Xx
U. flammula Crane X
U. signata (Hess) Xx
U. longzdigita (Kingsley)
U. seismella Crane x xX
U. polzta Crane xX X
U. tetragonon (Herbst) X
x KX
~ xX xX
~ x XM
mx xX
KK KM xX
~
*
~*~
Pata va Km KM mM mM OM
ea KK MK
xx mK MK
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
114 FAUNA OF AUSTRALIAN MANGROVES
APPENDIX 2
CRUSTACEANS RECORDED FROM AUSTRALIAN MANGROVES
NT Qld NSW Vict. SA
Nth Cent. Sth
U. dampieri Crane x
U. vomeris McNeill x x 4 x Xx
U. triangularts (Milne-Edwards) x x
U. mjobergi Rathbun x x Xx Xx
U. perplexa (Milne-Edwards) Xx x X
Macrophthalminae
Australoplax trindentata (Milne-Edwards) Xx x XxX ».4
Cletstostoma wardi Rathbun xX Xx x
Letpocten sordidulum Kemp xX Xx xX
Macrophthalmus abercrombiez Barnes x
M. boscw Audouin & Savigny xX xX x
M. convexus Stimpson Xx Xx Xx
M. crassipes Milne-Edwards XxX D4 x x x
M. darwinensis Barnes D4
M. japonicus (De Haan)
M. latifrons Haswell x
M. latrezllec (Desmarest) Xx Xx Xx ».«
M. pacificus Dana Xx x x x
M. punctulatus Miers xX xX Xx xX
M. setosus Milne-Edwards x x Xx
M. telescopicus (Owen) xX xX x xX
Paracleistostoma mcneilli (Ward) ».4 x x
Mictyridae
Mictyris liungstonez McNeill ».« Xx XxX Xx
M. longicarpus Latreille xX xX xX xX xX Xx
M. platycheles Milne-Edwards Xx Xx Xx
Grapsidae
Grapsinae
Metopograpsus frontalzs Miers xX x Xx Xx
M. latzfrons (White) xX x
M. quadridentatus Stimpson x
Metopograpsus sp. Xx x x x
Ilyograpsus paludicola (Rathbun) xX Xx
?Ilyograpsus sp. Xx
Varuninae
Varuna litterata (Fabricius) xX xX x x
Sesarminae
Clistocoeloma merguiense De Man Xx
Helice leachzi Hess
wm eM
x x xX
xx XK
~
Pad
Helograpsus haswellianus (Whitelegge)
Nannosesarma sp. A
Nannosesarma sp. B Xx Xx x Xx
Nannosesarma sp. C Xx x x
Paragrapsus gaimardz Milne-Edwards
P. laevis (Dana) x Xx
P. quadridentalus Milne-Edwards
x x xX
Sarmatium crassum Dana x x x
Sesarma (Chiromantes) brevicristatum x
Campbell
S. (Chzromantes) darwiniensis x
Campbell
’ Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
P. A. HUTCHINGS AND H. F. RECHER
MOLLUSCS RECORDED FROM AUSTRALIAN MANGROVES
S. (Chiromantes) liidum
Milne-Edwards
S. (Chiromantes) messa Campbell
S. (Chiromantes) semperz
longicristatum Campbell
S. (Chiromantes) semperz
2semperz Burger
S. (Holometopus) sp. 1
S. (Holometopus) sp. 2
S. (Holometopus) sp. 3
S
S
. (Parasesarma) erythrodactyla Hess
. (Parasesarma) leptosoma
Hilgendorf
(Parasesarma) ?moluccensis
De Man
(Parasesarma) sp. 1
. (Parasesarma) sp. 2
. (Bresedium) sp. (aff. brevepes)
(Bresedium) sp.
(Neoepzsesarma) sp. (aff. brockzz)
(Neoepisesarma) sp.
(2Neoepisesarma) sp.
. (Neosarmatium) fourmanozri
Séréne
)
ANNHAHAHHAHY
. (Neosarmatium) meznertz De Man
. (Neosarmatium) sp.
. (Neosesarma) sp.
. (Sesarmozdes) kraussz
borneenszs Tweedie
S. (Tiomanium) indica Milne-
Edwards
Xanthidae
Epixanthus dentatus (White)
E. frontalis Milne-Edwards
Heteropanope glabra Stimpson
H. serratifrons (Kinahan)
AnHnAN
?Heteropilumnus sp.
Myomenippe fornasinz (Bianconi)
Unident. sp.
Goneplacidae
?Rhizopa sp.
Speocarcinus sp.
Portunidae —
Scylla serrata (Forskal)
Hymenpsomatidae
Elamenopsis aspinifera Lucas
E. lineata Milne-Edwards
E. octagonalis (Kemp)
E. torrensica Lucas
Elamenopsis sp. 3
Elamenopsis sp. 4
Halicarcinus bedford: Montgomery
APPENDIX 2
NT Qld
Nth Cent.
xX
x x
x xX
x
x
x x
X x
x xX xX
xX
x xX
xX
x x xX
xX
xX
xX
Pr
xX x
xX
x xX xX
x
xX xX xX
xX
xX xX xX
xX
x
xX
xX
X xX xX
xX
xX MK
xX xX
xX
x
XK xX
xm x
Xx
NSW Vict.
x
Nth
x xX
xX x
xX
SA
115
WA
Xx
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
116 FAUNA OF AUSTRALIAN MANGROVES
APPENDIX 2 (continued)
Amarinus laevis (Targioni Tozzetti) Xx x
A. paralacustris (Lucas) Xx Xx xX Xx
A. latinasus Lucas x
Hymenosoma hodgkint Lucas x x Xx x
Sources of References:
Aust. Litt. Soc., 1977; Barnes, 1967, 1971; Butler, 1973, 1974; Butler et al., 1975, 1977; Campbell, 1967;
Campbell and Griffin, 1966; Crane, 1975; Davie (unpub.); Ellway, 1974; George and Jones (in press) ;
George and Knott, 1965; Graham e¢ al., 1975; Hale, 1927; Hegerl and Tarte, 1974; Hegerl et al., 1979;
Holdich and Harrison, 1980; Hutchings and Recher, 1974; Hutchings et al., 1977; Lucas, 1980;
McCormick, 1978; McNeill, 1926; Shanco and Timmons, 1975; Shine et al., 1973; Saenger, 1977; Ward,
1933; Weate, 1973 (unpub.), 1977; Wescott, 1976.
Davie in his unpublished manuscript has verified many of the decapod identifications but the other
crustaceans are based entirely on the literature. So some of the identifications and their distributions may be
inaccurate.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
P. A. HUTCHINGS AND H. F. RECHER 117
APPENDIX 3
MOLLUSCS RECORDED FROM AUSTRALIAN MANGROVES
NT Qld NSW Vict. SA WA
Nth Cent. Sth
POLYPLACOPHORA
Acanthozostera gemmata Iredale & Hull xX x x
GASTROPODA
Fissurellidae
Montfortula rugosa (Quoy & Gaimard) x Xx
Patellidae
Cellana tramoserica (Sowerby) Xx Xx x x
Acmaeidae
Patelloida mimula (Iredale) x x x x xX
Trochidae
Astrocochlea adelaidae (Philippi)
A. concamerata (Wood) XxX
A. constricta (Lamarck) x Xx Xx Xx Xx
A. torr¢ Cotton & Godfrey
Euchelus atratus (Gmelin)
Montodonta labio (L.)
Phastanotrochus sp.
Prothalotia comtessz Iredale
Neritidae
Nerzta atramentosa Reeve
N. albicilla L.
N. chamaeleon L.
N. lineata Gmelin
N. planospira Anton
N. plicata L.
N. ualenensis Lesson
Nerztena crepidularza Lamarck
Littorinidae
Bembicium auratum (Quoy & Gaimard) x
B. melanostomum (Gmelin)
Littorina acutispira Smith
L. irrorata Say
L. scabra (L.)
L. undulata Gray
Peasvella tantilla (Gould)
Hydrobiidae
Hydrobia buccinoides (Quoy & Gaimard)
Posticobia braziert (Smith) x
Tatea kestevenz Iredale
T. rufilabris (Adams)
Stenothyridae
Stenothyra sp. x X x
Iravadiidae
Travadza sp. ».« xX x
Assimineidae
Assiminea relata Cotton 4 Xx
A. tasmanica (1. Woods) Xx
Assiminea sp. Xx x
Turritellidae
Turritella terebra (L.) Xx x
Xx
xX
m~ X
mm
wm
*
ra
*
x xX “KM
mK K mm KK
rad ms x x MX
~*~ xX me Mm Mh Mm OM
mr xX x Kx
~
x KM
mm ra ~ mM mM
Planaxidae
Planaxis sulcatus (Born)
Quoyza decollata (Quoy & Gaimard)
Rissoidea
Rissozna sp. x xX
Vitrinellidae
~ x
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
118 FAUNA OF AUSTRALIAN MANGROVES
APPENDIX 3 (continued)
Pseudoliota sp.
Potamididae
Cerithidea largilliertz (Philippi)
C. obtusa (Lamarck)
C. cingulata (Gmelin)
Pyrazus ebeninus (Bruguiére)
Telescopium telescopium (L.)
Terebralia palustris (L.)
T. sulcata (Born)
Velacumantus australis (Quoy &
Gaimard)
Cerithiidae
Clypeomorus carbonarius (Sowerby)
Strombidae
Lambis lambis (L.)
Strombus luhuanus (L.)
Naticidae
Polinices conicus (Lamarck)
P. melanostoma (Swainson)
P. sordida (Swainson)
Muricidae
Bedeva hanley: (Angas)
Cronia aurantiaca (Hombron & Jacquinot)
Homalocantha secunda (Lamarck)
Lepszella vinosa (Quoy & Gaimard)
Morula marginalba (Blainville)
Naguetia capucina (Lamarck)
Buccinidae
Cominella eburnea (Reeve)
C. lineolata (Lamarck)
Melongenidae
Volema cochlidium (L.)
Nassariidae
Nassartus burchard: (Dunker zn Philippi)
N. cf. dorsatus (Roding)
N. oltvaceus (Bruguiére)
N. melaniozdes Reeve
N. pauperatus (Lamarck)
N. jonasiz (Dunker)
N. pullus (Linné)
Collumbellidae
Columbella duclostana
Pyrene scripta (Lamarck)
Zafra sp.
Mitridae
Mitra retusa Lamarck
Bullinidae
Bullina lineata (Gray)
Haminoeidae
Atys naucum (L.)
Haminoea sp.
Ellobiidae
Cassidula angulifera (Petit)
C. nucleus (Gmelin)
C. rugata Menke
C. sowerbyana (Pfeioffer)
Melampus striatus (Pease)
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
NT
ta
em KM KM
axe KM
Qld
Nth Cent.
x x
x
x
Xx xX
x x
XxX
x ».4
>.< xX
xX
x
xX x
x
x
XxX x
x
x
x
x
x
x
x
Xx X
x x
X x
x X
x
x x
NSW Vict. SA WA
Sth
Xx
x
x Xx
x
x x Xx
x Xx
x
x x
x x x X
x x
x x x x x
x x x
x xX x
x x x x
x x
x
x x
x
x
x
x
x x x
x
x
x
x
x
X
x
x
x x
x
x
P. A. HUTCHINGS AND H. F. RECHER 119
APPENDIX 3 (continued)
NT Qld NSW Vict. SA WA
Nth Cent. Sth
M. castaneus (Muehlfeldt)
Ellobium aurisjudae (L.) Xx
Melosidula granulosa Iredale
M. zonata (Adams & Adams)
Ophicardelus ornatus (Férussac)
O. quoy: (Adams & Adams)
O. sulcatus (Adams & Adams) xX
Pythia scarabaeus (L.) x
Amphibolidae
Salinator burmana (Blanford)
S. fragilis (Lamarck)
S. solida von Martens
Siphonariidae
Stphonaria bifurcatus Reeve
S. dentzculata Quoy & Gaimard
S. marza (Iredale)
Onchidiidae
Onchidina australis Semper
O. beutschlw Stantschinsky
O. daemelu Semper
O. verruculatum Cuvier xX xX
Oncis chameleon (Brazier)
Oncis sp. x
Aplysiidae
A plysza sp. xX
Bursatella leachi (Blainville)
BIVALVIA
Solemyidae
Solemya teraereginae (Iredale) x
Arcidae
Anadara trapezia (Deshayes) ».4 x D4
Arca sp. x
Barbatia sp. x x
Mytilidae
Amygdalum beddomez Iredale x
Modiolus inconstans (Dunker) Xx Xx
Musculus variocosus (Gould)
Mytilus edulis L.
Trichomya hirsuta (Lamarck)
Xenostrobus pulex (Lamarck) x
X. securis (Lamarck)
Isognomonidae
Isognomon isognomon (L.)
Tsognomon’ nucleus (Lamarck) Xx x
Isognomon cf. wtrea (Reeve) X
Isognomon sp. Xx
Pinnidae
Pinna bicolor (Menke) x Xx x
Pteriidae
Pinctada margaritifera (L.) xX x
Ostreidae
Ostrea nomades Iredale xX
Saccostrea amasa (Iredale) Xx x xX
S. cuccullata commerczalis
(Iredale & Roughley) xX x x Xx Xx
S. echinata (Quoy & Gaimard) x
Anomiidae
Xx
maxx KM KM
~*~
mm mM MK
mx mM
mx Xs
xx KK mx
xx KK mw
~
em
~*~ x
mx x
xxx KM
~*~ x
*
xx x
x KM eM OM
Km x KM
Proc. Linn. Soc. N.S.W.,106 (1), (1981) 1982
120 FAUNA OF AUSTRALIAN MANGROVES
APPENDIX 3 (continued)
Enigmonia aenigmatica (L.)
Patro australis (Gray)
Lucinidae
Arthritica helmsz (Hedley)
Cavatidens omissa Iredale
Leptonidae
Lasaea australzs (Lamarck)
Geloinidae
Batzssa violacea (Deshayes)
Geloina coaxans (Gmelin)
Veneridae
Circe tumidum (Bolten)
Paphia hiantina (Lamarck)
Tapes wathngi Iredale
Venerupis crenata (Lamarck)
V. crebrelamellata Tate
Tellinidae
Tellina deltordalis Lamarck
T. australis Deshayes
T. capsoides Lamarck
Mesodesmatidae
Mesodesma altenai de Rooij Schuiling
Mactridae
Notospisula trigonella (Lamarck)
Glauconomidae
Glauconome cf. cumingz (Prime)
G. virens (L.)
G. plankta (Iredale)
Teredinidae
Bankia australis (Calman)
. campanelata Moll & Roch
. carinata (Grey)
. bipalmulata (Lamarck)
. graciles Moll
. nord Moll
. rochi Moll
Bactranophorus thoracitzes (Gould)
Bactranophorus sp.
Dicyathifer manni (Wright)
Lyrodus massa (Lamy)
L. medilobata (Edmonson)
L. pedicellatus (Quatrefages)
L. bipartita (Jeffreys)
Nausitora dunlope: Wright
N. globosa Sivickis
Nototeredo edax (Hedley)
Teredo bartschi Clapp
T. johnsoni Clapp
T. navalis (L.)
T. furcifera Von Martens
T. clappi Bartsch
T. poculzfer Iredale
T. princessae (Sivickis)
T. trranularis Edmonson
Hiatellidae
Fluviolanatus armarus Laseron
Pholadidae
Bowswwns
xx KK
*
x KKK
Proc. LINN. Soc. N.S.W., 106 (1), (1981) 1982
me
~
~*~ Xs
ead wm mK
AK KKK KM
x Kx
AK mK mM KM KK OM
we
ax x x
~~ xX
mK mK
~
x x xX
nm x
~
axe KKM KM KX *
wx x
~*~ x
P. A. HUTCHINGS AND H. F. RECHER 12]
APPENDIX 3 (continued )
Martesza striata L. x
Laternulidae
Laternula creccina Reeve axe x
L. tasmanica (Reeve) x
L. vagina (Reeve) x x
Sources of references:
Aust. Litt. Soc., 1977; Butler, 1973, 1974, et al. 1975, 1977; Ellway, 1974; Graham et al., 1975; Hegerl
and Davie, 1977, et al. 1979; Hutchings and Recher, 1974; Hutchings et al., 1977; McCormick, 1978;
Shine et al., 1973; Saenger et al., 1976; Turner et al., 1972; Weate, 1973 (unpub.), 1975; Wells and
Slack-Smith, 1981; also unpublished data of Hutchings and Australian Littoral Society. The species in this
appendix are based entirely on the literature and the material has not been verified so some of the
identifications and their distributions may be inaccurate.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
Notes and Discussion
A. W.H. HUMPHREY, HIS MAJESTY’S MINERALOGIST IN NEW
SOUTH WALES,
1803-12 — ACOMMENT
R. J. FORD
Department of Geology, University of Tasmania, Box 252C, G.P.O., Hobart, 7001.
In his paper concerning the activities of the mineralogist, A. W. H. Humphrey,
particularly with respect to his mineral collecting in Van Diemen’s Land, Vallance
(1981) was puzzled by the lack of any subsequent knowledge of the occurrence of the
‘green garnets’ that were collected by Humphrey and Robert Brown during an
expedition to the Huon River district in 1804 and may have been like those offered for
sale in 1812 by Heuland.
The source of the ‘garnets’ and also of the ‘pitchstone’ mentioned by Humphrey as
associated with them has until recently been obscure. Observations now reported
indicate a distinctive provenance and, furthermore, show that Humphrey’s ‘green
garnet’ is an epidote with unusual habit.
A short summary of the geology of Port Cygnet is in order here, at least with
respect to the Cretaceous alkaline igneous rocks for which the district is noted. There
are three distinct groups of such rocks in the area:
(a)a group referred to as syenite porphyries that contain as main minerals
plagioclase as well as potash feldspar,
(b) rocks usually bearing phenocrysts of sanidine, and termed sanidine porphyries,
in which the feldspar is almost exclusively potash feldspar, and
(c)hybrid rocks, produced by reaction between potassium-rich magma (from
which the sanidine porphyries have formed) and pre-existing Jurassic dolerite.
The alkaline rocks occur at Port Cygnet as small dykes, sills and irregular intrusions
into Permian and Jurassic country rocks.
The rock of major significance here is the garnet trachyte described by Macleod
and White (1900). This is related to the sanidine porphyries, but lacks the porphyritic
texture due to absence of sanidine phenocrysts. It is most unusual in bearing
phenocrysts of light-brown spessartine garnet (Ford, 1967), represented by sporadic
euhedral crystals but more usually rounded though with some faces well developed,
and also less abundant phenocrysts of rounded, dark-green epidote grains. The usual
habit of crystallized epidote is columnar and striated with the crystal elongated
parallel to the ‘b’ crystallographic axis which is in contrast to that observed for this
rock (Fig. 1) where crystal faces are not developed. Massive and granular epidote is
more typical of skarns and contact metamorphic zones. Epidote in this rock from Port
Cygnet was not reported by Macleod and White, who were concerned with the garnet,
and, indeed, is not present in their specimen (No. X 1435), now in the collection of
the Tasmanian Museum. Both sets of phenocrysts range in size from less than 1 mm to
about 10 mm in diameter and commonly bear white rims of potash feldspar which has
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
124 NOTES AND DISCUSSION
Fig. 1. Garnet trachyte specimen from Port Cygnet showing rounded phenocrysts of epidote with white rims
of potash feldspar. One cent coin (17.5 mm diam.) added for scale.
apparently crystallized onto the phenocrysts (Fig. 1). The groundmass of the rock is
dense, dark grey in colour. It is made up of closely-packed fine-grained potash
feldspar laths showing a trachytic flow structure and also some patches of iron chlorite
and occasional biotite.
This rock, unique to the area, occurs as a near-vertical dyke, approximately
0.6 m thick and exposed for about 20 m on the shoreline of Port Cygnet north of
Langdon’s Point at the locality having co-ordinates 5217 750N, 506 800E on the
D’Entrecasteaux sheet of the Tasmanian 1:100 000 topographic series.
The origin of the epidote in the garnet trachyte is enigmatic. It shows no reaction
relationship to its environment and it is difficult to determine whether the phase is a
primary crystallate or a relict from older rocks. Bulk chemistry, particularly trace
elements and an MnO content of about 2%, suggests the trachyte formed from a
parent potassic magma which had been modified by assimilation of other material.
This is certainly the source of Humphrey’s ‘green garnet’. One would like to think
that he correctly identified the brown garnet by crystal form and then concluded that
the epidote was a green variety since these two minerals are texturally very similar in
the host rock. The fine, dark-grey groundmass could also constitute the pitchstone
matrix referred to by Humphrey (Vallance, 1981: 139).
The locality of the outcrop is marked on the accompanying map (Fig. 2). This
would confirm Vallance’s deduction of their having encountered the Port Cygnet
estuary on their way back to Hobart. It is quite possible that Humphrey and Brown
came to Port Cygnet at the point where the garnet trachyte outcrops, because within
about 100 metres north and south of the above, there are prominent outcrops of
sanidine porphyry dykes with spectacular euhedral phenocrysts of (010) tablets of
yellow-white sanidine crystals up to 40 mm maximum dimension, and having a swirl
structure in a green aegirine-bearing matrix. In contrast the outcrop of the garnet
trachyte is much more subdued. For someone like Humphrey who was looking for
minerals it would seem to be incredible that he would fail to comment on the sanidine
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
NOTES AND DISCUSSION
MAP OF HUON DISTRICT
Basalt centres
Garnet trachyte
RANELAGH
HUONVILLE
i
<=
w
=
&
w
Q
GRID) NORTH
DrENTRECASTEAUL
Fig. 2. Map of the Huon district showing localities referred to in the text
dykes had he seen them. Even today it is difficult to walk along the western shore of
Port Cygnet unless the tide is very low. Hence it is most likely that Brown and
Humphrey happened to come to the Port Cygnet shore at the garnet trachyte outcrop
and skirted the estuary by walking along the tops of the shoreline banks.
If Brown and Humphrey reached the Huon River by crossing the summit of Mt
Wellington and following the Mountain River as indicated by Vallance (1981: 115),
the only rock type they could have seen apart from sediments and dolerite would have
been basalt. On this route there are two small outcrops of basalt — one being at the
Mountain River bridge on the Ranelagh to Hobart road, the other near the summit of
Mt Wellington close to the Australian Broadcasting Commission TV transmission
tower. If a separate source for pitchstone is required these may qualify although
Humphrey’s letter states that the ‘garnet’ has an intimate association with its
pitchstone matrix. The Mountain River locality does not contain olivine nodules but
these do occur in some abundance at the Mt Wellington locality. If Humphrey had
collected from here because of its particular location, I am sure that he would have
made special note of it. These latter points have been added for sake of completeness
as the author believes the evidence supports the Port Cygnet locality as the source of A.
W.H. Humphrey’s ‘green garnets’.
ACKNOWLEDGEMENTS
Following earlier correspondence with Professor Vallance I would express my
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
125
126 NOTES AND DISCUSSION
appreciation to the Society for inviting me to present these comments in the
Proceedings and to Professor T. Vallance and Dr M. R. Banks for suggesting
improvements to the manuscript.
References
Forp, R. J., 1967. — A re-appraisal of johnstonotite. Pap. Proc. Roy Soc. Tasm. 101: 11-12.
Mac eop, W. A., and WuitE, O. E., 1900. — On the occurrence of a new species of garnet at Port Cygnet.
Pap. Proc. Roy. Soc. Tasm. for the years 1898-1899: 74-76.
VALLANCE, T. G., 1981. — The start of government science in Australia: A. W. H. Humphrey, His
Majesty’s Mineralogist in New South Wales, 1803-12. Proc. Linn. Soc. N.S.W. 105 (2): 107-146.
Communicated by T. G. Vallance ; accepted for publication 16 December 1981.
Proc. Linn. Soc. N.S.W., 106 (1), (1981) 1982
PROCEEDINGS
of the
LINNEAN
SOCIETY
NEW SOUTH WALES
VOLUME 106
_ NUMBER 2
Significance of Late
Cambrian (Idamean) Fossils
in the Cupala Creek
Formation, northwestern
New South Wales
€. MCA. POWELL, G. NEEF, D. CRANE, P. A. JELL,
and I. G. PERCIVAL
PowELL, C. McA., Neer, G., CRANE, D., JELL, P. A., & PeRcivat, I. G. Significance
of Late Cambrian (Idamean) fossils in the Cupala Creek Formation,
northwestern New South Wales. Proc. Linn. Soc. N.S.W. 106 (2), (1981) 1982:
127-150.
Late Cambrian (Idamean) fossils occur about 22 km southeast of Nuntherungie
station, northwestern New South Wales, in the upper part of a terrigenous,
transgressive succession (Cupala Creek Formation — new name) lacking carbonates.
The succession, about 1 km thick, consists of a basal conglomeratic unit overlain by
red and grey sandstone, the latter containing brachiopods, which in turn is overlain by
olive-grey siltstone containing trilobites and brachiopods. The basal part of the
formation represents one of the first pulses of clastics derived from the rising
Delamerian mountains to the south. Sedimentation was probably rapid from streams
intermediate between braided and meandering sinuosity. The middle and upper parts
of the formation are paralic and neritic, respectively.
The Cupala Creek Formation unconformably overlies the ?Vendian or Early
Cambrian Copper Mine Range Beds, and is unconformably overlain by the ?Late
Devonian Mulga Downs Group. The nearby Kandie Tank Limestone has the same
stratigraphic relations as the Cupala Creek Formation, but is probably younger (post-
Idamean — pre-Payntonian).
Six trilobite species, including one new genus, Notoaphelaspis Jell, gen. nov.,
and one new species, N. orthocephalis Jell, sp. nov., two brachiopods and a mollusc
are described. The discovery of fossils in the Cupala Creek Formation demonstrates
that the Copper Mine Range Beds were folded before the Late Cambrian. The Cupala
Creek Formation was subsequently folded about southeasterly-trending axes between
Late Cambrian and Late Devonian, and the whole succession was again folded,
probably in the Carboniferous.
C. McA. Powell and D. Crane, School of Earth Sciences, Macquarie University, North
Ryde, Australia 2113; G. Neef, W. S. and L. B. Robinson University College,
University of New South Wales, P. O. Box 334, Broken Hill, Australia 2880; P. A.
Jell, National Museum of Victoria, 285-321 Russell Street, Melbourne, Australia
3000, and I. G. Percival, Department of Geology and Geophysics, University of
Sydney, Australia 2006; manuscript received 7 July 1981, accepted for publication in
revised form 18 November 1981.
INTRODUCTION
The White Cliffs 1:250,000 geological map (Rose et al., 1964) shows a north-
northwesterly-trending syncline on the northeastern side of, and truncated by, the
more westerly-trending structure of Copper Mine Range. Sediments in the north-
northwesterly trending syncline are marked as ?Proterozoic on the map, but suggested
to be Lower Devonian in the explanatory notes (Rose, 1974) on the basis of cor-
relation with rocks about 20 km south near Mt Daubney and Old Gnalta homestead
where fragmentary plant remains had been found (Brown zn Freeman, 1965; Neef et
al., in prep.). The overlying, more shallowly-dipping and westerly-trending quartzose
sandstone and siltstone in Copper Mine Range were correlated by Rose (1974) with
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
128 SIGNIFICANCE OF LATE CAMBRIAN FOSSILS
the Middle to Upper Devonian Mulga Downs Group further east. The structure thus
indicated is a mid-Devonian unconformity (Webby, 1972; Evans, 1977) , which two of
us (Crane and Powell) set out to investigate as part of a more regional study of mid-
Devonian tectonism.
Upper Silurian
to Lower Devonian
xX eae
Cretaceous to Recent Alluvium -” _[5"
White, mature quartz MULGA DOWNS"
sandstone and siltstone GROUP pe
Angular Unconformity
={ Green/Grey siltstones and
=| fine sandstones CUPALA
Maroon to grey lithic sandstones SR are
Lithic conglomerate
Angular Unconformity
Micaceous siltstones and shale
of flyschoid aspect
Bedding
Bedding, younging determined
—— Anticline.
Syncline (with plunge)
RR Minor folds
— 2_ Mapped
——_ Probable Lees
(with thro}
Inferred indicated
COPPERMINE
RANGE BEDS
--~—~sB Airphoto trend
“Measured section
A~ ©1 Marine fossil locality
== Access track
S| i
A B Section line
Thickness of conglomeratic
unit in metres
Fig. 1. Geological map and cross section of the Cupala Creek Formation.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
400
300
200
100
sea level
C. McA. POWELL, G. NEEF, D. CRANE, P. A. JELL ANDI. G. PERCIVAL 129
Late Cambrian (Idamean) trilobites and brachiopods were found by Crane and
Neef thus ruling out the correlation with the plant-bearing part of the Mt Daubney
Beds (also known as the Cootawundy Beds, Pogson and Scheibner, 1971), which lie
west of the Koonenberry Fault. Previous correlation of the two successions was
tentative, on the basis of photogeological similarity (Rose et al., 1964; Wilson, 1967).
Three of us (Crane, Neef and Powell) mapped the area, examining the stratigraphy,
sedimentology and structure, and searching for fossils. Jell described the trilobites and
mollusc, and Percival the brachiopods. The main purpose of this paper is to document
the occurrence of Idamean fossils and to point out their significance in determining
the timing of regional deformation in northwestern New South Wales.
CUPALA CREEK FORMATION
The strata, herein named Cupala Creek Formation, are well exposed over an area
of 4.5 km? in the dry tributaries at the headwaters of Cupala Creek, and there are also
exposures in the low hills between the tributaries (Fig. 1). The structure (discussed in
detail below) is a faulted syncline that plunges gently south-southeast and is
erosionally truncated by mature quartzarenites of the overlying Mulga Downs Group
(Fig. 2). Approximately 1 km of sediment is present in the southern part of the area,
where the type section was measured (Fig. 3, A-B on Figs 1 and 2). The stratigraphic
succession consists of a basal conglomeratic unit overlain by a sandstone unit in which
a pale-grey sandstone is sandwiched between a lower and upper red sandstone. The
top of the formation is an olive-grey siltstone interbedded with very fine sandstone. In
the northern part of the area, only the conglomeratic and sandstone units are present.
Fig. 2. Oblique aerial photograph, viewing north, shows steeply dipping Cupala Creek Formation overlain
by gently dipping Mulga Downs Group. Type section is A-B, and Cupala Creek is in the centre background.
The most fossiliferous locality containing numerous trilobites (1 in Fig. 1) is at C.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
130 SIGNIFICANCE OF LATE CAMBRIAN FOSSILS
Fossil Localities
MULGA DOWNS 9! and 92
GROUP Metres
silt
very fine
fine
Ly)
Trilobites
9)
Outcrop of Cambrian
© Station Homestead
~——= Major Fault trends
0 10 20 30
———$-
km
Fig. 8. Distribution of Cambrian strata in northwestern New South Wales.
CAMBRIAN DEPOSITS IN NORTHWESTERN NEW SOUTH WALES
Cambrian strata are known from at least seven other areas in northwestern New
South Wales (Rose and Brunker, 1969; Warris, 1967; Wopfner, 1966; and, more
recently, Neef, unpub. data) (Fig. 8). A precise age to zone or stage level is known for
two of these areas, and the age of the others is inferred by lithological correlation. Fig.
9 summarizes the present stage of knowledge.
Cambrian deposits are most extensive near Mt Wright where Early to earliest
Middle Cambrian limestone, shale and volcanics are separated from latest Cambrian
strata by an unconformity (Opik, 1975; Kruse, 1978). Early Ordovician strata of the
Mt Wright district can be traced southward to the Scopes Range (near Bilpa station)
so that the underlying conformable strata are probably Late Cambrian (Rose and
Brunker, 1969). The stratigraphic relationship of Early or Middle Cambrian
Proc. LINN. Soc. N.S.W., 106 (2), (1981) 1982
ORDOVICIAN
CAMBRIAN
LATE
BOOMERANGIAN Leiopyge Loevigato<—I7
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C. McA. POWELL, G. NEEF, D. CRANE, P. A. JELL ANDI. G. PERCIVAL 137
STAGES
COMARTO
MT. WRIGHT
CUPALA CK
KANDIE TANK
KAYRUNNERA
AIRSTRIP
KOONENBERRY
MT. ARROW-
SMITH
PAYNTONIAN
PRE - PAYNTONIAN
POST - IDAMEAN
IDAMEAN s Erixanium sentum
MINDYALLA
UNDILLAN 4
FLORA
TEMPLETONIA
ORDIAN
OLENELLIAN
Stage
?
2
Undesignated :
{ Irvingella tropica
Stigmatoa diloma
Proceratopyge cryptica
Glyptagnostus reticulatus
Glyptagnostus stolidotus
N4 Cylagnostus quasivespa
Erediaspis eretes
‘A
Goniagnostus nathorsti
Ptychagnostus punctuosus
Sf Evagnostus opimus
Ptychagnostus atavus
N Ptychagnostus gibbus
N
k <"\y Zo-nowm wm 4n-—Frvuc/\\>
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Known
Turbidite dips eee precisely Shale +siltstone
E==] Limestone | inferred Sandstone
Volcanics Angular uncon- 3] Conglomerate
formity with
Metamorphic maximum Breccia
complex time range
Fig. 9. Cambrian biostratigraphy of northwestern New South Wales, from Henderson (1976), Shergold
(1971), and Opik (1968). Age of Cambrian strata from Opik (1968), Shergold (1971), Warris (1967),
Wopfner (1966) and Jell (unpub. data). Circled numbers refer to the following formations. 1, Mount
Wright Volcanics; 2, Cymbric Vale Formation; 3, Coonigan Formation; 4, Nootumbulla Sandstone; 5,
Wonaminta Complex; 6, Copper Mine Range Beds; 7, Cupala Creek Formation; 8, Kandie Tank
Limestone. Middle Cambrian is used in the sense of Opik (1968) , and Late Cambrian in the sense of Daily
and Jago (1975).
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
138 SIGNIFICANCE OF LATE CAMBRIAN FOSSILS
fossiliferous sediments near Mt Arrowsmith to the underlying ?Early Cambrian
volcanics is unknown (Wopfner, 1966; Warris, 1967). The Yandaminta Quartzite
(Warris, 1967), which underlies the fossiliferous Early Ordovician Tabita Formation
(Shergold, 1971), is separated from the Early or Middle Cambrian strata by an
unconformity at Mt Arrowsmith.
Between Mt Wright and Mt Arrowsmith, in the Nuntherungie-Kayrunnera area,
there are three isolated areas of Cambrian strata (including the Cupala Creek
Formation described herein). About 13 km southeast of Nuntherungie, the Kandie
Tank Limestone (Pogson and Scheibner, 1971) contains the latest Cambrian or Early
Ordovician brachiopod, Lznnarssonella sp., and distacodont conodonts (Warris,
1967). The Cupala Creek Formation lies about 9 km southeast of the exposed Kandie
Tank Limestone. Trilobites collected by Neef from near Kayrunnera airstrip are of
Mindyallan age (Jell, unpub. work) and are probably from the same outcrops from
which Opik (1966) reported Mindyallan trilobites.
Rose (1974) reported Lzngula sp. in the lower beds of a small outcrop of possible
Cambrian rock at “The Bunkers’ (GR557194, 1:250,000 White Cliffs geological
map); however, these lower beds are apparently continuous and conformable with
quartzose sandstone and siltstone of the Mulga Downs Group, so that their age
remains uncertain. Cambrian strata are present in a narrow graben formed by the
bifurcation of the Koonenberry Fault north of Wonaminta station at Koonenberry
Gap. Here a ?Late Cambrian conglomerate and a ?Late Cambrian hyolithid-bearing
limestone are known (Warris, 1967).
The flyschoid Copper Mine Range Beds, which unconformably underlie Cupaia
Creek Formation and Kandie Tank Limestone, are considered by Pogson and
Scheibner (1971) to be either Adelaidean or Early Cambrian. Edwards (1978)
suggested that the mafic rocks of the Wonaminta Block are Early Cambrian, and
reasoned that the Copper Mine Range Beds are also Early Cambrian in age — i.e.
deeper-water equivalent of Early Cambrian shelf deposits at Mt Wright — an
interpretation consistent with that of Scheibner (1973, fig. 7).
EARLY PALAEOZOIC HISTORY OF NORTHWESTERN NEW SOUTH WALES
The Wonaminta Complex (Wonaminta Block, 1971 Tectonic Map of Australia)
is a regionally metamorphosed succession (up to biotite grade) of sediments and some
volcanics considered to be Precambrian (Rose and Brunker, 1969). The mafic rocks
intruding the Wonaminta Complex have been shown on the tectonic map of New
South Wales as Early Palaeozoic island-arc intrusives, and a similar interpretation was
suggested by Edwards (1978) as one of the possibilities. In this interpretation
(Scheibner, 1973) a volcanic arc in the west (Mt Wright volcanic arc) is fringed with
shelf sediments (Gnalta Shelf) and a fore-arc terrace (White Cliffs Deeper Terrace)
to the east, and here the Copper Mine Range Beds would have accumulated.
An alternative hypothesis is that the mafic igneous rocks represent basaltic
outpourings associated with the Vendian or earliest Cambrian rifting of the Palaeo-
Pacific margin (Veevers, 1976). The alkaline nature of the basalts on the western
margin of the Wonaminta Block (Edwards, 1978) is consistent with this hypothesis, in
which case the Copper Mine Range Beds would be turbidites on a passive, divergent
continental margin.
No matter which model of the Early Cambrian palaeogeography is correct, the
Copper Mine Range Beds (clasts of which occur in the basal Cupala Creek
Formation) were highly folded and eroded before the Idamean. The age of this
folding cannot be constrained more tightly than latest Precambrian to earliest Late
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
C. McA. POWELL, G. NEEF, D. CRANE, P. A. JELL ANDI. G. PERCIVAL 139
Cambrian from the Cupala Creek area, because of the uncertain age of the Copper
Mine Range Beds. Other stratigraphic successions in northwestern New South Wales
(Fig. 9) show a paucity of Mindyallan and Idamean deposits. Admittedly, the lacunae
in each of the stratigraphic successions are of differing magnitudes, but if, for
argument’s sake, we assume that the uplift associated was broadly synchronous for this
region (50 km X 200 km), the uplift occurred either in the Undillan to Mindyallan
stages (the Mootwingee Movement of Webby, 1978) or in the latest Precambrian. The
importance of the Cupala Creek Formation in this interpretation is that it marks the
upper limit of the age of the uplift.
Scheibner (1976, p.141; 1978) concluded that deformation in northwestern New
South Wales (Delamerian Orogeny) started in late Early Cambrian time. Further
afield, in southeastern South Australia, deformation and uplift began in the Late
Cambrian and continued with increasing intensity into the Early Ordovician
(Thomson, 1969). In Tasmania, uplift with deformation (Tyennan Orogeny) began
in the Middle Cambrian and continued into the Early Ordovician (Campana zn
Banks, 1962; Williams, 1978). The possible late Middle to earliest Late Cambrian
deformation in northwestern New South Wales would correlate with the early
movements in the Delainerian Fold Belt of Rutland (1976).
In the Mt Wright area, the contact between the latest Early to earliest Middle
Cambrian Coonigan Formation and the conglomeratic Late Cambrian Nootumbulla
Sandstone, is paraconformable north of the Lawrence Fault (Fig. 8). South of the
fault, however, the conglomeratic base of the Nootumbulla Sandstone rests
disconformably on the Early Cambrian Cymbric Vale Formation, the Coonigan
Formation having been removed by mid-Cambrian erosion (Leu, 1980). Abundant
grey-green sandstone clasts (possibly derived from the Early Cambrian Cymbric Vale
Formation of the Mt Wright area) in the, basal conglomeratic unit of Cupala Creek
Formation, suggest that the Cymbric Vale Formation may have been deeply eroded in
the Idamean. Clasts of Cymbric Vale-type sediment, and limestone clasts bearing
brachiopods identical to those of the overlying Coonigan Formation, also occur at
Koonenberry Gap (Warris, 1967).
The Kandie Tank Limestone appears to be Early Ordovician because it contains
distacodont conodonts. However, the fossil evidence is not strong, and in view of the
terrigenous nature of Early Ordovician strata at Mt Arrowsmith and Mt Wright in
contrast with the limestone facies inferred for the Late Cambrian in the Comarto Hills
and Koonenberry areas, Kandie Tank Limestone may be Late Cambrian.
The timing of the southeast-trending folding of the Cupala Creek Formation
cannot be constrained from the Copper Mine Range area any more accurately than
post-Idamean and pre-Mulga Downs Group (assumed to be Middle or Late
Devonian). Webby (1978) suggested that, during the latest Cambrian and Early
Ordovician, clastics were deposited from northward- and eastward-flowing rivers in a
delta complex. The Cupala Creek Formation represents an early pulse of these
clastics, derived from the Delamerian Mountains rising to the south and west.
According to Webby (1978) these clastics were probably deformed in the Middle
Ordovician Dulingari Movement.
The last folding occurred after the Mulga Downs Group was deposited. These
open folds, commonly with nearly vertical to slightly overturned dips on the western
limb of the syncline adjacent to the Koonenberry Fault, and gentle (15° to 50°) dips
on the eastern limb, probably occurred during the Carboniferous when the rest of the
Lachlan Fold Belt was similarly folded (Powell et al., 1980). Tilting, rotation and
minor folding associated with the steep faults trending 100° occurred in the Cupala
Creek Formation at this time.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
140 SIGNIFICANCE OF LATE CAMBRIAN FOSSILS
SYSTEMATIC PALAEONTOLOGY
The trilobites and mollusc were collected from locality 1, and brachiopods from
localities 2-5 (Fig. 1). Trilobite debris is also present at locality 3. All described
material is housed in the Palaeontological Collection of the National Museum of
Victoria (numbers prefixed P).
ARTHROPODA (P.A.J.)
Class TRILOBITA Walsh, 1771
Order MIOMERA Jaekel, 1909
Superfamily AGNOSTOIDEA McCoy, 1849
Family DIPLAGNOSTIDAE Whitehouse, 1936
Subfamily PSEUDAGNOSTINAE Whitehouse, 1936
Genus PSEUDAGNOSTUS Jaekel, 1909
Pseudagnostus sp. aff. cdalis Opik, 1967
Fig. 10, 1,2
Material: One cranidium P61493 and one pygidium P61494.
Discussion: This species is related to P. zdalzs in the squarish glabellar anterior and
weak median preglabellar furrow, in the shape of the basal glabellar lobes and in the
wide border furrow. In the pygidium the shape of the posterior of the deuterolobe is
particularly reminiscent of P. zdalzs and the extremely wide border furrow and narrow
pleural area are also similarities.
The preservation of this material is too inferior for positive specific identification
but enough morphology is revealed to indicate a resemblance to P. zdalzs.
Order POLYMERA Jaekel, 1909
Superfamily PTYCHOPARIOIDEA Matthew, 1887
Family EULOMATIDAE Kobayashi, 1955
Genus STIGMATOA Opik, 1963
Stigmatoa tyson? Opik, 1963
Fig. 10, 3-6
1963 St?gmatoa tysoni Opik, p.92, pl. 4, fig. 3
1976 Stigmatoa tysoni Henderson, p.354, pl. 51, figs 8, 9
Materzal: One well-preserved (P61496) and three damaged (P61495, 61497 and
61498) cranidia.
Dimensions: Cranidial lengths range from 4 to 7 mm.
Diagnosis: As given by Henderson (1976) and Opik (1963).
Discussion: There can be no doubt that this material belongs to S. tysonz; glabellar
furrows, occipital ring and palpebral areas are all distinctive.
Fig. 10. 1, 2, Pseudagnostus sp. aff. cdalis Opik, 1967. 1, internal mould of cephalon, P61493, x7.2; 2,
damaged internal mould of pygidium, P61494, x7.2.
3-6. Stigmatoa tysoni Opik, 1963. 3, badly damaged and deformed cranidium P64195, x7.2; 4a, b, dorsal
and right anterolateral oblique views of internal cranidial mould, P61496, x5.4.; 5, internal mould of
damaged cranidium, P61497, x3.6; 6, internal mould of damaged cranidium, P61498, x5.4
7, 8. ? Aphelaspis sp. aff. A. australis Henderson, 1976. Internal moulds of damaged cranidia, P61499 and
61500, x6.3 and x7.2 respectively. :
9-12. Prismenaspis? sp. nov. 9, latex cast of damaged and tectonically shortened cranidium P61501, x2.7;
10-12, internal moulds of damaged and distorted cranidia, P61502, 61503 and 61504, x2.7.
13, 14. Genus and species indeterminate. Internal moulds of damaged distorted cranidia P61505 and 61506,
x3.6.
15-17. Proplina sp. Right and left anterolateral and dorsal views of internal mould, P61507, x1.8.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
14]
C. McA. POWELL, G. NEEF, D. CRANE, P. A. JELL ANDI. G. PERCIVAL
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
142 SIGNIFICANCE OF LATE CAMBRIAN FOSSILS
Family PTEROCEPHALIIDAE Kobayashi, 1935
Subfamily APHELASPIDINAE Palmer, 1960
Genus 4 PHELASPIS Resser, 1935
Type species: A. walcotti Resser 1938, p.59, pl. 13, fig. 14.
Diagnosis: As given by Palmer (1965, p.58) with the addition made by Henderson
(1976, p.345 in remarks on Eugonocare) that the interocular width approximately
equals cranidial length.
Discusston: In Australia this genus is represented by A. australis Henderson (1976,
p.342, pl. 49, figs. 5-7) and by a species from limestone in eastern Victoria (Thomas
and Singleton, 1956). A case could be made to exclude australzs from A phelaspis on
the basis of its very much shorter frontal area, its well impressed glabellar furrows, its
longer palpebral lobes, and its more rounded glabellar anterior. I consider that it
should remain in A phelaspis but that it is a distant relative of the North American
species described to date including A. brachyphasts Palmer, 1962 which Henderson
considered the ‘nearest match’. Future detailed work on Australian faunas of this age
may well show that australis belongs to a lineage quite separate from the North
American Aphelaspis. For the present it is placed in Aphelaspis as is the material
compared with it herein.
2A phelaspis sp. aff. A. australis Henderson, 1976
Fig. 10, 7,8
Material: Two poorly preserved, damaged internal moulds of cranidia, P61499 and
61500.
Description: Cranidium approximately as long as interocular width which is slightly
compressed. Glabella slightly tapering, anteriorly truncated, with almost straight
sides. Axial furrow well impressed but slightly shallower anteriorly. Frontal area short
(approximately 40% of glabellar length). Border furrow shallow. Border short and
convex, of uniform length except where the facial sutures cut across it. Interocular
cheeks narrow. Palpebral lobes elongate, slightly raised and opposite middle of
glabella.
Discusston: These specimens resemble A phelaspis australis in their short frontal area
measuring 40% of glabellar length and in their elongate palpebral lobes. They also
resemble 24 phelaspis sp. B. of Opik, 1963 in the type of border, glabellar shape, short
frontal area and narrow interocular cheeks. However, their poor preservation and
ignorance of the rest of the exoskeleton make closer identification impossible.
Family PTEROCEPHALIIDAE Kobayashi, 1935
Genus NOTOAPHELASPIS gen. nov.
Etymology: Notios Gr. southern, with North American genus name A phelaspis (from
apheles Gr. smooth).
Fig. 11. 1-10. Notoaphelaspis orthocephalis gen. et. sp. nov. la, latex of cranidium, Ib internal mould of
same cranidium P61508, x3.2 (approx.); 2a, right anterolateral oblique view and, 2b, dorsal view of
internal mould of holotype cranidium P61509, x2.2; 3, internal mould of cranidium with fixed cheeks only
slightly displaced and three thoracic segments P61510, x1.4; 4, internal moulds of damaged thorax and
pygidium, P61511, x1.8; 5, 6, internal moulds of damaged cranidia P61512 and 61513, x1.8 and x1.4
respectively; 7a internal mould and 7b, latex cast from external mould of damaged cranidium and partial
thorax P61514, x1.8; 8, 9, internal moulds of damaged cranidia P61515, 61516, x2.7; 10, internal mould
of free cheek broken down genal spine to reveal the external mould of the ventral surface P61517, x3.2.
11. Notoaphelaspis sp. cf. N. orthocephalis gen. et sp. nov. internal mould of incomplete cranidium
P61518, x2.7.
Proc. LINN. Soc. N.S.W.,106 (2), (1981) 1982
C. McA. POWELL, G. NEEF, D. CRANE, P. A. JELL ANDI. G. PERCIVAL 143
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Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
144 SIGNIFICANCE OF LATE CAMBRIAN FOSSILS
Type species: N. orthocephalis gen. et sp. nov.
Diagnosis: Aphelaspid with rectangular glabella, wide interocular cheeks, transverse
eye ridges, anteriorly placed palpebral lobes, long downsloping preglabellar field, well
impressed anterior border furrow, medially elongate anterior border, non-spinose
pleural tips, and transverse, non-spinose pygidium.
Discussion: Although this species could, with some difficulty, be placed in A uae
the combination outlined above is quite unlike any known species of that genus and
the eye lines, glabellar shape and deep border furrow are particularly distinctive. It
has no close relatives in North America. Eugonocare Whitehouse, 1939 is
distinguished by its broader glabella with sloping lateral furrows, anterior border,
more sloping eye ridges and longer pygidium. Its glabella and the furrows thereon are
extremely similar to those of Erzxanium which are themselves very distinctive among
trilobites. The major differences between N. orthocephalis and E. sentum are the
glabella shape, cephalic border, pleural tips and the pygidium. In view of the marked
changes in pleural tip and pygidial morphology along an aphelaspid lineage (Palmer,
1965) the possibility that N. orthocephalis is an earlier member of the Erixanzum
lineage (and therefore a member of the Erixaniidae) should not be overlooked. Such
placement would require considerable expansion of the Family concept given by Opik
(1963).
Notoaphelaspis orthocephalis sp. nov.
Fig. 11, 1-10
Etymology: Orthos Gr. right (angled) ; kephale Gr. head; for the angular glabella.
Holotype: P61509.
Other material: Several cranidia, two thoraxes, one free cheek and two pygidia
including the paratypes P61508 and P61510-61517.
Diagnosis: Up to 8 cm in cephalic length. Cranidial length and interocular width
approximately equal. Glabella rectangular, two-thirds cephalic length, with angular
anterolateral corners and truncated anterior, with occipital furrow poorly impressed
axially but not reaching axial furrow, with three pairs of very weakly impressed
glabellar furrows as circular depressions on either side of axis, well separated from
axial furrow. Preglabellar field relatively long, down sloping and slightly swollen in
front of glabella. Anterior border furrow well impressed. Anterior border convex and
expanded medially. Each interocular cheek as wide as glabella at midlength of
relatively long arcuate slightly upturned palpebral lobes. Eye ridges transverse; very
slightly posteriorly directed laterally in a few specimens. Facial sutures diverging
anteriorly from palpebral lobe at about 45° posteriorly running laterally at a low
angle to the transverse line so that the posterior part of the fixed cheek is short and
wide. Posterior border furrow distinct, of almost constant length throughout, defining
a short convex border also of constant length. Free cheek with convex genal field, well
impressed border furrow, and long slender slightly incurved genal spine that continues
the curve of the lateral margin. Thorax of at least 9 segments (no complete thorax
available). Axis highly arched and relatively narrow. Axial furrow simply a change of
slope. Articulating line about mid-width of each pleural area. Articulating facet very
wide, steep and flat. Pleural furrows prominent occupying most of the pleural area.
Anterior and posterior pleural margins parallel. Pygidium transverse, twice as wide as
long, moderately convex with axis standing above pleural areas. Axis of three rings
and a posteriorly rounded terminus that interrupts the border furrow. Pleural areas
convex, crossed by three pleural and one interpleural (including anterior border
furrow) furrows. Border furrow weakly impressed but not impressed at all behind the
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
C. McA. POWELL, G. NEEF, D. CRANE, P. A. JELL ANDI. G. PERCIVAL 145
axis. Border widening anteriorly, not present behind the axis. The margin forms a
smooth arc, except behind the axis where it is excavated to the extent that it comes
very Close to the axis.
Discussion: The material figured here is in a coarse matrix, is tectonically distorted to
some degree in each specimen and is mainly of damaged internal moulds but the
morphology available is so distinctive as outlined under discussion of the genus as to
demand a separate taxonomic name. Comparison with any existing species is
superfluous. The sample is too small to determine any intraspecific variation and no
morphogenetic variation is visible over the size range available. As discussed below the
small cranidium (Fig. 11, 7) could possibly be a juvenile of this species although this is
a remote chance.
Notoaphelaspis sp. cf. N. orthocephalis sp. nov.
Higa tet
Material: One fragmentary cephalon P61518.
Discusston: This specimen differs from N. orthocephalis in that its anterior border
does not taper laterally, its preglabellar field is shorter and its interocular cheeks are
relatively narrower.
Family UNCERTAIN
Genus PRISMENASPIS Henderson, 1976
Prismenaspis? sp. nov.
Fig. 10, 9-12
Materval: Six cranidia in various degrees.of distortion due to deformation, including
P61501-61504.
Description: Moderately large cranidium (up to 1.5 cm long) approximately as wide
as long with broad squat anteriorly tapering glabella just over half cranidial length.
Glabellar furrows not well impressed but 1P connected adaxially to 2P in at least one
specimen (Fig. 10, 11). Occipital ring tapering laterally and with an almost
imperceptible median node. Axial furrow well impressed with a marked pair of
fossulae in front of the glabella adaxial to the rounded anterolateral corners.
Preglabellar field slightly inflated medially, short and not distinguished from an
almost indiscernible, long, shallow, border furrow which in turn is not separated from
a long, flat, medially inflated border that tapers laterally and bears a low but distinct
transverse ridge that may or may not correspond to the edge of the doublure. Eye
ridges well defined and running posterolaterally to short wide, kidney-shaped
palpebral lobes. Facial sutures diverging slightly forward and sloping posteriorly
behind the palpebral lobe. Posterior portion of fixed cheek triangular, crossed by long
border furrow that runs forward laterally to leave a lengthening and flattening
posterior border.
Discussion: The state of preservation and small number of specimens make
identification of this with any existing species impossible but it does not appear to be
similar to any existing taxon. Even at the generic level the assignment to Przsmenaspis
is very tentative and requires expansion of that genus to include apparently
unornamented species. This is in conflict with Henderson’s concept of the genus as he
stressed the pustulose ornament on both internal moulds and external surfaces.
However, related genera such as Dunderburgza encompass smooth and ornamented
species. Moreover, Henderson described four pairs of lateral glabellar furrows in
Prismenaspts but material of the New South Wales species is too poorly preserved to
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
146 SIGNIFICANCE OF LATE CAMBRIAN FOSSILS
see any but the 1P glabellar furrows. Glabellar shape, well-impressed axial furrow
with prominent fossulae, narrow interocular cheeks, wide posterior portion of the
fixed cheeks, position and length of palpebral lobes, and most importantly, the flat
frontal area relate this species to Prismenaspis. Prismenaspis propinquum is the most
closely related species but differs in ornament, glabellar shape and lack of transverse
ridge on anterior border. As with Prismenaspis there are several younger Siberian
genera described by Rozova (1968, 1977) that appear similar, including Ketyna,
Monosulcatina, Mansiella, and Maduzya but the comparisons are obscured by the
states of preservation and incomplete information.
Genus and species indeterminate
Fig. 10, 13, 14
Materzal: Two damaged cranidia P61505 and 61506.
Description: Small cranidium longer than wide with subrectangular glabella having
only very slightly tapering sides, angular anterolateral corners, and no furrows
impressed. Axial furrow well impressed, without fossulae. Preglabellar furrow
transverse. Preglabellar field short, shorter than border. Border furrow shallow
throughout and indistinct medially where the border is expanded into a poorly-
defined plectrum. Border relatively long, tapering gradually laterally. Interocular
cheeks narrow. Palpebral lobes elongate, weakly curved and defined by shallow
palpebral furrows. Posterior part of fixed cheek rather long and narrow with a
prominent laterally lengthening border furrow running across it.
Discussion: These two specimens are distinctive in their clearly truncated glabellar
anterior, and broadly convex anterior border. They may belong to a genus that has
not yet been named but a new name has not been assigned because of the small
number of poorly-preserved internal moulds available and because of the lack of
knowledge on the remainder of the exoskeleton. Perhaps the most distinctive aspect of
this material is the long, convex anterior border, that is unique among this type of
trilobite of this age. As with Prismenaspis the closest morphology is to be found among
younger trilobites from the northwestern Siberian Platform. Monosulcatina laeve
Rozova, 1963, particularly the cranidium, figured by Rozova in 1968 (pl. 9, figs 11,
12) is very close in morphology to the N.S.W. specimen.
MOLLUSCA (PeASTs)
Class MONOPLACOPHORA Wenz in Knight, 1952
Order TRYBLIDIIDA Lemche, 1957
Family TRYBLIDIIDAE Pilsbry in Zittel-Eastman, 1899
Genus PROPLINA Kobayashi, 1933
Proplina sp.
Fig. 10, 15-17
Materzal: One shell P61507.
Description: A large shell 14 mm high, 21 mm wide and 30 mm long with the shape of
an inverted asymmetrical cone. Apex overhangs the margin of the oval aperture.
Internal and external surfaces of shell exhibit fine closely-spaced growth lines parallel
to the apertural margin. Dorsal surface of internal mould with a variety of curving
impressions but none of these are symmetrical nor could they be interpreted as muscle
scars.
Discussion: Ignorance of the muscle pattern and of the apex prevent specific
assignment but the exterior surface, position of the apex and apertural shape make
generic placement clear. The apex is presumably pointed but the infilling has broken
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
C. McA. POWELL, G. NEEF, D. CRANE, P. A. JELL ANDI. G. PERCIVAL 147
on the surface shown in Fig. 10, 16. As there is only one specimen available and the
infilling does not easily separate from the external mould it has not been fully
prepared out.
BRACHIOPODA (iGaPs)
Class ARTICULATA Huxley, 1869
Order ORTHIDA Schuchert & Cooper, 1932
Suborder CLITAMBONITIDINA Opik, 1934
Superfamily BILLINGSELLACEA Schuchert, 1893
Family BILLINGSELLIDAE Schuchert, 1893
Genus BILLINGSELLA Hall & Clarke, 1892
Billingsella sp.
Fig. 12, 1-5
Material: Eleven specimens (P61939-P61949) from locality 2; five specimens
(P61950-P61954) from locality 4; three specimens (P61955-P61958) from locality
5; two specimens (P61959 and P61960) from locality 3.
Description: Exterior of valves. Outline subquadrate; maximum width at about
midlength of pedicle valve, and occurring in posterior half of brachial valve; hingeline
equal to, or slightly less than, maximum width; cardinal extremities obtuse to right-
angled. Profile biconvex, with deeper brachial valve (depth about one-third valve
length) bearing shallow posteromedian sulcus. Ornament poorly preserved on
available specimens, but apparently finely multicostellate with prominent widely-
spaced concentric growth lamellae. Ventral interarea long, equal to between 0.2 and
Fig. 12. 1-5. Billingsella sp. All x2.9. 1-2. Pedicle valve internal mould and corresponding latex cast,
P61939. 3. Pedicle valve internal mould, P61940. 4. Brachial valve internal mould, P61941. 5. Brachial
valve external (latex cast), P61942. 6. obolid indet., P61938, x3.5.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
148 SIGNIFICANCE OF LATE CAMBRIAN FOSSILS
0.3 valve length, flat to weakly curved, orthocline to apsacline; delthyrium covered for
about one-third its length by strongly arched pseudodeltidium. Dorsal interarea much
lower, less than one-third length of ventral interarea; planar, anacline.
Pedicle valve intertor. Teeth stout, extending from hingeline to floor of valve; dental
plates lacking. Muscle field large, extending to between 0.4 and 0.5 valve length, and
occupying approximately one-quarter valve width; elliptical diductors divided
posteriorly by short, thin median ridge extending forward from apical callus; central
adductor impression expands rapidly in front of median ridge. Mantle canal
impressions prominent, of saccate pattern with thick vascula media weakly divergent
medially, strongly arcuate anterolaterally. Broad shallow groove-like depression
extends from hingeline adjacent to valve margin.
Brachial valve interior. Cardinalia simple, comprising fine ridge-like cardinal
processes, and short widely divergent rod-like brachiophores, flanking cup-shaped
sockets. Low, broad, median ridge extends from notothyrial platform to about one-
third valve length, bisecting faintly quadripartite muscle field. Mantle canal
impressions not discernible.
Measurements (in mm) Figured specemens. P61939 (pedicle valve): length 13.1,
width 12.4, length of muscle field 6.5; P61940 (pedicle valve): length 9.5, width
10.8, length of muscle field 4.1; P61941 (brachial valve): length 8.8, width 11.7;
P61942 (brachial valve) : length 7.2, width 9.2.
Pedicle valves, 8 specomens. Range of lengths: 9.5-13.1, range of widths: 10.8-14.0,
range of length/width ratios: 0.88-1.08, average length/width ratio: 0.99.
Brachial valves, 9 specimens. Range of lengths: 6.3-10.4, range of widths: 7.7-14.0,
range of length/width ratios: 0.73-0.82, average length/width ratio: 0.78.
Discussion: Billingsella is restricted to strata of Middle Cambrian to Early Ordovician
age, being particularly abundant in the Late Cambrian. The species found in the
Cupala Creek Formation is most similar to (and may prove to be conspecific with)
Bullingsella sp. from the Middle to Late Cambrian Mariner Formation (Cooper et al.,
1976) of Northern Victoria Land, Antarctica, described and illustrated by
MacKinnon (zm Shergold et al., 1976). Relatively minor differences between these
forms include a proportionately shorter pseudodeltidium and slightly less pronounced
dorsal median sulcus on New South Wales specimens. In other details such as outline,
profile, dimensions and internal structures, species from these two areas are virtually
indistinguishable. In common with the Antarctic species, the New South Wales form
resembles several North American species, such as B. perfecta Ulrich and Cooper and
B. texana Bell, typical of the Late Cambrian. Trilobites associated with the Antarctic
Billingsella indicate its age as late Idamean, Erzxaniwm sentum Zone (Shergold et al.,
WOTOE
Class INARTICULATA Huxley, 1869
Order LINGULIDA Waagen, 1885
Superfamily LINGULACEA Menke, 1828
Family OBOLIDAE King, 1846
Obolid indet.
Fig. 12,6
Discussion: A single poorly-preserved valve (P61938) was found at locality 3. The
available material does not permit precise identification or age determination.
Measurements: Length 9.2 mm, width 7.0mm.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
C. McA. POWELL, G. NEEF, D. CRANE, P. A. JELL ANDI. G. PERCIVAL 149
ACKNOWLEDGEMENTS
Powell and Crane were supported by the Australian Research Grants Com-
mittee. David Larsen and Jeff Vaughan (W. S. & L. B. Robinson University College)
helped in the field and Elaine Prendergast (Macquarie University) point-counted the
sandstone samples. Mr Bill Gall (Nuntherungie station) is thanked for hospitality and
he kindly piloted the light aircraft from which Fig. 2 was taken. Ian Percival
acknowledges financial support from a Rotary Foundation Graduate Fellowship, and
use of facilities at the Museum of Invertebrate Paleontology, Department of Geology,
University of Kansas. Erwin Scheibner, John Shergold, John Talent and John Veevers.
offered helpful comments on the manuscript.
