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PROCEEDINGS

OF THE

LINNEAN SOCIETY

NEW SOUTH WALES

VOLUME
109
(Nos 477-480; for 1986-87)

Sydney
The Linnean Society of New South Wales
1987

von ae a

Contents of Proceedings
Volume 109

NUMBER 1 (No. 477)
(Issued 20th March, 1987)*

WRIGHT, R. V. S. Sir William Macleay Memorial lecture 1984. New light on
Jae QA MOveOrn Orr Horo ANU VizvA TANECAN ANNO 5 ob yoo ocean ase ocd peo ead ace
KING, R. J. Aquatic angiosperms in coastal saline lagoons of New South Wales.
line ne cetanionioileake, Miaccuanies tar pean ieee eae eae eee
KING, R. J., and HOLLAND, V. M. Aquatic angiosperms in coastal saline la-
goons of New South Wales. II. The vegetation of Tuggerah Lakes, with
specific comments on the growth of Zostera capricornt Ascherson..........
KING, R.J., and BARCLAY, J. B. Aquatic angiosperms in coastal saline lagoons of
New South Wales. III. Quantitative assessment of Zostera capricorni.......
KING, R. J., and HODGSON, B. R. Aquatic angiosperms in coatal saline lagoons
OfINewsSouthevValestINe cong term changes sneer y arene nn

NUMBER 2 (No. 478)
(Issued 20th March, 1987)*

CARR, P. F., and JONES, B. G. Non-contemporaneity in the Marulan Batholith
BERGSTROM, D. M. An atlas of seeds and fruits from Macquarie Island.......
HOWDEN, A. T. Notes on the biology of adult and immature Amycterinae
(Colcopteray Curculionidae). tases sae ne EN ale ee erate ee
ANDERSON, D. T. The circumtropical barnacle Ytraclitella divisa (Nilsson-
Cantell) (Balanomorpha, ‘Tetraclitidae): cirral activity and_ larval
Gl EVE OMT Hit ees oe eh ye Rew Cotten ty cnt Ls itn adn gui A Nie cia genteel sal otsck ae
CARTER, J. J. Metagenesis as a possible key to animal form.................
Annexure to Numbers 1 and 2. The Linnean Society of New South Wales.
Record of the Annual General Meeting 1985. Reports and balance sheets.

* Publication of this issue, intended for December 1986, was delayed by a serious fire at the printery that
month. In consequence, the whole work of preparation had to be done again; fortunately, the original

figures were saved.

NUMBER 3 (No. 479)
(Issued 22nd December, 1987)

WATERHOUSE, J. T. Presidential Address 1979. The phylogenetic significance of
iDracaena=ty pe: Growth swale satya: e.g A oobi Me irs ape) Sere een Ne a
QUINN, C. J. Memorial Series no. 27. John Teast Waterhouse 1924-1983......
BARLOW, C. G., MCLOUGHLIN, R., and BOCK, K. Complementary feeding
habits of golden perch Macquaria ambigua (Richardson) (Percichthyidae)
and silver perch Brdyanus bidyanus (Mitchell) (Teraponidae) in farm dams.
PARNABY, H. E. Distribution and taxonomy of the long-eared bats Nyctophilus
gould: Tomes, 1858 and Nyctophilus bifax Thomas, 1915 (Chiroptera: Ver-
spertilionidae) imveastern) Australialy cra cite nie ea eee
MCMINN, A. Late Pleistocene dinoflagellate cysts from Bulahdelah, northern
New. Souths Wallésiiiht. 2 Cee Mae rat SI ntact eee eee
ROWE, F. W. E., and ALBERTSON, E. L. The echinoderm genus Henncia Gray,
1840 (Asteroidea: Echinasteridae) in southern and southeastern Aus-
tralian waters, with the description of a new species...................
ROWE, F. W. E., and ALBERTSON, E. L. A new species in the echinasterid genus
Echinaster Miller and Troschel, 1840 (Echinodermata: Asteroidea) from
southeastern Australia and) Norfollalslande ss se sins 79 ee
SHEA, G. M. Two new species of Delma (Lacertilia: Pygopodidae) from north-
eastern Queensland and a note on the status of the genus Aclys..........

NUMBER 4 (No. 480)
(Issued 22nd December, 1987)

MARTIN, H. A. Presidential Address 1982. Cainozoic history of the vegetation
and climate of the Lachlan River region, New South Wales............
AULD, T. D. Post-fire demography in the resprouting shrub Angophora hispida
(Sm.) Blaxell: flowering, seed production, dispersal, seedling establish-
ment and survival sie! orca ae a Gee LE rt aoe
MORRISON, D. A. A review of the biology of Acacia suaveolens (Smith) Willd.
(Mlimosaceae) iio ai oP ae EG as eee Se oe ea OE ee
FERGUSSON, C. L. Multiple folding of the Ordovician sequence, Tambo River,
CAstenn'e VICtOriame 2) ined n seni, Mga een glia ahah Tes eee aan
TIMMS, B. V. Geomorphic and physicochemical features of floodplain water-
bodies'of the lowersblunter ValleyaIN'SIWe oe eee
WATSON, J. E. Records of Eudendrium (Hydrozoa: Hydroida) from New Zealand
IVANTSOFF, W., CROWLEY, L. E. L. M., and ALLEN, G. R. Description of a
new species of freshwater hardyhead Craterocephalus katlolae (Pisces:
Atherinidae) trom satia,"Papua New Guinean. 24.) 05 ee ee
INDEX to Proceedingsvol, OQ. sec tails atales atop ce 2 Pol oc ste hen keg ee

1,
139

143

193

7S)

183

203

Zila

259

279

293
311
325

331
38)

PROCEEDINGS
of the

LINNEAN"
SOCIETY

NEW SOUTH WALES

VOLUME 109
NUMBER 1
NUMBER 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.

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 $35.00.

The current rate of subscription to the Proceedings for non-members is set at $45.00 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 pecan): (/

OFFICERS AND COUNCIL 1986-87

President: P,. M. MARTIN

Vice-presidents: HELENE A. MARTIN, GRO PHIPPS, C.Ni SMITHERS San. T.
WRIGHT

Honorary Treasurer? A. RITCHIE

Secretary) BARBARA J. STODDARD

Council: A. E. J. ANDREWS, JUDITH H. K. EASTMAN, R. A. FACER, L. W. C.
FILEWOOD, MARILYN D. FOX, L. A. S. JOHNSON, HELENE AS
MARTIN, P. M. MARTIN, J. R. MERRICK, P. J. MYERSCOUGH,
I, G. PERGIVAL, G. RK: PHIPPS, A. RITCHIE, (Ci\N: (SMIMHER SG. seG:
VALLANCE, KAREN L. WILSON, A. J. T. WRIGHT

Honorary Editor: T. G. VALLANCE — Department of Geology & Geophysics,
University of Sydney, Australia, 2006.

Linnean Macleay Fellow: R. W. JOHNSTONE

Linnean Macleay Lecturer in Microbiology: K.-Y. CHO

Auditors: W. SINCLAIR & CO.

The postal address of the Society is P.O. Box 457, Milson’s Point 2061, N.S.W.,
Australia. Telephone (02) 929 0253.

Cover motif: Transverse section (x 2.4) of the
Devonian coral described by A. J. T.
Wright as Melrosia rosae gen. et sp. nov.
From the Mount Frome Limestone,
Mudgee district, N.S.W.
Adapted by Len Hay from Proc. Linn.
Soc. N.S.W. 90, 1966, p. 266 (fig. 3).

PROCEEDINGS
of the

LINNEAN
SOCIETY

NEW SOUTH WALES

pa EE 2 ea ec eres
' Marine Biological Laboratory y |
LIBRARY H

APR 17 1987

Woods Hole, Mass. ;

VOLUME 109
NUMBER 1

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THE SIR WILLIAM MACLEAY MEMORIAL LECTURE 1984

New Light on the Extinction of the
Australian Megafauna

RICHARD WRIGHT

Department of Anthropology, University of Sydney

[Delivered 18 July 1984]

Coming as I do from the University of Sydney I frequently encounter the name of
Sir William Macleay. The energies of the staff of the Macleay Museum ensure that his
name is kept in front of us. ‘

The subject of Oceanic prehistoric archaeology has an important place within the
Macleay Museum. Thus my sort of archaeology is, within the University of Sydney,
associated with the name of Macleay. By contrast archaeology has not been so
associated, at least in this century, with your Society. I have looked at the contents of
your Proceedings and suspect that mine may be the first contribution in prehistoric
archaeology for some eighty years. I therefore feel that it is a double honour for me to be
giving the Sir William Macleay Memorial Lecture to your Society. The honour is both a
personal one, and one for the discipline of prehistoric archaeology within Australia.

Of course, much of Australian archaeology is not relevant to the aims of your
Society. I would not expect you, for instance, to find a place for research into the
manufacturing sequence of stone tools, or a study of their functions. However the study
of megafaunal extinctions, incorporating as it does such studies as zoology and botany,
is clearly relevant to your Society.

Extinctions is so manifestly a topic of what used to be called natural history that
some of you may be wondering why I should be claiming it as also a topic for study by
archaeologists. Let me give you two reasons why it should be. The first reason is chrono-
logical, and the second methodological. The chronological evidence that megafauna
and prehistoric humans overlap is increasing. The period of overlap was evidently some
tens of millennia. The methodical reason, related to the chronological one, is that
archaeologists, in their approach to studying past events in the field, are accustomed to
paying minute attention to stratigraphic detail. Of course such attention is not restricted
to prehistorians: indeed we have recently seen some palaeontologists, examining the ex-
tinction of dinosaurs at the end of the Cretaceous, reducing their research objective to
the study of the chemical properties of a band of dust 10mm thick. Nevertheless, though
they do not possess a technique peculiar to themselves, archaeologists have a strong
tradition of relishing the fine detail of the context in which their specimens are found, as
much as they relish the specimens. We do not dig for specimens as we might dig for
potatoes.

Field work designed to establish the relationship between humans and megafauna
will always require fine attention to contextual detail. From this point of view it is for-
tunate that the question of Pleistocene extinctions, in both the Americas and Australia,
is being answered by the use of archaeological methods.

It has not always been thought that the question of Pleistocene extinctions in Aus-
tralia was one for archaeologists. Until a decade ago in Australia one could respectably

PROC. LINN. SOC. N.S.W., 109 (1), 1986

EXTINCTION OF THE AUSTRALIAN MEGAFAUNA

argue, though with some opposition, that megafaunal extinctions pre-dated the first
settlement of this continent by the ancestors of the Australian Aborigines. Where were
the archaeological sites, with unambiguous stratigraphy and radiocarbon dates, at
which the sceptical excavator could repeatedly find artifacts and megafauna in associ-
ation? After decades of armchair speculation and (I would say) desultory field work
there were none. Indeed it was questionable whether there was any overlap in time. For
example a Holocene date for a tooth of Diprotodon from Orrorroo, in South Australia,
was about 7,000 years old: less frequently reported, by the supporters of recent mega-
faunal extinctions, was a date of greater than 40,000 years for the contents of the
stomach of the same beast (Jones, 1968). Another critical site, for the adherents of early
extinctions, was Lake Mungo. Archaeologists, studying in detail Lake Mungo’s artifacts
and food remains at stratified locales older than 20,000 years, were absolutely unable to
find any extinct species (Bowler et al., 1970).

Yet in spite of its attractiveness an extinction that pre-dated human arrival was not
a view that was strongly argued, largely because it left unanswered the question of what
else might have caused the extinctions. So a different explanation came into favour.
Rhys Jones (influenced by Paul Martin, that doyen of the American debate about ex-
tinctions) provided us with a respectable hypothesis of human causation. The argument
that gained strength in the 1960s was an extinction caused by the initial human settle-
ment of Australia towards the end of the Pleistocene (Jones, 1968). It was thought to be
sudden and ecologically catastrophic.

There were several observations that made plausible the hypothesis of extinctions

soon after first human settlement:

1. Species stable since the Tertiary (notably Diprotodon, short-faced macropods
such as Procoptodon and Sthenurus, and the carnivorous phalanger Thylacoleo)
suddenly became extinct at the end of the Pleistocene. There might be room for
argument about how synchronous the extinction of the various species was, but
by the vast scale of Cainozoic chronostratigraphy there could be no doubt that
the extinctions were sudden.

2. The species that became suddenly extinct had survived numerous climatic os-
cillations, of precipitation and temperature, that had earlier taken place in the
2-3 million years of the Pleistocene. They not only survived these oscillations
but, by their fossil distribution across latitudes and coastal-inland gradients,
showed themselves not to be sensitively adapted to specialized environmental
conditions. They were, in other words, species tolerant of diverse environmen-
tal conditions, in the same way that the grey kangaroo is tolerant. The largest
species (those allied to the genera of Diprotodon and Zygomaturus) have been col-
lected as fossils from Papua-New Guinea to Tasmania, and out west to the lakes
of the Lake Eyre basin. Even allowing for diachronic variations in palaeo-
climates it would be hard to argue that Diprotodon was anything but a very toler-
ant animal.

3. While not precisely knowing the habits of the extinct megafauna it is possible,
from a study of the functional anatomy of the teeth, to identify the majority as
browsers. Of especial interest was Diprotodon, evidently a riverine browser.
Thylacoleo was plausibly interpreted in a quite different way, namely as a
leopard-sized carnivore. In these two cases (and in others) species became ex-
tinct in niches that were then left unfilled. The principle of competitive ex-
clusion could not be invoked as a cause of extinction.

4. It was argued that climate could also not be invoked. Palaeoclimatologists the
world over had not detected, towards the end of the Pleistocene, a climatic
event that was novel. True, we saw the severely cold and dry pleniglacial event,

PROC. LINN. SOC. N.S.W., 109 (1), 1986

RICHARD WRIGHT 3

centred on 18,000 years ago, with particular clarity, but we saw it in that way
because it was so recent and had left so complete a record of its effects. However
even a cursory study of older landforms indicated that events just as severe had
been recurring events in the Australian Pleistocene.

5. There was only one discernibly novel event at the end of the Pleistocene in Aus-
tralia, namely the arrival of human beings. The question was whether we were
to see them as ecologically neutral or as analogous to a new disease.

All the considerations outlined above applied to the Americas as well, where
ground sloth and mastodon became extinct at the end of the Pleistocene. Again we know
that it was at that time that humans arrived.

Thus developed the idea of the extinction of a naive fauna faced by a novel predator
in the form of humans. The fauna was thought to be naive in the sense manifested by
animals that live today in areas not previously occupied by humans, such as certain
ocean islands, areas of the Arctic and all of the Antarctic. The absence of humans in
those places has meant that the animals evolved no behaviour specific to humans as
predators.

In 1973 Martin, writing about the Americas, sharpened up the general theory of
human predation to a specific one. He argued that newly-arrived humans, and their
descendants, formed an advancing predatory front. This front had become adapted to
the naive resource of megafauna. Martin argued that the first Americans swept the
Western Hemisphere and killed off the megafauna (the largest species being the most
economical part, for such a human adaptation, of the total mammalian biomass) within
1,000 years (Martin, 1973). This new and extreme theory for the precise mechanism of
extinction fitted with the fact the radiocarbon dates in Tierra del Fuego were as early as
dates from the north of North America, namely around 11,000 years old.

Though not explicitly extended to Australia, this blitzkrieg model attracted Aus-
tralian archaeologists. Here too dates in the south were as early as dates in the north,
and no sites associating artifacts and megafauna, in reliable and repeatedly observable
stratigraphic association, had been found.

It is perhaps not immediately obvious to you why the absence of blitzkrieg sites was
expected by the proponents of this model for megafaunal extinctions. The answer is
(and it is a reasonable one, even though it might be thought to retain overtones of special
pleading) that in terms of the probability of discovery we cannot expect to find the short-
lived blitzkrieg event materialized in the archaeological record. After all, and on a vastly
broader timescale, millions of individual Diprotodon must have lived in New South Wales
alone, yet the specimen from Tambar Springs is the only individual for which we have
more than half the bones in the skeleton. Indeed we can note in passing that when the
blitzkrieg argument was being put together we did not have even the Tambar Springs
specimen. This marvellously unique skeleton, that is now on display in the Australian
Museum, was found only in 1979. The argument in brief is that since we have so few
fossils, compared with the individuals that lived, why should we expect to be fortunate
enough to find those dating from the short-lived blitzkrieg. The implications of the blitz-
krieg model for field research are depressing.

My field research into the question of megafaunal extinctions did not start until
1974, well after the opposing armies of climatic and humanly-induced extinctions had
taken to the field. Had I earlier joined the theoretical fray I have no doubt that I would
have favoured the arguments for humanly-induced extinction. I was certainly attracted
by the bold explanatory power of the blitzkrieg model.

I now have to stress that, regardless of its theoretical appeal, the blitzkrieg model
for Pleistocene extinctions is seriously threatened by the stratigraphic evidence

PROC. LINN. SOC. N.S.W., 109 (1), 1986

EXTINCTION OF THE AUSTRALIAN MEGAFAUNA

emerging from the archaeological sites that 1 am going to make the subject of the rest of
this lecture.

In order to set this current field work in context I shall briefly describe some earlier
excavations carried out, by myself and several colleagues, near Melbourne in the 1970s
(Gillespie et al., 1978). At the site of Lancefield we were able to date the megafauna to
26,000 years ago. This is a date not for extinction itself, but rather for a still-extant
megafauna. Extinction in Victoria took place at some still unknown date after the
deposits at Lancefield were laid down.

Now a date of 26,000 years ago is a suspiciously late date if the blitzkrieg model is to
hold. We knew before we dated Lancefield that human occupation at Lake Mungo was
at least 34,000 years old. Furthermore after we published Lancefield we were to learn
that there is, for the Western Austraiian site of Upper Swan, reliable evidence for
humans at, or earlier than, 40,000 years ago (Pearce and Barbetti, 1981). By ‘reliable’ I
mean stones that are irrefutably artifacts, 7m sztu, and with repeated radiocarbon dates.

In brief, then, a period of several thousand years of overlap between humans and
megafauna was fatal for the blitzkrieg model. Yet the implications of Lancefield went
further than the mere overlap of radiocarbon dates with archaeological sites elsewhere.
We found at Lancefield two stone artifacts that could be stratigraphically related to the
bone bed — one, under conditions of impeccable control, was found in a little channel
under the bone bed and was therefore earlier than an occurrence of megafauna on
grounds of relative dating alone.

Though it was (and still is) a site that seriously challenges the blitzkrieg model,
Lancefield had a severe deficiency. It had no extended stratified sequence in which arti-
facts and megafauna could be found in association in successive layers. The deposits at
Lancefield happened during a short period around 26,000 years ago when the spring-fed
swampy depression was filled with shallow water. The depression filled up and, as the
pollen record shows, no more free water was available. There was therefore no attraction
for animals, or rather there were no depositional processes that preserved their visita-
tions. So the archaeologist at Lancefield excavates down through two metres of recent
deposits of sandy clay, across an unconformity and into the bone bed. Importantly the
charcoal used for dating was recovered from just under the undisturbed bone bed, with
(as I have already mentioned) an artifact. So in spite of the absence of a continuous
association I found the stratification beyond reproach as regards the late date for
megafauna. Furthermore the two dates of 26,000 years were obtained from separate
parts of the site. Nevertheless we did depend entirely on the reliability of the two radio-
carbon dates for the recent age of the megafauna, and we had no continuous sequence,
associating humans and megafauna, that could be dated from top to bottom and pro-
vide some chronological history of the asscciation.

One might have supposed that a site with better stratigraphic properties than
Lancefield would have been easy to find. Yet it has been the case that since we published
our findings the discovery of a stratified sequence on the mainland of Australia has
proved elusive, but, if the dates from Lancefield are correct, the discovery of such a
stratified sequence is to be expected.

As an improvement on our work at Lancefield our recent work on the Liverpool
Plains of New South Wales looks encouraging. It is to this work that I shall now turn.
The Tambar Springs Project, as it is informally called, has for the first time provided
direct stratigraphic evidence for a prolonged and continuous overlap of humans and
megafauna at a single site. Thereby the evidence from Lancefield has been markedly
improved upon.

In providing you with an account of the Tambar Springs Project I stress that there
are many loose ends to the work so far done. This lecture is therefore a report on work in

PROC. LINN. SOC. N.S.W., 109 (1), 1986

RICHARD WRIGHT 5

progress. It is also a report on work done in collaboration with others, currently David
Horton (of the Australian Institute of Aboriginal Studies) and Judith Fethney of the
University of Sydney. Students of the University of Sydney have provided the labour
that these labour-intensive sites require. The work has been supported in part by the
Australian Research Grants Scheme.

We have been excavating at two sites on the Liverpool Plains, called Lime Springs
and Trinkey. Both sites are spring-fed swamps, unimpressive depressions in the land-
scape. Each is less than 100m in diameter. Neither is identifiable as a site on aerial pho-
tographs, yet their unimpressive appearance belies their scientific importance. We
discovered both sites by ground survey, and into the banks of the swampy depressions we
dug blind, since no archaeological evidence outcrops on the surface.

The sites are 3km apart. Both have essentially the same stratigraphy. The pre-
historic stratigraphic units, from the top down, are: —

1. The Grey Silt, an aeolian dust dating to 6,000 years old: up to 0.8m thick.

2. The Black Swamp, an organic-rich black sandy clay dating from 6,000-c.

20,000 years old: up to 1.3m thick.

3. The Buff Silt, an aeolian dust c. 26,000 years old: up to 0.5m thick.

In substantiating a recent date of megafaunal extinctions it is the Black Swamp that
holds the clues. Wherever we have dug, and at both sites, the Black Swamp contains
stone artifacts and megafauna. All the mammalian remains (both extinct and extant
species) have been identified from fragments of the enamel of their teeth. Bits of bone
are fragmented, and nearly one third are burned, suggesting that they are the remains
of human activity around the spring-fed swamps.

At Lime Springs (by far the richer of the two sites) extinct species present include
Diprotodon, Macropus titan, Protemnodon, Procoptodon and Sthenurus (Gorecki et al., 1984:118).
In the same levels we found thousands of artifacts, including 1988 that were greater than
10mm in minimum dimension. From the point of view of the age of extinctions it is criti-
cal to note that the upper levels of the Black Swamp at Lime Springs (dated by analogy
with Trinkey to 6,000 years old) have as much extinct fauna as the lower levels.

When we published the site of Lime Springs we had only one date, and that was for
the base of the Black Swamp unit. Now we have several dates and they are all consistent.

We have the remarkable evidence of the megafauna living through to the Holocene.
As my late mentor Louis Leakey used to say, the textbooks will have to be rewritten. In-
deed a Holocene megafauna is not the only remarkable attribute of the Black Swamp
unit at both sites, since we also find high groundwater discharge from the springs
through the pleniglacial period centred on 18,000 years ago and which, to the south and
west of the Liverpool Plains, has been shown to be exceptionally arid. Another unex-
pected discovery is cultural: at the top of the Black Swamp (and therefore dating to
about 6,000 years ago) we find the first occurrence at our sites of the horsehoof cores of
the Kartan industry. This industry (previously undated with any precision) is found in
arid areas to the west; the Liverpool Plains represent the most easterly occurrence of this
fascinating prehistoric industry (Lampert, 1983). I will comment again on the sig-
nificance of the Kartan industry when I describe the uppermost unit called the Grey
Silt.

The sandy clays of the Black Swamp unit, though below the water-table, retain
well-differentiated cultural stratigraphy. We were able to use correspondence factor
analysis of the excavated units, taking as data the counts of the rock types used to make
artifacts, to demonstrate that the site was not disturbed (Gorecki et al., 1981:119). Since,
in addition, we cannot differentiate the state of preservation of extant and extinct species
we have very strong prima facie evidence for a Holocene megafauna on the Liverpool
Plains.

PROC. LINN. SOC. N.S.W., 109 (1), 1986

EXTINCTION OF THE AUSTRALIAN MEGAFAUNA

These two sites are better than Lancefield for the simple reason that each has a
sequence, covering some 14,000 years, during which artifacts and megafauna are
associated. For the first time claims for coexistence do not depend on casual and unre-
peatable observation. It is possible to invite a sceptic to excavate a square metre and
guarantee that the association will reveal itself again. Repeatability is one of the essen-
tials of good science.

It is important to note that our findings, surprising though they are, do not fly in
the face of findings from other archaeological sites on the central and western slopes.
There are no other sites yet excavated. rue, extinctions in the fragile ecosystems of the
lakes in western New South Wales were complete by 27,000 years ago (Hope é¢ al., 1983).
Yet we must remember that our area is (today at any rate) so well-watered as not to be so
easily disturbed by climate or humans. Let us consider an environmental parallel.
Giraffes and rhinoceroses went extinct in the Sahara following minor climatic changes,
but flourish to this day in the well-watered Serengeti. We do not think of African en-
vironments in a monistic way, yet we tend to fall victim to the cliche of Australia being
the driest continent. The driest continent it may well be, but this statement of average
overlooks the hundreds of thousands of fertile square kilometres in the well-watered east.
My point is that the fragile ecosystems to the west give us no reason to predict synchro-
nicity of extinctions in the different ecosystems of the east. The only theory that requires
synchronicity is blitzkrieg: Lancefield, Lime Springs and Trinkey indicate that
continent-wide extinctions are not attributable to initial human impact.

RAIN mm.

10! 102 102 104 10°
RETURN INTERVAL - YEARS

Fig. 1. Return intervals of droughts, computed from the rain falling in Year J + Year J-1. Gunnedah, on the
Liverpool Plains is proofed against drought in a way that Menindee, representing the western lakes of New
South Wales, is not.

We published (Gorecki et al., 1984:117) a graph showing the theoretical importance,
when studying extinctions, of summer rainfall on the Liverpool Plains. Fig. 1 extends
this line of thinking into the statistics of the return intervals of droughts. The graph is
calculated from the rainfall statistics of the last hundred years (in lieu of statistics from
the Pleistocene!). The algorithms for computing return intervals are taken from

PROC. LINN. SOC. N.SW., 109 (1), 1986

RICHARD WRIGHT 7

Gumbel (1958). ‘To make the statistics more ecologically meaningful I have computed
the return intervals not for annual rainfall but for the rainfall of Year J + Year J-1. ‘hus
we are looking at the return intervals of droughts calculated from the sum of two year’s
rainfall.

How can we make use of Fig. 1 to illustrate the stability of the Liverpool Plains? Let
us suppose that we assume a total of only 200mm of rainfall, spread over two years, as
locally catastrophic to large mammals and hunter-gatherer humans in any environ-
ment. Fig. 1 indicates that the average return interval of such an amount at Menindee is
in the order of once every one hundred years. At Gunnedah the return interval exceeds
one hundred thousand years. Thus we can see that the Liverpool Plains not only have
better absolute rainfall than the western lakes (including a critical peak of summer rain-
fall) but they are far better proofed against crippling droughts.

The enduring stability of the Black Swamp unit was destroyed 6,000 years ago
when the Grey Silt was blown in. This remarkable unit is best represented at Trinkey,
where nearly a metre of calcareous aeolian dust sits conformably on the Black Swamp
unit. At Lime Springs, where the silt lies below the water-table, it has evidently been
decalcified so that only the siliceous component remains. The deposition of Grey Silt
represents a rude interruption to the swampy ecosystem since no more swamp deposits
formed until post-European times, when the clearing of the land caused a rise in the
water-table.

This calcareous dust is foreign to the area. Certainly it is not a source-bordering
dune, since weathering products of the Pilliga Sandstone surround the two swamps.
Moreover the Grey Silt is found at its deepest within the Trinkey depression and to the
west, not as a dune on the eastern margin.

The theory that I am working on is that the Grey Silt was blown off fluviatile sedi-
ments along the Darling River and was trapped by the dense vegetation (such as the
reed Phragmites) growing in the swamps.

Having been so trapped the calcareous material in the dust was in part eluviated
down the profile of the Grey Silt. At its base, and in the top of the underlying unit of
Black Swamp, calcareous nodules have formed — chemically aiding the preservation of
bones and teeth in what would otherwise have been an acid black sandy clay.

Whatever the final explanation for this extraordinary dust may be, it is clearly a
rare event. The underlying deposits of the Black Swamp show no sign of dust. In scale it
is an event not to be compared with the trivial dust storms of today. The absence of soils
developed within it suggests that it was a single event. If it was blown from the Darling
River it perhaps represents either the onset of an arid period, with destabilization of
alluvial materials previously stabilized by wet-loving vegetation, or it represents the oc-
currence of a flood of rare magnitude followed by a rare hot northwesterly wind.

That there occur floods with a magnitude vastly exceeding anything we have seen
in historic times is implicit in Fig. 2. Again I have used Gumbel statistics, this time to es-
timate the return intervals for floods, of specified discharges, on the Darling at Bourke.
The mind boggles at the implications, for deposits of alluvium on the flood plain of the
Darling, of the flood with a return interval of 1,000 years.

Detailed sedimentological work must now be done on the Grey Silt and an attempt
made to estimate its source and climatological significance. In many ways I would like to
be able to interpret it as an arid event, because we could then link the aridity with the
arrival of the Kartan industry — aridity pushing to the east people whose industrial tra-
ditions had, for some thousands of years, been stably settled well to the west.

Some answers to the environmental implications of the Grey Silt may come from a
study of pollen spectra immediately before, during and after its deposition. Alas, no
swamp deposits formed after its deposition at Lime Springs and Trinkey. Moreover

PROC. LINN. SOC. N.S.W., 109 (1), 1986

EXTINCTION OF THE AUSTRALIAN MEGAFAUNA

MAX. DISCHARGE - 100k megalifres

2. el Oo aad 2 3 i
REDUCED VARIATE - y

(an 7 4 a
2 10 100

RETURN INTERVAL - YEARS

Fig. 2. Return intervals of floods on the Darling at Bourke. Note the implications, in the steepness of the line,
for the magnitude of the thousand year flood. See text for relevance of flooding to origins of the aeolian Grey
Silt unit at the site of Trinkey.

there is no pollen in the underlying Black Swamp unit. Pollen is to be expected in such
organic-rich deposits, so evidently it was destroyed when the swamps dried 6,000 years
ago, a process of destruction that would have been abetted by the calcareous contribu-
tion from the Grey Silt. I regard it as a primary objective of our field work to find a
swamp that has remained damp and thereby preserved pollen from these critical pre-
historic periods.

Meanwhile it is indeed tempting to rush in and explain the extinction of
megafauna, on the Liverpool Plains, as ‘caused’ by the Grey Silt. However at Trinkey,
underlying the Black Swamp, we have another dust, called the Buff Silt and tentatively
dated to 26,000 years. The megafauna clearly survived this event — not only its environ-
mental effects (whatever they may have been) but also the co-occurrence of humans at
that time. For we have undoubted artifacts in the Buff Silt at Trinkey.

We can now seen an unexpected twist emerging from these two sites. Not only have
we threatened the blitzkrieg hypothesis, but also the climatic explanation for extinction.
For how can one appeal to an arid event at 6,000 years ago when an earlier event has sur-
vived unscathed? Moreover we have also threatened any hybrid model, which seeks to
explain extinctions as a bit of climate with a bit of human interference, for humans were
also on the spot when the dust blew 26,000 years ago.

Does all the work we have done leave us with no explanation for extinctions? Yes,
for the Liverpool Plains. Certainly a crude explanation (in terms of climate, humans or

PROC. LINN. SOC. N.S.W., 109 (1), 1986

RICHARD WRIGHT 9

both combined) will not work. I will therefore leave you with some speculation. Let us
remember the important theoretical point, put forward by the proponents of humanly-
induced extinction, whereby they ask what was new at the end of the Pleistocene. For the
Liverpool Plains we must ask what was new after 6,000 years.

Two answers emerge. The first is the small-tool tradition, reasonably interpreted as
an improved hunting technology. The other is the appearance of the dingo. Both events
are thought to have happened around 5,000 years ago. It may well be hard to bring these
events into archaeological focus and to relate them, in a mechanistic sense, to the
process of megafaunal extinctions. Yet, over the next decade we shall seek to do just this
in our field work.

As an audience you represent various disciplines in the natural sciences. I invite
you to visit the critical sections we are establishing at our archaeological sites and to cast
a constructive, if sceptical eye, over the diverse sorts of evidence we are uncovering. The
question to which we are seeking answers is not scientifically trivial and certainly needs
the attention of diverse experts. The cause of megafaunal extinctions is one of the most
challenging ecological questions facing science in Australia.

References

BOWLER, J. M., JONES, R., ALLEN, H., and THORNE, A. G., 1970. — Pleistocene human remains from Aus-
tralia: A living site and human cremation from Lake Mungo, western New South Wales. World
Archaeology 2: 39-60.

GILLESPIE, R., HORTON, D. R., LADD, P., MACUMBER, P. G., RICH, I. H., THORNE, R., and WRIGHT,
R. V. S., 1978. — Lancefield Swamp and the extinction of the Australian megafauna. Science 200:
1044-1048.

GoRECKI, P. P., HorTON, D. R., STERN, N., and WRIGHT, R. V. S., 1984. — Coexistence of humans and
megafauna in Australia: improved stratified evidence. Archaeology in Oceania 19: 117-119.

GuMBEL, E. J., 1958. — Statistics of Extremes. New York: Columbia University Press.

Hope, J. H., DARE-EDWARDS, A., MCINTYRE, M., 1983. — Middens and megafauna: stratigraphy and dat-
ing of Lake Tandou lunette, western New South Wales. Archaeology in Oceania 18: 45-52.

JONES, R., 1968. — The geographical background to the arrival of main in Australia and Tasmania. Archae-
ology and Physical Anthropology in Oceania 3: 186-215.

Manrr«in, P. S., 1973. — The discovery of America. Science 179: 969-974.

PEARCE, R. H., and BARBETTI, M., 1981. — A 38,000-year-old archaeological site at Upper Swan, Western
Australia. Archaeology in Oceania 16: 173-178.

PROC. LINN. SOC. N.S.W., 109 (1), 1986

Aquatic Angiosperms in Coastal Saline Lagoons
of New South Wales.
I. The Vegetation of Lake Macquarie

R. J. KING

KING, R. J. Aquatic angiosperms in coastal saline lagoons of New South Wales. I. The
vegetation of Lake Macquarie. Proc. Linn. Soc. N.S.W. 109 (1), 1986: 11-23.

The distribution and relative abundance of the four aquatic angiosperms in Lake
Macquarie have been mapped. The seagrasses occupied an estimated 14.17km? or
12.3% of the lake surface area with Zostera capricorni (11.57km?2) and Halophila ovalis
(5.59km2) most widespread. There were some 2.01km? of Posidonia australis in the cen-
tral lake region nearest the ocean entrance. The halotolerant Ruppia megacarpa was
found in small amounts, and only in Chain Valley Bay.

R. J. King, School of Botany, University of New South Wales, Kensington, Australia 2033;
manuscript received 13 November 1985, accepted for publication 23 July 1986.

INTRODUCTION

Barrier estuaries or estuarine lagoons are characteristic of much of the south-east
Australian coast (Barnes, 1980). These barrier estuaries range in size from large estu-
aries such as Wallis Lake and Lake Macquarie, which are over 100km2, to small estu-
aries on the south coast e.g. Wallaga, Merimbula and Pambula Lakes. All are
characterized by narrow, elongated entrance channels with broad tidal and backbarrier
sand flats (Roy, 1984). Away from these active channels the lakes are shallow, low-energy
environments with the margins often densely covered by seagrasses.

Three of these barrier estuary systems (Lake Illawarra just south of Wollongong
and Lake Macquarie and Tuggerah Lakes on the N.S.W. Central Coast) are of special
interest. These lakes all support some commercial fishing and prawning, but are in
areas of rapid urbanization with increasing recreational use. Proximity to major popu-
lation centres and coal deposits has made these lakes suitable sites for steam-generating
power stations (Fig. 1). All of these power stations use steam-driven generating units and
lake water is continuously drawn from an inlet channel to cool the condensers.

This paper is part of a series reporting the results of a long-term study on the
aquatic vegetation in the Central Coast lagoons. An introductory paper on Zostera
capricornt in Illawarra Lake has already been published (Harris et.al., 1980). The broad
aim of these studies is to provide detailed qualitative and quantitative data on the
aquatic vegetation in these lakes, to monitor long term changes, and to assess the effects
of cooling water discharges from the power stations.

LAKE MACQUARIE

Lake Macquarie (Fig. 1) is a large barrier estuary some 90km north of Sydney, and
just south of the industrial city of Newcastle. The lake has a surface area of 110km? and
an irregular foreshore of some 166km. It extends 22km in a north south direction and
the maximum width is 9km. The easterly projection of Wangi-Wangi Point, and the
western sandy shallows from Swansea divide the lake into two natural parts. The lake
was formed by the inundation of coalescing river valleys, but the ocean entrance which
was initially much wider is now narrow and shallow due to sand deposition (Baas Beck-
ing et al., 1959). Roy (1984) considers that the lake represents a youthful stage of a barrier
estuary with characteristic rocky and highly irregular shoreline. The average depth of

PROC. LINN. SOC. N.S.W., 109 (1), 1986

THE VEGETATION OF LAKE MACQUARIE

Crowdace
Bay

OWER. STATION NX a
C Fishin
POWER STATION 330MW Sy Station Pt

264 0MW NY SSH wangi Wangi Bay ~

\S< Ba SS < Wangi N
\Y Lake \\ Achy ; i BN \

Eraring \ t
Dora LLY Shingle Swanse:
Creek OS & Splitters Pulbah f
s¢ Bonnells N. t island
Y « Bay . Youth
\ \\ Point Cc
\N ) Wolstoncroft
A
Buffy, iS
Spat Yc
\ NS
Pacific
SPt vales Pt Frying ee .
iw ASSESS Ocean

\
\,

MAY
\ \ SX

Fig. 1. Lake Macquarie, showing localities mentioned in the text and position of power stations.

the lake is 6.7m with a maximum depth of 11m east of Pulbah Island. At the entrance the
spring tidal range is 1.25m but at the western end of the entrance channel it is only
0.15m. Average tidal range in the lake is only 6mm (State Pollution Control

PROC. LINN. SOC. N.S.W., 109 (1), 1986

R. J. KING 13

Commission, N.S.W., 1983) and wind-induced currents are more important in produc-
ing water level changes (Roy and Peat, 1975). It is estimated that only one per cent of the
lake volume is exchanged on each tidal cycle (State Pollution Control Commission,
N.S.W., 1983) and salinity levels are thus highly dependent on catchment run-off and the
flushing action of storm flows. Salinity is usually in the range 25-33.

PREVIOUS STUDIES

The seagrass communities of Lake Macquarie were first discussed in relation to an
alleged depletion of fish (Wood, 1959a; MacIntyre, 1959). The former dealt with the
seagrasses in only two pages and included a single map (scale approx. 1:180,000) show-
ing the distribution of seagrasses within the entire lake. The latter made some cursory
comments on seagrasses in relation to the benthic macrofauna. Since that time there
have been a number of unpublished reports dealing with the seagrass distribution in
various parts of Lake Macquarie, and often with specific reference to faunal community
structure (State Pollution Control Commission, N.S.W., 1983). The only publications
since Wood (1959a) surveying the total lake are those of Evans and Gibbs (1981) and
West e¢ al. (1985). Both surveys were based on aerial photographs.

The aim of the present study is to provide a detailed set of maps showing the
present (1985) distribution and composition of the seagrass beds within Lake Macquarie
with an indication of the relative contribution of the various species.

METHODS

The lake was surveyed during late February/early March (1985). Observations on
the extent, pattern of cover and species composition of seagrasses were made by use of
transects run perpendicular to the shoreline at intervals of 50-200 metres, depending on
the uniformity of the vegetation. The transects were extended until the deepest limit of
the vegetation was reached, or in some cases were continued to the opposite shoreline.
The methods of observation included direct viewing from a boat, raking with a long-
handled rake, snorkelling, and wading in very shallow areas. The methods used at any
site depended on turbidity and depth. The distance from the shore was measured using
a rangefinder [Rangematic MK5: Ranging Inc. for distances over 50m (accuracy 99%
at 100m, 95% at 500m, 90% at 1000m) and a Ranging Optimeter 620 for shorter
distances].

The distribution of seagrasses was plotted on maps at an initial scale of 1:25,000.
Two subjective scales were used: a scale of abundance and a measure of growth or socia-
bility. Each scale has three categories:

Abundance 1. Sparse growth (< 15% cover); 2. Moderate growth (15-50%); 3. Abundant
growth (> 50%).

Sociability. a. individual strands or clumps; b. patches of growth up to 10m; c. beds of
relatively even distribution.

Thus the designation of a weed bed as ZlaH2c indicates a mixed bed of Zostera
(sparse in individual strands or clumps) and Halophila (moderate growth and relatively
evenly distributed). The area covered by weed beds was measured from enlarged copies
of the maps using an Apple 2E microcomputer coupled to a graphics tablet.

RESULTS AND DISCUSSION

This paper provides the basic distribution of seagrasses in Lake Macquarie in
Summer 1985.

PROC. LINN. SOC. N.S.W., 109 (1), 1986

THE VEGETATION OF LAKE MACQUARIE

wo ya eLZ

o (Sdujow) 3}e9S

qzHg7z >
0001 00S 0
%ELH9 2Z

wo?
W06
G2delz

WOL

ez—¥
WOE—Q

Id
47d92Z N woz Wwo9 epuoos

eS ws
= Sh ss 21H 412 \ aH A __ =yeiano

aN LHa2zz PLHELZ QZzHa1z AS qzHazz

woe \ ye1UI hog
eLHqz@Z—-* oUuNAW woz)

ELHELZ
SESS 191100 SS
®LH97Z__ 49 SS QYy

SS
qzHq2zz <S Sg zH922
U0!I}e}G 18M0d I6ueM eLHIEZ

uoljeyg Burysiy 40

o
\doowet ¢=08

S SUSAN
NAN a

PLHGZ

Posidonia

_F
Ruppia Yy

Halophila ee

Fig. 2. Distribution of seagrasses in Lake Macquarie: Fishing Station Point to Rocky Point, western margin of
and sociability see text.

lake.
PROC. LINN. SOC. N.S.W., 109 (1), 1986

For key to abundance

Key to Figs 2-8:

“J
Zostera SS \

R. J. KING 15

The results of the main survey are given in Figs 2-8. A full account of the survey is
available in King (1986). The total area occupied by seagrass was 14.7km?. The area
occupied by each species, is presented in Table 1 along with information for the major
regions and bays in the lake.

The magnitude and predictability of seasonal change and differences from year to
year are discussed in King and Hodgson (1986). The data give some indication of the
relative abundance of each species. For Zostera capricorn: this information can be con-
verted to estimates of total and above ground biomass (King and Barclay, 1986).

TABLE 1
Areas (km?) of seagrasses in Lake Macquarie, survey of January-February 1985

Reference
figure Zostera Halophila Posidonia Ruppia ‘Total

Lake Macquarie 11.57 5.59 2.01 0.15 14.17
(all areas)!

Myuna Bay 2 0.25 0.29 — — 0.29
(Eraring Outlet — Goonda Point)

Goonda — Fishing Station Point 2 0.47 0.26 0.01 — 0.48

Northern Lake 3 1.30 0.46 = _ 1.58
(Fishing Station Point — Cardiff Point) &4

Central eastern 3 3.69 0.52 1.67 — 5.10
(Cardiff Point — “Youth Camp’)

Crangan Bay 5 1.05 0.31 0.02 = 1.07
(‘Youth Camp’ — Pt Wolstoncroft

Chain Valley Bay 7 1.21 1.22 — 0.15 1.25
(Frying Pan Point — Vales Pt)

Wyee Bay inc. 7 0.16 0.83 — — 0.83
Mannering Bay

Wyee Point — Bluff Pt 7 0.62 0.60 = = 0.74

Bonnells Bay 8 2.00 1.08 = = 2.00
(Shingle Splitters Point — Rocky Point)

1total includes several small areas not listed separately

There are three major species of seagrass in Lake Macquarie:
Zostera capricornt Ascherson is the only species of Zostera present. References by Wood
(1959a) to Z. muellert in Lake Macquarie are incorrect and result from the fact that
Wood (1959b) did not appreciate the wide phenotypic variation which occurs in Z.
capricornt (see Robertson, 1984).
Halophila ovalis (R. Brown) Hooker f.
Posidonia australis Hooker f.
In addition, the salt-tolerant genus Ruppia has been recorded.

Following Jacobs and Brock (1982) this material is referred to R. megacarpa Mason.

Zostera capricorni was the most widely distributed and abundant of the seagrasses in Lake
Macquarie. It covered 11.57km? or 10 per cent of the lake area either in monospecific
stands, or mixed with other seagrasses. Halophila ovalis occurred in 25% of the area of
Zostera. The largest Zostera beds are in Belmont Bay and the areas just south of the en-
trance. Only 12 per cent of Zostera occurs in the northern half of the lake. The values

PROC. LINN. SOC. N.S.W., 109 (1), 1986

THE VEGETATION OF LAKE MACQUARIE

BELMONT
BAY

Fishing
Station Pt

ISS

SURE TSINS
RESUS SNS RN

(ul

KF
\}

Scale (Km)

Se ——— or
wassese— See,

Sos
fo.

—.
Lip, ——s.
Os
3

Lake Entrance
Swansea

Youth Camp

Fig. 3. Distribution of seagrasses in Lake Macquarie: central region of the lake: western margin (Dewey Point
to Fishing Station Point); eastern margin (Cardiff Point to ‘Youth Camp’). For key see legend to Fig. 2.

obtained in this study are comparable with data from West et al. (1985) and Evans and
Gibbs (1981) (Table 2). In the latter study the total for all seagrasses was greater than in
the present study but the area occupied by Zostera was estimated to be 12.24km‘’, cf.
11.57km? here. |

PROC. LINN. SOC. N.S.W., 109 (1), 1986

R. J. KING 17

TABLE 2

Comparison of the areas of seagrasses in Lake Macquarte with values recorded tn published surveys

Area of seagrass

Reference Time of survey (ha) Percentage of lake area!
Wood (1959a) August 1953 2331 20
MacIntyre (1959) August 1953 2548 22
Evans and Gibbs Aerial photographs:

(1981)2 Nov., 1971; May, 1975 1788 15.5
Field surveys:
May, 1978
West et al. Aerial photographs:
(1985) June, 1981 and
March, 1982
(West, pers. comm) 1339 11.6
Present study? Feb-March (1985) 1417 1263
Notes

1 Various estimates of lake area are available. Values calculated here are based on a mapped water area of
11511ha (West et al., 1985).

2 Areas quoted for Lake Macquarie in Evans and Gibbs need to be increased by a factor of four (P. Gibbs;
pers. comm). This correction has been applied to figures in the table above.

3 Lake Eraring was not included in this study but is included in all the other studies.

Halophila ovalis occurred over an area of 5.59km? often intermixed with Zostera. It was
most common in shallow bays and especially in Mannering Bay and the southern end of
Wyee Bay where it formed substantial monospecific beds. Evans and Gibbs (1981)
reported only 0.55km? of Halophila in the lake but given the nature of their survey using
aerial photographs it is likely that mixed beds were recorded as Zostera only, since the
Zostera dictates the overall appearance of the beds. Wood (1959a) considered that
Halophila was never in sufficient quantity to make it an important member of the
seagrass community, a conclusion which is contradicted by the results obtained here.

Posidonia australis has a limited distribution and covered an area of only 2.01km?, some
1.67km? on the eastern shore near the lake entrance. This compares with a total area of
0.53km? recorded by Evans and Gibbs (1981). It occurred alone in patches or beds of
relatively even distribution offshore from Zostera and mixed Zostera/Halophila beds, with
which it was sometimes interspersed (Fig. 3). There were small isolated patches of
Posidonia in Crangan Bay just south of Cams Wharf; on the east side of Pulbah Island;
and on the western lake shore of Wangi-Wangi Point.

Ruppia megacarpa was recorded in only 0.1km? in Chain Valley Bay, though Wood (1959a)
showed Ruppia to be the dominant seagrass in backwaters (Mannering Bay and the
southern portion of Chain Valley Bay) as well as the flats to the west of Swansea. The
abundance of Ruppia shows great fluctuations in other coastal saline lagoons such as
Smiths Lake and Tuggerah Lakes and this aspect is discussed in detail in King and
Hodgson (1986).

In addition to aquatic angiosperms, benthic algae, especially Gracilaria verrucosa
(Hudson) Papenfuss, Microdictyon umbilicatum (Velley) Zanardini and Cystophyllum
onustum (Mertens) J. Agardh, may be locally abundant.

PROC. LINN. SOC. N.S.W., 109 (1), 1986

THE VEGETATION OF LAKE MACQUARIE

35m Cockle
Z1bH1a—R¥& Bay

10m

Z1bH1b

Fennel Bay fe ZibHib

6 10m
Ay iN Z2bH2a f 20m Warners
f Omg H

WI —22b

Z2bH1a .
Kooroora ff 25m

Bay }

my g
\ f— 22b
Z2bH1a i} 2bHla Ff 15m
80m ,
Z2b

\
4

4 q 25m
) Broughton}
Point #

Z2b J2H 25m

Scale (Km)

) Cardiff Pt

Fig. 4. Distribution of seagrasses in Lake Macquarie: northern portion of the lake (Dewey Point to Cardiff
Point). For key see legend to Fig. 2.

The method of survey adopted in this study has proved to be very satisfactory for
broad scale vegetation mapping and estimations of abundance. It has several advan-
tages over mapping from aerial photographs:

— the surveys can be undertaken at an appropriate time, e.g. at the time of max1-
mum biomass rather than based on photographs generally taken for some
other purpose;

— species and mixed-species populations can be recognized at the time of survey,
and hence changes within small areas will be recorded;

— measures can be made of species abundance as well as distribution. The
scheme of recognizing three levels of abundance and three levels of sociability
is far simpler than, for instance, a direct Braun-Blanquet type scale of 0-9 of
the type used by Kirkman (1978). This is especially so where comparable
results are required from different field workers.

PROC. LINN. SOC. N.S.W., 109 (1), 1986

R. J. KING

\_—__) Pulbah Island

i Point Wolstoncroft

\
\
A

Z2cH1a

4 so 8 #095

Sa
>
SS

90m

Diamond Drill Point

Fig. 5. Distribution of seagrasses in Lake Macquarie: southeastern section (Crangan Bay). For key see legend

PROC. LINN. SOC. N.S.W., 109 (1), 1986

to Fig. 2.

5 7

Z1bH1b

Co» >

>
>

CRANGAN
BAY

Q
<P>

ZicH1a

on >»

oint Morisset

Ww
=)
(=)

Z3c

Cams Wharf
200m

Z2bH1a
Nords Wharf

100m
100m

Z3bH1a

xX
SSW
NS 3 Z3bH2a

WSs

Browns Pt 200m

g
A
N
Ny
N
\

Scale (metres)

20 THE VEGETATION OF LAKE MACQUARIE

Shingle Splitters
Pulbah Island

Point Wolstoncroft

Bluff Pt

Frying Pan Point

0 al 2
[eee ear eae) Mee |
Scale (Km)

Fig. 6. Distribution of seagrasses in Lake Macquarie: southern region of the Lake (Point Wolstoncroft to Fry-
ing Pan Point and Bluff Point to Shingle Splitters Point). For key see legend to Fig. 2.

— since the accuracy of the final estimates of area occupied by seagrass is ulti-
mately dependent on the scale of the original maps, this type of survey can be
much more accurate. For example reading areas from maps of the scale of that
in Wood (1959a) is impossible, even assuming that the map itself is reasonably
accurate. The problem is compounded by the difficulty of measuring in areas
where the seagrasses occur as narrow fringing beds. The provision of the abso-
lute values for the width of the beds as in the present survey can be useful for
later detailed comparisons in specific areas.

— the scale or intensity of the survey can be readily adapted to suit specific re-
quirements of the user.

PROC. LINN. SOC. N.S.W., 109 (1), 1986

R. J. KING

(S8syaw) ajeIS

Ret PZHICZ

ek | XY
NYY e7HazZ
b uolyeys
BRN 1@MOd

julog seer yy

SASS eee
PESSSSISY

P2H97Z

QZH41LZ
PZHIEZ

Hae QZHOlz
QZHaLz

wozt Keg Aayjean usey
qzueLH gez—tp 2
WO6L RRA QLHQLz

e2HQ2Z
SS WUIOd SPeN
SSN ee
SSS SYS SSI yoelg

SSS
WOEL WOL wo0z wos Ay

&2HIZZ

yulog ued BulAiy

wo#
®&2HQ7Z W06

woe

Aeg B6ulsouuew

&7HG2Z

TPT

LEV,

Q2H42Z

922 wood o¢7
EZRA W®Q
RSQ)
yulod VY
e0hM 4

Ors,

Gre

IF woz wos q
WOLL

QZHatz

Us co

lad.

PZH42Z

ohh.

SS FEI Z

W004

Fig. 7. Distribution of seagrasses in Lake Macquarie: southwestern portion of the lake (Wyee Bay and Chain

Valley Bay). For key see legend to Fig. 2.

The methods adopted for surveys of seagrass beds will depend on the use to which
the data will be put. Data to provide a base-line against which future change might be
measured should be at least as detailed as this method allows. However there are still
considerable problems with interpretation of changes over time (King and Hodgson

PROC. LINN. SOC. N.S.W., 109 (1), 1986

22 THE VEGETATION OF LAKE MACQUARIE

LAKE ERARING

channel markers

Dora Creek

SANS
TS = a

shy

Y Shingle
Splitters
Point

500 1000

Scale (metres)

Fig. 8. Distribution of seagrasses in Lake Macquarie: western portion of the lake, Bonnells Bay (Shingle Split-
ters Point to Rocky Point). For key see legend to Fig. 2.