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Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
Origins and Relationships
among the Animal Phyla
D. T. ANDERSON
ANDERSON, D. T. Origins and relationships among the animal phyla. Proc. Linn.
Soc. N.S.W. 106 (2), (1981) 1982: 151-166.
Conceptual and factual advances in the study of animal phylogeny have
emerged from recent work in palaeontology, functional morphology and comparative
embryology. Many earlier proposals on the origin and relationships of the animal
phyla can now be seen to be erroneous.
The fossil record of the Precambrian and early Cambrian has not revealed links
between phyla, but it places the time of origin and radiation of the metazoan phyla at
earlier than 700 million years ago. Functional morphological studies have raised many
possibilities of the convergent or parallel evolution of phyla, especially among those of
simpler body construction. In particular, these studies have called into question the
validity of hypothetical ancestors and of a monophyletic origin of the Metazoa.
Comparative embryology remains a major source of positive information about
the relationships between the animal phyla. Two major assemblages, the spiral
cleavage assemblage and the deuterostome assemblage, can be identified by this
approach. More information is needed on a number of phyla.
A synthesis of all three lines of investigation emphasizes the likelihood that
metazoans arose from protozoans on several occasions.
D. T. Anderson, School of Biological Sciences, The University of Sydney, Australia,
2006; manuscript recetved 15 October 1981, accepted for publication 18 November
1981.
INTRODUCTION
A student once wrote in an essay for me, ‘Phylogenetic relationships is a very
doubtful area of zoology’. This a sentence of delightful ambiguity. I think all zoologists
would agree with it, but for different reasons. To some, the word doubtful would
mean dubious, questionable, verging on the disreputable. To others, quite a strong
group, the doubt would be as to whether phylogenetic relations is a part of scientific
zoology at all. A third group, however, would take it that there are still matters of
doubt and unresolved questions concerning animal phylogeny, and would regard this
as an important challenge for zoologists to struggle with. I subscribe to the last of these
views, but I am well aware that there are plenty of dubious phylogenies as well as some
spectacularly unscientific ones in the recent literature. I shall mention some of these as
we proceed. I am also aware that a lack of suitable factual knowledge leaves many
questions still unanswered concerning phylogenetic relationships, but I think we have
now reached the stage where some firmly-based statements can be made.
Many attempts have been made over the years to evaluate the origins of the
metazoan phyla and their phylogenetic relationships. Until quite recently there was a
strong tendency among animal phylogeneticists to vie with one another in the erection
of monophyletic trees for the Metazoa, branching in various ways. Figs 1 and 2 show
two examples of this.
Preconception was the basic philosophical stance in these attempts, with the
known facts being used as hooks from which to suspend the imagined tree. This
approach has not lost its vogue. Papers are still being published which contain trees of
this type, replete with nonfunctional hypothetical ancestors. A good example of this,
first published in 1976, is shown in Fig. 3. Such works are of the same conceptual
status as land bridges in biogeography.
In the last twenty years, however, and especially in the last decade, very
substantial conceptual and factual advances have been made by many workers
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
152 ORIGINS AND RELATIONSHIPS AMONG ANIMAL PHYLA
Arthr. s
Annelids
Platyhelminths
Nemerteans
Brachiopods
Se eos Hemichordates
lophophorates
Coelomate Animals
Radiate Animals
Scyphozoans
Hydrozoans Anthozoans
Sporozoans
Protozoans
Sarcodinians
Flagellates
Fig. 1. The phylogenetic tree of Marcus (after Barnes, 1968).
interested in the problem of metazoan phylogenetic relationships. It may seem
perhaps surprising that this problem, which was at the forefront of zoological thinking
in the latter part of the nineteenth century, should receive a revival of interest in the
latter part of the twentieth century; but I think this can be explained by the
importance of phylogeny to zoology as a whole. Zoology, the science of animals, is
couched in terms of generalizations based properly and entirely on an acceptance of
the view that evolution has occurred. The better we understand this evolution in terms
of origins and lineages, the better will be our generalizations about its products.
CONCEPTUAL ADVANCES
What then are the conceptual advances which are assisting in this endeavour?
There are three. The first is a greater emphasis on the notion that facts take
precedence over preconceptions. Paraphrasing two famous pragmatists, Sidnie
Manton and Henry Ford, hypothetical animals are bunk! The hypothetical
urcrustacean shown in Fig. 4 and the hypothetical ancestral mollusc well known
amongst zoologists (Yonge and Thompson, 1976), while of value in comprehending
the phyla to which they are ascribed, have no place in thinking about the relationships
between phyla. Gaps in the evidence must be accepted for what they are and not filled
with inventions to satisfy the urge for a monophyletic oversimplification.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
D. T. ANDERSON 153
Molluscs Arthropods
Chordates
ee ee Hemichordates
lophophorates _—_ Brachiopod
seudocoelomat¢s
Echinoderms
Flatworms
Coelenterates
Acoeloid
Ancestor
Sporozoans Ciliophorans
Sponge s \
Rhizoflagellat
Sarcodinians
Flagellates
Fig. 2. The phylogenetic tree of Hanson (after Barnes, 1968).
Secondly, deriving from advances in many aspects of zoology, it is now realized
that animals exist as functionally integrated complexes of dynamically interacting
components, in which there are intrinsic constraints on change as well as the
possibilities of change over evolutionary time. Putting this more simply, evolution
must be functional and function must be viewed holistically. Any phylogenetic
proposal which ignores the requirements of functional gradualism (Anderson, 1967;
Dullemeijer, 1980) must be wrong. This is not to say that any proposal based on
holistic functional thinking must be right. There are other ways in which error can
potentially creep in. Functional thinking may, for example, be employed spuriously to
give credence to imaginary evolutionary sequences. Gutmann (1981) treats the
deuterostomes in this way, expounding the evolution of all deuterostomes from a
segmented chordate stem, itself evolved by an entirely hypothetical ‘functional’ route.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
154
ORIGINS AND RELATIONSHIPS AMONG ANIMAL PHYLA
Arthropoda
\ Annelida
a Chordata
Articulata
Mollusca :
d es
rer Heihordela — Ae
Pogonophora
Spiralian
Archicoélomate
Cnidaria Ctenophora
Porifera
Blastaea
Fig. 3. The phylogenetic relationships of the Metazoa according to Siewing (1976), redrawn and simplified.
Nowadays, however, most zoologists and many palaeontologists have begun to include
functional thinking in their phylogenetic interpretations (e.g. Bock, 1981; Szalay,
1981), though some palaeontologists still proceed in blissful unawareness of this need.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
D. T. ANDERSON 155
antennule
antenna
mandible
maxillule
maxilla
first trunk limb
Fug. 4. The hypothetical urcrustacean of Hessler and Newman (redrawn after Hessler ang Newman, 1973).
The speculations of Jefferies (Jefferies and Lewis, 1978; Jefferies, 1979, and earlier)
on the Calcichordata as ancestral chordates exemplify this well, as Philip (1979) has
remarked.
Curiously enough, this change of approach has led in turn to a third conceptual
advance with a somewhat opposite effect. This is the realization that for the simpler
kinds of animal organization, with few interacting components, it is very difficult to
exclude the possibility of parallel or convergent evolution. Simple metazoans which
may seem alike are not necessarily related. I shall return to this later.
FACTUAL ADVANCES
We are thinking about animal phylogeny with a new kind of honesty and
understanding as a result of these conceptual changes, but this new approach derives
basically from the availability of new facts about animals and their history. During the
last twenty years, factual knowledge has advanced on three fronts pertinent to the
problem of phylogenetic relationships. The early part of the fossil record of animals in
the later Precambrian and Cambrian has become much better known (e.g. Brasier,
1979). The functional morphology of the modern phyla has become much better
understood; and the embryonic development of many of the modern phyla has been
more deeply analysed in functional terms. The interplay of fact and concept has now
reached a stage at which the evidence of all three of these lines of investigation can be
brought together in a more comprehensive way. I shall briefly review each area and
then attempt to establish a synthesis from them.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
156 ORIGINS AND RELATIONSHIPS AMONG ANIMAL PHYLA
1 Jellyfish 2 Charniodiscus 3 Tribrachidium 4 Dickinsonia 5 Spriggina
Fig. 5. Some representatives of the Ediacara fauna (after Glaessner) .
The fossil evidence
For the metazoan phyla, the fossil record of the later Precambrian is sparse but
revealing (Durham, 1979; Paul, 1980; Runnegar, in press). It is well exemplified by
the famous Ediacara fauna of South Australia and elsewhere, shown in Figure 5
(Glaessner, 1969, 1971, 1972, 1976, 1979, 1980; Glaessner and Wade, 1966; Wade,
1970, 1972; Ford, 1980; Birket-Smith, 1981). Beginning about 680 million years ago,
cnidarians, polychaetes, echiuroids, arthropod-like animals and various other
metazoans were already present. Evolution had thus reached the stage of complex,
soft-bodied coelomate animals by the time metazoan fossils began to appear. This is
further borne out by the appearance of pogonophorans before the onset of the
Cambrian (Brasier, 1979) and a variety of complex skeletonized forms such as
molluscs, echinoderms, brachiopods and trilobites (Figs 6, 7) in the early Cambrian
(Runnegar, 1980a, and in press) . This evidence tells us that the important separations
of the metazoan phyla took place more than 700 million years ago, before any fossil
metazoans were laid down. It also tells us that the fossils that are present all belong to
discrete phyla, some modern, some extinct, and include no animals that can be
interpreted as missing links. No new phyla are directly known to have evolved since the
onset of the Cambrian, although some groups that might be accorded the status of
phyla (Archaeocyatha, Hyolitha, Trilobita and others; see Fig. 25) became extinct in
the Palaeozoic.
Fig. 7. The Cambrian evolution of the trilobites (after Bergstrom, 1979).
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
D. T. ANDERSON 157
Confirmation of these conclusions arises from the recent studies of the Middle
Cambrian Burgess Shale fauna (Conway Morris, 1979a; Whittington, 1979, 1980).
In this work we find extraordinary evidence of the early presence of a variety of soft
bodied animals belonging to modern phyla. The Burgess Shale fauna includes
priapulids, polychaetes, marine uniramians and even chordates (Figs 8-11). The
phylum Crustacea is also represented in the Burgess Shale by Canadaspis, and in the
Upper Cambrian by ostracods (Fig. 16) and cephalocarids (Briggs, 1977; Miller,
OOM LO Sis):
Ji (G Ze
12
Figs 8-12. 8, A polychaete of the middle Cambrian Burgess Shale (after Conway Morris, 1979b). 28 mm. 9,
A priapulid of the middle Cambrian Burgess Shale (after Conway Morris, 1979a). 180 mm. 10, The middle
Cambrian marine uniramian Aysheaza (redrawn from Whittington, 1980). 60 mm. 11, The middle
Cambrian chordate Prkaza (based on Conway Morris and Whittington, 1979). 40 mm. 12, The middle
Cambrian Opabznza, phylum unknown (after Whittington, 1980). 50mm.
In addition, the Burgess Shale fauna contains many representatives of groups of
animals that have not survived and do not belong within the modern phyla. These
include such forms as Opabinia (Fig. 12), Hallucigenia (Fig. 13) and strange
arthropods like Yohora (Fig. 14), Marrella (Fig. 15) and Odorata (Conway Morris,
1979a; Whittington, 1979, 1980, 198la; Briggs, 1981). It seems that there were
several (perhaps many) distinct groups of arthropods in the Cambrian fauna in
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
158 ORIGINS AND RELATIONSHIPS AMONG ANIMAL PHYLA
addition to the Trilobita, Crustacea, Chelicerata and Uniramia (Whittington, 1979,
1981b; Manton and Anderson, 1979). Whether Opabinia, Hallucigenza, the strange
arthropods and the other unique animals of the Burgess Shale should be interpreted
as members of discrete extinct phyla is debatable, but they are certainly indicative of
the existence of Cambrian groups of animals that do not fit the definitions of any of
the modern phyla. Like the Precambrian and early Cambrian metazoans, none of the
Burgess Shale animals is intermediate between or directly ancestral to any of the
recognized phyla. Their structural complexity, elucidated now in impressive detail,
gives further evidence of a long prior history of metazoan evolution.
AS
NS IES
=
(Soaat erie ae
14 UA XQ
Figs 13-16. 13, The middle Cambrian Halluczgenca, a fossil worthy of a Bestiary (after Whittington, 1980).
18mm. 14, The middle Cambrian arthropod Yohoza, phylum unknown (after Whittington, 1980). 23 mm.
15, The middle Cambrian arthropod Marrella, phylum unknown (after Whittington, 1980). 20 mm. 16,
The upper Cambrian ostracod Vestrogothia (after Muller, 1979). 0.5 mm.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
D. T. ANDERSON 159
From the point of view of the origins and relationships of the animal phyla, then,
the fossil evidence provides nothing directly, except a confirmation of the timing of
events. This puts the onus directly on functional morphology and embryology to
provide some answers within this context.
Functional morphology
Curiously enough, what this discipline has done for phylogenetic relationships
among the animal phyla is mostly negative. The recognition that animals are highly
functionally integrated and adapted in relation to habit and habitat has led to the
further realization that they are often more different than they seem. Common
ancestries then become much less plausible. The separation of the arthropods into
three modern phyla and probably several extinct phyla (Anderson, 1973; Manton,
1973, 1977; Schram, 1978; Manton and Anderson, 1979; Whittington, 1979, 1981b)
is perhaps the most spectacular example of how functional morphology has
demolished old beliefs, but there are others. The fact that structure is functionally
related to habit and habitat is well illustrated in a broad sense by the case of the
coelomic worms. Functional studies (Clark, 1979) have demonstrated a clear
relationship between:
1. an unsegmented coelom and burrowing slowly in compact substrata,
2. asegmented coelom and burrowing actively in looser substrata,
3. a trimeric coelom and tubicolous life.
Each of these conditions could have evolved independently more than once and none
needs any other as a functional prerequisite. For example, there is no reason to
suppose on functional morphological ground that the Sipunculida and the Priapulida,
both unsegmented coelomate burrowers, are related to one another. The Phoronida
and the Pogonophora both show a trimeric tubicolous condition, but the
REPART pre annular TRU postannular A
FOREPART TRUNK OPISTHOSOMA
Fig. 17. The adult and larva of a pogonophoran (after Southward, 1980).
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
160 ORIGINS AND RELATIONSHIPS AMONG ANIMAL PHYLA
Pogonophora (Fig. 17) are now thought to be specialized descendants of the annelids
(Southward, 1980; Jones, 1981), while phoronids are clearly lophophorates (Zimmer,
1973; Hermann, 1980). The trimeric coelomate condition, therefore, has evolved
more than once and of itself is not evidence for phylogenetic affinity, only of
adaptation to tubicolous life. Even tubicolous polychaetes such as serpulids tend to a
trisomic condition, while retaining metameric segmentation.
This case epitomizes the general effect of functional morphology on phylogenetic
thinking. It has made us more wary of overemphasizing general resemblances and
more honest about the gaps in the story. It has also made us realize the likelihood of
convergent evolution among complex phyla and even more, the likelihood of con-
vergent evolution among the animals of the simpler phyla. On the other hand,
functional morphology has not assisted in the positive identification of interphylum
relationships, except perhaps in relating ectoprocts to endoprocts (Nielsen, 1977) and
pogonophorans to annelids (Southward, 1980).
Fig. 18. Typical stages in polychaete spiral cleavage, (a) Arenzcola (b) Amphitrite (after Anderson, 1973).
apical tuft
stomach
protonephridium
Fug. 19. A polychaete trochophore larva.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
D. T. ANDERSON 161
Fig. 20. Stages in the spiral cleavage of the egg of the pogonophoran Nerezlinum murmanicum (after
Gureeva, 1979).
Comparative embryology
Where, then, are we to turn for evidence of phylogenetic relationships between
phyla? Fortunately, for a number of phyla, an answer still lies in embryology. Detailed
functional analyses of development have shown that developmental patterns can
remain astonishingly conservative despite wide divergences in adult form and
complexity. It is necessary, of course, to make allowance for functional specializations
of the developmental process in relation to its own requirements such as
planktotrophic adaptations and/or the presence of yolk in the embryo. Taking these
into account, we can discern certain well-established cases where embryological
studies strongly indicate phylogenetic affinities. Spiral cleavage development is one
such case.
The pattern of spiral cleavage development which includes 4d mesoderm and a
subsequent larva of the trochophore type (Figs 18-20) links many phyla, including
molluscs, annelids, pogonophorans and uniramians, with the platyhelminths
(Anderson, 1973, 1979; Gureeva, 1979; Bakke, 1980). Mostly these phyla cannot be
positively linked on other evidence. Other kinds of spiral cleavage development occur
in other phyla, linking them more remotely to the 4d group. These include (Figs 21,
22) rotifers, gastrotrichs, nematodes and crustaceans (Anderson, 1973, 1979, in
press ; Joffe, 1979). Another major grouping on embryological grounds is recognizable
from the radial cleavage, deuterostome, trimeric pattern of development which links
the hemichordates, chordates, echinoderms and chaetognaths (Philip, 1979;
Hermann, 1980, Pross, 1980). This kind of evidence works well for complex animals
where enough information is available, though a number of groups of complex
animals have still not been investigated sufficiently to allow phylogenetic inferences to
be drawn about them in embryological terms.
de aren
. caaee if
1A ay
2B 2D
Fog. 21. Stages in the spiral cleavage of the egg Fig. 22. Stages in the cleavage of the egg of a
of a rotifer, Neogossea (after Joffe, 1979). nematode, Prionchulus (after Joffe, 1979).
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
162 ORIGINS AND RELATIONSHIPS AMONG ANIMAL PHYLA
Embryological evidence is not satisfactory, on the other hand, for the simpler
groups, where development offers less information. Sponges, ctenophores and
cnidarians cannot be related to each other or to other phyla on embryological
grounds. The need for an integrated functional interpretation of the sequence of
development must also be emphasized. Evidence of this kind, when incomplete, is
easily misused. Attempts to relate the Pycnogonida to the Chelicerata and the
Pentastomida to the branchiuran Crustacea (Riley et al., 1978; Schram, 1978) on the
basis of inadequate description of a few embryo stages (Fig. 23) must remain totally
unconvincing, though there are good functional morphological reasons why the
pycnogonids should be included in the phylum Chelicerata (Manton, 1978;
Bergstrom et al., 1980).
dorsal organ
ganglion somite
Fig. 23. Two stages in the embryonic development of the pentastomid Rezghardza sternae (after Riley et al.,
1978).
PORIFERA * CNIDARIA
9O4/
4 X CTENOPHORA xf
iy
Ui
Can stage
Blastuloid
Choanoflagellate colony
Fig. 24. The phylogeny of the Radialia, as interpreted by Salvini-Plawen (1978)
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
D. T. ANDERSON 163
o jes) ~ | < (@) a Oo
fo) fo) xn > oD
; : | ate Pe ale
S 2y <
Zz | = i)
?4— —Archaeocyatha Zz >
CHOANOFLAGELLATES -|— — — > ?-—} ——Porifera a mneee
ZOOFLAGELLATES — —|— —@»> ?-—-—} —-Cnidaria
? — —-Ctenophora pe
_ Platyhelminthes [ +
7
~ _-Entoprocta
- a 7 ao
GAG) > SK ~>Ectoprocta |
/ SPIRAL \_ ee
| CLEAVAGE | ~ —Nemertea {
\ WORMS AP ~
SL U---\NSV IX. > Sipunculida |
4 \\ Se
/ \ JA. > Echiurida
/ yw
/ SS
i \ \ \ Annelida
/ \\ Ppogonophora
/ NNN
/ * ‘“Uniramia
/ NGA |
Ba ee \ Tardigrada
ao aS \
CILIATA / _ SPIRAL . Mollusca
Ei is So CLEAVAGE y= anes
E \. WORMS L& wl nmi cmUstaced
> - ROSES
i aa Se pei Chelicerata |
Ss XS
~ ~
x SRS ~~ Rotifera
a ~Nematoda
~ Nematomorpha
Gastrotricha
— — Kinorhyncha
G TRIMERIC i
~( COELOMATE Pee Et ean
ea WORMS _/ Sieh
~Brachiopoda
pease
S
( DEUTERONT
\\ STOMES x ) Hemichordata
Se See Se
_— Chaetognatha
~ Chordata
Ny
‘Echinodermata
?-|— —-Priapulida
— —-Acanthocephala lies === = —— alas
— — Pentastomida ies + 7 yo
——Trilobita hee ——_
2+ — —Hyolitha ee
27 —— OTHER | a Fal ge
Q4+—— — —)
) _ BURGESS —s igeser
Pam SHIAUE Conglomerate y
<. This paper &
This fauna Elmside Formation Philip (1971)
: ;
nN
SR Qe! pb} ¥
= as Silt one tities Cowridge
cw Ue Mudstoneqy~ JU Siltstone i :
ah See Las Rormation Link & Druce
2 (1972)
Graptolite faunas
of Jaeger (1967) Rosebank Shale
Black Bog Shale Formation
Silverdale Formation
Laidlaw Formation
Cliftonwood Limestone
O’Briens Creek
Sandstone
Douro Volcanics
Fig. 1. Stratigraphic position of fauna, in relation to other graptolites and conodonts. Column modified
after Link and Druce (1972).
Cowridge
Siltstone 4
Formation /::
“SPRING
MOUNT’ XY
Fig. 2. Location of fauna. Map modified from Link and Druce (1972).
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
C.J. JENKINS 169
For the greater part, the section is structurally uncomplicated and dips on the
bedding are at 35° towards 130°. Only at the northern end do small faults and gentle
folds appear.
The preservation of the graptolites is good. They are flattened but not
tectonically deformed, and the carbonized periderm is usually still present. A few
sparsely-distributed trilobites, brachiopods, conularids and _ scolecodonts are
associated with the graptolites. Some beds contain abundant broken plant material.
THE GRAPTOLITES
Linograptus posthumus Richter 1875.
This form is exceedingly abundant in all collections through about 20 m
stratigraphic thickness. Several specimens with sicular cladia have been recovered
(Figs 3A,B). As in typical forms of this species in other regions the thecal spacings are
about 9-10 th/cm. The stipes widen imperceptibly from 0.3-0.4 mm at th 1 to 0.7-0.8
mm distally. Minimum widths between the distal thecal apertures are 0.3 mm.
Broken, separated sections representing just the metasiculae are 1-1.2 mm long and
rather tubular in shape (0.2 mm wide). Distal thecal apertures occasionally show
evidence of cortical thickening and low, rounded lateral lappets (Fig. 3D). The
longest observed stipe portion is 3 cm (not including the long extended virgula). L.
posthumus is a long-ranging species from the Ludlovian into the Lochkovian
(Devonian) and is of little use for dating the fauna precisely.
Monograptus cf. angustidens Pfibyl 1940.
The graptolites represented in Figs 3 F, I-L, O-Q appear to constitute a single,
morphologically variable species. This is indicated by the slight continuous variation
that affects virtually all dimensions. In each case the variation coefficients are about
15%. The variation is also seen in their general appearances. Thus, while some
specimens are like Monograptus boucekz Pribyl in having a straight dorsal margin to
the thecal series at the proximal end, others have a tapered stipe proximally, which is
more like M. angustidens. The collections are thus apparently intermediate between
these two species.
Jaeger (1967) has already described M. boucekz from the stratigraphically lower
Cowridge Siltstone of the Yass syncline. These older specimens, however, show no
tendency to adopt the characteristics of M. angustzdens.
The following dimensions are representative of the collections. The rhabdosomes
may reach 40 mm in length, but are usually less than 20 mm. The siculae average 1.2
mm in length (range: 0.9-1.8 mm) and their apices reach to levels between the th 1
aperture and just above the th 2 aperture. In specimens with a straight dorsal margin
to the thecae, the 0.25 mm wide sicula produces a strong dorsal deflection of the
rhabdosome. In others that are related more closely to M. angustzdens this deflection
only corrects for the strong curvature between th 1 and th 10. A virgellar spine usually
projects down and ventrally at 20-35" from the axis of the rhabdosome. The thecae are
strongly hooked, though in a few individuals this form may weaken very slightly for the
thecae beyond th 10. Stipe widths are 0.6-0.9 mm at th 1, 1.0-1.3 mm at th5, 1.25-1.5
mm at th 10 increasing steadily to (at most) 1.7 mm distally. Stipe widths between the
thecal apertures grow from 0.4 mm just above th 1, to 0.9 mm distally. Thecal
spacings are 13-14/cm proximally and 10-11/cm beyond th 10. The distal interthecal
septa measure about 0.8 mm in length and are inclined at about 20° to the dorsal
margin.
Some of the dimensions differ consistently from the values found in overseas
specimens of M. angustidens and M. bouceki, perhaps reflecting geographic
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
LATE PRIDOLIAN GRAPTOLITES
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
C. J. JENKINS 171
subspeciation. The siculae are generally shorter and do not extend up as far as th 3.
The thecae are less closely spaced proximally and the stipes are wider at full
development.
Monograptus cf. formosus Boutek 1931.
A single incomplete specimen (Fig. 3E) represents a 9 mm long portion of
strongly dorsally curved stipe that is only 0.25 mm wide between the thecae and 0.7-
0.8 mm wide at the hooked, partly-isolated thecae. The interthecal septa are about
0.5 mm long. The fragment appears to represent a middle portion of the graptolite,
where the thecae are spaced at 10/cm. These dimensions compare well with those of
the M. formosus from the Rosebank Shale in the Yass syncline, (Jaeger, 1967, pl, 14,
figs. b, c) and from overseas successions. The form is not Monograptus paraformosus
Jackson and Lenz (1969) as the thecae are only moderately isolated.
Monograptus transgrediens Perner 1899.
This graptolite is present, but never common in the beds. It attains 30 mm in
length and 1.8 mm in width. Except for th 1 which appears to be slightly hooked or to
have lappets, all thecae are simple. Distal thecae are spaced at 9th/cm, proximal
thecae at about 12th/cm. The sicula is about 1 mm long and has a ventral virgella.
The rhabdosome is strongly tapered at the proximal end, causing a pronounced
ventral curvature of the dorsal margin.
Monograptus transgrediens cf. praecipuus Pribyl 1940.
Only two specimens are available (Figs 3 G-H). They possess the distinctive
ventral curvature of the proximal parts in the stipes, the rather narrow stripe width
(1.2-1.7 mm) in fully-developed overseas examples), and the mainly pristiograptid
thecae, except for th 1 and 2 which Lenz and Jackson (1971) describe as having a
‘beak-like profile’. This may be due to the presence of substantial lappets at the
apertures. The Yass specimens reach up to 23 mm in length and have stipe widths of
0.5-0.7 mm at th 1, 0.7-0.8 mm at th 5, and 0.8-1.0 mm at th 10. One possible distal
fragment (not figured) has a width of 1.8 mm. One sicula is 1.3 mm long, with its
apex lying level with the aperture of th 1. The primary difference between this and
Overseas representatives is the lesser stipe width at th 10.
FAUNAL RELATIONSHIPS
Late Silurian and Early Devonian graptoloid faunas show no detectable
geographic provincialism and can be used for very accurate worldwide
biostratigraphic correlation (Jaeger, 1978).
The fauna in the lower Elmside Formation is clearly late Pridolian in age. It is a
natural continuation of the graptolite faunas occurring lower in the Yass sequence and
it contains several species in common with the older faunas.
M. angustidens (as M. unzformis angustidens) is reported as a short-ranged
species characteristic of the very base of the Lochkovian (Jaeger, 1977) . However, it is
also recorded from the very uppermost Pridolian, apparently below the first
appearance of Monograptus uniformis uniforms Pfibyl — as for instance in Canada,
Fig. 3. Graptolites from the Elmside Formation, near Yass, NSW. A-D. Linograptus posthumus (Richter) .
A, 35971. B, 35907. C, 35965. D, 35967. E, Monograptus formosus Boucek, 35910. F, I-L, O-Q.
Monograptus cf. angustidens Pribyl. F, 35920. I, 35920. J, 35895. K, L, 35964. O, 35896. P, 35899. Q,
35908. G-H. Monograptus transgrediens cf. praecipuus Pribyl. G, 35911 & 35916. H, 35915. M-N, 35925.
All specimens held in Australian National University Palaeontology Collection. All magnifications are X
5.0 except C, D, F, K and L which are X 10.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
Nei LATE PRIDOLIAN GRAPTOLITES
Poland and probably Czechoslovakia (Lenz and Jackson, 1971; Tomcezyk et al., 1977;
Teller, 1964; Jaeger, 1967). Representatives in the lower Elmside Formation, appear
to represent an early intermediate of this form with the older species M. boucekz,
which is usually regarded as typical of graptolite horizons from lower to middle
Pridolian. M. unzformzs — the primary species which indicates the base of the
Devonian — has not appeared in any of the graptolite collections. A further indication
that the faunas are not Devonian is the presence of M. transgredzens (Jaeger, 1977).
The Elmside Formation graptolite assemblages reinforce the conodont-based
biostratigraphy of Link and Druce (1972) which dated limestones bearing Icriodus
‘woschmzdti in the upper Elmside Formation as basal Lochkovian. However, they have
made it possible to locate the boundary of the Devonian more accurately in the
succession, and show that it lies in the middle part of the Elmside Formation.
ACKNOWLEDGEMENTS
The first discovery of the fauna was made during a Geological Society of Australia
Sunday picnic (November 1980). I am indebted to Dr K. A. W. Crook for drawing
attention to the occurrence and checking a draft of the manuscript.
References
Jackson, D. E., and Lenz, A. C., 1969. — Latest Silurian graptolites from Porcupine River, Yukon
Territory. Bull. geol. Surv. Can. 182: 17-29.
JAEGER, M., 1967. — Preliminary stratigraphical results from graptolite studies in the Upper Silurian and
Lower Devonian of southeastern Australia. J. geol. Soc. Aust. 14: 281-286.
——, 1977. — Graptolites, pp. 337-345 zn MartTINssoN, A., (ed.), The Silurcan-Devonian boundary.
(International Union of Geological Sciences, Series A, number 5.). Stuttgart: Schweizerbart’sche
Verlagsbuchhandlung.
———, 1978. — Late graptoloid faunas and the problem of graptoloid extinction. Acta pal. Pol. 23: 497-
521.
LENz, A. G., and JAcksOoN, D. E., 1971. — Latest Silurian (Pridolian) and Early Devonian Monograptus of
northwestern Canada. Bull. geol. Surv. Can. 192: 1-24.
Link, A. G., 1970. — Age and correlations of the Siluro-Devonian strata in the Yass Basin, New South
Wales, J. geol. Soc. Aust. 16: 711-22.
———, and DruceE, E. C., 1972. — Ludlovian and Gedinnian conodont stratigraphy of the Yass Basin, New
South Wales, Bull. Bur. Miner. Resour. Aust. 134: 1-136.
Puitip, G. M., 1971. — The Silurian-Devonian boundary in Australia. Geol. Newsletter (1971) 1: 12-15.
Strusz, D. L., 1980. — The Encrinuridae and related trilobite families, with a description of Silurian
species from southeastern Australia. Palaeontographica (A) 168: 1-148.
TELLER, L., 1964. — Graptolite fauna and stratigraphy of the Ludlovian deposits of the Chelm borehole,
eastern Poland. Stud. geol. Pol. 13: 1-88.
Tomczyk, H., PajcHLowA, M., and Tomczykowa4, E., 1977. — Poland, pp. 65-83 7m MARTINSSON, A.,
(ed.), The Stlurtan-Devonian boundary. (International Union of Geological Sciences, Series A,
number 5). Stuttgart: Schweizerbart’sche Verlagsbuchhandlung.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
Darriwilian (Middle Ordovician) Graptolites
from the Monaro Trough sequence
east of Braidwood, New South Wales
C. J. JENKINS
JENKINS, C. J. Darriwilian (Middle Ordovician) graptolites from the Monaro Trough
sequence east of Braidwood, New South Wales. Proc. Linn. Soc. N.S.W. 106
(2), (1981) 1982: 173-179.
Darriwilian graptolites have been found in hemipelagic mudstones interbedded
with quartz-rich arenites of turbidite origin, in the Monga area, 13 km ESE of
Braidwood, N.S.W. This is the oldest reliably-dated sediment of the Monaro Trough
sequence, in the easternmost part of the Lachlan Fold Belt of New South Wales. The
fauna is diverse and is correlated with assemblages close to the boundaries of the Da 2
and Da 3 Zones in Victoria and the Paraglossograptus tentaculatus and Diplograptus?
decoratus Zones of New Zealand.
C. J. Jenkins, Department of Geology, Australian National University, P.O. Box 4,
Canberra City, Australia 2600; manuscript received 10 November 1981, accepted for
publication 16 December 1981.
INTRODUCTION
This short article describes a find of Middle Ordovician graptolites from the very
easternmost part of the Lachlan Fold Belt in southeastern New South Wales. The
locality lies 13 km directly ESE of Braidwood, in a region of geology that has been
termed the Monaro Trough by Crook and Powell (1976) and others.
The discovery is a valuable one because potentially it indicates the environments
of deposition at a very early stage in the development of the Monaro Trough, in the
eastern part of the Lachlan Fold Belt.
Furthermore, it marks a considerable extension of the extent of Middle
Ordovician sedimentation into that broad region of the Lachlan Fold Belt which lies
east of Canberra. Previously the nearest proven occurrence of Middle Ordovician
strata was at Canberra, 75 km to the west (Nicholl, 1980). The existence of beds of
this age in the region has been suspected (Webby, 1976), but only on very speculative
stratigraphic grounds. The Wagonga Beds, for example, have been thought to range
through the Middle Ordovician, but are now believed to be entirely Upper Ordovician
in age (Jenkins et al., in prep.). The locality lies well to the east of the strike of the
Middle Ordovician graptolite localities in Victoria — for instance those at
Tabberabbera (Beavis et al., 1976, p. 34).
Other Ordovician graptolite localities lie scattered around Braidwood (Sherrard,
1954, 1962) but none of these faunas is older than the Eastonian division of the Upper
Ordovician.
GEOLOGICAL SETTING
The locality occurs in the southern bank of a road cutting on the highway, 650 m
northwest of the crossing of the Mongarlowe River (Fig. 1). It lies 15.5 km by road
east of Braidwood, near Currajugg homestead and at grid reference 655673 on the
Monga 1: 100,000 sheet. At the present time there is no formal lithostratigraphic
name for the sediments.
The graptolites occur in two of the four isolated lumps of black mudstone that are
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
174 DARRIWILIAN GRAPTOLITES
GOULBURN
e
YASS
© COOMA
Fig. 1. Location of fauna.
exposed in the road cut. All lie within 20 m of the western end of the outcrop. Dips are
generally at 35° to the east but open folding, faulting and the intrusion of a number of
1-3 m thick (?)dolerite dykes causes variations. The sediments are all slightly
hornfelsed.
The mudstones appear to be of hemipelagic origin and are surrounded by well-
bedded grey- and purple-toned, coarse- to fine-textured greywackes, siltstones and
mudstones which are interpreted as turbidites.
Most beds are very continuous laterally, but some of the turbidite-siltstones and
all of the hemipelagic black mudstones rapidly pinch out laterally, in one case through
150 mm thickness in only 1-3 m distance. This may have been due to ‘boudinage’
effects in the soft-sediments (McCrossan, 1958) or to penecontemporaneous and
small-scale slumping of the deposits — a common process on continental slopes
(Nardin et al., 1979). Four separate occurrences of the black mudstone have been
discovered, each less than 1 m in extent and 150 mm thick. The westernmost has
disrupted bedding and disoriented fragmented graptolites, both suggesting a degree of
post-depositional fluidization or remobilization. The lenses clearly do not represent
clastic blocks within the greywackes.
One of the greywackes has been examined in thin section (ANU slide 13110).
The estimated proportions of the minerals are: 30% quartz (angular, unstrained
grains) , 20% altered coarse detrital micas (?originally biotite, now sericite) and 50%
groundmass (probably mainly of sericite). Grains of reworked black mudstone and
particles of green detrital chlorite and green euhedral tourmaline were also present,
but altogether constituted far less than 1% of the rock.
THE GRAPTOLITES
The following species have been identified in the mudstone bearing the most
prolific fauna, about 12 m from the western limit of the outcrop and about 2 m above
the roadside gutter level: Didymograptus cognatus Harris & Thomas, Tetragraptus
sp., ?Pseudobryograptus sp., Isograptus sp., ?Apiograptus crudus (Harris &
Thomas), Glossograptus acanthus Elles & Wood, Paraglossograptus cf. tentaculatus
(J. Hall), Cryptograptus cf. schaefert Lapworth, Glyptograptus intersitus Harris &
Proc. LINN. Soc. N.S.W., 106 (2), (1981) 1982
C. J. JENKINS 175
Fog. 2. A-D, Didymograptus cognatus Harris & Thomas; A, B, proximal ends, specimens 35869, 35887, C,
D, distal stipe portions, 35837, 35837; E-G, Tetragraptus sp., 35858, 35877, 35878; H-J, Isograptus sp.,
35880; 35883; 35882; K-N, Glossograptus acanthus Elles & Wood, 35876, 35855, 35847, 35886; O,
scalariform Glossograptus indet. 35874; P, ?Apiograptus crudus Harris & Thomas, 35888; Q-T,
Paraglossograptus cf. tentaculatus (J. Hall), 35862, 35846, 35856, 35851; U-Y, Cryptograptus inutzles
Hall, 35871, 35885, 35863, 35867, 35868.
All at x5 magnification. All specimens in the Australian National University Geology Department
Palaeontology Collection.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
176 DARRIWILIAN GRAPTOLITES
Thomas, Diplograptus? decoratus (Harris & Thomas). A brachiopod — perhaps
Palaeoglossa — has also been collected.
The other black mudstones are apparently barren except for one about 2 m from
the western end of the cutting, which has yielded one small specimen of each of D.
cognatus and ?Glossograptus sp.
The state of preservation of the graptolites is poor and tectonic strain has
produced the thin and flattened specimens which are now replaced by phyllosilicates.
The bars drawn on Figs 2 and 3 represent the direction of maximum relative
elongation as seen in the bedding.
In some cases taxonomic problems and the poor preservation have led to some
difficulties with the identifications. The remainder of this section discusses the details
of the identifications.
The specimens of D. cognatus (Fig. 2 A-D) are fragmentary. They are close in
form and in their dimensions to several described species including Didymograptus
Fig. 3. A-F, 2G, J. Glyptograptus intersitus Harris & Thomas, specimens 35844, 35849, 35875, 35842,
35839, 35873, 35879, 35865; ?H, I. Pseudoclimacograptus differtus (Harris & Thomas) 35854, 35848; 2K,
L. Diplograptus? decoratus (Harris & Thomas) 35838, 35864.
All at x5 magnification. All specimens in the Australian National University Geology Department
Palaeontology Collection.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
C. J. JENKINS 177
compressus Harris & Thomas and Didymograptus nicholsoni Lapworth so the specific
identification is not entirely certain. Small Tetragraptus species similar to that of Fig.
2 E-G are common in Middle Ordovician strata worldwide, but few have been
formally named or distinguished from Lower Ordovician forms. The name ‘T.
clarkfielde’ of Thomas (1960) which probably represents conspecific Darriwilian
material from Victoria is unfortunately a nomen nudum. The identification of
2Pseudobryograptus sp. is especially tentative, being based only on a few stipe
fragments. The small Isograptus sp. (Fig. 2 H-J) is a common form, close to the
Darriwilian Tsograptus sp.’ of Cooper (1973, p. 89). It may represent a population of
small-sized I. forczpiformzs Ruedemann or I. wictoriae divergens (Harris) which are
both common large species at this level (Cooper, 1973).
?Apiograptus crudus is identified on the basis of only one specimen (Fig. 2 P)
which is clearly not Glossograptus as the thecae are simple and dichograptid.
Furthermore, the long spines and produced ventral apertural processes of that genus
are clearly not present and the rhabdosome is bluntly rounded proximally. In spite of
the poor preservation and the tectonic distortions, an identification as Phyllograptus
or Pseudotrigonograptus can be discounted for the distal end is untapered, a nema is
present, and there is no evidence of the scalariform 3rd and 4th stipes. Although ?4.
crudus is generally regarded as restricted in range to the Yapeenian (Cooper, 1979, p.
80; Cooper and McLaurin, 1974) it appears to be present in the Darriwilian in both
N.S.W. and Victoria. Harris and Thomas (1935, fig. 1, no. 13) record a specimen in
association with Darriwilian faunas east of Bendigo, Victoria and this specimen was
included in the synonymy of A. crudus in Cooper and McLaurin (1974).
The identifications of the glossograptids and cryptograptids are fairly free of
problems and follow those of Cooper (1979). G. acanthus (Fig. 2 K-N) is an
especially common form. Some small specimens (Fig. 2 U-Y) probably represent
Cryptograptus inutzlis, a species that is commonly recorded at this horizon. The
laciniae on the specimens identified as P. cf. tentaculatus (Fig. 2 Q-T) are often
difficult to see, but a few instances have been arrowed on the figures. The
rhabdosomal shape and the dimensions of the specimens agree with those of P.
tentaculatus. Another specimen in the fauna (Fig. 20) probably represents one of the
above glossograptids in scalariform view.
The details of thecal form on the diplograptids have been badly affected by poor
preservation and the tectonic strains, particularly where the specimens have
undergone a strong degree of shortening (e.g. Fig. 3 B, F). Nevertheless P. dzffertus,
G. intersitus and D.? decoratus can be recognized from a number of the better-
preserved specimens (Fig. 3). None of the specimens of D.? decoratus has the distal
vane structure which is so often present in the species.
FAUNAL RELATIONSHIPS AND CORRELATIONS
The fauna is obviously very similar to those of the Darriwilian in Victoria and
New Zealand. These similarities are the basis of proposed stratigraphic correlations.
The zonation of the Victorian faunas is based on a succession of diplograptid species
and the presence together of G. zntersttus and D.? decoratus in the Braidwood fauna
indicates that it represents a level close to the boundary between the Da 2 and Da 3
zones of Victoria (Thomas, 1960). However, the greater abundance of G. zntersitus
suggests that it lies within the Da 2 zone. The remainder of the species are consistent
with this conclusion, except perhaps ?A. crudus which is generally regarded as
significantly older, and also P. dzffertus which Thomas (1960) depicts as a Da 4 zone
species. However as noted already D. crudus is known from the Darriwilian in
Victoria. Furthermore, P. differtus was recorded from the “D 1 zone” by Harris and
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
178 DARRIWILIAN GRAPTOLITES
Thomas (1935) and in Harris (1935); (their now superseded ‘D 1 zone’ equates with
the present Da 2 and Da 3 zones; Webby 1976, p. 422).
The zonation of the Middle Ordovician graptolites from the South Island of New
Zealand recently described by Cooper (1979), is based on the first appearances of a
few diverse forms of graptolites. Thus, the occurrence of D.? decoratus in the
Braidwood assemblage indicates the D.? decoratus Zone. However its association with
P. cf. tentaculatus, G. acanthus and G. intersztus is suggestive of horizons rather low
in the zone. The general species composition confirms this as it matches that of the
underlying P. tentaculatus Zone more closely than the Zone of D.? decoratus. The
only species inconsistent with such a level is, again, ?A4. crudus.
Middle Ordovician faunas of similar aspect are known also from Texas (Zone 9 of
‘Hallograptus etheridge: (= P. tentaculatus) ; Berry, 1960) and Peru (Lemon and
Cranswick, 1956).
Comments on the relations between the Braidwood fauna and the two other
Middle Ordovician graptolite assemblages known in N.S.W. from Canberra (Opik,
1958) and near Narrandera (Sherrard, 1954) must await re-examinations of these
collections, though it is clear from the listed faunas (Packham, 1969, pp. 89, 102)
that levels in the Darriwilian are again represented.
CONCLUSIONS
The locality extends the known extent of Middle Ordovician deposition in south
eastern Australia. It is a significant one, being the lowest reliably-dated horizon in the
Lachlan Fold Belt east of the Canberra-Molong High, in the area termed the Monaro
Trough. The sediments are not significantly different in character from the younger
flysch, being quartz-rich turbidites and black shales, indicating that similar,
reasonably deep ocean conditions prevailed at this early stage in the region’s
development.
ACKNOWLEDGEMENTS
Dr M. J. Rickard (ANU) first discovered the locality. Peter Ward and Dr K. A.
W. Crook (ANU) have helped with subsequent collecting. Both Dr Rickard and Dr
Crook kindly checked an early draft of the article.
References
BEAvIs, F. C., FERGUSON, J. A., TALENT, J. A., TATTAM, C. M., and Tuomas, D. E., 1976. — Ordovician.
In Douctas, J. G., and Fercuson, J. A., (eds), The geology of Victoria. Spec. Publ. Geol. Soc.
Aust. 5: 24-44.
Berry, W. B.N., 1960. — Graptolite faunas of the Marathon Region, West Texas. Publ. Bur. Econ. geol.
Univ. Texas 6005: 1-179.
Cooper, R. A., 1973. — Taxonomy and evolution of Isograptus Moberg in Australasia. Palaeontology 16:
45-115.
———, 1979. — Ordovician geology and graptolite faunas of the Aorangi Mine area, north-west Nelson,
New Zealand. Palaeont. Bull. geol. Surv. N.Z. 47: 1-127.
———, and McLaurin, A. N., 1974. — Apiograptus gen. nov. and the origin of the biserial graptoloid
rhabdosome. Spec. Pap. Palaeont. 13: 75-85.
Crook, K. A. W., and PowELL, C. Mc. A., 1976. — The evolution of the southeastern part of the Tasman
geosyncline. 25th Int. Geol. Congr. Field Guide 17A.
Harris, W. J., 1935. — The graptolite succession of Bendigo East, with suggested zoning. Proc. R. Soc.
Vict. 47: 314-37.
———, and Tuomas, D. E., 1935. — Victorian graptolites (New Series) Part III. Proc. R. Soc. Vict. 47:
288-313.
JENKINS, C. J., Kipp, P. R., and MILts, K. J., 2m prep. — Eastonian age graptolites from the Wagonga Beds,
near Batemans Bay, N.S.W.
Proc. LINN. Soc. N.S.W., 106 (2), (1981) 1982
C. J. JENKINS 179
Lemon, R. R. H., and Cranswick, J. S., 1956. — Graptolites from Huacar, Peru. Publ. Mus. Hist. nat.,
Lima, C. (Geol.) 5: 1-31.
McCrossan, R. G., 1958. — Sedimentary ‘Boudinage’ structures in the Upper Devonian Ireton Formation
of Alberta. J. sedim. Petrol. 28: 316-20.
Narpin, IT. R., Epwarps, B. D., and Fors.in, D. S., 1979. — Santa Cruz Basin, California Borderland:
Dominance of slope processes in basin sedimentation. Spec. Publ. Soc. Econ. Paleont. Min. 27: 209-
21.
Nico, R. S., 1980. — Middle Ordovician conodonts from the Pittman Formation, Canberra, A.C.T.
B.M.R. J. Aust. Geol. Geophys. 5: 150-153.
Opik, A. A., 1958. — The Geology of the Canberra City district. Bull. Bur. Miner. Resourc. 32, 1-99.
PAckHAM, G. H. (ed.), 1969. — The geology of New South Wales. J. geol. Soc. Aust. 16: 1-654.
SHERRARD, K. M., 1954. — The assemblages of graptolites in New South Wales. J. Proc. R. Soc. N.S.W.
87: 73-101.
———, 1962. — Further notes on assemblages of graptolites in New South Wales. J. Proc. R. Soc. N.S.W.
95: 167-78.
Tuomas, D. E., 1960. — The zonal distribution of Australian graptolites. J. Proc. R. Soc. N.S.W. 94: 1-58.
Wespy, B. D., 1976. — The Ordovician System in South-eastern Australia, pp. 417-446 in BAssETT, M. G.
(ed.). The Ordovician System: proceedings of a Palaeontological Association symposium,
Birmingham, September 1974. Cardiff: Univ. Wales Press and National Museum of Wales.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
H aah an ray PV ha
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scalars ; ’ ; Wd
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106 Ci/2):
The Fauna of Australian
Seagrass Beds
PAT HUTCHINGS
Hutcnines, P. A. The fauna of Australian seagrass beds. Proc. Linn. Soc. N.S.W.
486-65, (1981) 1982: 181-200.
Australia has a rich seagrass flora (26 species), widely distributed in estuaries
and shallow coastal embayments. The fauna of the seagrass beds is diverse and consists
of infaunal, epifaunal, epiphytic and epibenthic species. This community is composed
of temporary and permanent residents exhibiting tidal, diurnal and seasonal
fluctuations. The literature dealing with the fauna of Australian seagrass beds is
reviewed and possible food-web relationships and energy-flows within seagrass
communities are postulated.
P. A. Hutchings, Curator of Marine Invertebrates, Australian Museum, P.O. Box
A285, Sydney South, Australia 2000; manuscript recetved 19th May 1981, accepted in
revised form 16 December 1981.
INTRODUCTION
Seagrass beds occur in estuaries and shallow coastal embayments throughout
mainland Australia (Larkum, 1976a). In many estuaries they form extensive beds. In
eastern temperate Australia, seagrass beds often occur with mangrove and salt marsh
communities which together constitute .coastal wetlands (Hutchings and Recher,
1974; AMSA, 1977). In tropical regions this relationship rarely occurs. Instead,
seagrass beds may be associated with coral reefs.
Recently in the Caribbean, Ogden and Zieman (1977) have shown the close
relationship between coral reefs and seagrasses. Herbivorous coral reef fish migrate
from the reef to nearby beds of Thalassza testudinum (Banks ex Konig) to feed.
Carnivorous fish and grazing sea urchins move at night from the reef to the seagrass
beds to feed. Such migrations have not been documented in Australia.
The role of seagrass beds in estuaries and shallow coastal embayments has become
apparent during the past decade in Australia. Seagrass beds provide important
habitats for juvenile stages of commercial fish species (Bell et al., 1978a, b) and
prawns (Young, 1978), They stabilize and trap sediments improving water clarity
(Orth, 1977). A rich fauna is associated with the beds. There are few herbivores of
Australian seagrasses and seagrass beds also produce large amounts of organic matter
which is then broken down by the fauna of seagrass beds to detritus on which estuarine
food chains are dependent (Kirkman and Reid, 1979).
Overseas, considerable information on the fauna of seagrass beds exists. An
overview of seagrass faunas throughout the world is provided by Kikuchi and Péres
(1976). They describe the fauna of each region, and then attempt to assess the
importance of seagrasses in providing a structural habitat, shelter and a food resource.
Food web relationships and energy flows in the seagrass ecosystems are discussed.
Other studies have concentrated on regional seagrass communities (Orth, 1973, 1977;
Heck, 1977, 1979). Heck studied in detail the epibenthic invertebrates of Thalassza
beds in Panama. Considerable seasonal fluctuations in both species number and
abundance occurred. Sites varied in species composition and abundance and Heck
attributed this to the proximity of surrounding habitats especially coral reefs which
Proc. LINN. Soc. N.S.W., 106 (2), (1981) 1982
182 FAUNA OF AUSTRALIAN SEAGRASS BEDS
contain a number of species that utilize the seagrass meadows. Other workers like
Nelson (1979) have studied in detail one faunal component, such as the amphipods.
Other workers have experimentally manipulated seagrass beds to assess the value
of habitat complexity in determining species richness and abundance (Heck and
Wetstone, 1977). Young and Young (1977, 1978) have studied the effects of
predation at various times of the year by caging, adding dense populations of
suspension feeders and providing organic enrichment.
Breakdown of seagrass into detritus has been investigated by Robertson and
Mann (1980). Isopods and amphipods are important in shredding the intact dead
seagrass leaves. The decomposition of organic detritus has been extensively reviewed
by Fenchel (1977). Utilization of this detritus and export of this productivity out of
the seagrass beds has been considered by Ogden and Zieman (1977; Ogden, 1976)
who have studied fish feeding in Caribbean seagrass beds.
OBJECTIVES OF THE REVIEW
To collate and synthesize all the available information on Australian seagrass
faunas. This will reveal the dominant species whose life histories and trophic status
merit detailed investigations. Such information is essential for estimates of secondary
productivity to be made for although it is widely recognized that Australian seagrass
beds are important, this is largely based on inference from overseas data. Data on
secondary productivity of Australian seagrass beds are non-existent.
From overseas data and limited Australian data there are sound financial and
ecological reasons for maintaining seagrass beds (Larkum, 1981). Maintenance
requires management but we lack the necessary information for implementation.
Instead, as the use of estuaries increases (Burnley, 1974), seagrass beds are being
degraded. Larkum (1976b) has shown that the seagrass beds on the northern shores of
Botany Bay, New South Wales, have disappeared because of pollution from Cooks
River. Seagrass beds on the southern shores of Botany Bay are affected by increased
wave action and decreased water clarity caused by dredging associated with the port
development. Similar port and industrial developments in Cockburn Sound, Western
Australia, have also caused widescale disappearance of seagrass beds during the 1970s
and this is well documented by Cambridge (1979).
As estuarine resources are limited in Australia it may be necessary to sacrifice
some seagrass beds to keep others. This necessitates the ranking of beds in terms of
ecological importance. This can only be done if detailed information exists on a
variety of seagrass communities in differing environments.
Finally, it is hoped that this review will reveal the major gaps in our knowledge
and stimulate further research.
FLORISTICS OF SEAGRASSES
Seagrasses are angiosperms which are able to live permanently in seawater. Not
only are they able to fulfil their normal metabolic activities when fully submerged in
seawater but can also complete their generative cycle. All seagrasses have in common a
well developed anchoring system. In addition, they can compete successfully with
primarily marine organisms such as algae. This combination of properties is so rare
among angiosperms that only fifty two species of seagrass have been described (den
Hartog, 1970, 1979). Other species are currently being described (Larkum, pers.
comm.). Twenty six of these species occur in Australia (Larkum, 1976a). Since 1976
three new species have been described Pos¢donza senuosa, Cambridge and Kuo, P.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
P. HUTCHINGS 183
augustifolia, Cambridge and Kuo, and Halophila tricestata, Greenway. Larkum
suggests that latitude 25°S on the west coast and 30°S on the east coast of Australia
form a boundary between temperate and tropical species. South of these latitudes the
most successful community is formed by Poszdonza australis Hook f. This species forms
extensive monospecific beds in sheltered conditions. In New South Wales and Western
Australia the beds rarely extend below -10m (depth below mean low water mark),
but in the Great Australian Bight, they occur in more exposed conditions to depths of
-40m. Poszdonia is not a colonizing species and only appears after the substrate has
been stabilized by species such as Amphibolis antarctica (Labill) in Western and
South Australia and Zostera capricornt Aschers in New South Wales. The
development of P. australis is accompanied by the formation of a tough, wave resistant
rhizome ‘matte’ (cf. Molinier and Picard, 1952).
Larkum (1976a) has summarized on a state-wide basis the distribution of
seagrasses. Since Larkum’s review, Shepherd and Womersley (1976) have described
the seagrass communities around St. Francis Island in the northern Great Australian
Bight. New South Wales State Fisheries are currently documenting seagrass
distributions along the coast (Evans and Gibbs, in press) .
Considerable gaps in distribution data occur, especially in tropical regions where
many species occur. Den Hartog (1970) provides the only information on tropical
seagrass communities, none of which reach the density or areal size of Poszdonza
australis in temperate waters. a map detailing the distribution and extent of
Australian seagrasses is given by McComb et al., 1981.
Associated with the seagrasses may be rhizophytic companion species such as
Caulerpa and Halimeda. Such species occur mainly in sub-tropical or tropical
communities. A film of microalgae, especially diatoms, occurs on the muddy or sandy
substratum in between the seagrasses. Algae are often trapped by the seagrass and may
continue to grow in the shelter of the beds (den Hartog, 1979). This associated flora
has been neglected in Australia.
Primary Productivity Studies
McRoy and McMillan (1977) have suggested that such common plants as
seagrasses with such a high primary production must be of immense ecological
importance. Consequently, ‘seagrass ecosystems’ have become a popular focus for
research throughout the world, including Australia. Recently, considerable advances
have been made in measuring the rates of primary production of some temperate
species here in Australia. ;
The preferred method of measuring primary productivity amongst seagrass
workers seems to be measuring the production of organic matter directly, i.e. by
measuring the increase in blade length. This method has the distinct advantage that it
can give an integrated measure of primary productivity over a number of days. The
method has drawbacks, in that it measures only the productivity of part of a plant, the
frond, and it does not take into account any losses of dissolved organic carbon
compounds (Larkum, 1981), which may be significant (Kirkman and Reid, 1979).
Rates of leaf blade production of Posidonza australis range from 0.7-5.5 g dry wt.
m~ day, depending on site and season. It is significantly higher in clear waters, such as
Jervis Bay, than in disturbed areas like Botany Bay, New South Wales. Maximum
rates occur in the middle of the geographical range of this species, which is Spencer
Gulf, South Australia (West and Larkum, 1979).
Leaf production is markedly seasonal with a maximum production between
November and February, and a minimum in July to August. However, if flowering
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
184 FAUNA OF AUSTRALIAN SEAGRASS BEDS
occurs in July, then leaf blade production is severely curtailed. In Botany Bay three to
four crops of leaves are produced annually. In areas of frequent storms, biomass
figures may be relatively constant (West and Larkum, 1979).
In some areas such as Westernport Bay, Victoria, nitrogen limits the growth of
Heterozostera tasmanica during spring and early summer (Bulthuis and Woelkerling,
1981). This species and some other seagrasses, including Zostera marina, can absorb
nutrients from the sediments and translocate them to the leaves (Twilley e¢ al., 1977).
Zostera capricorn shows a similar growth pattern (Larkum et al., in press) which
seems to be more closely related to water temperature than to solar radiation. New leaf
production continues fairly uniformly throughout the year with a slight increase in
the warmer months. Turnover times for leaves vary from about 33 days in the summer
months to 67 days for the winter period (Kirkman et al., 1982).
In Port Hacking, New South Wales, Zostera grows 3-4 times faster than Poszdonia
australis. As these two seagrasses occur in varying densities, comparisons of production
per unit area of seagrass beds are more meaningful. Zostera production varies from
0.56-1.82 g m7? day" while P. australis ranges from 0.57-1.44 g m? day! —
throughout the year (Kirkman and Reid, 1979; Kirkman et al., 1982).
SEAGRASS FAUNAS
STRUCTURE
The animal communities of seagrasses are considered here in four major
categories :
a. infauna — animals living in the sediment associated with the rhizomes of the
seagrasses,
b. epifauna — animals living on the surface of the mud at the base of the seagrass,
c. sessile communities on the blades of the seagrass and,
d. epibenthic animals, swimming in the water column between the seagrass.
SAMPLING METHODS
When designing sampling programmes the three-dimensional nature of the
habitat must be considered, for each of the dimensions has an associated fauna.
Seagrass communities which are exposed to air during low tide, lose one of the
dimensions as the blades collapse eliminating the space between them. This dimension
is restored on a rising tide. As discussed later, the fauna shows distinct diurnal, tidal
and seasonal variations in composition.
The fauna can also be divided into permanent and temporary residents.
Permanent residents spend their entire lives within the seagrass beds. Temporary
residents spend only part of their life within the community. This element is rather
heterogeneous and consists of :
1. fishes and crustaceans which spend part of their juvenile life in the beds,
2. tidal migrants, which move into the beds on a rising tide and leave on an ebb
tide and terrestrial species which feed in the beds during low tide,
3. vertebrate grazers, such as ducks, turtles and dugongs.
Thus a seagrass community is in a constant state of flux as the relationship of the
community and its environment change. The community can be considered as having
a central core of residents whose numbers fluctuate seasonally, and migrants whose
composition varies according to the time of day, tide and season.
To devise one sampling programme to study seagrass faunas is impossible. Each
group of fauna necessitates a specific sampling programme, with various periodicities
and intensities.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
P. HUTCHINGS 185
a. Infauna:
The dense rhizome ‘matte’ of seagrass beds which is well developed in Poszdonza is
the major problem to overcome in sampling infauna. Collett et al. (in MS) used a
hand-operated corer with a good cutting edge to penetrate the ‘matte;. Poiner (1980)
successfully used a 0.1 m? Smith — MacIntyre grab in beds of Halophila and
Hutchings et al. (1978) used a similar grab in Zostera and patchy Poszdonia. This
grab is not efficient in dense Poszdonza.
Collett et al. (in MS) found that 15-20 samples (0.03 m*) in Poszdonza beds in
New South Wales were needed to sample 70% of the smaller infaunal species. The
samples were collected from an area of visually uniformly dense beds. Hutchings e¢ al.
(1978) sampling Zostera and patchy Poszdonza found that the number of samples
required to reach the upper asymptote of the respective species-volume curve varied
from 4-10 samples.
All infaunal studies should ascertain the minimum number of samples necessary
to collect 70% or more of the species present. This has not been standard practice.
The most commonly used mesh size is 1 mm (Collett ef al., in MS; Poiner, 1980;
Rainer, 1981; Rainer and Fitzhardinge, 1981). To facilitate the sorting of animals
from in amongst the large amount of plant matter, add Rose Bengal to the 7%
formalin solution, until a deep pink colour develops. All living, animal material
becomes dark pink in colour, considerably facilitating sorting. Micro-crustaceans can
be collected from the sieved fractions by running a steady stream of water over the
samples. The micro-crustaceans float off the samples which are gently agitated and
are retained in a sieve.
Poiner (1980) determined the density of seagrasses in the area of his infaunal
samples by counting the number of nodes per 0.25 m?. He also analysed the sediment
using standard methods of granulometric analysis (Buchanan, 1971). Such associated
data are invaluable in interpreting the benthic fauna and should become standard
practice.
Lewis and Stoner (1981) have recently compared macrofaunal sampling methods
in seagrass beds. Three different-sized core samples were tested for their relative
efficiencies. Core samples were sieved with both 0.5 and 1.0 mm mesh screens. The
smaller core was the most efficient and this was attributed to the problems of
penetrating the dense root ‘matte’. The 0.5 mm sieve retained approximately 50%
more individuals than the 1.0 mm sieve.
For sampling larger infauna such as bivalves. Anadara and Trapezza, 0.25 m®*
quadrats are laid and the sediment searched within each quadrat by a diver.
Hutchings and Recher (1974) used 40 such quadrats.
b. Epzfauna:
Techniques that have been used include visual observations by divers along
transects (Rainer, 1981; Shepherd, 1974) and small seines pulled through the
seagrass (Hutchings and Recher, 1974). Both methods are qualitative.
No quantitative study of this community has been undertaken in Australia.
c. Epiphytes :
No quantitative survey has been carried out on this fauna. Hutchings and Recher
(1974), while measuring density of seagrasses, scraped off the associated fauna. The
polychaetes are described by Hutchings and Rainer (1979).
d. Epzbenthic:
Young (1978) sampled with a roller beam trawl between fixed markers
Proc. LINN. Soc. N.S.W., 106 (2), (1981) 1982
186 FAUNA OF AUSTRALIAN SEAGRASS BEDS
fortnightly over a 12 month period. All stations were sampled within 2 days of the new
and full moon. All samples were collected within 2 hours of the night high tide. The
trawls were made towing into the current. Various mesh sizes were tested and
optimum size for crustaceans was 2 mm for the body of the net and a cod end of 1 mm.