1986). The major disadvantage of the method used here is that it is time consuming.
Hence it would be inappropriate to the provision of a broad scale inventory of seagrass
resources of the type provided by West et al. (1985).

ACKNOWLEDGEMENTS

This research is part of a long-term project on aquatic vegetation supported by the
Electricity Commission of New South Wales as part of its environmental monitoring
program (Grant B250-429 to Dr R. J. King, University of N.S.W.). I am grateful to
G. C. Coulter (Head, Power Development Division) and Dr B. R. Hodgson (Scientific
Officer) for continued support and help.

I am pleased to acknowledge technical staff involved with this survey, N. Jacobs,
S. McOrrie and L. G. Watson.

During the course of this project I have discussed many of the ideas with colleagues.
I thank them all, particularly Dr P. Farrant, Dr P. Adam, and Dr B. Hodgson.

PROC. LINN. SOC. N.S.W., 109 (1), 1986

R. J. KING 23

References

BaaS BECKING, L. G. M., THOMSON, J. M., and WOOD, E. J. F., 1959. — Some aspects of the ecology of Lake
Macquarie, N.S.W., with regard to an alleged depletion of fish. I. General introduction. Aust. J. mar.
Freshwat. Res. 10: 269-278.

BARNES, R. S. K., 1980. — Coastal Lagoons. Cambridge: Cambridge University Press.

Evans, P., and G1sss, P. J., 1981. — Distribution of seagrass in five N.S.W. coastal lagoons. N.S.W. State Fisheries
Technical Report, 34 pp.

Harris, M. MCD., KING, R. J., and ELLIS, J., 1980. — The eelgrass Zostera capricorni in Illawarra Lake, New
South Wales. Proc. Linn. Soc. N.S.W. 104: 23-33.

Jacoss, S. W. L., and BRocK, M. A., 1982. — A revision of the genus Ruppia (Potamogetonaceae) in Aus-
tralia. Aquat. Bot. 14: 325-337.

KING, R. J., 1986. — Aquatic vegetation surveys, 1980-1986. Unpubl. report: copy held in the Biomedical

Library, University of New South Wales, Kensington.
, and BARCLAY, J. B., 1986. — Aquatic angiosperms in coastal saline lagoons of New South Wales. III.
Quantitative assessment of Zostera capricorni. Proc. Linn. Soc. N.S.W. 109: 41-50.

——, and Hopcson, B. R., 1986. — Aquatic angiosperms in coastal saline lagoons of New South Wales. IV.
Long-term changes. Proc. Linn. Soc. N.S.W. 109: 51-60.

KIRKMAN, H., 1978. — Decline of seagrass in northern areas of Moreton Bay, Queensland. Aquat. Bot. 5:
63-76.

MacINTYRE, R. J., 1959. — Some aspects of the ecology of Lake Macquarie, N.S.W., with regard to an
alleged depletion of fish. VII. The benthic macrofauna. Aust. J. mar. Freshwat. Res. 10: 341-353.
ROBERTSON, E. L., 1984. — Seagrasses. In WOMERSLEY, H. B.S., ed., The Marine Benthic Flora of Southern Aus-

tralia. Part 1. Adelaide: Govt Printer S.A.

Roy, P. S., 1984. — New South Wales estuaries: their origin and evolution. Jn THOM, B. G., ed., Coastal Geo-

morphology in Australia: 91-121. Sydney: Academic Press.

, and Peat, C., 1975. — Bathymetry and bottom sediments of Lake Macquarie. Rec. Geol. Surv. N.S.W.

17: 52-64.

STATE POLLUTION CONTROL Commission, N.S.W., (SPCC), 1983. — Environmental Audit of Lake Macquarie.
Sydney: State Pollution Control Commission. 159 pp.

WEST, R. J., THOROGOOD, C. A., WALFORD, T. R., and WILLIAMS, R. J., 1985. — An estuarine inventory
for New South Wales. N.S.W. Dept Agriculture, Fishertes Bulletin 2.

Woop, E. J. F., 1959a. — Some aspects of the ecology of Lake Macquarie, N.S.W., with regard to an alleged
depletion of fish. VI. Plant communities and their significance. Aust. J. mar. Freshwat. Res. 10: 322-340.

— , 1959b. — Some east Australian seagrass communities. Proc. Linn. Soc. N.S.W. 84: 218-226.

PROC. LINN. SOC. N.S.W., 109 (1), 1986

Oe Ra ak RM ary

aT aN as ies ar Pe ie a ¥

Aquatic Angiosperms in Coastal Saline Lagoons
of New South Wales.
II. The Vegetation of Tuggerah Lakes, with
Specific Comments on the Growth of
Zostera capricornt Ascherson

R. J. KING and V. M. HOLLAND

KING, R. J., & HOLLAND, V. M. Aquatic angiosperms in coastal saline lagoons of New
South Wales. II. The vegetation of Tuggerah Lakes, with specific comments on the
growth of Zostera capricornt Ascherson. Proc. Linn. Soc. N.S.W. 109(1), 1986: 25-39.

The distribution and relative abundance of the three aquatic angiosperms in the
Tuggerah Lakes system have been mapped. The seagrasses occupy an estimated
19.11km? or 25% of the area of the lakes. Halophila ovalis occurs over an area of 10.40km2
and Ruppia megacarpa over 8.24km?. The growth rate of Zostera capricorni was measured
in the field using a hole-punching technique. Growth rates measured over a year ranged
from 0.8mg.shoot!.day? in winter to a summer maximum of 2.6mg.shoot?!.day?.
There were marked seasonal changes in the storage and use of starch in the rhizomes,
from a maximum at the end of summer (226mg.g"dwt of rhizome) to a minimum value
of 20mg.g" at the end of winter.

R. J. King and V. M. Holland, School of Botany, University of New South Wales, Kensington,
Australia 2033; manuscript recewved 13 November 1985, accepted for publication 23 July 1986.

INTRODUCTION

The aquatic vegetation of Tuggerah Lakes was surveyed by Higginson (1965) and
some ecological effects of pollution documented (Higginson, 1971). Since that time there
have been a number of detailed surveys of the aquatic vegetation of the lakes but these
have not been published in full or in generally available literature. A single value for the
area of seagrass beds is given by West e¢ al. (1985).

This paper provides a full survey of the distribution of seagrasses in Tuggerah
Lakes in summer 1985. In addition data are provided on the seasonal aspects of growth
of Zostera capricornt Ascherson which is the most abundant marine angiosperm in the
lakes. Basic data are provided on seasonal variation in shoot length, shoot weight,
reproductive status and rhizome starch reserves. Seasonal change in the rate of growth
of the vegetative shoot was estimated by the rate of production of new leaf tissue.

TUGGERAH LAKES

Tuggerah Lakes, approximately 100km north of Sydney on the east Australian
coast (Fig. 1) are a series of three interconnected coastal lagoons: Lake Munmorah,
7.8km?; Lake Budgewoi, 11.2km?; Tuggerah Lake, 58km? (total area 77km?) (Interdept.
Comm., 1979). West e¢ al. (1985) measured the water surface area at only 70.29km? and
this reflects the shallow topography: a small fall in the lake level exposes much of the lake
bottom especially on the Budgewoi Flats on the eastern margin of Lake Budgewoi. The
lakes extend 16km in a north—south direction and 5km east — west with a perimeter of
approximately 110km. The lakes are a barrier estuarine system at a relatively youthful
stage (Roy, 1984) and are thought to have been formed by long shore currents building a
series of sand bars across an indentation in the coastline. The eastern foreshores are thus
of a sand-dune nature (Bird, 1984). The lakes are shallow, the average depth being

PROC. LINN. SOC. N.S.W., 109 (1), 1986

26 THE VEGETATION OF TUGGERAH LAKES

\ MUNMORAH S
Ry \ POWER <
STATION

Ocolwolrolong Pt

Gorokan

TUGGERAH

Scale (Km)

Fig. 1. Tuggerah Lakes showing localities mentioned in the text, and the principal study site for growth
studies.

1.9m, and the greatest depth 4.9m under the Toukley bridge between Budgewoi and
Tuggerah Lake. The lake system is connected to the sea by a narrow channel at “The
Entrance’ and this connection is only rarely completely closed. Tidal flushing is low,
with an exchange of only about one per cent of lake volume; tidal action within the lakes
is negligible. Salinity is thus dependent on rainfall in the catchment of 670km?

PROC. LINN. SOC. N.S.W., 109 (1), 1986

R. J. KING AND V. M. HOLLAND

(Interdept. Comm., 1979). Salinity is generally in the range 17-28 though salinities as
low as 5 after heavy rainfall, and 49 after drought, have been recorded (Higginson,

1971).

Seagrass distribution survey

The seagrass distribution was surveyed and mapped using the method outlined in
King (1986b), then plotted using the same scale: abundance [1= sparse growth, (<15%
cover); 2= moderate growth (15-50%); and 3= abundant growth (>50%) | and so-
ciability [a= individual strands or clumps; b= patches to 10 m; and c= beds of rela-

tively even distribution].

METHODS

Gorokan

Z1aH1a
Z1bH1a

Z2bH1bR2b

m
Z1bH1bR2b Z1aH3c
Z2bH3c

Ourimbah
Ck

1b
ER ESAS ..
LESS SSS

yy
&

a 600m mares? wae

Z2bH1aR2b Z1aH1a

Z2c H2a
165m

* Z
NS
2 NY Z3bH1aR2b
ee
N 100m
NA

EN
4 ds

Z1aR1a

Z2bH1aR2b
Z23bH2a

7" 250m
Z2bH2b ——

Z2bR2b

R11] Gil RK"? R1b Ish
Z1aH3c WIZ SN At
900m

Z2b

Z1b

TUGGERAH
LAKE

Z2bH2b

NEF é 7

N Z2bR3b

Peli

PACIFIC
OCEAN

"9 OPCS,

thie

5965:

3c } Z1aR3b

The Entrance

310m

0 1 2
ih
Scale (Km)

Z1aR1b

R3b
GZ Z2bH1aR1a

Killarney

Fig. 2. Distribution of seagrasses in Lake Tuggerah.

NW Zostera

For key to abundance and sociability see text.

Halophila

Ruppia

PROC. LINN. SOC. N.S.W,, 109 (1), 1986

28 THE VEGETATION OF TUGGERAH LAKES

Budgewoi
Channel

Scale (metres)

Z1aH1aR1a

Z
4

Z2bH1bR1b

ere i,

ie)
>
WW
o
a
>
a

Gorokan

Z1bH1b R1b

Z3cH2bRi1a
Goobarabah Pt \

Wallarah Ck
Ocolwolrolong

Fig. 3. Distribution of seagrasses in Lake Budgewoi (see legend for Fig. 2).

B. Seasonal studies on Zostera capricorni
Study site:

The principal study site for seasonal growth studies was located in a depth of 0.5m
in the northwestern end of Lake Munmorah. This site was selected as representative of
relatively undisturbed Zostera beds. It was well beyond the thermal influence of the Mun-
morah power station and urban pressure in terms of housing construction, and hence
increase in siltation and nutrients, is low. There is little recreational use of these shallow
flats. Extensive beds of Zostera capricorni up to 500m wide, in some places mixed with
Halophila ovalis, occurred in this area of the lake (Fig. 4).

PROC. LINN. SOC. N.S.W., 109 (1), 1986

R. J. KING AND V. M. HOLLAND 29

1. Leaf growth

Leaf-blade growth rates were estimated using a modification of the leaf-marking
technique of Zieman (1974). In these experiments the datum mark in a shoot was a small
hole punched through the group of leaves using a pair of ‘tongue forceps’, the prong of
which had been ground to a sharp point. The scissor-like instrument minimizes damage
to the leaf blade, which is not crushed. In a preliminary experiment to test the use of this
method, holes were punched along the Zostera leaves at 2cm intervals, the first hole being
about 2mm from the shoot — rhizome junction. The plants were then grown in perspex
tanks in a growth cabinet at 25°C for 7 days. Individual leaves on each shoot were then
examined to determine possible effects of the marking method, elongation between suc-
cessive holes, and the relationship between leaf age and elongation rate.

Growth experiments were carried out at 2-month intervals (Oct. 1978 — Feb. 1979)
and then at monthly intervals from March 1979 to February 1980. Samples of the Zostera
were collected from the study site using a 20cm corer, ‘planted’ in plastic containers,
marked and returned to the field. Plants were then harvested after an interval of 12 days
and the dry weight (constant weight at 105°C) obtained for new tissue and leaf material
above the datum mark. All epiphytes were removed. Growth rates were calculated as in-
crease in biomass per shoot, per day. Each growth experiment was replicated four times.

2. Population data

The following data were obtained in conjunction with the growth experiments:

Shoot length — the length of the longest leaf on each of 50 randomly-selected in-
tact shoots was measured.

Shoot weight — data on shoot weight were obtained directly from the leaf growth
study.

Reproductive shoots — only vegetative shoots were used in the calculation of
growth rates. The frequency of reproductive shoots was estimated
for all core samples.

3. Rhizome starch reserves

Living rhizome segments were collected from September 1978 to February 1980.
Each sample consisted of segments from 10-15 plants. The rhizome segments were
washed free of sediment, the roots and shoots were removed, and the samples were then
frozen and stored at -15°C until extraction. The quantity of starch in the rhizomes was
determined using the extraction and assay procedure of Hassid and Neufeld (1964).

RESULTS

A. Seagrass distribution

Three aquatic angiosperms occurred in the Tuggerah Lakes in sufficient amounts
and widely distributed enough to be mapped at the scale used. These were Zostera
capricornt Ascherson; Halophila ovalis (R. Brown) Hooker f.; and Ruppia megacarpa Mason.
The maps constructed from the vegetation survey are presented as Figs 2-4. A full copy
of the survey is available in King (1986a). The total area occupied by seagrass was
19.11km? and the area occupied by each species is presented in Table 1. Data are given
separately for each lake in the Tuggerah Lakes system, as well as for the Budgewoi Flats
region.

B. Seasonal studies

Preliminary growth studies
In the preliminary experiments on material grown in the growth cabinet all leaf
elongation occurred within 4cm of the shoot base. Over 90% of growth occurred in the

PROC. LINN. SOC. N.S.W., 109 (1), 1986

30 THE VEGETATION OF TUGGERAH LAKES

H1aR2b Z2cH1b
Z2bH2bR1a

Budgewoi
Channel Scale (metres)

Fig. 4. Distribution of seagrasses in Lake Munmorah (see legend for Fig. 2).

TABLE 1
Areas (km2) of seagrasses in Tuggerah Lakes, survey of January-February 1985

Total area
Zostera Halophila Ruppia of seagrass
Tuggerah Lakes 12.26 10.40 8.24 19.11
(all areas)
Tuggerah Lake 9.58 6.43 5.48 12.69
Budgewoi Lake 1.19 2.49 2.22 4.03
Budgewoi Flats 0.59 1.34 1.71 2.63

(within L. Budgewoi)
Lake Munmorah 1.49 1.48 0.54 ° 2.39

PROC. LINN. SOC. N.S.W., 109 (1), 1986

R. J. KING AND V. M. HOLLAND 31

100
80 8
5 ie
a T
> 60 8 6
° ;
o E
= 40 aay
3 6
i} —_—
i°2)
— Cc
° o
ao) 70 Rie
2 AeaBeaC Onli e2unsue4 aS
Region Age Class

Fig. 5. a (left), Elongation between successive holes in Zostera capricorni. A represents the growth below the
datum (0 cm), B (0-2 cm), C (2-4 cm) and D (above 4 cm). b (right), Elongation rates of leaves of different age
classes. The youngest marked leaf is referred to age class 1; leaves arising during the course of the experiment
as age class ‘0’.

region below the first hole (at 2mm) and 99.5% below the 2cm datum mark (Fig. 5a). A
datum mark 2cm from the base was selected therefore for use in all further experiments.
The number of leaves per shoot varied from 3 to 6, and the rate of growth was directly
related to leaf age (Fig. 5b). Leaves which arose during the harvest interval are indicated
as age class ‘0’, and the elongation rate was estimated on the assumption that these new
leaves were present on average for half the experimental growth period (Sand-Jensen,

1975).

Leaf growth

The growth rate of new leaf tissue ranged from 2.6+0.5mg. shoot! .day? in sum-
mer to 0.8mg. shoot?. day? in late autumn (Fig. 6). Leaf tissue production was closely
related to the weight of the shoot. Shoot viability, (i.e. the percentage of shoots alive at
the end of the harvest interval) was generally high (90-100%) except in November and
December 1979 when it was 72 and 57% respectively.

Population data

Shoot length

Mean shoot length was at its lowest in late summer, and it ranged from 22.7 +0.8
cm in March to a winter maximum of 53.9+1.8cm in July (Fig. 7). Analysis of variance
showed that monthly differences were significant (P <0.005) and application of Tukey’s
test (Zar, 1974) showed that a difference in shoot length of 7.3cm was statistically
significant.

Shoot weight
Mean weight of shoots was highest in January 1980 (251 +12mg. shoot’ and lowest
in May 1979 (89+ 8mg. shoot"). The data are plotted in Fig. 8.

PROC. LINN. SOC. N.S.W., 109 (1), 1986

32 THE VEGETATION OF TUGGERAH LAKES

Reproductive shoots

Flowering commenced in October 1978, and in 1979 in early August. Seed set
occurred soon after flowering and in both years reproductive shoots persisted in the
population until December. The density of flowering shoots (shoots.m™) differed
markedly in the two years: in 1978 densities of 188 (Oct) and 113 (Dec) were recorded
and for the months Aug-Dec 1979 densities were 24, 63, 90, 35, and 31 respectively.

Rhizome starch reserves

Starch made up to 22.6% of the dry weight of the rhizome of Zostera capricorni, and
there was a marked seasonal change in the quantity stored (Fig. 9). There was a rapid
build up in summer with a maximum value of 226mg.g dwt? in February 1979 and a
decline in autumn to a minimum of 20mg.g dwt!. The method of starch extraction was
accurate but time consuming and therefore replicate samples were analysed in only four
months. The variability between replicates in these samples was low (coefficients of
variation between 7.1 and 18.2%).

DISCUSSION

A. Seagrass distribution

In this survey seagrasses were recorded as covering 19.11km? or 25% of the area of
Tuggerah Lakes. Zostera capricornt was the most widely distributed seagrass occurring
over 12.26km? or approximately 16% of the lake area. Halophila ovalis occurred over
10.40km? (14%) and Ruppia megacarpa over 8.24km? (11%). The only other recent figure
is that of West et al. (1985) who estimated from aerial photographs that seagrass covered
11.62km? of the whole lake system. This considerably lower area may result from the fact
that Halophila beds were not detected from aerial photographs whereas Zostera which
forms conspicuous beds was. Also West et al. (1985) measured a surface area for the lakes
6.7km? less than that in the Interdept. Comm. (1979). If this represents a real difference
due to the lowering of the water level in the lakes this reduction would be in the marginal
area normally occupied by seagrasses, and particularly the Budgewoi Flats would have
been not included.

The areas of seagrass recorded in this survey are considerably lower than those
found by Higginson (1965; 1968). The areas of seagrass which he recorded in 1963, 1965
and 1966 were 42%, 31% and 28% of the lake area respectively. Not only has the area
occupied by seagrass decreased but also the relative importance of the different species.
Higginson’s maps, based on a survey in May 1963 (Higginson, 1965), showed Lake
Budgewoi almost filled with seagrass (76.2% of lake covered cf. 36% in the present
study). The whole central region was occupied by Ruppia (42% of the lake area). Ruppia
was also dominant in deeper water in the southern part of Lake Tuggerah so that
approximately one-third of the beds were Ruppia-dominated communities. By 1966
there was no Ruppia in the lake system (Higginson, 1968). The results presented here
show that Ruppia had again become established in Tuggerah Lake but it was not found in
deeper water; rather in shallow areas inshore of the Zostera beds. The places where
Ruppia was dominant were west of Chittaway Point and just south of Toukley in Tug-
gerah Lake, and on the Budgewoi Flats. The central portion of each of the three lakes
was devoid of seagrasses. Details of long period changes in the vegetation are discussed
in King and Hodgson (1986).

B. Seasonal Studies
The modification of Zieman’s (1974) method of measuring seagrass leaf growth was
shown to be an appropriate method.

PROC. LINN. SOC. N.SW., 109 (1), 1986

R. J. KING AND V. M. HOLLAND 33

100

% Viability ~—

OND MJJASOND JF

1978 1979 Month 1980

Leaf Growth (mg.shoot-l.day-1) -e--

Fig. 6. Seasonal changes in leaf growth recorded on Zostera capricorn: in the field. Data for early 1979 lost due to
vandalism.

Length (cm)

Shoot

OUD FA J iA 0 DF
1978 Month 1980

Fig. 7. Seasonal variation in mean shoot length of Zostera capricorni. The standard error is shown; the vertical
bar (Tukey’s ‘W’) represents a significant difference in shoot length of 7.3 cm.

In the seagrass genera Thalassia, Posidonia and Zostera the leaf-initiating meristem is
at the base of the shoot, just above the rhizome. The meristem is enclosed and protected
by the older leaves and leaves are produced distichously from the leaf primordium (den
Hartog, 1970). Measurement of leaf growth by marking leaves (Zieman, 1974) assumes
that vegetative growth occurs only at the base of the shoot, below a fixed point — the
datum mark. Growth involves both the production of new cells and their subsequent ex-
pansion, and these two processes are spatially separated. Growth measured as increase
in size or weight is mainly due to the process of cell expansion, which occurs in an area
adjacent to the meristem (Preston, 1974). In Zostera this zone occurs immediately above
the meristem and results principally in elongation, since leaf width changes very little
during growth (Mukai et a/l., 1977). This means that leaf growth can be estimated by

PROC. LINN. SOC. N.S.W., 109 (1), 1986

34 THE VEGETATION OF TUGGERAH LAKES

measuring the single parameter of leaf length, but it is imperative that holes are
punched above the zone of cell expansion.

Preliminary experiments indicated that a point 2cm above the shoot —rhizome
junction was appropriate as a datum as 99.5% of leaf growth elongation occurred below
this point. No deleterious effects due to the hole-punching technique were observed, and
all shoots remained healthy over the seven day harvest interval. The holes, though small,
were easily located in all leaves. There are advantages in using a punched hole rather
than other datum marks such as staples. All leaves can be marked simultaneously, in-
creasing both accuracy and speed. Also the datum mark can be placed below the ligule,
through the sheath. In Zostera the younger leaves are successively enclosed by the leaf
sheaths of older leaves and this restricts alternative types of datum marks to areas above
the sheath (Sand-Jensen, 1975). This creates difficulties because of the need to estimate
the growth of new unmarked leaves.

250

200

Shoot Weight (mg.shoot-1)

ONDJFMAMJJASONDJFMA
1978 1979 month . 1980

Fig. 8. Seasonal variation in the mean shoot weight of Zostera capricorni (+ standard error).

In studies of the leaf growth of Posidonia australis in altered salinity, Tyerman et al.
(1984) showed that damage to the leaf sheath killed or inhibited leaf growth. In the
present experiments the hole made by the tongue forceps is small and the sheath may
reseal. Certainly, there was no visible damage, but it is possible that all new leaf growth
was inhibited.

The comparison of growth rates of Zostera measured here with those measured in
other studies on seagrasses is hampered by the lack of uniformity in the units in which
growth rates are expressed, coupled with the fact that the magnitude of the differences is
a function of the unit in which growth is measured. In Table 2 data from Botany Bay and
Port Hacking are summarized. In so far as generalizations may be drawn from such
data one can conclude that the relative differences in growth rates between summer and
winter are similar at all these study sites and that even though the measures of relative
growth rate are calculated on different bases they are comparable.

The viability of shoots was low during November and December (Fig. 6) and this is
reflected in a reduced growth rate during the period.

The range of shoot lengths (22.7cm, late summer—53.9cm in winter) is compar-
able to that reported by Larkum et al. (1984) for Zostera capricorn in Botany Bay except
that there the pattern is virtually the reverse with a maximum of 45cm in late summer

PROC. LINN. SOC. N.S.W., 109 (1), 1986

R. J. KING AND V. M. HOLLAND 35

and a minimum of 17.2cm in winter. A similar pattern to that found by Larkum e al.
(1984) in Botany Bay was reported by Harris eé¢ al. (1980) for Lake Illawarra, and by
Higginson (1968) for Lake Budgewoi. The length of the longest leaves has been used to
indicate the seasonal growth cycle (Harris et al., 1980) and the marked seasonal die back

TABLE 2
Comparative Data for Zostera capricorni on the New South Wales Coast

Summer Winter

‘Tuggerah Lakes (present study)

Measured shoot growth

mg. shoot?! .day? 2.6 0.8

Calculated growth rate*

g. g! day! 0.0104 0.0089
Botany Bay (Larkum et al., 1984)

Leaf growth

cm. day! 1.62 0.38

Eimojrorotorne! growth rate’

g. g! day? 0.0308 0.0213
Production rate

g. m7 day"! 8.92 1.37

Calculated shoot growth**

mg. shoot! day 4.1 0.8
Port Hacking (Kirkman et al., 1982)

Production rates

g. m}.day?! 2.5 0.3

‘Relative growth rate’

g. g! day"! 0.028 0.014

* rate calculated using mean shoot biomass data for Munmorah Lake.

** rate calculated on the basis of mean shoot density.

which is reported to occur in winter (Wood, 1959b; Larkum et al., 1984). Comparison of
the data on maximum shoot length with the leaf growth rates presented above makes it
clear that in this case there is no relationship between the two. If Zostera capricorni were al-
ways to show the marked seasonal fluctuations reported by Wood (1959b) the relation-
ship might be expected but rapid changes in seagrass beds may be caused by factors
other than normal seasonal ones, e.g. heavy rainfall (Harris et al., 1980), depredation by
swans (Wood, 1959a,b), and wave damage (Kirkman e¢ a/., 1982). These factors may be
more important in shallow protected lakes such as Lake Munmorah than in open bays,
such as Botany Bay. In Lake Illawarra full leaf development has been maintained in
some years into early spring, by which time new leaf growth had commenced. Given this
it would be better to use the leaf length frequency rather than longest leaf length to
establish comparative growth rates (see for example Kirkman et al., 1982).

The data on vegetative shoot weights are not directly comparable with other pub-
lished data where biomass is generally expressed on the basis of area. Conversion of the
data in Larkum et al. (1984) gives values ranging from 36mg. shoot" in winter to 133mg.
shoot! in summer (cf. 89 and 251 in this study). On the basis of average shoot weights
the growth rates in the present studies can be expressed in g.g.!.day! and thus be com-
pared with the proportional growth rate of Larkum et al. (1984). The values obtained are
0.0089 in winter (comparable values for Botany Bay and Port Hacking 0.0213 and 0.008
respectively) and 0.0104 in summer (0.0308 and 0.035). The relative similarity between

PROC. LINN. SOC. N.S.W., 109 (1), 1986

36 THE VEGETATION OF TUGGERAH LAKES

the rates for summer and winter obtained in this study reflects the less marked seasonal
change in above ground biomass compared to that in the other two studies.

Zostera capricornt commences flowering in spring or early summer. Various times for
flowering appear in the literature (November — Wood, 1959b; September — Harris et
al., 1980; Larkum et al., 1984). In 1978 flowering commenced in October but in 1979 it
commenced in early August. The factors initiating flowering have not been investigated
but they are possibly related to temperature. In 1979 winter temperatures were rela-
tively mild with the lowest temperature being 13.8°C in July, whereas Electricity Com-
mission data show a minimum generally in the range 11-12°C. Studies on the northern
hemisphere Zostera marina have implicated temperature in anthesis and seed production
(Setchell, 1929; McRoy, 1970; Churchill and Riner, 1978) though Jacobs (1979) sug-
gested that light intensity may be the controlling factor.

200

100

Rhizome Starch (mg.g-1)

SONDJFMAMJJASONDJF
1978 Month 1979 1980

Fig. 9. Seasonal variation in the quantity of starch in the rhizomes of Zostera capricorn.

The data on the prevalence of flowering are of the same order as those of Larkum et
al. (1984) who recorded up to 312 flowers.m?. The presence of reproductive shoots is
apparently not related to the ‘success’ of Zostera capricorni in the Tuggerah Lakes, since
establishment from seedlings has not been observed.

The principal storage product in seagrasses is starch, and starch grains are promi-
nent in the cortical cells of rhizomes of Zostera capricorni (Fig. 10). Soluble carbohydrates,
principally sucrose, are also found in quantity in both the rhizomes and leaves of
seagrasses. Drew (1980) reported that Zostera capricorni leaves contained 4.7% sucrose,
1.6% myo-inositol and 0.4% fructose while the rhizomes contained 17.1% sucrose.

The extraction method used here (Hassid and Neufeld, 1964) produces a pure
starch fraction. This represents stored reserves which need to be enzymatically mobi-
lized before they are available to the plant. Rapid incorporation of starch occurred in
summer (approx. 1.5-2.0mg.g!.day') leading to a maximum in early autumn after
which the shoot weight began to decline. During autumn and winter the starch level fell
rapidly, presumably due to mobilization and use of the reserves by the plant. This fall in
the quantity of starch occurred at the same time as the seasonal drop in light intensity
and temperature.

PROC. LINN. SOC. N.S.W., 109 (1), 1986

R. J. KING AND V. M. HOLLAND 37

Marked seasonal changes in the quantity of both soluble and insoluble carbo-
hydrates are known especially in alpine plants where high levels are essential to initiate
rapid growth in the spring growing season (Mooney and Billings, 1960; Bannister and
Ward, 1981). The seasonal pattern in aquatic plants differs with low levels of carbo-
hydrates in the rhizomes during winter/spring with rapid incorporation during summer
(Haag and Gorham, 1977; Dawes and Lawrence, 1980; Livingston and Patriquin, 1981).
The reserves accumulated in summer allow the species to survive low winter light inten-
sities (Dawes and Lawrence, 1980). A comparable situation has been demonstrated in
Arctic kelps (Chapman and Lindley, 1980). Defoliation studies with the tropical
seagrass Thalassia testudinum suggested that rhizome reserves also support leaf-blade
regeneration (Dawes and Lawrence, 1979) and Drew (1980) has shown the utilization of
endogenous soluble carbohydrates, especially sucrose, by Zostera angustifolia leaves
starved in the dark.

Fig. 10. Transverse section of the rhizome of Zostera capricorni showing starch grains in the cortical cells.
Material collected late summer, 1985.

ACKNOWLEDGEMENTS

The project was funded by the Electricity Commission of New South Wales (Grant
B250.429 to Dr R. J. King, Univ. of N.S.W.). We are pleased to acknowledge the support
and help of G. C. Coulter, Head, Development Section, Dr B. R. Hodgson (Scientific
Officer), and especially thank Mr W. Jefferson for his willing help with field work. We
thank those technical staff involved in field work in this project: in particular, thanks are
due to B. Kertesz, S. McOrrie and L. Watson.

PROC. LINN. SOC. N.S.W., 109 (1), 1986

38 THE VEGETATION OF TUGGERAH LAKES

References

BANNISTER, P., and WARD, G., 1981. — Changes in non-structural carbohydrate contents of Chionochloa rigida
and C. macra during winter. N.Z. J. Bot. 19: 233-239.

BIRD, E. C. F., 1984. — Coasts. An Introduction to Coastal Geomorphology (3rd edn). Canberra: ANU Press.

CHAPMAN, A. R. O., and LINDLEY, J. E., 1980. — Seasonal growth of Laminaria solidungula in the Canadian
High Arctic in relation to irradiance and nutrient concentrations. Mar. Biol. 57: 1-5.

CHURCHILL, A. C., and RINER, M. I., 1978. — Anthesis and seed production in Zostera marina L. from Great
South Bay, New York, U.S.A. Aquat. Bot. 4: 83-93.

DaweEs, C. J., and LAWRENCE, J. M., 1979. — Effects of blade removal on the proximate composition of the

rhizome of the seagrass Thalassia testudinum Banks ex Konig. Aquat. Bot. 7: 255-266.
, and , 1980. — Seasonal changes in the proximate constituents of the seagrass Thalassia testudinum,
Halodule wrightit, and Syringodium filiforme. Aquat. Bot. 8: 371-380.

DREw, E. A., 1980. — Soluble carbohydrate composition of seagrasses. Jn PHILLIPS, R. C., and McRoy, C.
P., eds, Handbook of Seagrass Biology: An Ecological Perspective. New York: Garland STPM Press.

Haac, R. W., and GORHAM, P. R., 1977. — Effects of thermal effluent on standing crop and net production
of Elodea canadensis and other submerged macrophytes in Lake Wabamum, Alberta. J. Appl. Ecol. 14:
835-851.

Harris, M. MCD., KING, R. J., and ELLIs, J. 1980. — The eelgrass Zostera capricorn: in Illawarra Lake, New
South Wales. Proc. Linn. Soc. N.S.W. 104: 23-33.

HARTOG, C. den., 1970. — The Seagrasses of the World. Amsterdam: North-Holland.

HassIpD, W. Z., and NEUFELD, E. F., 1964. — Quantitative determination of starch in plant tissues. Jn
WHISTLER, R. L., ed., Methods in Carbohydrate Chemistry. London: Academic Press.

HIGGINSON, F. R., 1965. — The distribution of submerged aquatic angiosperms in the Tuggerah Lakes sys-
tem. Proc. Linn. Soc. N.S.W. 90: 328-334.

——., 1968. — The ecology of submerged aquatic angiosperms within the Tuggerah Lakes system. Sydney:
Unviersity of Sydney, Ph.D. thesis, unpubl.

—, 1971. — Ecological effects of pollution in Tuggerah Lakes. Proc. Ecol. Soc. Aust. 5: 143-152.

INTER-DEPARTMENTAL COMMITTEE, 1979. — Tuggerah Lakes Study Report. Sydney: Ministry for Public Works.

Jacoss, R. P. W. M., 1979. — Distribution and aspects of the production and biomass of eelgrass, Zostera
marina L. at Roscoff, France. Aquat. Bot. 7: 151-172.

KING, R. J., 1986a. — Aquatic vegetation surveys, 1980-1986. Copy held in Biomedical Library, University
of New South Wales, Kensington.

——, 1986b. — Aquatic angiosperms in coastal saline lagoons of New South Wales. I. The vegetation of Lake

Macquarie. Proc. Linn. Soc. N.S.W. 109: 11-23.
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Long-term changes. Proc. Linn. Soc. N.S.W. 109: 51-60.

KIRKMAN, H., REID, D. D., and CooK, I. H., 1982. — Biomass and growth of Zostera capricornt Aschers. in
Port Hacking, N.S.W., Australia. Aquat. Bot. 12: 57-67.

LARKuUM, A. W. D., COLLETT, L. C., and WILLIAMS, R. J. 1984. — The standing stock, growth and shoot
production of Zostera capricorni Aschers. in Botany Bay, New South Wales, Australia. Aquat. Bot. 19:
307-327.

LIVINGSTONE, D. C., and PATRIQUIN, D. G., 1981. — Below ground growth of Spartina alterniflora Loisel:
Habit, functional biomass and non-structural carbohydrates. Estuarine, Coastal and Shelf Science 12:
579-587.

McRoy, C. P., 1970. — Standing stocks and other features of eelgrass (Zostera marina) populations on the
coast of Alaska. J. Canad. Fish. Res. Bd 27: 1811-1821.

Mooney, H. A., and BILLINGS, W. D., 1960. — The annual carbohydrate cycle of alpine plants as related to
growth. Am. J. Bot. 47: 594-598.

Mukal, H., Alol, K., KoIKE, I., 11zum1, M., OHTSU, M., and HaTTor1, A., 1979. — Growth and organic
production of eelgrass (Zostera marina L.) in temperate waters of the Pacific coast of Japan. I. Growth
analysis in spring-autumn. Aquat. Bot. 7: 47-56.

PRESTON, R. D., 1974. — Plant cell walls. In RoBARTS, A. W., ed., Dynamic Aspects of Plant Ultrastructure. Lon-
don: McGraw-Hill.

Roy, P. S., 1984. — New South Wales estuaries: their origin and evolution. Jn THOM, B. G., ed., Coastal
Geomorphology in Australia. Sydney: Academic Press.

SAND-JENSEN, K., 1975. — Biomass, net production and growth dynamics in an eelgrass (Zostera marina L.)
population in Vellerup Vig, Denmark. Ophelia 14: 185-201.

SETCHELL, W. A., 1929. — Morphological and phenological notes on Zostera marina L. Univ. Calif. Publs Bot.
14: 389-452.

TYERMAN, S. D., HATCHER, A. I., WEST, R. J., and LARKuM, A. W. D., 1984. — Posidonia australis growing in
altered salinities: leaf growth, regulation of turgor and the development of osmotic gradients. Aust. J.
plant Physiol. 11: 35-47.

PROC. LINN. SOC. N.SW., 109 (1), 1986

R. J. KING AND V. M. HOLLAND 39

WEST, R. J., THOROGOOD, C. A., WALFORD, T. R., and WILLIAMS, R. J., 1985. — An estuarine inventory
for New South Wales. N.S.W. Dept Agriculture, Fisheries Bulletin 2.

WOobD, E. J. F., 1959a. — Some aspects of the ecology of Lake Macquarie, N.S.W., with regard to an alleged
depletion of fish. VI. Plant communities and their significance. Aust. J. mar. Freshwat. Res. 10: 322-340.

——, 1959b. — Some east Australian sea-grass communities. Proc. Linn. Soc. N.S.W. 84: 218-226.

ZaR, J. H., 1974. — Brostatistical Analysis. New Jersey: Prentice Hall.

ZIEMAN, J. C., 1974. — Methods for the study of the growth and production of turtle grass, Thalassia testudi-
num Konig. Aquaculture 4: 139-143.

PROC. LINN. SOC. N.S.W., 109 (1), 1986

Gai) aygth

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Aquatic Angiosperms in Coastal Saline
Lagoons of New South Wales.
III. Quantitative Assessment of Zostera capricorni

R. J. KING and J. B. BARCLAY

KING, R. J., & BARCLay, J. B. Aquatic angiosperms in coastal saline lagoons of New
South Wales. III. Quantitative assessment of Zostera capricornt., Proc. Linn. Soc.
N.S.W. 109 (1), 1986: 41-50.

Data on Zostera capricorni (total biomass, below-ground, detrital leaf, flowering
stems and living shoot) and shoot measurements (percentage cover, density, leaflength
and width) and biomass of Halophila ovalis and Ruppia megacarpa for summer and winter
1978-79 are presented for 7 sites in Lake Macquarie and 5 sites in Tuggerah Lakes.
Regression equations are established relating Zostera capricorn total biomass and living
shoot biomass to percentage cover for these sites in winter and summer and these equa-
tions applied to field survey data. The total biomass of Zostera in Lake Macquarie in
summer 1985 was estimated as 1454 tonnes over an area of 11.57km?; living shoot bio-
mass was 330 tonnes. Comparable figures for Tuggerah Lakes are 1255 tonnes total bio-
mass and 453 tonnes living leaf biomass over an area of 12.26km?.

R. J. King and J. B. Barclay, School of Botany, University of New South Wales, Kensington, Aus-
tralia 2033; manuscript received 13 November 1985, accepted for publication 23 July 1986.

INTRODUCTION

The distribution of aquatic angiosperms in the estuaries and coastal lagoons on the
central and southern coast of New South Wales has been the subject of a number of
papers but these have been quantitative only in the sense that the broad areas occupied
by seagrasses have been mapped and in some cases surface area covered has been esti-
mated (Wood, 1959, for Lake Macquarie; Higginson, 1965, for Tuggerah Lakes; Harris
et al., 1980, for Lake Illawarra; Evans and Gibbs, 1980, for 5 lagoons including Lake
Macquarie and Lake Illawarra). West et al. (1985) provided an estuarine inventory for
New South Wales but in this the only information on seagrasses was a value for the total
area of seagrass and a list of those seagrasses occurring in each estuarine system.
Detailed information on the distribution, relative abundance and the area occupied by
seagrasses is available for Lake Macquarie (King, 1986b) and Tuggerah Lakes (King
and Holland, 1986).

Despite the information in these publications and in the plethora of unpublished
reports by State Government authorities until now there have been no biomass data
available for these estuarine ecosystems except for two isolated values for the maximum
total biomass of all species in Tuggerah Lakes in 1964 and 1967 (Higginson, 1971). In-
deed it is only in the last few years that any data have been published for Botany Bay
(Larkum eét al., 1984).

In both Lake Macquarie and Tuggerah Lakes the most abundant and widespread
macrophyte is the seagrass Zostera capricorni Ascherson. It covers 11.57km? of Lake Mac-
quarie (total seagrass area 14.17km?) (King, 1986b), and 11.66km? (total seagrass area
20.44km?) of the Tuggerah Lakes (King and Holland, 1986). Zostera commonly occurs
with Halophila ovalis (R. Brown) Hooker f. but while Halophila is widespread (5.59km?
and 9.82km? of Lake Macquarie and Tuggerah Lakes respectively) it is not such a sig-
nificant contributor to biomass. The fibre-weed Poszdonia australis Hooker f. does not
occur in Tuggerah Lakes and is of restricted distribution in Lake Macquarie. Ruppia
megacarpa R. Mason occurs in both systems but at the commencement of this study was

PROC. LINN. SOC. N.S.W., 109 (1), 1986

42 QUANTITATIVE ASSESSMENT OF ZOSTERA CAPRICORNI

relatively unimportant. Changes in the vegetation of Tuggerah Lakes since that time in-
dicate that more attention should be paid to this species.

In this report we present basic data on biomass of Zostera capricorn: of selected sites in
both Lake Macquarie and the Tuggerah Lakes system. This information is then used to
establish the relationship between various biomass attributes (total biomass, root, living
leaf, detrital leaf and flowering stem) and shoot measures (percentage cover, shoot den-
sity, leaf height and leaf width). Relationships between percentage cover, and both total
biomass and biomass of standing leaf stock are applied to data collected in the general
surveys (King, 1986a,b; King and Holland, 1986).

TABLE 1

Scale used to rate seagrass distribution, with estimations of percentage leaf cover

Sociability
a b
Abundance Individual strands Patches up to
or clumps 10 m diameter

c
Beds of relatively
even distribution

1 5% 10% 15%
Sparse growth (<15%)

2 15% 25% 35%
Moderate growth (15-50%)

3 inappropriate 60% 65%
Abundant growth (>50%) measure

Pacific
Ocean
33°20'S

2
Scale (Km)

Fig. 1. a (left), sampling sites in Lake Macquarie: b (right), sampling sites in Tuggerah Lakes. See also Tables
2 and 3.

PROC. LINN. SOC. N.S.W., 109 (1), 1986

R. J. KING AND J. B. BARCLAY 43

METHODS

Sampling for the quantitative analysis

Twelve sampling locations were selected to cover a range of seagrass density: sites
1-7 in Lake Macquarie and sites 8-12 in Tuggerah Lakes (Fig. la,b). These included sites
in power station cooling water discharge plumes and sites near the edge of the plumes.
Sites 1 to 5 are in the Vales Point power station cooling water plume, and sites 8 to 10 are
in the Munmorah power station plume. Seagrasses at each locality were sampled in
winter of 1978 and summer 1978/79 except for site 6 (Lake Macquarie) and sites 10 and
11 (Tuggerah Lakes) which were sampled in winter only.

In winter at least 10 quadrats were sampled at each site. These were located
randomly within a grid 20m x 5m parallel to the shore, and at approximately 1m depth.
In shallower water in these lakes seagrasses are sometimes physically damaged due to
wave action created by both wind and motor boat activity, and this may upset any
general relationships involving the leaf component. Samples were collected complete
with sediment using a diver-held corer with an internal diameter of 15cm.

Percentage cover of Zostera capricornt was visually estimated in quadrats
25cm x 25cm using a scheme similar to that of Kirkman (1978) but with 7 cover grades:

% Cover
Grade Range Midpoint Description
6 >80 90 occupying almost the entire quadrat
5 61-80 70 approx. three quarters quadrat
4 41-60 50 approx. half quadrat
3 25-40 33 approx. third quadrat
2 12-24 18 approx. fifth quadrat
1 Pall 7 1/10-1/20 quadrat
+ <2 1 very sparse, occasional leaves

The random location of quadrats in winter proved somewhat unsatisfactory since
in areas of mosaic weed growth some quadrats contained no seagrass at all (see sample
numbers in Tables 2 and 3 which summarize the data for all quadrats containing
seagrasses). In summer the samples were taken in areas subjectively assessed (Kirkman,
1978) as having intermediate weed cover for that particular locality: the size of the core
samples was increased to 18.5cm diameter, and the number of samples was standardized
at 5 at each locality. Localities 10 and 11 in Tuggerah Lakes were sampled in winter only.

At the field site all plant material was washed free of sediment in a 1.5mm mesh
sieve, bagged and then either preserved in formalin or frozen. In the laboratory the
samples were treated following the recommendations of Wetzel (1965) and Vollenweider
(1974). The macrophytes were separated into the different species, washed to remove
salt and physically cleaned of macrophytes. The Zostera was sorted into four components:
root and rhizome, detrital leaf, living leaf, and flowering stems. Fresh weights were ob-
tained after a standard spinning of each component in a simple kitchen ‘salad dryer. Dry
weights were obtained by drying to constant weight at 105°C. Percentage ash-free
weights of subsamples were obtained after oxidation to constant weight in a muffle fur-
nace at 550°C.

The following measurements were made for each core sample: number of upright
living shoots per unit area (‘density’); average length of the two longest living leaves per
shoot (‘leaf height’); average width of leaves, 10cm from the base of 10 mature living
leaves (‘leaf width’). A high level of correlation was found between biomass levels and leaf

PROC. LINN. SOC. N.S.W., 109 (1), 1986

QUANTITATIVE ASSESSMENT OF ZOSTERA CAPRICORNI

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PROC. LINN. SOC. N.S.W., 109 (1), 1986

R. J. KING AND J. B. BARCLAY

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PROC. LINN. SOC. N.S.W., 109 (1), 1986

46 QUANTITATIVE ASSESSMENT OF ZOSTERA CAPRICORNI

measurements (Barclay, 1983). This suggests the potential of using regression analysis
to estimate biomass with non-destructive sampling. The most relevant to broad scale
surveys are those relating biomass to estimates of percentage cover.

Data for total biomass of Zostera (excluding detrital material) and biomass of stand-
ing leaf stock were related to percentage cover through regression analysis. Using the
data on area and abundance of Zostera (King, 1986a,b; King and Holland, 1986) these
data have been used to give biomass figures for the entire lake systems. The categories of
sociability and abundance used in the field surveys have been combined in a two-way
table. For each category the cover was visually estimated, as above, and the percentage
cover (to the nearest 5%) assigned to each (Table 1). The category 3a is unused since
Zostera communities in which plants grow as individual strands or clumps cannot exhibit
abundant growth.

RESULTS

The full data set on seagrass biomass and Zostera shoot measurements for Lake
Macquarie and ‘Tuggerah Lakes are given in Tables 2 and 3 respectively. The biomass
figures are expressed in terms of per metre square but since the data refer only to quad-
rats in which plant material occurred they cannot be used in any comparative sense or to
indicate biomass typical of an area. All data are expressed in terms of dry weight. The
ratios of dry weight to fresh weight, and the organic contents as a percentage of dry
weight for Halophila and Zostera are given in Table 4.

TABLE 4

Dry weight as a percentage of fresh weight of Zostera and Halophila for sites in Lake Macquarie. Mean + s.e;n=82
Organic weight as percentage of total dry weight of Zostera and Halophila in both Lake Macquarie and Tuggerah Lakes.
Mean + s.e.;n=110

Zostera Be
below ground __ detrital leaf live leaf

Dry weight as Lake 9.6+0.3 8.2+0.5 10.5+0.7 8.8+0.5
percentage fresh weight Macquarie

Lake
Organic contents as Macquarie 76.541.5 78.541.7 85.9+1.0

percentage dry weight | Tuggerah 65.442.0 CIGLI6 B82 7
Lakes

69.7+1.0

The relationship between percentage cover with both total biomass and leaf stand-
ing stock is described by the series of equations given in Table 5. Separate equations are
provided for each season and for both Lake Macquarie and Tuggerah Lakes. Site 2
(Wyee Bay in Lake Macquarie) differs from other sites in a number of ways (Barclay,
1983) and is treated separately. In these equations biomass is expressed as a logarithmic
scale since there appeared to be a logarithmic relationship between percentage cover
and biomass (cf. Larkum et al., 1984).

DISCUSSION

There are relatively few studies which deal with the estimation of seagrass standing
stock or biomass in broad surveys, yet such data are important for management pur-
poses and especially so if vegetation change is to be monitored. In regions where there 1s

PROC. LINN. SOC. N.S.W., 109 (1), 1986

R. J. KING AND J. B. BARCLAY 47

TABLE 5

Linear regression equations relating log 19 (total biomass) and log 19 (living leaf biomass) to percentage cover for sites in southern
Lake Macquarie and Tuggerah Lakes.
X71 = percentage cover

SOUTHERN LAKE MACQUARIE

All sites except those in Wyee Bay
Summer

log; (total biomass) = 0.012%, + 0.525 r = 0.933 (n = 20) p<0.001
log;g (living leafbiomass) = 0.014X, - 0.200 r = 0.941 (n = 20) p<0.001
Winter ;
log; (total biomass) = 0.016X, + 0.697 r = 0.686 (n = 26) p<0.001
log;g (living leafbiomass) = 0.026X, - 0.586 r = 0.833 (n = 25) p<0.001
Wyee Bay (Site 2)
Summer
log jo (total biomass) = 0.010X, - 0.223 r=0.806(n= 5) 0.02<p<0.5
log;o (living leaf biomass) = 0.017X, - 0.434 r=0.986(n= 5) p<0.001
Winter
log jo (total biomass) = 0.035X, - 0.063 r=0.894(n= 9) p<0.001
log,g (living leafbiomass) = 0.030X, - 0.638 r=0.917(n= 9) p<0.001
TUGGERAH LAKES
Summer
log;g (total biomass) = 0.010X, + 0.417 r= 0.910 (n= 10) p<0.001
log;o (living leaf biomass) = 0.011X, - 0.037 r = 0.959 (n = 10) p<0.001
Winter
logo (total biomass) = 0.016X, + 0.240 r= .836(n= 18) p<0.001
log;9 (living leaf biomass) = 0.023X, - 0.501 r= .826(n=31) p<0.001

a marked seasonal growth pattern peak biomass may also be used as an indicator of
productivity (Nienhuis and de Bree, 1977). In broad-scale survey work normal destruc-
tive methods of vegetative sampling are rarely appropriate, not only because they are
time-consuming, but also, because the removal of vegetation may itself affect the result,
particularly if the survey area is ecologically sensitive or has only a sparse vegetation
cover.

Percentage cover has been used successfully to estimate ‘above-ground’ biomass of
aquatic angiosperms by a number of workers: Rorslett et al. (1978) in studies of fresh-
water macrophytes; Kirkman (1978) in monitoring the decline of Zostera capricorni in
Moreton Bay, Queensland; and locally by Larkum et al. (1984) in assessing total above
ground stock of Zostera capricorni in Botany Bay. A more sophisticated population density
index was used by Sheldon and Boylen (1978) to estimate cover and subsequently bio-
mass in a large freshwater lake in N.Y. State. In a broad-scale survey Mukai et al. (1980)
used a similar approach to that here to estimate the biomass of Zostera marina in Odawa
Bay, central Japan. They used somewhat fewer data, 9 samples only in an area of 68
hectares.

In the study of Larkum et al. (1984) percentage cover was related logarithmically to
both underground biomass and shoot biomass. A similar association was found in this
project in Lake Macquarie and Tuggerah Lakes. In other studies, (Nienhuis and de
Bree, 1977), in the Netherlands; McRoy, 1970, in Alaska) linear relationships were es-
tablished between the standing stock of Z. marina and percentage cover. Larkum ¢¢ al.
(1984) suggested that the logarithmic relationship may indicate a ‘synergistic effect of the
presence of one plant on the growth of another’, but whether this is caused by a more

PROC. LINN. SOC. N.S.W., 109 (1), 1986

48 QUANTITATIVE ASSESSMENT OF ZOSTERA CAPRICORNI

favourable redox potential and/or enhanced nutrient availability in dense stands (Orth,
1977) was not determined.

Non-destructive methods of estimating abundance were generally restricted to
above-ground material and in seagrass studies root biomass has often been ignored; as
indeed it is in most ecosystem studies (Caldwell, 1979). If the root to shoot ratio is any in-
dication of the energy investment in root systems then it is clear that the importance of
the root component has been underestimated. There are, however, several reports on
below-ground productivity in seagrasses which indicate that it is much less than would
be predicted by this ratio (see West and Larkum, 1983). The root/living shoot ratio of Z.
capricornt was in the range of 1.16-2.94 in summer (data from Tables 2 and 3). Such ratios
are compatible with observations made on a wide variety of communities where below-
ground productivity has been shown to account for 50-80% of total net production
(Caldwell, 1979). In winter when the living shoots die back the ratio of root/shoot is
much greater and more variable, but the picture is complicated by the impossibility of
distinguishing between living and non-living components and the root biomass.

The regressions established in this survey indicate that total biomass and standing
stock of living leaves can be estimated from percentage cover. Separate regressions are
required for the seasons (winter and summer). In this study separate regressions were
required for data from site 2 (Wyee Bay, 1.5km from the Vales Point power station out-
let). These regressions differed significantly from those at all other sites in Lake
Macquarie.

When considering the calculated biomass data the following qualifications should
be borne in mind:

(i) it is assumed that the relationship between biomass measures and percen-
tage cover at various sites and in various seasons has remained constant
during the period of the surveys,

(11) there is a compromise between breadth and intensity of survey such that
only 8 categories of percentage cover of Zostera are mapped. Hence there is
a built-in error in the estimation of biomass for any category of abun-
dance/sociability even assuming a particular area is accurately identified.
This error could be especially critical when small areas of the lake are con-
sidered separately,

(111) the equations do not take into account any possible variation in the re-
lationship between percentage cover and biomass with depth (cf. Larkum
et al., 1984).

There are few relevant data with which to compare the biomass figures calculated
using these relationships (Table 6). Larkum et al. (1984) recorded a total above-ground
biomass of 81 +4.2 tonnes for the 309ha of Zostera capricorni beds in Botany Bay. They
pointed out that this figure was considerably less (by a factor of 6-10) than would have
been estimated by taking the product of the area of the beds and biomass from a typi-
cally healthy bed. Their estimate took into account the patchy distribution of the beds.
The average biomass figure in tonnes per square kilometre of 26 for Botany Bay is com-
parable to the average values of 28 tonnes.km™ for Lake M..cquarie and 37 tonnes.km?
in Tuggerah.

The only published estimates of the dry weight standing biomass for either Lake
Macquarie or Tuggerah Lakes are those of Higginson (1971) for Tuggerah Lakes. A
single maximum value for 1964 and a single minimum value (1967) were given: 21000
tons (21333 tonnes) and 2300 tons (2337 tonnes) respectively. These figures include all
species and related to the much larger area of the lake which was then colonized by
plants. The figures were said to represent the equivalent of 2.5 tons.acre! (627
tonnes.km”™) and 0.4 tons acre (100 tonnes.km™). These values seem inordinately high

PROC. LINN. SOC. N.S.W., 109 (1), 1986

R. J. KING AND J. B. BARCLAY 49

TABLE 6

Total biomass (tonnes) and shoot biomass of Zostera capricorni in summer 1985 calculated using the linear regressions in

Table 5 in conjunction with the area and relative abundance of Zostera shown on maps for Lake Macquarie (King, 1986b)

and Tuggerah Lakes (King and Holland, 1986). Note that the total of Lake Macquarte includes seagrass from several small
areas not listed separately

Zostera capricorni

Total biomass (T) Shoot biomass (T) Area (km?)
LAKE MACQUARIE
Myuna Bay 26 6 0.29
Goonda — Fishing Station Pt 52 11 0.26
Northern Lake
(Fishing Station Pt — Cardiff Pt) 161 35 1.30
Central eastern Lake 364 78 3.69
Crangan Bay 227 55 1.05
Chain Valley Bay 171 39 1.21
Wyee Bay inc. Mannering Bay 4 4 0.16
Wyee Pt — Bluff Pt 72 16 0.62
Bonnells Bay 289 67 2.00
Total — Lake Macquarie 1454 330 11.57
TUGGERAH LAKES
Tuggerah Lake 1052 376 9.58
Budgewoi Lake 89 34 1.19
Munmorah Lake 114 43 0.59
Total — Tuggerah Lakes 1255 453 12.26

when compared with the range (10-55g.m~ or 10-55 tonnes.km”) for Zostera species in
Australia (see review of McComb et al., 1981) and values in the range 70-156g.m~ for
mature stands of Zostera capricorni in summer in Botany Bay (Larkum et al., 1984). The
highest values for any site in this survey were 172 +14g.m~? (n=5) for site 5 (Summer-
land Point) in summer. Higginson (1971) included all plants in his biomass but again
published data for ‘apparently healthy growing stands of plants’ are 49.9g.m~* for
Halophila (see McComb et al., 1981) and 403g.m~ for Ruppia; considerably less than the
values anticipated throughout the lakes. Unfortunately Higginson (1971) did not indi-
cate the way in which he derived his values. The method of estimating biomass
described here is appropriate to broad-scale surveys but it could be readily adapted to
more detailed surveys. It is especially useful when there is a need to embrace wide varia-
bility in both time and space but resources are limited. Although correlations are high
the field data are still prone to subjective assessment of the percentage cover. A multiple
regression based on several measured leaf characters (Barclay, 1983) is potentially more
accurate but the general applicability of such equations would need to be investigated.

ACKNOWLEDGEMENTS

The project was funded by the Electricity Commission of New South Wales (Grant
B250.429 to Dr R. J. King, Univ. of N.S.W.). We are pleased to acknowledge the support
and help of G. C. Coulter, Head, Development Section, Dr B. R. Hodgson (Scientific
Officer), and especially thank Mr W. Jefferson for his willing help with field work. We
thank those technical staff involved in field work in this project: in particular thanks are
due to B. Kertesz, S. McOrrie and L. Watson.

PROC. LINN. SOC. N.S.W., 109 (1), 1986

50 QUANTITATIVE ASSESSMENT OF ZOSTERA CAPRICORNI

References

BaRCLaY, J. B., 1983. — The distribution and abundance of seagrasses in estuarine lagoons on the central
New South Wales coast. Kensington: University of New South Wales, M.Sc. thesis, unpubl.

CALDWELL, M. M., 1979. — Root structure: the considerable cost of below ground function. Jn SOLBRIG et
al., eds, Topics in plant population biology: 406-427. New York: Columbia University Press.

EVANS, P., and Gisss, P. J., 1981 — Distribution of seagrass in five N.S.W. coastal lagoons. N.S.W. State Fisheries
Technical Report, 34 pp.

Harris, M. MCD., KING, R. J., and ELLIS, J., 1980. — The eelgrass Zostera capricorn: in Illawarra Lake, New
South Wales. Proc. Linn. Soc. N.S.W. 104: 23-33.

HIGGINSON, F. R., 1965. — The distribution of submerged aquatic angiosperms in the Tuggerah Lakes sys-
tem. Proc. Linn. Soc. N.S.W. 90: 328-334.

—., 1971. — Ecological effects of pollution in Tuggerah Lakes. Proc. Ecol. Soc. Aust. 5: 143-152.

-KING, R. J., 1986a. — Aquatic vegetation surveys, 1980-1986. Copy held in Biomedical Library, University
of New South Wales.

——, 1986b. — Aquatic angiosperms in coastal saline lagoons of New South Wales. I. The vegetation of Lake
Macquarie. Proc. Linn. Soc. N.S.W. 109: 11-23.

, and HOLLAND, V. M., 1986. — Aquatic angiosperms in coastal saline lagoons of New South Wales. II.
The vegetation of Tuggerah Lakes, with specific comments on the growth of Zostera capricorni Ascher-
son. Proc. Linn. Soc. N.S.W. 109: 25-39.

KIRKMAN, H., 1978. — Decline of seagrass in northern areas of Moreton Bay, Queensland. Aquat. Bot. 5:

63-76.

LARKUM, A. W. D., COLLETT, L. C., and WILLIAMS, R. J., 1984. — The standing stock, growth and shoot
production of Zostera capricornt Aschers. in Botany Bay, New South Wales, Australia. Aquat. Bot. 19:
307-327.

McComs, A. J., CAMBRIDGE, M. L., KIRKMAN, H., and Kuo, J., 1981. — The biology of Australian
seagrasses. In PATE, J. S.. and McComs, A. J., eds, The biology of Australian plants: 258-293. Nedlands:
University of Western Australia Press.

McRoy, C. P., 1970. — Standing stocks and other features of eelgrass (Zostera marina) populations on the
coast of Alaska. J. Canad. Fish. Res. Bd 27: 1811-1821.

Mukal, H., Atol, K., and IsH1pa, Y., 1980. — Distribution and biomass of eelgrass (Zostera marina L.) and
other seagrasses in Odawa Bay, Central Japan. Aquat. Bot. 8: 337-342.

NIENHUIS, P. H., and DE BREE, B. H. H., 1977. — Production and ecology of eelgrass (Zostera marina L.) in
the Grevelingen Estuary, the Netherlands, before and after the closure. Hydrobiologia 52: 55-66.
OrTH, R., 1977. — Effect of nutrient enrichment on growth of the eelgrass Zostera marina in the Chesapeake

Bay, Virginia USA. Mar. Biol. 44: 187-194.

RORSLETT, B., GREEN, N. W., and KVALVAGNPES, K., 1978. — Stereophotography as a tool in aquatic
botany. Aquatic. Bot. 4: 73-81.

SHELDON, R. B., and BOYLEN, C. W., 1978. — An underwater survey method for estimating submerged
macrophyte population density and biomass. Aquat. Bot. 4: 65-72.

VOLLENWEIDER, R. A., (ed.), 1974. — A manual on methods for measuring primary production in aquatic environments.
Oxford and Edinburgh: Blackwell Scientific Publ.

WEST, R. J., and Larkum, A. W. D., 1983. — Seagrass primary production — a review. Proc. Linn. Soc.
N.S.W. 106: 213-223.

——, THOROGOOD, C. A., WALFORD, T. R., and WILLIAMS, R. J., 1985. — An estuarine inventory for New
South Wales. N.S.W. Dept. Agriculture, Fisheries Bulletin 2.

WETZEL, R. G., 1965. — Techniques and problems of primary productivity measurements in higher aquatic
plants and periphyton. Jn GOLDMAN, C. R.., (ed.), Primary productivity in aquatic environments: 249-267.
Berkeley: University of California Press.

Woop, E. J. F., 1959. — Some aspects of the ecology of Lake Macquarie, N.S.W., with regard to an alleged
depletion of fish. VI. Plant communities and their significance. Aust. J. mar. Freshwat. Res. 10: 322-340.

PROC. LINN. SOC. N.S.W., 109 (1), 1986

Aquatic Angiosperms in Coastal Saline Lagoons
of New South Wales.
IV. Long-term Changes

R. J. KING and B. R. HODGSON

KING, R. J., & Hopcson, B. R. Aquatic angiosperms in coastal saline lagoons of New
South Wales. IV. Long-term changes. Proc. Linn. Soc. N.S.W. 109 (1), 1986: 51-60.

The area occupied by aquatic angiosperms and the biomass of Zostera capricorni
have been documented for southern Lake Macquarie and the Tuggerah Lakes from
1980 to 1985. The area of seagrasses in Lake Macquarie ranged from 4.81 to 6. 75km’,
and the total biomass of Zostera from 543 to 1099 tonnes. Winter biomass was signifi-
cantly greater than summer biomass. In the Tuggerah Lakes the area was 13.13-
19.11km* and total biomass of Zostera 840-1888 tonnes. Differences between years were
not significant because of the wide variation shown in both parameters. The data show
no time-related trends or predictable changes. When vegetation survey is to function as
baseline data this inherent variability in seagrass populations must be recognized.

R. J. King, School of Botany, University of New South Wales, Kensington, Australia 2033, and
B. R. Hodgson, Electricity Commission of N.S.W. Development Division, Hyde Park Tower,
Sydney, Australia, 2000; manuscript received 13 November 1985, accepted for publication 23 July
1986.

INTRODUCTION

The disturbance of seagrass beds by natural and artificial causes and the recog-
nition of the potential of such disturbances on commercial and recreational fishing have
led to the recent increased interest in the biology of these plants (McRoy and Helfferich,
1977; Phillips and McRoy, 1980; McComb et a/., 1981). In Australia where some 11.4%
of the coastline consists of coastal lagoons (Cromwell, 1971, in Barnes, 1980), and with
the increasing residential and industrial development in proximity to the coastal water
ways, such studies have especial relevance. In investigations of seagrass communities on
the New South Wales coast there are few detailed early studies. Hence baseline studies,
in which data are collected and analysed in order to specify the present state of the com-
munity, are a prime requirement. Generally such studies anticipate some environmen-
tal changes. The so-called ‘baseline data’ are essential to monitoring since change can
only be detected in relation to the unimpacted state. When the impact has to be inferred
from temporal change alone the sampling and statistical analysis is suboptimal in that a
control area defined a priori is lacking (Green, 1979). In cases where any future impact is
potentially able to affect a wide area, or act over a long time, the sampling of baseline
data is critical. The problem is always to obtain sufficient quantitative information that
takes into account natural fluctuations so that later comparisons can be meaningful.

In the long term studies in the coastal saline lagoons of the New South Wales coast
there have been detailed studies relating seagrass growth to specific environmental fac-
tors (e.g. Harris et al., 1980; Higginson, 1965) and very broad distribution studies in
which the results are essentially qualitative (e.g. Wood, 1959a; Evans and Gibbs, 1981;
West et al., 1985). Studies of the former type at least allow some predictions to be made
on the basis of environmental tolerances: from studies in the latter category change can
usually only be confirmed when the seagrass is no longer present.

This paper provides data on areas of seagrass coverage and biomass of Zostera
capricornt in two seagrass-dominated coastal lagoons on the central coast of New South

PROC. LINN. SOC. N.S.W,, 109 (1), 1986

52 LONG-TERM CHANGES IN SEAGRASS VEGETATION

Wales (Lake Macquarie and Tuggerah Lakes). Long-term vegetation changes are
discussed.

STUDY AREA

The two study sites, Lake Macquarie and the Tuggerah Lakes, are on the central
New South Wales coast and their aquatic vegetation is described in King (1986a,b) and
King and Holland (1986). Both lake systems support commercial fishing and prawning,
but are in areas with rapid urbanization and increasing recreational use. Proximity to
major population centres and to coal deposits have made the lakes suitable sites for
steam-generating power stations: Wangi (330MW), Vales Point (21I95MW), and
Eraring (2640MW) power stations on Lake Macquarie, and Munmorah power station
(1400MW) on the Tuggerah Lakes. These power stations use steam-driven generating
units, and the lake water is continuously drawn from an inlet channel to cool the con-
densors. The Electricity Commission of New South Wales commenced funding of long-
term seagrass surveys in the early 1960s in the Tuggerah Lakes and in the early 1970s in
Lake Macquarie. These early surveys evolved to the present quantitative surveys. The
cooling-water discharges result in the production of artificial warm water effluents (5-
9°C above ambient) in localized areas of the lakes. Given the potential impact of these
cooling-water discharges as well as changes such as the increasing turbidity and levels of
phosphorus in Lake Macquarie (State Pollution Control Commission, 1983), and com-
parable environmental changes in the Tuggerah Lakes, there was a perceived need for
quantitative data on the seagrass communities.

METHODS

Since winter 1980 the aquatic vegetation in the southern half of Lake Macquarie
and in the Tuggerah Lakes has been mapped quantitatively on a regular basis. All areas
were mapped in winter 1980 and 1981 and all areas have been mapped every summer
since. In addition areas in Lake Macquarie, Myuna Bay and Wyee Bay, have been
mapped every winter.

The vegetation was mapped and the areas of seagrass calculated using the tech-
niques outlined in King (1986b) but in surveys prior to 1984 the distance from the shore-
line to the seagrass boundary was measured using a marked tape rather than optical
range-finders. The formulae relating percentage cover and biomass measures estab-
lished in King and Barclay (1986) were used to convert the data from the maps into esti-
mates of total biomass and living shoot biomass.

RESULTS AND DISCUSSION

The results are given in a series of tables:
Table 1 Area occupied by seagrasses in southern Lake Macquarie (1980-1985).
Table 2. Biomass, total and living leaf, for seagrasses in southern Lake Macquarie
(1980-1985).
Table 3. Area occupied by seagrasses in Tuggerah Lakes (1980-1985).
Table 4 Biomass, total and living leaf, for seagrasses in the Tuggerah Lakes
(1980-1985).
A complete copy of the vegetation surveys (1980-1986) is available (King, 1986a).

LAKE MACQUARIE
The range of estimates for total area of seagrass in southern Lake Macquarie for
winter 1980 — summer 1985 was 4.81km?-6.75km? (Table 1) and differences between

PROC. LINN. SOC. N.S.W., 109 (1), 1986

R. J. KING AND B. R. HODGSON 53

TABLE 1
Areas (km?) of seagrass in southern Lake Macquarie in the years 1980-1985

Season & Year W80 S81 W681 S82 W82 $83 W83 S84 W84 $85

LAKE MACQUARIE (Southern Lake) — areas listed separately below

Zostera 5.28 3,118) 6.66 3.85 4.02 4.65 5.81
Halophila 1.16 3.58 3.29 2.68 2.24 3.85 4.59
Ruppia - — = 0.07 = 0.19 0.15
Posidonia 0.03 0.05 0.02 0.01 0.07 0.02 0.01
Total 5.49 5.99 6.75 4.92 4.81 5.00 6.65
MYUNA BAY (Eraring Power Stn outlet to Goonda Pt excluding Whiteheads Lagoon)

Zostera 0.38 0.42 0.71 0.40 0.19 0.25 0.44 0.32 0.32 0.25
Halophila 0.16 0.37 0.35 0.24 0.08 0.27 0.37 0.27 0.30 0.29
Total 0.38 0.42 OL 71 0.41 0.20 0.27 0.46 0.32 0.33 0.29
GOONDA POINT — FISHING STATION POINT

Zostera 0.52 0.54 0.85 0.51 0.38 0.53 0.47
Halophila 0.02 0.09 — _ 0.19 0.32 0.26
Posidonia 0.01 0.02 0.01 0.01 0.01 0.02 0.01
Total 0.53 0.62 0.86 0.51 0.39 0.53 0.48
CRANGAN BAY (‘Youth Camp’ — Pt Wolstoncroft)

Zostera 0.79 1.21 1.08 0.34 0.87 0.72 1.05
Halophila — 0.31 0.33 - 0.09 0.19 0.31
Posidonia 0.02 0.03 0.01 + 0.06 + ~
Total 0.81 il 337 1.09 0.35 0.92 0.72 1.07
CHAIN VALLEY BAY (Vales Pt — Frying Pan Point)

Zostera 1.30 12) 1.53 1.05 0.87 1.02

Halophila 0.31 1.01 1.14 0.93 0.58 0.74

Ruppia = = = 0.07 — 0.19 ;
Total 1.43 1.33 1.58 1.33 1.04 1.05 1.25
WYEE BAY including MANNERING BAY (Vales Pt — Wyee Pt)

Zostera 0.05 0.05 0.07 0.05 0.01! 0.09 0.16 0.21 0.20 0.27
Halophila 0.09 0.38 0.09 0.45 O251 0.43 0.82 0.53 0.83 0.83
Total 0.10 0.40 0.09 0.60 ( 0.48 0.84 0.53 0.86 0.83
WYEE POINT — BLUFF POINT

Zostera 0.38 0.38 0.36 0.23 0.27 0.42 0.61
Halophila 0.23 0.40 0.35 0.36 0.17 0.32 0.60
Total 0.38 0.40 0.36 0.36 0.41 0.47 0.74
BONNELLS BAY (Shingle Splitters Pt — Rocky Pt)

Zostera 1.86 1.43 2.06 1.27 1.29 1.43 2.00
Halophila 0.35 1.02 1.03 0.70 1.09 1.43 1.08

Total 1.86 1.45 2.06 1.36 1.30 1.43 2.00

1 Data for Mannering Bay only

values for different years and different seasons were not significant. Even within specific
areas of the lake differences in the area occupied by seagrass were not significant. In
Table 5 the data for seagrass areas are summarized along with comparative figures for
1953. The values for 1953 were measured by planimetry using an original map from the
CSIRO 1953 survey. The error in measuring such small areas is great (approx. 10%) but
based on the values presented there appears to have been an overall reduction in
seagrass beds of about one third. This reduction is apparent in all areas of the lake. Part
of this loss may be attributable to sedimentation though the estimate for loss of lake area

PROC. LINN. SOC. N.S.W., 109 (1), 1986

54 LONG-TERM CHANGES IN SEAGRASS VEGETATION

due to this is only one to two hectares per year (SPCC, 1983). Secchi disc data indicate
that lake turbidity has increased, and since the lower limit of seagrasses is thought to be
limited by light this would also reduce the area of seagrass.

TABLE 2

Biomass (tonnes) of Zostera capricorni in southern Lake Macquarie, Winter 1980 — Summer 1985. Winter surveys from
1982 onwards have included only Myuna and Wyee Bays. Maps of individual areas are given in King (1986)

LAKE MACQUARIE W80 S81 # £Wé8i S82. W82 S83 £W83 S84 W84 S85

Total biomass

Myuna Bay 87 34 231 47 55 43 115 72 69 26
Goonda-Fishing Stn Pt 117 51 264 75 = 65 = 64 = 52
Crangan Bay 188 = 115 296 52 — 176 — 132 = 227
Chain Valley Bay 340 166 299 =: 153 — 142 — 229 — 171
Wyee Bay 13 1 2 urea 1 14 3 7 pais 4
Wyee Pt-Bluff Pt 62 40 115 29 — 74 — 112 = 72
Bonnells Bay 292 =: 136 Bysshe — 201 — 427 — 289

TOTAL 1099 543 1755 531 702 1039 841

Above-ground biomass

Myuna Bay 9 8 39 10 9 10 17 18 7 6
Goonda-Fishing Stn Pt 13 11 41 17 - 15 - 14 — 11
Crangan Bay 25 24 40 12 = 42 = 31 — 55
Chain Valley Bay 38 38 29 35 = 34 = 56 — 39
Wyee Bay 2 1 1 <1 <1 1 3 2 3 4
Wyee Pt-Bluff Pt 5 9 17 6 — 19 — 28 — 16
Bonnells Bay 24 29 74 39 = 46 _ 107 = 67

TOTAL 116 8120 241 = 120 167 256 198

There are no data with which to compare the values for areas occupied by in-
dividual seagrass species (Table 2) nor the values for Zostera capricorni biomass.

While the area occupied by seagrasses remained relatively constant the biomass
changed considerably. Part of this is probably due to the marked seasonal change in the
biomass of Zostera. In winter the ratio of above-ground biomass to total biomass was
about 1:8 and in summer it was about 1:4. If the number of living leaves on the plants
varies and is influenced by non-seasonal factors such as King and Holland (1986) sug-
gested then the biomass estimates will be affected accordingly. Another factor influenc-
ing the changes between years is the relative contribution of other seagrass species to the
area occupied.

TUGGERAH LAKES

In Tuggerah Lakes the range of values for the area occupied by seagrasses is
13.13km? to 19.11km? and again there are no trends in the data (Table 3). A comparison
of the percentage of the total area of the lakes occupied by each of the three seagrass
species over the period 1980-1985 and 1963-1966 (Higginson, 1968) is given in Table 6.
This indicates a marked reduction in the area occupied by Zostera capricorn.

The area occupied by Halophila ovalis appears to have increased dramatically but in
Higginson’s surveys Halophila would have been recorded only when it grew in single
species stands. When Halophila occurs in mixed communities it is almost always the
other species which give the beds their overall physiognomy. In the period 1980-1985
Ruppia megacarpa had more than trebled in the area occupied, from 2.52km? to 8.24km?.
In the period during which Higginson observed the distribution of weeds in the

PROC. LINN. SOC. N.S.W,, 109 (1), 1986

R. J. KING AND B. R. HODGSON 2)5)

TABLE 3
Areas (km?) of seagrass in Tuggerah Lakes in the years 1980-1985

Season &
Year W80 S81 W681 $82 $83 S84 $85
TUGGERAH LAKES — (Total)
Zostera 10.92 8.66 15.96 12.85 16.69 14.61 12.26
Halophila 4.10 9.14 10.98 9.71 13.36 7.51 10.40
Ruppia 2.52 1.76 3.13 2.20 2a, 5.01 8.24
Total 14.34 13.13 17.60 14.19 18.64 16.28 19.11
TUGGERAH LAKE
Zostera 7.31 5.60 11.05 8.93 13.12 10.27 9.58
Halophila 0.67 6.05 7.16 4.53 8.15 3.77 6.43
Ruppia 2.50 1.76 3.12 1.71 ells) 3.75 5.48
Total 9.93 9.76 12.06 9.04 13.12 11.43 12.69
BUDGEWOI LAKE
Zostera 2.86 2.17 3.61 2.12 2.75 3.21 1.19
(1.91)* (1.24) (2.01) (1.06) (0.98) (2.12) (0.58)
Halophila 2.55 2.19 2.20 2.94 3.96 2.62 2.49
(1.83) (1.27) (1.89) (1.76) (2.19) (1.93) (1.34)
Ruppia 0.02 = 0.01 0.33 = 0.98 Dae
(-) (-) (-) (0.06) (-) (0.44) (1.71)
Total 3.05 2.28 3.82 3.00 4.19 3.43 4.03
MUNMORAH LAKE
Zostera 0.75 0.89 1.30 1.80 0.82 1.13 1.49
Halophila 0.88 0.90 1.61 2.24 1.25 ol? 1.48
Ruppia = = = 0.16 — 0.28 0.54

Total 1.36 1.09 Ae 2) 2.25 1.33 1.42 2039

*Data in brackets are for the Budgewoi Flats.

Tuggerah Lakes (1963-1966) the percentage of the area which was occupied by Ruppia fell
from 13.3% to zero. Such wild fluctuations in the populations of Ruppia have also been
observed in Smiths Lake (Myall Lakes system) but the factors involved have not yet been
recognized. From the data available on the total area occupied by seagrass it does not
appear that the increase in the area occupied by Ruppia is completely at the expense of
Zostera and Halophila (Fig. 2).

The only biomass figures available for Tuggerah Lakes are not directly comparable
with the data in Table 4 since they are for all seagrasses. The values were a maximum, in
1964, of 21,000 tons, and a minimum, in 1967, of 2,300 tons. These values seem to have
been arrived at by multiplying the dry weight values obtained for quadrat samples by
the area of the lake occupied by Zostera and the upper limit is considerably more than
could be anticipated (see King and Holland, 1986). The range is far wider than that
recorded here, 840-1888 tonnes (Table 4). In Tuggerah Lakes the ratio of above-ground
biomass to total biomass was about 3 at all seasons.

The justification for many environmental surveys in the marine and coastal zones
is that they are base-line studies against which man-induced change can be measured.
Gray (1976) has noted that many such surveys are inadequate to the task of monitoring
subtle, naturally-occurring changes and that they generally neglect the majority of the
living constituents in the ecosystem.

Examination of the published data on the seagrasses of the Central Coast lakes
reveals three problems:

PROC. LINN. SOC. N.S.W., 109 (1), 1986

56 LONG-TERM CHANGES IN SEAGRASS VEGETATION

TABLE 4
Biomass (tonnes) of Zostera capricorni in Tuggerah Lakes, Winter 1980-Summer 1985

TUGGERAH LAKES W80 S81 Ww8i1 $82 $83 S84 $85
Total biomass
Tuggerah Lake 627 747 1514 945 1159 1258 1052
Budgewoi Lake 169 127 236 219 242 456 89
Munmorah Lake 44 45 138 157 67 88 114
TOTAL 840 919 1888 1321 1468 1802 1255
Above-ground biomass
(living leaf)
Tuggerah Lake 213 291 642 369 443 499 376
Budgewoi Lake 50 47 78 86 92 183 34
Munmorah Lake 12 16 53 60 25 33 43
TOTAL 275 354 773 515 560 715 453
TABLE 5

Areas of seagrass in Lake Macquarie (km*) 1980-1985, compared with values measured from CSIRO maps, 1953
(R. J. MacIntyre, pers. comm.)

Year of Survey
1980-85 1953

Lake Macquarie (all areas) 14.17* 21.34
Myuna Bay (Eraring Power Stn outlet to Goonda Pt excluding Whiteheads

Lagoon) 0.20-0.71 0.78
Goonda Pt to Fishing Station Point 0.39-0.86 1.22
Northern lake (Fishing Station Point to Cardiff Point) 1.58* 3.22
Central eastern part of lake (Cardiff Point — ‘Youth Camp’) 5.10* 7.28
Crangan Bay (“Youth Camp’ — Pt Wolstoncroft) 0.35-1.37 0.65
Chain Valley Bay (Frying Pan Point to Vales Point) . 1.04-1.58 2.05
Wyee Bay including Mannering Bay (Vales Point to Wyee Point) 0.09-0.86 1.56
Wyee Point to Bluff Point 0.36-0.74 1.53
Bonnells Bay (Shingle Splitters Point to Rocky Point) 1.30-2.06 2.49

* value for 1985 summer only.

1. All surveys are a compromise between the area covered and the detail of informa-
tion collected. Earlier surveys of both Lake Macquarie and Tuggerah Lakes showed
presence and absence of the seagrass species and these were presented as maps for
each of the lake systems, but at very small scale [ (1:c.180,000 for Lake Macquarie
(Wood, 1959a) and 1:c.190,000 for Tuggerah Lakes (Higginson, 1965) |. These sur-
veys also included global figures for the area covered by seagrass, and Higginson
(1971) presented biomass figures for all aquatic angiosperms in the entire Tuggerah
Lakes system.

From these sorts of data absolute conclusions can only be drawn when seagrass
has entirely disappeared from an area which it formerly colonized. The scale of the
published maps makes it impossible to draw conclusions about specific areas of the
lakes which may have been ‘impacted’.

2. ‘The seagrass beds in the Central Coast lakes occur in an area undergoing rapid ur-
banization. As well as thermal effects associated with power stations there are en-
vironmental changes due to other factors, especially increasing sedimentation, toxic
metal pollution, and increased nutrient levels from fertilizers and sewage (Higgin-
son, 1971; Interdept. Comm., 1979). In the absence of information on species inter-

PROC. LINN. SOC. N.SW., 109 (1), 1986

R. J. KING ANDB. R. HODGSON Dy/

ao)
Oc
S 33
28D
— 2000
”
o
i=
(=
2
~ 1000
”
”
oO
£
2
ma fe)
20
qn
E Zostera
lO ~~e~ x _-® Halophila
: ~Ng--7 ..# Ruppia
o sor
® ue
<

ws WS WS WS W S
80 81 82 83 84 85

Season and Year

Fig. 1. Areas (km?) occupied by the three aquatic angiosperms: Zostera capricorni; Halophila ovalis; and Ruppia
megacarpa in Tuggerah Lakes, and biomass for Zostera.

actions, between even the common species and the environment, it is not possible to
attribute community change to any one of the concomitant environmental
perturbations.

Optimal impact study design requires the selection of a control area, or spatial
control (Green, 1979). Whether such a control area, which is uninfluenced by the
‘impact’ under study but in other ways is similar to the area under investigation, can
be designated a prior is a matter which can be generally argued (Gray, 1976). On the
basis of the data available for the present study it seems unlikely.

3. In the absence of an adequate control area impact must be inferred from temporal
change alone. In order to interpret such changes long-term systematic sampling and
recording is necessary, and it must be assumed that observed changes would not
have occurred if the area had not been environmentally altered.

With vegetation which changes as rapidly, as unpredictably and by such a mag-
nitude as the present data indicate such an assumption may be unwarranted.
Changes would need to be extensive and long lasting before firm conclusions could
be drawn.

Wood (1959b) was able to state — ‘the taxonomy of the Australian sea-grasses has
been dealt with by several workers, but no ecological studies have been made’. In the
same year he published his preliminary study on the plant communities of Lake

PROC. LINN. SOC. N.S.W., 109 (1), 1986

58 LONG-TERM CHANGES IN SEAGRASS VEGETATION

—s3—

a
E -_
= n
=
oe ax
(«>) —/
bh
< 5°]
[>)
bh
<q
|
3 t
= al
())
2 e
~ i=
(o)
=
” ~
77)
a ”
£ 7)
2 E
cs 9
oO

81 82 83 84 85
Year

Fig. 2. Area (km?) and total biomass occupied by aquatic angiosperms in Myuna Bay compared wtih value
for the total area of southern Lake Macquarie.

Macquarie (Wood, 1959a) and their significance, and in an enviable display of honesty
stated — ‘only generalities are possible regarding the requirements of the sea-grasses’.
Since that time seagrass ecology has received a great deal of attention but the un-
explained variability in species composition, distribution, and biomass of seagrass beds
in the Central Coast lakes still leaves us in a position where we could state very little
without qualification. There is certainly some evidence from Australian studies and
elsewhere that seagrass beds undergo drastic changes even under natural (non-
impacted) conditions (Larkum and West, 1983). One specific area which remains un-
studied is the means by which populations cope with environmental change. What is the
importance of acclimatization in environmental response, and/or ecotypic variation?

In North America Phillips (zn Phillips and McRoy, 1980: 29-40) has reported the
use of leaf width to indicate environmental stress: narrow leaves indicate stress, broader
leaves more optimum conditions. There have been no such morphometric studies in
eastern Australia, though Larkum et al. (1984) recorded leaf widths at various sites for
Zostera capricorn in Botany Bay.

PROC. LINN. SOC. N.SW., 109-(1), 1986

R. J. KING AND B. R. HODGSON 59

TABLE 6
Area of seagrass in Tuggerah Lakes expressed as a percentage of the total lake area, compared with values given by Higginson
(1968). Note that the values given in Higginson are for the community dominants only whereas data in the present study record
all species in mixed beds

Higginson (1968) Present study
May ’63 Aug’65 Aug’66 Period: Winter ’80/Summer’85
Zostera capricornt 27.8 27.5 oad) 1)1)2-21).7
Halophila ovalis 0.8 0.5 0.6 5.3-17.4
Ruppia megacarpa 13.3 3.3 0.0 2.3-10.7
Total seagrass 41.9 31.2 28.2 17.1-25.0
CONCLUSIONS

Long-term studies in southern Lake Macquarie and the ‘Tuggerah Lakes, of both
the area covered by seagrasses and the biomass of the major species, Zostera capricorni,
showed that marked fluctuations are inherent in the nature of the communities. Over
the period 1980-1985 these fluctuations did not appear to follow any regular or even
predictable pattern. Data such as those presented by Higginson (1965, 1968), King
(1986b) and King and Holland (1986) cannot be directly used as baseline data since they
present the results of only single surveys and therefore fail to take into account this vari-
ation in time. Long-term changes can be recognized by comparing earlier data with
those here but most earlier information is less detailed. Relative changes in abundance
and distribution of seagrass can be detected by following long-term change, in which
case the extent of change must be greater than the magnitude of the changes due to in-
herent variability, or by comparison against a control area. The selection of a control
area 1s difficult since apart from the habitat both the species and their distribution must
be similar. It is doubtful whether such control areas exist let alone can be recognized. In
the case of a specific impact such as the discharge of cooling water effluent into Myuna
Bay since early 1982 it may be valid to use the combined data for all other sites in
southern Lake Macquarie as a control. When this was done there were no significant
differences between Myuna Bay and the ‘control’ area for either area of seagrass or bio-
mass of Zostera capricorn (Fig. 2) but such areas require monitoring to see if the trends
which are apparent from the graph are continued.

Even large changes in seagrass distribution must be interpreted with caution. The
disappearance of Ruppia megacarpa from large areas of the Tuggerah Lakes in 1965 could
have been interpreted as resulting from drought conditions which existed at that time
and its reappearance in quantity since 1984 may be due to subsequent environmental
change. Such marked fluctuations are not a peculiarity of this lake system. Personal ob-
servations in Smiths Lake (Myall Lake system) show that Ruppia populations there
exhibit changes of a similar magnitude.

On the basis of what few data are available from earlier studies it is apparent that
the area of seagrass recorded in earlier studies of both Lake Macquarie and Tuggerah
Lakes has decreased. In both cases this may be attributed to increasing turbidity, but
this has not been demonstrated.

ACKNOWLEDGEMENTS

The continued support of the Electricity Commission of New South Wales has
allowed us to pursue this project for over 8 years. We are grateful to G. C. Coulter
(Manager: Power Development Division) for his long-term support and his recognition
of the value of such studies.

PROC. LINN. SOC. N.S.W., 109 (1), 1986

60 LONG-TERM CHANGES IN SEAGRASS VEGETATION

In preparing this review of the project we have drawn on the work of several
honours and graduate students and to them we are indebted: J. B. Barclay; M. McD.
Harris; V. M. Holland; K. Sweaney and L. G. Watson. We are also pleased to
acknowledge the help of technical staff of the Electricity Commission, especially Bill
Jefferson and Peter Smith, and from the Botany School, University of New South Wales:
Dr P. Farrant, M. A. Alderton, S. Francis, and B. L. J. Kertesz. Special thanks are due
to those technicians directly involved in field surveys: G. Allen, N. Jacobs, G. Jenkins,
S. McOrrie, J. Merrin, J. Nixon, L. Poole-Warren, R. Schneider, J. T. van der Velde,
and L. G. Watson. Dr R. J. MacIntyre kindly made available the copy of the vegetation
survey of Lake Macquarie: CSIRO Aug. (1953).

During the course of this project we have discussed ideas with many colleagues but
Dr P. Adam deserves our special thanks.

References

BARNES, R. S. K., 1980. — Coastal Lagoons. Cambridge: Cambridge University Press.

CROMWELL, J. E., 1971. — Barrier coast distribution: a world-wide survey. Abstr. Vol. 2nd Nat. Coast. Shallow
Water Res. Conf., p. 50.

EVANS, P., and Gbps, P. J., 1981. — Distribution of seagrass in five N.S.W. coastal lagoons. N.S.W. State Fisheries
Technical Report, 34 pp.

Gray, J., 1976. — Are marine base-line surveys worthwhile? New Scientist 70: 219-221.

GREEN, R. H., 1979. — Sampling Design and Statistical Methods for Environmental Biologists. New York: John
Wiley & Sons.

Harris, M. McD., KING, R. J., and ELLIS, J., 1980. — The eelgrass Zostera capricorn: in Illawarra Lake, New
South Wales. Proc. Linn. Soc. N.S.W. 104: 23-33.

HIGGINSON, F. R., 1965. — The distribution of submerged aquatic angiosperms in the Tuggerah Lakes sys-
tem. Proc. Linn. Soc. N.S.W. 90: 328-334.

——., 1968. — The ecology of submerged aquatic angiosperms within the Tuggerah Lakes system. Sydney:
University of Sydney, Ph.D. thesis, unpubl.

——,, 1971. — Ecological effects of pollution in Tuggerah Lakes. Proc. ecol. Soc. Aust. 5: 143-152.

INTER-DEPARTMENTAL COMMITTEE, 1979. — Tuggerah Lakes Study Report. Sydney: (N.S.W.) Ministry for Pub-
lic Works, 91 pp.

LARKUM, A. W. D., COLLETT, L. C., and WILLIAMS, R. J., 1984. — The standing stock, growth and shoot
production of Zostera capricorn: Aschers. in Botany Bay, New South Wales, Australia. Aquat. Bot. 19:

307-327.
, and WEST, R. J., 1983. — Stability, depletion and restoration of seagrass beds. Proc. Linn. Soc. N.S.W.
106: 201-212.
KING, R. J., 1986a. — Aquatic vegetation surveys, 1980-1986. Copy held in Biomedical Library, University
of New South Wales.

——, 1986b. — Aquatic angiosperms in coastal saline lagoons of New South Wales. I. Vegetation of Lake
Macquarie. Proc. Linn. Soc. N.S.W. 109: 11-23.

, and BARCLAY, J. B., 1986. — Aquatic angiosperms in coastal saline lagoons of New South Wales. III.
Quantitative assessment of Zostera capricorni. Proc. Linn. Soc. N.S.W. 109: 41-50.

, and HOLLAND, V. M., 1986. — Aquatic angiosperms in coastal saline lagoons of New South Wales. II.
Vegetation of Tuggerah Lakes. Proc. Linn. Soc. N.S.W. 109: 25-39.

McComs, A. J., CAMBRIDGE, M. L., KIRKMAN, H., and Kuo, J., 1981. — The biology of Australian
seagrasses. In PATE, J. S., and MCComg, A. J., eds, The biology of Australian plants: 258-293. Nedlands:
University of Western Australia Press.

McRoy, C. P., and HELFFERICH, C., 1977. — Seagrass Ecosystems, New York: Dekker.

PHILLIPS, R. C., and McRoy, C. P., 1980. — Handbook of Seagrass Biology. New York: Garland STPM.

STATE POLLUTION CONTROL COMMISSION (SPCC), 1983. — Environmental Audit of Lake Macquarie. Sydney:
(N.S.W.) State Pollution Control Commission.

WEST, R. J., THOROGOOD, C. A., WALFORD, T. R., and WILLIAMS, R. J., 1985 — An Estuarine Inventory
for New South Wales. N.S.W. Dept Agriculture, Fisheries Bull. 2.

——, 1959b. — Some east Australian seagrass communities. Proc. Linn. Soc. N.S.W. 84: 218-226.

PROC. LINN. SOC. N.S.W., 109 (1), 1986

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PROCEEDINGS
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LINNEAN
SOCIETY

NEW SOUTH WALES

VOLUME 109
NUMBER 2

Non-contemporaneity in the Marulan Batholith

PAUL F. CARR and BRIAN G. JONES

CarR, P. F., & JONES, B. G. Non-contemporaneity in the Marulan Batholith. Proc.
Linn. Soc. N.S.W. 109 (2), 1986: 63-67.

The Lockyersleigh Adamellite and Chapmans Creek Granodiorite crop out near
Brayton New South Wales and form part of the composite Marulan Batholith. A Rb-Sr
whole-rock isochron for these two plutons indicates an emplacement age of 326+6Ma
with an initial ®’Sr/®°Sr ratio of 0.7049. These isotopic data are very similar to pub-
lished data for the Carboniferous Bathurst Batholith (mean age of 310Ma) but differ
significantly from published data which indicate a Devonian age (approximately
400Ma) for other plutons in the Marulan Batholith. The low initial ®’Sr/*°Sr ratio im-
plies that the two plutons at Brayton were derived from an isotopically relatively un-
evolved I-type source.

P. EF. Carr and B. G. Jones, Department of Geology, University of Wollongong, PO. Box 1144,
Wollongong, Australia 2500; manuscript received 24 March 1986, accepted for publication 21 May
1986.

INTRODUCTION

The Marulan Batholith and comagmatic igneous rocks of the Bindook Volcanic
Complex crop out in the eastern part of the Lachlan Fold Belt of New South Wales (Fig.
1) and comprise a series of plutonic rocks and associated lavas and pyroclastics which
have an outcrop area of approximately 1350km?. Naylor (1939) considered that the
batholith was intruded during Devonian time and isotopic dating by Carr et al. (1980)
and Flood et al. (1982) has confirmed this age of emplacement. Two plutons of the
batholith which occur near Brayton (Fig. 1) were assigned a Devonian age by Naylor
(1939) whereas Brunker and Offenberg (1968) regarded these intrusions as Carbonifer-
ous. Isotopic data for the Brayton plutons have not been published previously and the
present investigation, which is part of a larger project on the Marulan Batholith and
Bindook Volcanic Complex, was undertaken to resolve the age of these intrusions by
using Rb-Sr isotopic techniques.

MARULAN BATHOLITH

Plutons of the composite Marulan Batholith form approximately 36% of the out-
crop area of the southern half of the Bindook Volcanic Complex and show a compo-
sitional range from tonalite and granodiorite through adamellite to granite and alkali
feldspar granite. Most plutons are relatively small and are elongate meridionally.

AGE CONSTRAINTS

The youngest stratified units with age-diagnostic fossil assemblages which are in-
truded by plutons of the Marulan Batholith are the Bungonia Limestone and the over-
lying Tangerang Formation. The Bungonia Limestone contains Ludlovian (Late
Silurian) fossils in the lower part and a Lochkovian (earliest Devonian) fauna near the
top of the formation (Jones et al., 1981), and Lochkovian faunas have been recorded from
the basal part of the Tangerang Formation (Jones et al., 1984; Jones et al., 1986; Mawson,
1975). The oldest rocks which unconformably overlie plutons of the batholith are Late
Devonian marine and terrestrial strata which crop out to the west of Bungonia (Naylor,
1939). These stratigraphic data indicate that the batholith in general is younger than
earliest Devonian but older than Late Devonian.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

MARULAN BATHOLITH

LACHLAN

SYDNEY BASIN STRATA

BATHURST BATHOLITH

BINDOOK VOLCANIC COMPLEX
sj and MARULAN BATHOLITH

is
WANly

ee LACHLAN FOLD BELT STRATA

LiL

TERTIARY ROCKS ae BINDOOK VOLCANIC
V/A CEECUANS CREEK [5 Z] suunan rocks
fe Paper ire rca ORDOVICIAN ROCKS
Fig. 1. Simplified geological map showing location of the Bathurst Batholith and the Marulan Batholith
together with the comagmatic Bindook Volcanic Complex. Sample locations and detailed geology of the

Brayton district are shown in the inset.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

PAUL F. CARR AND BRIAN G. JONES 65

Evernden and Richards (1962) obtained a Carboniferous age (313Ma; as recalcu-
lated using the constants of Steiger and Jager, 1977) for a biotite separate from a pluton
of the batholith and commented on the discordance between the stratigraphic age and
the isotopic data. Vallance (zn Packham, 1969: 198) and O’Reilly (1972) were also aware
of the anomaly.

K-Ar dates on biotite separates from three different plutons of the southern part of
the batholith indicate a mean age of emplacement of 398Ma (Carr et al., 1980). The
average Rb-Sr biotite age for the Marulan Batholith is 400Ma (Flood et al., 1982) and
both the K-Ar and Rb-Sr biotite data are consistent with a 12-point whole-rock Rb-Sr
isochron age of 419 + 33 (Flood et al., 1982).

BRAYTON PLUTONS

The oldest rocks in the Brayton district (Fig. 1) are a Late Ordovician sequence of
isoclinally folded slate, quartzite and phyllite which is unconformably overlain by
sedimentary strata and basic volcanic rocks of Silurian age (MacRae, 1978). These basic
volcanic rocks are overlain by, or faulted against, dacite and tuff of the Bindook Volcanic
Complex which Carr et al. (1980) and Jones et al. (1984) have equated with the Early
Devonian Tangerang Formation of the Bungonia region.

The Ordovician to Early Devonian sequence at Brayton was intruded by two
plutons of the Marulan Batholith, the Lockyersleigh Adamellite and Chapmans Creek
Granodiorite. O’Reilly (1972) referred to the latter intrusion as the Towrang grano-
diorite but as the name Towrang Beds has priority (Brunker and Offenberg, 1968) the
intrusion is named herein as the Chapmans Creek Granodiorite. Tertiary basalts,
dolerites and sedimentary strata post-date the adamellite and granodiorite (O’Reilly,
1972).

Stratigraphic criteria impose only broad limits on the age of the Lockyersleigh
Adamellite and Chapmans Creek Granodiorite. The intrusive relationship with the
rocks of the Bindook Volcanic Complex indicates a post-Early Devonian emplacement
whereas the non-metamorphosed Tertiary rocks indicate intrusion prior to the Tertiary.
The petrography of the various rocks and the contact metamorphic effects associated
with the intrusion of the adamellite and granodiorite have been described in detail by
O’Reilly (1972). Both plutons are composed of holocrystalline, granular rocks contain-
ing perthitic alkali feldspar, plagioclase, quartz, biotite and accessory sphene, iron-
titanium oxides, zircon and apatite. Hornblende occurs in the adamellite but is absent
from the granodiorite. Aplite veins composed of alkali feldspar and quartz occur in both
plutons.

TABLE 1
Rb-Sr data for Lockyersleigh Adamellite (LA) and Chapmans Creek Granodiorite (CCG)

Rb Sr 87Rb/86Sr 878 1r/86Sr
SAMPLE PLUTON (ppm) (ppm) (+20)
1120 LA 167 534 0.905 .70899 + 19
1121 LA 50 1029 0.141 .70553 + 11
1122 CCG 140 671 0.606 .70802 +05
1170 CCG 140 164 2.471 .71638+05
1188 LA 152 615 0.715 .70827 +05

Rb-Sr data have been determined for five whole-rock samples from the two plutons
at Brayton and the results are presented in Table 1 and Fig. 2. Regression of the data for
all five samples yields an age of 328+6Ma (Table 2) but the high value for the mean

PROC. LINN. SOC. N.S.W., 109 (2), 1986

66 MARULAN BATHOLITH

square of the weighted deviates (MSWD = 15.00) indicates that the scatter is in excess
of that attributable to analytical uncertainty alone. Exclusion of sample 1122 from the
regression reduces the MSWD to an acceptably low value of 2.48 (Table 2) but does not
make a significant difference to the date (326+6Ma). The resultant initial ®7Sr/®®Sr
ratio 1s 0.7049.

TABLE 2
Regression analyses for Lockyersleigh Adamellite and Chapmans Creek Granodiorite

INITIAL 87Sr/86Sr AGE (Ma)
SAMPLES FOR MODEL 1 ISOCHRON MSWD
(+ 20) (+ 20)
1120, 1121, 1122, 1170, 1188 .7049 +1 32846 15.00
1120, 1121, 1170, 1188 .7049 +1 326+6 2.48
Age calculated using \ = 1.42 x 1071! yr}
DISCUSSION

The Rb-Sr isochron for the four samples indicates that the two plutons at Brayton
were emplaced during the Carboniferous and are not contemporaneous with other plu-
tons from the Marulan Batholith which have been dated by Carr et al. (1980) and Flood
et al. (1982). The age of the Lockyersleigh Adamellite and Chapmans Creek Grano-
diorite does imply a temporal correlation with the Bathurst Batholith which crops out

718

.706

5 1.0 1.5 2.0 2.5

87 pp /86 ee

Fig. 2. lsochron diagram for the Lockyersleigh Adamellite and Chapmans Creek Granodiorite.

over an area of at least 1600km? in the Bathurst region of New South Wales (Fig. 1) and
has a mean age of emplacement of 310Ma (Facer, 1979). In addition, the initial

PROC. LINN. SOC. N.S.W., 109 (2), 1986

PAUL F. CARR AND BRIAN G. JONES 67

87Sr/86Sr ratio of 0.7049 for the Brayton plutons is indistinguishable from the initial
ratio of 0.7047 for the Bathurst Batholith (Flood et al., 1982) but is significantly different
from the initial ratio of 0.7061 for the Marulan Batholith and the Bindook Volcanic
Complex (Flood et al., 1982).

The K-Ar biotite age of 313Ma (recalculated) obtained by Evernden and Richards
(1962) for a pluton of the Marulan Batholith is similar to the mean age of the Bathurst
Batholith (310Ma) and the plutons at Brayton (326Ma). This similarity in age may be
fortuitous due to the loss of radiogenic argon or it may indicate the presence of another
Carboniferous pluton in the Marulan Batholith.

Mineralogical, chemical and isotopic properties of the granitic rocks of the Lachlan
Fold Belt generally permit subdivision into one of two groups (White and Chappell,
1983) which reflect derivation by partial melting of igneous rocks (‘I-type granitoids’) or
sedimentary rocks (‘S-type granitoids’). The low initial §’Sr/®°Sr ratio (0.7049) obtained
in the present study indicates that the plutons at Brayton were generated from an iso-
topically relatively unevolved source and the ratio is consistent with the I-type minera-
logical and chemical characteristics which White and Chappell (1983) have documented
for the Marulan and Bathurst Batholiths.

ACKNOWLEDGEMENTS

The University of Wollongong provided financial assistance for the project and we
are grateful for the analytical facilities provided by the University of Leeds and Carleton
University. We also gratefully acknowledge the continued help of the support staff in the
Department of Geology, University of Wollongong. Dr A. J. Wright is thanked for his
constructive criticism of the manuscript.

References

BRUNKER, R. L., and OFFENBERG, A. C., (compilers), 1968. — Goulburn 1:250,000 geological series, sheet
SI 55-12. Geol. Surv. N.S.W. Sydney.

Carr, P. F., JONES, B. G., and WRIGHT, A. J., 1980. — Dating of rocks from the Bungonia district, New
South Wales. Proc. Linn. Soc. N.S.W. 104: 113-117.

EVERNDEN, J. F., and RICHARDS, J. R., 1962. — Potassium-argon ages in eastern Australia. /. geol. Soc. Aust.
9: 1-49.

Facer, R. A., 1979. — New and recalculated radiometric data supporting a Carboniferous age for the em-
placement of the Bathurst Batholith, New South Wales. /. geol. Soc. Aust. 25: 429-432.
FLOOD, R. H., SHAW, S. E., and RILEY, G. H., 1982. — The relationship between the Bindook Volcanic
Complex and the Marulan Batholith: Sr isotopic evidence. Geol. Soc. Aust., Abstracts Ser., no. 7: 5.
JONES, B. G., Carr, P. F., and HALL, C. G., 1984. — The Early Devonian Tangerang Formation of the
Marulan-Windellama region, N.S.W. — definition and palaeoenvironmental significance. Aust. /.
Earth Sci. 31: 75-90.

—, , and WRIGHT, A. J., 1981. — Silurian and Early Devonian geochronology — a reappraisal, with
new evidence from the Bungonia Limestone. Alcheringa 5: 197-207.

——,, HALL, C. G., WRIGHT, A. J., and CARR, P. F., 1986. — The geology of the Bungonia-Windellama
area, New South Wales. Proc. Linn. Soc. N.S.W. 108: 267-286.

MacRae, G. P., 1978. — The geology of the Brayton district, N.S.W. Wollongong: University of Wollongong
BSc (Hons) thesis, unpubl.

Mawson, R., 1975. — The geology of the Windellama area, New South Wales. J. Proc. Roy. Soc. N.S.W. 108:
29-36.

NAYLOR, G. F. K., 1939. — The age of the Marulan Batholith. 7. Proc. Roy. Soc. N.S.W. 73: 82-85.

O'REILLY, S. Y., 1972. — Petrology and stratigraphy of the Brayton district, New South Wales. Proc. Linn. Soc.
N.S.W. 96: 282-296.

PacKHaM, G. H., (ed.), 1969. — The geology of New South Wales. J. geol. Soc. Aust. 16: 1-654.