A larger mesh size of 2.5 mm for the body of the net was used in a subsequent study
(Young, 1981).
Robertson and Howard (1978) sampled the zooplankton in Zostera and
Heterozostera beds, by using plankton tows of 10 minutes duration. The mesh size was
254 um and the net had a 30 cm mouth diameter. The upper zone of the water column
was sampled by maintaining the top of the net at 10 cm. The volume of water sampled
was measured by a digital flow meter.
For fish, Scott (1981) used a mesh of 1.5 mm and trawls of 1-2 minutes duration
covering a distance of 30-50 m. Sampling was considered sufficient when no
additional species were found.
Robertson and Howard (1978) used a 30 m seine net (1 cm mesh bunt and 1.25
cm mesh wings) to catch large, fast swimming*and mid-water living fish. Smaller
benthic fish were obtained by using a hand laid 10 m (1 mm mesh) seine net. Samples
were taken on the rising and falling tide, by day and by night. Robertson measured the
bias introduced into density estimates through fish avoiding the seine net in the study
area. The results indicate no apparently significant difference in the efficiency of the
net in capturing the major pelagic species, but efficiency values were much lower for
benthic species. Although the efficiency of capture differed, the nettings accurately
reflected the relative abundance of species within each group.
DESCRIPTION OF THE FAUNA
a. INFAUNA
Infauna has been the best studied faunal component of seagrass beds probably
because of the relative ease of collecting quantitative data. Even so, quantitative data
are restricted to temperate beds of Posedonza, Ruppia, Halophila and Zostera.
A recent survey of Poszdonza beds along the New South Wales coast by Collett et
al. (in MS) has shown that a large number of benthic invertebrates occur (363
species) dominated by polychaetes, crustaceans and molluscs. A large number of
species (211 or 54.5%) are restricted to one site and of these 102 (26.9%) are
represented by single individuals. Only two species occur at all sites. No characteristic
Posidonia fauna occurs. Rather, the fauna is determined by the hydrological and
sediment characteristics of a particular bed. Latitudinal gradients seem unimportant.
Analysis of the fauna shows that marine-dominated sites are more similar than
estuarine sites. The number of species and individuals occurring at sites varied greatly
as did community indices. Many of the species occurring in Poszdonza beds also occur
in Zostera beds or in seagrass-free sediments, in the same area.
Other seagrass communities studied quantitatively include those of Tuggerah
Lakes, New South Wales (Powis and Robinson, 1980). In the lakes, Zostera
capricornt, Ruppia spiralis and Halophila ovalis occur typically as discrete
monospecific stands. At only one site, does a mixed community of Zostera and Ruppia
occur. Powis and Robinson found that the sites could be split on the basis of sediment
and associated vegetation, in terms of number of species, and individuals, diversity
and characteristic species. Seagrass communities are generally more diverse than mud
communities. The Zostera sites share some species in common, although the
numerical dominance varies between sites. Ruppia sites have lower numbers of species
than Zostera sites. Mixed communities of Zostera and Ruppza have the highest
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
P. HUTCHINGS 187
number of species and diversity. Halophila beds are intermediate between Ruppia and
Zostera in terms of number of species and diversity.
Beds of Halophila ovalis and H. spinulosa have been extensively studied in
Moreton Bay by Poiner (1980). He found 137 species, of which 51% are polychaetes,
33% crustaceans (including 16% amphipods and 8% decapods) and 11% molluscs
(9% of which are bivalves).
The majority of these studies have shown that a richer infauna is associated with
seagrasses than with bare substrate. In Moreton Bay, one of the beds of Halophila
ovalis, which Poiner (1980) studied, disappeared during the winter, as a result of
storms and strong wave action, and reappeared during the calm, summer months.
Poiner found with an increase in plant growth during the summer, diversity of the
infaunal community increases significantly. The rise is due to an increase in the
number of individuals rather than additional species.
Several reasons have been advanced as to why seagrass communities have richer
benthic faunas than bare sediment. Seagrasses dampen the sediment water interface
by stabilizing sediments, baffling currents and damping wave action (Zieman, 1972;
Orth, 1977). This stabilizing effect of seagrasses could allow increased settlement of
larvae, prevent adults or juveniles from resuspension and reduce disruptive
disturbances by, for instance, nektonic predators, surface scavengers and wave action
(Poiner, 1980). Orth (1977) reported the impact of the blue crab (Callinectes
sapidus) on the infauna of seagrasses. Powis and Robinson (1980) suggest that
seagrass beds may provide a habitat of high structural complexity which, according to
MacArthur (1965), would have a high species diversity (Heck and Wetstone, 1977).
However, the abundance of infauna seems to vary according to the type of
seagrass. Powis and Robinson (1980), as already mentioned, found that Zostera
capricornz has a richer fauna than either Halophila or Ruppia. Hutchings and Recher
(1974) sampled the infauna of Posedonza australis and Z. capricorni and found that
Zostera supports a more diverse community, both in terms of number of individuals
and species than Poszdonza. Powis and Robinson suggest that the well-developed
‘matte’ of Poszdonza restricts the development of an infaunal community. By contrast,
some of the Posidonza beds sampled by Collett et al. (in MS) yielded large numbers of
individuals and species (e.g., Towra Point yielded 148 species with 4,480 individuals
per m’). Powis and Robinson (1980) further suggest that the differences in infaunal
diversity of different seagrasses may be related to the efficiency of individual species in
binding sediment. They suggest that Zostera is more efficient than Ruppza. Orth
(1977) also stresses the stabilizing properties of Zostera marina. Stable substrates
according to Sanders (1969) can support communities of high diversity. Ruppza, with
its poor root system and small blades, is not as efficient as Zostera in either of these
roles and hence has lower diversity communities. Powis and Robinson do not speculate
on Halophila beds which have intermediate values of diversity. This species does not
form dense beds in Tuggerah Lakes.
There is some contradiction in the literature, regarding the endemicity of
seagrass faunas. Kikuchi and Péres (1976) developed the concept of parallelism which
suggests that seagrass beds share many species in common or a ‘species’ is replaced by a
closely related equivalent in a different geographical locality. They give several
examples from the epifauna of Zostera beds in Japanese waters and imply it would be
valid for infauna. The data from Pos¢donza beds along the New South Wales coast do
not support this concept. For Collett e¢ al. (in MS) were not able to define a
characteristic fauna. Each site has a unique assemblage of species. Powis and
Robinson (1980), similarly, did not record any species totally restricted to a particular
species of seagrass in Tuggerah Lakes. Also, in Careel Bay many species occurring in
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
188 FAUNA OF AUSTRALIAN SEAGRASS BEDS
TABLE 1
Species of Polychaetes recorded from nine Zostera capricorni beds in N.S.W.
Harmothoe praeclara
Parahalosydna chrysostichtus
Paralepidonotus ampulliferus
Sigalion ovigerum
Sthenelazs boa
Eumida sanguinea
Gyptis sp.
Phyllodoce novaehollandiae
Sphaerosyllis sublaevis
S. nr. semzverrucosa
Typosyllis varvegata
Australonerets ehlersi
Ceratonereis erythraeensis*
C. mirabilis
Platynerets dumerilu antipoda
Nephtys australienszs
Glycera americana
Marphysa sanguinea
Lumbrinerts latrezlli
Leztoscoloplos bifurcatus
L. normalis
Natneris grubez australis
Phylo felix
Polydora soczalts
Prionospio malmgrenz
Pseudopolydora kempi
Magelona dakini
Chaetopterus variopedatus
Cirriformia filigera
C. tentaculata
Hyboscolex longiseta
Armandia intermedia
Polyophthalmus pictus
Barantolla lepte
Mediomastus californzensts
Notomastus torquatus
Scyphoproctus djiboutiensis
Arenicola bombayensis
Owenia fusiformis
Amphictets dalmatica
Lysilla apheles
L. pacifica
Rhinothelepus lobatus
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
Merimbula
Towra Point
x x
Weeney Bay
mx x
x KK
Bonnet Bay
Sites
Oo
BY)
sj
eee
ah
DO)
ae
O
xX
><
xX
x
xX
xX
>
XG gt XE
xX
XG RX
xX
XK XxX
xX
xX OX
xX
XxX
xX
xX
xX
x
XxX
xX
xX
XOX
XxX
xX XxX
>.<
><
x OX
xX
xX
X
peas
xx x x
Lower Myall River
Wallis Lake
xx x
Wollomba River
Number of sttes
COO OU mT DDT OO i OT Om OR OOOO OO ODT OO OO 8
P. HUTCHINGS 189
TABLE 1 (continued)
Species of Polychaetes recorded from nine Zostera capricorni beds in N.S.W.
Sites
n g sl
2 28
oes ew Stein eas a 5
Sali Oe Wea aging 9 omen cata eS
GO Bs ty, Ce ice Oy ells ea
Bee) cee Gur steel a etc mene mais
Se academe
at ES aes OP\ ie) Seles
Streblosoma acymatum x 1
Pista typha Xx xX 2
Amphiglena pacifica Xx 1
Janua (D.) foraminosa x 1
J. (D.) brasiliensis x 1
Total number of species 1 1 WG as 1s7y 7 9 We 4
Sources of Data:
Careel Bay — Hutchings and Recher (1974) and Hutchings and Rainer (1979)
Towra Point and Weeney Bay — ALS (1978)
Wallis Lake — Hutchings et al. (1978)
Lower Myall River — Weate (1975) ; Atkinson et al. (1981)
Wollomba River — Day (1975)
Bonnet Bay — Branagan et al. (1976)
Tuggerah Lakes — Powis and Robinson (1980)
Merimbula — Hutchings and Murray (in prep.)
*See footnote p. 190.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
190 FAUNA OF AUSTRALIAN SEAGRASS BEDS
the Zostera beds also occur in Postdonza (Hutchings and Recher, 1974). No
characteristic species are associated with the Zostera mueller? and Heterozostera
tasmanica communities in Westernport Bay, Victoria (Littlejohn et al., 1974).
Similarly, the Zostera communities in Gippsland Lakes (Poore et al., 1977) lack
characteristic species. However, Poiner (1980) found 35 species restricted to beds of
Halophila ovalts and H. spinulosa but does not list them.
Even if the concept of Kikuchi and Pérés (1976) is rejected, there is evidence that
similar trophic levels operate within seagrass communities. Collett et al. (in MS)
found that the very different faunal communities in the Poszdonza beds are all
dominated by detritus feeders, either feeding on detritus within the sediment (deposit
feeders) or in suspension (filter feeders). Similarly, the infauna of Zostera muelleri
and Heterozostera tasmanica in Westernport Bay, Victoria (Littlejohn et al., 1974)
are dominated by detritus feeders. Of the 44 species present, 14 are selective deposit
feeders, 10 are non-selective deposit feeders and 3 are filter feeders. Polychaetes
represented 50% of the biovolume.
As polychaetes are often the dominant group of animals in the infauna, I have
extracted from the literature species occurring in Zostera in New South Wales. Many
of these studies were carried out by myself or else I confirmed the identifications so
that consistent names have been given to the species (for several of these species pose
taxonomic problems). The sites in Table 1 have been arranged latitudinally. Zostera
capricorn flourishes in a variety of salinities, from almost fully marine, as in Careel
Bay, to upper estuarine areas such as Wollumba River which flows into Wallis Lake or
Bonnet Bay on the Woronora River, where considerable fluctuations in salinity occur.
Fully marine sites such as Careel Bay have the richest polychaete fauna (37 species)
and upper estuarine areas such as Bonnet Bay and Wollumba River have relatively few
species (3 and 4, respectively) which may be abundant, however. Such data reinforce
the concept that although seagrasses provide some benefits in stabilizing sediment,
factors such as salinity regimes have an overriding effect on the faunal community.
The polychaete fauna varies considerably between sites, and 30 species (62.5%) occur
at one site only; no species is shared by all 9 sites. However, certain families of
polychaetes are commonly present in infaunal communities, for example, all sites
possess at least one species of nereid (Australonerezs ehlersi, Ceratonerezs
erythraeensis*, C. mirabilis or Platynerets-dumerilu antipoda) and all sites except
Bonnet Bay have at least one species of capitellid (Barantolla lepte, Medzomastus
californiensis, Notomastus torquatus, Scyphoproctus djiboutiensis). These two
families are well represented in other seagrass beds (Collett e¢ a/., in MS) and
Hutchings and Turvey (in MS) have collected 10 species of nereids from seagrass beds
in South Australia. Capitellids are non-selective deposit feeders. Nereids are often
assumed to be carnivores, as they have well developed jaws but are probably
scavengers or omnivores.
Fluctuations in seagrass infaunal communities
i. Seasonality
Seagrass beds exhibit strong seasonality of growth, maximum growth occurs
during the spring and summer. In the autumn and winter, much of this growth dies
and is broken off and may be washed out of the seagrass beds. Severe storms may cause
loss of seagrass leaves, and even rip out the seagrass ‘matte’.
*Ceratonereis erythraeensis is a complex of species currently being described by Hutchings and Turvey (in
MS). Ceratonerezs erythraeensis Fauvel probably does not occur in Australia.
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
P. HUTCHINGS 191
Hutchings and Recher (1974) recorded strong seasonality of infauna in both
Posidonia and Zostera beds. Both showed fluctuations in the number of species, with
maximum numbers occurring in June. Numbers of individuals also fluctuated with
maximum numbers in November in Zostera beds and in June in Poszdonza.
Mass settlement of some species occurs. Two polychaetes Scoloplos (S.) simplex
(originally referred to as Haploscoloplos n. sp.) and Ceratonerezs erythraeensts
(originally referred to as Nerezs diversicolor but see footnote here) settle in large
numbers between June and November and, similarly, the mollusc Macoma deltozdalis.
Many species occur sporadically in the seagrass beds (Hutchings and Recher, 1974).
Poiner (1980), working on Halophila communities, measured the community
flux at 2-monthly intervals. Community flux is a measure of the degree to which
species contributions within a particular site (measured as weighted change in
abundances of individual species) are exchanged or given up altogether between
sampling periods. The community flux is low for permanent beds of Halophila,
whereas the flux in beds showing marked seasonal changes, fluctuate widely. In these
beds the number of species remains relatively constant, but an increase in the number
of individuals is strongly correlated with an increase in the density of Halophila.
i. Flooding
Severe floods occurred in Tuggerah Lakes while Powis and Robinson (1980) were
sampling the infauna. Halophila beds were severely reduced. Associated with the rain
was a severe storm with strong winds. This resulted in the devastation of some beds of
Zostera and Ruppra. After the floods limited re-establishment of fauna in muddy
areas occurred; this did not happen in the seagrass areas. Powis and Robinson suggest
that the loss of Halophila roots, resulted in a decrease in habitat availability.
In the lower Myall River and Myall Lakes, New South Wales, widespread
destruction of Zostera occurs periodically, probably as a result of salinity reductions.
The effect on the associated benthic faunas has not been substantiated (Atkinson et
ail, Ns),
b. EPIFAUNA
No quantitative data are available on this community. Qualitative studies have
been carried out in Cockburn Sound, Western Australia. Echinoderms (Marsh and
Devaney, 1978), and molluscs (Wells, 1978, and Wilson et al., 1978) were identified.
With the demise of seagrass beds (Cambridge, 1979) one of the echinoids
Temnopleurus michaelsenz is also declining in numbers. This species was abundant in
the early 70s. Similar studies have been carried out in the upper Spencer Gulf of South
Australia by Shepherd (1974). He found an inverse correlation between the density of
Posidonia and the abundance of filter-feeding molluscs, Penna, Malleus and
Trichomya, the sea urchin Helzoczdaris and the ascidian Polycarpa.
Water movement under the seagrass canopy is about 30% of that above the
canopy. This may explain the relatively small numbers of filter-feeding organisms
occurring in dense seagrass beds.
The bivalve Anadara trapezia recruits successfully only periodically so that within
a seagrass bed one age or size class dominates (Hutchings and Recher, 1974; Dixon,
pers. comm.). Since 1974, there has not been a successful recruitment to Careel Bay,
New South Wales, and widespread mortality (perhaps naturally as the population
ages) is currently occurring (Recher, pers. comm.) .
Cc. SESSILE COMMUNITIES ON THE BLADES OF THE SEAGRASSES
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
192 FAUNA OF AUSTRALIAN SEAGRASS BEDS
The epiphytic fauna of Australian seagrass beds has not been studied as a
separate component.
Anink (1980) reports that the number of epifaunal species so far observed is far
fewer than the number reported settling on artificial substrates.
d. EPIBENTHIC COMMUNITIES
Young (1978) has extensively studied the distribution of one component of
epibenthic communities, namely prawns in Moreton Bay, Queensland. Four species of
penaeid prawn were studied in detail. They utilize all available littoral areas in the
bay, only some of which contain seagrasses. The distribution of prawns differs but is
related to the prevailing salinity and temperature regime and the seagrasses.
The maximum abundances of post-larvae of all four species occur within two
seagrass communities (Zostera capricornt and Halophila ovalis or Z. capricorni,
Halodule uninervis and H. ovalzs). Three of the four species show increase in
abundances in summer which coincides with the increased abundance of Z. capricornz
and H. ovalzs. However, the other species has reduced numbers at this time suggesting
no simple relationships. Young was unable to isolate or attribute the relative
importance of these individual factors.
A similar complex pattern exists among the entire epibenthic community (Young
and Wadley, 1979). The type of sediment and the presence of seagrasses are again
important but the major factor is salinity. Salinity regimes are major determinants in
seagrass distribution, but depth is also important (Young and Kirkman, 1975).
Young and Wadley (1979) were unable to quantify the importance of specific seagrass
beds in determining their associated epibenthic fauna.
In a subsequent study, Young (1981) attempted to eliminate some of these
variables. He selected two adjacent beds of Postdonza australis and Zostera capricornt,
in similar depths of water in Port Hacking, New South Wales. Significant differences
in the species composition and abundances of the epibenthic fauna occur, with
Posidonia being the richer community. Young suggests that the differences in the
fauna of the two seagrasses are controlled by external events leading to variable
recruitment success, although he also suggests that the resources offered by the
seagrasses present a habitat which many species recognize and utilize. Wadley (1981),
working in the same area, showed that temporal heterogeneity of the epibenthos,
although significant, is small compared with spatial heterogeneity. Differences in the
fauna associated with the month and season of sampling show strong interaction with
the type of habitat. The absence of change in the major components of the fauna
during flood and the overall temporal homogeneity in samples indicate relatively
stable communities despite the unpredictable environment. Young (1981) further
suggests that the more diverse community of Poszdonza australis is not significantly
more stable than the Zostera community. This greater diversity of epibenthic fauna is
the reverse of infaunal communities, where the Zostera is more diverse (Hutchings and
Recher, 1974).
Fish
Relatively few studies have been carried out on the fish communities of seagrass
beds. Scott (1981) sampled the fish fauna of Posedonia beds in Geographe Bay,
Western Australia. Nineteen species occur, and the most abundant species is the
odacid Neoodax radiatus (weed whiting) .
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
P. HUTCHINGS 193
Hoese (1978, pers. comm.) has studied the fish communities occurring in
seagrass beds in coastal lagoons in New South Wales. These lagoons are periodically
closed to the sea. The communities fluctuate both in terms of number of individuals
and species. Recruitment appears to be solely from the sea, when the lagoons are
open, rather than from other parts of the lagoon. This is in contrast to Scott’s (1981)
study where the fish complete their life cycle within the seagrass.
Other fish studies by Bell et al. (1978a, b), Conacher et al. (1979) and Littlejohn
et al. (1974) have been related to investigating the feeding patterns of the fish. This
will be discussed in the energy flow section.
Zooplankton
Holoplanktonic calanoid copepods and meroplanktonic decapod larvae are
numerically dominant in the water column during the day in Zostera beds in
Westernport Bay, Victoria (Robertson and Howard, 1978). However, the number of
such animals is significantly lower than that collected at night. This pattern of low
catches near the surface during the day and high numbers at night has been recorded
on a number of previous occasions (Howard, unpublished data). These obligate
planktons have also been taken in planktonic trawls made close to the substrate during
the day. Thus estimates of zooplankton samples taken near the surface underestimate
the true density of these organisms throughout the whole water column during a 24-
hour period.
Amphipods are benthic during the day, at night they form a conspicuous
component of the plankton.
Ostracods, which are infaunal by day also enter the water column at night but at
lower densities than amphipods.
ENERGY FLOWS WITHIN THE SEAGRASS COMMUNITIES
Kirkman and Reid (1979) have constructed a carbon budget for Poszdonza
australis in Port Hacking, near Sydney. They measured biomass, calculated grazing
rates by herbivores, estimated the amount of floating seagrass detached from the beds
and collected the sinking Poszdonza detritus below the seagrass beds.
Small seasonal variations in biomass occur. Three species of Monacanthidae
(leatherjackets), Monacanthus chinensis, Meuchenza freycineti and M. trachylepszs
consume large amounts of Poszdonza. Bell et al. (1978b) calculated that adult M.
chinensis have a diet consisting of 41.7% P. australis by estimated volume. However,
looking at the entire Poszdonia beds, herbivores do not consume an appreciable
amount of attached Poszdonza (Conacher et al., 1979).
The amount of floating Poszdonza is related to winds and, unlike Zostera, is not
washed up onto beaches in New South Wales. However, in South Australia large beds
accumulate on beaches adjacent to seagrass beds. Pos¢donza occurs typically in greater
depths than Zostera. Leaves that are added to the organic detrital cover of sediments
make up the majority of particulate organic C lost from the seagrass beds. Only
heavily epiphytized leaves sink. Between 27-71% of the biomass collected in the traps
is epibiota.
Dissolved organic carbon accounted for the greatest loss of organic C from the
system.
A less extensive study of the Zostera beds in Port Hacking by Kirkman e¢ al.
(1982) has shown that the growth of Zostera is 3-4 times that of Poszdonza in the same
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
194 FAUNA OF AUSTRALIAN SEAGRASS BEDS
environment. The production figures are for Zostera 0.56-1.82 g m™ day while P.
australzs shows a production rate of 0.57-1.44 g m7” day" through the year.
It seems likely that a similar breakdown of the C budget occurs in Zostera beds as
Kirkman and Reid (1979) have described for Poszdonza in the same environment.
Larkum (1981) has estimated the percentage contribution of each of the primary
producers in the estuarine ecosystem of Botany Bay. Although seagrass beds do not
occur as extensive beds, they contribute 25% to the total budget of the bay. This
percentage includes the contribution made by the epiphytic algae which occur on the
seagrass blades.
Assuming that Kirkman and Reid’s (1979) data from Port Hacking are of
general applicability, the important contribution of seagrasses to the estuarine
ecosystem is the continual supply of leaves to the organic detrital cover of the
sediments. Attiwill and Clough (1974) have also stressed the importance of this
production of organic matter by the seagrass beds in Westernport Bay, Victoria.
Little work has been done on the breakdown or shredding of this continual supply
of leaves. Brand (1977) has experimentally investigated the production of detritus
from the breakdown of Zostera by the amphipods Parhyalella spp. This was to
quantify part of the model of the C budget proposed by Brand et al. (1974) for
estuarine ecosystems. The amphipods ingest the epiphytes on the blades of the Zostera.
Brand does not provide any evidence that the amphipods actually obtain nutrients
from the plant matter but implies that nutrition is obtained mainly from the attached
biota. The amphipods produce nitrogen-enriched faecal pellets which facilitate
further breakdown of the Zostera fragments. Seagrass leaf tissue is extremely refractile
(Godshalk and Wetzel, 1978) and the decay of the structural carbohydrates may be
the rate limiting step in the transfer of energy through seagrass ecosystems (Harrison
and Mann, 1975). Animals which can utilize these fragments appear to be mainly
amphipods (Robertson and Mann, 1980), although some herbivorous polychaetes
may be involved. Overseas, attempts have been made to trace the origin of detritus in
the diets of estuarine animals by measuring the ratio of '?C:'*C. These ratios differ
sufficiently in seagrasses and phytoplankton to allow the source of the detritus to be
traced (Fry and Parker, 1979; McConnaughey and McRoy, 1979; Thayer et al.,
1978). As mentioned earlier, few fish of seagrass beds in Australia are herbivores. The
leatherjackets do obtain some nutrients from ingested P. australis fragments
(Conacher et al., 1979). Most of the fish occurring in seagrass are carnivores
(Littlejohn et al., 1974; Scott, 1981) feeding on the detritus-feeding crabs, molluscs
and polychaetes (Robertson, 1980). Although Robertson and Howard (1978) have
documented the importance of zooplankton for several fish species, fish diets may shift
during the year as prey species fluctuate (Littlejohn et al., 1974).
The importance of detritus feeders (Tenore, 1977) which dominate the benthos
is related to the further breakdown of the plant matter by mechanical abrasion as it
passes through the guts of the detritus feeders (Fenchel, 1970, 1977). This increases
the surface area to volume ratio, and hence the bacterial and algal populations on the
particles (Newell, 1965). Many of these detritus feeders are probably obtaining
nutrients from the associated flora and fauna rather than from the seagrasses. During
the production of detritus some organic compounds are produced which can be taken
up directly by the seagrasses themselves (Bulthuis and Woelkerling, 1981) so that a
close relationship between the infauna and seagrasses occurs.
Where they occur, vertebrate grazers of seagrass beds may have considerable
local impact on the beds. Dugong dugon, a marine mammal, occurs in large herds in
coastal waters of northern Australia. The dugong feeds exclusively on seagrasses
(Heinsohn and Birch, 1972). In the Townsville region they feed selectively on
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
P. HUTCHINGS 195
Diplanthera and Cymodoce. Two other common seagrass species in the region,
Enhalus acorovdes (L.f.) Royle and Syringodzum isoetifolium, are not eaten. Dugongs
eat the entire plant (leaves, rhizomes and roots) , and ingest some animals and algae.
They graze most heavily in beds of low density (biomass 10-30 g dry matter/m7’) ,
producing trails of a depth of 3-5 cm. They remove on average 63% of the seagrass
from the trails with a maximum of 86% (Heinsohn et al., 1977). Two captive
dugongs ate 50-55 kg of seagrass per day (equivalent to 2 tonnes dry weight/annum)
(Jones, 1967). During feeding some seagrass is damaged although not ingested.
Assuming a grazing efficiency of 63% and a seagrass community biomass of 94 kg dry
matter/m* (Heinsohn, unpublished data from Cleveland Bay, Townsville) 3.5 ha is
required to support a dugong for a year. The area over which dugongs graze is not
known but should be if the impact of their grazing on seagrass beds is to be assessed.
Turtles also feed on seagrasses, but not exclusively. The green turtle Chelonza
mydas feeds on marine algae and seagrasses, including species of Zostera, Enhalus,
Thallasstna, Postdonia and Halodule. Loggerhead turtles Caretta caretta, also eat
seagrasses, but are primarily carnivores. Similarly, Hawksbill turtles Eretmochelys
zmobricata, eat seagrasses but mainly as juveniles (Rebel, 1974).
Wading birds are important temporary residents during low tide. Water fowl
such as black swans, Cygnus atratus, are estimated to eat 230 tonnes of Ruppia and
the freshwater Potamogeton a year in the Blackwood River estuary in Western
Australia (Hodgkin, 1978). Delroy (1974) calculated that 90% of food of black ducks
(Anas supercil), musk duck (Bzzzura lobata) and grey teal (Anas gibberifrons)
consisted of tubers and seeds of Ruppia and Lamprothaminium in the Coorong, South
Australia. They feed by pulling plants to the surface, thus damaging more than they
eat. They feed only during the summer on seagrases in this estuary. Ferguson Wood
(1959) suggested that black swans caused great damage to Zostera capricorni beds.
Seagrass beds are also used by a variety of water birds including pelicans
(Pelecanus conspicillatus), cormorants, etc. Pelicans and cormorants feed on
schooling fish within the seagrass beds. For birds such as herons (Ardeidae) which
stalk their prey, the presence or absence of seagrass beds probably determines the
number of individuals in an area. In dense seagrass beds, water clarity is improved,
more sedentary fish are present, giving herons ample opportunity to feed during low
tide. In areas of sparse or no beds, the water is more turbid, fewer sedentary species of
fish are present and herons must expend far more energy in catching fish. Such areas
will have fewer herons than areas with dense seagrass beds. Recher (pers. comm.)
points out that although herons and egrets are predominantly fish eaters, in marine
habitats crustaceans are an important part of their diet.
Migrant waders, are less dependent on seagrass beds. They are limited to low tide
and will often feed in non-grass situations. They feed on molluscs, annelids and
crustaceans, although a particular species may be highly selective. For example, the
knot feeds on surface-living molluscs whereas the bartailed godwit Limosa lapponica
feeds preferentially on polychaetes and the eastern curlew Numenius madagascariensis
on Callzanassa sp. (Recher, pers. comm.).
DISCUSSION
Concluding this review, I would like to consider the direction of future studies of
seagrass communities. Recent trends in ecological studies of these communities
overseas have been to manipulate the system experimentally.
Orth (1977) has been able to quantify the importance of seagrass beds in
Proc. Linn. Soc. N.S.W., 106 (2), (1981) 1982
196 FAUNA OF AUSTRALIAN SEAGRASS BEDS
stabilizing sediments and the production of organic material by artificially removing
Zostera. He could then compare intact Zostera with clipped beds. Such methods could
be extended to quantify the relative importance of different species of seagrasses in
Australia.
Other experiments have tested the hypothesis that seagrass provides shelter from
predation, by caging experiments (Heck and Thoman, 1981). Such information is
essential if the much-quoted statement that seagrasses act as nursery grounds and
provide shelter and protection is to be substantiated. How do different seagrasses
compare? Are some species more efficient than others? Young and Young (1978) have
experimentally investigated the factors important in regulating species densities and
diversities in seagrass associated benthos.
Such studies urgently need to be conducted here in Australia: for this
experimental approach seems to be a productive way of determining the factors
important in maintenance of seagrass communities. Unless we know these factors, the
management of seagrass communities cannot be soundly based. We also need more
integrated studies like those of Kirkman and Reid (1979) but these should be
extended to include the faunal components. Rates of turn-over of the dominant
animals must be determined.
The data from Australian seagrass communities clearly show the importance of
seagrasses. Some of the primary productivity studies indicate the variations between
sites (Larkum, 1981). Are similar fluctuations occurring in the animal communities?
Larkum (1981) has suggested that these variations are sometimes caused by urban
stress. Unless we understand the entire community we will not recognize the early
symptoms of a community under stress, and thus be able to take remedial action. Loss
of seagrasses will have serious impacts on the fishing, oyster and prawning industries
(AMSA, 1977; Young, 1978).
Oil spills and the use of oil dispersants and other types of industrial pollution
severely affect the growth of seagrass beds (Larkum, 1976b) and hence affect the
associated fauna (Jacobs, 1980). The number of oil spills and incidents of industrial
pollution are increasing and unless checked will have major effects on the seagrass
ecosystems and subsequently on estuarine and shallow coastal ecosystems.
ACKNOWLEDGEMENTS
I should like to thank Drs Arnold, Collett, Hodgkin, Rainer and Young and Mss
Marsh and Wadley for providing unpublished material and reports, or papers in press.
Drs Larkum, Rainer and Recher criticized an initial draft for which I am most
grateful. The paper also benefited from the discussions which occurred after the
presentation of this paper at the Seagrass Workshop held at Sydney University,
February, 1980. Mr Glasby and Ms Handley helped in the compilation of the
bibliography.
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PROCEEDINGS of LINNEAN SOCIETY OF NEW SOUTH WALES
VOLUME 106
Issued October 27, 1982
CONTENTS: |
NUMBER 1
1 By JONSELE
Linnaeus and his circumnavigating Apostles
21. B.A.FOSTER
Two new intertidal balanoid Barnacles from eastern Australia
33. D.K.McALPINE and D.S. KENT
Systematics of Tapeigaster (Diptera: Heleomyzidae) with Notes on Biology and larval
Morphology
59 M.A.SCHNEIDER
A comparative morphological Study of the reproductive Systems of some poses of
Tapeigaster Macquart (Diptera: Heleomyzidae)
67 D.E.SHAW, B.K. CANTRELL and K. J. HOUSTON
Neurochaeta inversa McAlpine (Diptera: Neurochaetidae) and Seed Set in Alocasia
macrorrhiza (L.) G. Don (Araceae) in southeast Queensland
83. P.A. HUTCHINGS and H. F. RECHER
The Fauna of Australian Mangroves
123 Notes and Discussion
~R.J. FORD
A. W. H. Humphrey, His Majesty’s Mineralogist in New South Wales. 1803-12 = A
Comment
NUMBER 2
127 C.McA. POWELL, G. NEEF, D. CRANE, P. A. JELL and |. G. PERCIVAL
Significance of Late Cambrian (Idamean) Fossils:in the Cupala Creek Formation,
northwestern New South Wales
151. D.T. ANDERSON
Origins and Relationships among the Animal Phyla
167 C.J.JENKINS
Late Pridolian Graptolites from the Elmside Formation near Yass, New South Wales
173. C.J. JENKINS
Darriwilian (Middle Ordovician) Graptolites from the Monaro Trough Sequence east of
Braidwood, New South Wales
181 P. A. HUTCHINGS
The Fauna of Australian Seagrass Beds
LINNEAN SOCIETY OF NEW SOUTH WALES
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VOLUME 106
NUMBER 3
Stability, Depletion and Restoration of
Seagrass Beds*
’ A. W. D. LARKUM and R. J. WEST
LARKUM, A. W. D., & WEST, R. J. Stability, depletion and restoration of seagrass
beds. Proc. Linn. Soc. N.S.W. 106 (3), (1982) 1983: 201-212.
The hypothesis that seagrasses are well adapted to the environment in which
they live is questioned. Evidence for instability in seagrass beds is documented. Good
evidence for depletion of seagrass beds from natural causes exists. In addition it is
clear that man-induced depletion has been very severe in many populated coastal
areas. The causes of instability in seagrass beds need to be understood both as regards
the prevention of further depletion and the restoration of seagrasses in depleted areas.
Several techniques have been used successfully for the restoration of seagrasses in
small scale experiments but these are expensive and labour-intensive.
A. W. D. Larkum, School of Biological Sciences, University of Sydney, Sydney, Australia 2006,
and R. J. West, N.S.W. State Fisheries, 211 Kent Street, Sydney, Australia 2000 (formerly
School of Biological Sciences, University of Sydney); manuscript received 19 May 1981, accepted
for publication in revised form 16 December 1981.
INTRODUCTION
Seagrasses are angiosperms with a rooted rhizome system which limits their
distribution generally to unconsolidated sedimentary substrates in relatively calm
waters (Thalassodendron pychyrizum which grows on rocky reefs in Western Australia
(Cambridge, 1975) is an exception to this rule). This distribution, in turn, has a special
significance for both depletion and restoration, for it means that the favoured sites for
seagrasses — estuaries and sheltered embayments — are also the favoured sites for
human settlement and activity. As a result there has in the past been much depletion of
seagrass beds (Phillips, 1978). However it should be pointed out that it is only recently
that man has had any real impact on seagrasses and it is really only the current life
styles of world civilizations that are inimical to co-existence at such sites of seagrasses
and people.
With the growing awareness of the biology of seagrasses has also come an
awareness of the economic importance of seagrasses in a variety of ways (see e.g.
Thayer et al., 1976; Phillips, 1978). And out of this have come a number of attempts to
restore seagrasses to depleted areas (see Table 1).
However, perhaps the best place to start is on the subject of the stability of
seagrass ecosystems in their natural state, since our ideas on the ability of seagrass
communities to withstand man-induced impacts depend on this crucial point.
STABILITY OF SEAGRASS BEDS
Introduction
Seagrasses it has been claimed are well adapted to the environment in which they
live (e.g. den Hartog, 1979). The supporting evidence for this statement is not com-
pletely convincing. Resolution of this matter is very important since on it rests the
degree of stability which may be expected from ‘natural’ seagrass beds and on this in
turn rest any assumptions that may be made as to the cause of depletions of beds
subjected to man-induced impacts.
* A paper presented at the Australian Seagrass Workshop, University of Sydney, 1981.
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
202 STABILITY, DEPLETION AND RESTORATION OF SEAGRASS BEDS
Seagrasses are certainly adapted to their environment otherwise they would not be
there. The real question to ask is ‘are they well adapted?’ This is a difficult question to
answer. It could be argued that the lack of speciation within seagrasses is a sign of poor
fitness. Seagrasses probably evolved early in the evolution of angiosperms; but whereas
there are approximately 300,000 species of flowering plants there are only 50 odd
species of seagrasses. The counter-argument would be that seagrasses are so well
adapted to their environment that further adaptation and speciation has been un-
necessary. However this sort of argument is unsatisfactory in the light of the vast
speciation in terrestrial grasses, freshwater macrophytes and reef fish (Ehrlich, 1975) to
take but a few examples. Furthermore the recent documentation of speciation in
Posidonia (Cambridge and Kuo, 1979) in south western Australia indicates that
speciation has taken place in an area which appears to be ideal for the growth of
seagrasses.
A number of inherent weaknesses may exist in the strategy of a submersed marine
angiosperm. These weaknesses include: —
1) sexual propagation
2) vegetative propagation
3) occlusion of leaves by epiphytes
4) internal aeration
5) salinity control and turgor regulation
6) susceptibility to microbial attack
7) requirement for moderately high light environments.
At present there are few hard facts about any of these in relation to the ‘fitness’ of
seagrasses. Johnston (1979) has made the interesting suggestion that the high leaf
productivity of seagrasses in comparison with other plants is a result of the need to
provide new photosynthetic tissue as the older parts of the leaves become overgrown by
epiphytes and epizoans. Despite the fact that seagrass leaf productivity is not ex-
ceptional (terrestrial tropical grasses such as sugar cane and napier grass have a
productivity several fold higher — see Boardman and Larkum, 1974), the suggestion
has some merit and if it is true leads to the constraint that seagrasses have less energy to
devote to sexual and vegetative propagative mechanisms. As regards salinity control,
Tyerman (1979) has shown that the leaf meristem tissue is hypotonic to seawater and
further experiments (unpublished) have shown that exposure of the meristem to
seawater leads to its death. Thus mechanical damage of the sheath due to wave action,
disease, or animal attack (and of course human activities such as bottom trawls or army
‘ducks’) can have serious consequences for seagrasses. Internal aeration does not
apparently pose any problems for seagrasses; freshwater macrophytes which in terms
of speciation are much more successful than seagrasses seem to cope with this con-
straint. Nevertheless the necessity of having rhizomes and roots in an anaerobic en-
vironment must place constraints on the growth and productivity of such a plant and
must make it more susceptible to microbial attack. Presumably the high concentration
of phenolics and other aromatic compounds in the roots and rhizomes of seagrasses
(Cariello and Zanetti, 1979) is an antibacterial response.
Evidence for instability in seagrass beds
The best example of instability in seagrass beds is the ‘wasting disease’ of Zostera
marina which caused a near catastrophic decline in stocks of this seagrass in the early
1930s. This example is dealt with further in the next section. However in summary it
appears that a slight climatic rise in temperature may have set off the demise
(Rasmussen, 1977). Apart from this example there is a great deal of evidence to
support recent cyclical changes in stocks of Zostera marina (Orth, 1976; den Hartog
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
A.W.D. LARKUMANDR. J. WEST 203
1979). In fact McRoy and Bridges (1974) suggested that the ‘wasting-disease’ of the
1930s was an extreme cycle of a rhythmic oscillation. Evidence for cyclical changes in
other seagrasses is not available but this probably reflects the absence of investigations
rather than the absence of the phenomenon.
Further evidence for instability can be found in a few studies of various transient
phenomena, usually induced by wave-action. Patriquin (1975) has described the
‘migration’ of blowouts in beds of Thalassia testudinum in the West Indies. Storm-
generated, crescentic holes in the beds were observed to move shorewards at the rate of
ca 0.5 cm/day. It was estimated that any one point would be ‘recurrently eroded and
restabilized at intervals of the order of 5-15 years’. Syringodium filiforme was involved in
the recolonization of the blowouts on the leeward slope. Similar blowouts have been
described in Western Australia (Cambridge, 1975) in P. australis beds (there Amphibolis
antarctica is involved in the recolonization). In both examples the bed is in a dynamic
equilibrium i.e. the seagrass system is able to cope with the factor causing the in-
stability. However it does indicate a rather delicate balance in the system: in both
examples the dominant seagrass is absent or very much depleted in more exposed sites.
Thus as pointed out by den Hartog (1979) we generally see today the end result of past
Fig. 1. Rings in Zostera capricorni bed in Botany Bay. Scale bar represents 20 m.
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
204 STABILITY, DEPLETION AND RESTORATION OF SEAGRASS BEDS
evolutionary and successional events and assume that the systems we see are stable.
This does not mean however that such events are not going on but they will un-
doubtedly be more difficult to detect.
Recently we have detected what appears to be a cyclical event in the growth of
Zostera capricornt (Larkum et al., in prep.). Rings of Z. capricornt were observed by
means of high-resolution aerial photographs of the northern shore of Botany Bay (Fig.
1). The rings occurred in a shallow region which previously had been an area of largely
bare, shifting sand. However after the establishment of the armoured wall for Port
Botany, the area became stabilized and wave action was almost eliminated. Under
these conditions Z. capricorni recolonized most of the area within a few years. It was
under these conditions that the rings developed. As can be seen the middle of the rings
was bare with only loose sand. Inspection of the rings revealed no apparent cause for
the phenomenon. A possible cause for the die-back in the centres could be the depletion
of an essential mineral. However this seems unlikely. Another explanation could be
that apical meristems, which are critical for seagrass production (Tomlinson, 1974),
grow initially in an outward radial direction; eventually the oldest shoots on the
rhizomes die and cannot be replaced unless seedlings establish in the centre region or
unless one ring grows into another. If this explanation is correct then it will be a further
example of the ‘apical meristem dependence’ of seagrasses and illustrates another
possible weakness in seagrass systems. It may well be that the inability of seagrasses to
generate secondary apical meristems results in the slow regrowth of eroded areas
(West, 1980).
The causes of natural depletion of Seagrass Beds
Research into the natural depletion of seagrass beds has not been extensive in the
past and the cause(s) of depletion in any one instance has never been fully established.
A number of possible causes have been put forward including disease (Rasmussen,
1977), climatic changes (Rasmussen, 1977; Orth, 1976), natural cycles (McRoy and
Bridges, 1974) sediment movement (Kirkman, 1978), salinity changes (Orth, 1976),
sea level changes (den Hartog, 1977) and faunal influences (Orth, 1976; Ferguson
Wood, 1959).
The most famous example of natural depletion of seagrasses is the ‘wasting
disease’ of Zostera marina which resulted in a nearly catastrophic decline in the stocks of
this seagrass on both sides of the Atlantic in the early 1930s. Although it is generally
agreed that dead plants were attacked by a fungus (Ophzobolus halimus) and a slime mold
(Labyrinthula macrocystis) Rasmussen (1977) has put forward good arguments to suggest
that the real cause of the ‘disease’ was a rise in sea level temperatures in the early 1930s
and that the weakened plants were then susceptible to attack by the fungal and slime
mold pathogens. Rasmussen (1977) also suggested that seed germination was inhibited
at the higher temperatures and that plants growing in brackish water were immune to
the ‘disease’. Orth (1976) documented a 70% decline in Zostera marina in Chesapeake
Bay (U.S.A.) from 1972 to 1974 and a virtual disappearance in 1975. This period was
accompanied by a rise in average temperatures and a marked rise in water tem-
peratures (Orth, 1976, fig. 7).
If the arguments of Rasmussen are accepted then the growth of Z. marina must be
seen as a finely balanced process in which temperature, salinity and susceptibility to
pathogen attack are finely balanced.
There are two examples of apparently natural depletion of seagrasses in Australia.
One is at Corner Inlet, Victoria, which is the only site in continental Victoria where
Postdonia australis occurs. Large beds have existed there in the past, as evidenced by
fibre remains, but in recent years the extent of the beds has decreased (Poore, 1978).
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
A.W. D.LARKUM ANDR. J. WEST 205
Sedimentation in the region has caused a general rise of the seabed in the region and
this in turn may have altered the salinity regime of the area (P. australis is a strictly
stenoholine species). However the exact cause of the decline is not known. One might
more profitably enquire as to why P. australis is absent from the remainder of the coast
of Victoria but occurs at Eden in N.S.W. and after Port MacDonnel in South
Australia.
The second example is the depletion of Zostera capricorni in northern areas of
Moreton Bay (Kirkman, 1978). This depletion seems to have occurred as a result of
sand inundation at a rate faster than the upward growth of the plants. At the same time
adjacent healthy beds were subject to increased grazing which contributed to the
overall depletion.
TABLE 1
Man-induced impacts affecting seagrass communities
Impact Seagrass community Place Effect Reference
Turbidity Posidonia oceanica S. France Large-scale Meinesz &
associated depletion Laurent, 1976
with at lower
eutroplication limit
Posidonza australis Cockburn Large-scale McComb ef al. ,
Sound, W.A. depletion 1981
Posidonia australis Botany Bay Loss of Larkum, 1976b.
deeper beds
Thalassia testudinum Florida Large-scale Taylor et al. ,
depletion 1973
Thalassia testudinum Virgin Local Dong et al. ,
Islands depletions 1972
Nichols e¢ al. ,
1972
Zostera capricorni/ Tuggerah Loss of Higginson, 1971
Ruppia spiralis Lakes, NSW deeper beds
Turbidity Thalassia testuainum Virgin Depletions Van Eepoel et al. ,
associated Islands, 1971
with West Indies
dredging
Thalassia testudinum St. Thomas, Depletions Grigg, 1970
West Indies
Hydraulic Thalassia testudinum/ Tampa and Extensive Godcharles,
clam dredge Halodule beaudette Tarpen losses 1971
Syringodium filiforme Springs,
Florida
Dredging and Thalassia testudinum Boca Ciega Local Taylor &
filling Bay, Florida depletions Saloman, 1968
Thalassia testudinum Redfish Bay, Local Odum, 1963
Texas depletions
Sewage Thalassia testudinum Biscayne Bay, Local McNulty, 1970
effluent Florida depletions
Thalassia testudinum Florida Local Hammer, 1972
depletions
Posidonia oceanica S. France Large-scale Peres &
depletions Pickard, 1974
Posidonta australis Adelaide Severe local Shepherd, 1970
S.A. depletion
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
206 STABILITY, DEPLETION AND RESTORATION OF SEAGRASS BEDS
Hot water Thalassia testudinum Florida Large Zieman, 1976
effluents depletion
Florida Large Thorhaug et al. ,
depletion 1978
Salinity Thalassia testudinum Florida Loss of McMillan &
changes Halodule beaudetter leaves and Moseley, 1967
Syringodium filtforme some
depletion
Thalassia testudinum Florida Die-back Zieman, 1975
but recovery
Overgrazing Thalassia testudinum Florida Local Camp et al. ,
by Lytechinus depletion 1972
variegatus
Oil pollution Thalassia testudinum Puerto Local Diaz-Piferrer,
Rico losses 1962
Phyllospadix torreyt Santa Local Foster et al.,
Barbara depletion 1971
Physical Thalassia testudinum Florida Small-scale Zieman, 1976
disturbance losses but
by boats spreading
Industrial Posidonia australis Cockburn Local Cambridge, 1975
effluents Sound losses
MAN-INDUCED DEPLETION OF SEAGRASS BEDS
There are a great number of examples of man-induced depletions of seagrass beds
(Table 1) and there are a great number of possible or known causes amongst which are
the following: —
1) turbidity increases associated with dredging,
2) turbidity increases associated with industrial or urban influences,
3) turbidity increases associated with eutrophication,
4) toxic chemicals,
5) hot water effluents,
6) oil spills,
7) activities of commercial fishermen using bottom trawls,
8) changes in salinity,
9) sewage.
Much of the evidence on these factors comes from overseas studies as can be seen
from Table 1, and much of it is less than fully convincing. It relies on field observations
and circumstantial evidence. Many of the sites where depletion occurred were subject
to multiple impacts and it may not be justified to single out one factor. Clearly careful
experiments with proper controls are needed.
In Australia there are a number of published reports on man-induced depletion of
seagrass beds (Shepherd, 1970; Higginson, 1971; Cambridge, 1975; Larkum, 1976 a,
b; Shepherd and Van der Borch, 1977). In no case, so far has the cause been
established categorically except for the mechanical removal of P. australis beds in the
Spencer Gulf, South Australia. In fact in all of the other examples multiple impacts
existed. The most clear-cut case would seem to be the disappearance of P. australis near
sewage outfalls off the coast of Adelaide (Shepherd, 1970) yet even this has been
disputed (Steffensen, unpublished). It would seem that transplant experiments with
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
A.W.D.LARKUM ANDR. J. WEST
207
proper controls should be able to provide a definitive answer. In Cockburn Sound
(McComb etal., 1981), in Botany Bay (Larkum, 1976a, 1976b and unpublished) and in
the Tuggerah Lakes increased turbidity would seem to be the most important factor. In
Botany Bay wave-action is also a clear-cut factor at some sites (Larkum, 1976b).
However at all these places a number of impacts are present. In the Tuggerah Lakes
some depletion is almost certainly due to hot-water effluents from electric power
stations (Weiner, personal communication).
Poorly documented evidence exists also for the following:
1) depletion due to oil and dispersants in Botany Bay and,
ii) depletion due to hot water effluent in Botany Bay (Bunnerong Power Station).
Depletion of seagrasses has almost certainly occurred in Port Phillip, Sydney
Method
Plugs
Plugs
Plugs
Plug
Plugs
Plugs
Plugs &
Turfs
Turfs
Turfs
Turfs
Turions
Turions
Turions
Seeds
Seeds
Seeds
Worker and
Publication
Date
Kelly et al.
(1971)
Breedveld
(1976)
Breedveld
(1975)
Kirkman
(1976)
Phillips
(1974)
Phillips
(1974)
Phillips
(1980)
Ranwell et al.
(1974)
Ranwell et al.
(1974)
Larkum
(1976)
Kelly et al.
(1971)
Eleuterius
(1975)
Phillips
(1974)
Phillips
(1974)
Thorhaug
(1974)
Thorhaug &
Hixon (1975)
TABLE 2
Transplantation of Seagrasses by Various Workers using Various Techniques
(Adapted from Thorhaug and Austin, 1976)
NAA = Naphthylacetic acid
Place
Anchoring
Method
Tampa Bay,
Florida
Tampa,
Florida
Tampa,
Florida
Moreton Bay,
Australia
Tampa Bay,
Florida
Tampa Bay,
Florida
Redfish Bay,
Texas
Norfolk,
England
Norfolk,
England
Botany Bay,
Australia
Tampa Bay,
Florida
Biloxi,
Mississippi
Whidbey
Isle,
Washington
Friday Harbor,
Washington
South Biscayne
Bay, Florida
North Biscayne
Bay, Florida
Tin can or Bag
30 cm substrate
into hole
30 cm substrate
into hole
Dug holes
Buried with
soil
Buried with
soil
Buried with
soil
Spaded into
hole
Spaded into
hole
Dug holes
1. Construction
rods
2. Brick pipes
Wire-mesh
anchor and
Construction
rod
Iron pipes and
trenches
Iron rods and
trenches
Plastic
Plastic peat-
pots
Chemical Dimensions of Success
Additive Transplant
NAA 20 cm? Control 40%
Exper. 15%
5% NAA 30cm 0-100%
5% Root Deep post-hole
dip digger
5% NAA 30cm 100%
5% Root Deep post-hole
dip digger
None
None 10 cm? Some
None 10cm? None
None 60 cm” 80-100%
None ca22 x 15 Varied
x 10cm deep
None ca22 x 10 100% year 1
x 15cm deep 35% year 2
None 50 x 50cm 50-100 %
10% NAA Short shoot 18% in some
and rhizome
None 45 x 45cm 3%
5%
None
None Short shoot Dependent
and rhizome on depth
portion 100% to
none
None Single seeds None
10% NAA Single seeds 80%
10% NAA Single 15%-55%
Species
Thalassia
testudinum
Thalassia
testudinum
Syringodium
filiforme
Zostera
capricornt
Halodule
wright
Thalassia
testudinum
T. testudinum
A. wright -
Zostera
noltir
Zostera nolti
Z. marina
Zostera
capricornt &
Posidonia
australis
Thalassia
testudinum
T. testudinum
H. wright
Cymodocea
manatorum
Zostera
marina
Zostera
marina
Thalassia
testudinum
Thalassia
testudinum
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
208 STABILITY, DEPLETION AND RESTORATION OF SEAGRASS BEDS
Harbour (and the Parramatta River) and the Brisbane River, but good documentation
of these events and their causes is lacking.
It is worth noting in passing that there has been no published work anywhere in
the world on the effect of toxic wastes such as heavy metals and chlorinated
hydrocarbons on seagrasses or seagrass communities.
Considering the economic importance attached to seagrass beds (see e.g. Thayer
et al., 1976; and Phillips, 1978) it is surprising that more public money is not being
spent on investigating the causes of the various depletions of seagrasses in Australia.
The lack of funding can partly be seen as a result of ignorance and the great expanse of
completely untouched seagrass beds in southern and western Australia.
RESTORATION OF SEAGRASS BEDS
Introduction
Restoration of seagrass beds does occur naturally at times, as for instance after the
‘wasting disease’ of the 1930s. However artificial restoration is an important
development that has come about in recent years mainly as a result of man-induced
depletions of seagrass beds. For instance in Florida (U.S.A.) real estate development
involving dredging of Thalassia testudinum meadows resulted in large scale depletion of
this seagrass in the 1960s and had the Florida State Board of Conservation setting up a
programme to restore this seagrass to several areas (Phillips, 1974). Also in Florida,
hot water effluents led to damage of Thalassia beds and restoration by transplanting was
effected (Thorhaug and Roessler, 1976), In fact the history of transplants goes all the
way back to the ‘wasting disease’ period of the 1930s when rather ineffectual attempts
were made to revegetate areas with Zostera marina (see Phillips, 1974).
Table 2 lists all published attempts to date to transplant seagrasses.
Techniques
a) Vegetative
Two basic techniques exist. They are: —
al) transplantation of seagrass plants without accompanying sediment. The
plants (including rhizomes and several shoots) are replanted in sediment of
the new site (Phillips, 1974; Larkum, 1976a),
a2) transplantation of seagrass turfs, i.e. plants are removed zn situ in un-
disturbed sediment placed into trays or bags and relocated into excavated
holes at the new site (Ranwell et al., 1974; Fuss and Kelly, 1969; Breed-
veld, 1975). Anchoring may be a problem. Dislodgement may occur as a
result of storms or through the activities of fishermen or prawners. A
number of devices have been tried. Kelly e al. (1971) mention cans,
sacking polyethylene, pipes, construction rods and breeze bricks. To this
list can be added steel mesh for reinforcing concrete (Larkum, 1976a) and
plastic netting (Ranwell et al., 1974).
b) Seedlings. This method has been pioneered with success by Thorhaug (1974,
1976, 1979).
Results and Discussion
Transplanting of seagrasses by the vegetative method has in most cases met with
reasonably good initial success. A great deal depends on the species of seagrass.
Pioneer and rapidly growing species may quickly spread out and cover a large area at
the new site, e.g. Zostera nolti: (Ranwell et al., 1974), Zostera marina (Phillips, 1974). In
this way an area may be ‘seeded’ with the minimum of work and money. On ihe other
hand with ‘climax’ species such as Thalassia testudinum and Posidonia spp. growth rate
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
A.W.D. LARKUM ANDR. J. WEST 209
may be much slower and replanting might mean the ‘sod by sod’ coverage described by
Thorhaug (1979). In fact Kelly et a/. (1971) note that regression may occur at trans-
plant sites because of death of older shoots and lack of replacement. This results from
the ‘meristem dependence’ of seagrass populations (Tomlinson, 1974). Few seagrasses
have dormant meristems and horizontal vegetative spreading depends on the presence
of an apical meristem. In an established seagrass bed there may be many shoots but few
apical meristems. Thus in vegetative transplants no apical meristem may be present
and expansion growth cannot then occur. Apical meristems can generate from older
(short) shoots but this is not common.
Seedlings generate an apical meristem directly and are thus ideal for transplant
work providing that large quantities of seeds can be obtained and germinated and
provided that these can be sown effectively. With Thalassia testudinum this can be done
easily and economically (Thorhaug and Austin, 1976). This species also has the ad-
vantage that seedlings grow rapidly producing a ‘long’ shoot and several ‘short’ shoots
within six months. With Poszdonia australis seedling growth is much slower and
production of an apical meristem (within this species the second shoot) does not occur
for at least a year (McComb et al., 1981; West, 1980).
Austin in Thorhaug and Austin (1976) has costed the operation of seagrass
transplants and for Thalassia seedling transplants this works out to be about $U.S.
13,000 per ha. Ranwell et a/. (1974) costed their vegetative process at £1,000 per ha.
(1973 prices). It would therefore appear that restoration of at least some depleted sites
is possible and not outrageously expensive. However in all the successful experiments
the sites have not been subject to stresses or impacts (for example the hot water
discharge involved in the area of depletion used by Thorhaug (1974) had been
discontinued before the transplant experiments were conducted). In areas where man-
induced impacts occur and where the most depletions have resulted restoration of
seagrass beds may be much more difficult, at least until the major causal factor has
been identified and controlled. Nevertheless such situations do provide the opportunity
to study the effects of a variety of impacts on seagrasses.
CONCLUSION
Australia is richly endowed with seagrasses both in terms of species and in terms of
the extent of seagrass beds. This affords many opportunities for the study of
stability/instability, evolutionary changes, successional changes and natural
depletions, all of which are in great need of much further research. Seagrass com-
munities may be less stable than they were previously held to be.
The overall wealth of Australian seagrasses may suggest that local depletion of
seagrass beds near to several state capitals is unimportant. This may be so but until we
know much more about the stability of natural seagrass systems, natural depletions,
man-induced depletions and the role of seagrass beds in the natural economy of coastal
waters there needs to be a note of caution in any acceptance of local depradations and a
positive attitude to the possibility of restoration of seagrass beds. In any event there will
probably always exist a need for the restoration of seagrass beds as a result of natural
depletions and activities such as dredging, harbour development and offshore mining.
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Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
Seagrass Primary Production — a Review*
R. J. WEST and A. W. D. LARKUM
WEST, R. J., & LARKuM, A. W. D. Seagrass primary production — a review. Proc.
Linn. Soc. N.S.W. 106 (3), (1982) 1983:213-223.
Seagrasses dominate many inshore benthic communities in Australia and
contribute a high proportion of the total fixed carbon of the inshore ecosystem. The
methods in use to measure primary production of seagrass communities with special
reference to Australia are reviewed, and some of the comparable results listed.
Seagrass productivity is a reflection of climate, local environmental conditions, and
individual growth strategies of particular species, but is high in comparison with most
other plant communities.
Light, temperature, salinity, nutrients, sediment and human influence are
reviewed as possible controls on seagrass production.
R. J. West, N.S. W. State Fisheries, P.O. Box N211, Grosvenor Street, Sydney, Australia 2001,
and A. W. D. Larkum, School of Biological Sciences, University of Sydney, Sydney, Australia
2006; manuscript received 19 May 1981, accepted for publication in revised form 16 December
1981.
INTRODUCTION
Production refers to the capacity of a biological system to form organic matter,
and the rate of this formation is called productivity. Primary production is the capture
of radiant solar energy by plants and subsequent conversion of carbon and water to
organic matter during photosynthesis.
The measurement of primary productivity is of importance as it can give an in-
dication of the capability of a biological system to support a food web of secondary
producers and leads to a better understanding of the growth strategies of plant species.
Productivity measurements have the units dry matter produced per unit area per
unit time, usually expressed as grams dry weight per square metre per day, or tonnes
dry weight per hectare per year. When the relationship between carbon content and
dry weight of the plant material is known, then another useful unit is grams carbon per
square metre per hour. Although dry weight has been used extensively in this review,
one should bear in mind that the variability in ash content (interspecific, and seasonal)
may make comparisons of productivity in terms of dry weight misleading. Wherever
necessary, the values of McRoy and McMillan (1977) have been adopted for the sake
of comparison. These authors report that ‘available data on eelgrass indicate an
organic weight of about 80% of the dry weight, 47% of which is carbon’.
There are two broad methods often used to measure primary productivity
(Larkum, 1981). One is based on the calculation of net photosynthetic rates, while the
other is a measure of growth. The former is a short-term method, the period of ex-
perimentation being from a matter of minutes, to perhaps as long as twenty-four
hours, and involves the measurement of radioactive carbon fixation or of net oxygen
production. The latter is a long term method, based on the increases in the dry matter
over a period of weeks, months or years.
In most seagrasses, at least three processes, photorespiration (Larkum, 1981), the
loss of dissolved organic carbon (Brylinsky, 1977) and formation and translocation of
carbon reserves (Dawes and Lawrence, 1979), will ensure that net photosynthesis and
growth are not equivalent over the short term. Despite this, direct comparisons of the
methods have been attempted (Bittaker and Iverson, 1976).
* A paper presented at the Australian Seagrass Workshop, University of Sydney, 1981.
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
214 SEAGRASS PRIMARY PRODUCTION
METHODS OF MEASUREMENT
The short-term methods of estimation of primary production include
measurement of oxygen production, or of radioactive carbon uptake (Vollenweider,
1971). Both of these methods may be affected by the recycling of gases in the extensive
lacunal system possessed by most seagrasses (Zieman, 1968; McRoy and McMillan,
1977).
Methods employed to measure oxygen release from seagrass communities may
involve the use of closed vessels, such as glass bottles or large acrylic chambers, or they
may involve measurement in open waters (Vollenweider, 1971). The principal method
of oxygen determination is the Winkler chemical method (Strickland and Parsons,
1972), although oxygen electrode techniques are also widely in use (Pijanowski, 1975).
Diurnal oxygen concentrations in a flowing water mass provided one of the
earliest short term measures of productivity of aquatic plants (Odum, 1956, 1957).
This method made no allowances for planktonic metabolism, and relied heavily on a
crude estimate of water volume and speed. Loss of oxygen through bubbling, and
changes in concentration due to reaeration were further problems. Several later studies
relied on this method (Odum, Burkholder and Rivero, 1959; Quasim and Bhattathiri,
1971). Improvements have been made to this technique by the measurement of the
planktonic contribution (through the use of light and dark bottles), and of the oxygen
reaeration (Nixon and Oviatt, 1972). A further improvement is in the isolation of a
known volume of water (Weiner and Kirkman, 1980).
A variation of the above technique is the collection of whole plants (or of leaf
sections) which are subsequently incubated in glass jars either zm sztu or under con-
trolled laboratory conditions, and assayed for oxygen production (Jones, 1968; Buesa,
1974; Drew, 1978).