STEIGER, R. H., and JAGER, E., 1977. — Subcommission on geochronology: convention on the use of decay
constants in geo- and cosmochronology. Earth Planet. Sci. Lett. 36: 359-362.

WHITE, A. J. R., and CHAPPELL, B. W., 1983. — Granitoid types and their distribution in the Lachlan Fold
Belt, southeastern Australia. Mem. Geol. Soc. Amer. 159: 21-34.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

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: ' f '

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An Atlas of Seeds and Fruits from
Macquarie Island

DANA M. BERGSTROM

BERGSTROM, D. M. An atlas of seeds and fruits from Macquarie Island. Proc. Linn. Soc.
N.S.W. 109 (2), 1986: 69-90.

Seeds and fruits of 30 members of the vascular flora from subantarctic Macquarie
Island are described and illustrated. The atlas was constructed to aid in identification of
fossils found in peat deposits on the island.

Dana M. Bergstrom, School of Biological Sciences, Macquarie University, North Ryde, Australia
2113; manuscript received 18 December 1985, accepted for publication 23 April 1986.

INTRODUCTION

Macquarie Island, (158°57’E, 54°30’S) in the Southern Ocean, is one of a num-
-ber of small isolated islands in the subantarctic zone. Like other subantarctic islands it
has a small vascular flora (Greene and Walton, 1975). The island’s flora consists of about
45 vascular plant species and 110 bryophyte species. Table 1 lists vascular species and in-
cludes all recent taxonomic revisions.

Climate during the Holocene has enabled extensive peat formations to develop on
the island. Fossil evidence of past vegetation, in the form of pollen grains, spores, seeds,
leaf and stem fragments, is preserved in these peat deposits. This atlas was constructed
to aid in the identification of seeds and fruits found in peat samples. Analysis of the fossil
record from peat deposits is providing valuable insight into vegetation dynamics and
tectonic processes on the island (Selkirk et al., 1983; Selkirk et al., 1984; Bergstrom,
1985). The usefulness of macrofossil analysis, in association with microfossil analysis, in
the reconstruction of past vegetation is becoming increasingly apparent (GreatRex,
1983; Griffin, 1977; Bergstrom, 1985; Huckerby and Oldfield, 1976; Campbell et al.,
1973). GreatRex (1983) reported that most seeds and fruit found in surface samples of
mires in Britain came from within 1m of the sampling point. Seeds coming from greater
distances were adapted for dispersal by wind or water. He suggested that reconstruction
of past communities from assemblages in a single sample would apply only to the im-
mediate vicinity of the sampling point.

There have been numerous comments in the literature on the stability of the mor-
phology of seeds (Montgomery, 1977; Corner, 1976; Berggren, 1969), with the last two
authors suggesting the value of the use of seed morphology as a tool in systematics.

METHODS

Collections

Seeds and fruits were collected from plants on Macquarie Island during the sum-
mers of 1979-80, 1983-84. The dry seeds and fruits are part of the Herbarium, School of
Biological Sciences, Macquarie University, but will be lodged with the National Her-
barium of New South Wales, Royal Botanic Gardens, Sydney, as voucher specimens. Of
the 40 angiosperm species known from the island, 10 have not been collected with either
seeds or fruits, or with mature seeds or fruit.

Form of Descriptions
All descriptions are for identification of seeds and fruits under a dissecting micro-

PROC. LINN. SOC. N.S.W,, 109 (2), 1986

70 SEEDS AND FRUITS FROM MACQUARIE ISLAND

scope. They are for the smallest dispersal unit, be it a seed or indehiscent fruit such as an
achene. On occasions when it was hard to determine whether seeds or fruit were dis-
persed, descriptions for both seed and fruit are given.

The descriptions are divided into a number of sections.

a) Dimensions
The position of the hilum or basal scar is taken as the base of the seed or fruit. The

TABLE 1

Extant Vascular Flora of Macquarve Island
Nomenclature after Copson (1984) and Seppelt et al. (1984) except where indicated

Lycopodiaceae — Lycopodium sp.
Blechnaceae — Blechnum penna-marina
Grammitidaceae — Grammitis poeppigeana
Hymenophyllaceae — Hymenophyllum peltatum
Aspidiaceae — Polystichum vestitum
Apiaceae — Azorella selago
Hydrocotyle sp.
Araliaceae — Stilbocarpa polaris
Asteraceae — Cotula plumosa
Pleurophyllum hookeri
Brassicaceae — Cardamine corymbosa
Callitrichaceae — Callitriche antarctica
Caryophyllaceae — Cerastium fontanum

Colobanthus muscoides
C. quitensis

Stellaria decipiens

S. media
Crassulaceae — Crassula moschata
Cyperaceae — Carex trifida

Isolepis aucklandicus(1)
* Uncinia divaricata

* U. hookert
Haloragaceae — Myriophyllum triphyllum
Juncaceae — Juncus scheuchzertoides
Luzula crinita var. crinita(2)
Onagraceae — Epilobium brunnescens var. brunnescens(3)
E. pedunculare(4)
Orchidaceae — Corybas macranthus
Poaceae — Agrostis magellanica

Anthoxanthum odoratum
Deschampsia chapmanii
D. penicillata

Festuca contracta

Poa annua

P. foliosa

P. hamiltoniz

P. litorosa

Puccinellia macquariensis
Polygonaceae — Rumex crispus
Portulacaceae — Montia fontana
Ranunculaceae — Ranunculus biternatus
Rosaceae — Acaena magellanica

A. minor
Rubiaceae — Coprosma pumila

Galium antarcticum

(1) = Scirpus aucklandicus (Wilson, 1981); (2) = Luzula campestris (Edgar, 1975); (3) = Epilobium nerteroides
(Raven and Raven, 1976); (4) = Epilobium linnaeoides (Raven and Raven, 1976).
* = specimens identified Karen Wilson (pers. comm., 1985).

PROC. LINN. SOC. N.SW., 109 (2), 1986

DANA M. BERGSTROM 71

L = Length
W = Width

T= Thickness

Fig. 1. Diagram illustrating how dimensions were determined. Three different seed/fruit types are shown.

length is measured from the base to the apex or, where a style is present, to the base of
the style. The width is perpendicular to this and the thickness, perpendicular to the 2
axes. All measurements are taken at the widest point of the seeds or fruits. Fig. 1 shows
how length, width and thickness measurements on three types of seeds and fruits were
determined. The measurements given are the means of measurements on 10 seeds +

standard error. The standard error by no means gives the range within the species, but
Montgomery (1977) suggests that although seeds may vary in size, depending on growth
conditions, the ratio of the measurements is usually constant. When 10 seeds were not
available the number of seeds measured is given in the form of n = x, where x is the
number measured.

b) Shape
The shape has been given in terms of longitudinal section (I.s.) and cross section
(c.s.). These sections have been considered as simple symmetrical plane shapes. The
shapes are delimited mathematically as ratios:
— in the case of|.s., length : width
— in the case of c.s., thickness : width.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

72 SEEDS AND FRUITS FROM MACQUARIE ISLAND

ik Og Beit 21ers 6:5 1:1 5:6 2:3 1:2 1:3 1:6 1:12

an oe ae
1 3 5 6 7 8 10 11
OBLONG | | | | Ce Qala —
iZeISHOeRIAe VTS Rr elORese riven! IBURT A aIO Re 20n Maia2l 237s
RHOMBIC () O > CO =
2526082) 2 B00. (eh 5,32)! Wh 33 See

SCS

NESE 36 38 40 41 42 43 44
WO
CEONATE Ay ig AS A BS Gp) ag
TRIANGULAR | \\ /\ iA is AX IS DANA
72 73 74 75 76 77 78 79 80 81 82 83
\w7 oS
OBTRIANGULAR | | Vy, iW) VV WY
84 85 86 87 88 89 90 91 92 93 94 95
Language equivalents:
ELLIPTIC OVATE
1-2 narrowly elliptic 36-37 narrowly ovate
3-4 elliptic 38-39 ovate
5 broadly elliptic 40-41 broadly ovate
6 circular 41-42 very broadly ovate
7 transversely broadly elliptic 43-44 depressed ovate
8-9 transversely elliptic
10-11 transversely narrowly elliptic OBOVATE
45-46 narrowly obovate
OBLONG 47-48 obovate
12 linear 49-50 broadly obovate
13-14 narrowly oblong 50-51 very broadly obovate
15-16 oblong 52-53 depressed obovate
17 broadly oblong
18 square TRIANGULAR
19 transversely broadly oblong 72 linear triangular
20-21 transversely oblong 73-74 narrowly triangular
22-23 transversely narrowly oblong 75-76 triangular
24 transversely linear 77-78 broadly triangular
78-79 very broadly triangular
RHOMBIC 80-81 shallowly triangular
25-26 narrowly rhombic 82-83 very shallowly triangular
27-28 rhombic
29 broadly rhombic OBTRIANGULAR
30 quadrate rhombic 84 linear-obtriangular
31 transversely broadly rhombic 85-86 narrowly obtriangular
32-33 transversely rhombic 87-88 obtriangular
34-35 transversely narrowly rhombic 89-90 broadly obtriangular

90-91 very broadly obtriangular
92-93 shallowly obtriangular
94-95 very shallowly obtriangular

Fig. 2. Chart of plane shapes and descriptive terminology (after Montgomery, 1977, after Systematics Associ-
ation, 1962).

PROC. LINN. SOC. N.S.W., 109 (2), 1986

DANA M. BERGSTROM 73

The numbers following the shape descriptions are serial numbers given by the Sys-
tematics Association Committee for Descriptive Biological Terminology (1962) to plane
shapes, shown in Fig. 2.

c) Comments
Comments are self-explanatory. A glossary is provided at the end of the
descriptions.

d) Colour

Colours of dry seeds and fruits were ascertained by use of the ‘Revised Standard
Soil Color Chart’ by Oyana and Takehara (1967). This standard was chosen as it is
widely available. The description method is based on a system in which colour can be
measured by three attributes: Hue which represents the dominant spectral colour such
as red or blue; Value — which represents the relative lightness of colour; Chroma — the
relative purity of spectral colour. A serial number is given. Thus, Hue 7.5YR 6/8 con-
sists first of the hue number (Hue 7.5YR), then the value number (6), followed by the
chroma number (8). A description of the colour is also given (e.g. orange) based on ter-
minology used by Oyana and Takehara (1967). All colours were assessed under the same
natural light conditions.

PHOTOGRAPHS

The photographs (Figs 3-10) show seeds and fruits against a background of grey
plasticine. The length-wise orientation of the figures has the hilum or fruit scar pointing
towards the caption. Where possible seeds and fruits were positioned so that both the Ls.
and c.s. could be viewed. The scale on each photograph indicates Imm.

DESCRIPTIONS OF SEEDS AND FRUITS
APIACEAE

Azorella selago (Fig. 3A)
Length: 1.61 +0.04mm
Width: 0.91 +0.05mm
Thickness: 0.68 +0.06mm
Shape: —

Longitudinal section: elliptic (3-4) or irregular.

Cross section: varied, due to distortion from other mericarps at the commissure.
Comments: Mericarps. Surface rough with 5 distinct irregular longitudinal ridges. Per-
sistent style 1.48 +0.5mm long. Floral remnant may be present.

Colour: Hue 10YR 7/6 bright yellow brown.

ARALIACEAE

Stilbocarpa polaris (Fig. 3B,D)
Length: 2.24+0.03mm
Width: 0.92 +0.05mm
Thickness: 1.14+0.05mm
Shape: —

Longitudinal section: broadly ovate (41-42).

Cross section: transversely elliptic (8-9) or irregular.
Comments: Seeds borne in black, shiny, spherical fruit, centre of which is hollow. Seed
surface coarse, often with fleshy endocarp still attached. No hilum. Cream, persistent Y-

shaped vascular trace on one surface. In arms of ‘Y’ there is a hole.
Colour: Hue 7.5YR 4/6 brown.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

SEEDS AND FRUITS FROM MACQUARIE ISLAND

Yes

"sa[eos WWI] ‘sauayoe vsownydg v/njJ07) = CD ‘spe2es

q ‘sina = g ‘sunjod ndins0q]145

= (9 q ‘sdieoiiaur odyas nyjasozp = Wg Sy

LINN. SOC. N.S.W., 109 (2), 1986

PROC

DANA M. BERGSTROM 75

fontanum seeds. D = Colobanthus muscoides seeds. 1mm scales.

tum

bosa, seeds. C = Cerast

ine corym

, achenes. B = Cardam

. 4. A = Pleurophyllum hooker

PROC. LINN. SOC. N.S.W., 109 (2), 1986

76 SEEDS AND FRUITS FROM MACQUARIE ISLAND

ASTERACEAE

Cotula plumosa (Fig. 3C)
Length: 2.04+0.04mm
Width: 1.18 +0.03mm
Thickness: 0.94 +0.03mm
Shape: —
Longitudinal section: obovate (47).
Cross section: transversely elliptic (8).
Comments: Achene. Longitudinal axis curved. Surface of achene reticulate and
coarsely punctate. Persistent style and tubular corolla.
Colour: Hue 10YR 7/4 dull yellow orange.

Pleurophyllum hooker (Fig. 4A)
Length: 3.0+0.14mm
Width: 0.88 + 0.05mm
Thickness: 0.54 + 0.02mm
Shape: —
Longitudinal section: narrowly obovate (45-46) to triangular (85-86).
Cross section: transversely oblong (21).
Comments: Achene with plumose pappus. Surface of achene velutinous, hairs white.
Stylar and perianth remnants often present. Pappus hairs approximately 6mm long, un-
equal in length and bristly.
Colour: Hue 2.5YR dull reddish brown.

BRASSICACEAE

Cardamine corymbosa (Fig. 4B)
Length: 1.340.02mm
Width: 1.0+0.02mm
Thickness: 0.4+0.02mm
Shape: —
Longitudinal section: elliptical to broadly elliptic (4-5).
Cross section: depressed ovate (44) or irregular.
Comments: Cotyledons accumbent with cotyledons and radicle indicated by a sulcus.
Surface undulating, puncticulate and shiny. In c.s. the seeds compressed at margins.
Funicular remnant light yellow orange (Hue 10Y).
Colour: Hue 10R 5/8 red, margins and hilum darker.

CARYOPHYLLACEAE

Cerastium fontanum (Fig. 4C)
Length: 0.7 +0.15mm
Width: 0.68 +0.01mm
Thickness: 0.54+0.01mm
Shape: —
Longitudinal section: broadly obovate (49-50) or irregular.
Cross section: transversely oblong (19-20).
Comments: Hilum within deep notch. Surface coarsely papillate. Papillae low and
rounded with ovoid stellate bases. Arrangement of papillae may be concentric, particu-
larly along margins. Small, hyaline, protoxylem remnant attached to hilum.
Colour: Hue 5YR 5/8 bright reddish brown.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

deh

DANA M. BERGSTROM

sa]BOs WW] ‘sauayoR vpifi4] xalDD = (‘SPOS DIpaw vILD]IAIG = DZ ‘Spaas suargrsap v1L0]]0;5

Spoas sisuajinb snyjungojo = WG diy

PROC. LINN. SOC. N.S.W., 109 (2), 1986

78 SEEDS AND FRUITS FROM MACQUARIE ISLAND

Colobanthus muscoides (Fig. 4D)
Length: 0.38 +0.002mm
Width: 0.65 +0.002mm
Thickness: 0.34 +0.002mm
Shape: —
Longitudinal section: depressed ovate (43-44) or irregular.
Cross section: transversely elliptic (8-9) or irregular.
Comments: Seeds slightly reniform. Margins often depressed forming sulcus (hence
irregular in 1.s.). Surface faintly colliculate and translucent. Minute white caruncle.

Colour: Hue 7.5YR 5/8 bright brown.

Colobanthus quitensis (Fig. 5A)
Length: 0.55 +0.02mm
Width: 0.62 +0.02mm
Thickness: 0.38 +0.0i1mm
Shape: —
Longitudinal section: depressed obovate (52-53), or irregular.
Cross section: transversely elliptic (8-9).
Comments: Margins often depressed forming sulcus (hence irregular in l.s.). Surface
translucent and very faintly colliculate. Minute white caruncle.
Colour: Hue 5YR 5/8 bright reddish brown.

Stellaria decipiens (Fig. 5B)
Length: 0.98+0.05mm n=7
Width: 0.97 + 0.05mm n=7
Thickness: 0.64+0.05mm n=7
Shape: —

Longitudinal section: circular (6) to irregular.

Cross section: transversely elliptic (8-9).
Comments: Hilum in deep notch. Concentric to irregular papillose surface. Papillae ir-
regular and elongate, especially along margin and towards the hilum/base.
Colour: Hue 7.5R 4/8, 3/4, 3/6 red to dark red.

Stellaria media (Fig. 5C)
Length: 1.25+0.04mm
Width: 1.26+0.04mm
Thickness: 0.8 +0.01mm
Shape: —
Longitudinal section: broadly elliptic (5-7) or broadly obovate (48-51) or irregular.
Cross section: transversely elliptic (8-9), or oblong (20-21) or irregular.
Comments: Hilum in deep notch. Concentric rings of low rounded papillae on surface.
Papillae arise from raised irregular or star-shaped bases.
Colour: Hue 7.5R 4/8 red.

CYPERACEAE
Carex trifida (Fig. 5D)
Length: 1.79+0.02mm n=6
Width: 1.05+0.06mm n=6
Thickness: 0.85 +0.06mm n=6

PROC. LINN. SOC. N.S.W., 109 (2), 1986

DANA M. BERGSTROM 19

,C = achenes. D = Juncus scheuchzerioides seeds. 1mm scales.

perigynium

,B =

tvaricata

wnia

cus seeds. B& C = Unc

kland

1S AUC.

. 6. A = Isolep

Fig

PROC. LINN. SOC. N.S.W., 109 (2), 1986

80 SEEDS AND FRUITS FROM MACQUARIE ISLAND

Shape: —

Longitudinal section: elliptic (3-4).

Cross section: triangular to shallowly triangular (80-82).
Comments: Achene. Longitudinal axis concave-convex. Surface puncticulate and lus-
trous. Persistent, slender and contorted stylar remnant or just stylar base present. Fruit
borne in papery perigynium which is ovate in |.s., transversely elliptic in c.s. (approxi-
mately 0.5mm long), somewhat fusiform with stipitate base.
Colour: Hue 10YR 8/6-8/8 yellow-orange.

Tsolepsts aucklandicus (Fig. 6A)
Length: 1.01 +0.01mm
Width: 0.73 +0.0imm
Thickness: 0.45 + 0.01mm
Shape: —
Longitudinal section: broadly obovate (49-50).
Cross section: shallowly triangular (81-82), tending to planoconvex.
Comments: Margins slightly ridged, base stipitate. Surface areolate. Stylar base obtuse.
Three loose, ligulate bristles from base, approximately 2x longer than achene.
Colour: Hue 7.5YR 4/6 brown.

Uncinia divaricata (Fig. 6B,C)
Length: 2.43 +0.02mm
Width: 1.21 +0.04mm
Thickness: 0.79+0.04mm
Shape: —
Longitudinal section: elliptic (3).
Cross section: shallowly triangular (80).
Comments: Achene. Sides slightly convex, edges rounded. Surface with profuse, low,
rounded papillae. Achene borne in perigynium. Stylar remnant that, if intact, protrudes
from perigynium and terminates in a hook. Surface of perigynium striated.
Colour: Hue 5YR 6/6-6/8 orange.

JUNCACEAE

Juncus scheuchzeriordes (Fig. 6D)
Length: 0.7 +0.01mm
Width: 0.25+0.01mm
Thickness: 0.25 +0.01mm
Shape: —
Longitudinal section: elliptic (3).
Cross section: circular (6) or irregular.
Comments: Seed fusiform. Coarsely striate, whitish membranous coating on surface.
There is often a prominent longitudinal ridge of tissue. Base is usually nodulous or

sometimes minutely pointed.
Colour: Hue 5YR 5/8-6/8 bright reddish-brown to orange.

Luzula crinita var. crinita (Fig. 7A)
Length: 1.03 + 0.01mm

Width: 0.53 + 0.0i1mm
Thickness: 0.43 + 0.01mm

PROC. LINN. SOC. N.S.W., 109 (2), 1986

(oc)
pare

DANA M. BERGSTROM

= Luzulacrinita var crinita seeds. B = Epilobium brunnescens var. brunnescens seeds. C = Epilobium pedunculare seeds. D = Corybas macranthus seeds. 1mm scales.

C
Fig. 7. A

PROC. LINN. SOC. N.S.W., 109 (2), 1986

82 SEEDS AND FRUITS FROM MACQUARIE ISLAND

Shape: —

Longitudinal section: elliptic (3).

Cross section: very broadly ovate (41-42).
Comments: Seed fusiform with minutely pointed apex and obtuse nodulous base, hilum
inconspicuous. A large (1.26 +0.01mm, n=10) whitish aril completely envelops the seed.
Surface of seed reticulate and glistening. Surface of caruncle faintly striated and
areolate.

Colour: Hue 7.5R 3/4-3/6 dark red, base darker.

ONAGRACEAE

Epilobium brunnescens var. brunnescens (Fig. 7B)
Length: 0.81 +0.01mm
Width: 0.62 +0.0imm
Thickness: 0.2 +0.01mm
Shape: —
Longitudinal section: narrowly obovate to obovate (46-47).
Cross section: transversely oblong (20-21).
Comments: Sides often depressed. Longitudinal sulcus, deepening towards apex, ter-

minating with cream coma. Surface longitudinally papillose.
Colour: Hue 5YR 6/8 orange.

Epilobium pedunculare (Fig. 7C)
Length: 0.70 +0.01mm
Width: 0.29 +0.0imm
Thickness: 0.21 +0.04mm
Shape: —
Longitudinal section: narrowly obovate (46-47).
Cross section: transversely oblong (20-21).
Comments: Sides often depressed. Surface longitudinally papillose. Longitudinal
sulcus, deepening towards apex, and terminating with cream coma. Base minutely
pointed.
Colour: Hue 5YR 6/8 orange.

ORCHIDACEAE

Corybas macranthus (Fig. 7D)
Length: 0.67 +0.03mm
Width: 0.12 +0.01mm
Thickness: approx. 0.1mm
Shape: —
Longitudinal section: elliptic (1) or irregular (e.g. twisted).
Cross section: circular (6) or irregular.
Comments: Small spherical embryo in transparent membranous reticulate seed coat.
Base tapering or blunt.
Colour: Hue 10YR 8/3 light yellow-orange.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

DANA M. BERGSTROM

‘sofeos wiuy ‘stsdoAie9 vnuun vog

q ‘sisdod1e9 0,904ju09 vIN SAT

9 ‘sisdoAre9 nunudoys visdunyssaq = g ‘sisdoAsed vaiunjpadow sysodpy = Wg di

ites
Ley

PROC. LINN. SOC. N.S.W., 109 (2), 1986

84 SEEDS AND FRUITS FROM MACQUARIE ISLAND

POACEAE

Agrostis magellanica (Fig. 8A)
Length: 1.5+0.01mm a7)
Width: 0.52 +0.03mm n=7
Thickness: 0.42 +0.02mm n=7
Shape: —

Longitudinal section: narrowly elliptic (2-3).

Cross section: broadly elliptic (5).
Comments: Caryopsis. Apex minutely pointed. Small coma present. V-shaped groove
near fruit scar. Surface sometimes concave.
Colour: Hue 7.5YR 7/8 yellow-orange.

Deschampsia chapmani (Fig. 8B)
Length: 1.02 +0.03mm
Width: 0.47 +0.02mm
Thickness: 0.45 + 0.01mm
Shape: —
Longitudinal section: obovate (47).
Cross section: circular (6).
Comments: Caryopsis fusiform. Nodulous apex terminating in small coma. Apiculate
base with lacerate fruit scar. Surface rugose.

Colour: Hue 7.5YR 5/8.

Festuca contracta (Fig. 8C)
Length: 3.19+0.01mm n=2
Width: 1.16+0 n=
Thickness: 0.58+0.04mm n=2
Shape: —

Longitudinal section: narrowly obovate (46).

Cross section: depressed obovate (43).
Comments: Caryopsis. Base minutely pointed, oblique fruit scar. Apex obtuse with ex-
ocarp extending into wing. Dark stripe on concave surface. Surface rugulose.
Colour: Hue 7.5YR 5/8 bright brown.

Poa annua (Fig. 8D)
Length: 1.48+0.06mm n=5
Width: 0.57 +0.02mm n=9
Thickness: 0.38 +0.02mm n=9
Shape: —

Longitudinal section: narrowly elliptic (2-3).

Cross section: irregular.
Comments: Caryopsis. Base blunt, apex terminating in a short white coma. Surface

rugose.
Colour: Hue 10YR 6/8.

Poa foliosa (Fig. 9A)
Length: 1.9+0.02mm
Width: 0.44+0.01mm
Thickness: 0.43 +0.01mm

PROC. LINN. SOC. N.S.W., 109 (2), 1986

DANA M. BERGSTROM

soyeos UW]

Spoos vuvjuof Duo

stsdoAie9 sisuatsonboou. D1JaUIIING$

stsdoAre9

LEX

ruoyrUDy Dog

stsdodie9 vs0170f Dog

6 ay

PROC. LINN. SOC. N.S.W., 109 (2), 1986

86 SEEDS AND FRUITS FROM MACQUARIE ISLAND

Shape: —

Longitudinal section: narrowly elliptic (1-2).

Cross section: broadly triangular (78).
Comments: Caryopsis. Sides often concave. Apex minutely pointed. Fruit-scar white,
rough surfaced and blunt. Surface translucent and rugulose.

Colour: Hue 7.5YR 5/8 bright brown.

Poa hamiltonu (Fig. 9B)
Length: 1.94+0.06mm
Width: 0.56 + 0.01mm
Thickness: 0.53 +0.02mm
Shape: —
Longitudinal section: narrowly elliptic (1-2).
Cross section: circular (6) or irregular.
Comments: Caryopsis. Exocarp extending beyond rest of fruit by approximately 0.5mm
and terminating in small white coma at apex.

Colour: Hue 7.5YR 5/8 bright brown.

Puccinellia macquariensis (Fig. 9C)
Length: 1.67 +0.04mm
Width: 0.56+0.02mm
Thickness: 0.47 +0.02mm
Shape: —
Longitudinal section: narrowly elliptic to elliptic (2-3).
Cross section: broadly elliptic to circular (4-6).
Comments: Caryopsis fusiform, apex minutely pointed. Seed coat forming cream fruit-
scar at base. Surface rugose.

Colour: Hue 2.5YR 4/8 reddish brown.

PORTULACACEAE

Montia fontana (Fig. 9D)
Length: 1.5+0.22mm
Width: 1.2 +0.0imm
Thickness: 0.9 +0.01mm
Shape: —
Longitudinal section: obovate to broadly obovate (48-49).
Cross section: transversely elliptic (8).
Comments: Embryo coiled. Compressed, keeled edge ascending from hilum. Surface
colliculose with regular, tending to concentric pattern. Very shiny (lustrous). Obvious
pale yellow caruncle (Hue 2.5Y 8/4) with areolate surface.
Colour: Hue 5RP 1.7/1 purplish black.

RANUNCULACEAE

Ranunculus biternatus (Fig. 10A)

Length: 1.9+0.03mm

Width: 1.8+0.04mm

Thickness: 1.3+0.02mm

Shape: —
Longitudinal section: very broadly ovate (41-42).
Cross section: broadly ovate (40-41).

PROC. LINN. SOC. N.S.W., 109 (2), 1986

DANA M. BERGSTROM 87

Fig. 10. A = Ranunculus biternatus achene. B = Acaena magellanica achenes. C = Acaena minor achenes. D

PROC. LINN. SOC. N.S.W., 109 (2), 1986

Coprosma pumila seeds. 1mm scales.

88 SEEDS AND FRUITS FROM MACQUARIE ISLAND

Comments: Achene. Deep sulcus ascending from hilum and fruit scar to base of lateral
persistent style. Style often recurved with terminal hook. Surface irregular and rugose.
Colour: Hue 10YR 8/1.2-8/4 light grey to light yellow-orange. Apex and style darker.

ROSACEAE

Acaena magellanica (Fig. 10B)
Length: 3.1+40.06mm
Width: 1.11 +0.03mm
Thickness: 1.11 +0.03mm
Shape: —
Longitudinal section: obtriangular (87).
Cross section: broadly oblong to square (17-18).
Comments: Achene enclosed in hardened villous calyx (hairs white) which is endowed
with 4 subulate spines approximately 8mm long. Spines barbed at tip. Corolla and
stylar remnant, or at least persistent stylar base, present.
Colour: 2.5Y 7/6 bright yellowish brown.

Acaena minor (Fig. 10C)
Length: 3.16+0.1mm
Width: 1.1+0.03mm
Thickness: 1.0 +0.03mm
Shape: —
Longitudinal section: obtriangular (87).
Cross section: broadly oblong to square (17-18).
Comments: Achene enclosed in hardened villous calyx (hairs white) which is endowed
with 4 subulate spines approximately 6mm long. Spines barbed at tip. Corolla and
stylar remnant, or at least persistent stylar base, present.
Colour: Hue 7.5YR 6/8 orange.

RUBIACEAE

Coprosma pumila (Fig. 10D)
Length: 2.5+0.1mm
Width: 1.58 +0.03mm
Thickness: 1.3 +0.01mm
Shape: —
Longitudinal section: obovate (48).
Cross section: broadly elliptic (5).
Comments: Seed fusiform. Longitudinal axis curved. The base is minutely pointed with
hilum inconspicuous. Surface rugulose.
Colour: Hue 10YR 7/6 bright yellow brown.

GLOSSARY

Accumbent Lying face to face

Achene A dry, indehiscent, one-seeded fruit

Areolate Having a distinct but fine network of spaces

Aril An appendage or outer covering of a seed, growing from hilum or funiculus
Bristle A stiff hair

Caruncle~ An aril at or about the hilum or funiculus

Colliculate Covered with small, rounded elevations, or hillocks

Coma _ A tuft of hairs covering apex

PROC. LINN. SOC. N.S.W., 109 (2), 1986

DANA M. BERGSTROM 89

Commissure A junction or seam

Fruit Scar Scar on fruit indicating point of attachment to parent plant
Fusiform Swollen in the middle and tapering towards the ends
Funiculus Stalk by which a seed 1s attached to ovary wall or placenta
Hilum Scar on a seed indicating point of attachment to funiculus
Indehiscent Not opening

Ligulate Strap-shaped

Mericarp _1-seeded portion of fruit which may/may not split at maturity
Nodulous With small knobs

Papillate Small, nipple-shaped projections

Perigynium Sheath which envelops achenes belonging to the Cyperaceae
Punctate Marked with dots or depressions

Puncticulate Finely punctate

Reticulate Netted, more distinct than areolate

Rugose Coarsely wrinkled

Rugulose With very fine wrinkles

Stellate Star-like

Stipitate Witha short stalk

Sulcus A groove or furrow

Velutinous Having fine straight hairs

Villous Having long silky hairs

ACKNOWLEDGEMENTS

Permission to visit Macquarie Island, granted by the Macquarie Island Advisory
Committee, and logistic support from the Antarctic Division, Australian Department of
Science, for visits in 1983 and 1984 are gratefully acknowledged.

Dr Patricia Selkirk (Macquarie University) and Geof Copson (Tasmanian
National Parks and Wildlife Service) helped with plant identification and collected
many plant specimens in fruit for the atlas. Ms Karen Wilson (National Herbarium of
New South Wales, Royal Botanic Gardens, Sydney) identified certain specimens. Dr
Tony Orchard (Director, Tasmanian Herbarium, Department of Botany, University of
Tasmania) allowed access to Macquarie Island herbarium material. Thanks are due to
Dr Patricia Selkirk and Dr Bob Selkirk for constructive comments on the manuscript.

Photographs were taken by Mr Ron Oldfield and printed by Ms Jenny Norman.

References

BERGGREN, G., 1969. — Atlas of seeds and small fruits of North West European plant species with morphological descrip-
tions. Part II Cyperaceae. Stockholm: Swedish Natural Science Research Council.

BERGSTROM, D. M., 1985. — The Holocene vegetation history of Green Gorge, Macquarie Island. North
Ryde: Macquarie University, M.Sc. thesis, unpubl.

CAMPBELL, E. O., HEINE, J. C., and PULLAR, W. A., 1973. — Identification of plant fragments and pollen
from peat deposits in Rangitaiki Plains and Maketu Basins. N.Z. j. Botany, 11: 317-310.

Copson, G. R., 1984. — An annotated atlas of the vascular flora of Macquarie Island. Australian national An-
tarctic research Expeditions, Research Notes, 18: 1-70.

Corner, E. J. H., — 1976. — The Seeds of Dicotyledons. First edition, 2 vols. Cambridge, London, New York:
Cambridge University Press.

Epoar, E., 1975. — Australasian Luzula. N.Z. J. Botany, 13: 781-802.

GREATREX, P. A., 1983. — Interpretation of macrofossil assemblages from surface sampling of macroscopic
plant remains in mire communities. J. Ecology, 71: 773-791.

GREENE, S. W., and WALTON, D. W., 1975. — An annotated checklist of the sub-antarctic and antarctic vas-
cular flora. Polar Record, 17(110): 473-484.

GRIFFIN, K. O., 1977. — Paleoecological aspects of the Red Lake Peatland, Northern Minnesota. Canadian _/.
Botany 55(2): 172-192.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

90 SEEDS AND FRUITS FROM MACQUARIE ISLAND

HUCKERBY, E., and OLDFIELD, F., 1976. — The Quaternary vegetation history of the French Pays Basque.
II. Plant macrofossils and additional pollen analytical data. New Phytologist 77: 499-526.

MONTGOMERY, F. H., 1977. — Seeds and fruits of plants of Eastern Canada and North-Eastern United States. Toronto
and Buffalo: University of Toronto Press.

Oyana, M., and TAKEHARA, H., 1967. — Revised standard soil color charts.

RAVEN, P. H., and RAVEN, T. E., 1976. — The genus Epilobium (Onagraccae) in Australasia: a systematic and
evolutionary study. Christchurch: New Zealand Department of Scientific and Industrial Research
Bulletin 216.

SELKIRK, D. R., SELKIRK, P. M., and GRIFFIN, K., 1983. — Palynological evidence for Holocene environ-
mental change and uplift on Wireless Hill, Macquarie Island. Proc. Linn. Soc. N.S.W. 107: 1-17.
SELKIRK, P. M., SELKIRK, D. R., and BERGSTROM, D. M., 1984. — Holocene vegetation history of Mac-

quarie Island. Tasmanian Naturalist 78: 21-23.

SEPPELT, R. D., COPSON, G. R., and BRowNn, M. J., 1984. — Vascular flora and vegetation of Macquarie
Island. Tasmanian Naturalist 78: 7-12.

SYSTEMATICS ASSOCIATION COMMITTEE FOR DESCRIPTIVE BIOLOGICAL TERMINOLOGY, 1962. — II. Ter-
minology of simple symmetrical plane shapes (Chart 1). Taxon, 11(5): 145-156.

TAYLOR, B. W., 1955. — The flora, vegetation and soils of Macquarie Island. Australian National Antarctic
Research Expeditions, Reports, series B, vol. 2, Botany. Publication No. 19. 192 pp.

WILSON, K. L., 1981. — A synopsis of the genus Scirpus sens. lat. (Cyperaceae) in Australia. Tlopea 2(2):
153-172.

PROC. LINN. SOC. N.SW., 109 (2), 1986

Notes on the Biology of Adult and
Immature Amycterinae
(Coleoptera, Curculionidae)

ANNE T. HOWDEN

(Communicated by C. N. SMITHERS)

HOowbDEN, A. T. Notes on the biology of adult and immature Amycterinae (Coleoptera,
Curculionidae). Proc. Linn. Soc. N.S.W. 109 (2), 1986: 91-105.

Adults of the endemic Australian weevil subfamily Amycterinae feed primarily on
monocotyledons but a few eat dicotyledons. The varied morphology of the adult
mouthparts is shown to correspond to the texture of the food plant. Species which feed
on wiry stems have stout, blunt mandibles and a pronounced ‘gular roll’; at the opposite
extreme, species which fced on the most tendcr lily leaves have thin-edged slicing man-
dibles and no gular roll, leaving the maxillae and prementum completely exposed.

Notes on the biology of the immature stages of ten genera of amycterines are
reported. Oviposition is apparently in the soil. Larvae of one species feed on the tender
new growth of Xanthorrhoea crowns; other species observed feed on underground stems,
tubers and rhizomes, and possibly roots. Pupation takes place in the soil.

The function of the modified 8th sternite (the ‘forceps’) of male Phalidura is
discussed.

Anne T: Howden, Research Associate, Biology Department, Carleton University, Colonel By Drive,
Ottawa, Ontario KIS 5B6, Canada; manuscript received 14 August 1985, accepted for publication
20 August 1986.

INTRODUCTION

The endemic Australian weevil subfamily Amycterinae includes an estimated 500
species. The weevils are favourites of collectors because of their unusual and often spec-
tacular structural modifications, but their biology was almost unknown as recently as
1970 (Britton, 1970: 619). The few published references to biology concern mainly the
weevils’ hiding places and supposed food sources. Many of the food records, such as Eu-
calyptus bark (Ferguson, 1921: 29, 30) and dry, dead wood (Macleay, 1865: 201) are spuri-
ous. So little is known concerning the biologies of the species that even precise localities
and dates of collection are worthwhile contributions.

Information is recorded herein concerning immature stages of ten genera of
Amycterinae and the adult food of another five genera. In addition, the morphology of
the adult mouthparts of amycterines is discussed as it relates to the texture of their food
plants, and the morphology of the ovipositor is discussed as it relates to method of
oviposition.

METHODS

E. C. Zimmerman, Curator of Weevils Emeritus, Commonwealth Scientific and
Industrial Research Organization, Canberra, assisted with the identifications of the
adults, and I have personally examined many of the types of Blackburn, Ferguson, Lea,
Macleay, Pascoe, Sloane, and Waterhouse. No nomenclatural changes are made in this
paper, because a comprehensive work including the Amycterinae is in preparation by
E. C. Zimmerman. At his suggestion, all references to higher classification are omitted
here because of his anticipated changes to the system. Genera are discussed using the

nomenclature and taxonomic sequence found in the Coleopterorum Catalogus (Schenkling
and Marshall, 1931).

PROC. LINN. SOC. N.S.W., 109 (2), 1986

92 ADULT AND IMMATURE AMYCTERINAE

Immature stages are being described in a separate publication by Brenda May,
Department of Scientific and Industrial Research, Auckland, New Zealand.

Voucher specimens of adult and immature stages will be deposited in the Aus-
tralian National Insect Collection, Canberra, and, where sufficient material exists, in
the New Zealand Arthropod Collection, Auckland, and the Howden collection.

Biological observations were made over a period of nine years during four trips to
Australia, totalling 16 months of field time. These trips were: (1) December 1974 to June
1975, in eastern Australia between Atherton, Queensland, and Canberra, Australian
Capital Territory; July 1975, in southwestern Western Australia. (2) July and August
1978, Sydney, New South Wales; Canberra, Australian Capital Territory; across the
Nullarbor Plain; and the Western Australian districts of South West, Murchison, Gas-
coyne, and Pilbara north to Point Samson. (3) August to November 1981, Sydney, New
South Wales; Western Australian districts of South West and Murchison; Yorke Penin-
sula, South Australia. (4) July and August 1983, Sydney, New South Wales; Western
Australian districts of South West and Murchison; Australian Capital Territory and
New South Wales between Canberra and northwestern New South Wales and adjacent
territory.

THE FOOD AND MOUTHPARTS OF ADULT AMYCTERINAE

Adults feed on a variety of plants, mostly monocotyledons, but also on some
dicotyledons, namely legumes and myrtaceous plants. According to the growth form of
these plants, amycterines feed at ground level (on Arthropodium, Bulbine, etc) to over a
metre above ground (on Acacia, Bosstaea). Table 1 summarizes the observed and sus-
pected food plants of adults and larvae. Note that the classification of these plant taxa
apparently does not closely follow the classification of the weevils at the generic level.
There is, however, a correlation between the texture of the plants and the weevil genera.
If the plant taxa were rearranged according to the relative tenderness of the tissues con-
sumed by the weevils and then compared to the type of mouthparts of the weevil, the
correlation would be striking.

TABLE 1

Plant-Amycterinae Associations

= s
Plant species Location Amycterine species and stage 3 8 si 5
s o = Q,
POACEAE
Astrebla pectinata Tibooburra, Eto 85kmW, NSW — Cubicorrhynchus taurus x
Cenchrus ciliaris 3km NW Pt Augusta, SA Cubicorrhynchus sp. xX
Enneapogon nigricans Pt Augusta and 3km NW, SA Cubicorrhynchus sp. x x?
Eragrostis eriopoda 100km SE Tibooburra, NSW Cubicorrhynchus sp. x?
70-100km NW Tibooburra, NSW = Cubicorrhynchus taurus x
Eragrostis eriopoda? Waka Hsd (90km W Tibooburra),
NSW Cubicorrhynchus taurus x x?
Warri Warri Gate and N, Old Cubicorrhynchus taurus x x?
Nassella trichotoma nr Yass, NSW Phalidura assimilis x x
Phalidura elongata x x
Cubicorrhynchus sp. x x?
Stipa sp. 56km E Kimba, SA Cubicorrhynchus calcaratus x »G
Damboring (N Ballidu), WA Cubicorrhynchus sp.’ x?
Stipa nitida 91km NW Bourke, NSW Cubicorrhynchus sp. nr taurus x x
87km E Wilcannia, NSW Cubicorrhynchus sp. nr taurus xX x4
125km E Wilcannia, NSW Cubicorrhynchus sp. nr taurus? =X x?
Triodia basedowit ikm S tip North West Cape, WA Notonophes auriger x x4
Unidentified grasses nr Yass, NSW Acantholophus spinifer x

PROC. LINN. SOC. N.S.W., 109 (2), 1986

Plant species

POACEAE
Unidentified grasses

CYPERACEAE
Lepidosperma sp. nr
gracile

Lepidosperma sp. nr
viscidum
ECDEIOCOLEACEAE
Ecdevocolea
monostachya
RESTIONACEAE
Lepidobolus
preissianus
LILIACEAE
Gen. indet.
Arthropodium
capillipes

Bulbine alata

Dianella revoluta
Stypandra imbricata
AMARYLLIDACEAE
Doryanthes excelsa
XANTHORRHOEACEAE
Lomandra sp.
Lomandra longifolia
Xanthorrhoea spp.

FABACEAE
Acacia sp.

Bossiaea linophylla
Daviesia teretifolia
Jacksonia folrosa
Jacksonia foliosa?
MYRTACEAE

Melaleuca sp.
Leptospermum sp.

ANNE T. HOWDEN

TABLE 1 (concluded)

Location

Oodlawirra, SA

Mt Horner, WA
Onslow, WA

36km W Yalgoo, WA
Yellowdine, WA
50km NW Yuna, WA

55km S Marvel Loch, WA

80km S Marvel Loch, WA

52km E Kalbarri, 42km NW
Yuna, 65km S Wurarga, WA

90 Mile Tank, WA

Thomas River, WA
28km W Yalgoo, WA

56km S Mullewa, WA

Tibooburra, Menindce, Bourke,
Wilcannia, NSW

Lake Bryde, Mt Madden, WA

52km E Kalbarri, 20km N North-
ampton, 18kmS Wurarga, WA

Mt Madden, WA

Engadine, NSW

Congo, NSW
Clyde Mt, NSW

Atherton, Qld
Engadine, NSW
Glenbrook, NSW

55km E Mullewa, WA
28km W Yalgoo, WA
30km W Pemberton, WA

Cape Le Grand, WA

18km S Wurarga, WA
Hopetoun, WA

90 Mile Tank, WA
Northcliffe, WA

* Identification of larva questionable.

Amycterine species and stage

Cubicorrhynchus sp.
Acantholophus planicollis
Cubicorrhynchus crenicollis
Chniotyphus tibialis
Notonophes sp.

Gen. indet.
Acantholophus maximus
Acantholophus maximus

Talaurinus sp.
Acantholophus transitus
Acantholophus transitus

Acantholophus maximus

Macramycterus draco

Talaurinus echinops
Dialeptopus echinatus
Polycreta metrica
Ennothus fallax

Bubaris pubescens

Mythites sp. indet.

Mythites basalis var. nodosus

Cucullothorax horridus
Acantholophus marshami

Talaurinus stmillimus
Mythites granulatus
Talaurinus rugifer
Talaurinus subvittatus
Acantholophus marshami
Acantholophus marshami

Dialeptopus echinatus
Dialeptopus echinatus
Acantholophus sp. A nr
aureolus
HAyborrhinus prodigus
Acantholophus sp. B nr
aureolus
Sclerorinus sp.
Acantholophus sp. B nr

aureolus

Acantholophus sp.
Gen. indet.

reese shin sd ddd od >t dd adult

RAK KK KM KR KK KM MMM

KK RK KM K

ess

mK RK

xe
ye

pupa

PROC. LINN. SOC. N.S.W., 109 (2), 1986

94 ADULT AND IMMATURE AMYCTERINAE

The mouthparts of adult amycterines exhibit a relatively wide range of form for one
subfamily. The base of the oral cavity and the mandibles especially show modifications
at the generic level. At one extreme, the mandibles of Polycreta metrica Pascoe (Figs 5, 6)
have a thin, blade-like cutting edge, and the ventral surface of the rostrum is not modi-
fied at the base of the oral cavity. The leaves of Arthropodium capillipes on which P. metrica
feeds are as tender as young onion leaves. At the other extreme, the mandibles of
Macramycterus draco Macleay (Fig. 3) are massive and blunt-edged. The base of the oral
cavity is grossly developed into a thick lip or ‘gular roll’ (Dohrn, 1872: 144) which ex-
tends forward 1mm or more to conceal the prementum. The leafless stems of Lepidobolus
preissianus, on which this Macramycterus feeds, are extremely tough and wiry. However,
the stout mandibles of M. draco do not prevent it from also feeding on tender plants such
as ryegrass. The neat little slices of the unknown plant in Fig. 2 were regurgitated by the
M. draco in Fig. 1.

It seems likely that the heavily reinforced base of the oral cavity provides structural
strength for the operation of the mandibles. The thickest lip is associated with the tough-
est food plants; moderate modification consists of a moderate lip with a sinuous edge
and is associated with leathery food plants; and so forth, until the extreme condition in
which there is no apparent modification of the base of the oral cavity associated with the
softest plants. Additional discussions of mouthparts in relation to texture of plants are
found under particular species, especially Yalaurinus, Acantholophus sp. nr. hypoleucus
(Boheman), and Mythites.

Table 1 shows also that all the dicot records refer to adults only. It is possible that
adults have a wider range of food plants and that immature stages are associated only
with monocots. For example, Dialeptopus echinatus Lea in captivity fed and bred on the
lily Arthropodium capillipes but was observed in the field on Acacza and other plants.

GENERAL OBSERVATIONS ON THE LIFE HISTORY OF AMYCTERINAE

Oviposition and morphology of the ovipositor

Oviposition apparently takes place in the soil, as the following evidence indicates.
Numerous attempts to rear confined adults of many species resulted in eggs and a larva
of Dialeptopus echinatus. The large, creamy yellow eggs were dropped on the surface of the
soil, and after two weeks of extreme conditions, the eggs were dead. However, one egg
had apparently been placed in the soil, and a larva was found in the container a month
after the adults were confined. This suggests that dropping the eggs on the surface was a
response to stress or infertile eggs and that placing the egg in the soil was the norm.

Newly eclosed larvae of Bubaris pubescens and Mythites spp. begin feeding on the ex-
terior surface of the underground stem of their food plant and not from a position
already within the stem which would be the case if the egg had been inserted into the
plant tissue.

The morphology of the ovipositor also seems to be consistent with oviposition in the
soil. In amycterines the stylus, near the apex of the coxite dorsally, has evolved into a
free, heavily sclerotized, outwardly-directed blade (Fig. 9). In most species the blade is
crescent-shaped, with or without a tooth on its dorsal edge, and ranges in shape from a
paddle to a simple scoop (Mythites). This same type of free sclerite is found in the South
African subfamily Hipporhininae and in at least some Tanyrhynchinae, but oviposition
has not been observed in these groups either. An analagous, but not homologous, ovi-
positor is found in some North American Leptopiinae which are known to oviposit in
the soil.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

ANNE T. HOWDEN 95

% poms 5 SO
Figs 1-6. Figs 1-3. Macramycterus draco Macleay, male: 1) sunning on twig, 2) slices of plant regurgitated by
specimen in Fig. 1, 3) oblique ventral view of mouthparts. Fig. 4 Cucullothorax horridus Ferguson, female, pro-

file of head and prothorax. Figs 5,6. Polycreta metrica Pascoe, male, mouthparts: 5) anterior view; 6) ventral
view. Scale line = 0.5mm.

Larval stage

With one exception (Acantholophus marshami (Kirby)?), all larvae found to date feed
underground. Species associated with grasses (e.g., Phalidura, Cubicorrhynchus, etc.) feed
on the subterranean crowns and there is no direct evidence of their feeding on roots.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

96 ADULT AND IMMATURE AMYCTERINAE

Figs 7-12. Figs 7, 8. Bubarts pubescens Lea: 7) dorsal view head and rostrum of female with typical dirt encrusta-
tion, 8) oblique ventral view mouthparts of reared male. Fig. 9. Macramycterus sp., apex of female genitalia,
dorsal view. Fig. 10. Talaurinus subvittatus Ferguson, male oblique ventral view of mouthparts. Fig. 11.
Acantholophus sp. nr. hypoleucus (Boh.), female, oblique ventral view of mouthparts. Fig. 12. Mythites basalis var.
nodosus Ferguson, female, ventral view of mouthparts. Scale line = 0.5mm.

Species that feed on Liliales feed on the subterranean stems and tubers or rhizomes, and
again there is no direct evidence of their feeding on roots.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

ANNE T. HOWDEN 97

Pupal stage

Larvae pupate where they have fed or only a few centimetres away (Cubzcorrhynchus
crenicollis Waterhouse, Bubaris pubescens Lea), or they may travel some distance to pupate.
On two occasions, Acantholophus sp. pupae were dug from the centre of a sand track a
metre or more away from any vegetation. The presence of the pupae was indicated by a
slight disturbance on the surface of the soil directly over the vertical cell.

BIOLOGICAL OBSERVATIONS BY SPECIES
Phalidura spp.

Male Phalidura are characterized by a pair of grotesque ‘forceps’ protruding from
the abdomen. These ‘forceps’ of the vernacular are the highly modified male 8th sternite
which is divided into a pair of massive, free, elongate sclerites up to half the length of the
weevil, and which together strongly resemble a pair of forceps. The function of the for-
ceps, . . . so far as one can reasonably infer, is to grip the female right round the hinder
end of her body . . ?(Gahan, 1918: cxxi); However, in repeated observations, copulating
pairs of P elongata Macleay and P. grandis Ferguson never used the forceps in this man-
ner, and furthermore, it seems physically impossible. The apices of the forceps can only
be moved apart by approximately 2mm; the minimum width of Phalidura females is
much greater. The sides of the 7th sternite confine the forceps at the base and prevent
greater lateral movement. The forceps extend into the abdomen by several millimetres
and are prevented from being pulled out any farther by a connecting vertical internal
disc which is approximately 3.5mm in diameter.

The copulatory posture observed in P elongata and P. grandis is as follows. The abdo-
men of the male is extended to the maximum amount so that the massive, heavily sclero-
tized penultimate tergite is directed ventrally and slightly anteriorly; the last tergite is
completely horizontal and anteriorly directed; the forceps are parallel to each other and
likewise extend the maximum distance, their apices almost reaching the female’s
metasternum and presumably pressing upwards. This pressure may be the only func-
tion of the forceps. The forceps also bear species-specific arrangements of setae which
suggests a recognition function (see Baker and Thompson, 1978).

Phalidura assimilis Ferguson and P. elongata Macleay

These species were reported by Barry Moore (1978: 138; and in litt.) breeding and
feeding on Nassella trichotoma near Yass, New South Wales. The larva of P assimilis is
illustrated in Moore (1980: 13).

Phalidura grandis Ferguson
A short series was taken in Tibooburra, New South Wales, in August feeding in a
small yard planted in Kikuyu and other grasses.

Talaurinus spp.

Two species of Talaurinus (rugifer and subvittatus) were observed feeding on Lomandra
longifolia, and a third species of Talaurinus (simillimus) is associated with a Lomandra sp.
The leaves of L. longifolia are very tough and the mouthparts of both species observed
feeding on them are heavily reinforced with a conspicuous gular roll (Fig. 10). The other
food plants listed below for Talaurinus spp. are likewise tough to very tough and the
mouthparts of the weevils are similar to those in Fig. 10.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

98 ADULT AND IMMATURE AMYCTERINAE

Talaurinus echinops (Pascoe)
One specimen was taken at Thomas River, Western Australia, in October in debris
of a Patersonia-like plant.

Talaurinus molossus Pascoe (Sclerorinus in Schenkling and Marshall, 1931)

A series was taken at Newdegate, Western Australia, in July walking on bare
ground at midday.

Talaurinus rugifer (Boisduval)
One male was observed at Clyde Mountain, New South Wales, in April feeding on
a leaf of Lomandra longifolia.

Talaurinus simillimus (Macleay)
One adult, eight larvae, and one pupa were collected by Murray S. Upton at
Congo, New South Wales, in March with Lomandra (B. May, zn litt.).

Talaurinus subvittatus Ferguson
A short series was observed at Atherton, Queensland, in February feeding on
Lomandra longifolia.