The most sophisticated method employed in the measurement of oxygen release
from seagrass communities is the simultaneous recording of oxygen, temperature, light
and salinity within a completely enclosed chamber, with circulating seawater (Weiner
and Kirkman, 1980; Clough and Attiwill, 1980). Clough and Attiwill (1980) have used
this method to predict the primary productivity of Zostera mueller: in Western Port Bay,
Victoria. These authors have calculated a photosynthetic response curve from several
diurnal incubations. This treatment could be substantially improved by increasing the
number of diurnal incubations to exclude seasonal variations in photosynthesis, and by
the continuous measurement of light at the plant-canopy level. However the problem
created by the existence of internal gas lacunae remains unresolved.
Wetzel (1964) and Drew and Larkum (1967) pioneered the in situ use of
radioactive isotopes to measure primary productivity of aquatic plants. These workers
used bell jars to incubate single plants which were later assayed for carbon-14. McRoy
(1974) adopted a similar method with the seagrass Zostera marina in Alaskan waters,
while Drew and Jupp (1976) measured uptake by Poszdonia oceanica in Mediterranean
waters. Penhale (1977) measured primary productivity of Thalassia testudinum and its
epiphytes by a carbon-14 method, this time improved by the reduction of unstirred
layers through the circulation of the seawater surrounding the plants. The use of radio-
isotopes in the field is somewhat difficult and does not overcome the problem of the
possible recycling of gases in the lacunae (McRoy and McMillan, 1977).
Long term methods of measurement of primary productivity were first used in
relation to seagrasses in Denmark in the early part of this century. Petersen (1913)
estimated the primary productivity of Zostera marina by harvesting this seagrass at
several stages of its growth cycle, and concluded that these plants made an important
contribution to production in the estuarine system. He did not however take into
account the rapid turnover of leaf material that is often characteristic of seagrasses, for
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
R. J. WEST ANDA. W. D. LARKUM 215
example, of many Zostera species (Sand-Jensen, 1975). Petersen’s estimates were
therefore very much lower than the true production rates. Such biomass figures are still
often used as an estimate of productivity of plants (Westlake, 1963), although turnover
of biomass is the more important concept. This can be defined as the number of times
mean biomass is replaced throughout the annual growing cycle:
Tusenowrae ite Annual increase in biomass
(crops per year) mean biomass
Zieman (1968, 1974, 1975) developed a marking technique to measure leaf
production in meadows of Thalassia testudinum in waters off Florida. This method was
described at the International Seagrass Workshop in the Netherlands as the only
reliable method to measure directly leaf production of seagrasses (McMillan et al. ,
1973). It has been used in many studies involving Thalassia testudinum (Zieman, 1975;
Patriquin, 1973; Greenway, 1977), Zostera marina (Sand-Jensen, 1975) and Posidonia
australis (West and Larkum, 1979; Kirkman and Reid, 1979).
Few workers have attempted to measure rhizome growth in seagrasses. Patriquin
(1973) introduced a method based on the time interval between new leaf production,
the dry weight of rhizome sections, and the number of leaf scars on the rhizome. This
method appears to work adequately for Thalassia testudinum and Posidonia australis
(West, 1980). Sand-Jensen (1975) has used the increase in shoot numbers in fixed
quadrats to estimate rhizome production for Zostera marina.
COMPARISON OF PRODUCTIVITY MEASUREMENTS
The results of many productivity studies have been summarized elsewhere
(McRoy and McMillan, 1977; Zieman and Wetzel, 1980). These authors have made
little or no reference to the many different methods adopted in the individual
productivity studies. However, in this review, the recommendation of the In-
ternational Seagrass Workshop held in the Netherlands (McMillan e¢ al., 1973) is
adopted, and only results based on leaf-making techniques are included.
Some representative values for leaf-production of several seagrass species are
shown in Table 1, and some available data concerning rhizome production shown in
Table 2.
There are often large differences in biomass between stands of the same seagrass
species, depending on localized environmental conditions, and this leads to variations
in the productivity estimates between such stands. The differences in biomass may be
due to different shoot densities, or variability in leaf length and width. West and
Larkum (1980) compared five study sites and found large variations in biomass,
densities and leaf characteristics between stands of Posidonia australis, however leaf
turnover was relatively constant (2.8-4.5 crops of leaves per year). This consistency in
mean leaf turnover is also apparent from a number of studies carried out on various
seagrass species (Table 1), although seasonal variations often mask the observation.
Thus it would seem that for many seagrass species, the number of leaves produced each
year by individual shoots may be quite consistent, although biomass, density and leaf
characteristics vary, depending on local conditions, such as depth, sediment, age of
bed, degree of wave exposure and exposure to salinity changes.
Thus care must be taken when assessing the productivity of a large area. It is not
often possible to make an estimate of the area of seagrass beds, as say by aerial
photography and then to multiply this area by the productivity found for plants in an
experimental area. Plant density is not always linearly related to plant cover (Larkum
et al., in press) and plant cover is what is usually estimated from aerial photography.
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
SEAGRASS PRIMARY PRODUCTION
216
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Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
217
R.J. WEST AND A. W. D. LARKUM
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Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
218 SEAGRASS PRIMARY PRODUCTION
Larkum ¢ al. (in press) found that a detailed analysis of the beds of Zostera capricorni in
Botany Bay gave a productivity value which was 5 fold lower than the estimate ob-
tained by the simple product of total area and a productivity value obtained in a dense,
shallow bed.
From the data available (Table 2) it would appear that rhizome production is
extremely variable between species. Rhizome production is obviously a reflection of
the growth habit of individual species, and the stability of a particular seagrass stand.
For example, West (1980) has shown that turnover of rhizome material is slow, but
depends markedly on site, in stands of Poszdonia australis in Botany Bay. Some erect
shoots may be more than 20 years in age (cf Kuo and Cambridge, 1978). Other
seagrasses must exhibit a high turnover of rhizomes, as these species appear to replace
rhizome annually (personal observation).
In general, it may be said that the production of organic matter in seagrass beds
occurs at a high rate when compared with most other plant communities (Westlake,
1963), and is a reflection of climate, local environmental conditions and individual
growth strategies of particular species.
CONTROLS ON PRODUCTIVITY
Although turnover of organic matter may be consistent in established stands of
seagrasses, there are many factors which can influence the biomass that can be
achieved or which influence the seasonality in growth at particular sites, and some of
these factors will now be discussed.
Light and temperature
Light is the most critical ecological factor for plant growth as it directly affects the
rate of photosynthesis. The lower depth limit to the establishment of seagrasses is
generally considered to be due to the minimum light intensity required by a particular
seagrass (Phillips, 1978), although few conclusive experiments have been performed.
Ostenfeld (1905) was one of the first workers to attempt to correlate distribution
patterns of a seagrass (Zostera marina) to underwater irradiance.
The short term methods of measuring productivity (see earlier) generally yield
information concerning the effect of light on photosynthetic rate, however the
variability in methods and light units measured makes comparison of this work dif-
ficult. For example, Jones (1968) and Buesa (1974, 1975) have used leaf sections
isolated in bottles, Drew (1978) used leaf slices in small jars, McRoy (1974) used whole
plants in jars, Penhale (1977) used whole plants in circulating seawater, and Clough
and Attiwill (1979) used whole plants (zn situ) in benthic chambers. Most of these
workers used different measures of light intensity.
Drew (1979) has compared the photosynthetic response of six seagrass species,
and found that, in all cases, a classic P-I curve for plants intermediate between sun and
shade adaptation could be demonstrated and that thermal damage occurred above 30-
35°C. Further this author has found that at least one species, Halophila stipulacea, is
capable of a chloroplast re-orientation that may be a protective mechanism in high light
situations. Trocine, Rice and Wells (1981) have also noted chloroplast clumping in a
Halophila species and suggest that this results in an increased photosynthetic tolerance
to ultraviolet-B radiation.
McRoy and McWilliam (1977) suggest that production can be expressed in terms
of light quantity. They report three separate studies giving estimates of between 5.3-
117.0 microgram carbon per langley per gram biomass, and propose that the similarity
in result may lead to a useful relationship from which productivity could be estimated
from light intensity.
PROC. LINN. Soc. N.S.W. 106 (3), (1982) 1983
R. J. WEST ANDA. W. D. LARKUM 219
Backman and Barilotti (1976) have shown that turion density of Zostera marina is
reduced when irradiance is reduced. Thus these authors present perhaps the best
evidence to show that a coastal development that affects turbidity will also affect
seagrass growth and production.
Temperature is also known to affect photosynthetic rates of seagrasses (Biebl and
McRoy, 1971), and the effect of high temperature on the growth of some species is well
documented (Zieman, 1975; Thorhaug, Blake and Schroeder, 1978). Zieman (1975)
found that leaf production in Thalassia testudinum was near zero at temperatures above
35°C or below 25°C.
The interaction of light and temperature in field situations makes the individual
effects of these environmental variables difficult to study, and has led to some
disagreement and confusion as to whether light, or temperature, is responsible for
seasonality of growth rates of seagrasses. The variation in method for the measurement
of seagrass production (i.e. leaf growth or photosynthetic rate) also adds to the con-
fusion. From our own studies with Poszdonza australis it would appear that temperature
controls individual leaf growth rates provided that light intensity has remained suf-
ficiently high to allow the plants to accumulate reserves in the rhizome (West, 1980;
West and Larkum, in preparation).
Salinity
There are few references in the literature to the effect of salinity changes on
seagrass productivity. Biebl and McRoy (1971) report a reduction in photosynthesis,
when salinity is reduced from seawater, for Zostera marina, while Ogata and Matsui
(1965) found for Zostera nana that a reduction in photosynthesis with increasing salinity
above seawater was due principally to carbon limitation.
Other laboratory studies have involved the transplantation of seagrasses to
aquaria at various salinities (McMillan and Mosely, 1967; McMahon, 1968; Mc-
Millan, 1974). McRoy and McMillan (1977) summarize these results: “Halodule shows
that broadest salinity tolerance . . . Halophila the narrowest ... and Thalassia and
Syringodium show intermediate tolerance’ .
Zieman (1975) found that productivity decreased above and below the optimum
salinity of 30 parts per thousand for Thalassia testudinum.
West (1980) found that large short term changes in salinity could cause mortality
of Posidonia australis, but there was no significant effect on leaf growth of surviving
plants.
The effect of salinity on establishment of seagrass species requires further studies
so that an attempt to explain distribution patterns can be made.
Sediment and nutrients
Little is known concerning the sediment requirements of most seagrasses although
some general observations have been listed by Den Hartog (1970). For example,
Posidonia australis generally occurs on sandy substratum, whereas Zostera capricorni is not
restricted to sand, and grows on a wide range of substrates (Harris, King and Ellis,
1979).
One advantage that seagrasses have over most algae is that they are not dependent ,
on the nutrient availability in the water column. Both anatomical studies and
physiological experiments indicate that seagrasses can take advantage of the often rich
nutrient supply offered by the sediment and interstitial water.
Kuo and Cambridge (1978) found a reduced vascular system in Poszdonta australis,
but all the elements required for nutrient uptake and transfer were present. Further,
McComb et al. (1981) suggest that the wide occurrence of bacterial colonies, fungi and
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
220 SEAGRASS PRIMARY PRODUCTION
other micro-organisms found in the rhizosphere and peripheral root tissue of Postdonia
australis may be involved in nutrient uptake and nitrogen fixation in seagrass roots.
Further evidence exists for the occurrence of nitrogen fixation in either the
phyllosphere or rhizosphere of other seagrasses (Patriquin and Knowles, 1972;
Goering and Parker, 1972). Capone et al. (1979) have found a relationship between
productivity (biomass) and nitrogen fixation in the Thalassia testudinum community.
McRoy, Blackburn and Klug (1981) have suggested that the level of nitrogen
fixation and sulphur reduction in sediments associated with seagrasses is related to the
successional stage of the seagrass community.
Phosphate absorption has been demonstrated for the roots and rhizomes of
eelgrass (McRoy and Barsdate, 1970; Penhale and Thayer, 1980), and for Poszdonia
australis (Larkum, unpublished data).
Human influence
The effect of man on seagrass productivity is a result of industrial and urban
development along shallow marine embayments and along coastal fringes. Thayer,
Wolfe and Williams (1976) have summarized many of the impacts of man on seagrass
systems. Light, salinity, water turbulence and nutrient levels are all affected by the
degree of foreshore development, and toxic contamination, dredging and ‘filling-in’
should be added as further possible impacts.
Evidence exists for a decline in Poszdonia australis beds near urban areas in New
South Wales, South Australia and Western Australia (Larkum, 1977). There has been
a continued reduction in the area colonized by Poszdonia australis in Botany Bay
(N.S.W.) through both increased turbidity and water turbulence (Larkum, 1976;
West, 1980). Aerial photographic evidence confirms the loss of 20% in the area
covered by seagrasses, in Botany Bay, in the last fifteen to twenty years. A decline of
Posidonia australis in Cockburn Sound, Western Australia, has also been described
(Cambridge, 1975, 1979). Similar situations have been documented for the meadows
of Posidonia oceanica off the French coastline (Meinesz and Laurent, 1978).
The recovery of seagrass communities in degraded areas may be extremely slow,
even if the environmental stress is relieved. This results from the fact that many
seagrasses appear to rely heavily on vegetative reproduction, and rhizome growth is
often very slow. Artificial revegetation is difficult and costly and thus careful
management of existing seagrass areas 1s imperative.
SECONDARY PRODUCTION
As stated, seagrasses produce large quantities of organic matter. Seagrass
meadows also offer a diverse habitat for secondary producers (Kikuchi and Peres,
OD):
Although leaf material may be grazed quite heavily in tropical areas by fish
(Ogden, 1976), turtles (Hirth, Klikoff and Harper, 1973) and dugongs (Heinsohn,
Wake, Marsh and Spain, 1977) there are few direct grazers in temperate regions
(Kikuchi and Peres, 1977). Conacher et al. (1979) have shown that the major fish
grazer of Postdonia australis, Monacanthus chinensis removes less than 1% of the
productive capacity. Nevertheless these fish do obtain some nutrients from ingested
leaf fragments. Kirkman and Reid (1979) have studied the role of a seagrass (P.
australis) in the carbon-budget of an estuary. Only 3% of the total seagrass biomass (in
terms of carbon) was consumed by herbivores, whereas the remainder was lost as
exuded dissolved organic carbon and detached leaves in approximately equal
proportions.
PROC. LINN. SOc. N.S.W. 106 (3), (1982) 1983
R. J. WEST ANDA. W. D. LARKUM 221
Thus, secondary production in temperate regions occurs primarily through the
detrital food chain (Fenchel, 1977):—
Leaching of dissolved organic and inorganic materials,
Mechanical breakdown of leaves (waves or ingestion),
Bacterial and fungal decomposition,
Attack by bacteriovorous microfauna and detritivores,
These small fauna are prey for large carnivores.
Oe OO Ne
GENERAL CONCLUSIONS
Care is required in the selection of a method to measure productivity and
Zieman’s leaf marking technique (Zieman, 1974) is still preferred.
Primary production of many seagrass species is high and related to growth
strategy of individual species and local environmental conditions.
The ability of seagrasses to inhabit sandy substrates, to use sediment nutrients,
and to accumulate carbon reserves are important factors which ensure a relatively
stable food supply and habitat, and this is reflected in a diverse and important faunal
community.
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Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
Ge
ah aie
Notes on the Biology of Australian Seagrasses*
J. Kuo
(Communicated by A. W. D. LARKUM)
Kuo, J. Notes on the biology of Australian seagrasses. Proc. Linn. Soc. N.S.W. 106 (3),
(1982) 1983: 225-245.
This contribution focuses on the anatomy, histochemistry and ultrastructure of
vegetative organs of Australian seagrasses in relation to their possible functions.
Detailed anatomical and ultrastructural structures vary with species. Some structures
are similar to those in terrestrial plants and others are unique in seagrasses for
adaptation to a marine environment. Some details of the taxonomy, distribution,
habitat and life history of Australian seagrasses are also included. The structure of the
phyllosphere and rhizosphere in seagrasses is discussed as well as the possible
significance of the role of epiphytes and epifauna.
J. Kuo, Electron Microscopy Centre, University of Western Australia, Nedlands, Australia 6009,
manuscript recewwed 19 May 1981, accepted for publication in revised form 17 November 1982.
INTRODUCTION
Seagrasses are aquatic angiosperms which are completely adapted to life in the
marine environment. According to Arber (1920) and den Hartog (1970) they require
at least the following adaptations to be able to colonize the marine environment suc-
cessfully: (1) they live fully submerged in seawater, (2) they have an anchoring system,
(3) they have a hydrophilous pollination, (4) they are able to cope with a high salinity.
Seagrasses are abundant in Australian waters and play a significant role in marine food
chains (Larkum, 1977; King, 1981; McComb et al., 1981). The ecology of seagrasses
including Australian species has been intensively reviewed (McRoy and Helfferich,
1977; Larkum, 1977; Phillips and McRoy,, 1980; McComb et a/., 1981) and will not be
dealt with here. Knowledge on other biological aspects of these marine vascular plants
is still fragmented. This contribution, which emphasizes anatomy, cellular structure
and function, is concerned with these adaptations and other biological aspects of
marine angiosperms in Australian waters. A comparative leaf morphology and gross
anatomy of seagrasses has been described by Tomlinson (1980).
TAXONOMY, DISTRIBUTION AND HABITATS
Taxonomy
Den Hartog (1970) contributed an excellent taxonomic study on seagrasses of the
world, listing 49 species belonging to twelve genera and four families of angiosperms
regarded as seagrasses. A brief taxonomic study of eastern Australian Zostera has been
made by Jacobs and Williams (1980). A comprehensive taxonomy of temperate
Australian seagrasses will be available soon (Womersley and Robinson, in
preparation). Australia is well endowed with seagrasses; there are 25 described species
belonging to all genera except the genus Phyllospadix represented in Australian waters
(Table 1). Three of these species have only recently been described (Greenway, 1979;
Cambridge and Kuo, 1979), and four new Posidonia species are to be described (Kuo
and Cambridge, in preparation). Dehydrated specimens, which were collected as drift
on the beach and deposited in herbaria, were used by den Hartog (1970). However,
herbaria have recently received an increasing number of seagrass specimens taken
from their natural habitats. Unfortunately, most herbaria have stored seagrasses in a
dehydrated state and the author found it difficult to study the morphology of these
dried seagrasses.
* A paper presented at the Australian Seagrass Workshop, University of Sydney, 1981.
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
226 BIOLOGY OF AUSTRALIAN SEAGRASSES
TABLE 1
The Australian Seagrasses
* 9)
Endemic to~
Genus! Species Australia Distribution?
Zosteraceae (Potamogetonaceae)
Zostera capricornt Aschers. C Qld., NSW
(12) mucronata den Hartog E S SA,WA
muellert Irmisch & Aschers. E SSA, Tas, Vic.
Heterozostera tasmanica (Martens ex Aschers.) E S SA, WA, Vic.,
(1) den Hartog NSW, Tas.
Posidoniaceae (Potamogetonaceae)
Posidonia angustifolia Cambridge & Kuo E S WA,SA
(5) australis Hook F. E S NSW, Vic., Tas.,
SA, WA
ostenfeldi den Hartog E S WA,SA
sinuosa Cambridge & Kuo E S WA,SA
Cymodoceaceae (Zannichelliaceae; Potamogetonaceae)
Halodule pinifolia (Miki) den Hartog N Qld.
(6) uninervis (Forsk.) Aschers. N WA, Qld.
in Boissier
Cymodocea angustata Ostenfeld E N WA
(4) rotundata Ehrenb. & Hempr. N Qld.
ex Aschers.
serrulata (R.Br.) Aschers. & Magnus N Qld.,WA
Syringodium isoetifolium (Aschers.) Dandy C WA, Qld.
(2)
Thalassodendron ciliatum (Forsk.) den Hartog N Qld.
(2) pachyrhizum den Hartog E S WA
Amphibolis antarctica (Labill.) Sonder et E S WA,SA,
(2) Aschers. Vic., Tas.
griffithu (J. M. Black) den Hartog E S WA,SA
Hydrocharitaceae
Enhalus acoroides (LL.F.) Royle N Qld.
(1)
Thalassia hemprichu (Ehrenb.) Aschers. N Qld.
(2)
Halophila decipiens Ostenfeld N Qld., NSW
(9) ovalis (R.Br.) Hook F. C WA, Vic., NSW,
Qld., NT
ovata Gaud. in Freycin. N WA, Qld.
spinulosa (R.Br.) Aschers. N WA, Qld.
tricostata Greenway 1B: N Qld.
1. Number in (_) represents total world species in the genus. Only the genus Phyllospadix (5) is not
represented in Australia.
2. E: species endemic to Australia.
3. N, occurring only in northern, tropical waters; S, occurring in southern, temperate waters; C,
cosmopolitan, occurring in tropical and temperate waters.
Qld., Queensland; NSW, New South Wales; Vic., Victoria; SA, South Australia; Tas., Tasmania; WA,
Western Australia; NT, Northern Territory.
Distribution
A brief biogeography of Australian seagrasses has been discussed by Specht
(1981). Australian seagrasses show a high degree of endemism as well as having a
strong affinity with those of other continents and island groups. Half of the Australian
species, and the genera Amphibolis and Heterozostera, are endemic to Australia, while
some species, including Enhalus acoroides (L.f.) Royle, Thalassodendron ciliatum (Forsk.)
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
J.KUO 227
Fig. 1. A typical temperate Australian seagrass Posidonia australis Hook F. Bd: Leaf blade; Sh: Leaf sheath;
Fr: Fibres; Rt: Roots; Ep: Epiphytes. Note the rhizome is covered by the fibres.
PRoc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
228 BIOLOGY OF AUSTRALIAN SEAGRASSES
den Hartog, Syringodium tsoetifolium (Aschers.) Dandy are also found elsewhere in the
Indo-Pacific region. Halophila ovalis (R.Br.) Hook.f. has a much wider distribution.
Within Australian waters, some genera (e.g. Enhalus, Cymodocea, Halodule, Thalassia
and most species of Halophila) are found only in tropical areas, others (e.g. Poszdonza,
Amphibolis, Heterozostera and most species of Zostera) occur only in temperate waters. On
the other hand, Syringodium tsoetifolium and Halophila ovalis are found in both temperate
and tropical waters. The latitudes 30°S on the east coast and 25°S on the west coast of
Australia appear to be transition points for a great number of tropical and temperate
seagrass communities (Larkum, 1977). No information is available at present on
seagrasses of nothern Australia, from Port Hedland in Western Australia to the Gulf of
Carpentaria in Queensland (McComb et al. , 1981).
Habitat
Different seagrasses grow on different substrates (den Hartog, 1970). Halophila is
found in all kinds of habitats from open sea to estuaries and inlets. Cymodocea, Thalassia
and Zostera can also be found on various substrates and the depth limit of most of the
species of these genera is 10 to 12 metres. Syringodium is restricted usually to the upper
sublittoral zone where it sometimes appears rarely to make up monospecific meadows.
Enhalus, Amphibolis, Thalassodendron and Posidonia are also restricted to the upper
sublittoral zone with some exceptions. In Western Australia, Cambridge (1975) found
that Thalassodendron pachyrhizum den Hartog occurs only on limestone and granitic reefs
and never on sands, and could be found to a depth of at least 40 metres. She also
noticed that Amphibolis usually inhabits sand and sand-covered reefs as either dense
meadows or small clumps, while Poszdonia ostenfeldi den Hartog is found in sandy
substrates on unsheathed sand banks. Posidonia australis Hook. f. forms vast meadows in
embayments enclosed by reefs, shoals and islands. P. australis appears to be quite
tolerant of salinity fluctuations, as it has been reported in Shark Bay where salinity can
be as high as oe (Cambridge, 1979, personal communication). The largest, densest
seagrass beds are found in St Vincent and Spencer Gulfs of South Australia and in
Shark Bay in Western Australia. The area of the latter embayment has been estimated
at more than 10,000 km? (Davis, 1970). There is no doubt that the most extensive
seagrass community in these beds is that formed by Posidonza spp.
VEGETATIVE STRUCTURE AND FUNCTION
Seagrasses show a high degree of uniformity in their morpological appearance;
most genera have well-developed rhizomes and linear or strap-shaped leaves, Halophila
being the only exception. However, the anatomical structure of the seagrasses exhibits
a considerable degree of diversity. Some seagrass genera are monomorphic (i.e. with
shoots bearing only one kind of leaf foliage leaves, Postdonia, Zostera, Heterozostera,
Cymodocea, Halodule and Enhalus) and others are dimorphic (i.e. with two types of shoot-
bearing scale leaves and foliage leaves respectively, Amphibolis, Thalassodendron,
Syringodium, Halophila and Thalassia). Species of Halophila have shoots of variable
length. A typical seagrass, P. australis (Fig. 1), has a creeping rhizome which at each
node produces two branching roots and a unit of three foliage leaves protected by a
sheath at the base.
1. LEAF BLADE (Figs 2-4).
A transverse section of leaf blade (Fig. 2, A-G) reveals a thin cuticle covering a
single cell layer of epidermis which lies above the parenchyma (mesophyll) or fibre cells
(mechanical tissue). Numerous vascular bundles and air lacunae are embedded among
the parenchyma cells.
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
J. Kuo 229
Fig. 2. The leaf blade anatomy of seagrasses and the leaf epidermal morphology of some Australian
Posidonia.
A-G. Transverse sections of the leaf blade of A, Syringodium isoetifolium; B, Zostera muelleri; C, P. australis; D,
Z. muellerti (SEM); E, Heterozostera tasmanica (SEM); F, P. ostenfeldu (SEM); G, Enhalus acoroides (SEM). A:
Air-lacunae; E: Epidermis; V: Vascular bundles.
H-J. The surface view of the Posidonia leaf epidermis (all same magnification). H, P. australis; I, P.
angustifolia; J, P. sinuosa.
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
230 BIOLOGY OF AUSTRALIAN SEAGRASSES
Cuticle: There are no stomata in seagrasses (Doohan and Newcomb, 1976; Kuo, 1978)
(Fig. 2, H-J). The appearance of the cuticle under electron microscopy differs among
species. With the exception of P. sznuosa (Cambridge and Kuo, 1982) the genus
Postdonia (Kuo, 1978; Fig. 3, D) has an unusual porous texture. The inner face of the
cuticle has oval cavities in Zostera muellert Irmisch & Aschers and Heterozostera tasmanica
(Martens ex Aschers.) den Hartog (Fig. 3, F). A similar structure has been observed in
Z. capensis and it has been suggested that it might have an association with iron ab-
sorption, secretion and storage (Barnabas et al. , 1977) or might even have an excretory
function (Barnabas ét al. , 1980). The cuticle may appear as a thin, electron-transparent
layer in P. sinuosa, Thalassia hemprichit (Ehrenb.) Aschers., Cymodocea serrulata (R.Br.)
Aschers. & Magnus, Cymodocea rotundata Ehrenb. & Hempr. ex Aschers. and
Syringodium isoetifolium (Fig. 3, E; Doohan and Newcomb, 1976). The cuticle has been
reported as absent in the northern hemisphere Thalassia testudinum Banks ex Konig
(Benedict and Scott, 1976). It is thought that the cuticle of the submerged leaves offers
little resistance to carbon diffusion from surrounding water in the absence of a func-
tional stomatal system.
Epidermal Cells: The shape of the epidermal cells is sometimes a useful taxonomic
character (Cambridge and Kuo, 1979; Fig. 2, H-J). The thickened walls of epidermal
cells are rich in pectin and celluloses but are not lignified (Kuo, 1978). Epidermal
transfer cells have been observed in Thalassia, Cymodocea, Halophila, Halodule, Zostera
(Fig. 3, G) and Heterozostera (Jagel, 1973, 1982; Birch, 1974; Doohan and Newcomb,
1976; Barnabas et al., 1977; Kuo, unpublished) but are absent in Poszdonia (Fig. 3, H),
Amphibolis, Syringodium and Thalassodendron (Kuo, 1978; unpublished). An unusual
structure has been observed in Halophila epidermis (Birch, 1974). Epidermal cells
contain most of the leaf chloroplasts as well as numerous mitochondria, Golgi bodies
and microbodies (Doohan and Newcomb, 1976; Kuo, 1978; Fig. 3, G-H). Phenolic
materials may also occur in the epidermal cells of certain species. The epidermal cells
appear to have active photosynthetic and mitochondrial metabolisms associated with
osmoregulation (Jagel, 1973; 1982). These cells may be the primary site of carbon
dioxide fixation and carbon dioxide uptake. The products of photosynthesis in the
epidermal cells may be transported to the mesophyll cells and to the vascular systems
via plasmodesmata (Kuo, 1978).
The !°C/!*C ratio, measured as 6 !°C values in the range - 3.0 to - 19.0%, have
been recorded in seagrass leaves (Smith and Epstein, 1971; Doohan and Newcomb,
1976; Benedict and Scott, 1976; Andrews and Abel, 1979; McMillan e¢ al., 1980).
These values are within the range usually associated with C, terrestrial plants that fix
carbon dioxide during photosynthesis with phosphoenolpyruvate carboxylase (Smith
and Epstein, 1971). The high variability in 6 '3C values might be associated with a
variable photosynthesis metabolism in seagrasses (McMillan et al., 1980) and it has
been thought that the C, pathway might operate in seagrass photosynthesis (Benedict
and Scott, 1976). However, recent biochemical studies of four Australian tropical
seagrasses by Andrews and Abel (1979) indicate that photosynthesis occurs via the C3
rather than the C, pathway. Studies of leaf anatomy, which do not show the ‘Kranz’
anatomy of terrestrial C, plants (Fig. 1, A-C) (Jagel, 1973; Doohan and Newcomb,
1976; Barnabas et al., 1977; Kuo, 1978; Tomlinson, 1980) and of photorespiration
(Hough, 1976) suggested that seagrasses may not be typical C3 or Cy plants. At the pH
of seawater, there is more bicarbonate and less carbon dioxide available in seawater
than there is in the air. Thus, it is still uncertain whether seagrasses take up carbon in
carbon dioxide or bicarbonate form from surrounding water for photosynthesis (Beer et
al., 1977; Sand-Jensen, 1977).
PROC. LINN. SOc. N.S.W. 106 (3), (1982) 1983
231
Fig. 3. The structure of the leaf epidermis and the cuticle of some Australian seagrasses.
A. The portion of the leaf blade in P. ostenfeldi. Note that the epidermal cells (E) contain tannins and fibre
cells (F) associated with epidermal and other hypodermal cells.
B and C. A pronounced cuticle (arrows) covering the epidermal cells (E) of P. australis, sudan black and
toluidine blue stained respectively.
D-F. Electron micrographs of seagrass cuticle.
D, P. australis cuticle (C) has a porous appearance. E, an electron-transparent cuticle (arrow) in Springodium
isoetifollum. F, cup-shaped invaginations (arrows) occur in the outer epidermal wall (W) of Z. muellert.
G. Electron micrograph of the epidermal cell of Z. muellert. Note that the cell has many chloroplasts; more
cell wall ingrowths (asterisks) occur in the basal than the distal tangential walls. Some plasmadesma (arrow)
are present between the epidermal cells.
H. Electron micrograph of the epidermal cell of P. australis. Note that the cell also has many chloroplasts
(CH) and a large nucleus (N) but is lacking cell wall ingrowths. Some plasmodesmata (arrow) also occur on
the walls between cells.
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
232 BIOLOGY OF AUSTRALIAN SEAGRASSES
It is believed that the leaves of seagrasses have the capacity to take up nutrients
from surrounding water (Arber, 1920; Sculthorpe, 1967; Raven, 1981). Using
radioactive isotopes the uptake of carbon (Harrison, 1978; Barbour and Radosevich,
1979), phosphorus (McRoy and Barsdate, 1970), cadmium (Faraday and Churchill,
1979; Brinkhuis e¢ a/., 1980) and manganese (Brinkhuis et al., 1980) by the leaf has
been demonstrated for several seagrasses. However, there are no similar studies on
Australian seagrasses.
Mesophyll Cells and Air-lacunae: Mesophyll cells of seagrasses are thin-walled but highly
vacuolated. The thin peripheral cytoplasm contains few chloroplasts with small starch
granules. The mesophyll cells surround air lacunae of varying size (Fig. 2, A-G) and in
some genera (e.g. Zostera) the surface area of air lacunae may occupy half of a leaf
surface area in cross section (Fig. 2, D). It has been shown that the air lacunae contain
oxygen and carbon dioxide (Zieman, 1974). The storage of these gases in the air
lacunae could swell the leaf blades of Thalassia testudinum up to 200-250% during the
day (Zieman, 1974). The presence of air lacunae has been considered important in
seagrass photosynthesis, particularly in re-fixation of carbon dioxide. Oxygen
produced in photosynthesis could build up in these lacunae, and respiratory and
photorespiratory carbon dioxide could diffuse into them as well. Zelitch (1971)
estimated that up to 50-67% of photorespired carbon dioxide in T. testudinum leaves
was recycled.
Z. muellert grows in both intertidal and subtidal mudflats in Westernport Bay,
Victoria. The surface area of air-lacunae in the leaves of the intertidal form is only
about one-third that in the leaves of the subtidal form (Kuo, in preparation). This
finding correlates well with the photosynthesis rate in this species. Clough and Attiwill
(1980) found that the photosynthesis rate of the subtidal form is about three times that
of the intertidal form.
There are many septa interrupting the air-lacunae along the leaf. Each septum
consists of a group of small parenchyma cells with minute pores at their intercellular
spaces. Numerous external wall ingrowths projecting into these pores are present in the
septa of Zostera muellert and Z. capricorni but are absent from those of more than twenty
seagrass species belonging to twelve genera examined so far (Kuo, in preparation).
Vascular System: In general, each vascular bundle is surrounded by a layer of sheath cells
(Fig. 4, B, E), though Benedict and Scott (1976) could not distinguish sheath cells in
Thalassia testudinum. The sheath cells of Posidonia have a thin but lignified wall (Kuo,
1978), those of Syringodium have suberin lamellae in the wall, those of Thalassodendron
and Amphibolis have a thick but not lignified wall and those of Zostera and Heterozostera
have cell wall ingrowths (Kuo, unpublished). The vascular sheath cells of seagrasses
therefore appear to have the potential to regulate solute movement between the
mesophyll and the vascular tissues, but this has never been demonstrated ex-
perimentally.
The number and size of xylem elements in the vascular bundles of seagrasses, as
well as in other aquatic plants, are much reduced in comparison with terrestrial plants
(Sculthorpe, 1967). Xylem walls in seagrasses have little lignification and secondary
wall thickening (Kuo, 1978; Fig. 4, D, G). The reduced xylem has led some resear-
chers to suggest that there is little xylem transport in seagrasses (e.g. Tomlinson,
1972), but precise experimental work on this point appears to be lacking. The structure
of the phloem tissues in seagrasses is similar to that of terrestrial plants. Sieve tubes of
Posidonia contain a plasmalemma, endoplasmic reticulum, mitochondria and plastids
but lack nuclei (Kuo, 1978; Fig. 4, G). Companion cells are rich in cytoplasm. Sieve
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
233
;
&
ie
Le
7Ou ’
:
Fig. 4. The leaf vascular structure in some Australian seagrasses.
A. A polarizing micrograph of P. australis leaf blade. Notice that the cell walls of the epidermal and fibre cells
are intensively birefringent but that of vascular bundle (V) is not so.
B. The sheath cells of vascular bundle (V) in P. sznuosa is not so intensively stained with PAS reaction. Fibre
cells (arrows) occur between the epidermal (E) and the hypodermal cells.
C.(SEM) and D. The vascular bundle (V) of Z. mueller’. Note that a single xylem lacunae (X) separates from
the phloem tissue (P). Fibre cells associated with vascular tissue but not with the epidermal cells (E).
E. Both the epidermal cell contents and the sheath cells of the vascular bundle (V) in P. australis leaf blade
are strongly autofluorescent.
F. Nacreous wall sieve elements (arrows) are found in the vascular bundles of H. tasmanica leaf sheath.
G. Electron micrograph of the vascular bundle in P. australis leaf blade. Note that the bundle is surrounded
by a layer of lignified sheath cells (B). The walls of xylem elements (X) are poorly secondary thickness and
weakly lignified and that of sieve tubes (S) and of parenchyma cells are normal.
H. Nacreous wall sieve tubes (N) and cell wall ingrowths in parenchyma cells (P) occur in the vascular
bundle of Z. mueller: leaf blade.
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
234 BIOLOGY OF AUSTRALIAN SEAGRASSES
tubes of the family Zosteraceae and the sole genus Halodule of the family Cymodoceacae
have nacreous or thick walls with reduced lumens and the companion cells of these
species have cell wall ingrowths (Kuo, 1983; Fig. 4, F, H). The reduction of lumen
area in nacreous sieve elements relates to the translocation rate and has received little
attention in terrestrial plants (Esau and Cheadle, 1958).
Mechanical Tissues: The prominent fibre cells (Fig. 3, A, C; Fig. 4, B) of the leaves are
of particular interest. Mechanical tissues occur only in the leaves of certain genera,
they may be associated with vascular bundles (e.g. Zostera) or distributed among
parenchyma tissue (e.g. Postdonia). They have thickened walls, which consist of pectin
and cellulose but do not contain lignin (Kuo, 1978). Thus they could provide tensile
strength but retain a high degree of flexibility, allowing the leaves to withstand
vigorous ocean wave action. The distribution of these fibre groups in the leaves of
Postdonia can be used as a taxonomic character (Sauvageau, 1890; Cambridge and
Kuo, 1979).
2. LEAF SHEATH (Fig. 5).
The leaf sheath encloses the basal portion of the leaves and usually lies beneath the
sediment surface (Fig. 5, A). The anatomy and cellular structures of the sheath of
Posidonia differ markedly from the blade (Kuo, 1978; Fig. 5, B, C). The cuticle is
electron-transparent and is not porous. A thin lignified or suberized layer is present on
the outer wall of abaxial epidermal cells. This layer or the cuticle may act as a physical
barrier between the inner developing leaf tissue and surrounding water. Tyerman
(1979, personal communication) reported that the sheath may function to maintain a
relatively low osmotic pressure for the leaves meristem of P. australis. In contrast to the
blade the sheath epidermal cells of Poszdonza lack chloroplasts and are highly vacuolated
with a thin peripheral cytoplasm. The structure of vascular bundles is similar to those
in the blade. Groups of fibre bundles are widely distributed among the parenchyma
tissues of the sheath (Kuo, 1978; Fig. 5, B) but in contrast to the fibre bundles of the
blade, they are lignified and because of this they persist on the rhizomes long after the
other tissues of the leaf sheath have rotted away (Fig. 5, A). The sheath fibres of
Posidonia can be deposited both beneath living seagrass meadows and on ocean floors
bordering the coast (Kuo and Cambridge, 1978). Deposits of these marine fibres in
Spencer and St Vincent Gulfs in South Australia are present in such quantities that the
fibres were harvested over a limited period (1905 to 1915) for grain bags, paper making
and insulation material (Winterbottom, 1917; Reid and Smith, 1919).
3. STEM AND RHIZOME (Fig. 5).
In transverse sections, both erect stem and rhizomes of Amphibolis, Zostera and
Thalassodendron exhibit similar morphological features (Ducker et al., 1977; Fig. 5, D,
E, G). The central stele has a central xylem surrounded by phloem bundles (Fig. 5, E).
The stele is surrounded by the cortex in Poszdonia or by aerenchyma in Amphibolss,
Thalassodendron and Zostera. Two distinct cortical zones can be recognized in Amphibolis
and Zostera based upon morphological and histochemical characters. Starch grains are
prominent in the cortical cells of Postdonia rhizomes, and groups of lignified fibre cells
are scattered through the cortex of Postdonia rhizomes (Kuo. and Cambridge, 1978; Fig.
5, G). A few peripheral vascular strands are distributed among the cortical tissues (Fig.
5, E). The distribution and number of vascular strands appear to be a species
characteristic (den Hartog, 1970; Ducker e¢ al., 1977). A distinct cuticle covers both
stem and rhizome. Cell walls of the epidermal and hypodermal cells are thickened and
lignified and sometimes these cells contain polyphenolic materials (Fig. 5, F). A few
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
235
4 28s
Fig. 5. The anatomy and the leaf sheaths and rhizomes in seagrasses.
A. Old leaf sheaths of P. australis contain numerous fibres.
B and C. Transverse sections of the leaf sheaths of P. australis and Z. muelleri respectively. Many air-lacunae
(A) occur in both species but P. australis has more fibre bundles among the parenchyma tissue. V: vascular
bundles.
D. A scanning electron micrograph of a P. australis rhizome in a transverse view.
E. A transverse section of Syringodium tsoetifolium rhizome shows numerous vascular bundles (V), air-lacunae
(A) and large tannins-containing cells (T) but there are no fibre bundles in the rhizome.
F. The cell walls of both the epidermal (E) and the hypodermal cells (H) of P. australis rhizome are
autofluorescent but those of the cortical cells (C) are not.
G. Transverse section of P. australis rhizome showing the central stele has a xylem element (X) surrounded
by numerous phloem bundles (P), many large fibre bundles scatter among the cortical tissue which contains
numerous minute starch granules.
H. A higher magnification of a fibre bundle in P. australis rhizome showing the middle lamellae of the fibre
cells are autofluorescent.
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
236 BIOLOGY OF AUSTRALIAN SEAGRASSES
bacteria and diatoms are often present on the surface of rhizomes and numerous
epiphytic algae occur on the stem.
4. ROOTS (Fig. 6).
Both rhizomes and roots serve as anchor systems for seagrasses. The external
morphology of roots may vary between species and appears to be associated with their
ecological habitat and mineral nutrient environment. For example, the roots of
Halophila and Zostera are creamy in colour and soft, with many root hairs, and appear to
be suitable for survival on soft substrates. Posdonia roots are thick but relatively soft
with many branches; the root hairs are sparse, and the plants live on the sandy ocean
floor (Figs 1, 6, A). Hard, wavy and branched roots are usually found in Amphibolis,
and appear suitable to anchoring on harder substrates such as rocks or marine fibres.
The roots of Thalassodendron are thick and strong, with shiny black surface roots, and
have few branches. These roots apparently penetrate the interstices of the rocky
substrates and anchor the plants against the turbulence of ocean waves. The anatomy
of the roots, however, appears to be very similar among the species. All roots have a
distinct root cap (Fig. 6, B). The central stele (Fig. 6, C-E) consists mainly of well
developed phloem and a central weakly lignified xylem, and is surrounded by an
endodermis which has a Casparian strip in the radial cell wall (Kuo and Cambridge,
1978; Fig. 6, H). Cortical cells are thin-walled and may have many large air-lacunae.
Cell walls of the epidermal cells and three to four layers of hypodermal cells are
thickened and lignified in harder roots, such as those of Thalassodendron, Amphibolis and
Posidonia (Fig. 6, F). The thickened hypodermal walls may have suberin lamellae (Kuo
and Cambridge, 1978; Fig. 6, 1). Nutrients are taken up by roots and rhizomes of
seagrasses from the sediment and are transported to all parts of the plant. Directions
and rates of nutrient transport are likely to be dependent upon the morphology, age
and physical condition of the individual plants as well as the species. This translocation
probably occurs in the xylem and phloem of the vascular system. However, McRoy
and Barsdate (1970) showed that the leaves and stems of Zostera marina L. absorbed
more phosphorus than did the roots and rhizomes. Wetzel and Penhale (1979) found
the rates of carbon uptake and transport from roots and rhizomes to the leaves of
Halodule wrghtii Aschers. are much higher than those for Zostera marina and Thalassia
testudinum. Thus it appears that both leaves and roots of seagrasses may have the
capacity to take up nutrients, although the quantitative role in nutrient uptake in a
natural environment is still unknown. Data on nutrient uptake by Australian
seagrasses are still not available.
LIFE HISTORY
Some seagrasses appear to show infrequent flowering. Cymodocea serrulata flowers
have been recorded once, and these were male (Kirkman, 1975) while Tomlinson
(1969) observed an average arrangement of male:female shoot ratio of 4:1 for Thalassza
testudinum in Florida. Other species show regular flowering and some have the male and
female flowers on the same shoot — e.g. Poszdonia australis whose flowers are initiated in
autumn when day lengths are shortening and temperatures are falling, and the flowers
open in spring (Cambridge, 1975).
Aspects of floral morphology and development in seagrasses have been in-
vestigated only for some members of the family Cymodoceaceae (Isaac, 1969; Kay,
1971; Ducker et al., 1978; Tomlinson and Posluszny, 1978). But this information is
lacking for other families of seagrasses, probably due to the difficulty of obtaining
material. Posidonia flowers (Fig. 7, A) are bisexual with a perianth. Mature pollen
grains of most seagrass species are thread-like with a pair of fine curved hooks at the
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
237
Fig. 6. The anatomy of P. australis roots.
A. A scanning electron micrograph of a mature root.
B. A median longitudinal section of a mature root tip with a pronounced root cap (RC).
C. A transverse section of a mature root showing the stele is surrounded by a series of air-lacunae (A), many
thin-walled cortical cells (C) and a layer of the thin-walled epidermal cells (E).
D. A scanning electron micrograph of a mature root in a transverse view.
E. A high magnification of a stele with a distinct layer of endodermis (EF).
F. A high magnification of the root periphery showing that the thin-walled epidermal cells (E) with thicker-
walled hypodermal cells (H).
G. Numerous starch granules (arrows) are located in the basal portion of the root cap cells.
H. A distinct Casparian strip (CS) is present in the radial walls of two adjacent endodermal cells.
I. Electron dense suberin lamellae (arrows) are present in thickened walls (W) of the adjacent hypodermal
cells.
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
238 BIOLOGY OF AUSTRALIAN SEAGRASSES
end (Pettitt and Jermy, 1975; Pettitt, 1976, 1980; Ducker et al., 1978; Fig. 7, B),
probably for attachment to the stigma surface. In contrast to terrestrial flowering
plants, the pollen of seagrasses lack an exine or it is poorly developed (Fig. 7, C, D).
The pollen is released under water and drifts about submerged. The mechanism of
seagrass pollination has been discussed elsewhere (Ducker and Knox, 1976; Ducker et
al., 1978; Pettitt, 1980). Enzymatic properties of the pollen wall and stigma pellicle in
seagrasses are comparable with those found in terrestrial flowering plants. Pettitt
(1980) suggested a similar mechanism of cuticle erosion in both pollen grain and stigma
might well follow compatible pollination both on land and in the sea.
The fruits of Postdonta mature about three months after anthesis (Fig. 7, E, F).
Cambridge (1975) estimated that P. australis in Cockburn Sound, Western Australia,
may produce 500 fruits per square metre of meadow. Each seed is enclosed by a thin
membrane which resembles the seed coat of terrestrial plants, and then by a pericarp
which has a well-developed aerenchyma system. The bulk of the seed consists of en-
dospermic tissue which serves as nutrient storage. Starch and lipid are the main
nutrient reserves and storage protein is lacking. Nitrogen, phosphorus and other
macro-and micro-elements occur in concentrations which are in general comparable to
those of terrestrial plants (Hocking e¢ al. , 1980). Mature fruits with their pericarps are
released in summer and float to the water surface. After a day or two the pericarp
ruptures and the seeds sink to the floor of the sediment (Fig. 7, G). Germination may
begin even before the seeds are released from the pericarp. The young seedlings (Fig.
7, H) may depend upon the nutrient storage of the seed up to nine months after
germination (Hocking et al. , 1981).
In Amphibolis and Thalassodendron the germination of the seed is viviparous. The
flowers of these seagrasses occur on separate plants in summer. The fruits develop at
the apices of the lower branches of female plants. Mature embryos burst through the
apices of their pericarps and appear as little seedlings, which continue their growth on
the parent plant for some time (Fig. 7, I). The seedlings, with cup-like pericarp
structures, are released from the parent plants in winter and float in the ocean. Finally,
the seedlings become anchored in the sediments or to the bases of plants in meadows by
the comb-like lobes of the upper pericarp. Then the stems begin to grow rapidly and
later shoots produce rhizomes with roots.
Cambridge and Kuo (in press) describe an unusual form of vegetative
propagation in Heterozostera tasmanica. Small propagules develop in the non-fertile erect
stems of the parent plant during spring and summer. Eventually, erect stems carrying
propagules break off from the parent plants and drift away. These propagules may then
find a suitable location for further development.
Although the rate of leaf production in an established seagrass meadow has been
estimated (West and Larkum, 1979; Cambridge, 1981), little is known about either the
rate of underground tissue production or the growth rate from a seedling to its maturity
in any seagrass. Old leaves of Postdonia plants usually break off at the junction between
the blade and sheath in autumn and leaf litter from seagrass may accumulate as high
as two metres along some parts of the south-west Australian coastline during the
winter. It is washed back into the ocean by late spring. There is no doubt that
seagrasses are playing an important role in marine food chains.
RHIZOME BRANCHING
Despite the large numbers of seeds produced, seagrass seedlings have been in-
frequently observed in the field and vegetative propagation by growth of rhizomes is
probably of greater importance in the maintenance and spread of seagrasses than is
seed production (Tomlinson, 1974).
PROC. LINN. SOc. N.S.W. 106 (3), (1982) 1983
J.KUO 239
je
SUH IU neers I ga fe
eee i HOA) ey a) j
2 U2 6 US 16 17 VR Ya 2ID 2 22a 2
Fig. 7. Flowers, pollen grains, fruits and germination in seagrasses.
A. Flowering in P. australis.
B. Thread-like pollen grains of P. australis.
C and D. Electron micrographs of pollen grains of P. australis showing that pollen has both sperm (SN) and
vegetative (VN) nucleus but lacks exine on the walls.
E. Fruits of P. australis.
F. Seeds of Poszdonza (left: P. ostenfeldi1; centre: P. sinuosa; right: P. australis).
G. Three germinating stages in P. australis. Note that there are no roots in the seedlings.
H. Six-months old seedling of P. ostenfeldi. Note it bears many leaves and roots, however, the seed (arrow)
appear still intact.
I. A young (viviparous) seedling of Amphibolis antarctica showing a cup-shaped modified pericarp (arrow)
attached to the stem of the parent plant.
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
240 BIOLOGY OF AUSTRALIAN SEAGRASSES
An excellent study on rhizome branching systems in seagrasses was made by
Tomlinson (1974). He found rhizome branching in some genera (e.g. Amphibolis,
Thalassodendron, Heterozostera) is sympodial and in others (e.g. Posidonia, Zostera,
Cymodocea, etc.) is monopodial. The branching sequence may be (a) continuous (i.e.
branch at every node, Syringodium, Halophila); (b) diffuse (a.e. branches produced at
irregular intervals, Postdonta, Zostera, etc.) or (c) periodic (i.e. branches produced at
regular intervals (Thalassza). In addition to relative uniformity of the marine en-
vironment, the success of vegetative growth rather than propagation from seed in
seagrasses could result in a low frequency of sexual recombination and, consequently,
in a slow rate of evolutionary change. It is possibly for these reasons that Poszdonia
oceanica (L.) Delile from the Mediterranean is similar to P. australis of Australia, and
Thalassia testudinum of the Caribbean 1s closely related to 7. hemprichi of tropical Indo-
Pacific regions. The species in both cases appear to have diverged relatively little since
they were originally separated from one another in the Miocene (den Hartog, 1970).
PHYLLOSPHERE AND RHIZOSPHERE
Phyllosphere. Seagrass leaves bear varying numbers of epiphytes, depending upon age,
season and environmental conditions. The epiphytes include bacteria, diatoms, algae
(Fig. 8, A-E), and animals or ‘zoo-epiphytes’ such as hydroids (Fig. 8, F), sponges,
etc. In general the older portions of the leaf support more epiphytes, and sometimes
epiphytes may occlude half of the leaf area of a seagrass standing crop (Fig. 8, I). Even
on the surface of newly emerged leaf blades there are numerous bacteria and diatoms
(Fig. 8, A, B). As far as the abundance of epiphytic algae in Australian seagrasses is
concerned, more than twenty species have been found on Poszdonia australis from
Kangaroo Island, South Australia (Womersley, 1956); more than fifty species were
found on Zostera capricornt Aschers., Heterozostera tasmanica (Martens ex Aschers.) den
Hartog, and P. australis in Botany Bay and Jervis Bay, New South Wales (May et al. ,
1978); Ducker et al. (1977) have recorded more than one hundred species of epiphytic
algae on Amphibolis around Australia. The level of salinity may affect the growth of
epiphytes. Den Hartog (1970) noted that P. australis from the high salinity areas such as
the sea inlets in the Sydney area and Shark Bay in Western Australia have considerably
fewer epiphytes than P. australis from an open sea.
The attachment of epiphytes to the host appears to be limited to the cuticle of the
host (Fig. 8, G) though Ducker and Knox (1978) have observed the red alga Heteroderma
cymodoceae to erode locally the cuticle and outer wall of the epidermal cells of Amphibolis
antarctica stems.
Epiphytes could reduce seagrass photosynthetic rates by acting both as a barrier to
carbon dioxide uptake and by reducing light intensity (Sand-Jensen, 1977; Cam-
bridge, 1979). On the other hand, seagrasses may be able to reduce their epiphyte
stands by producing new photosynthetic tissues, or perhaps by excreting toxic sub-
stances (Sand-Jensen, 1977).
The transfer of nutrients between epiphytic algae and Thalassia testudinum has been
demonstrated (Goering and Parker, 1972; Harlin, 1973; McRoy and Goering, 1974).
Rhizosphere. The roots of seagrasses could also provide an environment for micro-
organisms. The micro-organisms in the rhizosphere may play an important role in
aiding nutrient uptake, affecting pathogen invasion and in nitrogen fixation.
Numerous bacteria and other micro-organisms have been found in the rhizospheres of
several seagrass species (Kuo et al., 1981; Fig. 8, H), and appear to penetrate only the
periphery of the host tissue. However, it is still not known whether those micro-
organisms are able to fix nitrogen. Patriquin (1972) hypothesized that bacterial
PROC. LINN. SOc. N.S.W. 106 (3), (1982) 1983
J. KUO 241
eS
ba
‘oe
Fig. 8. Epiphytes and rhizosphere in seagrasses.
A-D. Scanning electron micrographs show various epiphytes associated with the leaf blade of P. ostenfeldit.
A, numerous bacteria; B, bacteria and diatoms; C, bacteria, diatoms and calcareous red algae; D, bacteria,
diatoms and unknown organisms had damaged the epidermal surface.
E. A fungal hyphae-like structure on the leaf blade of P. australis.
G. A transverse section of P. sinuosa leaf blade showing numerous epiphytes are present on the leaf surface
(V: vascular bundles; E: epidermal cells).
H. Electron micrograph of P. australis rhizosphere showing some bacteria located on the outer surface of root
epidermal cells (E).
I. Numerous algae epiphytes (Ep) in association with the leaf blades of a P. australis plant.
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
242 BIOLOGY OF AUSTRALIAN SEAGRASSES
nitrogen fixation probably operated in seagrass rhizospheres, and Patriquin and
Knowles (1972) provided experimental support. However, McRoy et al. (1973) could
barely detect C.H, reduction associated with sediments, roots, rhizomes or leaves of
Thalassia testudinum. Capone et al. (1979) found that dead leaves of T. testudinum have
only one third of the nitrogen concentration of green leaves, and they estimated
bacterial nitrogen fixation in the sediments supplied up to a quarter to half of the
nitrogen required for leaf production.
CONCLUDING REMARKS
Seagrasses are abundant in Australian waters and many of them are endemic.
Investigation of the biology of Australian seagrasses is still in an early stage. Natural
habitats of seagrasses should be more carefully studied. Seagrasses have yet to be
collected from the north-west of this continent to map Australian species completely. A
critical but comprehensive taxonomic study of Australian seagrasses, particularly in
the genera Halophila, Posidonia and Zostera is urgently needed. The storage of seagrass
specimens in liquid as well as dehydrated states in herbaria is highly desired. Some
structures of seagrasses are similar to those in terrestrial plants, while others are
specialized adaptations in a marine environment. The development of various
vegetative and reproductive structures as well as germination in seagrasses is not well
understood. A combined biochemical, physiological and structural study on seagrasses
in relation to nutrient uptake, photosynthesis, translocation and transportation is
required. The relationship between epiphytes, zoo-epiphytes, rhizosphere and
seagrasses also requires more attention.
ACKNOWLEDGEMENTS
I acknowledge with appreciation the continuing fruitful discussions with Dr M. L.
Cambridge and many encouragements and interest in the study by Dr A. W. D.
Larkum. Dr L. K. Abbott, Dr A. W. D. Larkum and Professor A. J. McComb made
helpful comments on the manuscript. Special thanks go to Drs K. Abel, J. W. An-
drews, M. L. Cambridge, H. Kirkman, S. V. Lewis, E. Nazaro and J. Stewart for
their supply of seagrass specimens. Many thanks to Mr S. Blackburn and Mr G. Lane
for their photographic assistance and to Mrs M. Sedlak for her patience and skill typing
many drafts of this manuscript. This study was supported by funding from the
Australian Research Grants Committee and the Australian Marine Sciences and
Technology Advisory Committee.
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Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
The ‘Taxonomy of the semi-communal Spiders
commonly referred to the Species Jxeutzcus
candidus (1. Koch) with Notes on the Genera
Phryganoporus, Ixeuticus and Badumna (Araneae,
Amaurobioidea)
M. R. GRAY
Gray, M. R. The taxonomy of the semi-communal spiders commonly referred to the
species Ixeuticus candidus (LL. Koch) with notes on the genera Phryganoporus,
Ixeuticus and Badumna (Araneae, Amaurobioidea). Proc. Linn. Soc. N.S.W. 106
(3), (1982) 1983: 247-261.
Phryganoporus Simon 1908 is synonymized with Badumna Thorell 1890. The
synonymy of Jxeutzcus Dalmas 1917 with Badumna is supported. Spiders previously
placed in the genus Phryganoporus or the species J. candidus (L. Koch) are referred to
either B. candida (L. Koch), B. gausapata (Simon) n. comb. or B. vandiemeni n. sp.;
these three species form the candida species group of the genus Badumna. Biological
notes are given.
M. R. Gray, Department of Arachnology, The Australian Museum, P.O. Box A285, Sydney
South, Australia 2001; manuscript recewed I September 1981, accepted for publication in revised
form 17 February 1982.
INTRODUCTION
Several authors have considered the taxonomic status of the semi-social
amaurobioid spiders described by L. Koch (1872) as Amaurobius candidus from
Queensland and Simon (1908) as the genus Phryganoporus from Victoria and Western
_ Australia. However, at present these spiders are usually referred to the genus Jxeuticus
as a single species, J. candidus (L. Koch). They are widely distributed in open forest and
woodland habitats where their communal webs are usually built in low tree or shrub
foliage. Occasionally, heavy infestations occur in orchards or shrubland pastures; the
extensive webbing can severely inhibit foliage growth and prevent grazing.
TAXONOMIC BACKGROUND
Three species were originally recognized within Phryganoporus Simon 1908: P.
gausapatus (Simon 1906) from Victoria (originally described in Amaurobius C. Koch)
with a subspecies, P. g. occidentalis Simon 1908 from Western Australia; P. nigrinus
Simon 1908 and P. tubicola Simon 1908 from Western Australia. Amaurobius candidus L.
Koch 1872 had previously been described from Queensland and was later transferred
to Ixeuticus Dalmas 1917 by Roewer (1954). This placement has been followed by most
subsequent authors (McKeown, 1963; Dondale, 1966; Hickman, 1967; Main, 1971,
1976). Dondale (1966) redescribed J. candidus but the description and figures given
were of a related species J. martius (Simon 1899). As a result of this confusion Leech
(1971, 1972) suggested that J. martius may be a junior synonym of J. candidus. In fact, as
previously noted by Lehtinen (1967), J. martius is a junior synonym of Amaurobius
longinquus (L. Koch 1872) also referred to Ixeuticus by Dalmas (1917).
Lehtinen (1967) retained the genus Phryganoporus to which he transferred J. can-
didus to create the new combination P. candidus (L. Koch). Lehtinen also synonymized
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
248 SEMI-COMMUNAL SPIDERS
two of Simon’s species, P. gausapatus and P. nigrinus, with P. candidus but retained P.
tubicola Simon.
Subsequently, Main (1971) synonymized all of the species formerly described in
Phryganoporus into a single widely distributed species, J. candidus.
THE GENERIC PROBLEM
Lehtinen (1967) synonymized Ixeuticus Dalmas 1917 (type species A. martius
Simon 1899 described from New Zealand but probably introduced from eastern
Australia) with Badumna Thorell 1890 (type species B. Azrsuta Thorell 1890 from Java,
Indonesia). Subsequent authors have not followed Lehtinen (Forster, 1970; Main,
1971, 1976; Leech, 1971, 1972). Only Leech (1972) supported his rejection of the
synonymy, arguing first that J. martius lacks functional tarsal claws on the female palps
whereas B. hirsuta has definite, pectinate palpal claws as noted by Thorell (1890) and
second that the anterior median eyes of his J. martius were largest whereas Thorell
indicated that the anterior and posterior median eyes of B. hirsuta were subequal.
However, all females of J. longinquus (syn. I. martius) examined by me possess well-
developed palpal tarsal claws with seven to ten pectinations; and while the A.M.E.
were usually the largest, this was not the case in all of the specimens examined or in
other species of Jxeuticus. Kulczynski (1908) provides an excellent description and
figures of B. hirsuta. These indicate clearly that the genitalic characteristics (male palp
and female epigynum) of this species are extremely similar to those of J. longinquus.
Consequently, Lehtinen’s synonymy of Jxeuticus with Badumna seems entirely valid.
One result of this has been a change in the name of a very common and widely
distributed species, the black house spider. Previously Jxeutscus robustus (L. Koch),
Lehtinen (1967) synonymized it with Amaurobius insignis L. Koch 1872, so creating the
new combination Badumna insignis (L. Koch).
Main (1971) noted the inadequacy of the characters used by Simon (1908) as a
basis for maintaining the separation of Phryganoporus and Ixeuticus. By placing all species
of Phryganoporus into I. candidus Main effectively synonymized the two genera but
retained the junior synonym, Jxeutzcus. ‘The position of Badumna was not considered
here.
In contrast, Lehtinen (1967) retained Phryganoporus while recognizing the
synonymy of Jxeuticus and Badumna. His criteria for the retention of Phryganoporus relate
mainly to spination and the structure of the male palp. However, specimens examined
by me do not support the spination differences cited; representatives of both
Phryganoporus and Badumna (sensu Lehtinen) have a rather constant pattern of ventral
metatarsal spination of 221, whereas ventral tibial spination varies widely in
Phryganoporus (010 to 222) and is not an adequate generic character.
The male palpal tibia of Phryganoporus is stated by Lehtinen to possess a single
basodorsal ( = retrodorsal basal) process. However, there is also a retroventral process
which is equivalent to that present in Badumna species. Two or three retrolateral to
retrodorsal palpal tibial processes are commonly present in Badumna. These may be
placed apically as in B. longinquus or basally as in B. inornata Simon. Lehtinen also cites
the basally protruding tegulum of the Phryganoporus male palp as a generic character;
however, this feature simply represents the accentuation of a character trend already
apparent in Badumna species. Unlike Badumna, Phryganoporus may possess a patellar
process but this is not significantly developed in all species.