Talaurinus sp.
One male from Marvel Loch, Western Australia, in September was associated with
Lepidosperma.

Molochtus gagates Pascoe
A series was taken 23km west of Yalgoo, Western Australia, in July under mulga
logs and under bark of mulga logs on the ground.

Chriotyphus tibsalis Ferguson
A series was taken at Onslow, Western Australia, in July in grass clumps growing in
sand near the beach. One amycterine larva (genus?) was in the same grass.

Chriotyphus sp.
A series was taken at Coral Bay, Western Australia, August, in grass clumps grow-
ing in sand near the beach.

Sclerorinus spp.
A series was taken 18km south of Wurarga, Western Australia, in August at night
after 2200 hours in matted stems of Jacksonia foliosa about 5-10cm above ground level.
Sclerorinus spp. were common on the Nullarbor Plain between Caiguna and
Madura, Western Australia, in August under glasswort.

Macramycterus spp.
Macramycterus species appear to be strictly diurnal in their activities, including
copulating (July and September) and sunning on a perch 45cm above ground (Fig. 1).

Macramycterus draco Macleay

A short series at 90 Mile Tank, Western Australia, in September displayed a typical
feeding behaviour consisting of walking up to a stem of Lepidobolus preissianus, chewing
off the stem near the ground and eating the stem. In captivity a weevil facing down ona

PROC. LINN. SOC. N.S.W., 109 (2), 1986

ANNE T. HOWDEN 99

stem of L. preissianus was observed to cut the stem off beneath itself (about 5cm above the
ground), to cling to the cut portion as it fell to the ground, and to continue feeding on the
freshly cut end, consuming about 2.5cm in 5 minutes. Mouthparts of Macramycterus are
discussed elsewhere in this paper (see p. 94).

Acantholophus spp.

The large genus Acantholophus is highly varied in its biology. There are many more
nocturnal than diurnal observations. Perambulating larvae of Acantholophus are
described below.

Acantholophus sp. A near aureolus (Boheman)

A large series was observed 30km west of Pemberton , Western Australia, between
1930 and 2200 hours in a light rain, feeding on Bosszaea linophylla and two other species of
shrubs at 0.5 to 1.2mm above ground.

Acantholophus sp. B near aureolus (Boheman)

A series was taken at Cape Le Grand, Western Australia, in October on cool over-
cast days under very dense Daviesza teretifolia, well-camouflaged on the ground in the
sparse debris of grey, dead leaves of the same size and colour as the weevils. All speci-
mens regurgitated green matter, presumably Daviesza foliage.

One adult at Esperance, Western Australia, in October was walking at night, in a
temperature near freezing, in the vicinity of Daviesza.

Two specimens were found at Hopetoun, Western Australia, in August on the
ground under Jacksonza sp.

Acantholophus sp. nr hypoleucus (Boheman)
A short series at Arrowsmith River, Western Australia, July, was taken under dead
grass trees; the mouthparts (Fig. 11) indicate food of moderate texture.

Acantholophus marshami (Kirby)

Described by Froggatt (1896: 77) as‘ . . the common Amycterid about the neigh-
bourhood of Sydney. Most of the members of this large genus live upon the grass, but
this one climbs up the leaves of the grass-tree, and clinging round them gnaws pieces
out.

A colony of amycterines at Engadine (40km south of Sydney), New South Wales,
has been monitored for several years by the G. A. Holloway family and is apparently this
species. The Holloways report (zn Jit.) larvae present from early May to mid November.
One male A. marshami (det. A. Howden) was taken 9 May on the crown of a Xanthorrhoea
containing amycterine larvae (det. B. May). The larvae are always in the crown, a metre
or more above ground, in head-up position in a column of chewed matter, and feed on
the innermost tender white leaves. Some larvae were of buff colour instead of white on 1
November; several weeks later all larvae were absent from Xanthorrhoea. Observations
are continuing.

The Holloways also report amycterine larvae from other species of grass trees and
from Glenbrook in the Blue Mountains.

Although the ectophytic behaviour of these larvae appears to be anomalous among
amycterines, it is possibly only an extension of feeding in ground-level crowns of plants.

Three males of A. marshami were taken at Engadine (same site as above) in August
under leaf scales of a standing dead flower stalk of the Gymea lily, Doryanthes excelsa.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

100 ADULT AND IMMATURE AMYCTERINAE

Acantholophus maximus (Macleay)

Single adults and series were taken at four sites between 52km east of Kalbarri and
65km south of Wurarga, Western Australia, in August in clumps of Ecdezocolea
monostachya. A single amycterine larva was excavated at two of the sites, each larva 7.5cm
below the surface of the ground in the crown of a plant.

Acantholophus planicollis Waterhouse
One adult was taken at Oodlawirra, South Australia, in August; nearby grass
yielded six amycterine larvae of unknown genus.

Acantholophus spinifer Macleay
Adults and larvae, according to Moore (1978: 138), are associated with grass near
Yass, New South Wales.

Acantholophus transitus Macleay

A short series was taken 55-80km south of Marvel Loch, Western Australia, in
clumps of Lepidosperma spp. near gracile and near viscidum; adults were also taken in the
vicinity of Lake King and Newdegate in July.

Cubtcorrhynchus spp.

All Cubicorrhynchus species collected to date have been associated with either native
or introduced species of Poaceae except for occasional instances in which the weevils
appeared to be hiding only. Cubicorrhynchus larvae taken from the crowns of grass plants
often regurgitated green material, indicating they had fed on underground stems and
not the roots. Adults are primarily nocturnal.

Cubicorrhynchus calcaratus Macleay
Adult fragments and a larva were taken 56km east of Kimba, South Australia, in
August in a clump of Stzpa sp.

Cubtcorrhynchus crenicollis Waterhouse

A colony near Mt Horner, Western Australia, was located in an unidentified
species of native grass growing in an isolated strip of relatively undisturbed coastal
heath. On 14 July larvae and fragments of adults were excavated from clumps of the
grass. On 9 September, seven larvae and 17 pupae (some with larval exuviae still
attached) were excavated. Pupae were 10-50mm deep in the soil in cylindrical ex-
cavations approximately 7mm in diameter, at the outer edge of the root mass of the
clump.

Three adults were reared from pupae collected 9 September, ecdysis occurring in 5,
34, and 52 days. The first two adults emerged from pupae which had remained on the
surface of the soil in the rearing tin. The third pupa was located in a cell in the bottom of
the rearing tin and was still white 11 days before its ecdysis was artificially accelerated by
increasing the moisture and temperature in the tin. Thus it appears that the duration of
the pupal period varies greatly, and pupae deeper in the soil might be expected to have a
longer pupation because warm temperatures and superficial rains would not reach
them.

The inferred life cycle is thus: larva — July to September, pupa — August to
November, adult emergence — September to November and later.

PROC. LINN. SOC. N.SW., 109 (2), 1986

ANNE T. HOWDEN 101

Cubicorrhynchus taurus Blackburn and related species

Several species may be included in this unit of specimens distinguished by having
the fore coxae contiguous and the hind tibia of the male nodose and produced into a con-
spicuous lobe on the inner edge medially. The range is throughout the northwest corner
of New South Wales and adjacent Queensland and South Australia, specifically,
Tibooburra, southeast to Wilcannia, southwest to Menindee, through Broken Hill and
northwest to Frome Downs. Specimens west of the Flinders Ranges are a different
species (Cubzcorrhynchus sp. from Port Augusta). Eighteen adults from a granite out-
cropping at Tibooburra were smaller and less developed in secondary sexual characters
than specimens from the clay or sandy areas and may be a different species.

Adults and a few larvae collected July through August were associated with a
variety of grasses especially Astrebla pectinata (Mitchell grass) and Eragrostis eriopoda
(woollybutt). Dissected females contained no eggs but an abundance of fat. A sample of
both sexes from various localities contained green plant matter in the digestive tract
although the grasses were only beginning to show green after recent rains.

Caged specimens fed on grass by climbing up the blade and while still facing up,
ate the blade from the tip down, backing down as necessary. One female observed cut
the top 2cm from a blade of grass and carried it back and forth, pausing to feed on it.
Many pieces of cut grass accumulated on the soil of the cage. This weevil cut the grass
blade not with a single cut, but with an up and down rasping motion not observed in
other amycterines. The mandibles and reinforced base of the oral cavity of this species
indicate the ability to cut very tough material.

Cubicorrhynchus sp. near taurus Blackburn

This species resembles C. taurus, but males lack the conspicuous lobe on the hind
tibia; the range is central New South Wales from 87km east of Wilcannia to 68km west
of Cobar and north to 91km northwest of Bourke. At the latter locality one pupa was
found 8cm deep in the ground under Stipa nitida. Larvae (presumably of this species)
were found at several sites in the crowns and root masses of Stipa nitida. Abundant
Eragrostis in the area yielded no amycterine adults or immatures.

Cubicorrhynchus sp. from Port Augusta

This species is related to C. taurus by the contiguous fore coxae, but the hind tibia of
the male is straight for the proximal two-thirds, then strongly curved forward, its inner
edge bearing a conspicuous tooth or prong at the distal fifth. The species was taken in
South Australia from Port Augusta southwest to 9km west of Iron Knob, and north of
Port Augusta to Quorn on the west side of the Flinders Ranges. Adults were taken in late
July and late August in clumps of Cenchrus ciliaris, Enneapogon nigricans, and Stipa nodosa;
larvae were taken in late July in Enneapogon nigricans and Stipa nodosa.

Notonophes auriger Ferguson

Adults were taken at Onslow, Western Australia, in July in debris washed around
the base of grass; adult carcasses were found at the North West Cape in late July near a
larva 3-5cm deep in soil under a tussock of Triodia basedow1t (spinifex).

Notonophes gascoynensis Baker (1972: 123)

Adults at Carnarvon, Western Australia, in August were sheltering under Salzcornia
australis (samphire); the weevils are similar in colour and form to the debris under the
plants.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

102 ADULT AND IMMATURE AMYCTERINAE

Hyborrhinus Marshall (1946: 94) (= Hyborrhynchus Macleay in Shenkling and Marshall,
1931)

Hyborrhinus prodigus (Macleay) was taken at Cape Le Grand, Western Australia, in
October, in debris under Daviesia teretifolia.

Mythites spp.

Two Western Australian species of Mythites (basalis var. nodosus Ferguson and sp. in-
det.) are associated only with Dianella revoluta, spreading flax-lily. The plant is discussed
here because of its importance to Mythites and other amycterines. Old leaves of the plant
remain stiff and upright, and old clumps of Dianella can be a metre or more wide, retain-
ing thick layers of their own litter or windswept debris of Eucalyptus leaves, etc. Occasion-
ally adults of other amycterines are found in the shelter of Dianella revoluta clumps, e.g.,
Acantholophus sp., Acantholophus maximus, Acherres mamillatus Pascoe, Cubicorrhynchus
bohemant (Boheman), Dialeptopus echinatus, Macramycterus sp., and Notonophes cichlodes
(Pascoe). However, all larvae excavated from D. revoluta are Mythites (B. May, zn litt. ).

Mythites adults usually feed by cutting out portions of the leaf margin, but caged
beetles cut off the flower stalk of one D. revoluta at 9cm above ground. Mouthparts of
M. basalts var. nodosus (Fig. 12) indicate a food plant of moderate texture.

Mythites basalts var. nodosus Ferguson

This species was observed and collected in a semicircle around Geraldton from
north of Northampton to the Kalbarri turnoff, east to Wurarga, southeast to 55km east
of Mullewa, south of Mullewa and west to Arrowsmith. Larvae were found 12 July to 10
September, pupae 7 September, adults 7 August through September. In the earliest ob-
servation, newly eclosed larvae 5-8cm below ground level had eaten into the under-
ground Dianella stem from without; in addition, one large larva was found in a rhizome.
In an August examination of a heavily infested Dianella clump, the rhizomes were found
to be extensively excavated by larvae. Two pupae were found 2.5cm deep in the soil of a
D. revoluta clump, one in a vertical cylindrical hole.

Mythites sp. indet.

Adults and larvae were taken at Lake Grace, Lake Bryde, and Mt Madden in
southwestern Western Australia, on four occasions between 22 September and 13
October in clumps of Dianella revoluta. Some larvae were just below the surface of the soil
above the roots and rhizomes. Other larvae had burrowed into the rhizomes and were
feeding in cavities they had excavated from the exterior. Adults were found in all clumps
of Dianella containing larvae.

Mythites granulatus Lea
One female at Clyde Mountain, New South Wales, on the morning of 8 April was
observed feeding on Lomandra longifolia.

Bubarts pubescens Lea

Adults and immature stages were found in August in the large area of red sand
plains of northwestern New South Wales ranging from Menindee north to Tibooburra
and the Queensland border, southeast of Tibooburra towards Wanaaring, from Wilcan-
nia to 87km east of Wilcannia and north to 35km north of Bourke.

Adults and larvae feed on the leek lily, Bulbine alata, a small lily with an onion-like
cluster of tender leaves and a short flower stalk. The roots do not bear tubers, nor is there
a swollen base or bulb.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

ANNE T. HOWDEN 103

The following synopsis is based on hundreds of observations at the many localities
listed above. Newly eclosed larvae eat directly into the base of the underground portion
of the stem, immediately above the roots, usually 4-25mm below the surface of the soil.
Older larvae feed externally on the underground stem while encased in a hollowed con-
cretion of sand grains which adheres to the stem. When injured, the lily stem bleeds a
slightly viscous sap and this seems to cause or contribute to the accretion of the sand.
Usually there is only one larva per plant; infestation by two or three larvae per plant is
uncommon and when it occurs involves only small larvae. By the time larvae leave a
plant, the plant is usually in seed and (1) showing no obvious above-ground evidence of
distress (infrequently), or, (2) at least the outer leaves are dead and pinkish in colour
(commonly), or, (3) the plant is completely dead (infrequently). Older larvae may travel
underground to feed or to pupate; large larvae are sometimes seen feeding on very
small, previously untouched lilies. Pupae and prepupae are found in an earthen cell as
far as 2.5cm from the host plant, but are usually directly beneath the plant and
0.6-15.0cm below the surface of the soil. Pupal cells are approximately 15-20mm long
and 5-6mm wide. No freshly emerged adults were found, but a pupa collected 24 August
was reared to adult in 21 days.

Since Bulbine alata responds very quickly to rain, local showers can have a strong in-
fluence on the availability of food for larvae. By waiting under bushes where the Bulbine
seedlings will first appear, adults can oviposit as soon as suitable lilies are available. As
with other species living in arid and semi-arid country with irregular rains, the life cycle
of Bubaris pubescens is probably influenced more by precipitation than by the calendar.

The naturalized onion weed, Asphodelus fistulosus, is similar in plant form to Bulbine
alata, and the two species may grow intermingled in one area. At Menindee, a large
number of both plants in equal numbers was examined; only the Bulbine was infested
with amycterine larvae. However, a large Bubaris larva confined for five days with only
an Asphodelus plant for food ate a typical spherical hole 3mm in diameter in the base of
the underground stem.

Adults of B pubescens have a dense coat of short, wiry, curled ‘setae’ distributed
everywhere except around the apex of the rostrum and ventral surface of the head (Figs
7, 8). These setae may produce a gummy exudate. The prothorax has closely spaced
cylindrical tubercles 0.3-0.5mm high, and the elytra have similar but fewer tubercles.
The result of this ornamentation is that the weevil becomes encrusted with particles of
soil firmly wedged and glued in place (Fig. 7). This crust is very good camouflage.

Dialeptopus spp.

Dialeptopus have long, slender legs and many of the Western Australian species can
be seen running quickly and conspicuously over the ground in the daylight hours of the
winter months.

Dialeptopus echinatus Lea

Adults are common east and southeast of Geraldton, Western Australia, in the
winter months. Copulating pairs were taken on Acacia at several localities and on a
‘sedge’ Caged beetles fed only on Arthropodium capillipes when given a sample of vegeta-
tion from the Yalgoo area which included Dianella revoluta and Acacia. Arthropodium
capillipes is a delicate lily with fleshy leaves and a tall flower spike appearing in the early
spring. The roots are fibrous and many terminate in small tubers. Leaves were eaten by
the weevils from the margin towards the centre of the leaf. One pair of weevils caged for
a month produced seven eggs, one of which eclosed, and the larva during that month ex-
cavated almost one-quarter of a tuber on the end of a root.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

104 ADULT AND IMMATURE AMYCTERINAE

Dialeptopus plantaris Pascoe
Two very active specimens were taken 82km north of Carnarvon, Western Aus-
tralia, in July on spinifex.

Cucullothorax horridus Ferguson

One female (Fig. 4) was taken at Mt Madden, Western Australia, in July on Sty-
pandra imbricata; additional specimens were taken at 90 Mile Tank (September) and car-
casses of specimens at Marvel Loch.

Sosytelus lobatus Pascoe
One specimen was taken near Sydney, New South Wales, in August walking during
the day.

Ennothus fallax Pascoe — Polycreta metrica Pascoe complex

These two monotypic Western Australian taxa are probably congeneric, Ennothus
being the senior name. Specimens from 56km southwest of Mullewa and from Tunney
are referable to Ennothus fallax. Specimens from 28-29km west of Yalgoo and 55km south
of Marvel Loch are referable to Polycreta metrica. This distribution suggests the range of
Polycreta metrica is in an area of lower rainfall, i.e., outside the wheat belt and with less
than 25cm of rain annually, where Ennothus fallax occurs in slightly higher rainfall areas.

These small, active amycterines, 5.0-8.6mm, are reminiscent of Dialeptopus with
their long legs and distinctly spider-like appearance.

All specimens were associated exclusively with Arthropodium capillipes. Usually the
weevils were concealed deep in the rosette of leaves at the base of the plant or under
debris very close to a lily. Weevils were observed feeding on leaves of the lily, once during
the day (P. metrica) and once at night (FE. fallax). In captivity the weevils ate only Arthro-
podium capillipes. Mouthparts of P. metrica are shown in Figs 5 and 6.

At the Mullewa site in July, three larvae were found 8-10cm deep in the soil around
an Arthropodium plant on which an E. fallax had been feeding. All tubers from the ex-
cavated plant were collected and examined under a microscope. One shell of a tuber
contained parts of an adult male E. fallax; other empty tubers contained pieces of head
capsules and an almost complete larval exuvia.

No immature stages were found in September, but adults were much more numer-
ous then.

CONCLUSIONS

Biological observations have helped to interpret some of the structural modifica-
tions of adult amycterines. The varied morphology of the mouthparts relates to the tex-
ture of the food plant. The unusual stylus of the ovipositor probably relates to
oviposition in the soil.

In addition, two findings indicate that amycterines are a relatively old group,
namely, placing the eggs in the soil (which is plesiomorphous in Curculionidae) and
larvae feeding on monocots (monocots being an older group of angiosperms than
dicots).

The inferred life cycle of the species observed south of the Tropic of Capricorn is as
follows. Eggs are laid in the soil in the winter. Larvae of the majority of species feed on
underground stems (Bubaris, Mythites), on the crown of grasses (Cubzcorrhynchus), on
tubers (Dialeptopus, Ennothus), and on rhizomes (Mythites); larvae of one species feed on
aerial crowns of grass trees (Acantholophus?). Larval development is completed in the
winter or early spring. Pupation occurs in the soil and the duration of the pupal stage in

PROC. LINN. SOC. N.S.W., 109 (2), 1986

ANNE T. HOWDEN 105

arid and semi-arid areas may be influenced more by moisture and temperature than by
the calendar.

ACKNOWLEDGEMENTS

My thanks go to E. C. Zimmerman, Curator of Weevils Emeritus, Commonwealth
Scientific and Industrial Research Organization, Canberra, for first introducing me to
Amycterinae and presenting the challenge of their unknown biology.

Field work especially benefited from the knowledge and assistance of K. and
E. Carnaby, Wilga, Western Australia; and G. A. Holloway, Australian Museum,
Sydney, and his wife and son. Assistance with transportation was received by
H. F. Howden (as principal investigator of a different project) from: the Australian
Museum, Sydney, in 1983; Commonwealth Scientific and Industrial Research Organiz-
ation in Canberra in 1975 and 1978 and in Perth in 1983; and the South Australian
Museum, Adelaide, in 1981.

Plant samples were identified by L. Adams and M. Gray, Commonwealth Scien-
tific and Industrial Research Organization, Canberra; and S. Jacobs and A. Rodd,
Royal Botanic Gardens, Sydney.

Permission to include original, unpublished observations by the following persons
is gratefully acknowledged: G. and J. Holloway, Engadine, New South Wales; B. Moore,
formerly of the Commonwealth Scientific and Industrial Research Organization, Can-
berra; and M. Upton, of the same address.

Scanning electron micrographs were taken by Lewis Ling, Biology Department,
Carleton University, Ottawa, Canada.

References

BAKER, F. H. U., 1972. — A revision of the genus Notonophes Sloane (Coleoptera: Curculionidae: Amy-
cterinae). J. Aust. ent. Soc. 11: 114-128.
, and THOMPSON, R. T., 1978. — A new genus of the tribe Acantholophini (Coleoptera: Curculionidae:
Amycterinae). J. Aust. ent. Soc. 17: 247-255.
BRITTON, E. B., 1970. — Coleoptera (Beetles). In The insects of Australia, pp. 495-518. Melbourne: Melbourne
University Press for CSIRO.
Donen, C. A., 1872. — 4. Zur Gruppe der Amycteriden. Stettin. ent. Ztg. 33 (1872): 143-154.
FERGUSON, E. W., 1921. — Revision of the Amycterides. Part VI. Acantholophus. Proc. Linn. Soc. N.S.W. 46:
19-75.
FRoGGaTT, W. W., 1896. — The entomology of the grass-trees (Xanthorrhoea). Proc. Linn. Soc. N.S.W. 21: 74-87.
Ganan, C. J., 1918. — (The president’s address.) Trans ent. Soc. London 1917: cix-cxxiii.
Mac eay, W. J., 1865. — The genera and species of the Amycterides. Trans ent. Soc. N.S.W. 1: 199-298.
MARSHALL, G. A. K., 1946. — Taxonomic notes on Curculionidae (Col.). Ann. Mag. nat. Hist. (11), 13: 93-98.
Mookrg, B. P., 1978. — Life on forty acres. Faringdon: E. W. Classey Ltd.
——,, 1980. — A guide to the beetles of south-eastern Australia. Aust. ent. Mag. Fasc. 1: 1-20.
SCHENKLING, S., and MARSHALL, G. A. K., 1931. — Amycterinae. In Coleopterorum Catalogus, Pars 116, pp. 1-
39. Berlin: Junk.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

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SMe COMET ia tne Cone

The Circumtropical Barnacle Ztraclitella divisa
(Nilsson-Cantell) (Balanomorpha, ‘letraclitidae):
Cirral Activity and Larval Development

D. T. ANDERSON

ANDERSON, D. T. The circumtropical barnacle Tetraclitella divisa (Nilsson-Cantell)
(Balanomorpha, Tetraclitidae): cirral activity and larval development. Proc. Linn.
Soc. N.S.W. 109 (2), 1986: 107-116.

T. divisa is reported for the first time from Australia (Magnetic Island, north
Queensland). The functional morphology and cirral activity of the species resemble
those of Tetraclitella purpurascens, with emphasis on prolonged cirral extension in response
to moderate water flow. Larvae brooded to the cyprid stage pass through 4 naupliar
stages in the mantle cavity, showing vestigial limb setation and enlarged caudal papilla
development. The cyprid undergoes further differentiation beforc release. The paradox
of circumtropical insular distribution in spite of abbreviated free larval life (as a cyprid
only) is discussed.

D. T. Anderson, School of Biological Sciences, University of Sydney, Australia 2006; manuscript
received 25 March 1986, accepted for publication 17 September 1986.

INTRODUCTION

Tetraclitella divisa is a small tetraclitelline found under and between rocks in low in-
tertidal habitats on tropical shores. The species, first described by Nilsson-Cantell (1921)
from material collected in Sumatra and the Java Sea, was notable as the first balano-
morph known to brood its embryos to the cyprid stage in the mantle cavity. Nilsson-
Cantell gave a brief description of a brooded metanauplius and a description and figure
of the cyprid.

Since its original discovery, 7? divisa has been found in the Hawaiian Islands
(Pilsbry, 1928; Edmondson, 1933; Ross, 1961), Formosa (Hiro, 1939), Kyusyu and
Ryukyu Islands (Utinomi, 1949), South China Sea (Zevina and Tarasov, 1963), tropical
west Africa (Bassindale, 1961; Stubbings, 1967), Dominica (Ross, 1968), India (Ross,
1971), Aldabra and Fii (Foster, 1974). It thus has a circumtropical insular and
occasional mainland distribution. Dense populations may occur, as in Fiji (Foster,
1974), where the species may cover up to 100% of available surfaces. B. A. Foster (pers.
comm.) has also found 7? divisa at Norfolk Island. The present paper reports the first dis-
covery of 7’ divisa from Australia. Brooding to the cyprid stage in the mantle cavity was
confirmed by Hiro (1939) and Ross (1961, 1968). Among other balanomorphs, this
phenomenon has been described only in Solidobalanus masignotus (Henry and
McLaughlin, 1967; Newman and Ross, 1971), and in a small tesseroporan from Hawaii
(Newman and Ross, 1977) and various other Pacific islands (B. A. Foster, W. A.
Newman, pers. comm.).

Some aspects of the anatomy of T° divisa were described by previous authors. The
external appearance, wall plates and opercular plates were variously presented and
figured by Nilsson-Cantell (1921), Hiro (1939), Ross (1961, 1968), Zevina and Tarasov
(1963), Stubbings (1967) and Foster (1974). The individual mouthparts, though not the
oral cone, were described by the same authors with the exception of Ross (1961) and were
well figured by Stubbings (1967). Brief descriptions of the cirri, including podomere
counts, were given by Nilsson-Cantell (1921), Hiro (1939), Zevina and Tarasov (1963)
Stubbings (1967) and Ross (1968). The functional morphology and cirral activities of the
species have not been investigated. Descriptions of development are also very brief,

PROC. LINN. SOC. N.S.W., 109 (2), 1986

108 TETRACLITELLA DIVISA

being confined to the two stages given by Nilsson-Cantell (1921). In the present work, the
opportunity is taken to describe the anatomy of the animal in more detail, to report on
its cirral activities and to examine the sequence of development based on a full series of
stages.

MATERIALS AND METHODS

Specimens were obtained from the undersurface of a coral boulder from the low
intertidal of Picnic Bay, Magnetic Island, north Queensland in August 1982. The boul-
der was collected by Dr John Collins of the James Cook University of North Queens-
land, who presented the animals to me as a seetuenilly occurring but unidentified
species from that habitat.

Observations were made on the living animals, including an investigation of cirral
responses to water currents generated across the aperture by means of a Pasteur pipette.
The specimens were then fixed in 0.5% formalin-seawater (V/V). Anatomical investiga-
tions were carried out by dissection following the methods of Anderson (1981). Develop-
mental stages found in the mantle cavities of 25 specimens were removed and
transferred through alcohol to methyl benzoate, benzene and finally liquid paraffin.
These specimens were studied microscopically as wet whole mounts in liquid paraffin.

RESULTS

The Anatomy of T: divisa

Wall and Operculum. Previous workers have described the wall plates and operculum of 7:
divisa in some detail. A resumé of these descriptions is given here, and illustrated from
the Queensland material, as a preliminary to a more detailed consideration of structure
and function.

The animal is of low conical shape in external view (Fig. 1A), with a basal diameter
of 6-1lmm. The basal margin of the wall is rounded in outline. There is a small,
diamond-shaped to hexagonal orifice with a length about one quarter of the basal
diameter. The four wall plates are distinct, with broad radii on the compound rostrum
and laterals. The radii have horizontal summits and exhibit dark striations indicative of
internal tubes. The external surface is covered by a pale brown chitinous cuticle marked
by close set, imbricating growth lines fringed with fine chitinous hairs. Each plate may
be strengthened by three or more ribs that protrude little or not at all beyond the mar-
gin. The parietes are penetrated by 2-6 rows of longitudinal tubes, quadrangular to hex-
agonal in cross section, filled with greyish mantle tissue. The basis is membranous,
sometimes calcareous around the periphery only, or calcareous throughout. The inter-
nal space enclosed by the wall is small and subcylindrical. The ratio of external to inter-
nal basal diameter is approximately 2.3:1.

The yellowish operculum, set superficially within a shallow sheath, is of typical
tetraclitelline form. In the closed position, the scuta occupy most of the orifice (Fig. 1A).
Each scutum is long rostrocarinally and short apicobasally (Fig. 1B). The external sur-
face has a broad median depression. The internal surface (Fig. 1B) shows poor defi-
nition of the adductor muscle scar and adductor ridge, and there are no crests for the
insertion of the rostral and lateral scutal depressor muscles. The articular ridge and fur-
row, however, remain broad and prominent. The tergum, as in other tetraclitellines, is
short and wide (Fig. 1B), with a broad, shallow tergal spur clearly separated from the
basiscutal angle and 4-8 crests for attachment of the tergal depressor muscles.

Little raising and rostral rotation of the opercular valves is involved in the opening
of the aperture. The main movement is a lateral upswing of the valves on the hinge
formed by the opercular membrane.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

D. T. ANDERSON 109

Fig. 1. T. divisa. A. — External apical view. B. — Right scutum and tergum, anterior view. C. — Basal view.
D. — Body and limbs, right lateral view. ad, adductor scutorum; ads, adductor scutorum scar; 67, branchia; c,
carina; co, oral cone; cr, compound rostrum; /, lateral; /sd, lateral scutal depressor muscle; /sds, lateral scutal
depressor scar; pe, penis; pr, prosoma; rsd, rostral scutal depressor muscle; td, tergal depressor muscle; tds, ter-
gal depressor muscle crests; I-VI, cirri I-VI.

Body, Mantle Cavity and Depressor Muscles. Commensurate with the low profile of the wall
and the small interior space, the body of 7? divisa shows the typical long, low configura-
tion of a tetraclitelline (Fig. 1D) and is small relative to the size of the animal as a whole
(Fig. 1C). The prosoma protrudes well forward of the level of the adductor scutorum, to
occupy the rostral part of the mantle cavity. A pair of long, pointed branchiae lies lateral
to the body and limbs. The paired tergal depressor muscles are large and fill the carinal
end of the mantle cavity. The lateral and rostral scutal depressor muscles are thin.

Cirrt. The cirri are not well described in previous accounts, but conform to the general
tetraclitelline pattern. The maxillipeds (cirri I-III) have short rami, while those of the
captorial cirri (IV-V1) are of moderate length.

Cirrus I (Fig. 2A) has a long, curved protopod and an exopod of 8-10 podomeres,
recurved carinally. The exopod carries apically directed, serrulate setae on the median
face and tip. The shorter endopod of 4-6 podomeres also has serrulate setae, but these
are directed posteriorly. Four long, serrulate setae project postero-apically from the
basis.

Cirrus II (Fig. 2B) has a long protopod but short rami, the exopod having 6-8 and the
endopod 5-7 podomeres. The serrulate setae of the protopod and both rami are directed
anteriorly, except for a sparse, postero-apically directed fringe along the posterior mar-
gin of each ramus. The apical podomere of each ramus also carries several forwardly

PROC. LINN. SOC. N.S.W., 109 (2), 1986

110 TETRACLITELLA DIVISA

PS Se

Fig. 2. T. divisa. A-E. — Left cirri in median view. A. — Cirrus I. B. — Cirrus II. GC. — Cirrus III. D. —
Cirrus V. en, endopod; ex, exopod.

curved, serrate setae with a double row of heavy setules along the concave margin. A few
pappose setae project from the postero-apical corner of the coxa and basis.

Cirrus III (Fig. 2C) is slightly shorter than cirrus II, with a broad protopod as well as
short rami. The exopod and endopod have 6-7 and 5-7 podomeres respectively. Setation
is similar to that of cirrus II, except for an additional intermixture of serrate setae
among the anteriorly pointing, serrulate setae of the endopod.

Cirri IV-VI (Fig. 2D) are typical captorial cirri. Representative podomere numbers are
13/11, 15/13 and 15/14 for the exopod and endopod. The podomeres carry 2 long pairs
and 1 short pair of serrulate setae anterolaterally. The protopod carries sparse anterior
spines.

Mouthparts. The general configuration of the oral cone has not been described for T
divisa. The individual mouthparts have been well described and figured by previous
workers (e.g. Hiro, 1939; Stubbings, 1967).

The oral cone (Figs 3A, 3B) is low and wide and the labrum is only weakly bullate.
In posterior view, the large, obliquely inclined mandibular palps obscure the labrum.
Curved setae on each palp fringe the entrance to the preoral cavity. The short maxillae
are set well on to the dorsal surface of the oral cone and point more posteriorly than
ventrally.

The labrum (Fig. 3C) has a shallow median indentation on the posterior margin,
with 3-4 low protuberances and a fringe of short hairs on either side. The long mandi-
bular palps have serrulate setae apically and a fringe of jointed setae along the median
edge. The mandible (Fig. 3D) is quadridentoid, with a prominent incisor tooth and sub-
sidiary cusps on the 2nd-4th teeth. The molar process carries a group of 4 sharp spines of
variable length. The maxillule (Fig. 3E) has two large and two small spines above a
small notch. 6-8 spines occupy the cutting edge below the notch, and a group of smaller
spines occupies the median angle. The maxilla (Fig. 3F) is typical, with an apical lobe
carrying serrulate setae and a proximoventral lobe with jointed setae.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

D. T. ANDERSON 111

0.2 mm

Fig. 3. T. divisa. A and B. — Posterior and right lateral views of oral cone. C. — Labrum, ventral view. D. —
Right mandible, lateral view. E. — Right maxillule, lateral view. F. — Left maxilla, median view. mb, man-
dible; mp, mandibular palp; mx, maxillule; mxa, maxilla.

Cirral activity in T. divisa

Prior to cirral activity, the small opercular valves swing open at a low angle, with
very little uplift. In still water, the opercular valves are held open and pumping beat
commences. The curled long cirri emerge slightly above the carinal end of the aperture
on each beat. The exopods of the first cirri also emerge, apposed and upright, in front of
the long cirri. The pumping beat becomes more active in response to mechanical vibra-
tion induced by light tapping on the rock surface adjacent to the animal.

The application of a gentle water current across the orifice immediately evokes
prolonged cirral extension. The cirral fan is held upright as long as the water current
flows. Rotation of the extended fan up to 70-80° in either direction occurs 1n response to
water currents impinging on the animal either laterally or carinally. Responses to
stronger currents were not investigated.

Larval development in T: divisa

As mentioned above, only Nilsson-Cantell (1921) described any of the developmen-
tal stages of the larvae brooded in the mantle cavity of 7’ divisa. He distinguished a
metanauplius stage, 0.84mm long, with a rounded body and well developed thoracic
limb rudiments. The larva retains the three pairs of naupliar limbs, but with reduced
setation. The second stage described by Nilsson-Cantell was the cyprid, 0.62mm long,

PROC. LINN. SOC. N.S.W., 109 (2), 1986

112 TETRACLITELLA DIVISA

with a typical bivalve carapace, eyes and antennules, but with a prominent thoraco-
abdominal process.

The present material has yielded a longer sequence of stages which reveal the major
progression of development. The embryos, deposited in the mantle cavity as paired ad-
herent masses, are not embedded in typical gelatinous lamellae. Early embryos are
brownish, yolk-filled and ovoid, with diameters of 450 x 350um. Within the egg mem-
brane, embryonic development proceeds with little increase in volume, yielding
naupliar limb rudiments and a small caudal papilla (Fig. 4A). These rudiments become
more conspicuous, and a median eyespot develops anteriorly. The embryo then hatches
from the egg membrane, freeing the naupliar limbs, and changes slightly in shape
(diameters 450 «x 330ym). The body of the hatched, embryonized nauplius remains
yolk-filled (Fig. 4B). The naupliar limbs are simple, with short, vestigial setae at the tip
of each limb. The antennae are slightly longer than the antennules and mandibles.

ps
fh ye
ne ye cp
A an2 B
ant 0.2mm
dts

Fig. 4. T. divisa. A-D. Developmental stages from the mantle cavity. A. — Embryo before hatching. B. —
Stage I nauplius; C. — Stage II nauplius. D. — Stage III nauplius. an/, antennule; an2, antenna; cp, caudal
papilla; dts, dorsal thoracic spine; fh, frontolateral horn; mb, mandible; fs, posterior shield spine.

The first stage nauplius increases in volume and undergoes a moult to a second
stage (Fig. 4C) in which the caudal papilla becomes more prominent. The antennae and
mandibles develop more clearly bifid tips. At the second moult, the third stage nauplius
(Fig. 4D) shows considerable enlargement of the caudal papilla and a dorsal thoracic
spine, reduction of the posterior shield spine and some reduction in the volume of yolk

PROC. LINN. SOC. N.S.W., 109 (2), 1986

D. T. ANDERSON 113

within the body. The lateral margins of the dorsal shield are now sharply delineated and
the setation of the limbs is reduced. During this stage, the caudal papilla begins to show
signs of segmentation. Another moult then follows, yielding a fourth stage nauplius
(Fig. 5A), the last before the cyprid. The caudal papilla is further enlarged, while the
naupliar limbs show a reduction in the number of vestigial setae. Paired compound eyes
develop anteriorly and paired, setose thoracic limbs are conspicuous in the caudal
papilla. The outline of the body changes towards that of a more typical, late stage
tetraclitid nauplius, with a broad straight front. Internally, the outlines of internal or-
gans become defined, although much yolk remains.

Fig. 5. T. divisa. A-C. — Later developmental stages from the mantle cavity. A. — Stage IV nauplius; B. —
Early cyprid. C. — Late cyprid. anJ, antennule; an2, antenna; ca, caudal appendage; cam, cyprid adductor
muscle; cg, cement gland; coe, compound eye; Ja, labrum; mb, mandible; mx, maxillule; mxa, maxilla; ¢/,
thoracic limbs.

The young cyprid, apart from the continuing yolk content, is of the usual cyprid
structure (Fig. 5B). As the cyprid continues its development and the yolk is used up,
(Fig. 5C), the 6 pairs of setose thoracic limbs and their musculature become fully
differentiated and the compound eyes and cement glands become more conspicuous,
but there is still little sign of development of the gut. The 3 pairs of mouthpart rudiments
are obvious but not functionally differentiated. Cyprids must escape the mantle cavity
at this stage, but it seems likely that free-swimming life is brief. Little reserve material
remains within the larva, and feeding is not possible.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

114 TETRACLITELLA DIVISA

DISCUSSION

Functional Morphology. Anderson and Anderson (1985), studying Tetraclitella purpurascens,
drew attention to the many generalized balanomorph features retained in the
Tetraclitellinae, some of which had already been recognized by Ross (1969). They in-
clude a low conical profile maintained by diametric growth: associated broad radii;
weak articulation of the wall plates; a flattish operculum with large tergal depressor
muscles, small scutal depressor muscles and no scutal depressor muscle scars; a small
mantle cavity with little free space for respiratory flow; an elongate prosoma and thorax
with limbs in serial array; short maxillipeds, but a high, narrow oral cone. Anderson
and Anderson also showed that the diametric growth of the tetraclitelline wall is based
on a unique specialization, open-sided tubiferous parietes capable of circumferential
growth. This feature was figured by Hiro (1939) but its significance in promoting rapid
diametric growth has only recently been appreciated.

The same pattern of morphological organization, except for a lower profile to the
oral cone, is displayed by Yetraclitella divisa, ata much smaller maximum size than T. pur-
purascens. Concomitantly, the two species share a common mode of opercular and cirral
activity. Anderson and Buckle (1983) observed that T’ purpurascens performs respiratory
pumping beat in still and slowly flowing water, and responds to moderate external water
flow by prolonged cirral extension. Anderson and Anderson (1985) found that the oper-
cular movements associated with these cirral activities in 7’ purpurascens are simple, in-
volving a hinge-like opening and closing action on a thick opercular membrane, with
only moderate uplift. 7’ divisa exhibits similar opercular movements, performs respir-
atory pumping beat in still water and enters into prolonged cirral extension in response
to slow to moderate external water flow. Both species also have a limited rotational
capacity of the extended cirral fan, to not more than 90° in either direction.

Thus in spite of obvious differences in size and distribution, 7? divisa and T. pur-
purascens epitomize a functional organization that is probably characteristic of all
tetraclitelline species, combining a generalized balanomorph pattern of respiration and
feeding with a hypobiontic habit and a capacity for rapid diametric growth. Further
studies on other species of VYétraclitella from this point of view would be of particular
interest.

Larval Development. In the light of the morphological conservatism of the tetraclitellines,
the unusual modifications of larval development in 7: divisa are particularly striking.
Embryonic and larval development described in some other species of the genus follow
the characteristic balanomorph mode, with eggs of moderate size (approx. 200um)
hatching as typical stage I nauplii and passing through the usual sequence of six plank-
totrophic naupliar stages followed by a cyprid stage (Anderson, 1969; Karande, 1974,
1982; Barker, 1976; Egan and Anderson, in preparation). 7! divisa has a secondarily en-
larged, yolky egg (450 x 350um), hatches as a lecithotrophic stage I nauplius with ves-
tigial setation, remains within the mantle cavity throughout its lecithotrophic
development to the cyprid stage and is free-swimming only as a cyprid. During this
brooded development, the number of moults is reduced, yielding only four naupliar
stages before the cyprid. The naupliar limb setation remains vestigial in all naupliar
stages, and the main emphasis is on development of a large caudal papilla. Some yolk
still remains when the moult to the cyprid takes place, and the cyprid undergoes con-
siderable further differentiation at the expense of this yolk, including functional elab-
oration of the antennules, the thoracic limbs and their musculature, and the cement
glands, before reaching the stage of release. Development of the cyprid organization 1s
thus a more direct and gradual process in 7? divisa than in typical planktotrophic balano-
morph larvae. Convergently similar modifications towards direct development through

PROC. LINN. SOC. N.SW., 109 (2), 1986

D. T. ANDERSON 115

a reduced number of lecithotrophic naupliar stages, followed by release from the mantle
cavity at the cyprid stage, have been noted in the lepadomorph Jb/a idiotica (Batham,
1945) and in most Acrothoracica (Turquier, 1972, 1985). The developmental sequences
in the two other balanomorphs known to release their larvae at the cyprid stage have not
yet been described.

Distribution. As pointed out by Newman and Ross (1977), the distribution of 7’ divisa
presents a paradox. The species is circumtropical and mainly insular, occurring as a
series of widely separated, geographically isolated, intertidal populations. At the same
time its planktonic larval phase is very abbreviated. The present work indicates that the
cyprid retains no yolk reserves when released, and is like other cyprids in being unable
to feed. Settlement in the vicinity of the adult population is thus favoured, resulting in
the known build-up of dense local populations; but dispersal as planktonic larvae over
long distances is not possible. The circumtropical insular distribution therefore suggests
oceanic transport on floating objects, for which some evidence was provided by New-
man and Ross (1977). On arrival at a suitable location, the mode of cyprid release would
facilitate rapid colonization, in a manner analogous to that of many ascidians with
brood retention and abbreviated larval life. The selective forces that led a few, scattered
balanomorph species into this mode, however, with its increased egg size and conse-
quent modifications towards direct development, remain a mystery.

ACKNOWLEDGEMENTS

I wish to thank Dr J. Collins for providing the specimens on which this study was
based; Assoc. Prof. R. Kenny of the Zoology Dept, James Cook University of North
Queensland for provision of laboratory facilities in that Department; Dr B. A. Foster for
confirmation of the specific identity of the material and my wife Joanne for technical
assistance in all aspects of the investigation. This research was supported by research
grants from the University of Sydney and the Australian Research Grants Scheme.

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PROC. LINN. SOC. N.SW., 109 (2), 1986

Metagenesis as a Possible Key to Animal Form

JEAN J. CARTER

CARTER, J. J. Metagenesis as a possible key to animal form. Proc. Linn. Soc. N.S.W. 109
(2), 1986: 117-128.

The origin of metazoan form is attributed to the evolutionary modification of a
metagenic life cycle, a model for which is proposed, based on two alternating diploid en-
tities, here termed trvoph and gone. These are expressed in the Hydrozoa and Scyphozoa
in their most readily identifiable form as independent polyps and mcdusae. Budding of
the troph and the strobilation of the gone are seen as characteristic expressions of asexual
reproduction, by which means each entity replicates its own generation. The formation
of the gone by the troph, however, involves a change in the nature of the unit and addi-
tional trophs can only be formed by way of a zygote. The integration of the two entities
(i.e. troph and gone or troph and gone plus strobilus) is considered as the underlying cause
of the development of thc triploblastic and coclomate condition and the role of the 1D
cell is interpreted as the ‘gone primordium’ This integration is termed syngenean.

Recognition of the troph- and gone-derived elements in segmented protostomes
identifies the adult mandibular segment as the first primary gone (G1). The effects on
cephalization of reduction of the tropA, of its invasion by gone tissue and of use of the troph
or gone mouth by the adult, are considered.

Jean J. Carter, 8 Scott Street, Maroubra Bay, Australia 2035, manuscript received 22 July 1986,
accepted for publication 17 September 1986.

INTRODUCTION

Metagenesis is the term for a life cycle in which an asexually reproducing (bud-
ding) generation, arising from a zygote, alternates with a sexually reproducing gener-
ation (Haeckel, 1866). In the animal kingdom, metagenesis is a phenomenon that
occurs so sporadically and in so small a number of seemingly unrelated groups of lower
metazoans, that little phylogenetic significance has been attached to it beyond its
relevance to the phylogeny of the cnidarians. Early writers on the subject (Steenstrup,
1845; Allman, 1864; Haeckel, 1866) recognized that, in its basic form, metagenesis con-
sisted of an alternation of a ‘vegetative’, ‘nurse’ or ‘feeding’ generation and a ‘fruiting’ or
gamete-producing generation. Later, others (Strasburger, 1894; Bower, 1908) character-
ized the alternating generations in plants in terms of chromosome numbers and identi-
fied the initiation of the gametophyte generation with the occurrence of meiosis and,
although its value as the basis for a generalization was challenged from time to time
(Svedelius, 1927), its precision and convenience have led to its wide acceptance as a
working hypothesis (cf. Weier et a/., 1974).

The discovery of apogamy in pteridophytes, although confusing the issue (Lang,
1898; Farmer and Digby, 1907), has since been attributed to certain irregular mitotic
figures (Steil, 1939) and accommodated as a secondary phenomenon. Although the
generations are sometimes viewed as subdivisions of a single cycle (Darlington, 1978),
there can be no doubt that a vegetative (or asexually reproducing) phase is followed by a
sexual phase in the life cycles of all plants.

Zoologists, confronted by alternating asexual and sexual stages in the lower in-
vertebrates, in which the haploid cells are restricted to the gonads and gametes (Camp-
bell, 1974), were unable to establish an equivalence between the life cycles of the lower
plants and animals. Many workers, following Brooks (1886), have favoured the view that
metagenesis in cnidarians is a secondary condition and discussion has centred on

whether the ancestral form was polypoid or medusoid, sessile or free (Thiel, 1966;
Campbell, 1974; see also Boudreaux, 1979: 17, 18).

PROC. LINN. SOC. N.S.W., 109 (2), 1986

118 METAGENESIS AND ANIMAL FORM

The interpretation of animal form presented in this paper depends largely on
recognition of the universality of potential polypoid and medusoid phases, or their
equivalents, in that order, in the life cycles of all metazoans and it attempts to follow
their separate evolutionary progress in representative types within the Metameria. It
could be argued that there is room for a new approach since no comprehensive theory of
the relationships between the invertebrate phyla has emerged from two recent symposia
of the Systematics Association (House, 1979; Conway Morris, 1985).

In hydrozoans, the polyps, gonostyles and medusae which succeed each other as
distinct morphological entities, retain continuity of the body layers as the polypoid
generation buds to form the medusoid generation. Though some doubt still exists about
the metagenic status of the (young) scyphopolyp and (older) scyphistoma, consensus of
opinion favours a polypoid interpretation (Campbell, 1974; Thiel, 1966; Chapman,
1966). The direct transformation of the strobilating scyphistoma into ephyrae (e.g. in
Aurelia) and the development of gonads in the lucernarian scyphopolyp (Hornell, 1893;
Hyman, 1940: 509), however, demonstrate that ultimately a specific region must be-
come medusoid in nature. In the normal process of ephyra production, the apical disc
or, on occasion, the first three discs, either degenerates or resumes a polypoid existence
(Thiel, 1966: 16-20). The pedal disc can also be regarded as part of the polypoid tissue,
since it reverts to a typical polyp after the liberation of the last ephyra and a scyphistoma
has been figured with a well-developed ring of polyp-like tentacles at the base of a stack
of developing ephyrae (Agassiz, 1860). As in hydrozoans, the scyphozoan medusa is
formed from a population of cells by transformation of part of the body wall. Campbell
(1974) has pointed out that because of this and because of the diploid nature of the
medusa as a whole, some workers feel that the polyp should be considered as a larval
phase. This continuity of body layers from one ‘generation’ to the next would be com-
parable with the condition in pteridophytes if the ‘gametophyte generatiom had been de-
fined to include the diploid tissue of the sporangia as well as the haploid prothallus
formed from spore germination. In higher plants, the equivalent of the gone generation
in animals would then be found in the stamens and carpels, rather than in the haploid
tissue alone. In other words, the haploid phase would represent only part of the ‘game-
tophyte generation. The implication of this alternative interpretation is that, whereas in
the plant kingdom the sporophyte or asexual generation forms the conspicuous vegeta-
tive unit, in animals it is the sexual generation that has become dominant in body form
(see below) and the analogy of the asexual polypoid phase should be sought in the pre-
metamorphic ‘primary marine larvae’ (Jagersten, 1972) or their embryonalized
equivalents.

The precocious formation of the sexual entity and the concurrent embryonic de-
velopment of both phases of the life cycle offer a simple explanation for both the trans-
formation of the vegetative polyp into part of an integrated and usually motile organism
and for the increasing complexity of some tissues and organ systems. It would appear
that, as in the higher plants, the identity of the two units in animals may have become
progressively obscured during evolution as their tissues became more intimately
associated. The relative dominance of the polypoid versus medusoid elements has sub-
sequently contributed a fundamental though simple source of variation at the higher
taxonomic levels.

THE SYNGENEAN MODEL

To challenge this concept, a new approach to morphology is proposed. In the
present discussion, the anatomy of selected types is related to a series of models for
which the following terminology is introduced (see Fig. 1). The term troph refers to the

PROC. LINN. SOC. N.S.W., 109 (2), 1986

JEAN J. CARTER 119

Fig. 1. Diagrammatic representation of methods of asexual reproduction. a. Troph. b. Asexual reproduction of
troph by budding. c. Asexual reproduction of troph by alternative budding method. d. Metagenic formation of
gone by troph. e. Gone independent of troph. f. Asexual production of gones by budding. g. Asexual production of
gones (strobes) by strobilation. h. Liberation and maturation of strobes and re-settling of apical troph. i. Stro-
bilated syngenean individual.

asexually reproducing, polypoid form or phase of the life cycle or to a primary larva of
_ the trochophore (or related marine group) or to its embryonalized equivalent. The term
gone is used for an individual of the gamete-forming, medusoid generation, or for an
equivalent entity in the sexual generation of the life cycle. The term syngenean refers to
single organisms formed by the integration of the troph entity with one or more gone enti-
ties and the term strobilus is used here for a segmented series and the term strobe refers to a
single unit in the series of gone entities that may or may not have secondarily lost its
primary reproductive function and may or may not have become tagmatized.

Hyman’s (1951) rejection of the scyphistoma as a candidate for the ancestry of the
Metameria was based partly on the difficulty of homologizing the head and anal

PROC. LINN. SOC. N.S.W., 109 (2), 1986

120 METAGENESIS AND ANIMAL FORM

‘segments with the other body segments and partly on the assumption that ‘a free-
swimming animal reproducing in this manner would lose its head at each fission’
(Hyman, 1951: 30). Clark’s account of the corm theory (Clark, 1964: 22), like Hyman’s,
does not recognize a metagenic difference between the oral and pedal discs and the in-
tervening ephyrae. Neither does it distinguish conceptually between the strobilus of a
scyphistoma, the stolons of syllids, the chains of ‘budding’ turbellarians and the serial
proglottids of cestodes. The metagenic status of each of these structures needs to be
better understood because they can only be homologous if each represents the same unit
or block of units.

THE COELOM AND PRIMARY GERM LAYERS

Before attempting to establish the equivalence or otherwise of these units or blocks
of units, the nature of the coelom and germ layers and the position of the mouth relative
to the blastopore are reconsidered. The presence of a coelom is a definitive adult charac-
teristic of all taxa with a primary marine larva (1.e. trochophore, nauplius, actinotrocha,
cyphonautes, tornaria and auricularia). By contrast, the acoelomate condition is so
characteristic of the carnivorous, coprophagous or parasitic platyhelminths and
nemathelminths that one must ask whether this is, in fact, the primitive condition or
whether it was initially related in some way to a more advanced or symbiotic way of life.

There has been considerable interest in the method of formation of the coelom (as a
schizocoel or an enterocoel) and its evolutionary origin (as a gonocoel, nephrocoel or en-
terocoel) (Clark, 1964). Discussion of these aspects, however, has failed to emphasize
that, despite the mechanics and phylogeny of its origin, its ultimate relationship to the
primary organ systems is remarkably constant. As with the formation of medusae in
cnidarians, the manner of their origin may vary (contrast Obelia and Aurelia), but the
results are clearly comparable.