Similarities in male palpal morphology between Phryganoporus and Badumna are
readily apparent. Both possess an S-shaped, spiniform embolus, a distally tapering,
marginally folded conductor and a spoon-shaped, membranous median apophysis.
The female genitalia also share a common pattern consisting of an anterior fossa
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
M.R.GRAY 249
bounded posteriorly by a prominent, transverse ridge with lateral teeth present. The
internal genitalia of Phryganoporus species show a consistent pattern that is very similar
to that of B. longinquus (Fig. 20).
Both genera share a similar tarsal organ and trichobothrial plate morphology (Figs
23-28). Both also possess a complex, strongly branched tracheal system confined to the
abdomen. This presents a marked contrast with another related, but distinct genus,
Forsterina Lehtinen, which possesses a simple (unbranched) tracheal system. The latter
finding is of wider interest because it suggests that for the Australian fauna this
character may be usable at generic level only, whereas it has been used at the super-
family level by Forster (1970) and Forster and Wilton (1973) to separate their Dic-
tynioidea and Amaurobioidea. One anomaly evident from this is their placement of the
related genera Jxeuticus and Reinga Forster, a close relative of Forsterina, into different
superfamilies.
In summary, the features used by Lehtinen to maintain Phryganoporus seem
inadequate to justify its separation from the large and rather variable genus Badumna.
The only unequivocal character at present available to Phryganoporus is its possession of
a single retrodorsal tibial process on the male palp compared to two or three in
Badumna. However, the tibial processes seem to be rather labile characters, both in
shape and number, in these spiders. Behavioural traits may be valid generic characters
but the social behaviour shown by members of Phryganoporus does not seem of special
significance — social tendencies are apparent in members of Badumna also e.g. B.
socialis (Rainbow).
Consequently, I think ‘it justifiable to consider Phryganoporus Simon, like Ixeuticus,
to be a junior synonym of Badumna Thorell. A generic revision of Badumna would
certainly require the delineation of either subgeneric or species group categories. Here,
the spiders formerly placed within Jxeutecus candidus (L. Koch) or Phryganoporus Simon
are regarded as forming the candida species group within the genus Badumna.
Badumna Thorell
Badumna Thorell 1890: 322.
Type species: Badumna hirsuta Thorell 1890.
Phryganoporus Simon 1908. N. syn.
Type species: Amaurobius gausapatus Simon 1906
Ixeuticus Dalmas 1917.
Type species: Amaurobius martius Simon 1899
Badumna candida species group
Three species are recognized here: B. candida (L. Koch), a widespread, variable
species found in Queensland, New South Wales and South and Western Australia. B.
gausapata (Simon) from southeastern Australia; and B. vandiement n. sp. from
Tasmania.
Diagnosis
Medium sized (carapace length 2.4 to 4.1 mm) cribellate spiders which live in
both communal and solitary webs. Carapace silvery brown in colour, white hairs
abundant. Abdomen light brown with a dark brown mid-dorsal stripe followed by
several light to dark brown chevron markings with white hair tufts laterally. Legs
banded brown and grey. Anterior median eyes or anterior lateral eyes largest.
Cheliceral teeth, retrolateral 2-4, prolateral 3-5. Cymbium large, broad. Embolus of
male palp sinuously curved (S-shaped), proximal part of tegulum strongly protuberant
basally. Male palpal tibia with a retrodorsal basal and a retrolateral ventral process;
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
250 SEMI-COMMUNAL SPIDERS
dorsal patellar process well developed or rudimentary to absent. Epigynum with a
prominent, subdistal, transverse ridge posterior to an unpaired fossa; lateral teeth
distal to subdistal. Tracheal system complex, confined to abdomen.
Repositories: Australian Museum (A.M.); Queensland Museum (Q.M.); Tasmanian
Museum and Art Gallery (T.M.); Zoologische Museum, Hamburg (Z.M.H.);
Museum National d’Histoire Naturelle, Paris (M.N.H.N.); Australian National
Insect Collection (A.N.I.C.).
Badumna candida (1. Koch), new comb. Figs 1-11, 23, 35-37
Amaurobius candidus L. Koch 1872
Phryganoporus gausapatus occidentalis Simon 1908
Phryganoporus nigrinus Simon 1908
Phryganoporus tubicola Simon 1908; Lehtinen 1967
Ixeuticus candidus Roewer 1954; Main 1971
Phryganoporus candidus Lehtinen 1967
Diagnosis
Patellar process on male palp rudimentary to absent; median apophysis directed
apico-laterally. Lateral teeth of epigynum distal; fossa widest in central to posterior
half. Cheliceral teeth, retrolateral 2-4 and prolateral 3-5. Ventral tibial spination, first
leg 010-222. Metatarsal trichobothria; first leg 4-5.
MALE (S 144, O.M.)
Measurements (mm): Body length 5.68. Carapace length 2.58, width 1.90. Abdomen
length 3.10, width 2.11.
Colour: Carapace silvery brownish-grey with numerous white hairs; brown patches
lateral and posterior to the fovea. Chelicerae and sternum dark brown, the sternum
with dark brown hairs only. Legs with silvery grey and brown bands; ventral surfaces
of coxae with white hairs. Abdomen light brown with a broad, dark brown mid-dorsal
stripe, paler centrally and less than half as long as abdomen, bordered by lateral
patches of white hairs. Immediately posterior to this stripe is a light brown patch partly
delimited anterolaterally and posterolaterally by four dark brown spots. Behind this is a
row of five to six dark brown chevron markings, paler brown centrally, which are
separated from each other by thin lines of white hairs which form white hair tufts
laterally. Dark brown flecks are present on the lateral abdomen particularly
lateroventrally where they form a more or less distinct longitudinal dark line. This is
separated on each side by a moderately broad line of white pigment from a broad, dark
brown midventral stripe running between the epigastric fold and the spinnerets.
Carapace: Longer than wide in ratio 1:0.74. Clypeus height about 1.5 times the
diameter of an A.M.E. Cephalic area well developed, fovea a narrow slit.
Eyes: A.L.E. > P.L.E. > A.M.E. > P.M.E. in ratio 1:0.86:0.73:0.71. Interdistance
ratios, A.M.E. - A.M.E. 0.50: A.M.E. - A.L.E. 0.69: A.L.E. - P.L.E. 0.27: P.L.E.
Figs 1-11. Badumna candida. 1-3 male palp; 1, retrolateral; 2, ventral; 3, prolateral. 4-7, female genitalia: 4,
epigynum, 9, internal genitalia (Girraween N.P., Qld); 6, epigynum, 7, internal genitalia (Brookton,
W.A.). 8, female, dorsal. 9, male abdomen, dorsal (dark colour morph). 10-11, male palp (TYPE of P.
tubicola): 10, ventral: 11, tibia and patella, retrolateral. Scale lines 0.2 mm.
PROG. LINN. Soc. N.S.W. 106 (3), (1982) 1983
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
252 SEMI-COMMUNAL SPIDERS
- P.M.E. 1:P.M.E. - P.M.E. 0.92. M.O.Q. length, anterior width, posterior width
ratio 0.97: 0.81:1. Lateral eyes slightly protuberant. From above, anterior eye row
slightly recurved, width 0.88 mm; posterior eye row slightly procurved, width 0.96
mm. The A.L.E., P.L.E. and P.M.E. all have broad, band-like tapeta, diffuse in
A.M.E.
Chelicerae: Boss present. Fang groove with 2-3 teeth on retromargin, 3-4 on promargin.
Mazxillae: Subparallel, slightly convergent, twice as long as wide.
Labium: Wider than long in ratio 1:0.79. Widest subbasally, shallowly notched apically
and basolaterally.
Sternum: Cordate, shortly pointed posteriorly; longer than wide in ratio 1:0.87.
Male palp: Cymbium short and broad, bulb large. Tegulum and proximal embolus
protrude strongly basally on the prolateral side. Embolus a large, sinuous, S-shaped
spine supported by a similarly sinuous, folded membranous conductor, both ending
retrolateral-ventral to the apex of the cymbium. Median apophysis membranous,
broad, spoon-shaped and, in ventral view, directed apicolaterally. Tibia with a blunt
retrolateral-ventral process directed ventrally and a pointed retrodorsal basal process
directed retro-dorsally. Patellar process indistinct to absent.
Legs: 1243. Spination: Leg 1, femur p 011, d 112, tibia p 11, r 11, v 122 or 022,
metatarsus p 11 or 12, r 11, d 02 or 12, v 221; leg 2, femur p 011, d 112, tibia p 11, r
11, d 012, metatarsus p 11, r11,d 12, v 221; leg 3, femurd 113, tibiap 11, r11,v 012,
metatarsus d 11, r11, d 112, v 221; leg 4, femur d 112 or 113, tibia p 11, r 11, v 112,
metatarsus p 011, r 001, d 222, v221. Calamistrum weak. Tarsal claws: superior with
9-11 pectinations; inferior with 2-3 pectinations. Hairs ciliate. Trichobothria: single
row on tarsus and metatarsus; tarsus of first and second legs with 5, others with 3,
placed in central half to third; metatarsus of third leg with 5, remainder with 4, placed
in distal three quarters to half, or distal quarter on fourth leg. Bothria collariform with
fine, longitudinal striae on proximal plate.
Tracheal system: Complex, consisting of four strongly branched tubes confined to the
abdomen; spiracle of moderate width, as wide as cribellum.
Crbellum: Bipartite, spinning area reduced, strongly sclerotized posteriorly.
Spinnerets: Six, short. Anterior lateral pair broad, conical, basally approximated with a
very short distal segment: posterior lateral pair thinner and slightly longer, the distal
segment one-third of the total length.
FEMALE (S 144, O.M.)
Similar to male except as indicated below.
Measurements (mm): Body length 7.00. Carapace length 2.84, width 2.04. Abdomen
length 4.42, width 3.43.
Eyes: A.L.E. > A.M.E. = P.L.E. > P.M.E. in ratio 1:0.92:0.92:0.87. Interdistance
ratios, A.M.E. - A.M.E. 0.40: A.M.E. - A.L.E. 0.61: A.L.E. - P.L.E. 0.16: P.L.E.
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
M.R.GRAY 253
- P.M.E. 1: P.M.E. - P.M.E. 0.89. M.O.Q. length, anterior width, posterior width
ratio 0.99:0.78:1.
Chelicerae: Fang groove with 4 teeth on retromargin, 5 on promargin.
Labium: Wider than long in ratio 1:0.84.
Sternum: Longer than wide in ratio 1:0.81.
Palp: Tarsal claw with 8-9 pectinations.
Legs: 1243. Spination: leg 1, femur p 011, d 112, tibia p 11, r 11, v 122, metatarsus p
011, r011,d 012, v 221; leg 2, femur p 011, d 112, tibiap 11, r11, v 112, metatarsus p
011, r011,d 112, v 220 or 221; leg 3, femur d 133, tibia p 11, r 11, v 012, metatarsus p
011, r011,d 212, v 221; leg 4, femur d 112, tibiap 11, r11, v 112, metatarsus p 011, r
001, d 122, v 121 or 221. Calamistrum well developed and occupying the proximal to
central half of the metatarsus. Tarsal claws: superior with 9-11 pectinations; inferior
with 2-3 pectinations. Trichobothria (legs 1 to 4): tarsus 5, 4, 4, 4; metatarsus 5, 3, 4,
4. Bothria collariform with several poorly defined ridges curving medially from the
lateral margins of the proximal plate and converging upon its base. Tarsal organ an
oval opening situation at the distal side of a low, mound ornamented with a few in-
distinct semi-circular folds; tarsal organ mound poorly delimited and about three times
longer than opening.
Cribellum: Bipartite, sclerotized posteriorly and at median partition, spinning areas well
developed, spigots strobilate.
Genitalia: Epigynal fossa a rounded depression about as long as wide, widest centrally to
posteriorly. Fossa bounded posteriorly by a broad, transverse, chitinous ridge; anterior
margin of ridge indented. Lateral teeth distal to ridge. Internal genitalia with broad,
singly coiled seminal ducts; receptacula small and adjacent near mid line; a broad,
curved fertilization duct extends posteriorly.
Holotype female: Amaurobius candidus L. Koch 1872 from Bowen, Queensland, Australia.
Zoologische Museum, Hamburg. Araneae type cat. no. 11. Museum Godeffroyi
cat. no. 7852.
Material examined
HOLOTYPE female, Bowen, Qld (Mus. Godeff. 7852, Z.M.H.). Male and female
(S 144, O.M.), Southwood, 30km west of Moonie, Old, R. Raven; 24.8.1973; from
communal web. 2 males and 4 females (Q.M.), Girraween National Park, nr.
Stanthorpe, Qld. Male (KS 6941, A.M.), ‘Burnside’, near Margaret River, W.A., M.
Gray, 26.1.1979; taken as juvenile in solitary web. Female (KS 6938, A.M.), Torbay,
W.A., B.Y. Main 10.10.1977. Female (KS 6939, A.M.), Brookton, W.A., B.Y.
Main, 5.5.1977. Male (Ar 811, M.N.H.N.), TYPE of Phryganoporus tubtcola Simon,
Denham, W.A. Male (KS 6937, A.M.), Wanaaring, N.S.W., 28.3.1977; taken as
juvenile from communal web. Female (KS 6940, A.M.), 21km east of Parkes,
N.S.W., M. Gray, 8.4.1972; from solitary web. 6 males, 10 females (KS 5089, A.M.),
6km east of Dubbo, N.S.W., M. Gray, 21.8.1980; from communal web. 7 females, 1
male (KS 8663, A.M.), Kimba, Eyre Peninsula, S.A.
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
254 SEMI-COMMUNAL SPIDERS
Variation
Measurements (mm): Males: carapace length 2.44-2.88, width 1.65-2.10. Females:
carapace length 2.84-3.80, width 2.04-2.54.
Colour: Sternum with brown or brown and white hairs.
In addition to normally pigmented spiders a colour form with increased melanic
pigmentation occurs in southwest Australia. As adults these spiders have a dark brown
carapace and the light brown patch normally placed immediately behind the mid
dorsal abdominal stripe is replaced by a dark brown chevron marking (Fig. 9). The
lateral abdominal areas are silvery grey in colour. These spiders correspond well with
Simon’s P. nigrinus from Boyanup, W.A., here synonymized with B. candida. Sub-
specific status may prove appropriate for this distinctive colour morph.
Chelicerae: Retrolateral teeth 2-4, prolateral teeth 3-5.
Spination: Leg 1, ventral tibia 010-222.
Genitalia: Epigynal fossa as long as wide or wider than long; lateral margins evenly or
unevenly curved.
Badumna gausapata (Simon 1906), new comb. Figs 12-18, 26
Amaurobius gausapatus Simon 1906
Phryganoporus gausapatus Simon 1908
Phryganoporus candidus Lehtinen 1967
Txeuticus candidus Main 1971
Similar to B. candida and agreeing with the description given for that species
except as indicated below.
Diagnosis
Definite patellar process on male palp; median apophysis directed apico-
laterally. Lateral teeth of epigynum subdistal; fossa widest in central to posterior half.
Cheliceral teeth, retrolateral 2-3 and prolateral 5-6. Ventral tibial spination, first leg
112-222. Metatarsal trichobothria, first leg 5-6.
MALE (KS 6942, A.M.)
Measurements (mm): Body length 7.05. Carapace length 3.45, width 2.61. Abdomen
length 3.60, width 2.31.
Colour: As for B. candida. Sternum with brown and white hairs.
Carapace: Longer than wide in ratio of 1:0.76. Clypeus height equals 1.25 diameters of
an A.M.E.
Eyes: A.M.E,. > P.L.E. > A.L.E. > P.M.E. in ratio 1:0.98: 0.96: 0.83. In-
terdistance ratios, A.M.E. - A.M.E. 0.38: A.M.E. - A.L.E. 0.53: A.L.E. - P.L.E.
Figs 12-18. Badumna gausapata. 12-14, male palp: 12, retrolateral; 13, ventral; 14, prolateral. 15-17, female
genitalia (SYNTYPES): 15-16, epigyna; 17, internal genitalia. 18, female abdomen, dorsal.
Figs 19-22. Badumna longinquus. 19-20, female genitalia: 19, epigynum; 20, internal genitalia. 21-22, male
palp: 21, ventral; 22, tibia and patella, retrolateral. Scale lines 0.2 mm.
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
M.R.GRAY 255
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
256 SEMI-COMMUNAL SPIDERS
0.23: P.L.E. - P.M.E. 0.89: P.M.E. - P.M.E. 1. M.O.Q. length, anterior width,
posterior width ratio 0.98: 0.80:1. From above anterior eye row recurved, width 0.97
mm, posterior eye row slightly procurved, width 1.09 mm.
Chelicerae: Fang groove with 2 retromarginal teeth, 4 promarginal teeth.
Maxillae: Parallel, twice as long as wide.
Labium: Wider than long in ratio 1:0.80; surface convex.
Sternum: Longer than wide in ratio 1:0.79.
Male palp: Median apophysis membranous, moderately narrow and spoon-shaped; in
ventral view directed apico-laterally. Patellar process a short, bluntly pointed, finger-
like projection, placed dorsally.
Legs: 1243. Spination: Leg 1, femur p 011, d 112, tibia p 101, r 101, v 122, metatarsus
p 011, r101,d012, v 221: leg 2, femur p 001, d 213, tibia p 11, r 11, v 112, metatarsus
p 101, r101,d 012, v 221: leg 3, femur d 113, tibia p 11, r 11, v 012, metatarsus p 011,
r011,d 212, v 221: leg 4, femur d 113, tibia p 11, r 11 or 111, v 112 or 122, metatarsus
p 011, r 001, d 222, v 221. Calamistrum very weak. Trichobothria (legs 1 to 4): tarsus
5, 4, 4, 3: metatarsus 5, 4, 3, 3.
FEMALE (KS 6086, A.M.)
Similar to male except as indicated below.
Measurements (mm): Body length 7.60. Carapace length 3.38, width 2.21. Abdomen
length 4.45, width 2.95.
Colour: Sternum with brown hairs, white hairs absent.
Eyes: A.L.E. > P.L.E. > A.M.E. > P.M.E. in ratio 1:0.95: 0.86:0.85. Interdistance
ratios, A.M.E. - A.M.E. 0.36: A.M.E. - A.L.E. 0.48: A.L.E. - P.L.E. 0.22: P.L.E.
- P.M.E. 1:P.M.E. - P.M.E. 0.76. M.O.Q. length, anterior width, posterior width
ratio 1:0.81:1.
Chelicerae: Fang groove with 3 or 4 retromarginal teeth, 4 or 5 promarginal teeth.
Labium: Wider than long in ratio 1:0.75.
Sternum: Longer than wide in ratio 1:0.81.
Palp: Tarsal claw with 7-8 pectinations.
Legs: 1423. Spination: leg 1, femur p 011, d 122, tibia p 11, r 101, v 222, metatarsus p
011, r011,d 112, v 221; leg 2, femur p 0111, d 122, tibia p 11, r11, v 122, metatarsus
pii,ri11,d112, v 221; leg 3, femur d 133, tibia p 11, r 11, d 212, v 221; leg 4, d 112,
tibia p 11, r11, v 112, metatarsus p 111, r 001, d 1012, v 221. Trichobothria (legs 1 to
4): tarsus 6, 4, 4, 4; metatarsus 6, 5, 4, 4. Bothria collariform, surface of proximal plate
with narrow, semi-longitudinal ridges. Tarsal organ opening oval, placed near the
distal margin of a poorly defined, elongate mound approximately four times as long as
opening.
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
M.R. GRAY 257
Figs 23-25. Tarsal organs, leg 1: 23, B. candida (similar in B. gausapata); 24, B. vandiemeni; 25, B. longinquus.
Figs 26-28. Trichobothrial bases, tarsus, leg 1: 26, B. gausapata; 27, B. vandiement; (both types present in B.
candida) 28, B. longinquus. Scale lines 5 p.
Genitalia: Epigynal fossa wider than long, widest in central to posterior area, margins
smoothly curved. Lateral teeth subdistal. Anterior margin of transverse ridge not
indented.
Material examined
SYNTYPE females (AR 810, Paris), Victoria, 1903. 1 male (KS 6942, A.M.),
Canberra, A.C.T., M.S. Upton, 24.5.1965. 5 females (KS 6086, A.M.), Black
Mountain, Canberra, A.C.T., M. R. Gray, 1.10.1980. 4 males, 3 females (KS 6090,
A.M.), Black Mountain, Canberra, A.C.T., 3.6.1965. 3 males, 4 females (KS 6088
A.M.), Wee Jasper, N.S.W., M. Gray 30.10.1980. 4 males (A.N.I.C.) Black
Mountain, Canberra, A.C.T., I. F. B. Common, 4.6.1965 (from light trap).
Variation
Measurements (mm): Males; carapace length 3.05-3.48, width 2.19-2.61. Females:
carapace length 3.38-2.44, width 2.21-1.22.
Chelicerae: Retrolateral teeth 2-3, prolateral teeth 4-5.
Spination: Leg 1, ventral tibia 122-222.
Badumna vandiemeni n. sp. Figs 24, 27, 29-34
Ixeuticus candidus Hickman 1967
Similar to B. candida and agreeing with the description given for that species
except as indicated below.
Diagnosis
Definite patellar process on male palp; median apophysis directed laterally.
Lateral teeth of epigynum subdistal; fossa widest in anterior half. Mid-dorsal ab-
dominal colour pattern light brown posteriorly. Cheliceral teeth, retrolateral 2 and
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
258 SEMI-COMMUNAL SPIDERS
Figs 29-34. Badumna vandiemeni. 29, male abdomen, dorsal. 30-32, male palp: 30, retrolateral; 31, ventral;
32, prolateral. 33-34, female genitalia: 33, epigynum; 34, internal genitalia. Scale lines 0.2 mm.
prolateral 4-5. Ventral tibial spination, first leg 121-222. Metatarsal trichobothria, first
leg 7-9.
MALE (KS 6976, A.M.), Holotype
Measurements (mm): Body length 8.55. Carapace length 4.06, width 2.80. Abdomen
length 4.58, width 3.70.
Colour: Dorsal abdominal stripe fairly long, about half as long as abdomen. Posterior
chevrons indistinct, light brown except for small, lateral, dark brown patches of
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
M.R.GRAY 259
pigment. Sternum with dark brown and white hairs.
Carapace: Longer than wide in ratio of 1:0.69. Clypeus height 1.25 diameters of an
A.M.E.
yee eNO. > ACL. > PE. > P.M_E. in‘ ratio of 1:0°95:0.92:0.84. In-
terdistance ratios, A.M.E. - A.M.E. 0.39: A.M.E. - A.L.E. 0.42: A.L.E. - P.L.E.
0.20: P.L.E. - P.M.E. 1.00: P.M.E. - P.M.E. 0.90. M.O.Q. length, anterior width,
posterior width ratio 1:0.82:0.96. From above anterior eye row slightly recurved,
width 1.20 mm; posterior eye row slightly procurved, width 1.34 mm.
Chelicerae: Retrolateral teeth 2; prolateral teeth, 4-5.
Mazxillae: Subparallel, slightly convergent, twice as long as wide.
Labium: Wider than long in ratio 1:0.80.
Sternum: Longer than wide in ratio 1:0.81.
Male palp: Median apophysis moderately narrow and spoon-shaped: directed laterally,
almost horizontal in ventral view. Patellar process dorsaily placed, short, bluntly
pointed and adorned basally with white, spatulate hairs.
Legs: 1243. Spination: leg 1, femur p 011, d 112, tibia p 11, r 11, v 222, metatarsus p
OMe OlpecdhOO2 ve22 leos?* femur pO0ll or 0111 d t2) tibiap ir 11 122,
metatarsus p 101, r 101, d 002, v 222: leg 3, femur p 0111, d 113, tibiap 11, r11v 022:
metatarsus p 0101, r 0101, d 112, v 221: leg 4, femur d 113, tibia p 11, r 11, v 112,
metatarsus p 111, r 001, d 112, v 221. Calamistrum weak. Trichobothria (legs 1 to 4):
tarsus 6, 5, 5, 5: metatarsus 7, 6, 6, 5.
FEMALE (KS 6977, A.M.), Paratype
Similar to male except as indicated below.
Measurements (mm): Body length 8.60. Carapace length 3.69, width 2.48. Abdomen
length 4.95, width 3.50.
Colour: Dorsal abdominal stripe less than half as long as abdomen.
Carapace: Longer than wide in ratio 1:0.67.
Eyes: A.M.E. > A.L.E. > P.L.E. = P.M.E. in ratio 1:0.95:0.85:0.85. Interdistance
ratios, A.M. — A.M_E. 0.42; A.M.LE,-— A.1L.E. 0.71: A.L.E. - P.L.E. 0.23: P.L.E.
- P.M.E. 0.86. P.M.E. - P.M.E. 1.00 M.O.Q. length, anterior width, posterior
width ratio 1:0.73:0.99. Eye row width, anterior 1.20 mm; posterior 1.34 mm.
Chelicerae: Retrolateral teeth, 2; prolateral teeth, 4.
Labium: Wider than long in ratio 1:0.84.
Sternum: Longer than wide in ratio 1:0.85.
Legs: 1243. Spination: leg 1, femur p 0011, d 112, tibia p 11, r 11, v 121; metatarsus p
011, r 011, d 002, v 221; leg 2, femur p 0011, d 112, tibia p 11, r 11, v 121, metatarsus
p 011, r011, d 002, v 221; leg 3, femur d 113, tibia p 11, r11, v 012, metatarsus p 011,
r 011, d 002, v 221; leg 4, femur d 112, tibia p 11, r 11, v 012, metatarsus p 011, r
0011, d 002, v 121. Trichobothria (legs 1 to 4): tarsus 5, 5, 4, 5; metatarsus 9, 5, 7, 5.
Bothria collariform with several, well defined, semi-circular ridges on proximal plate.
Tarsal organ a small, oval opening, acutely pointed proximally, placed near the distal
margin of a well delimited, oval mound six times as long as opening; surface of mound
ornamented by fine striae.
Genitalia: Epigynal fossa wider than long, widest anteriorly, lateral margins sloping
inwards to subdistal lateral teeth. Transverse ridge not indented anteriorly.
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
SEMI-COMMUNAL SPIDERS
37
Figs 35-37. Badumna candida, webs. 35, communal web, Dubbo, N.S.W.; 36, solitary web retreat, Goonoo
S.F., N.S.W.; 37, solitary web retreat of juvenile male, dark colour form, Margaret River, W.A.
Types
Holotype Male — KS 6976 (A.M.). Eaglehawk Neck, Tas., M. R. Gray, 3.7.1980; from
solitary web on shrub (Acacia uriczfolta).
Paratypes — Female, KS 6977 (A.M.), same data as holotype. Female, KS 6978
(A.M.), Eaglehawk Neck, Tas., V. V. Hickman, 6.3.1960. 3 females, J.763 (T.M.),
Lauderdale, Tas., April 1971.
Variation
Measurements (mm): Females: carapace length 2.95-3.69; carapace width 1.97-2.40.
Spination: Leg 1, ventral tibia 121-222.
BIOLOGICAL NOTES
Spiders of the B. candida species group make both communal and solitary webs. As
noted by Main (1971) communal web populations are made up mainly of juveniles,
but adults are also often present; some, at least, may complete their life cycles within
the communal web. However, many leave to take up a solitary existence soon after
they mature. Late instar juveniles as well as adults are involved in such dispersal as
both subadult and mature spiders can be found in solitary webs.
The webs (Figs 35-37) are built among the foliage of various low, sclerophyllous
trees and shrubs. Solitary webs are small, the retreat usually being fastened along a
stem while the irregular cribellate snare extends a short way into the surrounding
foliage. Communal webs vary greatly in size and may encompass much of the foliage
of a shrub or branch. Up to 95 spiders have been recorded from a single large web.
Counts of penultimate juveniles in communal webs often showed marked
disproportions in sex ratios; whether this simply represents differential dispersal or
involves some other factors is not clear. Though their retreat tubes are independent of
each other, juvenile spiders will hunt and feed together on the same prey animal in the
shared catching part of the web.
Main (1976, fig. 40e) noted that the structure of the solitary web in south western
populations of B. candida consisted of a short, bag-like tube of silk fastened onto a
branch, the small sheet web radiating out from it. Similar solitary nests are built by
members of the candida species group in eastern Australia (Fig. 36), though sometimes
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
M.R.GRAY 261
their retreat tubes are more elongated. In southwestern Australia long, horn-like
retreats containing juvenile males of the dark colour form of B. candida (reared to
maturity in the laboratory) have been collected (Fig. 37).
One to three egg sacs can be found embedded in the silk-plant-food detritus wall
matrix of the female retreats. The sacs are circular to oval, flattened spheres with a
definite circumferential seam varying in diameter from 4 to 7 mm. The outer silk is
flocculent and attaches the sac closely to the retreat wall: inside this is a more finely
woven, thin layer of silk. The eggs are non-glutinous and vary from 0.6 to 0.8 mm in
diameter. Each sac contains from 13 to 49 eggs. Sacs were found in retreats from both
solitary and communal webs.
ACKNOWLEDGEMENTS
My thanks go to Dr M. Hubert, Dr B. Y. Main, Prof V. V. Hickman, Dr V.
Davies and Mr M. Upton for the loan of material. Inking and shading of drawings was
done by Ms P. Greer.
References
DALMAS, C. de, 1917. — Araignées de Nouvelle-Zélande. Ann. Soc. Ent. France 86: 317-430.
DONDALE, C. D., 1966. — The spider fauna (Araneida) of deciduous orchards in the Australian Capital
Territory. Aust. J. Zool. 14: 1157-92.
FORSTER, R. R., 1970. — The spiders of New Zealand, Pt. III. Otago Mus. Bull. 3: 1-184.
, and WILTON, C. L., 1973. — The spiders of New Zealand, Pt. 1V. Otago Mus. Bull. 4: 1-309.
HICKMAN, V. V., 1967. — Some common spiders of Tasmania. Hobart: Tasmanian Museum and Art Gallery.
KULCZYNSKI, V., 1908. — Symbola and faunam aranearum Javae at Sumatrae cognoscendam. I.
Mygalomorphae et Cribellatae. Bull. Acad. Sci. Cracov. 1908: 527-581.
Koch, L., 1872. — Die Arachniden Australiens. Nuremburg: Ludwig Korn.
LEECH, R., 1971. — The introduced Amaurobiidae-of North America and Callobius hokkatdo n.sp. from
Japan (Arachnida:Araneida), Canad. Ent. 103: 23-32.
, 1972. — A revision of the Nearctic Amaurobiidae. Mem. Ent. Soc. Can. 84: 1-182.
LEHTINEN, P. T., 1967. — Classification of the Cribellate Spiders and some allied families with notes on the
evolution of the suborder Araneomorphae. Ann. Zool. Fenn. 4: 199-468.
MAIN, B. Y., 1964, 1967. — Spiders of Australia, (2 editions). Brisbane: Jacaranda Press.
1971. — The common ‘colonial’ spider Ixeuticus candidus (Koch) and its synonyms (Dic-
tynidae: Araneae). J. Roy. Soc. W.A. 54 (4): 119-120.
, 1976. — Spiders. Sydney: Collins. Australian Naturalist Library.
McKeown, K. C., 1963. — Australian spiders. Sydney: Angus and Robertson.
Roewer, C. F., 1942-54. — Katalog der Araneae, 2 vois. Bremen: Wissenschaften Paul Budy.
SIMON, E., 1899. — Ergebnisse einer Reise nach dem Pacific. Arachnoideen. Zool Jahrb., Syst. 12: 411-437.
SIMON, E., 1906. — Etude sur les Araignées de la section des Cribellates. Ann. Soc. Ent. Belg. 50: 284-308.
, 1908. — In Die Fauna Siidwest Australiens 1 (12): 359-446 (W. MICHAELSEN AND R. HARTMEYER
(eds) ). Jena: Gustav Fischer.
THORELL, T., 1890. — Studi sui Ragni malesie papuani. [V. Ann. Mus. Civ. St. Nat. Genova 28: 1-419.
?
Proc. LINN. Soc. N.S.W. 106 (3), (1982) 1983
PROCEEDINGS
of the
LINNEAN
SOCIETY
NEW SOUTH WALES
VOLUME 106
NUMBER 4
THE SIR WILLIAM MACLEAY MEMORIAL LECTURE 1982
Ecological Values of the Tropical Rainforest
Resource
L. J. WEBB
Honorary Fellow, School of Australian
Environmental Studies, Griffith University,
Nathan, Brisbane, Australia 4111*
[Delivered 8 September 1982]
‘Tf we knew enough no choice would be trivial, and it is our duty to acquire the knowledge which will
enable us to moralize our everyday actions, both by the study of available statistics and by encouraging
statistical inquiry elsewhere’ (Haldane, 1928).
In medieval times Wisdom was separated into Metaphysics, Morals, and Natural
Philosophy which was equivalent to natural science. In the seventeenth century,
Newton along with Galileo and Descartes established the prestige and paramountcy of
mathematics and physics, and the domination of scientific thinking was well under
way. The term ‘natural philosophy’ became restricted to the exact physical sciences,
which were joined by chemistry at the beginning of the 19th century. It remained for
Playfair in 1812 to identify Natural History simply as a descriptive and non-
experimental science that dealt with vegetable, animal, and mineral objects (Pantin,
1968). Much later, natural history became known as the biological and earth sciences,
parts of which became increasingly quantitative, and involved with finer grades of
structures and molecular events. In this hierarchical re-arrangement of the sciences,
which emphasized precision and experimentation, the so-called descriptive biological
sciences stayed at the bottom.
Thus within a century or two of western scientific progress, biology and the study
of organism-environment relationships that was to become known as ecology lost their
early links with morals and metaphysics, became ‘poor relations’ of the exact sciences,
and were obliged to renounce intrinsic human values in the promotion of objective
scientific knowledge.
It is relevant that William Macleay, that great Australian naturalist whose
memory we celebrate tonight, identified the Linnean Society of New South Wales,
when it was founded just over a century ago, as a ‘society of natural history’ that he
hoped would succeed. It was founded not only ‘to promote knowledge’, but also ‘for
the progress of this community, and for the welfare of humanity’. Even then, it seems,
the extraordinarily broad interests of early naturalists such as Macleay enabled them to
foreshadow the necessary integration of the social and natural sciences — an in-
tegration that only now, in international projects such as the Man and the Biosphere
programme, is being hesitantly and not too effectively approached.
While admiring what Sir Otto Frankel (1970) called Sir William Macleay’s
‘remarkable prescience’, we may wonder what Sir William would have thought about
* Formerly, Rainforest Ecology Unit, Division of Plant Industry, CSIRO Laboratories, Indooroopilly,
Australia 4068.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
264 ECOLOGICAL VALUES OF THE TROPICAL RAINFOREST RESOURCE
the future losses to human welfare when biological and physical sciences became so
rigidly separated, when science became so thoroughly insulated from ethics, and when
there was nothing left but utter disdain for metaphysics.
It is more helpful for the biologist today, who retains something of his natural
history heritage, to understand the fundamental contrast and separation as not be-
tween biological and physical sciences but, as Pantin (1968) put it, ‘between
unrestricted and restricted sciences’. Thus the ecologist who is obliged to follow the
analysis of problems into many other sciences, and even non-sciences, generally must
rely on ‘natural experiments’; and to gain a broad and integrative view must sacrifice
exactness.
Yet ecological observation is not mere description, because perception depends on
experience, training, and current concepts. It is hard to accept that observations can be
strictly neutral when research directions are selected beforehand, priorities are set by
economic standards, and the presentation and interpretation of data are coloured by
the requirements of a particular development. In other words, it may not always be
possible to separate questions of fact from questions of value. If we are honest about it,
I suggest that we all can find some favourite examples.
It has become clear to me that decisions about the use and management of natural
resources are unavoidably influenced by value judgements. Questions of fact and
questions of value become inseparable in the broad issues of ‘ecological problems’, as
distinct from narrowly-circumscribed technical ‘problems in ecology’ (Passmore, 1974;
Ashby, 1978). The ecologist as an unrestricted scientist is therefore continually im-
plicated with the ‘is’ of fact, and the ‘ought’ of valuation that is not intrinsic to his
study and rests on an ethical system of some kind.
I wonder what Macleay would have done, had he, with his broad humanist and
what we may call his wide ecological interests, been confronted as we are now by this
thorny path to applied ecology, to the resolution of choices impelled by knowledge of
ecological consequences, and to dawning insights about how certain resources have
more values than utilitarian.
In this lecture I shall try to identify some ecological values from a knowledge of
rainforests. It seems that understanding how valuable a resource really is must await
the accumulation of enough facts and experience: then crystallization occurs to reveal
unsuspected facets. First of all I give an international perspective to ecological
problems encountered in Australian tropical and subtropical rainforests. Then I briefly
examine the Australian forests: their history and interactions with other vegetation and
with man, which in turn provide a basis of comparison with rainforests of our northern
neighbours. Finally I return to the question of values — ecological and beyond — that
the rainforests project. The colligation of facts and intrinsic values prompts the
identification of fresh moral choices, and suggests new responsibilities for ecology that
has now become ‘the genesis of a science of man and nature’ (di Castri, 1981).
About the time that Macleay and T. H. Huxley met in Sydney in the middle of
last century, the world’s population had reached its first billion. Nowadays it takes only
14 years to add another billion. Of the people who will be added to the world’s
population by the end of the century, about 90 per cent (about 1.5 billion) will be in the
tropics, i.e. in the less developed, poor, third world countries of Asia, Africa, and Latin
America. Already, in the third world, about 2 billion of the rural poor, in their struggle
to survive, exhaust the soils, deplete the forests, destroy wildlife, and pollute the
waters.
As the direct and indirect result of technological developments and the ‘pullulating
millions’, the world’s tropical and subtropical moist forests are disappearing at an
accelerating rate. The rates of conversion vary from country to country, and among
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
L. J. WEBB 265
different land systems within a particular zone, so that average rates may be quite
misleading. They have been heavily dramatized, with estimates varying from 20 to 40
hectares per minute (Myers, 1980). Unfortunately, most of the forests that have so far
been cleared are the moister complex types with the greatest array of genetic resources
at lower altitudes. Jacobs (1978a) illustrates this for Malaysia by noting that there are 9
different types of dipterocarp forest below 300 m, as opposed to 6 types between 300
and 700 m, and only 3 types between 700 m and 1200 m altitude. In the tropical humid
region of North Queensland, this is more pronounced, as there are about 25 main
structural types of rainforest below 800 m altitude but only about 8 main structural
types in the 800-1600 m altitudinal zone. These facts indicate that proportionately
more diversity is lost in converting lowland rainforest types.
Of the estimated 3 million species of plants and animals in the tropics, only about
half-a-million have scientific names. It has been predicted that, at the present rate of
destruction of the tropical and subtropical moist forests throughout the world, one-half
to one million species of plants and animals will become extinct. Thus to retain as
much natural or near-natural tropical forests as possible usually means ‘conserving the
unknown’ (MAB Ecology in Action Exhibit, Paris, 1981). In stressing that less than
half of the world’s predicted threatened plants are known, we must realize that rare
plants are no longer ‘the playchild of a botanical elite’. In countries such as India,
where there is so much reliance on the wild plant resource, plant conservation is an
essential part of economic development (Synge, 1982).
Most people, especially biologists, would agree that species extinctions are a bad
thing, although people’s reasons vary. This is, however, a moral judgment. Since
morals are the practice of ethics, and ethics is the science of morals, any opinion and
action about species extinction must have ethical roots. Such judgment, which is freely
made by biological scientists and some others, is therefore worth examining in a little
more detail. The MAB Ecology in Action Exhibit in Paris (1981) claims that con-
serving both the known and the unknown is not just a question of altruism but of
human survival, and proposes three justifications: ‘to preserve the genetic information
lost through species’ extinctions; to keep open the options for facing our future
requirements; and to conserve a natural diverse environment which will satisfy us
morally and aesthetically’. This seems a fair proposition, and far from extremist.
While acknowledging expediency and the necessities of practical life, it recognizes that
less tangible values are of importance, and second to none.
There is thus a spectrum of ecological values for the conservation of nature, with
pragmatism and self-interest at one end and the rights of nature apart from man at the
other.
Myers (1978), while claiming that ‘the problem of disappearing species could
eventually be seen as one of the greater ‘sleeper issues’ of the late 20th century’, neatly
summarized the utilitarian reasons for conserving animal and plant genetic resources
in tropical rainforests: for many particular uses in modern agriculture, in medicine and
pharmaceutics, and in industrial processes.
At the other end of the continuum there are quasi-religious reasons for the con-
servation of so-called ‘non-resources’ (Ehrenfeld, 1976). Non-resources are species and
biotic communities that do not have an economic value, or demonstrated potential
value for use by man here and now. Some people try to justify non-resources to make
them competitive with economic resources. For example, that recreational and
aesthetic values are essential for mental health; species diversity and polycultures are
less risky than monocultures; natural communities may provide clues to organization
that ensures survival; certain species may be sensitive environmental impact in-
dicators; natural ecosystems could serve as models for reconstructing degraded ones;
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
266 ECOLOGICAL VALUES OF THE TROPICAL RAINFOREST RESOURCE
intact ecosystems are needed for teaching purposes. All these examples suffer from
being too remote and nebulous to be convincing as political arguments. The ultimate
justification is rarely articulated: a deep-seated fear that irreversible changes which
reach a significant scale ‘may carry a hidden and unknowable risk of serious damage to
humans and their civilizations’ (Ehrenfeld, 1976). Homocentric as this fear is, it is
clearly impossible to assign resource values and priorities to non-resources that can
rank with, and excel, the economic values of resources as treasured by a totally ex-
ploitative society. That neither development nor preservation can be total, however,
gives grounds for believing that fresh value systems will continue to emerge.
Elton (1958) understood the spectrum of human attitudes towards co-existence
with nature as a series of ‘three absolute questions . . . waiting patiently to be an-
swered’ (which a quarter of a century later is still the case). He identified the questions
as religious, aesthetic and intellectual, and practical. This is how he put the least
scientific of these questions:
‘The first, which is not usually put first, is really religious. There are some millions of people in the
world who think that animals have a right to exist and to be left alone, or at any rate that they should
not be persecuted or made extinct as a species. Some people will believe this even when it is quite
dangerous to themselves’.
Examples of this belief are multiplying everywhere in the world. We witness them
locally as campaigns and demonstrations in various rainforest areas in eastern
Australia from north Queensland to south-west Tasmania.
In recent times, in October 1978, the tropical botanist Jacobs (1978b) taunted the
8th World Forestry Congress at Jakarta, the theme of which was ‘Forests for People’,
by a paper quoting the contrary title: “People for Forests’. Jacobs sensed a growing
feeling in industrialized Europe that power over nature is no longer what it used to be,
and predicted a change in forest policy towards mixed natural forests as well as, or in
places instead of, monocultures. Jacobs argued that ‘People for Forests’ would have
carried ‘a more dignified appeal’, and implied ‘a notion of respect, and respect befits
people’. He cited his colleague P. van Heynsbergen, an Amsterdam legal scholar, who
went further, and published in 1977 a ‘Declaration of the Rights of Animal and Plant
Life’ with eight paragraphs, the first of which reads that, in principle, ‘Each living
creature on earth has the right to exist, independent of its usefulness to humans’. This
idea is echoed by the current Charter of Nature to be considered by the United Nations
(Kamanda wa Kamanda, 1980).
These sentiments are no longer the eccentricities of a cultivated few in the so-
called First World, nor are they restricted to philosophers, poets, oriental sects, and
primitive Aboriginal society. It is worth considering Ashby’s (1978) claim that the most
effective defenders of nature today are not humanists, but scientists — mainly natural
scientists aided by a few social scientists. To be effective — in a wider campaign —
does require a certain scientific objectivity, because this is a basic prerequisite for
communication by scientific workers who are not gifted like poets in this art, and
because objective data can be tested by any other scientist. Scientific concern that
nature be valued for its intrinsic worth reflects Elton’s (1958) passion for conservation
of the greatest possible ecological variety, as well as contemporary ecological intuitions
about what interdependence in ecosystems and other holistic principles mean in the
evolution of the human condition.
Somehow, despite the ingenuity of systems ecologists, and the synecologists’
vision of ‘emergent properties’ at different levels of organization, holism as a working
principle used by western scientists has done little to help solve the urgent problems of
conservation and development in the developing countries of the tropics. Moreover,
even in the most developed country of all, Odum (1977) noted that controversy had
PRoc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
L. J. WEBB 267
clouded the ‘holistic strategy for ecosystem development’, and suggested that the
disappointing performance of the United States in the International Biological
Programme could be attributed to lack of ‘unifying themes or concepts’.
Yet a holistic frame of reference — the concept of environment in its totality, the
habit of ‘looking around corners’, the prediction of inevitable effects in space and time
of present actions — was being established slowly and firmly in widening sections of
society by the ideas, whatever their media, of ‘planet earth’, ‘the global village’, and so
on. Similarly but in a quite different context, spokesmen in the developing countries
(e.g. Salim, 1982; Soemarwoto, 1982) were beginning to urge the use of traditional
knowledge, and of holistic rather than analytical methods as a basis of ecological
guidelines for development through conservation. Earlier ecological guidelines
produced by the International Union for the Conservation of Nature and Natural
Resources (Poore, 1976) had generally been applied from ‘top to bottom’ and had had
little impact among the people.
Given this international and partly philosophical background, we can examine the
values of the tropical rainforest resource to the Australian community. We should note
that the definition of ‘resource’ is surprisingly wide, and includes computable wealth in
materials; something in reserve; something to which one has recourse in difficulty;
provision of relief or recovery and a means of spending leisure time. Passmore (1974)
went much further with the definition of ‘nature’, teasing out its meanings in which
man ranges from despot to primitivist, and noting its tangents with metaphysics and
sentiment.
Throughout the four decades of my studies in CSIRO I have been exposed to an
expanding spectrum of uses and ‘non-uses’ of rainforests for people. The facts of
science — science that traditionally makes disinterested inventories of ‘nature out
there’ — were only part of the spectrum. .The spectrum was always being coloured,
rainbow-like, through the prism of different values.
Always there were the covert and unacknowledged moral choices. But even more
than this seems at stake. What of those people who believe, who somehow intuitively
know, that ancient and prolonged existence in nature of Australian rainforest relicts
has value? That it confers a right for these remnants to continue to exist as objects in
nature? That this is now a new kind of ‘justification’ for not exterminating or
damaging any more of them?
Let us examine some examples of the evolution of the less tangible values of
rainforests that are not readily separated as scientific facts i.e. as evidence free from
inference, opinion, and advocacy, and that are therefore bound to involve moral
choices. This examination will require brief reference to some recent developments in
tropical rainforest botany and ecology in Australia.
At the outset it should be acknowledged that any ecologist studying physical and
biological processes in rainforests of the tropics and subtropics of the world cannot
exclude human ecological processes from his chosen ecosystem. Often he cannot see the
trees for the people; or in places like Peninsular Malaysia, for the extensive food and
industrial monocultures into which nearly all the lowland rainforests have been con-
verted. Food crops are needed for subsistence, if they can keep pace with exploding
population. Industrial crops provide necessary income, although this may end up in
foreign hands. Wholesale conversions of rainforest such as wood-chipping, e.g. in the
Gogol Valley, Papua New Guinea, ignore ecological safeguards (Webb, 1977).
Economic development becomes the manipulation of things, and any increases in
human welfare ‘are seen as being spin-offs from a rapidly growing economic system in
which such things as income and consumption inequalities and natural resource
destruction are considered normal’ (De’ Ath, 1982).
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
268 ECOLOGICAL VALUES OF THE TROPICAL RAINFOREST RESOURCE
What are our historical perspectives as a nation about rainforests? Australia is the
only developed country largely in the tropics. The area of rainforests throughout the
whole of the continent when Europeans arrived was relatively small, about 1 per cent;
plus another half per cent or so if coastal sclerophyll-rainforest mixtures, and scattered
fragments in drier subcoastal north-eastern areas are added.
The rainforest area when Aboriginals arrived over 40,000 years ago was
somewhat larger than when the first Europeans settled 200 years ago. The area was
much larger during earlier moist warm interglacial periods. And long before man
arrived, beyond the onset of arid conditions in the Plio-Pleistocene, and stretching back
in time through the Tertiary to the Cretaceous and the origin of the flowering plants,
rainforests (or moist closed forests) covered most if not all of the soils of the land mass
that became Australia.
Current interpretations of the evolution of the rainforest and sclerophyll forest
floras have reversed traditional botanical doctrine and the ‘invasion theory’ (Keast,
1981; Barlow, 1981). Previously it was thought that the sclerophylls were the most
ancient, the only truly Australian (autochthonous) elements; whereas the rainforests
were ‘alien and invasive’ immigrants. However, the sharing of many taxa by the
tropical and subtropical rainforests in Australia and Indo-Malesia-Oceania is now
construed as indicating common origins of palaeotropical stocks in Gondwanic times,
as well as various migrations since (Webb and Tracey, 1981a; Webb, Tracey and
Jessup, 1982).
Since European settlement, about two-thirds to three-quarters of the area of
coastal and subcoastal rainforests and monsoon forests in the tropics and subtropics
have been cleared, leaving about 10,000 km?. Rainforest conversion has varied from
virtually 100 per cent on the lowlands and tablelands of the tropical and subtropical
coast, to virtually nil on high mountains.
We now view the last fragments of Australian rainforests through the eyes of fresh
scientific understanding that is most conveniently dated from last year, which marked
the publication of Keast (1981), and the first International Botanical Congress to be
held in Australia. The rare and scattered distribution of the rainforests is the legacy of
million-year-old climatic sifting, thousand-year-old Aboriginal burning, and century-
old impacts of Europeans. Their rarity has now become valuable.
The network of refugia for distinct community-types, and the abundance of
disjunct and different kinds of endemic plant species, are highly relevant to the location
of reserves for nature conservation, especially for scientific purposes. This is one of the
aims of the State National Parks and Wildlife Services, and a central aim of the
UNESCO Biosphere Reserves and the World Conservation Strategy.
For Australian rainforests, ‘small’ may also be ‘beautiful’ (with an efficient buffer
zone), so that a scattering of carefully selected smaller reserves — as well as larger ones
— is essential to preserve not only within-species variation, but also relictual species
and species-assemblages. There are virtually no data for minimum breeding
population sizes for different categories of plants in Australian rainforests. Many tree
species are restricted to very few sites (Tracey, 1981). These rare species are common
where they occur. Are their population sizes stable? If the refugia are ancient, and the.
species are of low vagility, are the rare species tending to extinction in the absence of
immigration? Australian rainforest distributions provide ready-made and natural
experiments in island biogeography. Elsewhere, as in Amazonia, scattered remnants
have to be preserved artificially for this purpose. The age, mutability, and diversity of
refugia, and the endemic and primitive species in them, should all be considered in
judgments about size, shape, and distribution of nature reserves (Webb and Tracey,
1981a; Kikkawa etal. , 1981).
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
L. J. WEBB 269
The transitions and interspersions with eucalypts and other sclerophylls of the mostly
closed, broad-leaved Australian rainforest and monsoon forest types that, unlike the
sclerophylls, have many genera in common with Indo-Malesia, have ecological
properties that are unique in the world. Boundaries in the tropics and subtropics are
often abrupt, as the result of a combination of factors dominated by surface fire.
Boundaries are generally gradual and extensive in temperate and montane regions
where crown fires are a dominant factor, and in wet tropical and subtropical regions on
poorer soils. The ‘tall dark wall’ of rainforest rising at the edge of the sunlit eucalypt
grassy woodland — a metaphor beloved by some early authors — 1s actually only one
of many types of transition. At least five types of transition of interspersion of the
rainforest and sclerophyll forest floras can be recognized (Webb and Tracey, 1981b;
Webb, Tracey and Williams, 1983). The evolutionary significance of the interspersed
taxa, both phylogenetically and syngenetically, has not been studied in detail. The
transitions and mixtures are nevertheless highly relevant to our understanding of
community development and stability involving the rainforest and sclerophyll floras. It
is certainly not helpful for this scientific purpose to discard or confuse ecological facts
and hypotheses by pragmatic forestry typology based on commercial timber volumes.
Commercial hardwoods such as Brush Box (T7istania conferta) reach relatively huge
volumes in these transitional forests or ‘mixed rainforests’ at advanced stages of
succession. Large Brush Box trees up to approximately 1200 years old were the subject
of confrontations in the forest at Terania, and in court at Sydney, New South Wales,
between forestry and the timber industry, and conservation groups (Prineas and
Elenius, 1980). The conflicting values placed on these trees, either as logs or as ancient
objects in nature, provide a classical example of the politics of ecology, and of the
power of popular support for ‘non-wood’ values of Australian moist forests.
Ulterior factors have also entered the botany of Australian rainforests at the
highest levels of vegetation classification. A comprehensive floristic analysis of
Australian rainforests, either by tree species or tree genera, separates the subtropical
formation-class at the third and fourth division respectively in the hierarchy (Webb and
Tracey, 1981a, 1981b; Webb, Tracey and Williams, 1983). The subtropical rain-
forests, often characterized by Araucarian emergents that are of great antiquity and of
high commercial value, should not, therefore, be regarded simply as impoverished
versions of the tropical types farther north. The separate ecological identity of the
subtropical rainforests has to be taken into account when judging the adequacy of
existing rainforest reserves and national parks, especially when compared with those of
the tropical and the montane types.
Species richness and the maintenance of diversity in natural and relatively un-
disturbed communities provides another example of the exercise of contrasting value
judgements by ecologists and others. Given the fact that in complex rainforest com-
munities, many species are represented by relatively few individuals in any one area,
and remembering our ignorance of the numbers of individuals that constitute a sur-
vival threshold for a particular species, what changes in species can be predicted in a
given area as the result of artificial disturbance, such as logging? Patchiness or
clustering of organisms at different scales in space and time — so-called intrinsic and
extrinsic diversity — is difficult to explain, and both stochastic and determinate
processes seem to be involved (Webb et al. , 1972; Letouzey, 1978; Connell et al. , 1982).
Regeneration and succession have become ecological problems that have sprouted
ideological roots. Several major types of succession have been described by different
models, beginning with Egler (1954), and extending to Slatyer (1977) and Noble and
Slatyer (1980). Alternative pathways and possible endpoints in reconstructive
secondary succession of tropical and subtropical rainforest in north-eastern Australia
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
270 ECOLOGICAL VALUES OF THE TROPICAL RAINFOREST RESOURCE
were clearly explained by Hopkins (1981). These and other ecological studies show that
regeneration and canopy development by secondary species in one rainforest type (e.g.
simple notophyll vine forest characterized by Coachwood), are not comparable with
another type (e.g. complex notophyll vine forest characterized by a mixture of species
including Booyong).
Data from long-term monitoring of species over representative areas are not
available as a basis for scientific judgment, which has wider terms of reference than
forestry decisions about adequacy of canopy closure by secondary species, and where
the restoration of an ecosystem similar to the original one controls the time scale since
logging. Scientifically, the long-term advantages of conserving diversity in natural
communities have been emphasized by many biologists with interests ranging from
population dynamics (e.g. Elton, 1958; Connell, Tracey and Webb, 1982) to
evolutionary genetics (Frankel, 1970).
So much for examples of ecological research which have harboured tacit value
judgments and have involved intuitive reference to ecological precepts. We have also
undertaken a project, with somewhat tenuous scientific links, to try to explicate, and
thus to a limited extent communicate, our impressions as experienced ecologists of the
rainforests. The attempt also has a practical basis: to determine criteria for selecting
and managing national parks and similar conservation areas in northern Australia.
Commercial values of material commodities, and protective values for soil and water
are already accepted as of demonstrable economic worth. Attention is focused rather
towards the other end — the cultural values — of the spectrum. The aim is to try to
identify and objectify the values of tropical rainforest and associated forest and land-
scape for scientific, educational, recreational, and aesthetic purposes (Kikkawa, Webb
and Tracey, 1974). A co-operative project with K. J. Polakowski, Landscape Ar-
chitecture, University of Michigan, to determine aesthetic values is of special interest.
It partly uses techniques for identifying relevant attributes at the macro-level of land-
scapes, by applying them to the micro-level of forestscapes. It describes perceptions of
the rainforest environment in terms of ecological site factors, about which we have a
great deal of quantitative information. The perceptions are then translated into a
limited number of perceptual effects and emotional states, using various physical
descriptors (vegetation structure, flora, fauna, sound, colour, light, etc.). The aesthetic
experience of an (interested) observer is an interaction between his cultural and
behavioural world and the intricacy and unfolding variety of the rainforest en-
vironment (Polakowski, Webb and Kikkawa, 1982). This experience is, from a
synecological point of view, a kind of ‘emergent property’ (cf. Odum, 1977). The
complexity and variability of natural ecosystems including humans cannot be captured
even by the most sophisticated statistical methods. It can be argued that the sense of
aesthetics in such situations is not a luxury, but a tool for survival. Aesthetic appeal
may play a central role as a human judgment of the viability of ecosystems, and thus be
somehow advantageous (Grossmann, 1982). Although it is appreciated that the
identification of very broad categories of aesthetic experience must remain sketchy, the
conceptual framework does bring together, in a very satisfying way, complex scientific
and other values that were otherwise isolated. The framework should provide a
stimulating approach to forest environments for children in schools, and for interested
adults in the community. It could also be used as a guide to Aboriginal environmental
perceptions that are part of the heritage of natural history.
Of wider ramifications geographically is our current attempt to match rainforest
habitats in northern Australia and Indo-Malesian countries. This attempt is of
theoretical and practical interest, and has social values. The aim of the project, partly
funded by the UNESCO Man and the Biosphere (MAB) Project, is to use structural
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
L. J. WEBB Zajfi|
typology, supplemented by elementary environmental information and floristic data
for common tree genera, to match forest types and habitats on a site-to-site basis.
Given the ancient floristic links already noted between Australia and other south-east
Asian countries, well-matched habitats with a high proportion of shared genera would
suggest parallel evolution (homology) through common descent. Intuitive comparisons
of forest types in the field during reciprocal visits of botanists to Thailand and Australia
indicate a remarkable extent of shared genera in certain deciduous monsoonal types
e.g. 66-90 per cent of Australian tree genera are shared with homologous habitats in
Thailand. Evergreen lower montane forest types share fewer genera e.g. 33-55 per
cent. Numerical analysis of the structural data is now being attempted, and will
hopefully yield a more objective measure of comparison. Northern Australian rain-
forest and monsoon forest patches, although small and scattered, provide the only
vegetation for structural and floristic comparison with other countries. This means that
the sclerophylls, that have mostly displaced the rainforests and monsoon forests, and
that have superior adaptations to fire, drought, and low soil fertility, provide a unique
reservoir for plant introduction elsewhere, once comparable habitat-types are
established by the systematic use of structural data. Besides enriching ecologic
biogeographical understanding, the comparisons could contribute to agricultural and
forestry practice. More than this, habitat-matching could help develop a ‘common
ecological language’ based on the biotic communities of the region, despite their many
man-imposed variations. Perhaps we should also say that tracing this ancient lineage of
the Australian flora contributes, in the best tradition of natural history, to the
enlargement of Australian cultural identity.
Although a common ecological language for the tropical region has yet to be
written, there is no doubt in my mind that it will contain many cultural values and
ethical overtones. I select only one recent scientific example, from the Man and the
Biosphere (MAB) Workshop in Kuala Lumpur, January 1982. The speaker is the
Director of the Forest Research Institute at Kepong, Peninsular Malaysia; and he is
urging multi-species planting and the retention of adequately-sized blocks of natural
forests in forestry areas:
*.. . It is acknowledged that any of the above practices suggested could result in reduction of profits to
the plantation owner. It is also true that the ‘pest’ population to the plantations could increase.
However, it is our contention that sacrifice is necessary for the sake of our environment . . . To limit
future generations to an environment of monotony is to limit their ultimate self-development. There is
a moral obligation of society to provide the necessary environment to maintain the myriads of floral
and fauna species of the world. Any loss and ultimate extinction of plant and animal life is a loss of
human society’ (Salleh and Hashim, 1982).
And so we return to where we began: to the meaning of values and their place in
the ecological language that we all share in some way. Conventional ethics provide
religious morality, and social morality and justice. These deal with man-man
relationships. Is ecological morality and ecological justice now to be seen as a third-
level development for man-nature relationships?
Thus the question about the intrinsic values of nature remains. I believe there are
signs that this question may be considered safely at the level of science. Environmental
values will continue to be ‘good’ or to be ‘bad’. There seems no possibility of con-
sensus, no definitive formulae, to decide how the protection of nature and the needs of
modern society can be reconciled. But it is becoming increasingly clear that we must
learn to live honestly with questions that will always remain open (Ashby, 1976). It
seems wise to reject the concept of human domination, and to change the old ecological
view from ‘man the outsider’ to ‘man the insider’ in ecosystems that involve nature (di
Castri, 1981).
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
272 ECOLOGICAL VALUES OF THE TROPICAL RAINFOREST RESOURCE
A re-assertion of ecological values cannot avoid involving feelings and sympathies
(Wright, 1973). Ecological value systems by themselves are not enough. Ecological
values are the results of valuing processes in relation to ecological principles (such as
interdependence, diversity, continuity and change, the long-term versus the short-
term, and so on). The separation of scientific research and problem-solving from
valuing, all of which are in turn integrated in decision-making, is becoming in-
creasingly questionable. It seems that novel and complex ecological situations must be
matched by revised ethics if man and his culture, and if nature (which he needs, but
which does not need him) are to survive.
We must find appropriate language, and appropriate values as part of it, for the
description of complex ecological situations to be resolved by choice. These are the
situations with tangled and seemingly incompatible strands: scientific values guided by
the demands of objectivity, and ecological values grown indistinguishable from the
responsibilities of care and preservation.
International forestry authorities are beginning to accept ethical responsibilities in -
forest management: for example, the recent statement by the President of the In-
ternational Union of Forestry Research Organizations:
“The post-war generation of foresters has witnessed the most savage and fast destruction of forests in
the entire history of man’s work in the world. Quite often foresters are taking part in such destruc-
tion . . . Research has become a part of those economic concepts which aim towards exploitation . . .
More than any other economic branch, forestry has been sat in the dock of the world, accused of
destroying nature, for which it is only partly to be blamed . . . Research must be based on ethical
principles of responsibility . . . Forestry and its research have to return to nature, which so far has not
been done. Tomorrow will take man back to nature, because there is no other way’ (Mlinsek, 1982).
What this forester, as a scientist, is asserting is that moral choices are inescapable
in research as well as its applications. We must learn to recognize and acknowledge
these choices. Such acknowledgment should not weaken our pursuit of objectivity, but
make us better scientists as we become aware of more and more of the components of
the ecological equation.
There are no simple solutions, but it seems clear that ethics and value judgments
may no longer remain private affairs for individuals or bureaucracies that make
decisions. Thus our values will evolve through the choices made. In other words, we
will discover that the last vestiges of Australian rainforests are not so good to cut as they
are ‘good to think’ (cf. Dwyer, 1982, p. 186).