By temporarily lessening the emphasis on the mechanism of formation in favour of
its ultimate expression, we could hypothesize a situation in which a tvoph and a gone, each
with its own ectoderm, endoderm and mesenchyme, are integrated in such a way that
the corresponding ectodermal and endodermal layers of each become confluent or inter-
mingled. In this condition, the mesenchyme of troph and gone would be equivalent,
despite possible differences in their origin and the primary cavity within the gone would
be equivalent to that of the troph and therefore to its blastocoel. Whereas the haemocoel
is recognized to be a persistent blastocoel (i.e. troph cavity), the coelom has not pre-
viously been considered as its counterpart in the gone generation. If the troph and gone are
separate entities or the troph has been reduced, a cavity in the gone mesenchyme will not
be recognized as a coelom and an acoelomate condition will result.

Although the relationship of the three primary layers of the troph to their counter-
parts in the gone unit will be influenced by the relative degree of development of each and
by their physical disposition, it is to be expected that some association or co-ordination
will occur between functionally equivalent units of the two generations. It will be evi-
dent that although this interpretation accepts the existence of three primary germ layers,
ectoderm, endoderm and mesenchyme (including mesoderm), it modifies the concept of
each by recognizing their possible dual origin. This may help explain the invasion of the
annelid prostomium and its appendages by the trunk mesoderm (cf. Akesson, 1968: 215)
and the histolysis and cell migration that characterizes the pre-oral tissues of arthropods
during metamorphosis (Anderson, 1966, 1973; Green, 1971; Manton, 1928), as well as
the dual embryological origin and physiological control of some organ systems.
Willmer’s (1970) concept of the two cell-types (epithelium and mesohyl), instead of the
traditional three, is equally applicable to the syngenean interpretation and reflects the

PROC. LINN. SOC. N.S.W., 109 (2), 1986

JEAN J. CARTER 121

CLEAVAGE NUMBER | ist. | 2nd.

NUMBER OF CELLS | 2 | 4 |

a '
ta “—— apical cell 1
. '

I falco annelidan cross cell

2 !
molluscan cross cell

! ta2! | | | 1 a2"
212
| | ! ! Hoa trochoblasts
149220 1 | lta
! ] 1 1 ' | 222
2a ' ' ; 1a
(ee D 2a2 left somatoblast |
? !
|

I
!
oo aie endomere
1
tb" apical cell ,
1b'!@ +— annelidan cross cell

G j
molluscan cross cell
1 !

trochoblasts

2b!

! ' !
median somatoblast
|

endomere

1 3b 1 2b

1°3—B +—
1 ! 1 1
ic'!!+— apical cell ,
4c 2 +— annelidan cross cell
1

molluscan cross cell

Fertilized ovum
1c"

jcl2
1c21
4c 212
1c 221
222

if
trochoblasts

I Ic
1 '

\
is ' ' !
tight somatoblast !
!

mt 0 '
1d —— apical cell ;
1d"? 4 annelidan cross cell
1d molluscan cross cell
Q : 421

I
I
!
|
\
3C —+—— endomere ! I
1
!
I

1
\ \ 1
! ! ' 4d trochoblasts
| I 14221

! I ! ' ! L teloblasts of

r ! I 1 1 x first
X xX EN Xx +——_<X R

I 1 I 1 X somatoblast
aa 1 se gS
'
3d second somatoblast
\ 1
7 , 1 4d(=M) QM , teloblasts of
3D 1 MR mesodermal bands

\ \ 1 4D -~— endomere

Fig. 2. Cell lineage of Nereis (Polychaeta) (after Okada, 1968). Note the idiosynchronous division of the 1D
cell, postulated here as the gone primordium.

dissatisfaction with the conventional interpretation of the germ layers (Oppenheimer,
1940).

Detailed studies of embryonic cell lineages in the protostomatous phyla (Wilson,
1892; Cather, 1971; Clement, 1971; Akesson, 1962; Kumé and Dan, 1968), which
characteristically exhibit spiral cleavage, have shown that while the behaviour and fate
of the A, B, C and 1d cells are comparable, that of the 1D lineage is unique (Fig. 2). Not
only is the timing of its divisions unrelated to the synchrony of the other cells, but its
derivatives contribute to the final structure in a unique manner. The attention that has
been directed to 4d (or M), as the ‘mother cell’ of the mesoderm and the sex cells, has ob-
scured the range and sum of the other 1D derivatives. In Nereis, 2d (=X) is the teloblast
of the ectoderm of the first trunk segment and also forms the proctodaeum; 3d con-
tributes to the posterior wall of the stomodaeum; 4d gives rise to some of the mesodermal
elements of the mid-gut as well as to the mesoblast (M) with its coelom, and the 4D con-
tributes endoderm to and behind the posterior pharyngeal region tissues. These 1D

PROC. LINN. SOC. N.S.W., 109 (2), 1986

122 METAGENESIS AND ANIMAL FORM

derivatives collectively fulfil the requirements of the first medusa-equivalent (or the
primary gone) and succeeding units of the strobilus.

Comparison of the cell lineages of the polychaete Nereis, the cirripede Mitella and
the phyllopod Polyphemus reveals a general similarity. From these examples, the differ-
ence between the annelid and crustacean lineages appears to be that the D lineage
achieves a ‘potentially self-sufficient’ identity, comprising ectoderm, endoderm, meso-
derm and germ cells after the third cleavage in Nereis (1.e. the 1D cell) (Okada, 1968) and
after the second, the D!! (i.e. the D cell) (Shiino, 1968), in the crustaceans Polyphemus and
Mitella. However, the origin of the endoblast and primordial germ cell of Polyphemus is
comparable with Nereis rather than with Mitella (see Fig. 2), suggesting that variation in
the details of cell lineage within a phylum (in this case Arthropoda) may be as great as
that between related phyla.

Prior to metamorphosis, the nereid trochophore (with the exception of the 1D
derivatives) can be compared with a ¢roph or a coelenterate polyp which has transformed
its mesogloea into mesenchyme. After metamorphosis, the original troph tissue is
separated by the vigorous growth of the 1D derivatives, so that the pre-oral region and
pygidium assume a relationship to the entire organism comparable with that of the oral
and pedal regions of a scyphistoma.

Let us, therefore, consider the consequences of the proposition that the triploblas-
tic, coelomate condition owes its origin to the development of the gone (identifiable in
Nereis as the 1D lineage, see Fig. 2) in intimate association with the troph, formed from
the derivatives of cells A, B, C and 1d.

If each gone has the metagenic status, function and anatomical potential of a
medusa and the strobilus is essentially a stack of such medusae, each strobe will retain a
potential self-sufficiency until this is irreversibly changed by evolution resulting in a
division of labour between strobes. With progressive differentiation of a strobilus, it is to
be expected that the leading units will have developed the gone-orientated sensing and
feeding systems to a greater degree than those following. By recognizing the first post-
metamorphic (i.e. the most anterior trunk or jaw-bearing) segment in Nereis as Gl (see
Table 1), the dual origin of the adult mouth and pharynx becomes clear. The cephaliz-
ation of G2 and G3 in arthropods makes the evolution of more specialized mouth parts
possible. The doubt that surrounds the homology of the peristomium in polychaetes
(Schroeder and Hermans, 1975) is largely due to conflicting definitions. These doubts
can be resolved, in part, by recognizing the errant polychaetes to be annelids with a well-
developed and persistent troph and the sedentarians to be forms in which the ¢roph is
reduced. The role of the peristomium as the sensory and oral region of Gl (cf. the
manubrium of the first medusa) is supported by the definition of Schroeder and Her-
mans (1975) and by embryological evidence. The peristomial appendages, together with
the sub-oesophageal ganglion and invasive mesohyl which supplements the pre-
metamorphic and pre-oral structures, can be regarded as elements of the first or
primary gone. The reciprocal growth of axons to and from the supra- and sub-
oesophageal ganglia also support the concept of this gone actively establishing oper-
ational partnership with the troph (cf. Henry, 1947).

Once the positional and functional relationships between troph and gone are recog-
nized, it becomes possible to interpret the proto- and deuterostomatous conditions, ir-
respective of assumed phylogenetic relationships. This will be deferred, however, until
certain aspects of segmentation, stolonization and tagmatization have been considered.

SEGMENTATION, TAGMATIZATION AND STOLONIZATION

The segmentation of annelids and arthropods is almost ubiquitous and widely

PROC. LINN. SOC. N.S.W., 109 (2), 1986

JEAN J. CARTER 123

regarded to be a primitive feature of both groups. Its partial obliteration in some arthro-
pods may occur during maturation, as in the parasitic crustaceans, or during the em-
bryonic stages, as in the acarine and araneid arachnids. In all these cases, the lack of
segmentation is clearly secondary. Tagmatization usually accompanies and possibly
precedes a loss of segmentation. It is characteristic of a specific functional specialization
such as feeding, locomotion or reproduction, the obliteration of segmentation being
commonly associated with the reproductive region of the body.

There is much variation and some uncertainty about the precise origin of the germ
cells in the various arthropod taxa. They have been found in representative polychaetes,
crustaceans and insects in all segments of the strobilus, including the mandibular (G1),
but not in the pre-oral or caudal ‘segments’ (Green, 1971; Anderson, 1973; Snodgrass,
1935; Bitsch, 1973). Because these exceptions coincide with the postulated troph tissues,
it is concluded that the gone or the strobilus itself is the potentially fertile unit and that
evolution has favoured the migration of the germ cells to restricted regions or meta-
meres, usually those posterior to the stomach and digestive glands. Variability in the
position of the genital ducts in the lower crustaceans and, therefore, in the extent of the
fertile (cephalo-) thorax, contrasts sharply with the rigid patterns that characterize the
malacostracans, insects, myriapods and chelicerates. In the oligochaetes, the sterile
region posterior to the fertile tagma is usually reduced or secondarily modified.

TABLE 1

Segmental equivalence of the anterior appendages of the Metamerta based on the syngenean interpretation of animal form

SYNGENEAN ANNELIDA ARTHROPODA
INIT POLYCHAETA CRUSTACEA INSECTA ARACHNIDA

ANTENNULES
PHOSTOMIAL

TENTACLES ANTENNAE ANTENNAE

AND PALPS
LARVAL MANDIBLES INTERCALARY SEGMENT

PERISTOMIAL TENTACLES
AND PHARYNGEAL JAWS | ADULT MANDIBLES ADULT MANDIBLES CHELICERAE

FIRST MAXILLA FIRST MAXILLA PEDIPALPS

SECOND MAXILLA SECOND MAXILLA

3
APPENDAGES
OF i ist.
TRUNK WALKING LEGS

: : 2nd. (OR :
SEGMENTS
MAXILLIPEDS i BEGS BES

PROSOMAL
THORACIC ( WALKING )

T: troph; G: gone

By identifying, as the essential feature of the onset of metamorphosis in crus-
taceans, the simultaneous histolysis of the pre-oral musculature of the nauplius, its
replacement by invasive somitic mesoderm and the development of the adult elements of
the mandibular segment, the nauplius can serve as a point of reference for the morpho-
genesis of the adult jaw-bearing segment of both annelids and the egg-nauplius. The
entity G1 (or the ‘primary gone’) and the subsequent strobes (G2, G3 etc.) can then be
identified and their tagmatization compared without the necessity for establishing
homology of the pre-oral tagmata. The advantages of this are evident in interpreting the
homologies of tagmata in arachnids and onychophorans where pre-oral (or ¢troph-
equivalent) structures appear to be reduced.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

124 METAGENESIS AND ANIMAL FORM

By adopting the nauplius in preference to the trochophore as a ‘standard’ or refer-
ence for the fully-developed ‘primary larva’ of the protostomatous phyla, the three paired
appendages (antennules, antennae and /arval mandibles), together with the proto-,
deuto- and trito-cerebra, the larval eyes and the larval alimentary system (stomodaeum,
larval stomach and digestive glands and the anterolateral region of the adult pharynx),
can be identified as parts of the mature troph. In Crustacea, the replacement of larval by
adult appendage musculature, like the development of pharyngeal jaws in the first trunk
segment of polychaetes, is due to the activity of cells derived from the mesodermal telob-
lasts and ultimately to the 1D lineage (cf. Manton, 1928; Green, 1971; Anderson, 1973).

The pharyngeal region of annelids and the (adult) mandibular segment of crus-
taceans appear to be structures of dual origin, derived in part from the troph and in part
from the gone, the development of each component varying with the taxon, but those of
the gone invariably developing after those of the troph.

In insects, the intercalary segment, now widely accepted as being innervated by the
tritocerebrum (Bitsch, 1973; Rempel, 1975) would, by the same reasoning, be part of the
troph and the equivalent of the larval mandibular region of a nauplius. In the
holometabolous insects, it seems to be the sudden burst of growth of gone tissue, initially
and precociously formed during the early embryonic period, but latent during the
secondary larval (‘caterpillar’) stage, which causes metamorphosis by completing the
differentiation of each strobe and by modifying and replacing some of the trop/’s charac-
teristic structures — e.g. the larval eye spots and simple pre-oral appendages. In the
mosquito, for example, the larval feeding brushes are replaced at metamorphosis by the
highly specialized structures that form the feeding mandibles. In orthopterans, cells
derived from a mesodermal somite migrate into the rudimentary antennae and prolifer-
ate to form the segmented adult antennae (Wheeler, 1893: 111). Although the 1D cell is
not identifiable as such in insects, the somitic mesoderm clearly belongs to the strobilus.
As such, it can be interpreted as another instance of the integration of equivalent tissues
of the two generations to form a single functional structure. Although the spiral cleavage
and, therefore, the possibility of recognizing the 1D lineage is replaced by superficial
cleavage in the peracaridean crustaceans and the insects, there seems to be no reason to
doubt the homology of the mandibular and strobilar regions of either group with their
counterparts in the lower crustaceans if the adult mandibular segment is G1 in all cases.
Here again, we see that although the details of differentiation may differ, the results are
closely comparable because no one doubts the homology of isopod structures with their
counterparts in other crustaceans.

Table 1 shows the strobilar patterns of certain metameric invertebrates, using the
post-metamorphic mandibular segment as the critical means of identifying Gl. This
homology has not previously been justified because of the difference in form between
the pre-oral appendages of the major groups. Once the pre-metamorphic appendages
are recognized as the homologues of the peri-oral tentacles of a troph, their initial radial
symmetry can be seen as secondarily modified to a paired bilateral arrangement. This
change in symmetry is best understood in relation to the establishment of the antero-
posterior axis of the typical syngenean metazoan, the establishment of a composite
mouth and increased ‘streamlining’ of the head.

A reduction of the troph appears to have occurred within most of the major phyla
during their early evolution and has possibly been a significant factor affecting variation
at class level and above. The scyphozoan coelenterates, the tubicolous polychaetes,
oligochaetes and hirudinean annelids as well as the chelicerate arthropods exemplify the
dominance of the gone within the larger taxa. In segmented forms, this results in greater
prominence of the derivatives of the first strobe in the pre-oral tissues and of the first two
or three strobes in the composition of the mouth and gnathocephalon.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

JEAN J. CARTER 125

The development of multi-segmental blocks, or epitokal ‘stolons’ in some annelids,
particularly syllids, though regarded by some earlier writers as a kind of alternation of
generations (cf. Potts, 1911), is now more widely interpreted as a form of regeneration.
Okada (1934) and Berrill (1952) have described the development of a new prostomium as
part of the stolon head and the regeneration of a typical head when the anterior seg-
ments are amputated. At first sight, this suggests the replacement of troph tissue by gone
tissue. However, the formation of the new head blastema is closely linked with the arrival
of multipotent neoblasts, formed as ‘syncitial nests of small mesenchyme cells’ (Berrill,
1952: 414), lying in the angles of the septum and nerve cord. If the neoblasts arise
directly from the larval mesenchyme or if ectoderm derived from A, B, C or 1d lineages
(e.g. 2a?, 2b?, 2c? — the right, median and left somatoblasts respectively: cf. Fig. 2) con-
tributes to the new head, there is no departure from the principle of head formation
from troph-derived tissue. The development of epitokal heads in certain polychaetes may,
therefore, be a consequence of the distribution of tvoph tissue in the adult worm.

Experimental studies (Wada, 1968; Clement, 1962) have shown that embryos of
protostome larvae cannot undergo metamorphosis without the D cell. However, the D
cell alone, although capable of producing a ‘normal’ post-trochal region, dies before it
can reach the stage of metamorphosis, suggesting that a normally-functioning troph is
necessary for the development of the gone.

PROTOSTOMATOUS AND DEUTEROSTOMATOUS TAXA

Grobben (1908) and some later authors (e.g. Beklemishev, 1969) have used the
method of mouth formation, inter alia, to distinguish what they believe to be two major,
fundamentally divergent stocks of metazoans: the Deuterostomia and the Protostomia.
Neither group, however, has been universally accepted. For example, the validity of the
deuterostomes has been under challenge in recent years (Brien, 1974; Ldvtrup, 1975)
and some undoubted protostomes (e.g. onychophorans; Manton, 1949) have been
shown to possess a mouth that arises de novo and not from the blastopore or its point of
closure. There seems little doubt, however, that the annelid-arthropod-mollusc-
turbellarian group does possess features that indicate a true relationship between them.

The syngenean interpretation offers a solution to the dilemma about homology in
regarding the mouth of the polyp in the cnidarians to be the equivalent of the mouth of
the troph in protostomatous metazoans. The Protostomia are then seen to be a group of
phyla in which the troph mouth remains as part of the functional adult mouth, however
formed. Any exceptions to this among protostomes can be accommodated as special-
izations comparable with the degree of variation in cnidarian gastrulation (ie. by
epiboly, ingression or delamination). The deuterostomes, collectively identified more by
the absence of protostome features, lack the homogeneity of the protostomes and appear
in most, if not all, cases to be taxa in which the distinctive features of the troph are
reduced and the oral aperture of the gone becomes the functional mouth.

In these deuterostomes, where the adult mouth arises independently of the blasto-
pore, the way in which the mouth originates lends itself to the interpretation that it is
derived from the gone while the blastopore is largely confined to the pygidial area. In
such cases, the troph is still able to contribute to the sensory and neural structures of the
head or lophophore. The extent to which it does so is an expression of the relative contri-
butions of troph and gone to the adult body form and appears to provide a simple expla-
nation for some of the differences that exist between the ‘lophophorate deuterostomes’
and the ‘true deuterostomes’. Close relationship between these groups would not have to
be assumed in order to explain their embryological similarities.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

126 METAGENESIS AND ANIMAL FORM

CONCLUSIONS AND SUMMARY

If the Metazoa evolved from the Protista (Sleigh, 1979: 50-52), there is reason to
argue that because metagenesis is so widespread in the Protista and Cnidaria, it could
well be a fundamental characteristic of the Metazoa. In an a prior interpretation of
metagenesis in the metazoans, the more primitive form is the free troph (larva) while the
embryonalized equivalent (e.g. the egg-nauplius; Shiino, 1968) is its phylogenetic suc-
cessor. The general trend appears to have been towards direct development, the pre-
cocious formation of the gone (sexual generation) and the syngenean state. A hypogenic
interpretation of the life cycle, on the other hand, offers no basis for seeking an evo-
lutionary progression from alternating generations towards the development of primary
larvae and thence to progressive embryonalization and permits the marine primary
larvae to be considered as a series of individual late adaptations.

When the metagenic model is applied to the higher invertebrates, the existence and
metamorphosis of the marine primary larvae become explicable without need to iden-
tify the direct or hypothetical ancestors. The introduction of the concept of troph and
gone, as models for the polyp and medusa equivalents respectively, also allows the two
forms to be considered as homologues each capable of their own maturity, modifications
and evolution. It also suggests that through integration they could form a syngenean
animal, the morphology of which would vary depending on the relative contributions
made by the troph and gone components. The degree of development and persistence of
each entity controls such features as the degree of cephalization, the presence or absence
of acoelom, segmentation and tagmatization, as well as the type of mouth development.
By this a priori approach and the avoidance of traditional assumptions, an alternative in-
terpretation of animal form can now be offered.

The recognition of metagenesis and the almost simultaneous and integrated de-
velopment of the two generations can explain the trend toward embryonalization of the
larval form as an evolutionary advance. By recognizing the contribution which the troph
and gone entities each make to the functional morphology of the adult, it is possible to
trace an increasing specialization of the trop/’s sensory role in the developing brain and
its decreasing role in food uptake and locomotion. Against this background, other func-
tional systems can be isolated in similar fashion by embryological criteria and the direc-
tion of any progressive morphological series can be determined independently of
existing theories about phylogeny.

References

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JEAN J. CARTER 127

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WEIER, T. E., et al., 1974. — Botany, an introduction to plant biology, 5th ed. New York: J. Wiley and Sons.

WHEELER, W. M., 1893. — A contribution to insect embryology. J. Morph. 8: 1-160.

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WILSON, E. B., 1892. — The cell-lineage of Nereis. J. Morph. 6: 361-480.

PROC. LINN. SOC. N.S.W., 109 (2), 1986

128 METAGENESIS AND ANIMAL FORM
NOTE ADDED IN PROOF

“Tagmatization is used here to describe the progressive differentiation of a block of strobes into a tagma. It is
distinct from tagmosis, in which the troph may be incorporated with gone units, as in cephalization.

‘Embryonalization’ is defined as the gradual suppression of the free-living primary marine larvae (the troph
generation) and the modification or loss of their morphological identity due to the precocious development of
the gone before hatching. This requires recognition of a distinction between larval (troph) ectoderm, endoderm
and mesenchyme and the gone-derived ectoderm, endoderm and mesoderm. Its range is exemplified in crusta-
ceans where hatching may occur at any stage from a nauplius to a miniature adult. The word is identified with
the syngenean interpretation of animal form and is suggested as the ‘unknown force’ or ‘adult pressure’ which
lies behind adultation and acceleration as discussed by Jagersten (1972: 6, 218).

PROC. LINN. SOC. N.SW., 109 (2), 1986

Annexure to Proceedings Vol. 109

THE LINNEAN SOCIETY OF NEW SOUTH WALES

RECORD OF THE ANNUAL GENERAL MEETING, 1985

The one hundred and tenth Annual General Meeting was held in the Maiden Theatre of the National
Herbarium, Royal Botanic Gardens, Sydney, on Wednesday, 27th March 1985, at 7.30 p.m.

The President, Mr G. R. Phipps, occupied the Chair. The minutes of the one hundred and ninth
Annual General Meeting were adopted by the members present.

REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR 1984-85
Publications

The Society’s Proceedings were published as follows: —
Volume 107 (1982), Nos. 3 & 4 December, 1984.

This larger-than-usual issue contained papers presented to an international symposium on the Evolution
and Biogeography of Early Vertebrates held in Sydney and Canberra, February 1983.

In addition, the Society also published in December 1984 a special volume containing 36 papers
presented to the First International Polychaete Conference.

Dr P. A. Hutchings edited the Polychaete volume; Prof. T. G. Vallance continued to edit the Proceedings.
The honorary service of these two members is gratefully acknowledged.

Newsletter

The Linnean Society News, edited by Dr Helene Martin up to and including No. 33 (July 1984), and
thereafter by Mr G. R. Phipps, has been issued quarterly and distributed to members. It continues to
spread details of our programme, reports of Council resolutions, titles and authors of papers accepted for
publication in the Proceedings, news of grants from Joyce W. Vickery Scientific Research Fund and other
items believed to be of interest to members. Dr Martin and Mr Phipps are thanked for the time and effort
they have devoted to the News.

Membership

During the year, 8 full members were elected. Two members transferred to associate membership, one
to senior membership and three members resigned. At 26th March 1985, the roll stood at 280 full members,
21 associate members and one senior member.

Dr Peter Stanbury of the Macleay Museum, University of Sydney, was congratulated by the Society
on his being awarded the Medal of the Order of Australia.

Meetings

On Sunday, 3rd June, members and friends met at Connell’s Hill to walk through the intertidal
wetlands of southern Botany Bay and to examine the characteristic communities there under the guidance
of Drs P. Adam and R. J. King and Prof. A. W. D. Larkum.

On Wednesday, 20th June, at the Australian Museum, Mr Alan Andrews of the Metropolitan Water,
Sewerage, and Drainage Board gave an illustrated talk on Mount Kosciusko and the nineteenth-century
scientists and others who explored and studied the area.

The Macleay Memorial Lecture for 1984 was heard on the 18th July. The lecturer, Prof. R. V. S.
Wright, of the Department of Anthropology, University of Sydney, entitled his address: ‘New Light on the
Extinctions of the Australian Megafauna. We await submission of the text for publication in the Proceedings.

An excursion led by Dr L. A. S. Johnson, Director of the Royal Botanic Gardens, Sydney, to the
gardens’ annexe at Mount Tomah was held on 23rd September.

On Wednesday, 24th October, at the Australian Museum, Dr Marilyn Fox (Royal Botanic Gardens)
gave an illustrated talk on the year-to-year variations in the vegetation of western New South Wales.

The Society joined the Institute of Biology in Australia on 21st November for a combined visit to the
Division of Entomology Research Station at Warrawee. The meeting included four short talks by members
of the station staff on the general theme of biological control of pest arthropods.

Our first meeting in 1985 was as a co-sponsor, with the Institute of Biology in Australia, the Royal
Zoological Society of New South Wales and the Australian Museum Society, of a symposium entitled: ‘In
Defence of Science: A Response to Creationism. Speakers at the symposium, held on Saturday, 9th March,
at the N.S.W. Conservatorium of Music, were Mr Ron Strahan (National Photo Index of Australian
Wildlife), Prof. Ron Brown (Monash University), Dr Alex Ritchie (Australian Museum), Prof. Mike
Archer (University of New South Wales), and Dr David Briscoe (Macquarie University).

The Joyce W. Vickery Scientific Research Fund

Grants totalling $2,413 were awarded during the year to assist seven research projects. Details were
announced in Linnean Society News nos. 34 and 35:

Linnean Macleay Fellowship

The appointment of Mr R. W. Johnstone as Linnean Macleay Fellow was noted in the last annual
report. Mr Johnstone has now supplied the following account of his investigation of ‘Detrital Fluxes of
Carbon and Nitrogen in Coral Reef Sediments: One Tree Island Lagoon’.

‘As the title suggests, this project examines the flow of nitrogen and carbon through sediments in a coral
reef lagoon on One Tree Island, southern Barrier Reef, Queensland’

‘Work in 1984 centred primarily on the collection of data relating to the seasonal variations in sediment
carbon and nitrogen levels, with emphasis on the different nitrogen species. The data collected show a
marked difference in the levels of sediment ammonia between the very fine and coarse sediments and a
significantly higher level of ammonia in sediments during the summer months’

‘In conjunction with this work, time was spent developing a polarographic oxygen sensor for measuring
oxygen micro-gradients within sediment types. The results from this show interstitial oxygen levels to be
negligible below 2cm in all sediment types. Such a result was not found using coarser techniques but does
comply with the discovery of high H.S levels within the sediment.

‘The later part of 1984 was also spent carrying out nitrogen enrichment experiments in the field. As
may have been expected, all sediment types showed varying levels of nitrogen uptake outside that due to
diffusion. This aspect of my work represents the main area of interest for 1985 and several different methods
will be developed to monitor the fate and uptake kinetics of nitrogen supplied to the different sediment
types.

At the Council meeting held on 24th October, it was resolved that Mr Johnstone’s Linnean Macleay
Fellowship be renewed for 1985.

Office
The Society’s office at 6/24 Cliff Street, Milsons Point, is open from 9.30 a.m. to 5 p.m. every Tuesday.
The telephone number is (02) 929 0253.

Linnean Macleay Lectureship in Microbiology

Dr Kai Yip Cho, of the Department of Microbiology, University of Sydney, reports progress in the
following terms.

‘The work this year is concerned with the introduction of a wide variety of edible mushrooms to
Australian growers. The use of liquid spawn to replace conventional grain spawn is extensively studied’

Dr Cho adds that he has received a research grant of $55,000 from the N.S.W. government, through
the Advanced Technology Development Assistance Fund. He hopes that his project will ‘boost production,
diversification and profitability for mushroom growers in New South Wales within the next few years’.

The Society congratulates Dr Cho on his success and notes, with pleasure, how far his efforts go to
confirm the vision of Sir William Macleay.

PRESIDENTIAL ADDRESS

Mr G. R. Phipps delivered his Presidential Address on the ornithology of the Chevert expedition to New
Guinea, sponsored and led by Sir William Macleay in 1875. Illustrating his address with specimens from
the Macleay Museum, Mr Phipps discussed his own recent systematic studies and related them to the
history of the expedition and its bird collection. It is hoped to publish the address in the Proceedings in due
course.

DECLARATION OF ELECTIONS

As the number of nominations did not exceed the number of vacancies, no voting was necessary.
The following members were therefore declared elected to the Council for the year 1985-86:

President — DrP. M. Martin
Members of Council — Prof. M. Archer

Mr L. W. C. Filewood

Dr P. M. Martin

Dr I. G. Percival
Auditor — W. Sinclair & Co.

Mr G. R. Phipps then introduced Dr P. M. Martin as the President for 1985-86 and invited him to
take the Chair. The meeting closed with a vote of thanks to the retiring President.

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PROCEEDINGS of LINNEAN SOCIETY OF NEW SOUTH WALES
VOLUME 109

Issued 20th March 1987

‘NOTE: Publication of this issue, intended for December 1986, has been delayed by a serious fire at
the printery that month. Editor and printer regret the inconvenience to authors and readers.

CONTENTS:

ae)

63
69

91
107

117

NUMBER 1

R. V. S. WRIGHT

Sir William Macleay Memorial Lecture 1984. New Light on the Extinction of the
Australian cen tlh

R. J. KING

Aquatic Angiosperms in coastal saline Lagoons of New South Wales.
I. The Vegetation of Lake Macquarie

R. J. KING and V. M. HOLLAND

Aquatic Angiosperms in coastal saline Lagoons of New South Wales.

ll. The Vegetation of Tuggerah Lakes, with specific Comments on the Growth of
Zostera capricorni Ascherson

R. J. KING and J. B. BARCLAY

- Aquatic Angiosperms in coastal saline Lagoons of New South Wales.

lll. Quantitative Assessment of Zostera capricorni

R. J. KING and B. R. HODGSON
Aquatic Angiosperms in coastal saline Lagoons of New South Wales.
IV. Long-term Changes

NUMBER 2

P. F. CARR and B. G. JONES
Non-contemporaneity in the Marulan Batholith

D. M. BERGSTROM
An Atlas of Seeds and Fruits from Macquarie Island

A. T. HOWDEN
Notes on the Biology of adult and immature Amycterinae (Coleoptera,
Curculionidae)

D. T. ANDERSON
The circumtropical Barnacle Tetraclitella divisa (Nilsson-Cantell) (Balanomorpha,
Tetraclitidae): cirral Activity and larval Development

J. J. CARTER
Metagenesis as a possible Key to animal Form

ANNEXURE to Numbers 1 and 2. The Linnean Society of New South Wales. Record
of the Annual General Meeting 1985. Reports and Balance Sheets’ -

Printed by Southwood Press Pty Limited,
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PROCEEDINGS
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LINNEAN
SOCIETY

NEW SOUTH WALES

VOLUME 109
NUMBER 3
NUMBER 4

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
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the stipend of the Linnean Macleay Lecturer in Micro-
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Proceedings. Meetings include that for the Sir William
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OFFICERS AND COUNCIL 1986-87

President: P. M. MARTIN

Vice-president: HELENE A. MARTIN, G. R. PHIPPS, C. N. SMITHERS, A. J. T.
WRIGHT

Honorary Treasurer: A. RITCHIE

Secretary) BARBARA J. STODDARD

Council: A. E. J. ANDREWS, JUDITH H: K. EASTMAN, R. A. FACER, L. W. C.
FILEWOOD, MARILYN D. FOX, L. A. S. JOHNSON, HELENE A.
MARTIN, P. M. MARTIN, J. R. MERRICK, P. J. MYERSCOUGH,
IG. PERCIVAL, G., R. PHIPPS, A. RITCHIE, GN. SMIBHERS, iG:
VALLANCE, KAREN L. WILSON, A. J. T. WRIGHT

Honorary Editor: T. G. VALLANCE — Department of Geology & Geophysics,
University of Sydney, Australia, 2006.

Linnean Macleay Fellow: R. W. JOHNSTONE

Linnean Macleay Lecturer in Microbiology: K.-Y. CHO

Auditors: W. SINCLAIR & CO.

The postal address of the Society is P.O. Box 457, Milson’s Point 2061, N.S.W.,
Australia. Telephone (02) 929 0253.

PROCEEDINGS
of the

LINNEAN
SOCIETY

NEW SOUTH WALES

VOLUME 109
NUMBER 3

Marine Biological Laboratory
LIBRARY

APR 21 1988

Woods Hole, Mass.

Patan eg oe Ane Le oo

at ats, aa ah al
tA Ree ie Sheng et Uy

PRESIDENTIAL-ADDRESS

The Phylogenetic Significance of
Dracaena-type Growth

JOHN T. WATERHOUSE tabs)
(Delivered 28 March 1979)

Edited from the author's notes by C. J. Quinn*

The lilies, as we know from an old comment, are arrayed in more glory than ever
was Solomon. In spite of their glory which, to a modern taxonomist is but a manifesta-
tion of character-states, the lilies seem to be awaiting someone with the wisdom of
Solomon to find themselves classified. Of course I use the name ‘lily’ rather extrava-
gantly; in a strict recourse to nomenclature the family taxon, Liliaceae, must accept the
genus Lilium. But the problems seem to be: what other genera might be placed with
Lilium in the family of lilies, and which other families have, say, an ordinal relationship
with the lilies?

In this comment to-night I am not going to delay with the history of the convo-
lutions in the classification of these Monocotyledons, but the historical candidates for
the Liliaceae have flowers that may be derived from a trimerous 5-whorled condition
P33 A3,3 Gig). The ovary may be superior or inferior, and a whorl of stamens is some-
times missing.

Amongst the various taxonomic treatments of these candidates for the Liliaceae we
can for the moment note that:

Brongniart (1813) disposed them in several families, including Liliaceae

and Lindley (1846) and Amaryllidaceae;

Engler (1897) grouped them ina few big families;

Hutchinson (1934) allocated them to many families in several orders
(Lihales, Agavales, Amaryllidales and Haemodorales
PP.)

Cronquist (1968) has more recently become confused and has gone back
to some larger-than-Engler families in a single order,
Liliales;

Huber (1969) and who are enjoying some present day popularity, have lots

Dahlgren and Clifford (1982) of small families arranged in several orders (e.g.,
Liliales, Asparagales, Amaryllidales and Haemodorales
Pp:):

The differences in the above systems are not the simple matter of tribes being
elevated to families and families becoming orders, for an inspection shows the content
and alliances keep altering. Hence we are faced with a real systematic problem, which
indicates that we have been doing something wrong — wrong in the sense that wide

* Copy received 22 April 1987. We thank Dr Quinn for realizing the late John Waterhouse’s presidential
address. Dr Quinn has also prepared the Memorial to our distinguished former president that is printed in
this issue (pp. 139-142).

PROG. LINN. SOC. N.S.W., 109 (3), (1986) 1987

130 PRESIDENTIAL ADDRESS

—_—— wiesclltion ne rt a tit aN

Fig. 1. Examples of pachycaulous monocotyledons.

a. Kingia australis with post-anthetic inflorescences: Albany, Western Australia. b. Beaucania sp. c. Xanthorrhoea
reflexa; box at base of trunk is 15cm high; Beverley Western Australia. d. Transverse section of 1.9m high
Xanthorrhoea arborea trunk at ground level. The primary tissues, including the basal woody cone, occupy the
central region to a diameter of 8.5cm; the remainder of the trunk is secondary, arising from a perimeristem
that is under the dark outer layer of leaf bases and bark (see Fig. 2a). Calga, NSW. e. Yucca elephantipes. f.
Dracaena draco. g. Nolina sp. b and e-g, Royal Botanic Gardens, Sydney.

PROC. LINN. SOC. N.SW., 109 (3), (1986) 1987

JOHN T. WATERHOUSE 131

acceptance or approval is not being given to any one system. What is it we are doing
wrong?

Now, one of the features of natural classification is that if we use the character-states
of A, B and C to set up family number 1, we see no necessity to use another set of
character-states of A, B and C to set up family number 2. It is, however, a fact of taxo-
nomic history that character-states of the flower have, by and large, been used very suc-
cessfully in setting up families and supra-specific taxa in the dicotyledons. Of course we
have seen plenty of tidying up with input from more recently available data sources such
as histology, karyology, biochemistry, palynology. But the tidying up was often already
seen to be required. There are indeed also many monocotyledonous taxa of wide and
long-time acceptance that can be well circumscribed on floral character-states.

It seems, therefore, that, following the comparative success of floral characters in
the classification of most dicotyledons and many monocotyledons, we have expected the
same approach to be successful over-all. This expectation, I believe, should be aban-
doned with reference to the Lilies and their relatives. I have come to this opinion as a
result of some studies contemplating the Australian family Xanthorrhoeaceae, which
consists of genera that have all been considered at some time or another candidates for
the Liliaceae. The Xanthorrhoeaceae (in its present content) was set up by Hutchinson
(1934), whose system for the ‘lilies’ in particular presented some novelty, in that attention
was paid to non-floral characters — habit and leaves — as well as to floral characters, in
setting up the numerous families and orders.

Hutchinson’s Xanthorrhoeaceae contained genera with species of gross habit —
small trees or ‘shrubs’ (as far as these terms can be applied to monocots; Fig. la, c). It in-
cluded, inter alia, the three genera, Xanthorrhoea (ca 15 spp.), Kingia (1 sp.) and Dasypogon
(2 spp.), and he put this family in the order Agavales along with his family Agavaceae,
which again contained species of gross habit (Fig. 1b, e-g) belonging to such genera as
Dracaena, Cordyline, Doryanthes, Yucca, Phormium and Nolina. In both families, therefore,
there are species which can be described as arborescent or at least frutescent — in
general, pachycaulous; so in this complex — Hutchinson’s Agavales — we are dealing
with big perennial robust monocotyledons with a habit that probably impressed a
botanist of the northern hemisphere where monocotyledons are spring-flowering shoots
produced annually from bulbs, corms and rhizomes.

In the development of a natural classification it has forever been unwise to fasten on
to one character-state for setting up a taxon. Moreover, such a character-state dis-
tributed in representatives of that taxon must be seen as an homologous expression of a
character. Thus, is it reasonable to postulate that pachycauly in Hutchinson’s Xan-
thorrhoeaceae and Agavaceae is the result of homologous developmental processes in all
the genera/species referred here?

Some years ago I started to investigate this point. It was relatively simple to show
that the pachycaulous habit in members of the Xanthorrhoeaceae is the expression of
two fundamentally different processes. In the course of the studies it also became ob-
vious that Hutchinson’s character of ‘dry perianth’ as typical of the family is also ques-
tionable; this uncertainty ultimately led to an investigation of the floral anatomy of some
of the members of the family. The precise results of these investigations are to be pub-
lished in full later. For the present, and to be succinct, Hutchinson’s family Xan-
thorrhoeaceae is monstrously unnatural. However, as a result of these studies I consider
there are two histological aspects in the ‘lilies’ (? and indeed in other monocots) that are
of systematic importance:

i) features associated with the development of pachycauly; and

il) features of the ovary wall — especially the septa.

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

132 PRESIDENTIAL ADDRESS

In searching through the literature I note that there is nothing particularly novel
about the features I am going to describe; it is simply that their systematic importance
has not been highlighted. Moreover, the details are fairly well recorded, so I can only
conclude that as systematists we are ignoring a whole pool of histological data already
scored by various workers.

Regarding pachycauly, the kind of development I wish to consider tonight is that
kind found in species of Dracaena. I am going to imply that other kinds of pachycauly in
monocotyledons are very different (e.g., that in Kingza; Fig. la), though I am not going
to describe them to you (see Staff and Waterhouse 1981). As you can see from the illus-
trations (Fig. 1b-g), many of the species that have the Dracaenoid kind of pachycauly are
quite arborescent.

The basic histological processes producing this type of pachycauly have been
known for a long time (since 1840-1870), but they really are not given much attention in
accounts of general plant histology, and, if mentioned at all, the histological picture is
usually described only in the transverse aspect of the stem, whereas it is in the longitudi-
nal aspect that some of the most interesting and fundamental features of the process are
to be seen.

In the transverse aspect of the primary stem (Fig. 2d) I want you to note:

a) the scattered vascular bundles, a typical monocotyledonous character; all tissues
are of course primary;

b) that the vascular bundles are cribricentric, or nearly so, a feature found in, but not
widespread in, monocotyledons;

c) that the tracheary elements are tracheids and the metaxylem ones have oblique
narrow-aperture pits, features found in, but again not widespread in
monocotyledons (these features are of course only ascertainable in macerated
tissue);

d) the peripheral cylinder of meristematic cells, which is probably more than one cell
thick.

The transverse aspect of a stem with secondary thickening 1s shown in Figs Id and
2a. The points to note are:

a) the perimeristem cuts off radial rows of parenchyma cells to the inside and a few to
the outside; these are appropriately termed secondary tissue;

b) proceeding centrally from the perimeristem —

Fig. 2. Diagrams of secondary thickening in Cordyline stricta. lg, mature lignified ground tissue; p, phloem; pd,
primary vascular bundle at outer limit of primary stele; ph, phellogen established under the leaf bases; pm,
perimeristem; sb, secondary vascular bundle; s7, secondary derivatives of the perimeristem formed on its
inner face; so, secondary derivatives of the perimeristem formed on its outer face; x, xylem.

a. TS sector of outer part of mature stem showing secondary tissues. The radiating broken lines indicate the
radial rows of secondary lignified ground tissue laid down by the perimeristen. Scale = Imm.

b. Schematic representation of the longitudinal aspect of intrusive growth during the differentiation of secon-
dary vascular bundles. Positions 1, 2 and 3 indicate successive stages in the differentiation of a vertical file of
perimeristem derivatives into secondary vascular elements. The arrow indicates a horizontal row derived
from a particular perimeristem initial, and members of this row in each bundle are labelled ¢;, t) and ¢3, and
are stippled. At formation (position 1), only one cell of the vertical file is present in the indicated horizontal
row (arrow). At position 2, file members have extended to twice their initial length, and intruded halfway into
the rows immediately above and below their original position; there are thus two cells at any level in the indi-
cated horizontal row. At position 3, each member of the file now extends through 7 vertical rows, and seven
cells will be found in the indicated horizontal row; only three members of vertical file 3 are shown in the
diagram for reasons of simplicity.

c. Schematic representation of the transverse aspect of differentiation of secondary vascular bundles. 6,, 6»
and 4, indicate three successive stages in the process.

d. TS sector of young stem with perimeristem just differentiated outside the primary vascular bundles. Scale
= Imm.

PROC. LINN. SOC. N.SW., 109 (3), (1986) 1987

133

JOHN T. WATERHOUSE

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PROC. LINN. SOG. N.S.W., 109 (3), (1986) 1987

134 PRESIDENTIAL ADDRESS

1) there is an apparent cutting-up of (1.e., lots of cells divisions in) scattered single

derivatives of the perimeristem;

ii) these strands of cells differentiate into cribricentric (or nearly so) vascular

bundles similar in appearance to the primary bundles;
c) again, the tracheary elements are tracheids of the type described above;
d) the secondary bundle is formed within one (? always) radial row of secondary
tissue;
e) the secondary parenchyma between the mature secondary bundles becomes ligni-
fied, an aspect of the process that will not be discussed further here.

In the longitudinal aspect (RLS) of the secondary tissue it is apparent that there is a
great difference between the length of the perimeristem initial and the length of the trac-
heid that developed from it — indeed from macerated tissue the tracheid is about 20
times the length of the perimeristem initial. Now it can only have attained this length by
intrusive growth. Let us look at the process more schematically. In the diagram (Fig.
2b), all the members of the vertical file 7 are destined to become vascular elements; they
may be considered protracheary elements. Imagine that we take up a position of inspec-
tion at the horizon marked by the arrow. At position 1 in this horizon there is one deri-
vative in the vertical file (¢;). In position 2 each protracheary element of the file has
started to elongate apically and basally, and consequently has started to intrude between
the cells above and below itself (see cell tz). The result is that at the given horizon there
are now two cells where only one existed before. In position 3 (cell t;) each protracheary
element has elongated to 7 times its original length. Thus at the given horizon our origi-
nal element (stippled) is now accompanied laterally by six other elements of the same
vertical file, three having intruded from above, and three from below; there would be
seven elements, therefore, in a transverse section of the file where there was formerly
only one. This is zntrusive growth; what looked like cell-division in the transverse aspect
(Fig. 2c) is not. It is this elongation of the tracheids that is responsible for the lateral ex-
pansion of the secondary vascular bundle as it matures (cf. 5;, 62 and 63 in Fig. 2c). In
macerated secondary tissue (Fig. 3) the tracheids can be seen to be quite twisty and to
have battered looking tips as a result of having had to force their way between adjoining
elements. Note also that the pitting on these elements is very similar to that on the
primary tracheids, being rather like the pitting one usually finds on fibres. The block-
shaped lignified parenchyma that occupy the regions between the bundles are also
visible in some macerates.

A comparison of macerates from a range of genera possessing this Dracaena-type
growth is shown in Fig. 3. These are chosen to indicate how similar the elements are, yet
they constitute 2-3 different families (depending on author) disposed in 2 different
orders.

The seven genera or groups of genera having Dracaena-type growth that are recog-
nized in Table 1 have sub-sets of characters that distinguish each from the other. The
Aristeae are probably the most out on a limb and are usually referred to the family
Iridaceae, which of all the groups we might be considering here has been the most taxo-
nomically stable. But as well as the Dracaena-type growth there are other shared
character-states that indicate affinity. And this form of growth of stems represents a set
of characters that is unlikely to have evolved more than once, and I suggest that it
indicates a common origin of those taxa that have it.

Let us look at the world distribution of the generic groups as set out in Table 1.
These distributions are compatible with a single origin when the continental masses
were together. That is, they appear to be an ancient group that has become differen-
tiated after continental separation.

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

JOHN T. WATERHOUSE 135

Fig. 3. Xylem tracheids from macerated secondary stem of Dracaenoid plants, showing battered looking tips
resulting from intrusive growth. Scale = 100um.
a. Cordyline stricta. b. Aloe sp. c. Yucca sp. d. Xanthorrhoea australis.

TABLE 1
Distribution of the taxonomic groups recognizable in the genera displaying dracaenoid thickening
GENERIC GROUP DISTRIBUTION
Agave and Yucca Central'America
Nolina North America
Xanthorrhoea Australia
Aloe South Africa
Lomandra Australia, New Zealand, Hawaii
Cordyline Australia, New Zealand, widespread in the tropics

and Dracaena

Aristeae (Iridaceae) South Africa, Australia

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

136 PRESIDENTIAL ADDRESS

Fig. 4. Median transverse section of ovary of Xanthorrhoea resinosa at anthesis. The dorsal or median (m) and the
ventral or marginal (/) vascular traces of each carpel are indicated in black; other vascular bundles (laterals of
the dorsal trace) are in outline (see Fig. 5). sg, septal gland. Scale = Imm.

As a group where do they fit is into the lilies sensu latzssiemo. This is where I believe an
aspect of floral histology is important. Let us look at some features of the ovary wall. Fig.
4 shows a transverse section of the ovary of Xanthorrhoea resinosa in which glands can be
seen clearly in each septum of the ovary. A generalized diagram is shown in Fig. 6. The
gland is a pouch opening at the top of the ovary and which may extend right to the base
of the ovary and even interconnect below the locules. Note also the three-armed stylar
canal leading from the stigma to the locules. Similar septal glands occur in the inferior
ovary of Agave and the superior ovaries of Aloe and Yucca.

Fig. 5. Diagrammatic reconstruction of a carpel of Xanthorrhoea resinosa opened out flat and viewed from the
adaxial side. The shaded region indicates the septal portion of the carpel. Note the highly vascularized condi-
tion with numerous laterals arising from the dorsal trace (m). /, ventral or marginal traces. 4 indicates the level
of the section illustrated in Fig. 4.

Septal glands appear to occur throughout such groups as Amaryllidaceae (of either
Engler or Hutchinson) and Iridaceae, although in some species they appear to have
been secondarily lost, and are characteristic of Zingiberaceae, Musaceae, Cannaceae
and the syncarpous Palmae. Within the lily family or families their distribution 1s ‘taxo-
nomically sporadic in current systems. Certainly, Huber (1969) has considered them,
but seems to be hesitant/capricious in his taxonomic use of them. Many of these same
families are also characterized by a highly vascularized carpel in which the ovary wall 1s

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

JOHN T. WATERHOUSE 137

traversed by laterals arising from the dorsal vascular bundle (i.e., from the mid-vein of
the ancestral fertile leaf; Figs 4 and 5). Both septal glands and the highly vascularized
carpel are fairly complex characters that indicate a common origin of the plants that
bear them.

Well, all the Dracaena-type genera have septal glands — this probably is also an an-
cient character typical of some of the so-called lilialian alliances. It is apparently com-
mon in Asphodeloid genera but not in Melanthoid genera. I have no great idea on how
systematically significant the presence or absence of septal glands is, but it might be
worth considering whether it has any bearing on the evolutionary hypothesis that the
monocotyledon flower is a synanthium in origin rather than a strobilus; under this
hypothesis each carpel plus 2 stamens is a floral unit subtended by bracts (= perianth).

Fig. 6. Generalized diagram of syncarpous gynoecia of Dracaenoid monocotyledons showing the septal
glands (sg) between the locules (/oc) in both longitudinal and transverse aspects.

There is, then, a large group of ancient plants which have Dracaena-type growth and
septal glands. How do they relate to other lilialian monocotyledons? The following
points are of relevance:

1. within the group there are already small species with a vestigial amount of secon-
dary growth

2. outside the group there are small species with vestigial amounts of secondary
growth, e.g., Chlorophytum

3. all such species have cribricentric bundles, or collateral bundles with U-shaped
xylem consisting of Dracaena-like tracheids, whether primary or secondary

4. there are other species with amphivasal bundles, or nearly so, with xylem consisting
of tracheids that more or less resemble Dracaena-type primary tracheids, but which
show no sign of a vestigial perimeristem. All these small species might have septal
glands.
In conclusion, there appears to be a series leading to the small perennials with

annual re-growth more typical of the temperate regions and much more common in

the general botanical literature. But when I speak of an evolutionary series from

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

138 PRESIDENTIAL ADDRESS

arborescent Dracaena-type plants to some of the smaller species I don’t mean to imply

that all the smaller species have come this way. I simply believe that the Dracaena-type

habit is ancient and the septal gland is probably also old in some of the Lilialian lines.

The implication is that other Lilialian genera that do not have these histological charac-

ters are not so closely related. The Dracaena-type plants are not the ancestors of the lilies

sensu latissimo. I regret we haven't found a Solomon with his wisdom yet. The problem of
elucidating the relationships within this group still remains. If we are to solve it over and
above determining the presence or absence of dracaenoid growth, we must:

1. describe properly the tracheary elements. There is pronounced variation from fern-
like tracheids in Dracaena to those shown in Fig. 3, but a recent publication has them
all scored simply as non-vessel elements. The distribution of the various kinds of
elements is still a puzzle in the monocotyledons

2. look critically at the bundle types in the stems. Cheadle and Uhl (1948) have too
many types, and we must be careful to distinguish between stems, rhizomes and
leafy inflorescence axes when making comparisons

3. look for septal glands and multi-veined carpels. Huber (1969) has acknowledged
these, but doesn’t use data on tracheary elements.

One of the problems that must be overcome is that we must stop thinking of our
taxonomy from the top of our hierarchies downwards, and start building up groups from
the bottom with what we have at hand. The number of genera involved is so large that no
one person can adequately survey them all. If 1 am going to start on the Sydney scene —
with what I have at hand — my groups are going to be incomplete, lack overall world
perspective and as a consequence cause, horror of horrors, later nomenclatural and cir-
cumscriptive chaos. But start we must, if our treatment of the lilies is to be improved.
The remedy, I suggest, is to divorce one’s groups from the code and launch them simply
so that others can add to or modify them before formalizing them under the code, a
procedure that has been adopted quite successfully by Pryor and Johnson (see Pryor and
Johnson, 1971) in grappling with the sub-generic taxa in the genus Eucalyptus.

References

BRONGNIART, A. T., 1843. — Enumération des genres de plantes cultivés au Muséum d'histoire naturelle de Paris. Paris:
Fortin, Masson et Cie.

CHEADLE, V. I., and UHL, N. W., 1948. — Types of vascular bundles in the Monocotyledoneae and their re-
lation to the late metaxylem conducting elements. Amer. J. Bot. 35: 486-496.

CRONQUIST, A. 1968. — The evolution and classification of flowering plants. London: Nelson.

DAHLGREN, R. M. T., and CLIFFORD, H. T., 1982. — The monocotyledons: a comparative study. London:
Academic Press.

ENGLER, H.G. A., and PRANTL, K. A. E., 1897. — Die naturlichen Pflanzenfamilien. Nachtrage 1. Berlin.

Huser, H. 1969. — Die Samenmerkmale und Verwandtschaftsverhaltnisse der Liliifloren. Mutt. Bot.
Miinchen 8: 219-258.

HUTCHINSON, J. 1934. — The families of flowering plants, Vol. 2. London: Macmillan.

LINDLEY, J., 1846. — The vegetable kingdom. London: Bradbury and Evans.

Pryor, L. D., and JOHNSON, L. A. S., 1971. — A classification of the Eucalypts. Canberra: ANU Press.

STAFF, I. A., and WATERHOUSE, J. T., 1981. — The biology of arborescent monocotyledons, with special
reference to Australian species. In PATE, J. S., and McComs, A. J., (eds), The Biology of Australian
Plants: 216-257. Nedlands (W.A.): University of Western Australia Press.

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

John ‘Teast Waterhouse 1924-19835

(Memorial Series No. 27)

Members were greatly saddened by the sudden death of Mr John T. Waterhouse at
his home in Gordon, Sydney, on the night of April 1, 1983, aged 58 years.

Born on December 2nd, 1924, he received nig secondary education at Tamworth
Boys High School and then attended the University of Sydney, where he obtained a
Bachelor of Science with First Class Honours in Botany in April, 1947. During his
honours year, which was supervised by the late Dr J. McLuckie, he carried out ‘A physio-
logical investigation of the fungus, Penicillium expansum Link.’ During the next three
years he held positions as demonstrator and teaching fellow in Botany at Sydney
University. In 1948 Dr A. J. Eames spent a period of leave in Sydney, and he was in-
strumental in directing John towards his life-long interest in arborescent
monocotyledons. In 1950, however, John left the university for life on a property which
he always referred to as “The Blue Duck’, a corruption of its aboriginal name, because of
the problems he faced with wild ducks following introduction of irrigation for the
production of lucerne. It was partly because of these problems that he returned to the
University of Sydney in 1953 as atemporary lecturer in Botany, but at the end of 1954 he
again opted for life on the land, this time on a grazing property, ‘Burragillo, in the
Collarenebri area. Throughout his time on the land he maintained and developed his

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

140 MEMORIAL SERIES

knowledge of the native flora, and started compiling a checklist for the area. It is hoped
to publish this in the near future. In 1957 John was seriously injured in a fall from a
horse while yarding stock, and this resulted in some lengthy periods in hospital over sub-
sequent years. He left the rural life he loved at the start of 1962 and joined the staff of the
Department of Botany at the University of New South Wales, as a Senior Tutor, just be-
fore the department moved to the Kensington Campus. Shortly after joining the depart-
ment he was appointed a lecturer. He resumed his studies on Australian Grass Trees,
and in 1967 presented a thesis entitled, “Some aspects of the status of the family Xan-
thorrhoeaceae’, for which he was awarded a Master of Science by the University of New
South Wales.

In 1971-72 he spent his first study leave with Professor V. H. Heywood in the
Department of Botany, University of Reading, during which he carried out a study on
the tribe Anthemideae (Asteraceae), and obtained a Master of Science in Pure and
Applied Taxonomy.

John joined the Linnean Society of New South Wales in 1947. He was elected to the
Council of the Society in August, 1975, and remained an active and valued member
until his death. He was President in 1978-1979, at a time of considerable controversy
over the accumulating debt on Science House, and chaired several torrid meetings with
great skill. He also devoted a great deal of time to researching the complexities of the
issues, and went out of his way to inform members. He continued to make a large con-
tribution to the solution of problems faced by the Society in his term as Vice President,
1979-1982.

In his years at the University of New South Wales John had a great impact on the
teaching of botany, and was particularly active in promoting field work. He was respon-
sible for raising taxonomy in the undergraduate syllabus above the level of plant iden-
tification, developing at first a part unit and later a full unit in the third year on the
principles and methods of taxonomy. He also made a large contribution to teaching at
first and second year levels. He organized the day excursion to Kurnell for First Year
Biology students until the pressure of numbers made the logistics of the occasion intoler-
able: the last excursion involved 13 double-decker buses loaded to the gunwales. He was
a mainstay of the annual second-year field camp which was held at Mount Boss State
Forest from 1966 until that too outgrew the facilities. He also conducted a third year
field camp in the same area for his taxonomy students for many years. John’s field camps
were always a happy combination of efficiently organized work by day and relaxed
socialization around the fire at night, always with a few interesting specimens in his
hand, and books and a plant press at his elbow. He felt that field work should be enjoy-
able. The combination of his quiet, easy manner and critical mind sharpened the wits of
a large number of students, and stimulated many to take a deeper interest in botany.
John also considered that preparation of food should receive proper attention; his third
year camps were an object lesson on just what gourmet delights could be produced over
an open fire, and word soon spread through the student grape vine.

Two of his research projects had their origins in student exercises on these field
camps. The study of the growth of the Bangalow Palm (Archontophoenix cunninghamiana)
was initiated on the 1967 second year camp, while his work, in collaboration with Dr
M. M. Hindmarsh, on a field key to the rainforest species south of the Macleay River
grew out of their key to the species of the Wilson River Primitive Area, which was tested
and upgraded with the assistance of successive groups of second year students. The final
key, which is based on personal examination of fresh specimens of all species, and uses
characters of petiole anatomy and exudate, as well as the more usual range of vegetative
characters, is now being completed by Dr Hindmarsh. It should prove a valuable aid to
identification in these complex communities.

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

JOHN TEAST WATERHOUSE 141

Another of his research interests centred on the Myrtaceae. Apart from his work
with Dr P. G. Wilson on Trstania and its allies, he also studied Eucalyptus and the
Syzygium-Acmena complex, and drew attention to the distinctiveness of Syzygium floribun-
dum. Only the week before he died he saw ‘Waterhousea’, the name for the segregate genus
Hyland (1984) erected to hold this and two Queensland species, in print in the new
Royal Botanic Gardens pamphlet entitled ‘A Rainforest Walk’ It is typical of him that he
should protest with a grin that the name had not then been published.

During his field work, John assembled a large body of data on the eucalypts, par-
ticularly relating to venation patterns, oil gland size and distribution, and bud and cap-
sule morphology. Although this is preserved on cards, it seems unlikely that it will ever
be put to use in the construction of a key as he had envisaged.

In 1980, he embarked upon an intensive floristic survey of the Magela Creek
Catchment, Northern Territory, in relation to the projected establishment of the
Jabiluka uranium mine by Pancontinental Mining Ltd. He studied the area throughout
the full monsoonal cycle, amassing a large collection and a wide range of field obser-
vations. Although he had produced a draft species list (Puttock and Waterhouse, 1981),
and had completed studies of Limnophila (Wannan and Waterhouse, 1985) and
Blepharocarya (Wannan et al., 1985; 1987), much working up of this collection remained to
be done, and he was looking forward to an early retirement which would allow him to
devote more time to it. It is a source of great satisfaction to his colleagues that, through
the good offices of the Office of the Supervising Scientist for the Alligator Rivers Region,
money has now been made available so that more can be realized from this important
collection of the northern Australian flora.

John considered it his duty to profess botany in the broadest sense; he was always
very willing to spend time assisting those who called on his expertise, whether they were
students, colleagues or members of the public who so often were directed to his door. He
also devoted a great deal of effort over the years to expanding the collection held by the
herbarium at the University of New South Wales, which now comprises some 45,000
specimens, and improving the level of its curation. In 1980 he registered the herbarium
with the acronym UNSW, and soon afterwards obtained a special development grant
from the university to enlarge the accommodation both for the collection and for
associated staff and students. This reconstruction was in hand at the time of his death,
and it is a suitable acknowledgement of his contribution to the teaching of taxonomy
that the enlarged facility was named “The John T. Waterhouse Herbarium at the official
opening ceremony in June 1983.

In 1973 John was elected a Fellow of the Linnean Society, London, and also became
a member of the british Systematics Association. He was a foundation member of both
the Australian Systematic Botany Society and a member of the founding committee of
the Friends of the Royal Botanic Gardens, Sydney. John T. Waterhouse will be remem-
bered particularly by generations of students from both the University of Sydney and
the University of New South Wales as a stimulating teacher of botany, and by the botani-
cal community as a valued colleague with a splendid sense of humour. He will also be
remembered for his contributions to monocotyledonous anatomy, the taxonomy of the
Myrtaceae and his work on the key to rainforest species of New South Wales. Not only
the Society but the Australian botanical community at large is much the poorer for his
passing.

Works by John Teast Waterhouse

WaTERHOUSE, J. T., 1947. — A physiological investigation of the fungus, Penicillium expansum Link. Sydney:
University of Sydney, B.Sc. honours thesis, unpublished.

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

142 MEMORIAL SERIES

WATERHOUSE, J. T., 1967. — Some aspects of the status of the family Xanthorrhoeaceae. Kensington:
University of New South Wales, M.Sc. thesis, unpublished.

WATERHOUSE, J. T., 1972. — Studies on the Anthemideae. Reading: University of Reading, M.Sc. thesis,
unpublished.

WATERHOUSE, J. T., and QUINN, C. J., 1978. — Growth patterns in the palm Archontophoenix cunninghamiana.
Botanical Journal of the Linnean Society, London, 77: 73-93.

PuTTock, C. F., and WATERHOUSE, J. T., 1981. — Floristic survey of the Jabiluka area. In MORELY, A.., (ed.),
A Review of Jabiluke Environmental Studies: Chapter 19. Sydney: Pancontinental Mining Ltd.

StaFF, I. A., and WATERHOUSE, J. T., 1981. — The biology of arborescent monocotyledons, with special
reference to Australian species. Jn PATE, J. S.. and McComs, A. J., (eds), The Biology of Australian
Plants: 216-257. Nedlands: University of Western Australia Press.

WILSON, P. G., and WATERHOUSE, J. T., 1982. — A review of the genus Tristania R.Br. (Myrtaceae): a hetero-
geneous assemblage of five genera. Australian Journal of Botany 30: 413-446.

WANNAN, B. S., WATERHOUSE, J. T., GADEK, P. A., and QUINN, C. J., 1985. — Biflavonyls and the affinities
of Blepharocarya. Biochemical Systematics G Ecology, 13: 105-108.

WANNAN, B., and WATERHOUSE, J. T., 1985. — A taxonomic revision of the Australian species of Limnophila
R.Br. (Scrophulariaceae). Australian Journal of Botany 33: 367-380.

WANNAN, B. S., WATERHOUSE, J. T., and QUINN, C. J., 1987. — A taxonomic reassessment of Blepharocarya
F.Muell. Botanical Journal of the Linnean Society, London, 95: 61-72.

WATERHOUSE, J. T., 1987. — The phylogenetic significance of Dracaena-type growth. Presidential address,
1979. QUINN, C. J., (ed.). Proceedings of the Linnean Society of New South Wales, 109: 129-138.

C. J. Quinn

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

Complementary feeding Habits of Golden Perch
Macquaria ambigua (Richardson) (Percichthyidae)

and Silver Perch Bidyanus bidyanus (Mitchell)
(Ieraponidae) in farm Dams

C. C. BARLOW, R. MCLOUGHLIN and K. BOCK
(Communicated by J. R. MERRICK)

BARLOw, C. G., MCLOUGHLIN, R., & BOCK, K. Complementary feeding habits of
golden perch Macquaria ambigua (Richardson) (Percichthyidae) and silver perch
Bidyanus bidyanus (Mitchell) (Teraponidae) in farm dams. Proc. Linn. Soc. N.S.W.
109(3), (1986) 1987: 143-152.

A comparison was made of the diets and morphology of the alimentary tracts of
golden perch Macquaria ambigua and silver perch Bidyanus bidyanus. Each species was
grown in monoculture in farm dams and their diets compared with the available food
sources using the prey-selection index C. The two species were then cultured together
and the dietary overlap measured using Schoener’s index. Golden perch proved to be a
macrophagic carnivore, eating insects and crustaceans, whereas silver perch was an
omnivore, feeding predominantly on zooplankton. The alimentary tract of golden
perch is typical of a carnivore, whereas that of silver perch is adapted to an omnivorous
diet, with a filtering mechanism on the gill rakers for capturing zooplankton. On the
basis of their dietary habits, these two species are ideally suited for polyculture.

Christopher G. Barlow (now Queensland Dept of Primary Industries, Research Station, Walkamin,
Australia 4872), Richard McLoughlin (now C.S.I.R.O. Marine Laboratories, G.P.O. Box 1538,
Hobart. Australia 7001) and Ken Bock, Inland Fisheries Research Station, Narrandera, Australia
2700; manuscript received 29 April 1986, accepted for publication 20 August 1986.

INTRODUCTION

Large numbers of golden perch (Macquaria ambigua (Richardson 1845): Percich-
thyidae) and silver perch (Bidyanus bidyanus (Mitchell 1838): Teraponidae) are stocked
annually in farm dams throughout eastern Australia, to provide fish for recreational
angling and domestic consumption (Rowland et al., 1983; Rowland, in press a,b).
Management for fish production is minimal, since the primary purpose of the dams is
for watering domestic stock. Consequently, fish stocking rates are usually low, in the
region of 150-350 fish/ha, and the carrying capacity of the dams 1s only 200-500kg/ha
(Barlow, in press). One method of increasing the production of fish in these dams 1s to
stock two or more species with complementary feeding habits, that is, polyculture.

Little is known about the diets of golden perch and silver perch, although limited
observations indicate that golden perch is a carnivore, feeding mainly on crustaceans,
insect larvae and molluscs, and that silver perch is an omnivore, consuming small
aquatic insects, molluscs, earthworms and plant material (Merrick and Schmida, 1984).

The aim of this study was to investigate the feeding habits of golden perch and
silver perch reared in farm dams and thus ascertain if these fishes are suitable for poly-
culture. This was done by determining the preferred foods of the two species when
grown separately, and then comparing their diets when grown together. The mor-
phology of their alimentary tracts was also examined.

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

144 FEEDING HABITS OF PERCH

MATERIALS AND METHODS

The fish used in the trials were artificially bred and reared at the Inland Fisheries
Research Station, Narrandera. Those used in the monoculture trial were one year old,
whereas those used in the polyculture trial were three years old. The fish were stocked in
earthern dams situated in flat or gently undulating pastoral country. The dams were
about 0.06ha and 2-3m deep. The dam stocked with golden perch in the monoculture
trial had a dense growth of red milfoil, Myrophyllum verrucosum, extending 2-3m from the
shore and occupying the entire perimeter of the dam. The other dams did not contain
macrophytes. The food available to the fish consisted of the organisms produced
naturally in the dams.

MONOCULTURE TRIAL

Twenty-eight golden perch were placed in one dam and 25 silver perch in another
in January 1980, and harvested with a seine net three weeks iater. The fish were trans-
ported live to the laboratory in plastic bags containing water and an oxygen atmosphere.
The total length and weight of each fish were recorded and the alimentary tract, from
the oesophagus to the anus, removed and measured. Examination of the transporting
medium showed that no regurgitation or defaecation occurred between the time of
capture and dissection.

Macroinvertebrates in the guts were identified and counted. In addition, the per-
centage of zooplankton in the stomach contents of each silver perch was estimated
volumetrically.

The available food sources, or potential prey species, were sampled with a 500um
dredge net. Samples collected with the dredge net provide an accurate estimate of the
relative abundance of epibenthic animals in farm dams (Barlow et al., 1982). To sample
the dam containing milfoil the net was modified by removing the kick chain and attach-
ing a rake to direct weed under the net (Topp, 1967). Ten samples were collected from
each dam six days before the fish were sampled. The macroinvertebrates in all samples
were later identified and counted. Three plankton samples were collected from the dam
containing silver perch using an 100um plankton net towed horizontally for 15m just
below the surface.

The diet of the fish was compared with the available foods using the prey-selection
index, C, which is statistically testable for any degree of selection at any sample size
(Pearre, 1982). C is zero valued for no selection and has the limits -1 for complete selec-

tion and -1 for complete avoidance. Statistical tests were conducted using x? tests
(method 3 of Pearre (1982) ).

POLYCULTURE TRIAL

The dam was stocked with 14 golden perch and 28 silver perch in November 1981
and harvested in April 1982. The fish were transported to the laboratory and dissected
as described above. The diets were analysed by determining the percentage volume
occupied by each food item in each stomach, as recommended by Wallace (1981).

In addition to determining the degree of interspecific overlap, the degrees of in-
traspecific overlap were also calculated to ascertain how well the diets of each species
were characterized (Wallace and Ramsey, 1983). Specimens of each species were
randomly divided into two sets and the dietary overlap calculated. This procedure was
repeated 25 times for each species, and the means and standard deviations computed.

The degree of dietary overlap was determined using Schoener’s index

a= 1-05( o-Ps)
i=]

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

Cc. G. BARLOW, R. MCLOUGHLIN AND K. BOCK 145

where f,; is the proportion of food item 7 in the diet of group x, f,; the proportion of food
item 7 in the diet of group y and n the number of food categories. Computed values range
from 0 for no overlap to 1 for complete overlap. Although there is no statistical method
for judging the reliability of overlap (Wallace and Ramsey, 1983), it is generally con-
sidered to be biologically significant when the value exceeds 0.60 (Zaret and Rand,
OTA

ALIMENTARY TRACT MORPHOLOGY

Analyses were conducted on all one year old fish from the monoculture experiment
and a further 15 two year old golden perch and 20 two year old silver perch. All measure-
ments were taken to the nearest millimetre on fresh specimens. ‘Total length was
measured, the digestive tract was then dissected out, laid on a dry enamel dish, and the
lengths of the stomach and intestine-rectum measured immediately. The relationships
between the gut length (oesophagus to anus) and total length were determined, and the
ratios of intestine-rectum length : stomach length were calculated. The dentition and
gill rakers were examined and illustrated.

RESULTS

MONOCULTURETRIAL

The food organisms sampled from each dam, the dietary analysis and C values for
golden perch and silver perch are given in Tables 1 and 2 respectively. The values of C
indicate selection or avoidance of a food type, but the reality of selection or avoidance is
shown only by the level of significance of C. Identification of the macroinvertebrates was
usually possible to the species level, with the exception of the Notonectidae which could
not be identified beyond family, and larval insects which usually could not be identified
below order. The greater diversity of insects in the dam containing golden perch was
probably due to the presence of macrophytes.

Golden perch consumed a wide range of macroinvertebrates. Corixid nymphs,
which comprised about 50% by volume of the diet, were the major food. The most
preferred, or actively selected, organisms were notonectids and the corixid Agraptocorixa
eurynome. Three other comparatively abundant corixids were avoided, even though one,
Agraptocorixa parabiopunctata, was similar in size to A. eurynome (approximate total length
of adults 7mm and 9mm respectively). No zooplankton was found in the stomachs of
golden perch.

The stomach contents of silver perch comprised 80% zooplankton, with the re-
mainder being macroinvertebrates, allochthonous plant material and gravel. The
stomach contents of 16 fish consisted entirely of zooplankton. There was comparatively
little zooplankton in the intestines compared with the stomachs, but this is to be ex-
pected because of the rapid digestion of zooplankton. The percentage composition of
zooplankton consumed by silver perch differed markedly from that collected from the
dam, as shown below:

Cladocera Copepods Ostracods
Consumed by silver perch 78% 22% trace
Collected from dam 26% 74% —

However, it is not known if these samples, collected from just below the surface of the
dam, accurately represented the relative abundance of the zooplankton groups.

PROC. LINN. SOC. N.S.W,, 109 (3), (1986) 1987

146 FEEDING HABITS OF PERCH

TABLE 1

Total number of each food type sampled from the dam (Na) and found in the diet (Nd) of golden perch (TL. 224+ 35mm, Wt.
188 + 100g) reared in a monoculture trial; the prey selection index C and the level of significance for C.
N.S. not significant, ** P<0.01, *** P<0.001

Significance
FOOD TYPE Na Nd C of C
Notonectidae 57 129 0.234 es
Corixidae
Sigara spp. 640 24 -0.056 eee
Agraptocorixa eurynome 321 345 0.317 toe
Agraptocorixa parabiopunctata 157 0 -0.039 se
Micronecta annae group 3983 3 -0.246 see
Nymphs 4099 712 0.101 se
Dytiscidae
Sternopriscus multimaculatus 543 1 -0.075 eed
Megaporus howutti 41 Ase) 0.120 bps
Antiporus gilberti 42 1 -0.014 N.S.
Necterosoma wallastoni 9 0 -0.005 N.S.
Hydrophilidae
Spercheus sp. 3 0 -0.003 N.S.
Laccobius sp. 0 1 0.011 N.S.
Unidentified sp. 0 1 0.011 N.S
Hydracarina 274 1 -0.051 ae
Hydraenidae 0 1 0.011 N.S.
Atyidae
Paratya australiensis 3 0 -0.003 N.S.
Atheriniformes
Gambusia affinis 2 0 -0.006 N.S.
Mollusca
Physa sp. 12 0 -0.007 N.S.
Larval insects
Trichoptera 35 22 0.059 ieee
Odonata 2 3 0.026 ee
Coleoptera a 8 0 -0.004 N.S.
Coleoptera, Hydrophilidae 12 4 0.013 N.S.
Diptera, Culicidae 35 2 -0.007 N.S.
Diptera, Chironomidae 6 0 -0.002 N.S.
Ephemoptera a 18 0 -0.010 N.S.
Ephemoptera b 62 0 -0.023 Se
Ephemoptera, Baetidae 205 5 -0.036 che
Lepidoptera, Pyralidae 3 1 0.001 N.S.
Plecoptera 94 57 -0.010 N.S.
TOTAL NUMBER 10666 1312

Silver perch selectively fed on notonectids, but avoided both crayfish, Cherax destruc-
tor and mosquitofish, Gambusza affinis. Of the macroinvertebrates eaten by silver perch,
45 were found in the intestines and 15 in the stomachs. The large proportion of macro-
invertebrates in the intestine possibly indicates a diel feeding pattern or perhaps dif-
ferent rates of passage through the stomach and intestine.

POLYCULTURE TRIAL
The intraspecific dietary overlap value for golden perch was 0.80+0.06 and for

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

C. G. BARLOW, R. MCLOUGHLIN AND K. BOCK 147

TABLE 2

Total number of each food type sampled from the dam (Na) and found in the diet (Nd) of silver perch (T:L. 205+ 23mm, Wt.
111 + 36g) reared in a monoculture trial; the prey-selection index C and the level of significance for C.
N.S. not significant, * P<0.1, ** P<0.01, *** P<0.001

Significance

FOOD TYPE Na Nd Cc of C
Notonectidae 29 41 0.305 ee
Corixidae

Sigara sp. 0 4 0.156 x

Nymphs 0 4 0.156 a
Dytiscidae

Antiporus gilberti 3 2 -0.027 N.S.
Larval insect 1 4 0.106 N.S.
Parastacidae

Cherax destructor 11 0 -0.230 a
Atheriniformes

Gambusia affinis 34 4 -0.391 aun
TOTAL NUMBER 78 59

silver perch 0.84+0.04. These values indicate that the diet of each species was well
characterized, even though there were comparatively few fish in the samples. Thus, it is
valid to use the present data to compare the diets of these species.

The interspecific dietary overlap value was 0.23, indicating that the diets of the two
species were significantly different. The major items consumed by golden perch were
trichoptera larvae (63%) and crayfish (14%). In contrast, the major foods of silver perch
were chironomid larvae (34%), cladocera (14%) and ostracods (10%), while trichoptera
larvae formed only 7% of the diet and crayfish were absent from the silver perch
stomach contents (Fig. 1).

ALIMENTARY TRACT MORPHOLOGY

The shape of the mouth and dentition of the two species are illustrated in Fig. 2.
Golden perch has a large mouth, and possesses teeth on the upper and lower jaws,
vomer, palatines and roof and floor of the pharynx. The teeth are numerous, tiny and
stout. All teeth are set in bony plates. In contrast, silver perch has a comparatively small
mouth, and possesses teeth on the upper and lower jaws and the roof and floor of the
pharynx. The villiform teeth are conical and pointed, and generally aligned to point
posteriorally. An exception to this is the outer band of larger teeth on the premaxillary,
which point ventro-anteriorally and in some instances protrude slightly beyond the lips.
The premaxillary and mandibular teeth are set in bony plates while the upper and lower
pharyngeal teeth are embedded in fleshy pads.

In both species, gill rakers form an anterior and posterior series on all four gill
arches. The anterior rakers on the first arch are elongated, whereas the posterior series
on the first arch and all rakers on the other arches are shorter (Fig. 3). The gill rakers of
golden perch are short and firm and covered with tiny tubercules which provide a rough
surface. The gill rakers of silver perch are finer and adorned with rows of villiform teeth
on both margins of the flat edge of the rakers facing the pharyngeal cavity (Fig. 3). The
arrangement of the rakers is such that those on adjacent arches are interposed when the
arches are brought together.

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

148 FEEDING HABITS OF PERCH

Golden Perch

volume

Trichoptera
Crayfish
Coleoptera
Odonata
Corixidae
Chironomidae
Cladocera
Ostracoda
Copepoda
Plant material
Unidentified

Percentage

30 Silver Perch

Food item
Fig. 1. Diets (average of the percentage volume of individual fish) of golden perch (T.L. 286+ 20mm, Wt
289 + 65g) and silver perch (T.L. 315 +12mm, Wt 429 + 60g) reared in a polyculture trial.

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

C.G. BARLOW, R. MCLOUGHLIN AND K. BOCK 149

The relationships between gut length and total length were linear for both species
(Fig. 4), and were described by the following equations:

Golden perch, total length range 155-292mm,
Giy— 04980 — 29:5) (er =0184n—435 <= 0:001)
Silver perch, total length range 175-328mm,
GL =1260L,— 106 (@=0:93, n=45, p< 0.001)
where GL = gut length and TL = total length.

The ratio of intestine-rectum length : stomach length for golden perch was
2.34 +0.34, and for silver perch it was 6.75 £1.31. That is, the alimentary tract of silver
perch was longer than that of golden perch; this difference was due to the relatively
longer intestine-rectum of the silver perch, rather than a difference in the size of the
stomachs of the two species.

DISCUSSION
The advantage of employing a statistically measurable prey-selection index for
comparing diets and potential prey species was evident in the monoculture trial. By

GOLDEN PERCH Roof of mouth Floor of mouth

u.ph
I.ph

SILVER PERCH

)

Fig. 2. Shape of the mouth and arrangement of teeth in golden perch and silver perch. (md = mandibular,
pmx = premaxillary, v = vomerine, p = palatine, l.ph = lower pharyngeal, u.ph = upper pharyngeal).

pmx

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

150 FEEDING HABITS OF PERCH

GOLDEN PERCH SILVER PERCH

First gill arch

Gill rakers

Fig. 3. The lower limb of the first gill arch of golden perch and silver perch, and magnified gill rakers showing
the typical shape of the posterior rakers on the first gill arch and the anterior and posterior rakers on the se-
cond, third and fourth arches.

using C it was possible to determine which prey species were being significantly selected
or avoided by the fish. Furthermore, realistic values of C could be computed even when
a particular food type was absent in either the diet or the environment.

Preference ratings, however, apply only at the time of sampling; the most preferred
species tend to be depleted first and the remaining species affected more or less severely
according to their preference ranking and the intensity and duration of cropping
(Petrides, 1975). One possible example of depletion of preferred species is the notonec-
tids, which were actively selected by both golden perch and silver perch in the mono-
culture trial. However, notonectids were absent from the diets of both species in the
polyculture trial, and thus presumably absent from the dam, even though notonectids
are the numerically dominant insect group in farm dams in the study area (Barlow and
Bock, 1981).

In the monoculture trial, golden perch fed on a wide variety of insects and the silver
perch fed predominantly on zooplankton. Strict comparison of the diets was not possible
because of the different available food sources. However, these apparent dietary differ-
ences were real, as indicated in the polyculture trial, in which golden perch fed mainly
on trichoptera larvae and crayfish, whereas silver perch ate zooplankton and
chironomid larvae. The diet of golden perch in this study agrees with published infor-
mation, but the diet of silver perch indicates greater consumption of zooplankton than
previous observations on the stomach contents of wild fish had indicated (Merrick and
Schmida, 1984).

The morphology of the alimentary tracts also suggests that the diets of the two
species are different. The large mouth of golden perch is obviously adapted for taking
large prey. The numerous, tiny teeth set in bony plates, together with the stout, hard gill
rakers, would aid crushing of the prey. The short intestine is also typical of a carnivore

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

C. G. BARLOW, R. MCLOUGHLIN AND K. BOCK Lay

300
Z ane Silver Perch
f=
ahd
=
Cc
o
od
>}
(O)
100
Golden Perch
O
100 200 300 400

Total length (mm)

Fig. 4. Regression of gut length on total length for golden perch and silver perch.

(Das and Moitra, 1956). In contrast, silver perch has a small, terminal mouth with pre-
maxillary teeth which are apparently capable of rasping aufwuchs from solid substrates.
Aufwuchs was not present in any of the dams in this study, but periphyton has often
been observed in the gut contents of silver perch from other waters (Barlow, unpublished
data). Of particular interest are the villiform teeth on the gill rakers of the silver perch.
These teeth form a very effective sieving device when the gill arches are brought
together. Presumably, this is the mechanism silver perch uses to capture zooplankton.
The comparatively long intestine of silver perch 1s also indicative of an omnivorous diet
(Das and Moitra, 1956).

In conclusion, golden perch can be classified as a macrophagic carnivore eating in-
sects and crustaceans, whereas silver perch is an omnivore eating mainly zooplankton,
insects and aufwuchs if available. Such dietary differences indicate that these species are
well-suited for rearing together in polyculture, at least in unmanaged farm dams.
Although both species have many of the biological attributes necessary for successful
aquaculture (Barlow, in press), research on production levels attainable in intensively

PROC. LINN. SOG. N.S.W., 109 (3), (1986) 1987

if2 FEEDING HABITS OF PERCH

managed ponds is necessary before any determination can be made regarding the
economic feasibility of farming these fishes.

ACKNOWLEDGEMENTS

This study was partly funded by the Reserve Bank’s Rural Credits Development
Fund. We thank staff at the Inland Fisheries Research Station, L. Halbisch and
J. McDougall for the use of their dams, H. Favier and J. Riches for preparing the
figures, Drs P.L. Cadwallader and S.J. Rowland for commenting on a draft of the paper,
and D. MacIntyre for typing the manuscript.

References

BARLOW, C. G., in press. — Fish in farm dams and implications for extensive aquaculture. Jn: L. F.
REYNOLDS, (ed.), Proceedings of the First Freshwater Aquaculture Workshop, Narrandera, NSW.

, and Bock, K., 1981. — Fish in Farm Dams. Sydney: N.S.W. State Fisheries, 172 pp.

——., LEEDow, M., and MCLOUGHLIN, R., 1982. — Macroinvertebrate sampling using a dredge net in a
farm dam in southwestern New South Wales. Aust. Zool. 21(1): 97-104.

Das, S. M., and Moitra, S. K., 1956. — Studies on the food of some common fishes of Uttar Pradesh, India:
Part II. On the types of fish-food and the variations in the relative length of the alimentary canal, with
a description of the latter. Proc. natl Acad. Sci. India 27(4): 213-223.

MERRICK, J. R., and SCHMIDA, G., 1984. — Australian Freshwater Fishes: Biology and Management. Sydney; John
R. Merrick.

PEARRE, S., 1982. — Estimating prey preferences by predators: uses of various indices, and a proposal of
another based on x2. Can. J. Fish Aquat. Sci. 39: 914-923.

PETRIDES, G. A., 1975. — Principal foods versus preferred foods and their relations to stocking rate and
range condition. Biol. Conserv. 7: 161-169.

ROWLAND, S. J., in press a. — The hormone-induced spawning and larval rearing of Australian native fresh-
water fish, with particular emphasis on the golden perch, Macquaria ambigua. In: L. F. REYNOLDS,
(ed.), Proceedings of the First Freshwater Aquaculture Workshop, Narrandera, N.S.W.

——., in press b. — Design and operation of an extensive aquaculture system for breeding warmwater fishes.
In: L. F. REYNOLDS, (ed.), Proceedings of the First Freshwater Aquaculture Workshop, Narrandera,
N.S.W.

——.,, DirROou, J. F., and SELOsseE, P. M., 1983. — Production and stocking of golden and silver perch in
N.S.W. Aust. Fish. 42(9): 24-28.

Topp, R. W., 1967. — An adjustable macroplankton sled. Prog. Fish-Cult. 29(3): 184.

WALLACE, R. K., 1981. — An assessment of diet-overlap indexes. Trans Am. Fish. Soc. 110: 72-76.

, and RAMSEY, J. S., 1983. — Reliability in measuring diet overlap. Can. J. Fish. Aquat. Sci. 40: 347-351.

ZARET, T. M., and RAND, A. S., 1971. — Competition in tropical stream fishes: support for the competitive
exclusion principle. Ecology 52(2): 336-342.

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

Distribution and ‘Iaxonomy of the Long-eared
Bats, Nyctophilus gould: ‘lomes, 1858 and
Nyctophilus bifax ‘Thomas, 1915 (Chiroptera:

Vespertilionidae) in eastern Australia

H. E. PARNABY

PARNABY, H. E. Distribution and taxonomy of the long-eared bats, Nyctophilus gouldi
Tomes, 1858 and Nyctophilus bifax Thomas, 1915 (Chiroptera: Vespertilionidae) in
eastern Australia. Proc. Linn. Soc. N.S.W. 109(3), (1986) 1987: 153-174.

Although recently synonymized with N. gould, N. bifax is shown to be a distinct
species separable on a number of external, cranial and bacula features. Cranial and ex-
ternal characters examined show that both species are similar in size, but can be readily
identified in the field on the basis of fur colour, relative ear size, and details of the snout.
Striking differences exist in external phallic morphology between each species.

Additional data on the distribution of N. bifax and N. gould: have resulted from
recent field work and a re-evaluation of material in Australian collections. The distri-
bution of both species is summarized. The known distribution of N. gould: is extended
north by about 1100km to Atherton, north Queensland. N. bifax is recorded from a
number of new localities south to Iluka, New South Wales, representing a southern
range extension of some 750km. These species are therefore sympatric over 1600km in
eastern Australia.

H. E. Parnaby, School of Zoology, University of New South Wales, PO. Box 1, Kensington,
Australia 2033, manuscript received 20 August 1986, accepted for publication 17 December 1986.

INTRODUCTION

Six species of the australo-papuan bat genus Nyctophilus are generally recognized
from Australia (Hall, 1984): N. geoffroy:, N. timoriensis, N. gouldi, N. bifax, N. arnhemensis,
and N. walker. This paper considers the distribution and status of eastern Australian
populations of N. gould: and N. bifax, and is part of a continuing taxonomic study of
Nyctophilus.

Historically, the taxonomic status of N. gould: has been subject to controversy.
Tomes (1858) described N. gould: from specimens from Moreton Bay, Queensland, and
Bathurst, New South Wales. Iredale and Troughton (1934) considered N. gouldi to be a
southeastern Australian subspecies of the widespread N. tzmoriensis. ‘Tate (1952) regarded
N. gould: to be a distinct species. However, until recently the arrangement of Iredale and
Troughton (1934) has been followed by most authors. There is now general agreement
that N. gould: and N. timoriensis are distinct species (Hall and Richards, 1979; Allison,
1982; Koopman, 1984) and are known to be sympatric at several localities in eastern
Australia (Parnaby, unpublished). Richards (1983) summarized the known range of N.
gould: as southwestern Western Australia, Tasmania, Victoria, eastern New South Wales
and southeastern Queensland as far north as Bundaberg.

Thomas (1915) described N. bzfax based on material from the Torres Strait islands
and north Queensland. As of 1983, N. bifax was recorded from northern Western Aus-
tralia, the Northern Territory and across to northeastern Queensland as far south as
Sarina (Allison, 1983). Within this range, two subspecies are commonly recognized: N.
bifax bifax from Queensland, and WN. 0. daedalus from the Northern Territory and Western
Australia.

Koopman (1984) believed that the criteria proposed by Thomas for the separation
of N. bifax from N. gould: were not of sufficient magnitude to warrant species status and,

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

154 LONG-EARED BATS

in view of the wide separation of their then known distribution, he tentatively placed N.
bifax as a subspecies of N. gould:. Based on the distribution data of Hall and Richards
(1979), Koopman (1984) believed that the distribution of N. bifax and N. gould: were
separated by approximately 1000km in eastern Australia.

Koopman (1984), uncertain about the status of daedalus, placed it with N. gouldz,
thereby tentatively proposing a single species with three allopatric subspecies: N. gouldi
gould: from southern Australia (but not Tasmania), N. gould: bifax from north Queens-
land and N. gouldi daedalus from northern Western Australia and the Northern Territory.

Churchill et al. (1984) record a considerable southern extension of the known range
of N. bifax, to the Pilbara region of Western Australia, and to Rockhampton in Queens-
land, thus closing the gap in the known range of this species and that of N. gouldi to
approximately 1000km in Western Australia, and some 300km in Queensland.

Field work conducted by the author and others over the last five years, combined
with a reexamination of existing Australian research collections, has resulted in further
insight into the taxonomic status and distribution of these taxa. Morphologically, N.
gould: and N. bifax have been found to be readily distinguishable, both in the field and by
examination of preserved material. This has facilitated the recognition of many new
locality records including major range extensions of both species. Significantly, elec-
trophoretic analysis of tissue samples collected from areas of sympatry in eastern Aus-
tralia has provided unequivocal evidence of the distinct separation of N. bifax and N.
gould: (M. Adams and P. Baverstock, pers. comm.).

In this paper, the distribution of each species is summarized and new distribution
records are presented which indicate large scale sympatry between N. bzfax and N.
gouldi. The major external, cranial and bacula features useful for distinguishing each
species from throughout their range in eastern Australia are discussed.

Abbreviations used for research collections from which material has been
examined are: Australian Museum, Sydney (AM); Australian National Wildlife Collec-
tion (CSIRO), Canberra (CM); Museum of Victoria, Melbourne (MV); South Aus-
tralian Museum, Adelaide (SAM); Queensland Museum, Brisbane (QM) and Western
Australian Museum, Perth (WAM).

DISTRIBUTION

N. gouldi

As noted above, this species is recorded from southwestern Western Australia, Vic-
toria through eastern New South Wales, to southeast Queensland (Richards, 1983).
Richards also records this species from Tasmania but the specimens on which this is
based belong with the large Tasmanian form of N. geoffroyi (see Discussion).

Previously, the most northern record of this species in eastern Australia was from
near Gin Gin, Queensland (Thomas, 1915). Examination of museum material, and
recent field collecting, have resulted in a substantial expansion of the known range to
Atherton, 1100km north from Gin Gin.

The known distributional limits of N. gould in eastern Australia are shown in Fig.
la and localities are listed in Appendix 1. The species is usually considered to be an in-
habitant of mesic eucalypt forests of the Great Dividing Range. Hall and Richards
(1979) record the species from inland southern Queensland. Additional records
presented here from inland Queensland and New South Wales and field work in New
South Wales indicates that N. gould: is widespread and probably more common through-
out lower rainfall regions than was previously realized. Thus this species appears to

PROC. LINN. SOC. N.SW., 109 (3), (1986) 1987

H. E. PARNABY 155

Fig. 1. Distribution of a, N. gould: and b, N. bifax in eastern Australia.

occur in a range of habitats, including semi-arid woodland (such as in the Charters
Towers region), brigalow country and sclerophyll forest.

N. bifax

The original description of N. bifax by Thomas (1915) was based on material from
localities ranging from ‘Torres Strait islands, south to Herberton and west to Cloncurry,
northwestern Queensland. In eastern Australia, there are few published records of WN.
bifax south of Townsville; Allison (1983) depicts this species as extending south to about
Sarina and Churchill et al. (1984) record N. bifax at Byfield, near Rockhampton. Exam-
ination of museum material, some of which had previously been misidentified, has
resulted in a number of additional locality records in near coastal areas extending south
from Townsville to southern Queensland.

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

156 LONG-EARED BATS

y Wy hh

Fig. 2. Dorsal and lateral views of the skull and lateral view of the dentary of a, N. gould: from Brisbane (OM
J]©M5366, female); b, N. bifax from Atherton, Old (AM M16181, female). Bar represents 4mm.

During field work in north coastal New South Wales, N. bifax was one of the most
frequently trapped species of insectivorous bat. It was captured at low elevations at a
number of localities as far south as Iluka (Parnaby, 1986), which is currently the most
southern record for the species.

PROC. LINN. SOC. N.SW., 109 (3), (1986) 1987

H. E. PARNABY

Fig. 3. Stereopairs of ventral view of a, N. gouldi from Brisbane (QM JM5366 female); b, N. bifax from Ather-
ton (AM M16181, female). Bar represents 4mm.

PROC. LINN. SOC. N.S.W., 109 (3), (198

158 LONG-EARED BATS

Fig. 4. Photographs illustrating differences in relative ear size between males of a, N. gould: from Broken
Head, N.S.W. (AM M13389) and b, N. bifax from Brunswick Heads, N.S.W. (AM M13388). Bar represents
lcm.

The updated distribution of N. dzfax in eastern Australia is shown in Fig. 1b, and
localities are listed in Appendix 1. With the exception of a specimen from the Cloncurry
area (20° 42’S, 140° 30’E) reported by Thomas (1915) and one from the Einasleigh
River (18° 11’S, 144° 00’E, AM M13356), most records are close to the coast. Allison
(1983) notes the occurrence of this species in a variety of habitats ranging from rain
forest to dry sclerophyll woodlands.

MORPHOLOGICAL DIFFERENCES

Substantial between-locality variation exists within each species throughout
eastern Australia, particularly so in N. gould: (see ‘Metric variation’ below). Despite this,
diagnostic external and cranial criteria discussed in the following sections appear to
hold, irrespective of locality.

Skull morphology

Compared with WN. bifax the skull of N. gouldi is relatively narrow for its length, and
is more slightly built (Fig. 2). In N. gould the bullae are relatively much larger, and con-
sequently appear to be closer together medially (Fig. 3). The hamular processes of the
pterygoids are slightly more ossified in N. gould:. The paroccipital processes are slightiy
more pronounced in WN. bifax and are more clearly distinct from the occipital condyles.

The upper canines are laterally splayed in N. bifax yet in N. gould: they are usually in
line with the upper tooth row, however, occasional specimens of N. gould: with splayed
canines have been observed.

Like the skull, the dentary of N. gould: is more delicately built and is shallower. The
upward inflection of the postero-ventral border of the ramus is more marked in N. bzfax
and the angular process is generally shorter than that of N. gould: (Fig. 2).

External Morphology

The most obvious external differences between these two species are in the relative
size of the ears and general fur colour. Relative to body size, the ears are very large in N.
gould: (Fig. 4), while in N. bifax they are distinctly shorter.

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

H. E. PARNABY 159

Fig. 5. Frontal and frontolateral views of the snout of a, and b, N. gould: from Border Ranges, N.S.W. (AM
M13412, male); c, andd, N. bifax from Brunswick Heads (AM M13388,male), showing differences in the post-

nasal prominence. Bar represents 4mm.

From field observations, the dorsal pelage of N. gould: is predominantly a slate grey
or grey-brown with ash-grey ventral fur often mottled with very light buff. In N. bifax,
the dorsum is light brown or tannish, with a lighter tannish undersurface.

There is a consistent difference in the shape of the post-nasal bump, as originally
noted by Thomas (1915). Relative to the noseleaf, this structure is more developed in N.
gould: than N. bifax (Fig. 5) and in N. gould has a faint vertical groove which is weaker or
absent in N. bifax. The specimens illustrated in Fig. 5 differ in the shape of the dorsal
margin of the noseleaf; there is a median concavity in N. gould: while the margin of N.
bifax is evenly convex. However this difference is not diagnostic as the shape of the
noseleaf is variable in both species.

Camera lucida drawings of a representative glans penis of an adult of each species
are shown in Fig. 6. Each species differs in a number of obvious features. The lateral
surfaces in N. gould are constricted in the mid-line, dividing the penis into a distinct

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

160 LONG-EARED BATS

a b Cc d

Fig. 6. Camera lucida drawings of the glans penis with prepuce removed. Lateral view of a, N. gould: from
Border Range, NSW (AM M13412), b, N. bifax from Brunswick Heads, NSW (AM M13388). Ventral view of
c, N. gould and d, N. bifax. Bar represents 1.0mm.

dorsal and ventral element. The dorsal element forms a very distinct distal ‘beak. In
marked contrast, the penis of N. bifax is approximately cylindrical, and the head of the
penis is more truncated. Consequently, each species has a very characteristic profile
which 1s clearly visible to the unaided eye.

Bacular morphology

As noted by Thomas (1915) N. bifax differs from N. gould: in bacular morphology: in
N. gould: the baculum comes to a simple point distally whereas in N. bzfax the tip 1s bifur-
cated by acrescentic notch.

Using X-ray photography, I have examined bacula from 17 specimens of N. bifax
from localities scattered throughout the complete eastern Australian range of this
species, from Iluka north to Lockerbie Scrub. The bifid condition is present in every
specimen. Likewise, the sharp distal point 1s present in all 25 examples of N. gouldi ex-
amined from widespread locations throughout Victoria, New South Wales and Queens-
land. Thus, the condition of the distal tip of the baculum appears to be monomorphic in
each species.

Metric Variation

A univariate and multivariate analysis of metric variation using analysis of vari-
ance, principal components analysis and canonical variates analysis of these and other
species of Nyctophilus is in progress, and preliminary findings are outlined here.

Ten external dimensions were taken from 130 spirit-preserved specimens of N.
gould: from central and northern New South Wales and Queensland, and 118 spirit speci-
mens of N. bifax bifax from throughout Queensland and northern New South Wales
(listed in Appendix 1). Twelve cranial measurements were made on 69 skulls of N. gouldi
and 52 of N. bifax bifax, all from throughout New South Wales and Queensland.

Sexual size dimorphism was found in the majority of dimensions in both species,
with females on average larger than males. Consequently, sexes were treated separately
in all analyses.

PROC. LINN. SOC. N.SW., 109 (3), (1986) 1987

H. E. PARNABY 161

(mm.)

DISTANCE

INTER-BULLAE

3.5 4.0 4.5

BULLA LENGTH (mm.)

Fig. 7. Plot of interbullae distance (mm) against bulla length (mm) for 65 N. gould: and 50 N. bifax from central
and northern NSW and Qld. Both dimensions were measured in ocular units with a dissecting microscope,
and the axes converted to mm. Circles represent males, squares females. Numerals indicate overlapping
points.

When specimens from all localities are pooled, mensural ranges overlap between
each species for all external and all but one of the 12 cranial and dental dimensions.
Measurements with overlapping ranges are: ear length, measured from the notch;
length of first digit; forearm length; third digit: length of metacarpal I, length of
metacarpal II, length of metacarpal III; fifth digit: length of metacarpal I, II, and III;
length of hind-leg; condylobasal length; greatest length of skull; length of upper tooth
row (C!-M?); zygomatic breadth; interorbital constriction; outer breadth of upper third
molars; height of braincase; skull breadth at mastoids; interbullae distance; bulla
length, and basicranial length (measured from the anterior margin of foramen magnum
to the anterior border of the posterior palatal emargination).

Interbullae distance is the only measurement in which the range for each species do
not overlap: the range for N. gould: being 2.0-2.5mm (n=65) and N. bzfax 1.3-1.8mm

PROG. LINN. SOC. N.S.W., 109 (3), (1986) 1987

162 LONG-EARED BATS

(n=50). A plot of this dimension against bulla length for all localities combined (Fig. 7)
results in a clear separation of N. bifax and N. gouldt.

In both species substantial size differences were found between localities. Although
analysis of these data is preliminary, observed variation seems to correspond with local
climatic conditions. This is evident in a plot of forearm length against longitude for sam-
ples of N. gould: from diverse sites throughout New South Wales and Queensland (Fig.
8), where increasing longitude approximates decreasing aridity. For example, mean
forearm length of a sample of female N. gould: from a dry inland site (Pilliga Scrub,
NSW) is 38.7mm compared with 45.8mm for females from a higher rainfall site in the
Great Dividing Range at a comparable latitude (Doyles River State Forest).

The between-locality size variation evident in both species might be expected to
obscure differences present within a given locality. In an attempt to reduce these locality

153

152

151
Ww
a
> 150
=
©
5
149
~—n
wi
wi
wi ««:148
©
bad
a

147

146

FOREARM LENGTH (mm)

Fig. 6. Forearm length (mm) of N. gouldi versus longitude for samples from throughout NSW and Old showing
a trend of increase in size with longitude. Circles represent female sample means, dots male samples. Lines
indicate sample range, sample size in brackets. Localities are, 1. Tweed Rge, northern NSW; 2. Gloucester
Tops, central NSW; 3. Doyles River State Forest, central NSW; 4. Olney State Forest, central NSW;
5. Cassilis region, central NSW; 6. Pilliga State Forest, northern NSW; 7. Tambo region, south-central Qld;
8. Capella, central Old; 9. Henty region, southern NSW, 10. Mt Leysham, north Old. (from Churchill et a/.,

1984).

PROC. LINN. SOC. N.SW., 109 (3), (1986) 1987

H. E. PARNABY 163

effects, measurements of specimens from a restricted geographic region were compared.
The only area of sympatry for which specimens are available in any numbers is north-
eastern New South Wales and far southeast Queensland. Twenty-six specimens of N.
gould: and 23 of N. bifax were examined from localities ranging from the Conondale
Ranges, south to Iluka in New South Wales and inland to Tooloom. Only three of these
specimens are from north of Brisbane.

Measurements of spirit specimens from within this region were pooled by sex for
each species. Summary statistics for external dimensions of each species are given in
Table 1, and dimensions of 19 skulls of N. gould: and 11 of N. bifax from the same region
are given in Table 2.

TABLE 1

Summary statistics for external dimensions (mm) of spirit specimens of N. bifax and N. gouldi from northern N.S.W.
and southeastern Qld
Sample sizes forall dimensions are: N. bifax (9 males, 14 females), N. gould: (10 males, 8 females)

MEAN SEs S.D. RANGE CV

EAR LENGTH MALES

bifax 24.78 .43 OD HO 9B7 5.20

gouldi 28.33 44 1.40 25.8-30.0 5.78

FEMALES

bifax 25 ee 100) 23) 529633 3.58

gouldi 28.55 .29 Bie 0 BBO. 2.83
FOREARM MALES

bifax 41.46 34 p10 39324205 2.49

gouldi 42.24 43 135) es Om 24409 3.62

FEMALES

bifax HDI} 98} 87 41.0-44.4 2.06

gouldi ABNG © 9 BY 1.46 41.2-45.2 3.38

DIGIT I MALES
bifax
gouldi
FEMALES
bifax
gouldi

DIGIT 3, METACARPAL I MALES
bifax
gould
FEMALES
bifax : : 37. 7-41.20
gouldi ‘ : BOR2— 4289)

DIGIT 3, METACARPAL II MALES
bifax ; : 14.9-16.3
gouldi : 3 12, JNO}
FEMALES
bifax ; : 1D, = G8)
gouldi 7 : 15.2-18.0

DIGIT 3, METACARPAL III MALES
bifax
gould
FEMALES
bifax
gouldi

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

164 LONG-EARED BATS

TABLE 1 (continued)

DIGIT 5, METACARPAL I MALES
bifax : 37.4-40.7
gouldi ; ; 37.2-42.6
FEMALES
bifax ; ‘ 38.8-41.4
gouldi : 5 38.9-42.8

DIGIT 5, METACARPALII MALES
bifax 5 : 10.0-11.2
gouldi ; ; NO2=N2
FEMALES
bifax i ; 10.4-11.4
gouldi : 10.6-12.2

DIGIT 5, METACARPALIII MALES
bifax
gouldi
FEMALES
bifax
gouldi

HINDLEG LENGTH MALES
bifax
gouldi
FEMALES
bifax
gouldi

TABLE 2

Summary statistics for skull and dental dimensions of N. bifax and N. gouldi from northern N.S. W. and far southeastern
Qld
All dimensions are in mm. Note that interbullae distance, bulla length and basicranial length were measured
in occular units (1 unit = 0.082mm), and converted to mm

MEAN

CONDYLOBASAL LENGTH MALES

bifax ) 15.30 - 13 29 11429 [Bo 1.89
gould 11 O33 Oommen oO) 15.9-17.0 2.26
FEMALES

bifax

gouldi

GREATEST LENGTH MALES

hifax PIO I Oe MIG OsLO i192
gouldi {i 17.30. 313 042)” 73-1
FEMALES

hifax 6 16.93 .080

16.6-17.1
gould =

cm! LENGTH MALES
bifax 3) 22 AO 7 084 6.1-6.3 1.35
gouldi 11 6.57 .060 .20 6.3-7.0 3.04
FEMALES
bifax

gouldi

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

c!_c! BREADTH

ZYGOMATIC BREADTH

INTERORBITAL CONSTRICTION

BREADTH M> >

BRAINCASE HEIGHT

MASTOID BREADTH

INTERBULLAR DISTANCE

BULLAR LENGTH

BASICRANIAL LENGTH

H. E. PARNABY 165

Table 2 (continued)

MALES

bifax
gould
FEMALES
bifax
gouldi

MALES
bifax
gould
FEMALES
bifax
gouldi

MALES
bifax
gouldi
FEMALES
bifax
gouldi

MALES
bifax
gouldi
FEMALES
bifax
gould

MALES
bifax
gould
FEMALES
bifax
gould

MALES
bifax
gouldi
FEMALES
bifax
gouldi

MALES
bufax
gould
FEMALES
bifax
gould

MALES
bifax
gouldi
FEMALES
bifax
gouldi

10.2-10.8
10.0-11.1

10.5-10.9
10.1-10.9

359) eer .092 34-3). 2.56
AO BK gil 3.8=4.3 2.95
D209) AB) 0H Sho 328 2.65
A089 2053 14 3.9-4.3 3243
5 M57 tS 5.4=5.7 2.70
Git oO) 1G) 5.8-6.6 Selb
DE Ommen OO, wiles De 7010) Deh}
6.00 .072 oY) IJ =0..3) BL i7/

PROG. LINN. SOC. N.S.W., 109 (3), (1986) 1987

166 LONG-EARED BATS

The ranges of ear length for females of each species do not overlap; this separation
is shown in Fig. 9. For the remaining nine external measurements however, there is ex-
tensive overlap between each sex of each species. For all external dimensions except ear
length, the two species have similar mean values for equivalent sex. As there is nearly
complete overlap in the ranges of forearm length between both species, a bivariate plot
of ear length against forearm length (frequently used in bat systematics) is not useful for
separating males of these species (Fig. 9).