ACKNOWLEDGEMENTS
I wish to thank Professor Jiro Kikkawa for several helpful suggestions after
reading the draft of this lecture. Helpful comments on the final manuscript were also
made by Mrs Judith Wright McKinney and Sir Rutherford Robertson.
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Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
A new Genus of Spiders of the Subfamily
Metaltellinae (Araneae, Amaurobioidea) from
southeastern Australia
M. R. GRAY
Gray, M. R. A new genus of spiders of the subfamily Metaltellinae (Araneae,
Amaurobioidea) from southeastern Australia. Proc. Linn. Soc. N.S.W. 106 (4),
(1982) 1983: 275-285.
The metaltelline genus, Austmusza gen. nov., from southeastern Australia and
three species, A. wilsoni sp. nov., A. kioloa sp. nov., and A. lindi sp. nov., are
described. This represents the first record of the subfamily Metaltellinae outside the
neotropical region. The presence of a simplified tracheal system in Austmusza is noted.
M. R. Gray, The Australian Museum, P.O. Box A285, Sydney South, Australia, 2000;
manuscript recewed 11 May 1982, accepted for publication in revised form 20 October 1982.
INTRODUCTION
The subfamily Metaltellinae has previously been recorded only from the southern
and central neotropical region (synanthropic in the southern nearctic (Leech, 1971) ).
Lehtinen (1967) recognized four genera in the subfamily: Metaltella Mello-Leitao 1931,
Ciniflella Mello-Leitao 1921, Calacadia Exline 1940 and Exlinea Lehtinen 1967. Leech
(1971) synonymized Exlinea with Metaltella and noted (Leech, 1972) that Ciznzflella
should be regarded as a nomen dubium (type material lost, description inadequate).
Metaltella is a cribellate genus while both Calacadia and the genus described below,
Austmusia gen. nov. from eastern Australia, are ecribellate. These genera are ap-
parently derived from a palaeoaustral fauna which occupied the southern parts of
Gondwanaland.
METALTELLINAE Lehtinen 1967
Diagnosis: Ecribellate or cribellate; cribellum bipartite. Chelicerae with 2-6 retrolateral
and 3-7 prolateral teeth. Labium longer than wide. Male palp with reduced primary
conductor. Secondary conductor a large, falciform, anteriorly-directed process en-
closing a long, curved, slender emboius. Tegulum more or less bulbous basally.
Median apophysis absent. Fixed anterior tegular apophysis present or absent.
Epigynum with a large, flat, well-sclerotized plate, lateral teeth present or absent.
Internal genitalia usually with convoluted ducts, sometimes simple, with associated
diverticula.
KEY TO GENERA
1. Ecribellate. Cheliceral teeth 2 (retro.) and 2-3 (pro.) Epigynum with
Sinomfalerbercl PeetmOIaMOMe: 25 s:5) 4 4.5. ec vik- ahs costae 4 ue ops Saaeeee nae eee eae 2
— Cribellate. Cheliceral teeth 4-6 and 5-7. Epigynum with elongate lateral
OSU Maal ohk Bee aie hae eS aE De PI ho SPO ac onic eto Metaltella
2. Male palp with large patellar apophysis. Epigynal plate unmodified .. Calacadia
— Male palp lacking patellar process. Epigynal plate with a pair of
depressions roofed anteriorly by thinlaminae .................. Austmusia
Austmusia gen. nov.
Diagnosis: Ecribellate. Cheliceral groove with 2 retrolateral, 3 prolateral teeth. Male
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
276 A NEW GENUS OF SPIDERS
Figs 1-4. Body pattern, females. 1 and 2, Austmusia wilsoni sp. nov. 1, lateral; 2, dorsal. 3, A. kzoloa sp. nov.,
dorsal abdomen. 4, A. lindi sp. nov., dorsal abdomen. Scale lines 0.45 mm.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
M.R.GRAY 277
palp with a vestigial primary conductor. Basal tegulum bulbous; tegular process hook-
like. Patellar process absent. Epigynum with two shallow depressions roofed anteriorly
by thin cuticular laminae. Two tracheal tubes confined to abdomen.
Description: Large spiders, 10 to 14 mm body length. Carapace dark brown, darkest
anteriorly; central cephalic and marginal thoracic areas lighter orange brown. Ab-
domen dark brownish-grey dorsally with well-defined fawn markings (Figs 1-4).
Ventral abdomen light brown with two lateral, dark grey, longitudinal stripes.
Chelicerae black with an orange boss. Legs banded dark and light brown.
Carapace (Figs 1, 2) almost glabrous, cephalic area prominent, arched
longitudinally, raised at fovea. Clypeus vertical, height three to four times the width of
an A.M.E. Anterior median eyes smallest. From above both eye rows straight to
slightly recurved; from in front both rows procurved (Fig. 8).
Cheliceral teeth, Fig. 7; maxillae, labium, sternum, Fig. 5. Sternum with three
prominent groups of lyriform organs near each lateral margin.
Legs 4123 (females) or 1423 (males). Trochanters weakly notched. Hairs plumose,
feathery hairs absent. Ventral spination, legs 1 and 2, tibia 222, metatarsus 221.
Tarsal organ tear-shaped, widest distally (Figs 18, 19). Tiichobothria in single row on
tarsi and metatarsi, double row on tibiae. Trichobothrial bases collariform; hood
striated, margin entire (Figs 16, 17). Superior tarsal claws with eight to thirteen teeth;
inferior claw with four to eight teeth. Toothed bristles present ventrally on distal tarsus.
Abdominal spiracle narrow and situated just in front of colulus. Colulus a large,
rectangular, pilose plate, slightly wider than long, distal margin slightly notched (Fig.
6). Spinnerets six, short. Epiandrous glands absent. Tracheal system consists of two
tubes only, oimined to abdomen (Fig. 31).
Palpal organ (Figs 9-11) with a vestigial primary conductor. Basal tegulum bulbous,
Figs 5-8. Austmusia wilsoni sp. nov. 5, maxillae, labium, sternum. 6, spinnerets, colulus, spiracle. 7,
cheliceral teeth. 8, eyes, anterior view. Scale lines 0.5 mm.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
278 A NEW GENUS OF SPIDERS
Figs 9-15. Austmusia wilsoni sp. nov. 9-11, male palp. 9, prolateral; 10, ventral; 11, retrolateral (e = em-
bolus, ta = tegular apophysis, 1° = primary conductor, 2° = secondary conductor). 12-13, male palpal
tibia and patella; 12, retrolateral; 13, ventral. 14-15, female genitalia. 14, epigynum; 15, internal
genitalia, dorsal, left hand side. Scale lines 0.2 mm.
PROC. LINN. SOc. N.S.W. 106 (4), (1982) 1983
M.R.GRAY 279
separated from apical part by a deep transverse furrow. Apical tegulum with a fixed
hook-like process arising subapically on retrolateral side. Ejaculatory duct visible on
lower tegulum. Male palpal tibia (Figs 12-13) with a prominent, retrolateral basal
process and a retrolateral ventral lamina which ends apically in a small blunt hook.
Patellar processes absent.
Epigynum (Fig. 14) broad and strongly sclerotized with a pair of large, shallow
depressions, open posteriorly but roofed anteriorly by thin cuticular laminae. Lateral
teeth absent. External foveae placed in the posterior ends of two deep, curved grooves
near each lateral epigynal margin. Internal genitalia (Fig. 15) with convoluted sper-
mathecal ducts (partly visible externally) into which a pair of diverticula open.
Spermathecae adjacent near midline, elongate, narrowest centrally. Fertilization ducts
short, broad.
Type Species: Austmusia wilsoni sp. nov.
Etymology: Exline (1960) named the Chilean metaltelline genus Calacadia after the
Californian Academy of Sciences. This provides a precedent for similarly naming the
closely related Australian genus, Austmusza, after its repository institution, The
Australian Museum.
Type Repository: The Australian Museum, Sydney (A.M.).
Austmusia wilsoni sp. nov.
(Figs 1-2, 9-15, 16, 18, 31)
Diagnosis: Male palp with apex of primary conductor level with highest part of tegular
process (Fig. 11). On the basal tegulum a broad segment of the ejaculatory duct loops
apically between two slender diverging segments of the duct (Figs 9, 10). Female
genitalia with long spermathecae, the anterior ends bent 45° laterally (Fig. 15).
Female holotype (A.M., KS 5736)
Measurements (mm) — Carapace length 5.57 (5.10-6.40), width 3.51 (3.19-3.98),
height 2.21 (2.02-2.50). Abdomen length 5.50 (5.25-6.10), width 4.25 (3.80-4.65).
Colour pattern — as for genus. Posterior paired abdominal markings often fused
medially.
Carapace — longer than wide in ratio 1: 0.63. Clypeus height 4 diameters of an A.M.E.
Eyes — ALE > PLE > PME > AME in ratio of 1: 0.87: 0.74: 0.52. Interdistances
(mm): AME-AME 0.14, AME-ALE 0.15, ALE-PLE 0.13, PLE-PME 0.38,
PME-PME 0.26. M.O.Q. (mm): length 0.58, anterior width 0.44, posterior width
0.64. Eye row width (mm): anterior 1.25; posterior 1.50. From above A.E.R. straight,
P.E.R. recurved; from in front A.E.R. gently procurved, P.E.R. strongly procurved.
Labium — longer than wide in ratio 1: 0.63
Sternum — longer than wide in ratio 1: 0.83
Legs: 4123. Lengths, legs 1-4 (mm): 18.28, 16.38, 14.94, 18.55. Spination: Leg 1,
femur p 01, d 213; tibia p 11, r 11, v 222; metatarsus p 012, r 012, v 221. Leg 2, femur
p 0111, d 01212; tibia p 011, r 001, v 222; metatarsus p 112, r 012, v 221. Leg 3, femur
p 0111, d 1202; tibia p 011, r011, d 010, v 222; metatarsus p 112, r 112, d 010, v 221.
Leg 4, femur p 101, d 1102; tibia p 101, r 101, d 01, v 212; metatarsus p 112 or 111,r
112 or 111, d 010 or 011, v 221. Tarsal organ a narrow, tear-shaped opening, widest
distally, not on obvious mound. Trichobothria, legs 1-4: tarsus 10, 9, 9, 9; metatarsus
9,9, 9, 9. Tarsal claws; superior 11-13 teeth; inferior 4-8 teeth.
Genitalia — Figs 14, 15. Spermathecae visible externally, elongate with the apical third
to half bent 45° laterally. Spermathecal length approximately half the width of the
epigynum (measured between the lateral foveal margins). Diverticulum duct narrow
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
280 A NEW GENUS OF SPIDERS
2
Bae
Figs 16-19. Sensilla, first tarsus. 16-17, trichobothrial base. 16, Austmusza wilsoni; 17, A. kioloa. 18-19, tarsal
organ. 18, A. wilsonz; 19, A. kioloa. Scale lines: 16-17, 0.006 mm; 18-19, 0.002 mm.
with a small, rounded, distal part which lies well lateral to and below the level of the
anterior end of the spermatheca. Fertilization ducts short and broad.
Male paratype (A.M., KS 7795)
Similar to female except as follows—
Measurements (mm) — Carapace length 5.49 (4.59-6.41), width 3.79 (3.20-4.21),
height 2.16 (1.98-2.45). Abdomen length 5.30 (4.85-5.75), width 3.50 (3.00-3.80).
Carapace — longer than wide in ratio 1: 0.67. Clypeus height 3 diameters of an AME.
Eyes PLE > ALE > PME > AME in ratio 1: 0.95: 0.81: 0.72. Interdistances (mm):
AME-AME 0.08, AME-ALE 0.14, ALE-PLE 0.06, PLE-PME 0.26, PME-PME
0.18. M.O.Q. (mm): length 0.58, anterior width 0.42, posterior width 0.58. Eye row
width (mm); anterior 1.10; posterior 1.29.
Chelicerae — cheliceral teeth: retrolateral 2; prolateral 3 with an additional microtooth.
Labium — longer than wide in ratio 1: 0.79.
Sternum — longer than wide in ratio 1: 0.78.
Legs — 1423. Length, legs 1-4 (mm): 19.04, 16.76, 14.12, 18.84. Spination: Leg 1,
femur p 0011, d 122; tibia p 112, r 011, v 222; metatarsus p 11, r 010, v 221. Leg 2, p
1011, d 122; tibia p 11, r 11, v 222; metatarsus p 112, r012, v 221. Leg 3 femur p 101,
d 122; tibia p 11, r 11, d (1)01, v 22; metatarsus p 112, r 112, d 010, v 221. Leg 4,
femur p 101, d 112; tibia p 11, r 011, d(1)01, v 212; metatarsus p 112, r 112, d 010, v
221. Trichobothria, legs 1-4: tarsus 9, 8, 8, 9; metatarsus 10, 9, 8, 8. Tarsal claws:
superior 11-12; inferior 4-8.
Palp — Figs 9-13. Apex of reduced primary conductor level with highest part of hooked
tegular process. Course of ejaculatory duct on basal tegulum — Figs 12, 13.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
M.R.GRAY 281
Types:
Holotype female — KS 5736 (A.M.), Cathedral of Ferns, Mt. Wilson, N.S.W., M.
Gray, 17.4.1974; under log, depauperate closed forest.
Paratypes — Male, KS 7795 (A.M.), Mt. Wilson, N.S.W., M. Gray and C. Hor-
seman, May 1978; in pitfall trap, tall open forest. Female, KS 2046 (A.M.), Mt.
Wilson, N.S.W., M. Gray and C. Horseman, October/November 1978; in pitfall
trap, tall open forest. Male, KS 3877 (A.M.), Waterfall Trail, Mt. Wilson, C.
Horseman, August 1979; in pitfall trap, closed forest. Male, KS 7796 (A.M.), Mt.
Victoria, 14.5.1972, G. S. Hunt; under log in valley, tall open forest. Male, KS 7797
(A.M.), Mt. Edwards, Boyd Plateau, N.S.W., M. Gray, 15.5.1971; under log, tall
open forest. Female, KS 7798 (A.M.), Blood Filly Creek, Boyd Plateau, N.S.W., M.
Gray 26.3.1976; in rotting log, tall open forest.
Austmusia kioloa sp. nov.
(Figs. 3, 17, 19, 20-26)
Similar to A. wilsoni except as follows:
Diagnosis: Male palp with apex of primary conductor clearly lower than highest part of
tegular process (Fig. 22). Ventrally, two slender segments of ejaculatory duct on the
basal tegulum first converge anteriorly, then diverge, the prolateral duct curving
laterally away (Figs 20, 21). Spermathecae long, apical third enlarged and laterally
protuberant (Fig. 26).
Female holotype (A.M., KS 4551).
Measurements (mm) — Carapace length 4.87 (4.48-5.25), width 3.15 (2.88-3.43),
height 1.89 (1.60-2.18). Abdomen length 5.70 (5.45-5.95), width 4.70 (4.50-4.85).
Carapace — longer than wide in ratio 1: 0.65. Clypeus height 3.5 diameters of an AME.
Eyes — ALE > PLE > PME > AME in ratio 1; 0.95: 0.82: 0.68. Interdistances
(mm): AME-AME 0.11, AME-ALE 0.16, ALE-PLE 0.08, PLE-PME 0.30,
PME-PME 0.23. M.O.Q. (mm): length 0.53, anterior width 0.39, posterior width
0.61. Eye row width (mm), anterior 1.09; posterior 1.37.
Labium — longer than wide in ratio 1: 0.80.
Sternum — longer than wide in ratio 1: 0.81.
Legs — 4123. Lengths, legs 1-4 (mm): 14.68, 12.79, 11.37, 15.36. Spination: Leg 1,
femur p 002, d 111; tibia p 101, r 101, v 222; metatarsus p 111, r 012, d 010, v 221.
Leg 2, femur p 101, d 1112; tibia p 101, r 101, v 222; metatarsus p 112, r012,d 010, v
221. Leg 3, femur p 101, d 1112; tibiap 11,r101,d 11, v 212; metatarsus p 112, r 112,
d 010, v 221. Leg 4, femur p 001, d 112; tibia p 101, r 101, d 01, v 212; metatarsus p
111,r111,d 012, v 221. Tarsal organ a tear-shaped opening, widest distally, placed on
alow, poorly delimited mound. Trichobothria, legs 1-4: tarsus 8, 8, 7, 8; metatarsus 8,
9, 7, 8. Tarsal claws: superior 8-11 teeth; inferior 4-7 teeth.
Genitalia — Figs 25, 26. Basal parts of spermathecae visible externally. Spermathecae
elongate, apical third laterally bulbous and rounded. Spermathecal length greater than
half epigynal width. Diverticulum duct narrow, distal part elongate, lying over or
adjacent to apical part of spermathecae. Fertilization ducts longer than in A. wzlsonz.
Male paratype (A.M., KS 1651)
Similar to female except as follows:
Measurements (mm) — Carapace length 4.67 (3.90-5.15), width 3.37 (2.93-3.60),
height 1.70 (1.45-2.03). Abdomen length 4.20 (3.80-4.30), width 3.10 (2.85-3.25).
Carapace — longer than wide in ratio 1: 0.73. Clypeus height 3 diameters of an A.M.E.
PROC. LINN. Soc. N.S.W. 106 (4), (1982) 1983
282 ANEW GENUS OF SPIDERS
Figs 20-26. Austmusia kioloa sp. nov. 20-22, male palp. 20, prolateral; 21, ventral; 22, retrolateral. 23-24,
male palpal tibia and patella. 23, ventral; 24, retrolateral. 25-26, female genitalia. 25, epigynum; 26,
internal genitalia, dorsal, left hand side. Scale lines 0.2 mm.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
M.R. GRAY 983
Eyes — ALE = PLE > PME > AME in ratio 1: 1: 0.95: 0.85. Interdistances (mm):
AME-AME 0.10, AME-ALE 0.14, ALE-PLE 0.08, PLE-PME 0.25, PME-PME
0.20. M.O.Q. (mm.): length 0.51, anterior width 0.44, posterior width 0.55. Eye row
width (mm): anterior 1.03; posterior 1.30.
Labium — longer than wide in ratio 1: 0.72.
Sternum — longer than wide in ratio 1: 0.83.
Legs — 1423. Lengths, legs 1-4 (mm): 17.42, 15.42, 13.53, 16.83. Spination: Leg 1,
femur p 002, d 1111; tibia p 101, r 101, v 222; metatarsus p 112, r 012, v 221. Leg 2,
femur p 101, d 1112; tibia p 101, r 101, v 222; metatarsus p 111, r 011, d 012, v 221.
Leg 3, femur p 101, d 1112; tibia p 11, r 101, d 01, v 212; metatarsus p 111, r111,d
012, v 221. Leg 4, femur p 101, d 112; tibia p 101, r 101, d 01, v 212; metatarsus p
111,r111,d 012, v 221. Trichobothria, legs 1-4: tarsus 10, 9, 8, 9; metatarsus 8, 8, 9,
9. Tarsal claws: superior 8-10 teeth; inferior 4-7 teeth.
Palp — Figs 20-24. Apex of reduced primary conductor well below the highest part of
the hooked tegular process. Course of ejaculatory duct on basal tegulum — Figs 20, 21.
Types
Holotype female — KS 4551 (A.M.), Kioloa Forest Drive, Kioloa S.F., N.S.W., M.
Gray and C. Horseman, May/June 1979; in pitfall trap, tall open forest.
Paratypes — Male, KS 1651 (A.M.), Kioloa S.F. (rest area on Princes Highway),
N.S.W., M. Gray, 14.8.1978; under litter, tall open forest. Male, KS 3115 (A.M.),
Kioloa Forest Drive, Kioloa S.F., N.S.W., C. Horseman, May/June 1979; in pitfall
trap, tall open forest. Female, KS 2810 (A.M.), same data as KS 3115, March 1979.
Male, KS 1730 (A.M.), same data as KS 3115, August 1978.
Austmusia lind: sp. nov.
(Figs 4, 27, 28)
Similar to A. wilsoni except as follows:
Diagnosis: Female genitalia with short spermathecae, visible externally, apical end
swollen; spermathecal length less than half the width of the epigynum (between lateral
foveal margins).
Female holotype (A.M., KS 1777)
Figs 27-28. Austmusia lindi sp. nov., female genitalia. 27, epigynum. 28, internal genitalia, dorsal, left hand
side. Scale lines 0.2 mm.
PROC. LINN. Soc. N.S.W. 106 (4), (1982) 1983
284 A NEW GENUS OF SPIDERS
3] 30 29
Figs 29-31. Abdominal tracheal system. 29, Badumna insignis (L.K.). 30, Metaltella stmoni (Keysl.). 31,
Austmusia wilsont sp. nov. Scale lines 0.4 mm.
Measurements (mm) — Carapace length 6.42, width 4.07, height 2.50. Abdomen length
6.7, width 4.2.
Colour pattern — dorsal, paired abdominal markings not fused posteriorly.
Carapace — longer than wide in ratio 1: 0.63. Clypeus height 3.5 diameters of an AME.
Eyes — ALE = PLE > PME > AME in ratio 1: 1: 0.83: 0.78. Interdistances (mm):
AME-AME 0.18, AME-ALE 0.19, ALE-PLE 0.12, PLE-PME 0.38, PME-PME
0.30. M.O.Q. (mm): length 0.64, anterior width 0.52, posterior width 0.70. Eye row
width (mm): anterior 1.35, posterior 1.73.
Labium — longer than wide in ratio 1: 0.77.
Sternum — longer than wide in ratio 1: 0.70.
Legs — 4123. Lengths, legs 1-4 (mm): 17.53, 15.72, 13.71, 18.13. Spination: Leg 1,
femur p 002, d 11(2); tibia p 101, r 001, v 222; metatarsus p 111, r 011, d 002, v 221.
Leg 2, femur p 101, d 122; tibia p 101, r 111, v 222; metatarsus p 111, r011,d012,v
221. Leg 3, femur p 101, d 122; tibia p 11, r 11, d(1)1, v 212; metatarsus p 1101, r
111, d 012, v 221. Leg 4, femur p 001, d 112; tibia p 101, r 101, d (1) 01, v 212;
metatarsus p 111, r 111, d 012, v 221. Trichobothria, legs 1-4; tarsus 8, 8, 7, 7;
metatarsus 8, 8, 7, 7. Tarsal claws; superior 9-13 teeth; inferior 4-8 teeth.
Genitalia — Figs 27, 28. Spermathecae relatively short, less than half the width of the
epigynum (between the lateral foveal margins); anterior third swollen. Diverticula
large, irregularly ovoid, apex lying just lateral to and above the anterior end of the
spermathecae; duct broad. Fertilization ducts longer than in A. walsonz.
Types
Holotype female — KS 1777 (A.M.), Dingo Creek, Lind National Park, Victoria, M.
Gray and C. Horseman, 24.3.1978; in rotting log, mixed closed/tall open forest.
NOTES ON TRACHEAL SYSTEMS
The tracheal respiratory system of Austmusia gen. nov., is unique within the
PROG. LINN. SOc. N.S.W. 106 (4), (1982) 1983
M.R.GRAY 285
Amaurobioidea (sensu Lehtinen, 1967) so far investigated. Within the Dic-
tynioidea/Amaurobioidea (sensu Forster, 1970; Forster and Wilton, 1973) two basic
tracheal patterns have been recognized: A ‘complex’ pattern consisting of four finely
branching tracheal tubes, often arising from a wide spiracle, either confined to the
abdomen (Fig. 29) or extending through into the cephalothorax; and a simple pattern
consisting of four unbranched tracheal tubes, usually arising from a narrow spiracle,
and confined to the abdomen (Fig. 30). In Austmusia, the pattern is further simplified
(Fig. 31), only the two medial tracheal tubes being present; they are unbranched,
confined to the abdomen and arise from a narrow spiracle placed immediately in front
of the spinnerets (Fig. 6). However, this pattern is not characteristic of all metaltelline
spiders. Metaltella smoni (Keyserling, 1877) has the simple pattern of four unbranched
abdominal tracheal tubes (Fig. 30). Unfortunately no specimens of Calacadia spp. were
available for dissection.
ACKNOWLEDGEMENTS
Type material of Calacadia Exline was made available by Dr F. Arnaud, California
Academy of Sciences. A male of Metaltella simoni (Keys.) was lent by Dr H. W. Levi,
Museum of Comparative Zoology, Harvard.
References
EXLINE, H., 1960. — Rhoicinine spiders (Pisauridae) from western South America. Proc. Calif. Acad. Sct.
4th Ser. 29 (17): 577-620.
FORSTER, R. R., 1970. — The spiders of New Zealand, Part III. Otago Mus. Bull. 3: 1-184.
, and WILTON, C. L., 1973. — The spiders of New Zealand, Part IV. Otago Mus. Bull. 4: 1-309.
LEECH, R., 1971. — The introduced Amaurobiidae of North America and Callobius hokkaido n.sp. from
Japan (Arachnida: Araneidae). Canad. Ent. 103: 23-32.
——.,, 1972. — A revision of the Nearctic Amaurobiidae (Arachnida: Araneida). Mem. Ent. Soc. Canad.
No. 84: 1-128.
LEHTINEN, P. T., 1967. — Classification of the cribellate spiders and some allied families, with notes on the
evolution of the suborder Araneomorpha. Ann. Zool. Fenn. 4: 199-468.
A Reappraisal of the Stratigraphy of the upper
Shoalhaven Group and lower Illawarra Coal
Measures, southern Sydney Basin, New South
Wales
PAUL F. CARR
CarR, P. F. A reappraisal of the stratigraphy of the upper Shoalhaven Group and
lower Illawarra Coal Measures, southern Sydney Basin, New South Wales. Proc
Linn. Soc. N.S. W. 106 (4), (1982) 1983: 287-297.
Recent mapping and reappraisal of previous work have clarified the stratigraphy
and nomenclature of the upper Shoalhaven Group and lower Illawarra Coal Measures
of the southern Sydney Basin. The interval between the Berry Siitstone and Illawarra
Coal Measures is termed the Broughton Formation which can be subdivided in the
Gerringong-Wollongong region where distinctive latite flows occur. In stratigraphic
sequence the members of the Broughton Formation are Westley Park Sandstone
(lowermost unit), Blow Hole Latite, Kiama Sandstone, Bumbo Latite, Jamberoo
Sandstone, Saddleback Latite, Dapto Latite and Cambewarra Latite (uppermost
unit). The Saddleback and Dapto Latite Members are considered to be separate flows
which occur at similar stratigraphic positions below the Cambewarra Latite. The
Pheasants Nest Formation of the Illawarra Coal Measures overlies the Broughton
Formation and contains four latite members: Five Islands, Calderwood (new unit),
Minumurra and Berkley.
P. F. Carr, Department of Geology, University of Wollongong, P.O. Box 1144, Wollongong,
Australia 2500; manuscript received 5 August 1981, accepted for publication in revised form 18
August 1982.
INTRODUCTION
The southern Sydney Basin contains Permian and Triassic marine and non-
marine sedimentary rocks which were subjected to four phases of igneous activity
between the Late Permian and the Late Oligocene (Carr and Facer, 1980). Permian
rocks are subdivided into two groups, the lower of which comprises the marine strata of
the Shoalhaven Group, the other consists of the overlying non-marine strata of the
Illawarra Coal Measures. Diversity of opinion concerning the mode of emplacement,
and the number, of Late Permian igneous rock-units which crop out in the
Wollongong-Gerringong region (Fig. 1) has been a major factor in the lack of
agreement on the stratigraphic status and relationships of many Late Permian units in
the southern Sydney Basin. Remapping by the author has clarified the stratigraphic
nomenclature of the upper Shoalhaven Group and lowermost Illawarra Coal
Measures. Rationalization of this confused terminology prompts the present author to
propose one new name and a number of changes in status. Important previously
published stratigraphic schemes and the scheme proposed herein are shown in Fig. 2.
STRATIGRAPHIC SETTING
The Permian sequence in the southern Sydney Basin unconformably overlies
earlier Palaeozoic strata. In the southern extremity of the basin minor coal measure
sedimentation (Clyde Coal Measures) occurred as a lateral western equivalent to the
shallow marine Conjola Sub-group. This latter unit is conformably overlain by the
Wandrawandian Siltstone which in turn underlies the Nowra Sandstone. Boundaries
between all the units in the lower Shoalhaven Group represent facies changes which are
slightly diachronous. All of the units so distinguished appear to grade westwards into
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
288 STRATIGRAPHY OF THE SOUTHERN SYDNEY BASIN
OSI [aa] sedimentary and SSS >: WOLLONGONG
PERMIAN igneous rocks =i f::
ILLAWARRA 3
= COAL MEASURES —— a FLINDERS
PERMIAN is KEMBLA * ISLET
SHOALHAVEN e a
GROUP
fo) fo)
fe}
_STOCKYARD 2
MOUNTAIN , 2 -~
ce)
KANGAROO [-
o VALLEY 5 : GERRINGONG
CAMBEWARRA °
MOUNTAIN
Fig. 1. Locality map and simplified geology of the Wollongong-Gerringong region.
the Megalong Conglomerate (McElroy e¢ al., 1969). The Berry Siltstone conformably
overlies both the Nowra Sandstone and the Megalong Conglomerate.
Joplin e al. (1952) grouped the marine and igneous rocks between the Berry
Siltstone and Illawarra Coal Measures into the Gerringong Volcanics and named the
unit above the Berry Siltstone the Broughton Tuff. They considered the Broughton
Tuff to be divisible in the Gerringong-Wollongong region because of the presence of
distinctive extrusive units. Rose (1966) included the Gerringong Volcanics as a Sub-
group of the Shoalhaven Group. He recognized the Budgong Sandstone Member (of
the Berry Formation) which he considered was stratigraphically partially equivalent to
the Broughton Tuff. The distinctive igneous units near the top of the Shoalhaven
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
PAUL F. CARR 289
Group had formation status (Harper, 1915; Joplin et al., 1952; Rose, 1966) but were
termed members of the “‘Gerringong volcanic facies’? by Bowman (1970, 1974).
Sandstones between these igneous rocks have been described as formations (Harper,
1915; Rose, 1966) and as members (Joplin e a/., 1952; Bowman, 1970, 1974). West of
the volcanic flows the interval between the Berry Siltstone and the Illawarra Coal
Measures has also had a varied nomenclature including Broughton Tuff (Joplin et al.,
1952; Rose, 1966) and Budgong Sandstone (Bowman, 1970, 1974). Recent discussions
on the ages of the marine faunas of the Shoalhaven Group may be found in Runnegar
and McClung (1975) and McClung (1978).
The Shoalhaven Group is conformably overlain by the non-marine deposits of the
Pheasants Nest Formation which is the lowermost unit of the Illawarra Coal Measures.
Harper (1915), Joplin e al. (1952) and Bowman (1970, 1974) have recognized two
distinctive igneous rock-units within the Pheasants Nest Formation.
BROUGHTON FORMATION
In the present study the interval between the Berry Siltstone and the top of the
Shoalhaven Group is termed the Broughton Formation which is subdivided into three
sandstone members and five latite members in the Wollongong-Gerringong area.
The basal unit of the formation is the Westley Park Sandstone Member which
conformably and gradationally overlies the Berry Siltstone at Mount Coolangatta (Fig.
1). The nature of this boundary was recognized by Card and Jaquet (1903) and
McElroy et al. (1969). Subdivision of the formation is entirely based on and, indeed, is
only made possible by the presence of interbedded latites. Problems in stratigraphic
terminology have arisen due to restriction, in large part, of some volcanic flows to the
coastal strip. The top of the Cambewarra Latite Member or equivalent boulder
horizon marks the top of the Shoalhaven Group (Harper, 1915, p. 223) and thus the
top of the Broughton Formation.
The name Broughton is chosen by the author in preference to Gerringong (equal
priority) as the former proposal appears to reflect most closely the original intention of
the nomenclature proposed by Jaquet et al. (1905, plate VII). In addition, usage of the
name Broughton Formation allows the term Gerringong volcanics to be used as an
informal name for the succession characterized by volcanic flows in the upper
Shoalhaven Group and lower Illawarra Coal Measures. The term Budgong is rejected
since it was introduced by Rose (1966) long after both Broughton and Gerringong had
been initially defined (Joplin et al., 1952).
Westley Park Sandstone Member. Jaquet et al. (1905) defined the Westley Park Tuffs which
were renamed the Westley Park Tuff Member, Westley Park Sandstone and Westley
Park Sandstone Member by Joplin ef al. (1952), Raam (1968) and Bowman (1970,
1974) respectively. The unit gradationally and conformably overlies the Berry Siltstone
at Mount Coolangatta and is conformably overlain by the Blow Hole Latite Member
(Fig. 2). The unit has been described by Raam (1968, 1969) as a shallow marine
arenite with minor siltstone and conglomerate, and attains a maximum thickness of 45
m.
Blow Hole Latite Member. Jaquet et al. (1905) named the Blow Hole Flow which was
renamed the Blow Hole Latite by Joplin et al. (1952). The member is stratigraphically
the lowest of the Permian flows in the Kiama-Wollongong region with present outcrops
confined to a narrow zone from Kiama southwards for 22 km to Mount Coolangatta
(Fig. 1). A maximum exposed thickness of 50 m occurs on the eastern side of Sad-
dleback Mountain (Fig. 1). Bowman (1974) has reported that the unit consists of three
flows.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
STRATIGRAPHY OF THE SOUTHERN SYDNEY BASIN
290
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Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
PAUL F.CARR 291
The unit contains pillows and breccias consisting of latite in a sedimentary matrix
showing soft-sediment deformation structures. Both pillows and breccias have been
described in detail by Raam (1964, 1969) who considered that these features indicate
that the Blow Hole Latite Member was contemporaneous with the associated
sedimentary rocks. Since the sandstone units both above and below the Blow Hole
Latite Member contain sedimentary structures and an abundant fauna indicative of a
near-shore marine environment (Raam, 1968), the Blow Hole Latite Member is in-
terpreted as a near-shore, submarine flow on to wet, unconsolidated sediments.
Locally the flow intruded into the sediments to show restricted intrusive contacts.
Kiama Sandstone Member. This unit conformably overlies the Blow Hole Latite Member
and was defined as the Kiama Tuff by Jaquet et al. (1905). Joplin et al. (1952) renamed
the unit the Kiama Tuff Member and Raam (1968, 1969) referred to this arenite as the
Kiama Sandstone. A maximum thickness of 53 m is developed at Jamberoo (Fig. 1).
The faunal assemblages indicate deposition in a shallow marine environment (Raam,
1968).
Bumbo Latite Member. The Bumbo Flow which overlies the Kiama Sandstone Member
was originally named by Jaquet et a/. (1905) and was renamed the Bumbo Latite by
Joplin e al. (1952). This unit is one of the most extensive and most voluminous Per-
mian flows of the southern Sydney Basin and outcrops over a total area of ap-
proximately 240 km?. A maximum thickness of 150 m is developed at Saddleback
Mountain (Fig. 1). Bowman (1974) has reported that the unit consists of three flows in
some areas.
The base of the Bumbo Latite Member is well exposed at Bombo Point where the
contact is either sharp or is marked by a basal breccia up to 3 m thick. Sub-vertical to
vertical pipe-like masses of breccia, 1 to 10 m in diameter, occur at many localities.
Contacts between the breccia pipes and the surrounding latite are normally sharp, with
a decrease in the size and abundance of vesicles in the latite away from the core of the
breccia zone. Both the basal breccia and pipe-like masses of breccia consist of clasts of
latite in a matrix of sedimentary material and cement of zeolite and hematite. The
sedimentary material is indistinguishable from the underlying Kiama Sandstone
Member and in places shows contorted bedding which has been interpreted by Raam
(1964) as due to soft-sediment deformation. The breccias appear to have formed as a
result of the interaction of magma with the underlying water-saturated sediment.
Steam produced at this contact fragmented the base of the lava flow and, in some areas,
dragged up wet sediment into the overlying volcanic rock (Raam, 1964).
Jamberoo Sandstone Member. The Jamberoo Sandstone Member conformably overlies the
Bumbo Latite Member and is conformably overlain by the Saddleback Latite
Member. Jaquet ef a/. (1905) defined the unit as the Jamberoo Tuffs but Joplin et al.
(1952) changed the name to the Jamberoo Tuff Member. Subsequently Raam (1968)
renamed the unit the Jamberoo Sandstone. A maximum thickness of 155 m is
developed at Jamberoo (Jaquet et al., 1905).
Saddleback Latite Member. Nomenclature of the Saddleback Latite Member has been
varied and problematical. The extrusion was defined as the Saddleback Flow by Jaquet
et al. (1905) and renamed the Dapto-Saddleback Flow by Harper (1915) who con-
sidered that the two unconnected outcrops in the vicinity of Port Kembla in the north
and Saddleback Mountain in the south resulted from the same flow. Joplin e¢ al. (1952)
accepted Harper’s opinion and redefined the two outcrops as one unit — the Sad-
dleback Latite. Bowman (1970, 1974) considered that the Port Kembla and Saddleback
Mountain outcrops resulted from two flows at different stratigraphic levels. Thus
Bowman (1970, 1974) recognized the Saddleback Latite Member in the south, and the
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
292 STRATIGRAPHY OF THE SOUTHERN SYDNEY BASIN
Dapto Latite Member in the north and he considered the latter unit to overlie the
Cambewarra Latite Member. In the present investigation the Saddleback and Dapto
Latite Members are considered to be separate flows which occupy similar stratigraphic
positions below the Cambewarra Latite Member. Stratigraphic relations between these
units are discussed in a later section of this paper.
Outcrops of the Saddleback Latite Member extend around the Illawarra escarp-
ment from the southern side of Saddleback Mountain to Woodhill and the eastern limit
of Kangaroo Valley. The unit does not occur on the northern slopes of Saddleback
Mountain but crops out further to the north along the scarp on the southern flanks of
Stockyard Mountain. The maximum thickness of 35 m is developed on the southern
side of Saddleback Mountain.
Harper (1915) interpreted the Saddleback Latite Member as a submarine ex-
trusion. This view was supported by Lowder (1964) who found probable pillow
structures at several localities. Also, the presence of vesicles near the top of the unit is
suggestive of emplacement at, or near, the earth’s surface. Thus the Saddleback Latite
is interpreted as being a submarine lava flow.
Dapto Latite Member. All outcrops of the Dapto Latite Member occur in the vicinity of
Port Kembla within an area of approximately 100 km?. Flagstaff Hill (Fig. 1) and its
vicinity provide the most extensive and thickest (85 m) outcrop and the unit thins to the
north, west and southwest away from this area.
Various features present in the Dapto Latite Member and surrounding
sedimentary rocks have been interpreted as indicating that the unit is a flat-topped
laccolith intruded into unconsolidated Permian marine sedimentary rocks (Wilshire
and Hobbs, 1962). However, since the time of publication of their article, drilling and
extensive road-making activities by the New South Wales Department of Main Roads
have provided access to much more information regarding the mode of emplacement of
the Dapto Latite Member. As the drill-core is not available (lost or destroyed), the
borehole logs of Cook (1966) were used to provide sub-surface data on the Dapto Latite
Member. Integration of the data from outcrops and the boreholes provides the
following interpretation:
(1) The top of the Dapto Latite Member is uneven and is overlain by approximately
3 m of sandstone containing abundant feldspar crystals and fragments of the
underlying latite. The fragments of igneous rock are usually rounded, are
concentrated into particular horizons, and are isolated from the main mass of
igneous rock. The clasts of the Dapto Latite Member in the sedimentary rocks
are undoubtedly detrital in origin.
(11) Although fine-scale individual horizons are difficult to follow in outcrop, near
horizontal bedding (dip 3° to north-northwest) is apparent in the sedimentary
rocks above the Dapto Latite Member. Apart from localized draping of bedding
over the detrital latite clasts there is no evidence of bedding distortion that would
be expected to accompany the intrusion of a large mass of igneous rock into
unconsolidated sediments.
(i) The Dapto Latite Member consists of two flows which, over much of the region,
are separated by a breccia. The borehole logs of Cook (1966) describe the breccia
as a complex succession of sandstone, siltstone and blocks of latite ranging from 5
cm to 3 m in vertical extent. The sedimentary rocks contain clasts of feldspar and
latite and, apart from steeply inclined contacts with the latite blocks, are
horizontally bedded with some cross-bedding.
Cambewarra Latite Member. The Cambewarra Latite Member is the uppermost unit of
the marine Shoalhaven Group and is the most extensive of the Permian extrusions in
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
PAUL F. CARR 29
Le)
the southern Sydney Basin. A maximum thickness of 70 m occurs at Woodhill. The
flow has a pronounced physiographic expression and forms steep slopes or cliffs ex-
tending from Browns Mountain to the northern side of Stockyard Mountain (Fig. 1).
Between Stockyard Mountain and Wollongong, the Cambewarra Latite Member is
represented by a boulder horizon (Harper, 1915).
Numerous amygdules and vesicles near the top of the Cambewarra Latite
Member indicate that crystallization occurred at or near the earth’s surface. A
subaqueous, extrusive mode of emplacement is indicated by the presence of pillow
structures with an inter-pillow matrix of shale and sandstone (Lowder, 1964). Marine
fossils both below and within the equivalent unit suggest deposition in a marine en-
vironment (Harper, 1915, p. 223).
A boulder horizon containing numerous vesicular and amygdaloidal clasts of the
Cambewarra Latite Member in a volcarenite matrix extends beyond the northern limit
of the flow to Wollongong. Harper (1915) considered that the boulder horizon
originated by contemporaneous erosion and deposition of clasts of Cambewarra Latite
Member by a strong marine current flowing towards the north. The direction of the
current was deduced from the absence of the boulder horizon south of Saddleback
Mountain and the decrease in size of the clasts in the northerly direction whereas
marine deposition is indicated by the presence of marine fossils in the horizon. A
southerly source is consistent with recent work on the sedimentology of the upper part
of the Shoalhaven Group and lower part of the Illawarra Coal Measures (Bowman,
1974; B. G. Jones, pers. comm. 1980). Harper (1915) traced the boulder horizon as far
north as the Wollongong Water Supply Service Reservoir which was situated 5 km
north-northeast of Flagstaff Hill (Fig. 1). Excavation of a 30 m deep cutting through
the western flank of Flagstaff Hill by the New South Wales Department of Main Roads
has exposed a conglomerate 3 to 5 m stratigraphically above the Dapto Latite Member.
The conglomerate, which contains rounded clasts of amygdaloidal igneous rock in an
arenite matrix, is considered by the author to represent the boulder horizon in this
region.
PHEASANTS NEST FORMATION
The Broughton Formation is conformably overlain by non-marine deposits of the
Pheasants Nest Formation at the base of the Cumberland Sub-group of the Illawarra
Coal Measures (Fig. 2). Where the Cambewarra Latite Member or the equivalent
boulder horizon is lacking the contact between the Shoalhaven Group and Illawarra
Coal Measures is transitional (Wilson, 1969, p. 372). Sedimentary strata of the
Pheasants Nest Formation consist of interbedded sandstone, siltstone and shale which
lack marine fossils but contain fossil tree stumps, logs and plants. Only the four latite
members interbedded with these sedimentary strata are described here.
Five Islands Latite Member. The Five Islands Flow was named by Harper (1915) but for
consistency of nomenclature is renamed herein the Five Islands Latite Member. The
unit is very restricted in outcrop and occurs only on Flinders Islet which is the nor-
thernmost of the Five Islands (Fig. 1). Pillow structures up to 1 m in diameter indicate
a subaqueous, extrusive mode of emplacement.
As the Five Islands Latite Member is the only unit which crops out on Flinders
Islet, the stratigraphic position of the flow is problematical. The sedimentary strata of
the Broughton Formation on the mainland nearest to Flinders Islet are almost
horizontal with a slight dip (2° to 3°) towards the northwest. Evidence for major
faulting is lacking. Also, the islands south and southeast of Flinders Islet are composed
of Dapto Latite Member. Assuming no major faulting occurs between Flinders Islet
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
294 STRATIGRAPHY OF THE SOUTHERN SYDNEY BASIN
and the mainland or the other islands, the Five Islands Latite Member must be
stratigraphically above the Dapto Latite Member. A lower limit on the stratigraphic
separation between the Dapto Latite Member and Five Islands Latite Member cannot
be inferred due to the lack of outcrops in proximity to Flinders Islet. However, an
upper limit on the stratigraphic separation may be inferred from the regional dip of 3°
to the northwest and the minimum distance between outcrops of the Five Islands and
Dapto Latite Members. The nearest outcrop of the Dapto Latite Member is Bass Islet
which is 1700 m southeast of Flinders Islet (i.e. down dip from the Five Islands Latite
Member) and implies a maximum stratigraphic separation of 90 m. Thus the Five
Islands Latite Member is taken to occur in the lower part of the Illawarra Coal
Measures. The Calderwood, Minumurra and Berkley Latite Members occupy similar
stratigraphic positions to that of the Five Islands Latite Member but the exact
relationship between the former three units and the Five Islands Latite Member is
unknown.
On the basis of similarity of petrography to some samples from Kiama and
Jamberoo, Chalmers (1941) mapped Flinders Islet as a remnant of the Bumbo Latite
Member. However, the Five Islands Latite Member is geochemically distinct from the
other latites (Carr, unpublished data) and on the basis of the inferred stratigraphic
relationships outlined previously, Flinders Islet is considered to be composed of the
Five Islands Latite Member and not the Bumbo Latite Member.
Calderwood Latite Member. The Calderwood Latite Member has not been recognized
previously. The name is here proposed for the unit which crops out around Calder-
wood (Fig. 1) and is 3 m stratigraphically above the Cambewarra Latite Member (i.e.
within, and near the base of, the Illawarra Coal Measures). The Calderwood Latite
Member is restricted in outcrop area and extends discontinuously from the north-
eastern side of Stockyard Mountain towards the north-northwest for a distance of 9
km.
A maximum thickness of 38 m occurs near Calderwood (GR 908728 Robertson
1:25,000 Topographic Sheet 9028-IV-N First Edition) but the unit thins rapidly
towards the north, south and west. A probably extrusive mode of emplacement is
indicated by the high proportion of volcaniclastic material in the sedimentary rocks
both above and below the unit. The volcaniclastic material above and below the
Calderwood Latite Member is possibly equivalent to Bowman’s (1974) Tappitallee
Mountain Tuff Member.
The outcrop on the northeastern side of Stockyard Mountain (GR 958685 Albion
Park 1:25,000 Topographic Sheet 9028-I-N First Edition) corresponds to the outcrop
which Jaquet et al. (1905) and Lowder (1964) mapped as Saddleback Latite Member
occurring at the top of the Cambewarra Latite Member. These authors were aware of
the anomalous stratigraphic position but considered that the Saddleback Latite
Member at this locality formed a topographic high at the time of eruption of the
Cambewarra Latite Member. The present investigation, however, indicates that the
outcrop on the northeastern side of Stockyard Mountain is separated from the un-
derlying Cambewarra Latite Member by 3 m of sandstone and shale of the Illawarra
Coal Measures.
Bowman (1970, 1974) did not record either the Saddleback Latite Member un-
derlying the Cambewarra Latite Member between Saddleback and Stockyard
Mountain, or the Calderwood Latite Member on the northeastern side of Stockyard
Mountain. However, the other outcrops of the latter unit were mapped as Dapto Latite
Member (Bowman, 1970, 1974). These outcrops occur above the Cambewarra Latite
Member and thus he concluded that the Dapto Latite Member is stratigraphically
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
PAUL F. CARR 295
above the Cambewarra Latite Member. In spite of Harper’s (1915) recognition of the
Cambewarra boulder bed as far north as Wollongong, Bowman did not mention the
possibility of another flow which was stratigraphically above the Cambewarra Latite
Member, and which cropped out in the Calderwood region.
Minumurra Latite Member. Jaquet et al. (1905) defined the Minumurra Flow which was
subsequently renamed the Minumurra Latite (Joplin e¢ a/., 1952). The incorrect name
Minnamurra Latite has also been used extensively (e.g. Lowder, 1964; Raam 1969;
Bowman, 1970, 1974). The unit occurs approximately 40 m above the base of the
Illawarra Coal Measures and extends as a discontinuous sheet from the northern side
of Saddleback Mountain to a point 15 km west of Port Kembla. A maximum thickness
of 35 m occurs at Minnamurra River.
The base of the Minumurra Latite Member is even but the top of the unit un-
dulates by up to 0.5 m in a horizontal distance of 3 m. Horizontally-bedded sand-
stones, shales and siltstones fill the depressions in the upper surface and individual
sedimentary units are thicker in the depressions than ove: the accompanying ridges
(Lowder, 1964) (GR 904647 Kangaroo Valley 1:25,000 Topographic Sheet 9028-IV-S
First Edition). An extrusive mode of emplacement is supported by the occurrence in
joints near the top of the unit of sedimentary material lithologically different from the
sedimentary rocks overlying the Minumurra Latite Member (Lowder, 1964). No
evidence in support of an intrusive mode of emplacement has been found previously or
in the present study. Since the Minumurra Latite Member occurs within the Illawarra
Coal Measures, the unit is considered to be an extrusion which erupted on to the
Permian land surface or into a shallow subaqueous environment.
Berkley Latite Member. Harper (1915) defined the Berkley Flow which was renamed the
Berkeley Latite (McElroy, 1952), and the Berkley Latite (Joplin e a/., 1952). The
incorrect spelling Berkeley has been used in many publications (e.g. McElroy, 1952;
Raam, 1969; Bowman, 1970, 1974).
The base of the unit ranges between 15 and 40 m above the top of the Shoalhaven
Group and thus occurs at approximately the same stratigraphic level as the Minumurra
Latite Member. This variation in stratigraphic position possibly reflects the
palaeotopography. The Berkley Latite Member has been considered to be intrusive
(McElroy, 1952) and extrusive (Harper, 1915; Rose, 1966; Raam, 1969; Bowman,
1970, 1974). The latter conclusion is supported by the present study for the following
reasons:
(i) clasts of Berkley Latite Member occur in the sandstone immediately above the
unit at Mount Kembla (GR 005876 Wollongong 1:25,000 Topographic Sheet
9029-II-S First Edition);
(ii) apart from localized draping of the bedding over the latite clasts, the sandstone
above the Berkley Latite Member shows no evidence of distortion of bedding as
would be expected to accompany intrusion of a large mass of igneous rock into
unconsolidated sediments (GR 012869 Wollongong 1:25,000 Topographic Sheet
9029-II-S First Edition); and
(iii) in the Figtree region, the Berkley Latite Member consists of two flows separated
by approximately 3 m of sedimentary rocks of the Illawarra Coal Measures (GR
026875 Wollongong 1:25,000 Topographic Sheet 9029-II-S First Edition).
The features described and cited as evidence for emplacement as a sill (McElroy, 1952)
possibly represent localized intrusive contacts produced by restricted intrusion of the
flow into unconsolidated sediments. The flow has a maximum thickness of ap-
proximately 35 m in the Figtree region and thins to the north and southwest.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
296 STRATIGRAPHY OF THE SOUTHERN SYDNEY BASIN
STRATIGRAPHIC RELATIONSHIPS BETWEEN THE DAPTO AND SADDLEBACK LATITE
MEMBERS
Harper (1915) considered the Dapto Latite Member to be a northern extension of
the Saddleback Latite Member. Since there is no evidence of continuity between the
outcrops at Saddleback Mountain and Port Kembla, and since the thickness of igneous
rock decreases away from a maximum for each area, the units are considered to have
been separate flows.
The top of the Cambewarra Latite Member or the equivalent boulder horizon
marks the top of the marine Shoalhaven Group (Harper, 1915). The contact between
the Saddleback Latite Member and the overlying Cambewarra Latite Member is
poorly exposed but no evidence has been found for the existence of sedimentary
material between the two latites. By comparison, the Dapto Latite Member is
separated from the Cambewarra boulder horizon by at least 3 m of sedimentary strata.
Thus both the Dapto and Saddleback Latite Members occur within the Shoalhaven
Group and occupy similar stratigraphic positions below the Cambewarra Latite
Member or the equivalent boulder horizon. Assuming that deposition of the boulder
horizon is essentially synchronous with eruption of the Cambewarra Latite Member,
then either (i) the Saddleback and Dapto Latite Members are of the same age; or (il)
the Dapto Latite Member is slightly older than the Saddleback Latite Member. The
implications of these equally plausible alternatives are discussed below.
Contemporaneous eruption of the Saddleback and Dapto Latite Members implies
that subsequent to the extrusion of these flows and prior to the emplacement of the
Cambewarra Latite Member and boulder horizon, sediment accumulated in the Port
Kembla region whereas no deposition occurred in the vicinity of Saddleback Moun-
tain. Alternatively, if the Dapto Latite Member is slightly older than the Saddleback
Latite Member, sediment accumulated on top of the Dapto Latite Member prior to,
and contemporaneously with, eruption of the Saddleback Latite Member. Subsequent
to the eruption of the latter flow, the Cambewarra Latite Member and boulder bed
were emplaced.
SUMMARY AND CONCLUSIONS
Strata of the upper Shoalhaven Group and lower Illawarra Coal Measures of the
southern Sydney Basin have had a varied nomenclature. Reappraisal of previous work
and recent mapping by the author have led to the recognition of nine distinct extrusive
igneous rock-units in the Wollongong-Gerringong region. These flows provide a
convenient basis for the subdivision of the strata of the upper Shoalhaven Group and
lower Illawarra Coal Measures. The uppermost formation of the Shoalhaven Group is
termed the Broughton Formation which contains five Latite Members — Blow Hole
(lowermost unit), Bumbo, Saddleback, Dapto and Cambewarra (uppermost unit). The
sedimentary strata between the Blow Hole and Bumbo Latite Members, and between
the Bumbo and Saddleback Latite Members are known as the Kiama Sandstone
Member and Jamberoo Sandstone Member respectively. Four Latite Members (Five
Islands, Calderwood, Minumurra and Berkley) are recognized in the Pheasants Nest
Formation of the Cumberland Sub-group of the Illawarra Coal Measures.
ACKNOWLEDGEMENTS
I am grateful to A. C. Cook, B. G. Jones and A. J. Wright for their helpful advice
and constructive criticism of the manuscript.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
PAUL F. CARR 297
References
Bowman, H. N., 1970. — Palaeoenvironment and revised nomenclature of the upper Shoalhaven Group
and Illawarra Coal Measures in the Wollongong-Kiama area, New South Wales. Rec. geol. Surv.
N.S. W., 12: 163-182.
—, 1974. — Geology of the Wollongong, Kiama and Robertson 1:50,000 sheets. Geol. Surv. N.S.W.,
Sydney.
CARD, G. W., and JAQUET, J. B., 1903. — The geology of the Cambewarra Range, New South Wales, with
especial reference to the volcanic rocks. Rec. geol. Surv. N.S.W., 7: 103-140.
CarR, P. F., and FACER, R. A., 1980. — Radiometric ages of some igneous rocks from the southern and
southwestern coalfields of New South Wales. Search, 11: 382-383.
CHALMERS, R. O., 1941. — The petrology of the Five Islands, Port Kembla, New South Wales. Rec. Aust.
Mus. , 21: 27-42.
Cook, A. C., 1966. — Report on the geology of Flagstaff Hill, Parish of Wollongong, New South Wales.
(Unpublished).
HARPER, L. F., 1915. — Geology and mineral resources of the Southern Coalfield. Part I. The south coastal
portion. Mem. geol. Surv. N.S. W., 7.
JAQUET, J. B., CARD, G. W., and HarpEr, L. F., 1905. — The geology of the Kiama-Jamberoo district.
Rec. geol. Surv. N.S. W., 8: 1-66.
Jopuin, G. A., HANLON, F. N., and Noakes, L. C., 1952. — Wollongong — 4 mile Geological Series.
Explan. Notes, Bur. Miner. Resour. Geol. Geophys. Aust.
LOWDER, G. G., 1964. — The geology of the Minnamurra Falls area. Sydney: University of Sydney, B.Sc.
(Hons) thesis, unpubl.
McCLunG, G., 1978. — Morphology, palaeoecology and biostratigraphy of Ingelarella (Brachiopoda:
Spiriferida) in the Bowen and Sydney Basins of eastern Australia. Pubs. geol. Surv. Qd, 365: 11-87.
McELRoy, C. T., 1952. — Evidence of the intrusive nature of the Berkeley Latite, Wollongong district,
N.S.W. Aust. J. Scv., 15: 100.
——,, BRANAGAN, D. F., RAAM, A., and CAMPBELL, K. S. W., 1969. — Shoalhaven Group. Jn PACKHAM,
G. H. (ed.), The geology of New South Wales. J. geol. Soc. Aust. , 16: 357-366.
RaAAM, A., 1964. — Geology of the Minnamurra-Gerroa area. Sydney: University of Sydney, B.Sc. (Hons)
thesis, unpubl.
——,, 1968. — Petrology and diagenesis of Broughton Sandstone (Permian), Kiama district, New South
Wales. J. sedim. Petrol. , 38: 319-331. :
——, 1969. — Gerringong Volcanics. Jn PACKHAM, G. H. (ed.), The geology of New South Wales. /. geol.
Soc. Aust. , 16: 366-368.
ROSE, G., 1966. — Wollongong 1:250,000 Geological Series Sheets SI 56-9. Geol. Surv. N.S. W., Sydney.
RUNNEGAR, B., and MCCLUNG, G., 1975. — A Permian time scale for Gondwanaland. Jn CAMPBELL, K. S.
W. (ed.), Gondwana geology, 425-441. Canberra: Australian National University Press.
WILSHIRE, H. G., and Hosss, B. E., 1962. — Structure, sedimentary inclusions, and hydrothermal
alteration of a latite intrusion. J. Geol. , 70: 328-341.
WILSON, R. G., 1969. — Illawarra Coal Measures. A. Southern Coalfield. Jn PACKHAM, G. H. (ed.), The
geology of New South Wales. /. geol. Soc. Aust. , 16: 370-379.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
Fossil Eucalyptus Remains from the Middle
Mauocene Chalk Mountain Formation,
Warrumbungle Mountains, New South Wales
W.B. K. HOLMES, F. M. HOLMES and H. A. MARTIN
HOLMES, W. B. K., HOLMES, F. M., & MARTIN, H. A. Fossil Eucalyptus remains from
the Middle Miocene Chalk Mountain Formation, Warrumbungle Mountains,
New South Wales. Proc. Linn. Soc. N.S. W. 106 (4), (1982) 1983: 299-310.
Eucalyptus fruits and leaves are preserved in the diatomite of the Middle Miocene
Chalk Mountain Formation (new name), Warrumbungle Mountains, New South
Wales. The fossil eucalypts show some features of advanced states in fruit and
venation and are compared with some extant species. Based on present knowledge of
Eucalyptus sensu lato, the fossils provide evidence of two separate phylogenetic lines
being present in Middle Miocene time. The microflora recovered from the diatomite
and associated lignite includes pollen grains of Myrtaceae (Myrtaceidites spp) of which a
few are referable to Eucalyptus.
W. B. K. Holmes, Hon. Research Fellow, Department of Geology, Unwwersity of New England,
Armidale, Australia 2351; F. M. Holmes, ‘Noonee Nyrang’, Gulgong Road, Wellington,
Australia 2820 and H. A. Martin, School of Botany, University of New South Wales, Ken-
sington, Australia 2033; manuscript received 8 June 1982, accepted for publication 18 August
1982.
INTRODUCTION
Despite the present day prominence of Eucalyptus and the great diversity of its
species, little is known of its evolutionary history from the fossil record (Martin, 1978;
Briggs and Johnson, 1979). For phytogeographical reasons alone, there can be little
doubt that the genus is of ancient Australian origin (Barlow, 1981); however, the
‘eucalypt-type’ pollen, Myrtaceidites eucalyptoides Cookson and Pike, which is probably
bloodwood-Angophora first appears in the Oligocene (Martin, 1981). Fossil leaves which
resemble eucalypts in gross form but which were inadequately described and figured,
have been recorded from many poorly-documented Tertiary localities in New South
Wales, Victoria, Tasmania, South Australia and Central Australia. References to
these occurrences are given in Duigan (1951). Lange (1978) reported a diverse
assemblage of Lepidospermae fruit, including a number of forms closely similar to
Eucalyptus, preserved as impressions on silicified slabs of unknown age from the arid
zone of South Australia. Ambrose et al. (1979) have also collected Eucalyptus fruit
similarly preserved to those of Lange. The age of these fossils from Stuart Creek,
northwest of Lake Torrens in South Australia is thought to be either Eocene-Oligocene
or Miocene.
Deane (1902) described several forms of eucalypt-like leaves from the Berwick
Quarry in Victoria. A re-investigation of this flora by Nelson and Webb (in prep.) has
shown that the cuticle of one of the leaves is similar to the cuticle of some extant
Eucalyptus. Impressions of a eucalypt-like fruit and of rainforest leaves are also present
but no Eucalyptus type pollen has been observed in the microflora which is dominated
by Nothofagus pollen (Nelson and Webb, pers. comm.).
Eucalyptus fruit and leaves and the microflora from the Middle Miocene diatomite
deposit at Bugaldie near Coonabarabran in northwestern New South Wales are
described below. Associated with the eucalypts are remains of leaves and flowers of
plants (Holmes, in prep.) which have affinities with extant plants growing in situations
with a much higher rainfall than the present Bugaldie average annual rainfall of 650
mm. Also in the diatomite are fossils of fish (Hills, 1946), a bird described as an owlet-
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
300 MIDDLE MIOCENE FOSSIL EUCALYPTUS
nightjar (Rich and McEvey, 1977), diatoms (Thomas and Gould, 1981 a and b),
insects and freshwater bivalves.
GEOLOGY AND AGE OF THE DIATOMITE
Chalk Mountain, which lies 6 km south of Bugaldie village, is a remnant of an
extensive dissected plateau of late Permian, Triassic and Jurassic freshwater sediments
capped by basalt associated with the Warrumbungle Volcano Complex to the south.
Lacustrine sediments which are interbedded between basalt flows on the Jurassic
sediments and a 25 metre thick overlying basalt flow, outcrop around the escarpment
of Chalk Mountain. Diatomite in these sediments was commercially exploited from
1919 until 1980. Griffin (1961) gave a detailed description of the geological and
commercial aspects of the deposit but he did not name the formation. Reports by
Herbert (1968a and b) deal with the diatomite.
Chalk Mountain Formation (Holmes and Holmes, new name). The Chalk Mountain
Formation is a lake deposit over 15 metres in thickness and is composed of almost
horizontal layers of mudstone, clay, diatomite, three interbedded horizons of tuf-
faceous clay and a thin band of lignite. The type locality is Quarry A on the western
face of Chalk Mountain immediately below the basalt flow capping the mountain (grid
reference 199148, Gilgandra 1: 250 000 Geological Series Sheet SH 55-16). The base
of the formation is obscured at this locality by the basaltic talus slope and by quarry
waste. On the roadway up the northern slope of the mountain the basal beds of
mudstone and clay rest on basalt. The section exposed in Quarry A is shown in Table
I,
The area of lake sediments that have disappeared due to erosion around Chalk
Mountain is unknown, but it is likely that the original lake was very much larger than
the 38 hectares of the existing deposit. Plant macro-fossils have been collected only
from the diatomite. Plant micro-fossils occur in both the diatomite and the thin band of
lignite near the top of the formation. For a horizontal distance of about thirty metres in
from the outcropping face, the diatomite has weathered to a pure white lightweight
material much of which has been removed by open-cut mining and some tunnelling.
Beyond the weathered zone the diatomite is of a grey to black colour due to the
presence of organic matter and is known as ‘black earth’. The ‘black earth’ was only
sparingly exploited due to the necessity to process it to remove the organic matter. In
the weathered zone of the diatomite the plant macro-fossils are preserved as colourless
or limonite-stained impressions and in the unweathered zone as colourless impressions
TABLE 1
Type Section of Chalk Mountain Formation exposed in Quarry A on Chalk Mountain, Bugaldie (after Griffin, 1961)
Basalt
Clay with lignite band
0.91 m Tuff, clay and grit
4.05 m Clay and diatomite
0.13 m Tuff, clay, diatomite and gritty sandstone
3.04 m Diatomite high grade
0.08 m Clay and tuff
Diatomite high grade
Mudstone, tuff and clay
Bottom Basalt, trachyte and andesite overlying
Jurassic Pilliga Sandstone
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
W.B.K. HOLMES, F. M. HOLMES AND H. A. MARTIN 301
or carbonaceous compressions. The diatomite exhibits seasonal varves each 0.33 mm
to 0.5 mm thick (Thomas and Gould, 1981b). A deposition time of between 14 000 and
21 000 years would have been needed to accumulate the total thickness of 7 metres of
diatomite, assuming the varves to be annual.
The bands of tuff within the deposit indicate contemporaneous eruptive volcanic
activity during the period of lacustrine sedimentation. K/Ar dating of igneous rocks
from localities around the Warrumbungle complex (Dulhunty and McDougall, 1966;
McDougall and Wilkinson, 1967; Wellman and McDougall, 1974) show that the
volcanic activity took place during the period 17 million years to 14 million years
before the present. The nearest sites to Chalk Mountain with K/Ar dates are Looking
Glass Mountain (16.6 + 0.6 m.y.), 15 km to the west-northwest, and Woorut
Mountain (15.5 + 0.4m.y.), 20 km to the south (Wellman and McDougall, 1974). As
no study has been made of the geology of the northern portion of the Warrumbungle
Volcano, the stratigraphic relationships of these localities to Chalk Mountain are not
known. Therefore, with present information, we can only date the diatomite as being
within the age limits of the active Warrumbungle Volcano — 17 million years to 14
million years, Middle Miocene.
THE FOSSIL EUCALYPTS
Family MYRTACEAE
Genus Eucalyptus L’ Herit.
Eucalyptus bugaldiensis Holmes and Holmes sp. nov.
Fig. 1 A, Fig. 2 A-D
Diagnosis: Fossil infructescence of a group of three umbellasters of seven or fewer small,
ES?
SS Z
Fig. 1. A Eucalyptus bugaldiensis sp. nov. AMF61713 holotype x 2. B Eucalyptus sp. Leaf form B AMF61721
x 2.C Eucalyptus sp. Leaf form A AMF61724 x 2.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
302 MIDDLE MIOCENE FOSSIL EUCALYPTUS
hemispherical, ribbed, woody fruit attached by tapering pedicels to peduncles. Rim of
fruit broad; exerted valves broad and low.
Description: Infructescence of three umbellasters on peduncles 5 mm - 10 mm in length,
and 0.8 mm in width attached at a common point to a stem. Umbellasters composed of
seven or fewer fruit attached to the peduncle apex by pedicels (anthopodia) 5 mm - 10
mm in length widening upwards to merge into the base of the fruit. Fruit
hemispherical, 4 mm - 5 mm in diameter. External surface ornamented by 7 - 10
longitudinal ribs which occasionally fork near the rim. Rim flat ca 1 mm in width.
Exerted valves forming a broad low triangular projection 0.5 mm above the rim.
Type material: Holotype AMF61713 and counterpart AMF61714. Paratypes
AMF61715-20. The Australian Museum, Sydney, N.S.W., Australia.
Type locality: Diatomite beds in Quarry A on Chalk Mountain, 6 km south of Bugaldie
village near Coonabarabran, New South Wales. Grid reference 199148, Gilgandra 1:
250 000 Geological Series Sheet SH 55-16. Chalk Mountain Formation, Middle
Miocene.
Discussion: The fossil fruits of Eucalyptus bugaldiensis are preserved as impressions in the
soft diatomite. The hollow moulds contain no organic material. The surface of the
diatomite was hardened with Bedacryl 122X and latex casts were then taken of the
holotype and its counterpart. When whitened with ammonium chloride, the latex casts
revealed some of the finer details not readily seen in the original specimens. The distal
ends of the peduncles show scars where additional buds or fruits have been aborted or
lost during development or preservation. The umbellaster on AMF61715 has five fruit
attached by pedicels to the peduncle. In addition to the still-attached fruit, the distal
end of the peduncle shows the abscission scars of another two pedicels. ‘This suggests
that the total flower number per umbellaster was seven, a number which may be ex-
pected in dichasial inflorescences in a family with primarily opposite and decussate
phyllotaxy. Extant species in subgenus Symphyomyrtus very often have seven flowered
umbellasters. The arrangement on the holotype of three umbellasters about a common
point, is found in the eucalypt conflorescence sub-types of T,; and 83 of Johnson (1972,
fig. 3). The T, sub-type characterizes the small tropical subgenus Telocalyptus (section
Equatoria) and the S3 sub-type occurs mainly in the subgenus Eudesmia, but the fossil
fruits are quite different from any in Eudesmia. However, three umbellasters can be so
attached in some variants of sub-type S, also (L. A. S. Johnson, pers. comm.) when the
phyllotaxy in the inflorescence region is opposite rather than disjunct-opposite (Briggs
and Johnson, 1979), as for instance in some inflorescences of Eucalyptus microtheca of
subgenus Symphyomyrtus section Adnataris. The infructescence and the individual fruits
of E. bugaldiensis are very close in form to some extant species of Eucalyptus. However,
due to the type of preservation and to the lack of supporting organs, this new species
cannot at present be placed with certainty in any particular subgenus.
The fruits of E. microtheca F. Muell. (Blake, 1953, pl. 30, figs 20-36) are closely
similar in size and form to those of E. bugaldiensis. E. microtheca is very widely
distributed throughout the continent (Blake, 1953, map 24) on ground that is subject to
seasonal flooding. E. raveretiana F. Muell. from areas near the coast in central and
northern Queensland has similar but rather smaller fruit.
None of the diverse forms of fossil Eucalyptus fruits from South Australia (Lange,
1978; Ambrose et al., 1979) appear to be similar to E. bugaldiensis but their different
aspect of preservation makes comparisons difficult. The cluster of fruits from Berwick,
Victoria (Nelson and Webb, in prep.) is superficially similar to £. bugaldiensis but lacks
evidence of valves.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
W. B. K. HOLMES. F. M. HOLMES AND H. A. MARTIN 303
S
aN
Fig. 2. A-D Eucalyptus bugaldiensis Holmes & Holmes sp. nov. Holotype. A, B AMF61713,A x 1;B x 4.6,
D AMF61714 counterpart of AMF61713,C x 1;D x 4.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
MIDDLE MIOCENE FOSSIL EUCAL YPTUS
C, D Eucalyptus sp. Leaf form B
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Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
W.B.K. HOLMES, F.M. HOLMES AND H. A. MARTIN 305
Eucalyptus sp. Leaf-form A
Fig. 1 C, Fig. 3 A-B
Description: Leaf lanceolate to narrow lanceolate, falcate, asymmetrical about lamina
base; margin scarious, entire, sometimes insect-damaged. Length 8 cm-14 cm; width
1.5 cm-2 cm, widest at one quarter of length from the base and tapering gradually to
an obtuse apex. Petiole curved, to 2 cm in length. Midvein prominent, tapering from
the petiole to the apex. Secondary veins parallel, ca 1 mm apart, sometimes with a
minor vein in between; attached to the midvein at 50°-60° and running straight to the
intramarginal vein which runs close and parallel to the leaf margin. Tertiary veins
form an irregular reticulum of about four rows of areoles between the secondary veins.
Material: AMF61721-3.
Discussion: The venation pattern of leaf-form A is of the Transversae class (Cambage,
1913; Maiden, 1922) (not to be equated with the section Transversaria of Pryor and
Johnson, 1971) and is present in the subgenera Corymbia and Blakella of Eucalyptus and
in Angophora which Pryor and Johnson (1971) have suggested is also of subgeneric rank
in Eucalyptus s.1. Extant species with venation similar to leaf-form A are the widespread
Angophora floribunda (Sm.) Sweet and Eucalyptus trachyphloia F. Muell., a bloodwood
tree, which grows on the drier sandy soils of the Upper Hunter River and Pilliga areas
of New South Wales and into Queensland.
Eucalyptus sp. Leaf-form B
Fig. 1 B, Fig. 3C-D
Description: Leaf lanceolate, slightly falcate; margin entire; apex obtuse; lamina base
asymmetrical. Width 16 mm-23 mm; length 8 cm-12 cm. Secondary veins sub-
parallel, spaced irregularly, 5-6 per cm, running a slightly undulate course to the
intramarginal vein at an angle of 55°-60° to the midvein. Secondary veins occasionally
fork and anastomose to form a very elongate reticulum. Petiole curved, to 2 cm in
length.
Material: AMF61724-5. MMF15284.
Discussion: MMF15284 (Fig. 3 C) shows portions of five leaves, one of which is at-
tached to a slender stem which has sub-opposite projections from where the other
leaves may have become detached. Leaf-form B differs from leaf-form A by the wider
spaced and irregular course of the secondary veins. The venation pattern is in-
termediate between the Transversae and Obliquae classes (Cambage 1913; Maiden
1922). This general pattern of leaf venation is present in extant species within the
subgenus Symphyomyrtus section Adnataria series Oliganthinae or series Pruinosae and in the
subgenus Telocalyptus series Degluptae. Leaves of Eucalyptus raveretiana F. Muell. (a
member of Telocalyptus) bear a close resemblance to this fossil form.
PALYNOLOGY OF THE CHALK MOUNTAIN FORMATION (H.A.M.)
Table 2 lists the taxa found in the ‘black earth’ diatomite and in the lignite from
near the top of the formation. Preservation is rather poor which limits identifications,
and this may be the result of initial poor conditions at the time of deposition or of
subsequent weathering on exposure. Nevertheless, the assemblages are workable and
invaluable because they can be dated by independent evidence.
The two assemblages are essentially the same, although there is a quantitative
difference between the diatomite and the lignite. This difference could result from the
different conditions of deposition. Diatomite is formed in lakes and so would ac-
cumulate more wind-blown pollen. Because lignite forms in swamps most of the pollen
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
306 MIDDLE MIOCENE FOSSIL EUCALYPTUS
TABLE 2
The Pollen Frequencies
Fossil Name
ALGAE
Botryococcus braunu Kitzing
FERN SPORES
Cyathea paleospora Martin
Deltoidospora inconspicua Martin
Laevigatosporites ovatus Wilson & Webster
Polypoditdites sp.
Reticuloidosporites minisports Martin
Unknown spore
GYMNOSPERMS
Araucariacites australis Cookson
Cupressaceae sp. indet
Dacrydium florinu (Cookson & Pike) Cookson
Phyllocladus palaeogenicus Cookson & Pike
Podocarpus australiensis (Cookson & Pike) Martin
Podocarpus elliptica (Cookson) Martin
ANGIOSPERMS
Cupanierdites orthoteichus Cookson & Pike
Encipites sp.
Graminidites media Cookson
Haloragacidites harrisit (Couper) Harris
Loranthaceae
Malvacipollis diversis Harris
Milfordia sp.
Monosulcites cf Palmae
Myrtaceidites eucalyptordes Cookson & Pike
Myrtaceidites cf M. eucalyptordes Cookson & Pike
M. parvus Cookson & Pike
Myrtaceae not identified further
Nothofagus aspera Cookson
N. emarcida Cookson
Polyporina chenopodiaceoides Martin
Proteacidites sp.
Quintinia psilatispora Martin
Tnicolporites scabratus Harris
Tricolporites cf Cunoniaceae
Tricolporites cf Goodeniaceae
Tricolporopollenites wvanhoensis Martin
Tniontes minisculus McIntyre
T. orbiculatus McIntyre
Tnorites cf Ulmaceae
Unknown tricolpate/tricolporates
Number of kinds of unknowns
Botanical Affinity
Botryococcus brauni
Cyathea spp.
Total Fern Spores
Agathis and Araucaria
Cupressaceae
Dacrydium Sec. B
Phyllocladus spp.
Podocarpus Sect. Dacrycarpus
Podocarpus, most other sections
Total Gymnosperms
Tribe Cupaneae (Sapindaceae)
Epacridaceae
Gramineae
Casuarina
Loranthaceae
Austrobuxus— Dissiliaria (Euphorbiaceae)
Restionaceae
Like Palmae
Bloodwood—Angophora, possibly other genera
Possibly other eucaly pts
Possibly T71stanta—Backhousta—Baeckia and other genera
Myrtaceae
Total Myrtaceae
Nothofagus, the menztesu type (e.g. N. moorez,
N. cunninghamii)
Nothofagus, the brassi type
Total Nothofagus
Chenopodiaceae and some Amaranthaceae
Proteaceae
Quintinia
like Cunoniaceae
like Goodeniaceae
probably Rutaceae
like Apananthe— Celtis
Notes:
(1) Subtotals may not add up exactly because individual percentages have been rounded off.
(2) + indicates a presence but not counted.
(3) The use of ‘like’ indicates a tentative identification.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
Diato- Lig-
mite nite
%o %
+
it 58) 29
i129) 2.4
1.5 5.6
0.5
0.2
paren Mae:
46 17.2
10.7. 18.1
0.8 DP
4.6 1.0
0.5
5.3 5.4
14) WO.)
29.0 34.8
+
1.2
0.2
13.0 24.0
0.2
a
+
0.7
0.5
0.8
6.8 0.2
S75 5.2
25.2 3.2
0.8 0.5
APIO oe
3 0.5
0.8
+
2.0
3.0
0.7
0.2
0.2
0.2
1.0
0.2
22.1 13.2
No. No.
7 6
W. B. K. HOLMES, F.M. HOLMES AND H. A. MARTIN 307
would come from the plants growing in and around the edges of the swamps. Fern
spores are more abundant and diverse in the lignite. The swamp environment would
be more suitable for fern growth. Gymnosperms are well represented in both
assemblages but are slightly more abundant in the lignites and of this group,
Araucariaceae are the most common. Casuarina is almost twice as abundant in the
lignites. Some species of Casuarina grow on the swamp edges today.
The Myrtaceae are more abundant in the diatomite than in the lignite. Un-
fortunately, the identification of pollen to genera within this family is not particularly
reliable. The bloodwood-Angophora type is only found in the lignite in a very low
concentration. Judging from surface samples, Rose (1981) found that this pollen type
does not disperse aerially far from the source of production. However, water transport
cannot be ruled out; but swamp vegetation would act as a filter and prevent the pollen
from travelling far (Birks and Birks, 1980). The other ‘possibly eucalypt’ type is
smaller than the bloodwood-Angophora type, and Rose found that it is more widely
distributed. This second type has been found only in the diatomite. Myrtaceous pollen
which is not like that of eucalypts is also present (see Table 2). The majority of the
pollen cannot be identified beyond the family level because of poor preservation or
crumpling which obscures diagnostic features.
Nothofagus is present but in very low concentration, indicating transport from a
distant source. The small amount of the brassi type may have come from plants
growing a long way from the catchment of this lake. Little can be said of the taxa
present in low frequencies, for these are usually low pollen producers. The in-
determinate tricolpate/tricolporate group is quite abundant. These pollen assemblages
must represent forests, judging from the low content of herbaceous elements such as
Gramineae and the absence of Compositae.
The high content of Myrtaceae in the diatomite together with the low content of
Nothofagus in both samples fits the Myrtaceae phase of the Pliocene (Martin, 1973).
Malvacipollis diversus and Triporopollenites ivanhoensis are usually found only in the lower
part of the older phase (Martin, unpubl.). The Pliocene age was based only on the
general geology of the Riverina region (Martin, 1973), so it may not be reliable. None
of the diagnostic species of the mid-late Miocene Trporopollenites bellus Zone, described
from the Gippsland Basin of south eastern Victoria (Stover and Partridge, 1973), has
been found at Chalk Mountain and the general description of this Zone is a poor fit for
the present assemblages. The base of this Zone contains abundant Nothofagus which
decreases upwards through it. Its upper limit possibly extends into the Pliocene but the
relationship to the overlying Myrtaceae phase (older) has not been documented. The
problem is one of geographic variation during the Tertiary. In time-equivalent
assemblages, Nothofagus is more abundant in southeastern Victoria than in New South
Wales. Conversely, Myrtaceae are more abundant in New South Wales, particularly
in the most westerly sites (Martin, 1983). Since the change from an assemblage with
high Nothofagus to one with high Myrtaceae depends on the climate becoming drier,
this change should have occurred earlier in the drier, inland regions than on the wetter
coastal southeastern part of Australia. The Chalk Mountain microfloral assemblages
show that the change had occurred in this region by the mid Miocene, some 14-17
million years ago.
THE MIDDLE MIOCENE CHALK MOUNTAIN PALAEOENVIRONMENT
The Tertiary pollen record is largely that of rainforest assemblages (Martin, 1978,
p. 200) and most deposits of plant macro-fossils show a preponderance of mesophyllous
leaves which constituted the ‘Cinnamomum’ flora (Sussmilch, 1937). The preservation
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
308 MIDDLE MIOCENE FOSSIL EUCAL YPTUS
of pollen and other plant remains as fossils usually depends on the presence of lakes,
swamps or bogs. Therefore, a strong bias exists against the fossilization of plants
adapted to growing in drier and harsher environments. Beadle (1981) has suggested
that scleromorphic plants began to differentiate at the margins of rainforests and
developed along declining soil-fertility gradients. Johnson and Briggs (1981) discussed
the possibility of differentiation of scleromorphic taxa during the Early Tertiary in
nutrient-deficient forest sites. The large tracts of low-fertility soils derived from the
underlying Jurassic Pilliga Sandstone surrounding the Warrumbungles would have
provided such an environment.
At the present time eucalypts characteristically occur in open forest and woodland
associations and only a few species have been successful in rainforest margins (Barlow,
1981). In addition to the eucalypt remains, the Chalk Mountain fossil flora includes
fern fragments, podocarp leaves, Ceratopetalum sp. flowers and myrtaceous, lauraceous
and other leaves that show similarities in gross form with present-day rainforest species
in high rainfall areas of northeastern New South Wales and eastern Queensland. This
mixed fossil flora should not be regarded as evidence that eucalypts in the Middle
Miocene were adapted to rainforest conditions. Two present-day environmental
situations provide a similar mixture of leaves. In moist sclerophyll forests on the North
Coast of New South Wales, rainforest will develop, in the absence of fire, beneath tall
Eucalyptus trees such as E. grandis W. Hill ex Maiden, E. saligna Sm. and occasionally
E. microcorys F. Muell. Along the coastal strip of New South Wales and Queensland,
rainforest species sometimes form a fringing forest along the banks of watercourses.
Sclerophyll forest abuts and often overhangs this rainforest. Leaf-litter in the stream
bed is a mixture of eucalypt and rainforest plant remains.
At Chalk Mountain in Middle Miocene time, the rich soils derived from the
basalt, the moist conditions adjacent to the lake, and the probable higher rainfall
(Kemp, 1978) would have favoured rainforest growth. Sandstone hills around the lake
and areas away from the soil-improving influence of the basalt would have supported a
sclerophyll forest or woodland to provide the eucalypt, herb and grass remains that
would have been carried by wind or water to mingle with the rainforest remains in the
lake sediments. The high percentage of gymnosperm and Casuarina pollen probably
indicates the proximity of these plants to the site of deposition.
CONCLUSION
On the basis of leaf venation, the Chalk Mountain fossils show that by the Middle
Miocene the Eucalyptus genus s.1. had differentiated into at least two of the phylogenetic
groups as proposed by Johnson (1972, fig. 1). The fossil eucalypts show some features
of advanced states in fruit and venation (L. A. S. Johnson, pers. comm.). This
suggests that a suitable environment for their development must have existed very
much earlier in the Tertiary. The paucity of eucalypt remains in the fossil record
probably reflects the lack of opportunity for scleromorphic plants to become fossilized.
The lack of similarity between the Miocene pollen assemblages from the Gipp-
sland Basin and Chalk Mountain is probably due to geographic and climatic factors
rather than a great difference in age of deposition.
K/Ar dating of the basalts above and below the Chalk Mountain Formation is
desirable to determine the exact age of the locality.
ACKNOWLEDGEMENTS
The authors wish to thank Mr V. Mills and Mr and Mrs G. Hawker of Bugaldie
for assistance and information on the Chalk Mountain area; Dr J. W. Pickett and Dr
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
W.B.K. HOLMES, F.M. HOLMES AND H. A. MARTIN 309
A. Ritchie for the loan of specimens from the Mining and Geological Museum and the
Australian Museum respectively; Mr R. S. Jones for preparing the latex casts; Mr P.
J. Althofer for making available his private collection; Professor N. C. W. Beadle, Mr
J.B. Williams, Dr A. G. Floyd, Dr L. A. S. Johnson and Mr D. F. Blaxell for helpful
discussion on the relationship of the fossils to extant species; Professor J. F. G.
Wilkinson for information on the Warrumbungle Volcano; Mr D. Nelson and Dr J. A.
Webb for useful comments and for information on their unpublished investigations of
the Berwick fossil eucalypts. W. B. K. Holmes gratefully acknowledges support from
the Science and Industry Endowment Fund.
References
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RICH, P. V., and McEvey, A., 1977. — A new owlet nightjar from the Early to Mid-Miocene of Eastern
New South Wales. Mem. Nat. Mus. Vict. 38: 247-253.
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Univ. N.S.W., B.Sc. (Hons) thesis, unpubl.
SToveR, L. E., and PARTRIDGE, A. D., 1973. — Tertiary and late Cretaceous spores and pollen from the
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Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
Long distance ‘Transport of Spores of Puccinia
graminis tritici in the southern Hemisphere
I. A. WATSON and C. N. A. DE SOUSA
WaTSsON, I. A., and DE Sousa, C. N. A. Long distance transport of spores of Puccinia
graminis tritici in the southern hemsiphere. Proc. Linn. Soc. N.S.W. 106 (4), (1982)
1983: 311-321.
Evidence is presented that viable uredospores of the wheat stem rust fungus
Puccinia graminis tritict reached Australia from Africa in 1968-69. Two strains 326-
1,2,3,5,6 and 194-1,2,3,5,6 both very dissimilar from any previously found in
Australia were collected in 1969 from Clinton, S. Australia and Tichborne, N.S.W.
respectively. These strains or derivatives from them were found to be
indistinguishable from their African counterparts. The weather systems responsible
for the transport of the spores across the Indian Ocean are discussed.
I. A. Watson, 49 Old Castle Hill Road, Castle Hill, Australia 2154, and C. N. A. de Sousa,
EMBRAPA, Passo Fundo RS 99100, Brazil (both formerly of the Department of Agricultural
Botany, The University of Sydney, Sydney); manuscript received 21st April 1982, accepted for
publication 21 July 1982.
INTRODUCTION
Rust pathogens of cereals are widely distributed throughout the world and survive
on native or introduced host plants. Regardless of the geographical area, these
pathogens exist as a multiplicity of strains differentiated by their pathogenicity,
morphology, physiological and other characters. The origin of this variability can be
attributed to a number of different causes. Both sexual and asexual processes may
proceed side by side in some countries but where the alternate host is insignificant,
mutation can explain most of the variation.
It is generally recognized that where geographical areas are isolated the rust flora
of those areas becomes characteristic of them. The extent of the differences between
areas will depend on the amount of recombination occurring within them due probably
to sexual reproduction in the fungus and on the survival of mutations. One of the most
outstanding examples of this geographical isolation is seen in the linseed rust fungus
Melampsora lini (Pers.) Lév. When comparisons are made between isolates from North
America and Australia, Bison is found to be susceptible in the former but highly
resistant to a group of strains in Australia. Strains avirulent on Bison in Australia
commonly survive on the native flax Linum marginale Cunn. which has not yet been
shown to have specific genes for resistance.
It has already been pointed out (Watson, 1977) that on a global basis the total
variation in Puccinia graminis f.sp. tritict Eriks. and Henn. that can be demonstrated in
any one area is related not only to sexual reproduction and mutation but to the extent
of wheat breeding activities especially where programs have made use of gene specific
resistance. Reports appearing in the literature however do not always give the true
picture of the range of variability, since the latter is revealed only when suitable single
gene differentials are used in the survey work. In Puccinia graminis tritict maximum
variability occurs in North America where the alternate host is common and a large
effort is expended on breeding resistant cultivars using the well documented Sr genes.
There is thus extensive variation around the genes of the standard set of differentials
and also around the Sr genes of the parents used by the breeders.
In Australia by contrast, there are few standard races, but a multiplicity of
components have evolved within each following mutation and the cultivation of
resistant wheats of diverse genotypes. Once these variants of the population have been
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
312 LONG DISTANCE TRANSPORT OF SPORES
catalogued and described any spectacular deviations from them can be easily
recognized and will indicate that something other than normal evolution has occurred.
‘TRANSPORTATION OF SPORES
(1) Over Land areas. Pathogen gene flow between regions will tend to have a unifying
effect on their rust flora. The effectiveness of wind transport of spores in contributing
to this uniformity will depend on the proximity of the regions and any impediments
offered by natural barriers. It is suggested that eastern and western Europe may be
different regions (Hogg et al. , 1969) but the rapid spread of race 77 of P. recondita Rob.
ex Desm. following the release of wheats with the IB/IR translocation indicate that
there is a ready exchange of spore material over most of this land area. The distances
covered by any one spore movement are probably relatively short and seldom exceed
500-700 km (M. Boskovic, personal communication).
It has been known for a long time that spores of P. graminis tritici are transported
on the North American continent along the Puccinia pathway, but the extent of the
distances covered in any one step is not clear from the reports. As in Europe it would
appear to be of the order of several hundred km (Hogg et al. , 1969).
On the Australian continent the movement of spores of P. graminis tritici over
much longer distances and in different directions has been well documented over the
last 25 years (Hogg et al., 1969). The results from earlier work were confirmed in 1973.
In that year practically no stem rust was found in Western Australia but there was an
epidemic in the eastern states. The composition of the flora in both the east and west
was well known. Sometime after this epidemic 12 strains of eastern origin were isolated
in Western Australia for the first time (Luig, 1977). Presumably the spores were
caught in the anticyclones which may extend for 6,000 km and which, in the
November-April period, feature easterly winds over the continent. Spores must have
been transported about 3,500 km from east to west over the harsh inland because there
are no congenial hosts between the areas which were the source of the spores and the
points of new infections.
(ii) Over Seas. Adjoining countries seldom present any barriers to the movement of
rust spores but as they become increasingly separated by seas such as the
Mediterranean, the Baltic and the Tasman, the possibility of spore transport from one
to the other is reduced. From the summary presented by Hogg ¢t al. (1969) it is clear
that movement of rusts across water is readily occurring in Europe and spores in transit
have been trapped in the atmosphere. While there is no reason to doubt this movement
of P. graminis tritici the designation of the strains involved has not been very precise as
only the broad categories of standard race numbers were known.
Hirst and Hurst (1967) believe that to establish proof of aerial dissemination of
spores there needs to be ‘incontrovertible association between the source and new
colonies’ or ‘interception of migrant spores’. Work in the Australia-New Zealand area
has adhered rigidly to the first of these requirements and where spore movement of
rusts is postulated, the extensive genetic markers in the material at the source and at
the new site have been identical as far as is known.
There are several reasons why, despite the extent of the land and water in this
geographical area, the rust organisms can be so accurately characterized. First, the
annual race survey has been in progress for 60 years and for much of that time the
classification of the strains has been very precise because the genes involved in dif-
ferentiation have included those of the standard series as well as those in the Australian
supplementals. Consequently it has been possible to relate to some extent the local flora
with that found overseas. Second, and very important, Australia is an isolated con-
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
I. A. WATSON ANDC.N.A.DESOUSA 313
tinent, politically uniform so that collections from a vast area can be made and
examined at will. Since wind currents transport spores around the country the national
wheat crop in years of favourable climatic conditions must serve as a huge biological
trap to snare not only local inoculum but an occasional viable spore that may reach the
continent from elsewhere. Because of this isolation it has been possible to construct an
evolutionary pathway for the rusts of the Australia-New Zealand area in which the
important events have been underlined (Watson, 1981; Burdon et al. , 1982).
A third factor in the build-up of knowledge on rust epidemiology is the continuous
documentation of the population structure of the New Zealand rusts. Arrival of a new
rust in Australia is followed after a short interval of some months by its isolation in
New Zealand (Hogg é¢ al., 1969; Luig, 1977). There is now irrefutable evidence that
cereal rust spores are transported across the Tasman Sea at frequent intervals, a
distance of at least 2,000 km.
(iii) Over Oceans. There are as yet no reports of wind-borne inoculum of P. graminis
tritict having crossed any of the world’s oceans. With the discovery of the coffee rust
organism (Hemuileia vastatrix Berk. & Br.) in Brazil in 1970 it was suggested (Bowden et
al., 1971) that since trade winds blow frequently and for long periods across the
Atlantic Ocean spores of this organism had been transported from the African con-
tinent. Since the strains observed on the two continents showed similar characters this
appeared likely but Ingold (1978) believes there is some doubt as to whether this
organism is normally dispersed by wind.
Australian workers have speculated on the possible overseas origin of several
strains of P. graminis tritict that have successfully colonized this country. Luig (1977)
discusses this in some detail but gives no data to pin point the origin of any strain
common to the area. The original six standard races found 60 years ago were
characteristic of Australia but no origin ‘for them has been suggested. Luig (1977)
described how these and other rusts were replaced with the passage of time by more
aggressive strains. Three periods 1925, 1948-54 and 1968-69 are important in this
replacement.
Entry of new rusts to the country at those times probably involved wind transport
of spores across the great distances of the Pacific or Indian Oceans. The possibility of
accidental or deliberate introduction by man must be largely discounted since com-
munication between the critical areas of Australia and Africa would be minimal.
In the period 1948-54 standard race 21 was isolated as a type completely new to
Australia. This race was at the time among the most common in the world but there
were no comparative data from overseas to suggest from where it came. Almost 20
years later in 1968-69 further dramatic changes occurred in the population of P.
graminis tritici in Australia. Intensive investigations were commenced to uncover the
possible origin of the strains involved in this latest event.
ISOLATION OF STRAINS FROM AFRICA AND A COMPARISON WITH THEIR
AUSTRALIAN COUNTERPARTS
(i) Early suggestions of Similarities. Luig (1977) has already outlined some of the details
concerning studies of two Australian strains of P. graminis tritict which differed
markedly from any previously isolated in this area. Both collections had been made in
1969 by the senior author, one 326-1,2,3,5,6 from Clinton, S. Australia, the other
194-1,2,3,5,6 from Tichborne, N.S.W. The material on which each was collected had
few large isolated pustules and carried the gene S76.
Although these collections were part of the annual race survey conducted by Luig
and Watson the identifications revealed something unique. The first clue that overseas
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
314 LONG DISTANCE TRANSPORT OF SPORES
inoculum may have been involved in the origin of these two rusts came from data sent
by Dr J. C. Santiago who was then at Elvas, Portugal. These data showed that stan-
dard races 194 and 222 were present in Mozambique and there was evidence that
pathogenicity on some lines of the Australian supplementals was the same as that of the
two new local rusts. In order to make detailed comparisons of rusts from the two
continents inoculum was requested from Angola, Rhodesia (now Zimbabwe) and
Mozambique.
(ii) Detailed studies of Pathogenicity of African rusts. Two collections were received from
Angola, one from Nova Lisboa the other from Caconda. A single collection was sent
from Salisbury, Rhodesia. Because these collections were from overseas, strict
quarantine precautions were taken during the studies with them. All tests were made in
glasshouses at the University of Sydney, Glebe, thus minimizing the possibility of
spores escaping and lodging on a congenial host. At the conclusion of each test the
plant material and the rust were collected and destroyed.
The strain components of the three collections were identified using the dif-
ferentials of Stakman et al. (1962), supplemented by the eleven local differentials
adopted in Australia by Luig and Watson (1977). After the initial multiplication of
spores, inoculum was placed onto certain key varieties vzz. Marquis (S77b), Reliance
(S75), Acme (S79g), Einkorn (S721), Vernal Emmer (S79e), McMurachy ($76), Yalta
($711) and C.1.12632 (SrTt,). Many isolations were made and finally from single
pustules five strains were identified from Nova Lisboa and one from Caconda. ‘Two
strains were found in the Salisbury material. According to the classification of Luig and
Watson these were designated as in Table 1 and the seedling infection types on certain
wheats are given in Table 2.
The two Rhodesian strains were differentiated on Acme and Barleta Benvenuto
and no differences could be found between Rhodesia 2 and Angola 4. Although all
differentials were tested, complete separation of the strains was possible using only
those five of Table 2.
(111) Studzes on the Australian counterparts. As indicated above large isolated pustules were
found in 1969 at Clinton, S.A. Laboratory tests showed this culture 69822 to be of
strain 326-1,2,3,5,6. As reported by Luig (1977) this combination 1(S76), 2(S711),
3(S79b), 5(S717), 6(S78) of virulences had not been previously reported in Australia.
Moreover and very spectacularly, this strain was avirulent on plants with S7r7b a host-
parasite interaction that had not been seen in Australia for about 30 years. The strain
was Clearly new to the population.
TABLE 1
Strains of P. graminis tritici zsolated from Angola and Rhodesia
Culture Number Location Australian Designation
Angola 1 Nova Lisboa 194 — 1,2,3,5,6,7
ee 2 ny 194 — 1,2,3,5,6
oe 3 ae 343 — 1,2,3,5,6
ie 4 me 222 — 1,2,3,5,6,11
re, 5 gi 194 — 1,2,3,5,6,*
2 6 Caconda 222 — 1,2,3,5,6
Rhodesia 1 Salisbury 34 — 1,2,3,5,6
4 2 He 222 — 1,2,3,5,6,11**
* Acme mesothetic
** As Angola 4
PRoc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
I.A. WATSON ANDC.N.A.DESOUSA 315
TABLE 2
Seedling infection types on key differentials selected from the standard and local sets when inoculated with seven strains of
P. graminis tritici from Angola and Rhodesia
Differential Rust Cultures from
Standard
Marquis S77b
Reliance S75
Acme S79g
Norka $715
Barleta
Benvenuto $7BB
mm MPP
Shortly after the collection at Clinton was made culture 691042 was found about
1,000 km east at Tichborne, N.S.W. It was classified as 194-1,2,3,5,6. Hence within a
short time two rusts vastly different from any previously seen in this country were
recovered. They differed only in the infection types found on plants with S77b. Luig
however, in unpublished work, has shown that although avirulence on plants with S715
had been found before in Australia these two strains produced a distinctly lower in-
fection type on plants with S715 than all other Australian strains with P15.
At the time of commencement of these studies in 1972 we had access to both these
strains, but on comparing the African rusts it was found that only one, Angola 2
(194-1,2,3,5,6), had an Australian counterpart even though six of them had the
numbers 1,2,3,5,6. Both 326 and 194 are avirulent on plants with $75 and a careful
search was made for variants of them with virulence. Strain 326-1,2,3,5,6 was soon
predominant in southern Australia and since Summit (575) was widely grown in
Victoria the appropriate mutation could be anticipated. This did not appear, however,
until the rust epidemic of 1973 when at Kerang, Victoria, one of us (I.A.W.) collected
rust (culture 73879) on a roadside plant of Hordeum leporinum Link. It was identified as
343-1,2,3,5,6 and represented the second counterpart of the African rusts, in this case
Angola 3. The third counterpart was of Angola 6 (222-1,2,3,5,6) and this was collected
at Willowie, S. Australia in 1974 (culture 74220). Luig and Watson later identified a
fourth counterpart (culture 74654) 194-1,2,3,5,6,7 (Angola 1) at Yacup in W.
Australia in 1974-75 and it was used in some of the tests.
(iv) Comparison of the Pathogenicity of the Angola strains and their Australian counterparts.
(a) On the standard and local differentials. When the components of the African
collections had been purified and when from the survey of Luig and Watson it was
found that in the Australian wheatfields counterparts of at least three of them were
present, detailed comparisons were commenced. The Angola numbers were
2(194-1,2,3,5,6), 3(343-1,2,3,5,6) and 6(222-1,2,3,5,6). The corresponding three
Australian cultures were 691042, 73879 and 74220.
The six cultures were used in pairs to inoculate seedlings of the standard and
supplemental differentials. The infection types of the Angola cultures are presented in
Table 3 and were identical with those of their respective counterparts. From these
readings it was apparent that the only genes important in the differentiation of the six
cultures were S75 and Sr7b. We concluded that this set of genetic material could not be
used to distinguish between the members of each pair of strains.
(b) On entries in the International Virulence Gene Survey. At the time of analysing
the African collections one of us (I.A.W.) was involved in testing the pathogenicity of
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
316
LONG DISTANCE TRANSPORT OF SPORES
TABLE 3
Seedling infection types at 17-22°C of 11 standard differentials and 11 local supplementals when tested with the three strains
of P. graminis tritici from Angola
Standard
Differential
Little Club
Marquis S77b
Reliance S75
Kota
Arnautka
Mindum
Spelmar
Kubanka
Acme S79g
Einkorn
Vernal
Australian Angola Culture and Strain
Supplemental
Angola Culture and Strain
2 3 6 2 3 6
194-1,2, 343-1,2, 222-1,2, 194-1,2, 343-1,2, 222-1,2,
3,5,6 3,5,6 3,5,6 3,5,6 3,5,6 3,5,6
4 4 4 McMurachy $76 4 4 4
4 2 4 Yalta S711 4 4 4
0 4 4 W2402 S77b Sr9b 4 2 4
4 4 4 C.1. 12632 SrTt, sil = ole ile
4 4 4 Renown Sr7b Sr17 4 2 4
4 4 4 Mentana S78 4 4 4
4 4 4 Norka S715 xX x x
oD = oD = oD) = Festiquay 5730 aD, a) a7)
i ;2= ;2= Agropyron
intermedium : : :
;1- ;1- ;1- Entrelargo 2+ 2+ 2+
de Montijo
;1- ;1- ;1- Barleta
Benvenuto S7BB x x xX
several Australian strains on seedlings of a collection of 117 wheats. These had been
selected as part of a survey to catalogue on a world basis the geographic location of
virulence genes and combination of them. The group had been divided into six sections
according to the alleles they carried.
Group 1
Group 2
Group 3
Group 4
Group 5
Group 6
Alleles with world wide usefulness as differentials. S7 genes such as
5,6,8,9b,11 and Tt, were included.
Alleles useful as differentials only in certain areas of the world.
S7 genes 7a,/b,9a,10,13,14,15,17,23 and several others as yet uncatalogued
were included here.
Combinations of known alleles.
These had two or more Sr genes per line.
Combinations of unknown alleles which were found to react differentially
from one region of the world to another. They were complex genotypes and
included such cultivars as Thatcher, Chris, Selkirk, Timgalen and Gamut.
Lines of unknown genetic constitution which were highly resistant to all
strains in many locations. The group included lines of Triticum turgidum L.
as well as many lines of T. aestivum L. from the International Spring Wheat
Rust Nursery.
Lines commonly used as susceptible parents in crosses and comprised Line
E W3498, W 2691 a Little Club derivative, Marquis and Chinese Spring.
All 117 lines of this collection had been tested with the available Australian strains
that closely resembled those from Africa. As well they were tested with Angolan rusts
numbers 1,3,4, and 6 and Rhodesia number 1. From the tests it was confirmed that the
only genes effective in differentiation were S75, S77b, S715 and S7BB. Angola 2 was not
compared
with culture 691042 in these tests but again no separation between members
of the pairs was possible despite the extent of the genetic variability in the host
material.
(v) Comparisons of other Characters in the African and Australian Rusts.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
I. A. WATSON ANDC.N.A.DESOUSA 317
(a) Shape and Size of Uredospores. The length of the uredospores of Angola 3,4 and 6
were compared with the Australian cultures 73879 and 74220. On the basis of measure-
ments on 950 spores of each there were no significant differences between Angola
3(33.48 + 1.75) and 73879 (34.56 + 2.18) and between Angola 6(34.32 + 2.99) and
74220 (34.08 + 2.91) at the P = 0.01 level although there was a marginal difference
between the first two at P = 0.05. Angola 4(30.08+ 2.06) was distinctly rounder than
any other culture but it had no Australian counterpart. In width of spore all cultures
examined fell within the range 17.644 1.84 — 18.684 1.25p.
(b) Uredospore Colour. There was no difference between Angola 2,3 and 6 and their
Australian counterparts. Angola 4 and Rhodesia 2 with the same pathogenicity both
differed from the other African rusts in having yellowish rather than brown
uredospores.
(c) Growth in Axenic Culture. Uredospores of Angola 3 and 6 along with their
Australian counterparts were sprayed separately onto Williams’s (1971) agar medium
in Petri dishes. The material was observed under the microscope 24 hours later and the
structures showing were identical in all four cultures. This comparison as with those for
pathogenicity, spore morphology and colour was of no value in demonstrating dif-
ferences between corresponding rusts from the two continents.
DISCUSSION AND CONCLUSIONS
(1) Pathogenicity and other Characters of the Cultures. It is apparent from the results obtained
that there is considerable diversity among the strains of P. gramuinis tritici in that part of
the African continent represented by Angola and Zimbabwe. It was not the purpose of
the study to examine in detail the extent of this variability but to compare African rusts
with those from Australia where there were known counterparts.
Four Angola cultures 1(194-1,2,3,5,6,7), 2(194-1,2,3,5,6), 3(343-1,2,3,5,6) and
6(222-1,2,3,5,6) each had counterparts in Australia which were respectively cultures
74654, 691042, 73879 and 74220. When the overseas cultures were examined in paired
comparisons with the corresponding local strains no differences in the members of each
pair could be detected, even though the host material was very diverse. The
uredospores were of the same size within reasonable limits of variation and although
one Angola culture was clearly different from the others in spore shape, no counterpart
for it was found in Australia. Growth in axenic culture which is sometimes useful in
distinguishing between strains revealed no differences between local and introduced
material. Although four counterparts were examined we are postulating that strains
326-1,2,3,5,6 and 194-1,2,3,5,6 arrived initially and the others 343-1,2,3,5,6,
194-1,2,3,5,6,7 and 222-1 ,2,3,5,6 evolved from them here by mutation.
(ii) Significance of certain genes for Avirulence in the Australian environment. Luig has already
pointed out (1977) that when found late in 1969 the two cultures 69822 and 691042
carried the formula 1,2,3,5,6 from their infection on the local supplementals. This was
a new combination for Australia. Virulence on plants with S76 (1) gave the first im-
portant hint in the field of this new material. In the previous four seasons 1965, 1966,
1967 and 1968 only seven isolates with this ability had been found east of the Nullarbor
Plain among 1684 examined during the period (Luig and Watson, 1970). In addition
both these new strains had another rare character, virulence on plants with 578. In the
five years prior to 1969 only 12 isolates from a total of 2603 from throughout Australia
were virulent on plants with this gene and not one of these could attack the Sr6 S78
combination. Thus to find these two cultures each with such a rare genotype £6 8 was
unexpected enough, but all the more so when one of them in addition had avirulence
on plants with S77b.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
318 LONG DISTANCE TRANSPORT OF SPORES
It has previously been pointed out that during the 15 years prior to 1970 new
patterns of pathogenicity arose in the Australian rust population due to the progressive
introgression of new alleles into it (Watson and Luig, 1966). About 1950 the fungal
alleles p5, £8, p15 and pBB predominated locally in P. graminis tritict. With the arrival of
21-0 all this changed since this strain brought with it P5, P8, P15 and PBB. It is clear
from our studies that of the eight alleles at these four loci, seven of them were found in
the African collections. P8 was missing. This all points to Africa also as the source of
standard race 21 in the 1948-54 period and we strongly suggest this as a real
possibility.
(iii) The origin of strain 343-1, 2,3,5,6. It may be argued that our failure to recover this
strain from the African material suggests some alternative origin for it. We pointed out
earlier however that only three collections were available and these may not have been
adequate to sample the area. If spores were windblown from Africa it would be ex-
pected that they would be of the most common strains. We know from Dr Santiago that
components of race 194 had been present in Central Africa and in 1970 Fonseca (1974)
found races 194 and 222 in Mozambique. To the best of our knowledge avirulence on
plants with S77b had not been identified in the area. However the ease with which its
presence was established in our collections demonstrates that it was part of the
population at the time of these introductions. During the interval between the
movement to Australia (1968-69) and the isolation in our tests (1972-73) the
population may have shifted from 326-1,2,3,5,6 to 343-1,2,3,5,6 by simple mutation
at the P5 locus. The demonstration of contrasting pathogenicity on plants with S77b
requires good control over environment especially temperature and without it the four
rusts of this family can easily be misclassified as shown in Fig. 1.
P5 to ps
320 ee
P7b |to p7b P7b to| p7b
P5 tops
194 S$ 222
Fig. 1. Changes in Standard race number with single gene mutations at the locus indicated.
We believe that 326-1,2,3,5,6 entered Australia about 1968 multiplied on
susceptible cultivates in southern states especially on Halberd (S76), mutated to
virulence at the P5 locus and under selection pressure resulting from the cultivation of
wheats such as Summit (S75) those mutants became apparent as 343-1,2,3,5,6 (73879).
(iv) Other similarities between African and Australian rusts. All the evidence we have
presented and the evidence of Luig and Watson (1970) and Luig (unpublished) show a
strong resemblance between the rust cultures we have compared. Unless gene flow has
been involved, the similarity is much greater than would be expected in cultures
originating in countries separated by the Indian Ocean. While this conclusion has been
reached mainly from comparisons of pathogenicity, Burdon e¢ al. (1982) have reached
the same conclusion using isozymes as genetic markers. This new approach can also be
used to trace evolution in rust populations. While these workers did not use material
identical with ours their results completely confirm the close similarity between the
strains of the two continents. No isozyme differences were found between the isolates of
standard races 21, 194, 222, 326 and 343. Moreover two isolates from Angola viz 2 and
PROC. LINN. Soc. N.S.W. 106 (4), (1982) 1983
I. A. WATSON ANDC.N.A.DESOUSA 319
Fig. 2. Initial trajectories of 3 identical constant level balloons released almost simultaneously from Men-
doza, Argentina on 20.9.71 (Partially redrawn from Morel and Bandeen, 1973).
C = Clinton, S. Aust., T = Tichborne, N.S.W., N = Nova Lisboa, Angola, P = Pretoria, S. Africa, R =
Rio de Janeiro, Brazil, B.A. = Buenos Aires, M = Mendoza, Argentina, L = Lima, Peru, SPR = South
Polar Region. Stippling indicates the probable source of spores.
4 were found to have the same phenotype as the following Australian rusts:
194-1,2,3,5,6 (S837 from Queensland), 326-1,2,3,5,6 (69822) and 343-1,2,3,5,6
(S1205 from Queensland). These workers have also proposed an African origin for
strain 21-0 and this supports the conclusion we have also reached.
(v) Weather conditions making spore transport possible. The shortest distance between the
wheat fields of Africa and Australia is approximately 12,000 km. To postulate natural
movement of viable spores across this distance of water and the establishment of
colonies from them in Australia we must assume ‘a delicate synchrony of biological and
physical processes’ (Hirst and Hurst, 1967). It has been demonstrated that the rust
strains from the two areas are indistinguishable, and it is known that physical processes
are available to elevate spores of them trom the crop to considerable heights. According
to Ingold (1978) rust uredospores are set free around midday, and under relatively dry
conditions vigorous thermals could carry the spores high into the air.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
320 LONG DISTANCE TRANSPORT OF SPORES
Schematic air paths shown by Ludlam (1967) suggest how they may be carried by
rain clouds into the high troposphere, and possibly to the jet stream.
Once into the atmosphere there are a number of meteorological systems which
could transport these spores to Australia. The first would be the anticyclones moving at
a height of up to 3 km and covering the distance in about six days. While there will be a
tendency for the spores to fall or to be washed out, circulation will have the effect of
lengthening the period when they are airborne. The fall speeds of the spores would be
of the order of 1 cms"! so that having reached an initial height of 5 km over the ocean,
drift could carry them easterly for another 10,000 km. If they are in winds of 10 ms"!
and falling at 1 cm s! they would cross the Indian Ocean at the rate of about 1000 km
each day (Pedgley, 1980).
As indicated above, some spores may reach the jet stream at a height of about
7-12 km and travel at much greater speed covering the distance in about two days. It
has become apparent from the work of Lalley et al. (1966) and of Morel and Bandeen
(1973) that jet streams in the southern hemisphere are much more continuous and
rapid than originally estimated. Monitored balloons released in New Zealand and
Argentina completed the global circuit in 10-12 days. Some of these trajectories are
partially redrawn in Fig. 2, and show that the balloons may pass over the critical areas
of both Africa and Australia. Presumably if spores reached the jet streams they could
do the same, and eventually subside following air current originating in local thun-
derstorms. Once at ground level the spores would need to reach a congenial host on
which to colonize. Apparently in 1968-69 the favourable climatic conditions which
enabled Australia to produce a record wheat crop were also those allowing migrant rust
spores to become established.
Intercontinental travel for spores of P. graminis tritict is hazardous and of rare
occurrence. It is for this reason that we believe Australia has been involved in this
migration only three times in the last 60 years.
ACKNOWLEDGEMENTS
The authors would like to thank Mr J. Colquhoun, Senior Research
Meteorologist of the Department of Science, Sydney, and Dr I. Watson of the School
of Earth Sciences at Macquarie University for help in interpreting the weather data.
The work is based on a thesis submitted to the University of Sydney by the junior
author for the degree of M.Agr. while holding a fellowship sponsored by the Food and
Agriculture Organisation. We are grateful to Mr L. N. Balaam, Dean of the Faculty of
Agriculture and Mrs Helen Keys for the typing of the manuscript.
References
BOWDEN, J., GREGORY, P. H., and JOHNSON, C. G., 1971. — Possible wind transport of coffee rust across
the Atlantic Ocean. Nature 229: 500-501.
BURDON, J. J., MARSHALL, D. R., Luic, N. H., and Gow, D. J. S., 1982. — Isozyme studies on the origin
and evolution of Puccinza graminis f. sp. tritici in Australia. Aust. J. Biol. Sct. 35: 231-238.
FONSECA, N., 1974. — Racas fisiologicas de Puccinia graminis f. sp. tritici (Eriks. and E. Henn.) em
Mocambique 11. — Resultados da prospecca feita em 1971. Inst. de Investigacao Agronomica de
Mocambique. Informacao Tecnica No. 77.
Hocec, W. H., Hounam, C. E., MALLIK, A. K., and ZADOKS, J. C., 1969. — Meteorological factors af-
fecting the epidemiology of wheat rusts. World Meteorologic Organisation Tech. Note No. 99.
Hirst, J. M., and Hurst, G. W., 1967. — Long-distance spore transport. In GREGORY, P. H., and
MONTEITH, J. L., (Eds.), Azrborne Microbes. Cambridge: Cambridge Univ. Press.
INGOLD, C. T., 1978. — Dispersal of micro-organisms. In SCOTT, P. R., and BAINBRIDGE, A., (Eds), Plant
disease Epidemiology. Oxford: Blackwell Scientific Publications.
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LALLEY, V. E., LICHFIELD, E. W., and SOLOT, S. B., 1966. — Global Horizontal Sounding Technique.
World Meteorologic Organisation Bull. 15: 124-128.
LupDLaM, F. H., 1967. — The circulation of air, water and particles in the troposphere. In GREGORY, P. H.,
and MONTEITH, J. L., (Eds), Azrborne microbes. Cambridge: Cambridge Univ. Press.
Luic, N. H., 1977. — The establishment and success of exotic strains of Puccinia graminis tritici in Australia.
Proc. Ecol. Soc. Aust. 10: 89-96.
, and WATSON, I. A., 1970. — The effect of complex genetic resistance in wheat on the variability of
Puccinia graminis f. sp. tritici. Proc. Linn. Soc. N.S.W. 95: 22-45.
, and , 1977. — The role of barley, rye and grasses in the 1973-74 wheat stem rust epiphytotic in
southern and eastern Australia. Proc. Linn. Soc. N.S. W., 101: 65-76.
MoREL, P., and BANDEEN, W., 1973. — The Eole experiment. Early results and current objectives. Bull.
Amer. Meteorolog. Soc. 54: 298-306.
PEDGLEY, D. E., 1980. — Weather and airborne organisms. World Meteorologic Organisation Tech. Note No.
173.
STAKMAN, E. C., STEWART, D. M., and LOEGERING, W. Q., 1962. — Identification of physiologic races of
Puccinia graminis var. tritict. Agr. Res. Service E-617 (Rev. ). 53 p.
WaTSON, I. A., 1977. — The national wheat rust control programme in Australia. Vestnik sel skochozja —‘
jstvennot Nauki No. 6, 123-133. (Russian).
——, 1981. — Wheat and its rust parasites in Australia. In EvANS, L. ‘T., and PEACOCK, W. J., (Eds),
Wheat science — Today and tomorrow. Cambridge: Cambridge Univ. Press.
, and Luic, N. H., 1966. — S715 — a new gene for use in the classification of Puccinia graminis var.
tritict. Euphytica 15: 239-250.
WILLIAMS, P. G., 1971. — A new perspective of the axenic culture of Puccinia graminis f. sp. tritict from
uredospores. Phytopathology 61: 994-1002.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
A Study of the Pollination of Alocasia macrorrhiza
(L.) G. Don (Araceae) in southeast Queensland
DOROTHY E. SHAW and B. K. CANTRELL
SHAW, D. E., AND CANTRELL, B. K. A study of the pollination of Alocasia macrorrhiza
(Araceae) in southeast Queensland. Proc. Linn. Soc. N.S.W. 106 (4), (1982) 1983:
323-335.
Flowering of Alocasia macrorrhiza (L.) G. Don in southeast Queensland was in-
vestigated and appears to be protogynous. Insects in the open spathal chambers, on
the spadix after closure of the constricting area, and in the sealed chamber of the
nearly mature infructescences were recorded to determine possible pollinators. The
number of seed set in an experiment involving various combinations of emasculation
and bagging of 143 inflorescences including controls appear to indicate cross
pollination by insects.
Dorothy E. Shaw, Plant Pathology, and B. K. Cantrell, Entomology, Department of Primary
Industries, Mewers Road, Indooroopilly, Australia 4068; manuscript received 26 June 1982,
accepted for publication 20 October 1982.
INTRODUCTION
Alocasia macrorrhiza (LL.) G. Don (Araceae) occurs in wet tropical Asia, eastern
Australia and the Pacific Islands; to some of the latter at least it was probably in-
troduced, as well as to other parts of the wet tropics and subtropics. In Australia it
occurs in the high rainfall coastal areas and adjacent plateau regions in Queensland
and New South Wales, from about 12° to 34°S latitude.
Hamilton (1898) reported the beetle, Brachypeplus murray: Macleay (Coleoptera:
Nitidulidae) as the pollinator of A. macrorrhiza, and that a species of Agaristidae
(Lepidoptera) also aids in fertilization. McAlpine (1978) described Neurochaeta inversa
(Diptera: Neurochaetidae) (whose life cycle is intimately connected with the flowers of
this plant growing in its natural habitat in eastern Australia) which he thought may act
as the pollinator. He observed, however, that as many insects visit the flowers, it is
probably not the sole agent.
Shaw et al. (1982) reported that A. macrorrhiza still set seed in gardens where no N.
inversa were recorded, and in remnant natural stands now surrounded by mainly non-
forested areas, where occurrence of N. inversa was sparse. It was also shown that seed
set in plants from rainforest areas was not related to the numbers of N. inversa recorded
in chambers of developing and matured infructescences. They concluded, therefore,
that pollination is probably independent of ovipositing females of N. znversa.
The present study records the stages of flowering and the insects associated with
the inflorescences or flowering heads (hereafter called ‘heads’) and in the nearly mature
infructescences. Also reported are the results of an experiment involving emasculation
and bagging of heads before anthesis, with controls, to provide information on the
possibilities of selfing or crossing, and of wind or insect pollination, as measured by
seed set.
SITES AND METHODS
Sites
A. macrorrhiza has been under study since 1980, but the work reported below was
mainly carried out in the flowering period October 1981 to March 1982 in southeast
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
324 POLLINATION OF ALOCASIA MACRORRHIZA
Queensland, at sites in rainforest areas along the coastal range west and southwest of
Brisbane, and in simulated rainforest areas (gardens) in Brisbane suburbs. Details of
the sites are as follows:
Rainforest: Three sites in small private holdings adjacent to large or relatively large
reserved areas of rainforest at Mt Nebo, Mt Glorious and Mt Tamborine, at about
520, 650 and 500 m above sea level (a.s.1.) respectively; plants at the first and third
sites were growing in and at the edge of streams; at each site plants occurred in scat-
tered clumps.
Garden: Three sites in the Mt Coot-tha Botanic Gardens, developed as simulated
rainforest in natural sclerophyll forest during the last four years, and one site at the
University of Queensland, St Lucia, surrounded by parklands; all sites about 50 m
a.s.l.; plants at one site at Mt Coot-tha and most plants at St Lucia, at the edge of
streams; at each site plants occurred in scattered clumps.
Methods
At each site inflorescences were examined throughout the entire flowering period
and observations made on the various stages of flowering, both in the field and on
excised heads removed to the laboratory. In some in situ (non-treatment) and other
excised heads, small hinged ‘windows’ about 1 cm square were cut into the wall of the
spathal chamber for stigma viewing, the flap being resealed with adhesive tape.
All sites were visited as frequently as possible after the treatments below were
applied. Any rotting heads were harvested when the rotting became evident, but heads
maturing were allowed to remain in situ until near maturity but before actual splitting
of the wall of the spathal chamber to ensure no seeds were lost. Each head was
examined at harvest, and if any fertilization had occured, the numbers of unfertilized
ovaries, fertilized ovaries (berries) and fertilized ovules (seed) were determined.
a. Treatments
Emasculation and bagging were carried out only on ‘buds’ when each spadix was
still enclosed by the furled spathe, prior to anthesis. The treatments were as follows:
1. Emasculated, mesh-bagged. This involved the excision of the distal portion of the
spathe (consisting of the furled spathal limb, the sterile terminal appendage, the
portion bearing the staminate flowers and the upper portion of the median sterile
florets) at the constricted zone. The excised part of the spadix was removed from the
excised spathal limb and checked to ensure that all the staminate flowers had been
removed. The portion of the spadix remaining on the plant (consisting of the lower
part of the median sterile florets and the pistillate flowers) was enclosed by a nylon
mesh bag fine enough to keep out insects larger than thrips — no thrips, however,
although they could physically enter, were found inside the bags.
Emasculated, unbagged. As above, but unbagged.
3. Non-emasculated, whole head paper-bagged. A white paper bag normally used in maize
pollination studies was employed.
4. Non-emasculated, whole head plastic-bagged. A commercially available clear plastic bag
was found to be suitable.
5. Non-emasculated, whole head mesh-bagged. Similar bag as used in Treatment 1. In some
cases the bags were distended by the insertion of a framework composed of two
metal rings maintained in position on a rigid metal upright attached to the
peduncle.
6. Non-emasculated, unbagged, labelled. Heads untouched, labelled as controls while still
in bud.
During each visit to the field, at least one set of the six above treatments were
prepared, heads being labelled individually. Occasionally additional heads of some of
NO
Proc. LINN. SOc. N.S.W. 106 (4), (1982) 1983
DOROTHY E. SHAW ANDB. K. CANTRELL 325
the treatments were included if extra heads were available. All the paper, plastic and
mesh bags were tied at the base with nylon net tape.
7. Non-emasculated, unbagged, unlabelled. Early maturing heads at each site, previously
unused in our experiments, were also harvested as further controls for comparison.
b. Insects
Records were made of all insects observed within the open spathal chamber when
the pistillate flowers were exposed and those later found around the staminate flowers
and on the accumulated pollen of the spathal ‘catchment area’ above the constricted
portion at the base of the spathal limb. An aspirator was used to collect representative
samples of all species of insects encountered (from non-treatment heads). Any insects
in the sealed spathal chambers of treatment heads at harvest, whether immature or
adult, were also recorded and retained.
RESULTS
Flowering
After the green spathal limb unfurls, the constricting portion at the base of the
limb gapes widely, with a gap of 3-4 mm around the spadix, so that the pistillate
flowers can be partially viewed from above. At this stage the subsessile stigmas appear
dull, white and rough, and the 2-3-(4) short lobes are seen microscopically to consist of
a cushion of short hyaline ‘hairs’ ca 138 to 227 um long and ca 13 to 23 wm wide (mean
157.1 by 16.2 um), to which the sticky pollen grains adhere. The chamber is open for
about one day in the field, after which the constricting portion of the spathe closes
tightly around the spadix in the region of the median sterile florets, sealing off the
chamber. Pollen is then extruded in white cirri from the synandria of the staminate
flowers for about two days, and fallen pollen accumulates in the pollen catchment area
formed at the base of the spathal limb abave the constriction. At this stage, stigmas in
the enclosed chamber are glistening, white and glutinous. (The only exceptions noted
to the above were a few heads from 650 m a.s.1., excised at the beginning of anthesis,
and then held in the laboratory at about 50 m a.s.1. Some pollen in each head began to
extrude, mainly from the lower staminate flowers, before the final closure of the
constricting portion. This may have been because the heads had not been visited by
insects, and the ‘unfertilized’ stigmas may have been receptive longer, perhaps
delaying the closure. Also, as Stanley and Linskens (1974) pointed out, altitude and
temperature affect pollen dehiscence, so that the warmer temperature in the laboratory
at the lower altitude may have caused extrusion of the pollen earlier than normal. This,
however, was an unnatural situation.)
A fragrance, present at the beginning of anthesis, was still present after pollen
extrusion, though less pronounced.
Details of pollen studies will be published separately, but are summarized here as
follows: When extruded in the cirri, the pollen is sticky and will adhere to a scalpel
blade, but the stickiness decreases with age, so that accumulated pollen, although still
mainly in clumps, could be launched into the air by deliberately jarring the head. In
the cirri, the pollen appears white, but when shaken out of the head is deep ivory,
about plate 4A2 in the Methuen Handbook of Colour (Kornerup and Wanscher,
1961). The grains are microscopically colourless, spherical, 32.3-47.54m long by
30.3-47.5 um wide (mean 41.2 x 39.3 mm) excluding ornamentation of surface
spinules ca 2 um long. Twenty-two samples of pollen shaken from 10 heads, examined
microscopically, revealed no pollen other than that of A. macrorrhiza. In germination
tests on modified Brewbaker and Kwack (1973) medium, pollen shaken from heads in
the laboratory at the beginning of each day was mainly viable during the first and
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
326 POLLINATION OF ALOCASIA MACRORRHIZA
TABLE 1
Presence ( +) or absence (—) of insects recorded in the open chambers (O) and around the spadix after sealing of the chambers
(AS) of non-treatment heads
HABITAT, Insects
locality, N. inversa T. Brachypeplus Staphyl- Taento- Other*
site carbonaria sp. inidae thrips
O AS O AS O AS O AS O AS O AS
RAINFOREST
Mt Nebo + + + + + + + + Sto 5
Mt Glorious + + + + + + + + - + 27
Mt Tamborine + + - - + + + + - + BaOaae)
GARDEN
Mt Coot-tha
Site 1 +# - + + + + + + ey 6
Site 2 - = ft + fs fi + + See 1
Site 3 +# - + + + + + + - + 8
St Lucia - - - = 4 f + ns SE
% Other = 1 A. mellifera #
2 Pteromalidae #
3 Curculionidae (5 specimens: 4 open chamber; 1 around spadix)
4 P. fimetarius #
5 Cyphon sp. (2 specimens)
6 T. ? minor #
7 Hippelates sp. (2 specimens)
8 Dolichopodidae #
# only single individuals
second days of production, with much lower and slower germination on the third day
and only rarely on the fourth day. Some sterile (non-staining) inviable pollen was
present in samples from all sites.
Insects
1. Within the open chamber
Four species of insects were commonly encountered within the open chamber
(Table 1). These were N. inversa, Trigona carbonara Smith (Hymenoptera: Apidae),
Brachypeplus sp. (Coleoptera: Nitidulidae) and an unidentified alaocharine staphylinid
(Coleoptera: Staphylinidae). The two beetles were present at all localities while N.
inversa was limited to rainforest localities except for two specimens (only one captured)
at Mt Coot-tha. Although common at most localities, T. carbonaria was not recorded
during any visit to Mt Tamborine or St Lucia.
Several other insects were occasionally recorded within open spathal chambers,
but usually only as single individuals. These were Taenzothrips (Isochaetothrips) sp.
(Thysanoptera: Thripidae), Cyphon sp. (Coleoptera: Scirtidae), Triphyllus ?minor Lea
(Coleoptera: Mycetophagidae), an unidentified derelomine weevil (Coleoptera:
Curculionidae), Hippelates sp. (Diptera: Chloropidae), an unidentified dolichopodid
(Diptera: Dolichopodidae) and an _ unidentified pteromalid (Hymenoptera:
Pteromalidae) (Table 1).
2. On the spadix after closure of the constricting portion
Four species of insects were commonly encountered on the spadix (or in the ac-
cumulated pollen) after closure of the spathal constriction. These were 7. carbonara,
Brachypeplus sp., the staphylinid and Taeniothrips (Isochaetothrips) sp. All were present at
each locality except for 7. carbonara which was absent as noted previously (Table 1).
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
DOROTHY E. SHAW ANDB. K. CANTRELL 327
TABLE 2
Insects other than Neurochaeta inversa recorded in sealed chambers of matured heads *
Insects”
HABITAT, Head Brachypeplus Chloro- Diptera Fermi- Collem-
locality designation sp. pidae Muscidae unidentified cidae bola
No. No. No. No. No. No.
RAINFOREST
Mt Nebo 7A.4 2L
2P
Mt Glorious 6A.1 1 L#
X.1 iL
X.2 1L
Mt Tamborine 1A.1 1L
1A.4 2L
9A.1 1L
9A.2 1L
X.1 1A
X.2 6P WIL
X.4 1A 1P DI
2L
GARDEN
Mt Coot-tha
Site 2 1A.1 iL
Chambers in 131 observed heads (including unlabelled controls) were without insects, other than
presence of NV. inversa in some.
Life forms: L = larva; P = pupaor puparium; A = adult.
# Larvae in one of two damaged ovaries.
= Dead.
TABLE 3
Numbers and fate of heads in seven treatments
Heads in the following classes
HABITAT, Emasculated Non-emasculated
locality, =
site Mesh- Unbagged Paper- Plastic- Mesh- Unbagged, Unbagged,
bagged bagged bagged bagged labelled unlabelled
M R M R M R M R ME TR EIR M R
No. No. No. No. No. No. No. No. No. No. No. No. No. No.
RAINFOREST
Mt Nebo 0 1 0 1 4 0 Se
Mt Glorious 0 5 1 2 0 2 0 3 0 7 2 3 3
Mt Tamborine 0 6 60 6 0 2 4 0 5 1 i}
GARDEN
Mt Coot-tha
Site 1 0 3. (0 3 0 2 3 1 2 3 1 Dh ok
Site 2 0 3 2 1 0 32> 2 0 Sie 33 1 1 ex
Site 3 0 1 0 2 0 Se) 3 0 1 3 1 Sk
St Lucia 0 3 0 1 0 3 4 0 2px
Total OP A1s eS SS as QO ts OQ ag LU} 28} 7 Bil sx
M = Matured; all heads with seed (details Table 4)
R = Rotted; all heads without seed except four with few seeds (details Table 4)
x = Some rotted heads occurring naturally, but uncounted.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
328 POLLINATION OF ALOCASIA MACRORRHIZA
TABLE 4
Numbers of seed set in treated heads
Heads in the following classes
HABITAT, Emasculated Non-emasculated
locality, Ri Sane hy en ES Pe EE TL a Ee ee ee
site Mesh- Unbagged Paper- Plastic- Mesh- Unbagged, Unbagged,
bagged bagged bagged bagged labelled unlabelled
No. No. No. No. No. No. No.
RAINFOREST
Mt Nebo 6* 0 66 310
229 381
191
260
Mt Glorious 0 37 0 0 0 0 131
0 0 0 0 0 0 140
0 0 0 0 2 103
0 0 148
0 0 138
0
0
Mt Tamborine 0 0 0 De a 33 223
0 0 0 0 0 69 183
0 0 0 0 152 434
0 0 0 0 41 180
0 0 0 0 147
0 0 5
64
101
GARDEN
Mt Coot-tha
Site 1 0 0 0 0 13 174 205
0 0) 0 0 0 0 106
0 0 0 0 130
173
Site 2 0 173 0 0 0 223 137
0 12 0 0 0 0
0 0 0 0 6
44
Site 3 0 0 0 0 0 173 109
0 0 0 155 18
0 0 0 98
0 14
0
St Lucia 0 0 0 101 2
0 0 137 40
0 0 288
44
Total heads, No. 18 21 15 19 19 30 21
Total seeds, No. 0 228 0 2 18 2649 3117
Seed, Mean 0) 10.9 0 0.1 0.9 88.3 148.4
* Heads rotted, although a few seeds were set.
Proc. Linn. Soc. N.S.W. 106 (4), (1982) 1983
DOROTHY E. SHAW ANDB. K. CANTRELL 329
Occasional encounters were Apis mellifera L. (Hymenoptera: Apidae), N. inversa,
Phalacrus fimetarius (F'.) (Coleoptera: Phalacridae) and the unidentified weevil (Table 1).
3. Within the sealed spathal chamber near maturity
The main species found within the sealed spathal chambers near maturity was N.
inversa, as recorded in Table 5. It occurred, however, only at the rainforest sites, not in
the gardens. A few occasional insects were also recorded, as listed in Table 2.
Inflorescence treatments
In all, 139 heads were involved in Treatments 1 to 6. Of these, 10 were lost
through accident, stealth, etc., and seven heads in Treatment 3 were discarded because
of perforation of the paper bags due to heavy rainfall and possibly pressure from the
growing heads, giving an actual total of 122 heads. In addition, 21 heads in Treatment
7 were included in the results for comparison, making a total of 143 heads.
The results are reported in Tables 3 to 6. Table 3 shows the number of heads
maturing or rotting for each treatment. No difference in behaviour occurred between
any site, locality or habitat, nor was any difference noted between heads bagged with
paper bags compared with those with plastic bags. Some rotted heads occurred
naturally, as shown in the labelled and unlabelled controls. Only those heads matured
where seed had set except for four, which between them had only 12 berries with 15
seeds. Conversely, the only heads which rotted were those where no seed had set, apart
from the four exceptions mentioned previously.
The details of seed set are given in Table 4. The only heads which set seed were
those of the controls, except for four unbagged, emasculated heads (one of which
rotted) and one plastic-bagged and two mesh-bagged non-emasculated heads (two of
which rotted).
Table 5 shows the numbers of fertilized ovaries (berries) and fertilized ovules
(seed) for all matured heads and the numbers (or absence) of N. imversa (mainly
puparia) recorded in the sealed chambers of these heads. Seed set varied considerably
at most sites, ranging from 1.5% (St Lucia) to 99.3% (Mt Nebo). The numbers of N.
inversa per head ranged from three (Mt Tamborine) to 77 (Mt Glorious) with averages
of 24.2, 26.5 and 17.5 respectively for the three rainforest localities (Mt Nebo, Mt
Glorious and Mt Tamborine). There was no relation between numbers of N. znversa
and the numbers of seed set per head, locality or habitat. No N. inversa were recorded
in heads from the garden areas, but up to 88.3% of seed was still set.
DISCUSSION
Pollination ecology
In A. macrorrhiza the pistils (megasporophylls) and stamens (microsporophylls) are
present on different parts of the flowering head of the same plant. The exposure of the
stigmas by gaping of the constricting areas at the unfurling of the spathal limb,
presumably at the time of receptivity of the stigmatic surface, and the closing of the
constricting portion of the spathe (therefore isolating the pistillate flowers) before the
extrusion of the pollen, indicate protogyny. The precise period and peak of stigma
receptivity, measured by the number and maximum of seeds set, need to be deter-
mined by controlled crossing at different times during anthesis.
The possibility of pollination by wind (anemophily) as against insect pollination
(entomophily) is examined below.
The pollen when first extruded in cirri above the surface of the staminate flowers
(and in the best position for pick-up by wind currents) is at its stickiest and so less likely
to be wind dispersed. Also, the extended, upright and slightly incurved spathal limb
would act as a protector and deflector of wind from the rear of the head. After the
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
330 POLLINATION OF ALOCASIA MACRORRHIZA
TABLE 5
Numbers and percentages of fertilized ovaries (berries) and numbers of seed and N. inversa per head, locality and habitat for
all matured heads
HABITAT, Fertilized Seed per head N. inversa per head
locality, ovaries
head (berries)
designation per head Locality Habitat Locality Habitat
No. % No. Mean Mean No. Mean Mean
RAINFOREST 156.9 21.4
Mt Nebo 239.5 24.2
6A.1 55 49.1 66 21
6A.2 131 89.1 229 26
7A.1 124229 191 46
7A.2 107 76.4 260 11
X.1 150 995.5 310 16
X.2 14) 99.3) 381 25
Mt Glorious 116.2 26.5
6A.1 87 = 558.8 148 11
6A.2 88 59.9 138 8
X.1 128 975.7 131 22
X.2 140) 9772 140 11
X.3 134 61.9 103 77
8B 262A 37 30
Mt Tamborine 136.0 17.5
1A.1 29 22.3 33 3
1A.4 46 =28.9 69 29
9A.1 117. 92.1 152 7
9A.2 S22 3ED 41 18
X.1 148) 728) 223 12
X.2 117 = 46.3 183 19
X.3 210 =98.1 434 31
X.4 85 61.6 180 6
X.5 96 49.0 147 29
X.6 5 28) 5 9
X.7 47 = 44.8 64 20
X.8 Ouro 101 27
GARDEN 107.3 0
Mt Coot-tha
Site 1 133.5 0
1A 132 60.6 174 0
5A.1 80 63.5 130 0
5A.2 145 87.9 173 0
X.1 136 = 88.3 205 0
X.2 Uy 43.9) 106 0
1D 10 6.5 13 0
Site 2 99.2 0
1A 143 77.3 223 0
4A.1 6 5.3 6 0
4A.2 33 56.9 44 0
X.1 117 += 83.0 137 0
1B 155 74.9 173 0
4B 10 5.8 12 0
Site 3 94.5 0
1A.1 92220 173 0
1A.2 119 58.3 155 0
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
DOROTHY E. SHAW ANDB. K. CANTRELL 331
5A 10 7.6 14 0
ll 62 30.7 109 0
X.2 14 8.6 18 0
X.3 75 42.9 98 0
St Lucia 102.0 0
3A.1 7A 538.0 101 0
3A.2 IQ FO 137 0
3A.3 Nile 7: 288 0
3A.4 94 19.5 44 0
Yen 2 185 2 0
xe? Sela 40 0
TABLE 6
Theoretical expectancies of selfing vs crossing and wind vs insect pollination and the actual results
Heads in the following classes
Emasculated Non-emasculated
Mechanism Paper- and
Mesh- plastic- Mesh-
bagged Unbagged bagged bagged Unbagged
SELFED! no seed no seed seed seed seed
CROSSED
Wind
Pollen reaching chamber
of emasculated heads” seed seed no seed seed seed
Pollen not reaching chamber ?
of emasculated heads” no seed no seed no seed seed seed
Insects
Attractant still present
in emasculated heads* no seed seed no seed no seed seed
Attractant excised
in emasculated heads* no seed no seed no seed no seed seed
Tf insect crossed, attractant
excised, no. of heads with:
(Theoretical)
Seed 0 0 0 0 30
No seed 18 21 34 19 0
(Actual)
Seed 0 4 1 2 24
(2 seeds) (few seeds)
No seed 18 17 33 17 6
' Normally protogynous, but selfing is perhaps possible in absence of cross pollen or pollinator(s) if some
residual stigma receptivity is still present at time of pollen extrusion, unless self-incompatible.
2 Cross pollen would need to find the small entrance to the spathal chamber.
3 Attractant (possibly fragrance) probably situated in the appendage/staminate portion of the spadix
removed with emasculation.
pollen has fallen, when it is less sticky, the grains are fairly well protected from wind
pick-up, being within the overlap at the base of the spathal limb in front (in the pollen
catchment area) and the spadix itself at the back. However, neither occurrence (that is,
stickiness when extruded and protection within the base of the overlapping spathal
limb) need necessarily completely exclude the possibility of dispersal of some pollen
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
332 POLLINATION OF ALOCASIA MACRORRHIZA
grains by wind. This may occur, as previously reported, by deliberately jarring the
head. Such jarring could occur in nature by falling branches, accidental encounters by
birds or ground animals and possibly by high winds.
Faegri and Pijl (1979) have stated that an obvious condition for effective wind
pollination is the presence of wind, as found in open, sparse vegetation or in the top
layer of closed, multi-layered vegetation types, and that in dense forest vegetation wind
is so slight and infrequent that anemophily is contra-indicated. At all the sites except
Site 3 at Mt Coot-tha the plants under study were part of the understorey, and at most
sites were well protected by topography (being in or on the banks of streams) and by
lush gully vegetation.
Provided pollen grains are fortuitously distributed, the total output of pollen
produced, N, gives a number of ‘effective’ grains, i.e. grains carrying out pollination,
n, which is a function of the areas of stigmatic surfaces, a, and the total area of the
surroundings, A: n _ a (Faegri and Pijl, 1979). The production of inflorescences on
Nig A
any plant of A. macrorrhiza is sequential, so that when a head is in anthesis, the previous
head is already in ‘pod’ (with the spathal chamber sealed) and the subsequent head is
still in bud, with the spathe still furled. Therefore pollen, if wind-borne, would need to
encounter heads on other plants at the right stage of stigma receptivity. Such heads
could occur on plants in neighbouring clumps, though these are often scattered and
sparse. Wind pollination is considered much less precise than biotic pollination, and to
achieve its objective it presumes a very high incidence of pollen grains near their
source, the dilution factor going by a power of 3 of the distance (Faegri and Pil, 1979).
While considerable pollen of A. macrorrhiza is produced in any one head, the overall
quantity in any one area would not be great, and also it is either sticky when at source
or when older and less sticky is protected from easy wind pick-up in the pollen catch-
ment area, as discussed previously. Even if wind-borne pollen were to encounter a
recently unfurled head with receptive stigmas, the pollen has to reach those receptive
surfaces. The chance of wind-borne pollen falling under gravity directly into an open
chamber through the gaping constricting area would seem rather remote. Alter-
natively, wind-borne grains could strike the inner surface of the extended upright
spathal limb, if the limb happened to be at an appropriate angle to the line of flight, but
again the grains would need to fall directly, or down the side of the spathal limb, onto a
receptive surface. Even if a successful landing on a receptive stigma (pollination) was
achieved, successful fertilization (germination on the stigma and fertilization of the
ovules) may not necessarily follow, because the less sticky the pollen and therefore
more readily available for wind dispersal, the older and less viable it is, and therefore
less capable of achieving germination and fertilization.
Faegri and Pil (1979) pointed out that one of the factors of major importance in
anemophiles is the buoyancy of the pollen grains, which increases as the size of the
grain decreases, and considered that pollen of anemophiles belongs to the smaller size
classes with a ‘typical’ diameter of 20-30-(60) um, even if equally small or even smaller
pollen grains are found in many entomophiles. Contributing to the effectivity of pollen
transport in anemophiles and to the buoyancy of the pollen is the fact that grains do not
adhere to each other, but are smooth and dry and are spread separately or in very small
groups. Entomophilous pollen, on the other hand, is ornamented and sticky and the
grains stick together (Faegri and Pijl, 1979). Thus the size of A. macrorrhiza pollen
(mean 41.2 x 39.3 wm exclusive of ornamentation) measured in this study is in the
larger part of the range for an anemophile, and their ornamentation of spinules and
their stickiness suggest an entomophile.
In A. macrorrhiza therefore, the extrusion of sticky pollen in cirri, the continued
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
DOROTHY E. SHAW ANDB. K. CANTRELL 333
though decreased stickiness of the pollen with age, coupled with decrease in viability,
the occurrence of spinules on the pollen grains and the short stigmatic hairs all suggest
pollination by insects (entomophily) rather than wind pollination (anemophily). In
addition, the occurrence of the fragrance at anthesis may also indicate entomophily.
Insects
N. inversa was present in all open chambers (Table 1) examined from the rainforest
localities, usually moving on the walls of the chamber, seldom near the stigmas. Flies
were seldom encountered on the spadix, either before or after closure of the spathal
constriction, and were much more often seen running over the spathal limb. On three
occasions copulating pairs were noted, once within an open chamber and twice on the
inside face of the spathal limb.
Brachypeplus sp. and the aleocharine staphylinid were usually moving over the
ovaries below the stigmas in open chambers. No beetles, when examined
microscopically, had pollen adhering, but this could have been dislodged during
capture. T. carbonaria collected from the open chambers were from the stigmas, often
with heavy pollen loads in their corbiculae. Macroscopically the corbicular pellets
appeared homogeneous in colour and texture. Microscopical examination of samples
from six pellets chosen at random showed that they were entirely composed of A.
macrorrhiza pollen, except for four foreign grains from one sample. 7. carbonaria was
therefore mainly ‘A. macrorrhiza constant’ and must be considered to be a possible
pollinating agent since they were also often observed collecting pollen on heads with
sealed chambers.
Of the insects occasionally recorded within the open chambers (Table 1) the thrips
were present in large numbers on the spadix of such inflorescences at all localities and
the presence of some in the chamber would be expected. The records of the remaining
species cannot be easily explained but is probably due to chance occurrence. In any
case, their rarity discounts any possibility of their being effective pollinators.
Considering the insects on the exposed spadix after closure of the spathal con-
striction (Table 1), 7. carbonaria were observed visiting and collecting pollen at most
sites, and their absence at others is probably due to lack of colonies in the area.
Brachypeplus sp. and the staphylinid were always present, often immersed in the ac-
cumulated pollen of the spathal catchment area above the spathal constriction.
However, no beetles were ever observed arriving or leaving and experiments on their
vagility are necessary to determine their ability as pollinators. The small size of the
thrips makes them unlikely pollinators despite their common occurrence, and like the
beetles, their vagility is unknown.
Normally ubiquitous, the honeybee was only observed visiting on a single oc-
casion, despite their presence on other flowers at all localities. The only other insects
encountered apart from N. inversa (already mentioned) (Table 1) were the phalacrid
and the unidentified weevil, both single individuals.
The numbers of N. inversa recorded in the sealed chambers (Table 5) confirms the
report of McAlpine (1978) and Shaw et al. (1982) that they occurred only in rainforest.
(It should be noted, however, that two adult N. inversa were found in two open
chambers at two of the Mt Coot-tha sites (Table 1). This may indicate that the fly is
now finding its way to this area, which has only been established for four years.) The
few occasional insects recorded in the sealed chambers (Table 2) show a similar basic
composition, though less extensive, to those recorded by Shaw et al. (1982, table 5).
In his studies on the pollination of A. macrorrhiza in the Illawarra District near
Sydney, Hamilton (1898) recorded the beetle, Brachypeplus murray: in the open
chambers. He also made some observations not substantiated by us, viz., beetles
feeding on liquid exuded by the pistils and beetles, becoming confined in sealed spathal
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
334 POLLINATION OF ALOCASIA MACRORRHIZA
chambers, later burrowing their way out. We noted no such feeding habit, nor any
perforated chambers.
Hamilton (1898) also observed an unidentified species of agaristid (Lepidoptera:
Agaristidae) in open chambers and claimed that they aided in fertilization. Larvae of
the agaristid Crurta donovani (Boisduval) were noted at all rainforest localities during our
study, where they were often seen inside buds and open chambers feeding on stigmas
and ovaries. However, as a single head contains sufficient tissue to allow a larva to
reach maturity they are unlikely to be effective pollinators.
If indeed A. macrorrhiza is insect pollinated as suggested by this study, then
Brachypeplus sp. and the staphylinid would appear to be the most likely candidates since
they occur at all localities examined. While N. inversa and T. carbonaria might appear to
be more likely to be effective in this role, their absence from some localities where good
seed set occurred does not support this, or at least suggests that a number of pollinating
agents are involved, and that a resolution requires further investigations.
Emasculation/bagging experiment
The results of the emasculation/bagging experiment given in Table 3 show that all
heads rotted if no seed was set (except four with a few seeds only) and conversely, the
only heads which matured were those where seed had set (other than the four ex-
ceptions).
The results in Tables 3-6 show that, as seed occurred (other than in seven heads)
only in unbagged, non-emasculated heads (Treatments 6 and 7), A. macrorrhiza is not
apomictic, because if it were, seed could have occurred in all heads, unless
pseudogamic.
Mesh-bagging allowed seed set (18 seeds) in only two heads out of 36 (Table 4)
and therefore wind pollination is probably unlikely as a general occurrence. The 18
seeds (set in 12 berries) may have resulted from chance wind-borne or self pollen.
Emasculation resulted in seed set (228 seeds) in four out of 39 heads, compared
with 2669 seeds in 27 out of 83 non-emasculated heads including labelled controls
(Table 4). The four emasculated heads which had some seed set were unbagged and
could have been fertilized by wind- or insect-borne pollen. In the non-emasculated
heads, only two seeds were set in one out of 34 heads in any imperviously-bagged head
(paper or plastic) and only 18 seeds in two heads out of 19 in the mesh-bagged heads
(Table 4); the two seeds may have been the result of self pollination while the 18 seeds
may have resulted from self or chance wind pollination. Twenty-four of the unbagged,
non-emasculated (labelled) controls (Treatment 6) set 2649 seeds against six heads
which did not (Table 4); the non-seed set in these heads may have been due to lack of
pollen at the time of stigma receptivity and/or lack of pollinating agent(s).
The results shown in Table 5 confirm data previously reported by Shaw et al.
(1982) viz., that there is great variation in the numbers of seed set per head (in the
present study from 1.5% to 99.3% in the 48 heads, all controls except two); that there
is no relation between numbers of seed set and numbers of N. inversa recorded in the
sealed chambers (which it is presumed reflects the numbers of ovipositing females in
the open chambers) at the rainforest sites; that no N. znversa were recorded in the sealed
chambers at the garden sites but despite this seed up to 88.3% was still set in garden
site heads.
Table 6 shows the theoretically expected seed set figures deduced from the
treatments applied in our experiment. The actual results, also shown in Table 6, most
nearly approximate those of crossing with insect pollination with the attractant excised
in the emasculated heads. Small differences from the theoretical results occurred in the
emasculated unbagged heads, where four heads set 228 seeds (possibly insect or chance
wind-borne pollination); in the imperviously-bagged non-emasculated heads, where
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
DOROTHY E. SHAW ANDB. K. CANTRELL 335
two seed set in one head (probably self pollen); and in the mesh-bagged non-
emasculated heads, where 18 seeds set in two heads (possibly self or wind-borne
pollen), whereas no seed was expected in any of these classes. Also, all control (labelled)
heads were expected to set seed but six heads out of 24 did not, a situation also found in
the unlabelled controls.
The results of the experiment therefore appear to indicate cross pollination by
insects, with little attraction for the insects if the appendage/staminate portion of the
spadix is removed.
ACKNOWLEDGEMENTS
The owners of the sites sampled and other individuals who assisted in obtaining
the collections, including Mrs J. and Miss L. Cantrell, Mr A. Hiller, Mrs J. Hope, Ms
K. Howdesell, Mr and Mrs L. J. Manning, Mr A. L. Marstella, Mr and Mrs W.
Scattini and also Mr H. Caulfield (Mt Coot-tha Botanic Gardens) and Dr A. B. Cribb
and Mr A. R. Steginga (University of Queensland) are thanked for their co-operation
in this study.
We are also grateful to Mr T. A. Weir and Dr E. C. Zimmerman, both of the
Commonwealth Scientific and Industrial Research Organization, Canberra, and Dr I.
D. Galloway and Mr K. J. Houston, both Department of Primary Industries,
Brisbane, for identification of insects, and to the Director, Plant Pathology Branch, of
the same Department, Indooroopilly, for facilities to D.E.S. during the study.
References
BREWBAKER, J. L., and KWAck, B. H., 1963. — The essential role of calcium ion in pollen germination and
pollen tube growth. Amer. J. Bot. 50: 859-865.
FAEGRI, K., and PYL, L. van der, 1979. — The principles of pollination ecology. Oxford: Pergamon Press.
HAMILTON, A. G., 1898. — On the methods of fertilization of some Australian plants. Report of the
Seventh Meeting of the Australasian Association for the Advancement of Science, held at Sydney,
4898. Published by the Association. (pp. 557-565).
KORNERUP, A., and WANSCHER, J. H., 1961. — Methuen handbook of colour. London: Methuen & Co. Ltd.
MCALPINE, D. K., 1978. — Description and biology of a new genus of flies related to Anthoclusza and
representing a new family (Diptera, Schizophora, Neurochaetidae). Ann. Natal Mus. 32: 273-295.
SHAW, D. E., CANTRELL, B. K., and Houston, K. J., 1982. — Neurochaeta inversa McAlpine (Diptera:
Neurochaetidae) and seed set in Alocasia macrorrhiza (L.) G. Don (Araceae) in southeast Queensland.
Proc. Linn. Soc. N.S.W. 106: 67-82.
STANLEY, R. G., and LINSKENS, H. F., 1974. — Pollen: biology, biochemistry, management. Berlin, New York:
Springer-Verlag.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
Rabbits, Vegetation and Erosion
on Macquarie Island
P. M. SELKIRK, A. B. COSTIN, R. D. SEPPELT and J. J. SCOTT
SELKIRK, P. M., Costin, A. B., SEPPELT, R. D., & ScoTT, J. J. Rabbits, vegetation
and erosion on Macquarie Island. Proc. Linn. Soc. N.S.W. 106 (4), (1982) 1983:
337-346.
Rabbits were introduced to subantarctic Macquarie Island in about 1880.
Selective grazing by rabbits has been important in changing the floristic composition
and structure of grassland and herbfield vegetation, and the effects of grazing and
burrowing activities on erosion have concerned scientific visitors to the island for some
years.
Re-examination in 1980 of sites first documented in 1958 shows increases in
erosion at some sites, and revegetation of some areas. Photographic documentation is
provided for both sets of observations, and changes noted in the 1980 photographs are
discussed. The role of rabbits in erosion of grassland sites is difficult to assess, and may
in the past have been overestimated.
P. M. Selkirk, School of Biological Sciences, Macquarie University, North Ryde, Australia 2113;
A. B. Costin, Centre for Resource and Environmental Studies, Australian National University,
Canberra, Australia 2600; R. D. Seppelt, Antarctic Division, Department of Science and
Technology, Channel Highway, Kingston, Australia 7150, and J. J. Scott, Geography Depart-
ment, Monash University, Clayton, Australa 3168; manuscript recewed 6 July 1982, accepted for
publication in revised form 20 October 1982.
INTRODUCTION
Macquarie Island (54°30'S, 158°57'E) is a small subantarctic island with a moist,
cool, windy, oceanic climate. The vegetation includes tussock grassland, herbfield, fen
and bog communities, and feldmark (Taylor, 1955).
Since their first introduction in 1879 or 1880 (Cumpston, 1968), rabbits have
spread over most of the island and their numbers have increased dramatically. Their
population was estimated at 50 000 in 1956, 150 000 in 1965-6 (Sobey e¢ al., 1973),
50 000 in 1974 (Skira, 1979) and up to 150 000 in 1977-78 (Copson e¢ al., 1981). The
introduction of rabbits appears to have affected a number of components of the en-
vironment. They form the major item of food for feral cats on the island (Jones, 1977).
Taylor (1979) described the role of rabbits in the demise of the Macquarie Island
parakeet. The effect of rabbit grazing on the floristic composition and structure of
tussock grassland and herbfield was described by Taylor (1955), and further discussed
by Costin and Moore (1960), Johnston (1966) and Jenkin ef a/. (1981). The effects of
rabbit grazing and burrowing on erosion have concerned scientific visitors to the island
for some years, and have been the subject of several reports. Taylor (1955) and Costin
and Moore (1960) concluded that selective grazing by rabbits of the stabilizing species
in grassland, aggravated by burrowing, has substantially accelerated erosion of steep
slopes. Johnston (1966) and Griffin (1980) concluded that rabbit-induced erosion is of
minor importance in the overall erosion picture on Macquarie Island. Griffin (1980)
considered that soil creep is the major erosion mechanism on the island, and that rabbit
grazing has had little influence on this.
COMPARISON OF PHOTOGRAPHS
Eight sites, initially photographed in connection with a study of the role of rabbits
in erosion in December 1958 (Costin and Moore, 1960) were revisited and
rephotographed during January and February 1980. The sites show some changes over
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
338 RABBITS, VEGETATION AND EROSION ON MACQUARIE ISLAND
the 21-year period between the two sets of photographs. There have been some in-
creases in erosion (deepening of gullies, increased stone exposure in bare peat areas)
and revegetation of some areas. Comparison of the photographs and a discussion of
changes in the sites follows. Vascular plant nomenclature follows that of Greene and
Walton (1975) except for Luzula crinita which follows Edgar (1966). Bryophyte
nomenclature follows Seppelt (1980).
Figure 1: These steep slopes on which Poa foliosa tussock occurs appear unchanged in
1980 compared with 1958. Immediately above the limit of the photograph, however,
rabbit grazing is evident at the top of the steep grassland slopes having spread
downwards from the gentler herbfield slopes above.
Figure 2: The area of bare peat in right foreground has enlarged by 1980, compared
with 1958. The Poa foliosa tussocks on the knoll are stunted. The area of white (dead) P.
foltosa in the 1958 photo is, in 1980, occupied by a Luzula crinita — Festuca contracta short
tussock vegetation amongst which shoots and small clumps of P. foliosa are growing
quite prolifically. Present vegetation on the knoll includes Acaena minor, Azorella selago,
Colobanthus muscoides, Epilobium linnaeoides, Festuca contracta, Luzula crinita, Poa foliosa,
Ranunculus biternatus, Breutelia pendula, Diucranoloma billardiert, Ditrichum strictum,
Rhacomitrium crispulum.
Figure 3: The dark area of stable tussock (Poa foliosa) in the 1958 photo appears un-
changed in 1980. The white areas are now grassland with Agrostis magellanica — Luzula
crinitta dominated vegetation forming more or less continuous cover.
Both the fresh ‘peat flow’ landslip area (p) and the tumbled area (t) below it in the
1958 photograph have, in 1980, a complete plant cover. The present vegetation in both
‘peat flow’ and ‘tumbled’ areas is similar (unpublished quadrat data, J. J. Scott),
including: Acaena minor, Agrostis magellanica, Epilobium linnaeoides, Luzula crinita, Poa
annua, Ranunculus biternatus, Breutelia pendula, Drepanocladus sp., Marchantia berteroana,
Thuidium furfurosum.
To the west of the peat flow area are a number of old tussock bases, (b)
presumably rabbit grazed, some now overgrown with Luzula crinita and Acaena minor.
There is abundant evidence of rabbits in the area of photograph 3: rabbits, burrows,
squats and grazed tussock are all to be seen. In the lower left-hand corner of the 1980
photo is an area of landslip which has occurred since 1958. The ridge (r) between the
old and the new slip areas is covered with ‘terracettes’ the origin of which is ascribed by
Fig. 1. From a few metres southeast of Hurd Point hut, looking to west. In ail figures, photographs (A) taken
in 1958 by A. B. Costin, previously published in Costin and Moore (1960), photographs (B) taken in 1980
by P. M. Selkirk. Letters on photographs are discussed in text.
Fig. 2. From about 150 m altitude on west side of scree-slope track above Hurd Point hut, looking down onto
knoll to west of scree slope.
Fig. 3. North of Windsor Bay, south of Hurd Point-Caroline Cove track, taken from base of rocky outcrop to
north of track.
Fig. 4. Above Hurd Point, west of scree-slope track, on slope above hill in Fig. 2.
Fig. 5. Above Hurd Point, east of scree-slope track, on plateau edge. Photograph B taken from closer to
background ridge than photograph A.
Fig. 6. On Hurd Point-Caroline Cove track, below rocky knoll from which Figs 3 and 7 were taken.
Fig. 7.* East of Petrel Peak, north of Windsor Bay, south of Fig. 6, immediately to east of Fig. 3.
Fig. 8.* In Jessie Niccol Creek valley, from point at which Overland Track crosses south arm of Jessie
Niccol Creek, looking northwest toward Mt. Blake.
* Note that in Costin and Moore (1960) captions for photographs 7 and 8 were reversed.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
339
P.M. SELKIRK, A. B. COSTIN, R. D. SEPPELT AND J. J. SCOTT
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
RABBITS, VEGETATION AND EROSION ON MACQUARIE ISLAND
340
Proc. LINN. SOc. N.S.W. 106 (4), (1982) 1983
P.M. SELKIRK, A. B. COSTIN, R. D. SEPPELT AND J. J. SCOTT
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
RABBITS, VEGETATION AND EROSION ON MACQUARIE ISLAND
342
PROC. LINN. SOc. N.S.W. 106 (4), (1982) 1983
P.M. SELKIRK, A. B. COSTIN, R. D. SEPPELT AND J. J. SCOTT 343
Griffin (1980) to creep, solifluction and/or small scale slumping processes. We believe
the terracettes are initiated by small-scale slumping, then may be aggravated by rabbits
using them for squats and territorial digs, and the whole ultimately overgrown by
vegetation (J. J. Scott, unpublished data).
Figure 4: The tongue of vegetation on the far side of the bare peat area in the 1958
photograph is, in 1980, an elongated island. The bare peat area has enlarged slightly,
with a greater area of bare stones, and appears to be forming into a stony erosion
pavement which should offer some protection from frost-heave, wind- and water-
erosion to the remaining peat (Wimbush and Costin, 1979; Totterdell and Nebauer,
1973) allowing its colonization by mosses and angiosperms.
Figure 5: The areas bare of vegetation on the background ridge are similar in area in the
two photographs although perhaps more stones are exposed in 1980. The 1958
photograph caption describes rabbit burrowing undercutting dead tussocks above the
soil slip in the left background, but by 1980 there is no evidence of tussock grass having
been there very recently. Rabbits were still active in the area in 1980. On the
background slope (25°), present vegetation includes: Acaena magellanica, Agrostis
magellanica, Azorella selago, Cardamine corymbosa, Epilobium nerterioides, Luzula crinita,
Ranunculus biternatus, Stellaria decipiens, Breutelia pendula, Drepanocladus sp., Psilopilum
australe, Rhacomitrium crispulum.
The present vegetation on the gentle slope (5°-15°) of the foreground ridge, much
less prone to erosion than the steeper ridge behind, includes: Acaena minor, Cardamine
corymbosa, Epilobium nertertoides, Luzula crinita, Ranunculus biternatus.
Figure 6: The total area of bare sites has not altered substantially between 1958 and
1980, but some changes are visible. In the centre of the 1980 photograph a larger area
of stony pavement is visible than in 1958, exposed presumably by wind- and water-
erosion of surrounding peat particles. There is in 1980 a more distinct drainage line (d)
connecting the two non-vegetated areas.
Figure 7: In the bare areas on the upper slopes, gullies have deepened and some
‘islands’ of vegetation (i) have disappeared since 1958, presumably by the action of
frost, wind and water. Lower down the slope, gullies have deepened, exposing a
greater area of bare soil along the gully sides. Other areas, bare in 1958, show some
revegetation (v) in 1980.
Figure 8: The creek gully in 1980 is somewhat infilled at its junction with the small
tributary from the northwest, compared with the 1958 photograph. The infill material
has presumably been brought down by the tributary creek. Since its deposition it has
been densely covered with bryophytes. Upstream of the junction with the tributary,
however, the creek gully still has quite steep sides as in 1958.
GENERAL DISCUSSION
In trying to assess the contribution of rabbits to soil erosion on Macquarie Island,
a number of factors must be considered: effect of rabbit activities (burrowing, squats,
territorial digs, grazing), size of rabbit population, extent of rabbit activities, and other
factors contributing to surface instability.
Undercutting of tussocks by rabbit burrowing was described by Costin and Moore
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
344 RABBITS, VEGETATION AND EROSION ON MACQUARIE ISLAND
(1960). The suggested role of rabbit squat and territorial digging activities in ac-
centuating terracettes has been mentioned above. It is by their selective grazing that
rabbits exert most effect on the Macquarie Island environment. Areas may be com-
pletely denuded of their vegetation when this was composed entirely of palatable
species. These areas may be subject to erosion until colonized by less palatable and/or
grazing-adapted species. An example is a steep slope above the beach just south of
Nuggets Point beach, previously clothed with Stilbocarpa polaris and a little Poa foliosa on
which rabbit grazing was severe during 1978-9 (G. Copson, pers. comm.). The area
was bared when the Stlbocarpa was grazed down to its rhizomes. In 1980, rapid
colonization of the soil surface was being achieved principally by Marchantia berteroana
(via its abundantly-produced gemmae), but also by Poa annua, Epilobium linnaeoides, E.
nertertordes and Cardamine corymbosa. Marchantia berteroana seemed only rarely to be
touched by the rabbits which were still present in the area. Erosion of the surface
appeared minimal.
The recolonization of areas bared by rabbit grazing appears similar in general
terms to the recolonization of areas bared of vegetation by landslips, but with some
important differences ( J. J. Scott, unpublished data). These arise because landslips
commonly remove all vegetation plus a peat layer, while selective grazing by rabbits
usually leaves some (possibly less palatable) vegetation on the site. For example, grazing
on the coastal slopes usually removes the dominant species Poa foliosa and Stilbocarpa
polaris, allowing remaining subsidiary species such as Ranunculus biternatus, Acaena minor
and Agrostis magellanica to achieve local dominance, in similar fashion to the infilling of
gaps between senescing Poa foliosa tussocks (Ashton, 1965). A landslip removes both
dominant and subsidiary species, as well as surface litter, seeds and underground
rhizomes in a peat layer of varying thickness. Revegetation proceeds much more slowly
than on areas bared by grazing. In the absence of further grazing once an area has been
bared, both landslip surface and areas grazed by rabbits can eventually be revegetated
by their former dominant species. However, if regeneration of Poa foliosa and Stilbocarpa
polaris is suppressed by continued grazing of landslip- or rabbit-bared surfaces, a short
tussock grassland characterized by Agrostis magellanica, Luzula crinita and Acaena spp.
develops.
In areas whose dominant vegetation includes a mixture of palatable and un-
palatable species, selective grazing by rabbits can have a very marked effect on species
composition. For example, the rabbit population of the Flat Creek valley, near Bauer
Bay, was dramatically reduced by myxomatosis during winter 1979. The quantities of
Pleurophyllum hookeri, Stellaria media and Cerastium fontanum, all preferentially eaten by
rabbits, were observed to increase noticeably in this valley between the beginning and
the end of the 1979-80 summer.
The floristic composition of the short tussock grasslands and herbfields has been
particularly affected by rabbit grazing. Pleurophyllum hookert now seems less widespread
in the herbfields than Taylor’s (1955) photographs and descriptions would suggest.
The highly palatable Pleurophyllum is badly damaged when rabbits graze the leaves right
down to the stem stump. Species which are now more abundant in the herbfield than
Taylor’s (1955) descriptions suggest may be less palatable to rabbits (e.g. Luzula crinita)
or able to withstand and thrive despite grazing, by virtue of abundant lateral shoot
growth (e.g. Acaena spp.), tillering (Poa annua, Agrostis magellanica) or seed set (Acaena
spp., Poa annua, Agrostis magellanica, Uncinia compacta). Copson et al. (1981) note that the
majority of rabbits live in the herbfield vegetation, including the short tussock
grasslands mentioned above. Few rabbits live in the tussock grassland, bog, fen and
feldmark formations as described by Taylor (1955), although these vegetation types are
grazed by rabbits from nearby herbfield areas (Copson et al., 1981).
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
P.M. SELKIRK, A. B. COSTIN, R. D. SEPPELT AND J. J. SCOTT 345
Early descriptions (Cumpston, 1968) and recent estimates (Sobey et al., 1973;
Skira, 1979; Copson et al., 1981) of rabbit numbers on the island are consistent with the
suggestion that, in common with introduced species elsewhere, the rabbit population
on Macquarie Island was low for a long period after its introduction, went through a
period of rapid expansion to a maximum, perhaps beyond the capacity of the habitat to
support the numbers, then fell again to a lower level, in balance with the capacity of the
habitat to support it. Within this general picture, numbers appear to have fluctuated:
for example, a peak between 1880 and 1906 is discussed by Taylor (1979) using data
from Cumpston (1968), and a peak occurred in the 1977-78 summer due to several
successful breeding seasons preceding it (Copson et al., 1981). It is suggested by
Copson é al. that annual population fluctuations are a result of seasonal changes in
weather, with extended drier periods leading to higher survival rate of kittens.
Dense rabbit populations have occurred in different places on the island at dif-
ferent times (maps, Sobey é al., 1973). Detailed rabbit counts have been made in
recent years by officers of the Tasmanian National Parks and Wildlife Service but the
size of the population at any time is not easy to estimate. The times of peak populations
in the past may never be known, but it may be assumed that maximal effect by rabbits
on an environment would correspond with peak populations in the area. Such an area
would be especially vulnerable to accelerated erosion, if there were a lag between the
initial damage to the taller-growing dominant plants (Poa foliosa, Stilbocarpa polaris) and
the subsequent succession of lower-growing herbs and mosses, as in Figs 2, 3 and 5.
The observations of Costin and Moore (1960) may have been made soon after a period
of maximal rabbit population in the Hurd Point area, and consequently maximal
deleterious effect on the local environment. Although rabbit activities may have
provided a trigger for more than usual peat instability, it is clear that the predicted
extension of damaged areas (Costin and Moore, 1960) is not occurring, except locally.
Griffin (1980) discussed a number of erosive forces active on Macquarie Island,
namely soil creep, landslips triggered both by unusually heavy rain periods and by
earthquakes, wind erosion and rabbit activities. In addition, royal penguin, gentoo
penguin and giant petrel activity on some of the lower coastal slopes contributes locally
to the erosion of peaty soil between Poa foliosa tussocks, thereby affecting the balance
and stability of the tussocks. After prolonged rain, slipping of these unstable situations
may occur.
In the presence of these other erosive forces on Macquarie Island, the role of
rabbits in initiating and contributing to continued erosion of grassland areas is difficult
to assess, and may in the past have been overestimated. Through their selective
grazing, however, rabbits have had major effects in changing the floristic composition
and structure of much of the original grassland and herbfield vegetation.
ACKNOWLEDGEMENTS
Permission from Tasmanian National Parks and Wildlife Service, and logistic
support from Australian Antarctic Division for visits to Macquarie Island by the
authors are gratefully acknowledged. Thanks are due to Mr G. Copson for help in
location of sites, to Dr J. Jenkin for valuable comments on this paper, to Dr N. Wace
and Dr B. Fennessey for helpful discussions, and to the editors of the Journal of Ecology
for permission to reproduce photographs originally published in that journal.
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
346 RABBITS, VEGETATION AND EROSION ON MACQUARIE ISLAND
References
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Corson, G. R., BROTHERS,, N. P., and SkiIRA, I. J., 1981 — Distribution and abundance of the rabbit,
Oryctolagus cuniculus (L.), at subantarctic Macquarie Island. Aust. Wildl. Res. 8:597-611.
CosTIN, A. B., and Moore, D. M., 1960 — The effects of rabbit grazing on the grasslands of Macquarie
Island. J. Ecol. 48: 729-732.
CUuMPSTON, J. S., 1968 — Macquarie Island. ANARE Scientific Reports, Series A(1), Narrative. Antarctic
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EpGaR, E., 1966 — Luzula in New Zealand. N.Z. J. Bot. 4: 159-184.
GREENE, S. W., and WALTON, D. W. H., 1975 — An annotated checklist of the Subantarctic vascular flora.
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GRIFFIN, B. J., 1980 — Erosion and rabbits on Macquarie Island: some comments. Pap. Proc. R. Soc. Tasm.
114: 81-83.
JENKIN, J. F., JOHNSTONE, G. W., and Copson, G. R., 1981 — Introduced animal and plant species on
Macquarie Island. Colloque sur les Ecosystémes Subantarctiques, Paimpont, C.N.F.R.A. 51: 301-313.
JOHNSTON, G. C., 1966 — Macquarie Island and its rabbits. Tasm. J. Agric. 37: 277-280.
JONES, E., 1977 — Ecology of the feral cats Felis catus (L.), (Carnivora: Felidae) on Macquarie Island. Aust.
Wildl. Res. 4:249-262.
SEPPELT, R. D., 1980 — A synoptic moss flora of Macquarie Island. Antarctic Division Technical Memorandum
Number 93, 8 pp.
SKIRA, I. J., 1979 — Studies of the rabbit population on Macquarie Island. Hobart: University of
Tasmania, M.Sc. thesis, unpubl.
SOBEY, W. R., ADAMS, K. M., JOHNSTON, G. C., GOULD, L. R., Simpson, K. N. G., and KEITH, K., 1973
— Macquarie Island: the introduction of the European rabbit flea Spzlopsyllus cuniculi (Dale) as a
possible vector for myxomatosis. J. Hyg., Camb. 71: 299-308.
TAYLOR, B. W., 1955 — The flora, vegetation and soils of Macquarie Island. ANARE Reports, Series B,
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TOTTERDELL, C. J., and NEBAUER, N. R., 1973 — Colour aerial photography in the reappraisal of alpine
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Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
Index
VOLUME 106
Page
Alocasia macrorrhiza, Crustaceans from Australian mangroves... . . 112
pollamationlot; fs Suk ha de sls tka ee 323. Cupala Creek Formation (Cambrian)....... 129
SECCbse tpl peusra tachi: oaAlens Mh. stot ule 67
Amaurobioidea, (Araneae) ........... 247, 275 Dapto Latite Member, of Broughton For-
Anderson, D. T., Origins and relationships ration (HEAVEN). o5cceces¢eeoccone. 292
among the animal phyla .............. 151 Darriwilian (middle Ordovician) graptolites
AUDIOS TOS co. 028 ROS ao. Hi Fegeo aenueat hE ER ee ME 142 from the Monaro Trough sequence east of
?Aphelaspis sp. aff. Braidwood, New South Wales, by C. J.
Al, GUSTS Ew fonts oe ae ee a ee 142 Henkin sie dihesye ts alee etre Ria eae 173
Araceae (Alocasia macrorrhiza) ........... ORS 2 3) eee Diptera Neunochachica cli lei aeer 67
Araneae, Amaurobioidea............. UE Pea Oipterawilele omilyzidac ein ie ee 33, 59
Archaeobalanidae, (Balanoidea)........... 22
AOMTISED GSDa WOonc 6 cae ea en dea ae Hoe 275 Ecological values of the tropical rainforest
Austmusia kiolbasp.nov. ............-.-.-. 281 resource, Macleay Memorial Lecture, by
Austmusia lindi sp.nov. ...........+.----. 283 IE Wich breast aes Sere atic eee 263
Austmusia wilsonisp.nov. ................ OE Eiliminvusicovertusts DOV Ae eee 24
A. W. H. Humphrey, His Majesty’s Hilminausynodes tus apie ee eee 23
mineralogist in New South Wales, 1803-12 Elmside Formation (Siluro-Devonian) ...... 167
—Acomment,byR.J.Ford.......... 123 Eucalyptus bugaldiensis sp. nov.............. 301
Eucalyptus spp. (middle Miocene) .......... 305
PRC LUUITNT Oleaeg Ne opp ea ae Bie ay Ric tea visas apse sah 249 Fauna of Australian mangroves, by P. A.
Badumna candidacomb. nov. .............. 250 Hutchings & H.F. Recher ............ 83
Badumna gausapatacomb. nov.............. 254 Fauna of Australian seagrass beds, by P. A.
Badumna vandiemeni sp. nov................ 257 Flutchinigss seh hee oer oes cence 181
Barnaclesmioalanoidyes se so 4 oe sae center 21 Five Islands Latite Member, of Pheasants Nest
Berkley Latite Member, of Pheasants Nest - Kormation| (Permian) se eee eee 293
Formation (Permian) ................ 295. Ford, R. J., A. W. H. Humphrey, His
[ETI RECALLS Ovaen cet eR Gt Siri. paren Ses er oa Ac 147 Majesty’s mineralogist in New South
Blow Hole Latite Member, of Broughton Wales, 1803-12 — Acomment ......... 123
Formation (Permian) ................ 289s Fossil Eucalyptus remains from the middle
Brachiopods, Cambrian................. 147 Miocene Chalk Mountain Formation,
Broughton Formation, of Shoalhaven Groap Warrumbungle Mountains, New South
(Renmian)) beaters. cen ns, getters. 28. 289 Wales, by W. B. K. Holmes, F. M.
Bugaldie, N.S.W., fossil Eucalyptus from..... 299 Folmme sya lel ay Ate Van; Gan 299
Bumbo Latite Member, of Broughton For- Foster, B. A., Two new intertidal balanoid
mation((Permian)),.3.°.>.-..-...- 2 291 barnacles from eastern Australia........ 21
Calderwood Latite Member (new name), of Graptolites 25) as28 ee ee e 167, 173
Pheasants Nest Formation (Permian) .... 294 Gray, M. R., A new genus of spiders of the
Cambewarra Latite Member, of Broughton subfamily Metaltellinae (Araneae, Amauro-
Formation (Permian) ................ 292 bioidea) from southeastern Australia. .... 275
Cambrian (Idamean) rocks and fossils, north- Gray, M. R., The taxonomy of the semi-
WWE SLOTIMVINS AV cee arues cog eo tas ai tsiad apal Geset ce oes = 136. communal spiders commonly referred to
Cantrell, B. K., see Shaw, D. E. the species Jxeuticus candidus (L. Koch) with
Carr, P. F., A reappraisal of the stratigraphy notes on the genera Phryganoporus, Ixeuticus
of the upper Shoalhaven Group and lower and Badumna (Araneae, Amaurobioidea).. 247
Illawarra Coal Measures, southern Sydney
Basin, New South Wales.............. 23 tleleomyzidae.y (Diptera) ys. eee 33599
Chalk Mountain Formation (middle Miocene) IE CERTTE TOS (GEN INO 20 oo gy 0002 2o58 5555 28
TENNPINEN IDSA o (oh aes Ubou alla suse eos ence aleenes eet OND 300 Hexaminius popeianasp.nov. ...........-.- 28
Comparative morphological study of the Holmes, F. M., see Holmes, W. B. K.
reproductive systems of some species of Holmes, W. B. K., Holmes, F. M., & Martin,
Tapeigaster Macquart (Diptera: Heleo- H. A., Fossil Eucalyptus remains from the
myzidae), by M. A. Schneider.......... 59 middle Miocene Chalk Mountain For-
Costin, A. B., see Selkirk, P. M. mation, Warrumbungle Mountains, New
Crane, D., see Powell, C. McA. SouthWralesiii sinus 6.0 eres Ase 299
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
348
Houston, K. J., see Shaw, D. E.
Jelnnanyoores7, AN. Wo lal, oc sccooncccocunde:
Hutchings, P. A., The fauna of Australian
seagrass beds
Hutchings, P. A., & Recher, H. F., The fauna
of Australian mangroves..............
Idamean (late Cambrian) fossils...........
Illawarra Coal Measures (lower), Permian, of
southern Sydney Basin
Txeuticus candidus syn... ........+005++005.
Jamberoo Sandstone Member, of Broughton
Formation (Permian)
Jell, P. A., see Powell, C. McA.
Jenkins, C. J., Darriwilian (middle-
Ordovician) graptolites from the Monaro
Trough sequence east of Braidwood, New
South Wales 0s 55 2s ois aie ciate:
Jenkins, C. J., Late Pridolian graptolites from
the Elmside Formation near Yass, New
SouthiWaalesi ira. sins cola suroei estes
Jonsell, B., Linnaeus and his two circum-
navigating apostles..................
Kent, D.S., see McAlpine, D. K.
Kiama Sandstone Member, of Broughton
Formation (Permian)
Kuo, J., Notes on the biology of Australian
SEAGT ASSES eNetreee aun sets her Npussaencons ©
Larkum, A. W. D., & West, R. J., Stability,
depletion and restoration of seagrass beds .
Larkum, A. W. D., see also West, R. J.
Late Pridolian graptolites from the Elmside
Formation near Yass, New South Wales,
byi@is Jienkinsiew see ice ee wacree aie a
Linnaeus and his two circumnavigating
apostles, by B. Jonsell................
Linnean Society of New South Wales. Record
of the annual general meeting 1981.
Reports and balance sheets..... annexure
Linograptus posthumus...................-.
Long distance transport of spores of Puccinia
graminis tritici in the southern hemisphere,
by I. A. Watson & C.N.A.deSousa ....
McAlpine, D. K., & Kent, D. S., Systematics
of Tapeigaster (Diptera: Heleomyzidae) with
notes on biology and larval morphology. . .
Macleay Memorial Lecture, 1982..........
Macquarie Island, rabbits, vegetation and
ELOSIONN jest ees teers Guages ae oe eee
Mangroves, faunaofAustralian...........
Martin, H. A., see Holmes, W. B. K.
Metaltellinae, Amaurobioidea............
Minumurra Latite Member, of Pheasants Nest
Formation (Permian)
Miocene Eucalyptus remains
Mollusc, Cambrian
Molluscs from Australian mangroves
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
INDEX
123
181
83
127
287
247
291
173
311
Monaro Trough sequence east of Braidwood,
graptolites
Monograptus cf. angustidens ................
Monograptus cf. formosus..................
Monograptus transgrediens..................
Monograptus transgrediens cf. praecipuus........
Neef, G., see Powell, C. McA.
Neurochaeta inversa McAlpine (Diptera:
Neurochaetidae) and seed set in Alocasia
macrorrhiza (L.) G. Don (Araceae) in
southeast Queensland, by D. E. Shaw, B.
K. Cantrell & K. J. Houston...........
New genus of spiders of the subfamily
Metaltellinae (Araneae, Amaurobioidea)
from southeastern Australia, by M. R.
Gray
Notes on the biology of Australian seagrasses,
bya)e Kuo cbsceet ne oes Stal om) caucenee ae
Notoaphelaspis gen. nov..................--
Notoaphelaspis orthocephalis sp.nov...........
Notoaphelaspis sp. cf. N. orthocephalis sp. nov... .
Obolid, (Obolidae), indet................
Ordovician (middle), graptolites...........
Origins and relationships among the animal
phyla, by D.T.Anderson.............
Palynology, Chalk Mountain Formation. ... .
Percival, I. G., see Powell, C. McA.
Permian stratigraphy, south Sydney Basin .. .
Pheasants Nest Formation, lower Illawarra
Coal Measures (Permian).............
Phryganoporus syn.............220+-204--
Pollination of Alocasia macrorrhiza...........
Polychaetes from Australian mangroves
Polychaetes from seagrass beds............
Powell, C. McA., Neef, G., Crane, D., Jell, P.
A., & Percival, I. G., Significance of late
Cambrian (Idamean) fossils in the Cupala
Creek Formation, northwestern New South
Pridolian, (Siluro-Devonian), graptolites ....
Prismenaspis?'sp. MOV. . =... 5-252 25:
PrOplina sp satel is, Sashes cies pita den curmaee te terere
Pseudagnostus sp. aff. idalis
Puccinia graminis tritici, transport of spores of. . .
Rabbits, vegetation and erosion on Macquarie
Island, by P. M. Selkirk, A. B. Costin, R.
D. Seppelt & J. J. Scott...............
Rainforest, tropical, ecological values.......
Reappraisal of the stratigraphy of the upper
Shoalhaven Group and lower Illawarra
Coal Measures, southern Sydney Basin,
New South Wales, by P. F.Carr........
Recher, H.F., see Hutchings, P. A.
Reproductive systems of Tapemgaster.........
Saddleback Latite Member, of Broughton
Formation (Permian)
Schneider, M. A., A comparative mer-
173
169
171
171
171
67
275
225
142
144
145
148
173
151
305
287
293
247
323
110
188
127
167
145
146
140
311
337
263
287
59
291
phological study of the reproductive
systems of some species of Tapeigaster
Macquart (Diptera: Heleomyzidae)...... 59
Seagrass and seagrass beds... .. 181, 201, 213, 225
Seagrass primary production — a review, by
R. J. West & A.W. D.Larkum......... 213
Selkirk, P. M., Costin, A. B., Seppelt, R. D.,
& Scott, J. J., Rabbits, vegetation and
erosion on Macquarie Island........... 337
Seppelt, R. D., see Selkirk, P. M.
Scott, J. J., see Selkirk, P. M.
Shaw, D. E., & Cantrell, B. K., A study of
pollination of Alocasia macrorrhiza (L.) G.
Don (Araceae) in southeast Queensland... 323
Shaw, D. E., Cantrell, B. K., & Houston, K.
J., Neurochaeta inversa McAlpine (Diptera:
Neurochaetidae) and seed set in Alocasia
macrorrhza (L.) G. Don (Araceae) in
southeast Queensland................ 67
Shoalhaven Group (upper), Permian, of
southern Sydney Basin............... 287
Significance of late Cambrian (Idamean) fossils
‘in the Cupala Creek Formation, north-
western New South Wales, by C. McA.
Powell, G. Neef, D. Crane, P. A. Jell & I.
(Gaeercivale yan Seay a. cierr Gs suacot oss) eet 127
Solandeny Daniel’. tcc. so) sa: cls a ae 1
Sousa, C. N. A. de, see Watson, I. A.
SpauumiankyAnders)-e1420-2 2) oe ons 1
Stability, depletion and restoration of seagrass
beds, by A. W. D. Larkum & R.J.West.. 201
Stigmatoa tysont) . 6. ee es 140.
Study of the pollination of Alocasta macrorrhiza
(L.) G. Don (Araceae) in southeast
Queensland, by D. E. Shaw & B. K.
(Clanntinellls cacceaseede pe peteens icc eens eer 323
INDEX
Sydney Basin, southern, stratigraphy of .....
Systematics of Tapeigaster (Diptera: Heleo-
myzidae) with notes on biology and larval
morphology, by D. K. McAlpine & D. S.
INE n bites sr gusty crpie ean Reese Ge apo Sts cee ee 33
TG PeUgas lenin Sedo t, 5 hye cn eneAS tg POPS LENORE 34, 59
Tapeigaster annulatacomb.nov. ....... . 48, 61, 63
Tapeigaster annulipes.............+.+... 42, 61, 63
Tapeigaster argyrospila..............+.+4+. 39
Tapeigaster brunneifrons..........-..+400-- 39
Tapeigaster cinctipes comb. nov.............. 38
Tapeigaster digitatasp.nov................ 45, 63
Tapeigaster luteipennis ................ 41, 61, 63
Tapeigaster nigricornis comb. nov. ...........- 37
Tapeigaster paramonovi sp.nov.............. 44
Tapeigaster pulvereasp.nov. ...........- 50, 61, 64
Tapeigaster subglabrasp.nov. .......... 52, 63, 64
Mrilobitess sje ree nes ee Se ates eee Se cee 140
Two new intertidal balanoid barnacles from
eastern Australia, by B.A. Foster....... 21
Watson, I. A., & Sousa, C. N. A. de, Long
distance transport of spores of Puccinia
graminis tritici in the southern hemisphere. .
Webb, L. J., Ecological values of the tropical
rainforest resource (Macleay Memorial
Lecture, 1982)
West, R. J.. & Larkum, A. W. D., Seagrass
311
primary production — areview......... 213
West, R. J., see also Larkum, A. W. D.
Westley Park Sandstone Member, of
Broughton Formation (Permian)........ 289
Yass, N.S.W., graptolitesfrom............ 167
Proc. LINN. Soc. N.S.W. 106 (4), (1982) 1983
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PROCEEDINGS of LINNEAN SOCIETY OF NEW SOUTH WALES
VOLUME 106
Issued May 10, 1983
CONTENTS:
NUMBER 3
201 A.W.D.LARKUM and R. J. WEST
Stability, Depletion and Restoration of Seagrass Beds
213 R.J.WEST andA. W. D. LARKUM
Seagrass primary Production — aReview
225 ..J. KUO
: Notes on the Biology of Australian Seagrasses
247 M.R.GRAY
The Taxonomy of the semi-communal Spiders commonly referred to the Species
Ixeuticus candidus (L. Koch) with Notes on the Genera Phryganoporus, Ixeuticus and
Badumna (Araneae, Amaurobioidea)
NUMBER 4
263 L.J.WEBB
Sir William Macleay Memorial Lecture 1982
Ecological Values of the tropical Rainforest RESOUICE
275 M.R.GRAY -
A new Genus of Spiders of the een Metaltellinae (Araneae, Amaurobioidea) from
southeastern Australia’
287 P.F.CARR
A Reappraisal of the Stratigraphy of the upper Shoalhaven Group and lower Illawarra
Coal Measures, southern Sydney Basin, New South Wales
299 W.B.K.HOLMES, F.M. HOLMES and H. A. MARTIN
Fossil Eucalyptus Remains from the middle Miocene Chalk Mountain Formation,
Warrumbungle Mountains, New South Wales
' 311 |. A. WATSON andC.N. A.deSOusA
Long distance Transport of Spores of Puccinia graminis tritici in the southern
Hemisphere
323 D.E.SHAW and B. K. CANTRELL
A Study of Pollination of Alocasia macrorrhiza (L.) G. Don (Araceae) in southeast
Queensland
. 337. +P.M.SELKIRK, A. B. COSTIN, R.D. SEPPELT and J. J. SCOTT
Rabbits, Vegetation and Erosion on Macquarie Island
347 INDEX to Volume 106
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