LENGTH (mm)

EAR

40 42 44 46

FOREARM LENGTH (mm)

Fig. 9. Plot of ear length (mm) against forearm length (mm) for N. gould: (circles) and N. bifax (triangles) from
northern NSW and far southeastern Old. Closed symbols represent males, open symbols females.

Skull dimensions overlap or abut in all but four characters: condylobasal length,
greatest length of skull, interbullae distance and bulla length (Table 2). Clear separation
of the two species results on a plot of zygomatic breadth against greatest length of skull
(Fig. 10). Thus in terms of absolute size specimens of N. gould: from northern New South
Wales have longer skulls with bigger bullae. It remains to be seen whether these differ-
ences are maintained in larger samples.

Specimens of N. gould: from this restricted region are more variable than WN. bifax for
the majority of external and cranial dimensions. This is reflected in sample coefficients
of variation (see Tables 1 and 2), and is evident in a bivariate plot of zygomatic breadth
against greatest length of skull (Fig. 10). This does not appear to be a site effect as

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

H. E. PARNABY 167

individuals from the same site have measurements at either end of the scatter for these
characters. The significance of this, as well as the greater variation in the majority of
other dimensions of N. gould: is not evident at present. Until such variation is more
clearly understood, it seems prudent not to test each species for statistically significant
differences between character means.

(mm)

BREADTH

ZYGOMATIC

16.0 17.0 18.0

GREATEST LENGTH (mm)

Fig. 10. Plot of zygomatic breadth (mm) against greatest length of skull (mm) for 19 N. gould: (circles) and 12
N. bifax (triangles) from northern NSW and southeastern Qld. Males represented by closed symbols, females
by open symbols.

DISCUSSION

There is little doubt that in eastern Australia, N. gould: and N. bifax are distinct
taxa. In the original description of N. bifax, Thomas distinguished it from N. gould: by its
relatively shorter ears, smaller bullae, more reduced post-nasal bump, and by the dis-
tally bifurcate baculum (compared to the simple point of N. gouldz). This study supports
the validity of all of these criteria.

Based on specimens pooled from different localities, ranges for each species overlap
for the majority of external and skull dimensions. Pooling localities, however, could ob-
scure size differences that might exist between species at a single locality. Specimens
pooled from a restricted geographic area (northern New South Wales and southeastern
Queensland) were collected from a particularly diverse range of environments, but
either sex of the two species could be separated using some dimensions. Adequate sam-
ples of both species from a single locality are obviously required to analyse this situation
further but are not currently available.

Skulls of either sex of both species are separable on the basis of interbullae distance
irrespective of locality, this character being a reflection of bullae size, a further distin-
guishing feature. This is significant given the extent of within-species variation and that
when specimens are pooled from all localities ranges of all other dimensions examined

PROG. LINN. SOC. N.S.W., 109 (3), (1986) 1987

168 LONG-EARED BATS

overlap between species. This apparently reflects differences in the auditory capabilities
of each species though audition in either species 1s little known.

As well as being broadly sympatric over a large part of their respective range, N.
bifax and N. gould: have been captured together at the same site in a number of areas.
Both species were trapped at the same site in the same evening at a number of localities
in northern New South Wales (Parnaby, 1986). N. bzfax was captured frequently at near
coastal localities up to 300m in elevation. By contrast, N. gould: was captured in much
lower numbers but at sites ranging from near sea level (such as at Broken Head Nature
Reserve) to 1120m at Bar Mountain in the Tweed Range. Churchill (fers. comm.) cap-
tured both species at the same site in the same evening at Mt Leysham near Charters
Towers, north Queensland.

I know of only three specimens of N. gould: from the Atherton region, which at
present represent the most northern record for this species. Cairn and Grant collected
two females (AM M/7024-25) from the Herberton area in 1889 while the third specimen
(QM JM5381, also female) was captured in a harp trap set near a stream in dry
sclerophyll forest at Atherton. Richards (1984) did not detect this species during an ex-
tensive bat survey of north Queensland rainforests, which included the Atherton region,
while he found N. bifax to be abundant. A variety of techniques were used but he relied
principally on identification of free flying bats by electronic monitoring of echolocation
calls. Although more effort was concentrated on rainforest, sclerophyll forest and wood-
land were also sampled (Richards, pers. comm.). Thus N. gould: appears to be uncommon
in the Atherton region where it is possibly restricted to schlerophyll forest.

Electrophoretic evidence based on samples of each species taken in sympatry at
several localities in northern New South Wales indicates clearly that they are good bio-
logical species. In addition, electrophoretic profiles between each species are such that
hybrids between them would be readily detected (M. Adams and P. Baverstock, fers.
comm.). However, only a small number of individuals of each species have been
examined electrophoretically from areas of known sympatry.

The affinities of a number of populations variously referred to N. gould, N. bifax
and N. timoriensis require clarification. Churchill et al. (1984) describe specimens of a
Nyctophilus occurring sympatrically with N. geoffroy: and N. bifax from Mt Leysham,
south of Charters Towers and while noting the similarity with N. gouldi, they were unable
to allocate these animals to any known form of Nyctophilus.

I have examined these specimens (Queensland Museum numbers JM5248 and
JM5358, not JM4361 and JM4362 as stated by Churchill et al. (1984), which are com-
parable with material from other areas of central Queensland. In view of the consider-
able size variation amongst N. gould: from different localities, which appears to reflect
local environmental conditions, there seems to be no reason for distinguishing these
animals from N. gouldi.

A further question concerns the nature of the relationship between N. bifax bifax
and N. bifax daedalus, and their distribution in north Queensland. These taxa are mor-
phologically distinct and both could occur in the Gulf region of northwest Queensland,
although sympatry has not been demonstrated. Churchill et al. (1984) record N. bifax
from the Lawn Hill area. Although they did not distinguish N. bifax bifax from N. bifax
daedalus, a specimen from Lawn Hill lodged by them in the Queensland Museum
(JM5246) is clearly referable to N. 6. daedalus. Of the material I have examined, the most
western record of N. bifax bifax in Queensland is a specimen (AM M13356) from Einas-
leigh River (18° 11°S, 144° 00’E), west of Mt. Surprise.

McKenzie ¢t al. (1977) identified a specimen of N. bifax from the Drysdale River
National Park, northern Western Australia. They assigned this specimen (WAM
M14097) to N. 6. bifax rather than N. 6. daedalus, on the basis of its bifurcate baculum. I

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

H. E. PARNABY 169

have examined this specimen and believe that it is distinct from N. 6. bzfax; in bacula and
dental features it is closest to N. 6. daedalus. Thus with the possible exception of a speci-
men in the British Museum from Cloncurry (Thomas, 1915) which I have not ex-
amined, I am not aware of any definite record of N. 6. bifax west of that collected from the
Einasleigh River.

The status of N. bifax daedalus remains confused. The arrangement of Johnson
(1964) is usually followed. He regarded it as the western race of N. bifax though its
affinities have also been placed with N. gould: by Troughton (1941) (as N. tamortensis gould:)
and Koopman (1984). Although the affinities of daedalus remain unclear, it does not
appear to belong with N. bzfax.

Large Nyctophilus from Tasmania have variously been referred to both N. gould: and
N. timoriensis. Hall and Richards (1979) record N. gould: from Tasmania for the first time.
The basis of this record (Richards, fers. comm.) is a specimen from Flinders Island,
registered in the Queen Victoria Museum (reg. number 1978.1.351). In skull and bacu-
lum shape, it resembles a large version of mainland WN. geoffroy:, and is unlike Victorian
N. gouldi. Thus, I am not aware of any valid record in the literature of N. gould: from
Tasmania; the relationships of Tasmanian Nyctophilus require clarification.

ACKNOWLEDGEMENTS

This study was enhanced by the collecting activities of the following people: Mark
Adams, Sue Churchill, Stan Flavel, Terry Reardon and Dr Chris Watts via an ABRS
grant to Dr Chris Watts; Dr Les Hall, Dave Milledge, Greg Richards and Ray
Williams.

Loan of museum collections was made possible by Dr John Calaby, Australian
National Wildlife Collection (CSIRO), who read an early version of the manuscript;
Joan Dixon and Linda Huxley, Museum of Victoria; Bob Green, Queen Victoria
Museum, Launceston; Dr Cath Kemper, South Australian Museum; Dr Darrell
Kitchener, Western Australian Museum; and Steve van Dyck, Queensland Museum.
Their patience during this part time study is appreciated. Linda Gibson and Dr Tim
Flannery, Mammal Department, Australian Museum, allowed free access to the collec-
tions and kindly provided space and facilities. Thanks to Kate Lowe, Photographic
Department, Australian Museum, who X-rayed numerous bat genitals.

Partial costs of a collecting trip to northern Australia during 1983 were met by an
Australian Museum Post-graduate Research Grant. The assistance and company of
Cathy Hill during the ten weeks of that trip is much appreciated. Anne and Greg
Richards of Atherton and Maree Fowlers and Alex Gilmore of Federal, N.S.W., were
generous hosts during several collecting trips.

The unflagging financial support from Shirl and Dave Parnaby enabled the
continuation of this work.

Mark Adams and Dr Peter Baverstock, South Australian Museum, made their
electrophoretic data freely available and allowed me to quote from their unpublished
inquiries into Nyctophilus.

Valuable assistance was given by staff at the Faculty of Biological Sciences, Univer-
sity of New South Wales; Ross Arnett and Lisa Jensen, Photographic Unit, who set up
equipment and gave advice; Ken Aplin, School of Zoology, read several versions of the
manuscript and provided many helpful suggestions; and I have also benefited from dis-
cussion of the problem and this manuscript with Dr Michael Archer, School of Zoology,
University of New South Wales, the supervisor of my postgraduate work.

PROC. LINN. SOC. N.S.W., 109 (3), (4986) 1987

170 LONG-EARED BATS

References

ALLISON, R. F., 1982. — Nyctophilus bifax; p. 194. In: HONACKI, J. H., et al., (eds), Mammal species of the world: A
taxonomic and geographic reference. Lawrence: Allen Press and Association of Systematic collections.

——,, 1983. — Nyctophilus bifax; p. 335 in STRAHAN, R., (ed.), The Australian Museum complete book of Australian
mammals. Sydney: Angus & Robertson.

CHURCHILL, S. K., HALL, L. S. and HELMAN, P. M., 1984. — Observations on long-eared bats (Vespertili-
onidae: Nyctophilus) from northern Australia. Aust. Mamm. 7: 17-28.

HALL, L. S., 1984. — And then there were bats. Jn ARCHER, M., and CLayTON, G., (eds), Vertebrate zoogeogra-

phy and evolution in Australasia: 837-852. Carlisle, W.A.: Hesperian Press.

, and RICHARDS, G. C., 1979. — Bats of eastern Australia. Queensland Museum Booklet No. 12. 66 pp.

IREDALE, T., and TROUGHTON, E. L., 1934. — A checklist of the mammals recorded from Australia. Mem.
Aust. Mus. 6: 1-122.

JOHNSON, D.H., 1964. — Mammals of the Arnhem Land Expedition. Jn: SPECHT, R. L., (ed.), Records of the
American-Australian scientific expedition to Arnhem Land, vol. 4, Zoology: 427-51.

Koopman, K. F., 1984. — Taxonomic and distributional notes on tropical Australian bats. Amer. Mus. Novit.
2778: 1-48.

MCKENZIE, N. L., CHAPMAN, A., YOUNGSON, W. K., and BURBIDGE, A. A., 1977. — The mammals of the
Drysdale River National Park, North Kimberley, Western Australia. Jn: Kapay, E. D., and
BURBIDGE, A. A., (eds), A biological survey of the Drysdale River National Park, North Kimberley,
Western Australia in August, 1975. Wildl. Res. Bull. West. Aust. No. 6: 79-86.

PARNABY, H. E., 1986. — A survey of bats of the rainforest of northeast New South Wales. Unpubl. report to
National Parks & Wildlife Service, N.S.W. 45 pp.

RICHARDS, G. C., 1983. — Nyctophilus gouldi; p. 330. In: STRAHAN, R.., (ed.), The Australian Museum complete
book of Australian mammals. Sydney: Angus & Robertson.

——,, 1984. — The conservation status of the rainforest bat fauna of North Queensland. Jn: WaRREN, G. L.,
and KERSHAW, A. P., (eds), Australian National Rainforests Study Report. Vol 1. Proceedings of workshop
on past, present and future of Australian rainforests at Griffith University, Dec. 1983: 292-300.
Clayton, Victoria: Geography Dept., Monash University.

Tate, G. H. H., 1952. — Results of the Archbold Expeditions. No. 66. Mammals of Cape York Peninsula
with notes on the occurrence of rainforest in Queensland. Bull. Amer. Mus. Nat. Hist. 98: 563-616.

THOma~S, O., 1915. — Notes on the genus Nyctophilus. Ann. Mag. Nat. Hist. ser. 8, 15: 493-499.

TOMES, R. F., 1858. — A monograph of the genus Nyctophilus. Proc. Zool. Soc. Lond. (1858): 25-37.

TROUGHTON, E. leG., 1941. — Furred animals of Australia. Sydney: Angus & Robertson.

PROC. LINN. SOC. N.S.W,, 109 (3), (1986) 1987

H. E. PARNABY 171

APPENDIX 1
SPECIMENS EXAMINED

All the following specimens have been examined by the author. A subset of these
has been measured and details are available from the author.

Nyctophilus gouldi

Queensland. Atherton, 17° 16’S, 145° 29’ E: If (JM5381); Herberton, ca 17° 23’S, ca 145°
23’E: 2f (AM M7024-25); Mt Leysham, 20° 15’S, 146° 15’E: 2m,lf (QM JM5248,
JM5358, AM M12967); Finch Hatton, 21° 09’S, 148° 38’E: lf (QM JM1113); Capella,
23° 05'S, 148° 01’E: 2m (AM M6312-13); Retro Downs Stn, Capella, 22° 52’S, 147°
54’E: 8m,lf (QM JM6184-85, JM6218-21, JM6301-03); Craigmore, 23° 55’S, 147°
- 53°E: lf (QM JM2946); Blackdown Tableland, Mimosa Ck, 23° 47’S, 149° 05’E: If
(QM JM5359); Pluto Timber Reserve, via Tambo, ca 25° 00’S, ca 147° 05’E: I(un-
sexed) (CM15593); Mt Pluto, 3km E, 25° 00'S, 147° 05’ E: Im (CM4393); Mt Moffatt,
225km N of Mitchell, 25° 01’S, 147° 57’E, Im (QM J20378); Mitchell, 26° 29’S, 147°
58’E: Im (CM2334); Babbiloora Stn, 25° 12’S, 147° 57’E: 4m (JM5360-5363);
Mioolangool Stn; via (Gin) Ginyca 24°" 45)S, (ca lol? 32 EK: 2f (AM: M5972;
M6096Gmm.) ); Hervey Bay, ca 25° 06'S, ca 152° 49’E: 2f (QM JM2577, JM2596);
Chinchilla, ca 26° 44’S, ca 150° 36’ E: Im, lf (AM M7634, QM J1763); Chinchilla, 32km
SW, ca 26° 55'S, ca 150° 25’E: Im (QM JM506); Conondale Rge, ca 26° 39'S, ca 152°
38’E: lf (AM JM5364); Pomona, 26° 22’S, 152° 52’ E: lm (QM JM991); Millmerran,
27° 93'S, 151° 16’ E: Im,lf (AM M3911-12); Caloundra, 26° 48’S, 153° 08’ E: lf (QM
J1762); Moggil Forest, 2f (QM JM5365-5366); Wallangara, 28° 55’S, 151° 56’E: lm
(AM Mi1841); Levers Plateau, NW edge, 28° 19’S, 152° 51’E: Im,lf (AM M13411-12);
North Tamborine, 27° 56’S, 153° 11’E: lm (QM J7095); Brisbane, 27° 28'S, 153°
01’E: lm (QM J10872).

New South Wales. Tweed River, Tyalgum 28° 22’S, 153° 12’E, lf (AM M5450); Tweed
Range, Paddys Mountain Tk, 28° 25’S, 153° 07’ E: 2m (AM M13171, M13392); Tweed
Range, Bar Mountain picnic area, 28° 27’S, 153° 19’E: lm (AM M13403); Tweed
Range, Airdrop Rd, 28° 24’S, 153° 03’E: 2m (AM M13393, M13406); McPherson
Rge, Palm Gully area, 28° 22’S, 152° 55’ E: lm,lf (AM M13178, M13181); Tooloom, 28°
37S, 152° 25’ E: lm,lf (CM573-574); Big Scrub Flora Reserve, 28° 38'S, 153° 20’E:
Im (AM M13243); Terania Ck, Nightcap National Park, 28° 34’S, 153° 17’E: lf (AM
M13235); Whiam Whiam State Forest, 28° 35’S, 153° 22’ E: Im,2f (AM M16029-16030,
M14184); Broken Head Nature Reserve, 28° 33’S, 153° 37’E: Im lf (AM M13380,
M13389); Woolgoolga Nature Reserve, 30° 07'S, 153° 09’E: Im,2f (AM M13227-28,
M13231); Dorrigo National Park, 30° 22’S, 152° 48’E: 2m (AM M13197-98); New
England National Park, 30° 30’S, ca 152° 27’E: 2m,2f (AM M13200, M13202,
M13208, M13213); Moree, 29° 28'S, 149° 51’E: lf (AM M5956); Pilliga Scrub, ca 31°
00'S, ca 149° 30’E: Im, lf (AM M5968, M8471); Pilliga Scrub, 30° 32’S, 149° 32’E: 3f
(AM M16023-16025); Spear Ck, 7km E of Apsley Rv., 30° 55’S, 152° 05’E: lf (AM
M14189); Mt Boss State Forest, 31° 11’S, 152° 27’E: 4m,2f (AM M14108-M14113);
Mooraduck Ck, Werrikimbie National Park, 31° 09’S, 152° 13’E: Im (AM M12548);
Doyles River State Forest, 31° 27’S, 152° 10’ E: Im,4f (AM M14114-M14118); Sea Acres
Nature Reserve, Port Macquarie, 31° 28’S, 152° 56’E: If (AM M13434); Gloucester
Tops, 3.8km WNW, 32° 02’S, 151° 34’E: Im,3f (AM M13425-26, M13429, M13432);
Upper Allyn via Eccleston, 32° 10’S, 151° 29’E: Im (AM M9394); Canningalla Stn,

PROC. LINN. SOC. N.S.W,, 109 (3), (1986) 1987

172 LONG-EARED BATS

7 ml NW of Dungog, 32° 24’S, 151° 45’ E: If (AM M10002); Weabonga, 31° 13’S, 151°
19’E: lf (AM M7631); Wheogo Stn, 20km N of Dunedoo, ca 31° 50’S, ca 149° 24’E:
Im, lf (AM M4284-4285); Turee, 31° 56’S, 149° 48’E: lm (AM M3746); Munghorn
Gap, NE of Mudgee, 32° 25’S, 149° 50’E: 2m,2f (AM M14104-M14107); Bulahdelah,
Skim SH 32°°257S;, 1522 12/7 Es mi (AM! Mil0312); Olney, State Forest, 33° 0o¢ss tole
18’E: 16m,10f (AM M15980-15989, M16007-16022); Anna Bay, S of Port Stephens, 32°
4609; 152° 040 Be 2th (AIM M4273") M4321) Wyong Gk) 332, 10 Sia oleasiOMaEraslis
(CM1575); Ourimbah, 33° 22'S, 151° 22’ E: lm (AM M4224); Gosford, 33° 26’S, 151°
20’E: 4f (AM M4433-4436); Ash Island, Hunter River, 32° 51’S, 151° 43’E: lm (AM
M2566); Dangar Island, 33° 32’S, 151° 14” E: lm (AM M9265); Sydney area: Botany,
Im (AM M2564); Carlingford, lm (AM M7618); Eastwood, lf (AM M8014); North
Wahroonga, 33° 42’S, 151° 07’E: Im (AM M9191); Gymea Bay, Im (AM M7482).
Hazelbrook, 33° 44’S, 150° 27’E: Im,2f (AM M3041, M3545, M3739); Lawson, 33°
4305, 1502) 26) Be lh (AM M1437); Abercrombie, Arch) Gave332) 90i09,9149 22am
(CM2323); Jenolan Caves, 33° 49'S, 150° 02’E: If (AM M1702); 3km S of Coco Ck
Cave, Capertree Valley, ca 33° 08’S, 150° 10’E: lm (AM M9846); Junction of Caper-
tree Rv. — Wolgan Rv., 33° 12’S, 150° 28’E: Im,lf (AM M11481-11482); Cob area,
Culoul Rge, ca 33° 13'S, ca 150° 36’E: lm (AM M11632); Grassy Hill Tk., W of Putty
Rd., ca 33° 22'S, ca 150°41’E: lm (AM M11090); Millamalong, nr. Mandurama,
33°15’S, ca 150° 40’ E: lf (AM M3414); Pheasants Nest nr. Picton, 34° 15’S, 150° 40’ E:
Im (AM M11557); Campbelltown, 9km S, 34° 08’S, 150° 47’E: lm (AM M14088);
Berrima, 34° 29'S, 150° 20’ E: Im (AM M3436); Robertson, 34° 35’S, 150° 35° E: lf
(AM M6272); Carrington Falls, 34° 38'S, 150° 41’E: lf (GM4588); Mt Keira, 34°
24S; 1002 (ol Es 2m) (AMG M9140) (GNIZ987)" Kemraville; 342025175; slo Otero 0mieaelin
(CM4589); Bungonia Caves, 34° 52’S, 149° 57’E: If (AM M7638); Narooma, 36°
13’S, 150° 09’E: lf (GM1745); Jervis Bay Nature Reserve, 35° 09’S, 150° 43’ E: lm,2f
(AM M13438-13440); Sussex Inlet, 35° 12’S, 150° 33’E: lm (AM M14188); Araluen,
39° 39'S, 149° 49 E: lf (M612); Mogo, 35° 477S, 150° 09” E; Im (M1895); 1-4km i
of Wollybut Tk.. Mumbulla State Forest, 36° 33'S, 149° 52°E: Im (AM M12752);
Argalong, 35° 18'S, 148° 24’ E: lm (AM M4534); Tumut State Forest, 35° 22S, 148°
12’E: Im (AM M11300); Yarrangobilly Caves, 35° 39’S, 148° 28’E: lm (CM6295);
Murrumbateman, 34° 58’S, 149° 02’E: lf (CM4705); Sutton, 35° 10’S, 149° 15°E: lm
(GM1457); Mt Tindery, 35° 42’S, 149° 16’ E: 1 (@M2062); Temora, 9km W, 34° 3957S;
1472 25 ES lt (AM) Mill716); demora; lokmy SWi34°"3iy S) 1472. 221s lanl NE
M13576-77); The Rock Nature Reserve, 35° 16’S, 147° 04’ E: lm (AM M11522); Wagga
Wagga, Livingstone State Forest, 35° 25’S, 147° 25’E: 2m,2f (AM M11217, M11521,
M11655, M11717); Gerogery, 35° 50’S, 147° 00’E: lm (CM4029); Wahgunya State
Forest, 35° 51°S, 145° 59” E: Im (AM M1494); Deniliquin, 35° 327S, 144° 57 “Ea lim
(MV C5159).

Australian Capital Territory. Captains Flat, 35° 35’S, 149° 27°E: lm (CM4747);
Bushrangers Ck, Brindabella Rge, lm (CM4361); Canberra, 13km N, 35° 17’S, 149°
13" E: 2m (CM2382, GM2387); Lake’ Burley ‘Griffith, 35° 1778; 1497913 Es slit
(CM2345, CM2386, CM2434); Black Mountain, 35° 16’S, 149° 06’E: Im (CM2077);
Yarralumla, lm (CM591).

Victoria. Junction Little Bog Ck-NSW border, 37° 19'S, 149° 05’E: lm (MV C26595);
Boundary Road, Wingham River, 37° 42’S, 149° 28’E: lf (C24907); Mitta Mitta, 36°
32’S, 147° 22’ E: lm (CM4359); Mt Buffalo National Park, 36° 45'S, 145° 48’E; lm,3f
(MV C26957-26960); Myrtleford, 24km S, 36° 45'S, 146° 44’E: If (MV C11467);
Bogong 36° 48’S, 147° 13’E: Im,lf (MV C11565-11566); Dargo, 30km NNW, 37° 11’S,

PROC. LINN. SOC. N.SW., 109 (3), (1986) 1987

H. E. PARNABY We

147° 12’ E: 2m,lf (MV C26961-26963); Nathalia, Goulburn River, 36° 10’S, 145° 06’E:
Im (MV C25643); Mt Warby, 1.6km NNW, 36° 20’S, 146° 12’E: 2m (MV C25671-
25672); Balook, 38° 26’S, 146° 34’ E: If (MV C26952); Mt Killawarra, 36° 09’S, 146°
14’E: Im (MV C25657); Edi, 4km WNW, 36° 38'S, 146° 24’E: Im,3f (MV C25186-
25187, C25194); Mt Macedon township, 37° 25’S, 144° 34’E: lm (MV C7452); Dayles-
ford area, 37° 21'S, 144° 09’E: 4f (MV C16020-16021, C16152, C26951); Maroondah
Reservoir, 37° 38'S, 145° 35’E: 2m (MV C€26968); Croydon, 37° 48'S, 145° 17’E:
Im,lf (MV C5419, C25476); Jindivick, 3km NNW, 38° 00’S, 145° 52’ E: (MV C24907);
Refuge Cove, Wilsons Promontory, 39° 03’S, 146° 28’E: lm (MV C17129); Mt Erica,
ilk SSW, 37° 537S, 146° 21°E: 8m,lf (MV €25350, €25355, €25360, C25366-
25367, C25372, C25374, C25386); Mt Baw Baw, 5.8km SSW, 37° 50’S, 146° 17’E: If
(MV C25336); Willow Grove 38° 05’S, 146° 11’E: lm (MV €C25324); Nowa Nowa area,
37° 16°S, 147° 58’E: Im,2f (MV €26093-26094, C26096); Bruthen, 8km NW, 37°
39'S, 147° 46’ E: lm, lf (MV C26965-26966); Lockup Ck, Waratah Spur Tk. 37° 24'S,
149° 06’E: lf (MV C25905); Yalmany Rd, Roger Rv., 37° 18’S, 148° 35’ E: 5m,lf (MV
C25934-25939); Cooagalah block, 37° 24’S, 149° 21°E: 2m,2f (MV C25969-25972);
Mt Noorinbee, 6km NNW, 37° 27'S, 149° 05’E: lf (MV C26964); Cann River, 37°
S449) 104 lm (NIV €22278). Bemm) River, 37°49" S, 1482998 E: Im (MV
C1434); Mt Ellery, 1.7km W, 37° 24'S, 148° 47’E: lm,lf (MV €26409-26410);
Teddington Reservoir, 1.5km S, 36° 52’S, 143° 16’E: 5m,6f (AM M16750, MV
C26968-26978); Zumstein, 37° 05'S, 142° 22’ E: lm (MV €C26967); Mt Rosea, Gram-
pians State Forest, 37° 12’S, 142° 30’E: lf( MV €C25691); Mt Eccles National Park, 38°
Oa Sa0142 700i: lm: (MV. C2605): Napier Forest, 37° (94S; 1429 047 BE: Imi (MV
C23676); Otway Ranges, 38° 41'S, 143° 36’ E: 3m,]f (MV C26953-26956); Grey River
Scenic Reserve, nr. Kennett River, 38° 40’S, 143° 51’E: lf (MV C25343); Irrewillipe,
Sem daswl4o 25) Elm OVE .C17906):

Nyctophilus bifax

Queensland. Moa Island, 10° 11’S, 142° 16’E: lf (MV C8800); Prince of Wales Is., ca 10°
41’S, ca 142° 09’ E: 2m, lf (GM11632-34); Lockerbie Scrub, 10° 48’S, 142° 27’ E: 4m,2f
(GM11626-31); Carnegie Rge, Cape York, 10° 46’S, 142° 31’E: lf (QM JM5367);
Captain Billy Ck. 11° 37’, 142° 50’E: 2m,3f (QM JM5368-5372); Iron Range, 12°
3S 2 Ou i 2imelf (@MUIIM53 73-0379) [ron Range, 122 40S) 1439 12 E42 th
(AM M16188-16193); Archer River, 13° 27’S, 142° 57’E: lm (QM JM5376); Archer
River, 5km S, 13° 29’S, 142° 58’ E: 2m,2f (AM M12958-12959, M16032-16033); Peach
Ck, Mcllwraith Rge, 13° 40’S, 143° 07’ E: 2m, lf (QM JM5377-5379); Buthen Buthen,
Cape York, 13° 21'S, 143° 28’ E: lm,2f (QM JM2429-30, JM2475); Rocky River, ca 13°
48'S, ca 143° 27’ E: lm (CM4486); Station Ck, 16km S of Coen, 14° 03’S, 143° 16’E: 2f
(AM M13352, M13369); Cooktown, Jones Lagoon, 15° 26’S, 145° 10’E: lm,If (AM
M12956-12957); Cooktown, Walker Bay, 15° 31'S, 145° 17’ E: 1f(AM M12955); Bloom-
field, 15° 56’S, 145° 21’E: lm (AM Mi11278 imm.) Cape Tribulation, 16° 10’S, 145°
25’E: 5m,2f (AM M13344-50); Kewarra Beach, 18km N of Cairns, 16° 37’S, 145°
41’E: Im (AM M17301); Clump Point, Tully, 17° 52’S, 146° 07’E: If (AM M8386);
Kuranda, 16° 497S, 145° 38 E: 2m (CM15043, OM J4409); Atherton, 17° 16’, 145°
29°E: 6m,2f (QM JM5380, JM5382, AM M13602-13605, M13611-13612); Severin
Creek State Forest, Atherton Tableland, 17° 11’S, 145° 40’E: 3m,4f (AM M16183,
M16185-16187, QM JM5401-5403); Wongabel State Forest, Atherton Tableland, 17°
19’S, 145° 29’E: lm,3f(AM M16181-16182, M16184, OM JM5404); Herberton district,
ca 17° 23'S, ca 145° 23’ E: lm,2f(AM M557-58, M560); Chillagoe, 17° 09'S, 144° 31’E:
Im (CM5891); Einasleigh River, Kennedy Highway, 18° 11’S, 144° 00’E: Im (AM

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

174 LONG-EARED BATS

M13356); Ravenshoe, ca 17° 36'S, ca 145° 29’E: lm (AM M8011); Innisfail, 17° 32’S,
146° 01’E: Im, lf (CM4834, AM M8005); Cardwell, Kirrama State Forest, ca 18° 10’S,
ca 145° 40’ E: lf (QM JM5383); Mt Spec, 64km N of Townsville, 18° 56'S, 146° 11’E: If
(CM4835); Palm Is., 18° 40’S, 146° 33’E: lf (QM J5274); Fanning River Stn, 19° 16’S,
146° 49’ E: lm (QM JM5262); Charters Towers, 18km E, 20° 06'S, 146° 26’ E: 2m (AM
M13353-54); Charters Towers, Mt Leysham, 20° 15’S, 146° 15’E: lm (QM JM5384);
Eungella, 21° 08’S, 148° 29’E: lm (AM M11175); East Funnell Ck, nr. Sarina, ca 21°
33'S, ca 149° 09’E: 2f (AM M/7038-39); Minga Mountain, Byfield, 22° 52’S, 150°
32’E: lm (QM JM5385); Rockhampton, 23° 22’S, 150° 32’ E: (QM JM5386); Hervey
Bay, ca 25° 06’S, ca 152° 49’E: 2f (QM JM2571, JM2574); Noosa Heads, 26° 23’S,
153° 07’ E: l(?sex) (MV M15368); Conondale Rge, ca 26° 39’S, ca 152° 38’ E: lm (QM
JM5387); Mt Nebo, 27° 24’S, 152° 47’ E: lm (QM JM5388).

New South Wales. Reserve Ck, Murwillumbah, ca 29° 39'S, ca 150° 43’ E: lm (CM4743);
Billinudgel Swamp, 28° 29'S, 153° 32’ E: lm,3f(AM M14090-14093); Brunswick Heads
Nature Reserve, 28° 33’S, 153° 33’E: lm (AM M13388); Broken Head Nature Reserve,
28° 33'S, 153° 37’ E: 5m,2f (AM M13378, M13381-83, M13385-87); Broken Head area,
28° 44’S, 153° 36’E: lf (AM M15330); Whiam Whiam State Forest, 28° 35’S, 153°
22’E: lf (AM M16031); Nightcap National Park, Terania Ck, 28° 34’S, 153° 17’E: lf
(AM M13234); Big Scrub Flora Reserve, 28° 38’S, 153° 20’E: 2m,2f (AM M13240,
M13247-M13249); Iluka Nature Reserve, 29° 24’S, 153° 22’E: 2m,]f (AM M13191-
M13192, M13232).

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

Late Pleistocene Dinoflagellate Cysts from
Bulahdelah, northern New South Wales

ANDREW MCMINN

(Communicated by H. MARTIN)

McMinn, A. Late Pleistocene dinoflagellate cysts from Bulahdelah, northern New
South Wales. Proc. Linn. Soc. N.S.W. (109(3), (1986) 1987: 175-181.

A Late Pleistocene dinoflagellate microflora has been recovered from the water
bore WRC 39275 near Bulahdelah, N.S.W. Species present include Protoperidinium (Pro-
toperidinium sect. Quinquecuspis) leonis (Pavillard), Tuberculodinium vancampoae (Rossignol),
Polysphaeridium zoharyi (Rossignol), Spiniferites mirabilis (Rossignol), Spiniferites ramosus
(Ehrenberg) and Spiniferites sp. cf. ramuliferus (Deflandre). This assemblage suggests a
warm, nearshore, marine depositional environment.

A. McMinn, Geological Survey of N.S.W., Building Blla, University of N.S.W., PO. Box 1,
Kensington, Australia 2033; manuscript received 20 August 1986, accepted for publication 17
December 1986.

INTRODUCTION

A Late Pleistocene sequence was intersected in the water bore WRC 39275, which
is located approximately 10km southeast of Bulahdelah, N.S.W. (Fig. 1). This sequence
consists of 42m of interbedded gravels, sands and clays which can be divided into three
depositional units (Fig. 2). Below 18.8m the sequence consists of fluvial lithic sands,
clays and gravels, between 18.8m and 11.5m it consists of estuarine/shallow marine clay
culminating in fine- to medium-grained quartz sand, and above 11.5m it is composed
solely of fine- to medium-grained quartz sand of the Pleistocene Inner Barrier system
(Pickett, 1983; Drury, 1982). By an analogy with nearby deposits of known age Pickett
(1983) assumed the sequence to be associated with the Last Interglacial and therefore to
be approximately 120,000 years old.

Four samples were investigated: from 9-l1lm, 14.2-18.8m, 28.0-38.8m and 40.5-
40.8m. The lower three samples yielded microfloras but only one, from 14.2-18.8m,
yielded a dinoflagellate cyst assemblage. This latter interval also contained foramini-
feral and molluscan assemblages (Pickett, 1983). The spore-pollen component of the
palynomorph assemblages is dominated by Casuarina (11.0-25.4%), Myrtaceae (37.5-
43.0%) and Cyathea (10.0-29.0%); dinoflagellates comprise less than 0.5% of the upper-
most assemblage and were not recovered from the underlying samples.

The samples were prepared according to standard palynological procedures,
although they were not oxidized as even mild oxidation has been observed to destroy
some cyst types (Dale, 1976). Palynological preparations are located in the palynological
collection of the Geological Survey of N.S.W.

DINOFLAGELLATES

Fossil dinoflagellate cysts have been used extensively both to determine the age and
to provide information on the depositional environments of Mesozoic and ‘Tertiary
marine sequences. Until relatively recently, however, the study of Quaternary dino-
flagellate cysts has been neglected and even now is virtually restricted to the northern
hemisphere. Pleistocene dinoflagellate cyst assemblages have previously been described
from Great Britain (Harland and Downie, 1969; Harland, 1977; Wall and Dale, 1968;
West, 1961), the North Atlantic Ocean (Harland, 1979; 1984a; 1984b), the Caribbean

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

176 LATE PLEISTOCENE DINOFLAGELLATE CYSTS

f=] Quaternary Sediment
@ wre 39275

“STEPHEN

Fig. 1. Location of the bore WRC 39275 (Map reference: Newcastle 1:250 000 sheet, G.R. 5279 9865).

Sea (Wall, 1967), the North Sea (Harland et al., 1978), the Black Sea (Wall et al., 1973;
Wall and Dale, 1973; 1974), Japan (Matsouka, 1976a; 1976b), Matsouka and Nishida,
1973), New Zealand (Wilson, 1973) and from Israel (Rossignol, 1962; 1964). There is no
published account of Australian Pleistocene marine dinoflagellate cysts.

Six species, however, were recorded from Bulahdelah; these are:

Protoperidinium (Protoperidinium sect. Quinquecuspis) leonis (Pavillard) Balech 1974; Figs
3A-F.

Tuberculodinium vancampoae (Rossignol) Wall 1967; Figs 3J-K,M-N.

Spiniferites mirabilis (Rossignol) Sarjeant 1970; not illustrated.

Spiniferites ramosus (Ehrenberg) Mantell 1854; Figs 3H,I,L.

Spiniferites sp. cf. ramuliferus (Deflandre) Reid 1974; not illustrated.

Polysphaeridium zoharyt (Rossignol) Bujak et al. 1980; Fig. 3G.

The relative abundance of each species in the dinoflagellate assemblage is shown 1n
Fig. 2. Absolute dinoflagellate abundance is low, being less than two cysts per gram of
sediment. This compares with abundances of up to 14,000 cysts per gram in Pleistocene
assemblages from Great Britain (Harland and Downie, 1969). The total number of
dinoflagellate cysts observed was 54.

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

ANDREW McMINN 177

Depth Lithology ere RELATIVE DINOFLAGELLATE

a) ABUNDANCE
fice P. leonis

10 EI Spiniferites spp.

T. vancampoae

Fig. 2. Stratigraphic column, sample positions and dinoflagellate cyst abundance in the bore WRC 39275.

RECENT DISTRIBUTION OF RECORDED DINOFLAGELLATE CYSTS

Cysts of Protoperidinium (Protoperidinium sect. Quinquecuspis) leonis have been recorded
from coastal areas around Great Britain, from the Caribbean and from off the coast of
West Africa (Harland, 1983), and from Japan (Matsouka, 1976a). Harland (1983) sug-
gested that this species has a tropical to temperate distribution in inner and outer neritic
environments. The thecate form of this species has been recorded extensively around
Australia (Wood, 1954).

Cysts of Tuberculodinium vancampoae have been recorded from the southern coast of
eastern U.S.A., Bermuda, Bahamas, Puerto Rico, Peru and the Mediterranean Sea
(Wall e¢ al., 1977), from the Persian Gulf and Red Sea (Bradford, 1973; 1975; Wall and
Warren, 1969), Israel and the eastern Mediterranean Sea (Rossignol, 1962; 1964),
Japan (Matsouka, 1976b; 1981; Harada and Matsouka, 1974; Shimakura et al., 1971).
Wall et al. (1977) noted that the maximum concentration of this species was in estuarine
environments. Harland (1983) also observed that 7’ vancampoae was more common in
tropical to subtropical areas. The highest latitude at which this cyst has been recorded is
approximately 40 degrees north, in Japan. In the Australasian region the thecate form
of this species (Pyrophacus vancampoae (Rossignol), Wall and Dale, 1971) 1s common in the
Coral Sea and other northern seas but also extends south along the N.S.W. coastline at
least as far as Sydney Harbour (new data) and Port Hacking (C. Hallegraeff, C.S.I.R.O.
Marine Laboratory pers. comm.).

Polysphaeridium zohary: has been recorded from Bermuda, Puerto Rico, southern
coast of eastern U.S.A. and the Mediterranean Sea (Wall et a/l., 1977), Israel and the

PROC. LINN. SOC. N.S.W,, 109 (3), (1986) 1987

178 LATE PLEISTOCENE DINOFLAGELLATE CYSTS

eastern Mediterranean Sea (Rossignol, 1964; Rossignol and Pastouret, 1971), Persian
Gulf and Red Sea (Bradford, 1973; 1975; Wall and Warren, 1969), the Black Sea
(Roman, 1969) and Japan (Harada and Matsouka, 1974). Harland (1983) suggested
that most of the reported northern European occurrences are probably misidentifica-
tions and that Polysphaeridium zoharyi is apparently restricted to tropical and subtropical
areas. It is also apparently more abundant in estuarine and nearshore environments. In
the Australasian region the thecate form of this species (Pyrodinium bahamense Plate 1906)
has not been recorded south of New Guinea (C. Hallegraeff pers. comm.).

Cysts of Spiniferites mirabilis, Spiniferites ramosus and Spiniferites sp. cf. ramuliferus each
have a cosmopolitan distribution. The thecate form of Spzniferites mirabilis and Spiniferites
ramosus (1.e. Gonyaulax spinifera (Claparede and Lachmann), Diesling, 1866; Wall and
Dale, 1970) is also widely distributed in the Australian region (Wood, 1954); the thecate
form of Spiniferites sp. cf. ramuliferus is not known.

DISCUSSION

The composition of the Bulahdelah assemblage bears little resemblance to any pre-
viously described assemblage. Wall et al. (1977) described 168 dinoflagellate cyst assem-
blages from a wide variety of modern marine and estuarine environments but they
recorded a maximum abundance of Tuberculodinium vancampoae, the dominant species at
Bulahdelah (66% of the assemblage), of only 11%. The unusual composition of the
Bulahdelah assemblage, therefore, creates difficulties in postulating possible depo-
sitional environments. The geographically closest described Pleistocene assemblages
are from the middle Pleistocene Te Piki bed, New Zealand (Wilson, 1973). These assem-
blages are all dominated by Bitectatodinium tepikiense Wilson 1973, a species usually
associated with temperate and cold temperate environments and not recorded from
Bulahdelah. The abundance of this species in modern environments (maximum 11%
(Wall et al., 1977) ) does not approach that reported in the Pleistocene of New Zealand
(43% to 100%). At this stage no explanation can be given for the dominance of these two
species which are usually only minor elements of an assemblage.

The absence of detailed Recent cyst distribution data for the Australian region pre-
vents a comparison of the Pleistocene Bulahdelah assemblage with modern cyst assem-
blages from known depositional and climatic environments; interpretations based on
cyst assemblages from the northern hemisphere will of necessity rely on extrapolations.
However, when those data are combined with data on the present distribution of thecate
dinoflagellates in the Australasian region (Wood, 1954) it can be inferred that the
Bulahdelah assemblage was deposited in a subtropical estuarine/shallow marine en-
vironment. This conclusion is consistent with results determined from foraminiferal
and molluscan faunas (Pickett, 1983).

ACKNOWLEDGEMENTS

I would like to thank the Secretary of the N.S.W. Department of Mineral Resources
for permission to publish this manuscript.

References

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Fig. 3. A,B Protoperidinium (Protoperidinium sect. Quinquecuspis) leonis MMMC 01771; C,F Protoperidinium
(Protoperidinium sect. Quinquecuspis) leonis MMMC 01772; D,E Protoperidinium (Protoperidinium sect. Quin-
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PROC. LINN. SOC. N.SW., 109 (3), (1986) 1987

ANDREW McMINN 179

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

180 LATE PLEISTOCENE DINOFLAGELLATE CYSTS

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PROC. LINN. SOC. N.S.W,, 109 (3), (1986) 1987

ice
Hs

nuaN a

Grabs}

ie!
iis

Wild ligt a a

The echinoderm Genus Henricia Gray, 1840
(Asteroidea: Echinasteridae) in southern and
southeastern Australian Waters, with the
Description of a new Species

FRANCIS W. E. ROWE and E. LYNNE ALBERTSON

Rowe, F. W. E., & ALBERTSON, E. L. The echinoderm genus Henricia Gray, 1840
(Asteroidea : Echinasteridae) in southern and southeastern Australian waters, with
the description of a new species. Proc. Linn. Soc. N.S.W. 109(3), (1986) 1987: 183-194.

Three species of Henmcia can be identified in southern and southeastern Australian
waters. These are: H. compacta (Sladen). H. obesa (Sladen), of which hyades: (Perrier) is a
junior synonym, and a new species, H. kapalae, from New South Wales. These species are
herein reviewed and described.

FW. E. Rowe and E. L. Albertson, Division of Invertebrate Zoology (Echinoderms), Australian
Museum, Sydney, Australia 2000; manuscript received 21 October 1986, accepted for publication 18
February 1987.

INTRODUCTION

Some 12 specimens of Henricia, collected during the ‘Endeavour’ Expeditions (1909-
1914) from between Gabo Island, Victoria and westward to the Great Australian Bight,
have been identified by H. L. Clark (1916) and echinoderm taxonomists in the Australian
Museum (unpublished) as H. Ayades: (Perrier). Clark (1916) commented on the‘. . . con-
siderable diversity in proportions and in the spinulation . . .’ of the material before him.
He concluded that he was unable to recognize more than a single species, H. hyadesi, pre-
viously recorded from South American waters. He noted the relatively deep water from
which material he had examined had been collected (91-365m) and suggested the South
American and Australian Henricia might be distinct. Although Clark reported 11 specimens
from the ‘Endeavour collections, only six of the 12 remaining in the Australian Museum
collections can be confirmed, by the senior author, as having been identified by Clark. The
remaining six specimens have been identified by other echinoderm taxonomists working
in the Museum, who compared them with those seen by Clark.

Fisher (1940), following the examination of three of H. L. Clark’s (1916) specimens,
declared that the specimen from south of Gabo Island, and similarly one from east of
Maria Island (Tasmania), appeared to represent H. sufflata (Sladen), a species described
from Kermadec Islands (north of New Zealand). The third specimen, from the Great Aus-
tralian Bight, appeared identifiable within a group of species including H. compacta
(Sladen), this species having been described from New Zealand waters. Fisher did not con-
sider any of these three specimens to be Perrier’s H. hyadest, a species he believed to be
conspecific with H. obesa (Sladen).

In 1946, H. L. Clark recorded Fisher’s re-identification of the three ‘Endeavour’ speci-
mens. However, on re-examining those specimens, and comparing them with other
material held in the Museum of Comparative Zoology at Harvard, notably two specimens
of hyades: from off Patagonia and a cotype of H. compacta var. aucklandiae Mortensen (1925)
from New Zealand waters, Clark was unable to reach a conclusion about the ‘Endeavour’
specimens. He determined the best course was to leave the ‘Endeavour’ material identified
as hyadesi, with the provisional note that the Great Australian Bight material probably

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

THE ECHINODERM GENUS HENRICIA

Fig. 1. Henricia compacta (holotype, BM(NH) 1890.5.7.830). a. abactinal view: b. actinal view. (R

PROC. LINN. SOC. N.S.W,, 109 ( 3), (1986) 1987

F. W. E. ROWE ANDE. L. ALBERTSON 185

Fig. 2. Henricia obesa (holotype, BM(NH) 1890.5.7.831). a. abactinal view; b. actinal view (R = 71mm).

PROC. LINN. SOG. N.S.W., 109 (3), (1986) 1987

186 THE ECHINODERM GENUS HENRICIA

represented a different species from that further east in Bass Strait and off eastern
Tasmania.

Fisher (1911, 1940) and Hayashi (1940) both expressed the great variability of species
within the genus Henricia, and the extreme difficulty in satisfactorily classifying them.

According to A. M. Clark (1962), Fisher (1940) had done much to clarify the inter-
relationships of species within the genus Henricia when he grouped the Southern Ocean
species into three superspecies. She (1962) noted Fisher’s (1940) re-identification of three
of H. L. Clark’s (1916) Australian specimens of Henricia and accepted Fisher's (1940)
synonymy of Ayadesi (Perrier) with obesa (Sladen). A. M. Clark (1962) did not examine

material from the Australian coast but suggested obesa‘. . . probably occurs off the south
coast of Australia. She also provided additional information about the holotype of H.
sufflata (Sladen).

Hayashi (1973) commenting again on the difficulty of classifying the species of Henricia
observed that the reproductive strategy of direct development might lead to limited distri-
bution and that‘. . . this local isolation might have exerted the important influence on the
differentiation of species’.

During the last decade some 250 specimens of Henricia have been collected from
southeastern Australian waters (coast of New South Wales and eastern Victoria) during
cruises of the N.S.W. State Fisheries Research Vessel ‘Kapala. These specimens have been
deposited with the Australian Museum (AM) collections. This material has been examined
as part of a Marine Sciences and Technologies Grants Scheme (MST) supported study of
the echinoderm fauna of NSW. The material has been compared with the remaining
‘Endeavour specimens of Henricia held in the Australian Museum, 8 specimens from
southern Australian (Victorian) waters held in the Museum of Victoria(MV), together
with the holotype specimens of H. compacta, H. sufflata and H. obesa which are housed in the
British Museum (Natural History) (BM(NH), London. Comparisons have also been made
with available material of New Zealand and South American species.

As a result of this study we conclude that both H. compacta and H. obesa occur in
southeastern and southern Australian waters. Additionally a new species is recognized
from New South Wales. Two ‘Endeavour’ specimens, which were identified as hyadesi by
H. L. Clark (1916), 16 specimens collected more recently from New South Wales waters
during ‘Kapala’ cruises and 7 from Victorian waters have features we recognize as
warranting the establishment of a new echinasterid genus. This material together with a
specimen from Japan, one from Washington State (west coast of North America), (United
States National Museum, Washington, D.C., U.S.A.), and two from South African waters
(BM(NRH) ), will be dealt with by us elsewhere.

SYSTEMATIC ACCOUNT
Family Echinasteridae
Genus Henricia Gray, 1840

Henricia compacta (Sladen)
Fig. la-b

Cribrella compacta Sladen, 1889: 543, p. XCVI, figs 1-2, pl. XCVIII, figs 3-4.

Henricia compacta, Mortensen, 1925: 307; Fisher, 1940: 163, 164, 166; H. E. S. Clark, 1970:
4,

Henricia hyadest, H. L. Clark, 1916: 60(part), 1946: 148(part). [Non H. hyadesi (Perrier) =
H.. obesa (Sladen) according to Fisher, 1940: 164].

Diagnosis: R up to 85mm, r up to 14mm, R/r = 4.4.-7.5; arms slender, tapering to a fairly
acute tip; abactinal skeleton compact, plates crescentic; papular areas small, rarely contain-

PROC. LINN. SOC. N.SW., 109 (3), (1986) 1987

FE. W. E. ROWE ANDE. L. ALBERTSON 187

“fy

Pgs

Fig. 3. Henricia kapalae sp. nov. (holotype; AM J19707). a. abactinal view (R = 56mm); b. spines on abactinal
surface of ray.

PROG. LINN. SOC. N.S.W., 109 (3), (1986) 1987

188 THE ECHINODERM GENUS HENRICIA

ing 1-2 small accessory plates, papulae 1-3 per area; abactinal spinelets stout (up to 0.2mm
long, 0.05mm wide), tapering only near the tip, terminating in a few points, up to 45 spine-
lets in triple, crowded rows across the plates; marginal plates not prominent unless cleaned
of spines, plates quadrilobed of similar size; intermarginal plates few, extending to 3rd mar-
ginal plate in holotype (R=15.5mm) or 15th marginal plate in largest specimen
(R=85mm); single row of actinal plates, extending almost to arm tip, plates squarish, with
up to 35 spinelets; adambulacral plates slightly wider than long, rectangular; 2-3 furrow
spines on vertical face of adambulacral plates, 2-3 prominent subambulacral spines on
adradial edge of plate, behind which are 2-3 slightly smaller spines, the remainder of the
plate bears up to 30-35 spinelets similar to those on the adjacent actinal plates; papulae
occur between the marginal, intermarginal and inferomarginal and actinal plates along the
arms.

Material examined: Holotype, BM(NH) 1890.5.7.830, 38°50’S, 169°20’E, off W coast
of North Island of New Zealand, 503m; 2 specimens, AM J3075, E3773(1), Great Aus-
tralian Bight, 146-274m; 1 specimen, E4716, S of Gabo Island, Victoria, 366m; 2 speci-
mens, J19700, 58km off Mount Cann, Victoria, 205m; 2 specimens, J12886, 33°18.8’S,
127°19.3’ Eto 33°20.8’S, 127°29.7’E, off Terrigal, N.S:W., 300-310m; 1 specimen, J18629,
32°58.8'S, 152°41.6’E, off Newcastle, N.S.W., 1150-951m; 1 specimen, J12875, NE of
Wollongong, N.S.W. 457m; 1 specimen J12863, 33°38’S, 151°57’E to 33°34’S, 152°01E,
E of Broken Bay, N.S.W., 786-805m; 1 specimen, J12860, 34°18’S, 151°26’E to 34°24’S,
151°23’E, E of Bulli (Wollongong), N.S.W. 457-476m; 2 specimens, J19701, 34°53’S,
151°08’E to 35°00’S, 151°06’E, off Lake Wollumboola, N.SW., 412m; 1 specimen,
J19699, 38°10’S, 149°52’E to 38°14’S, 149°43’E, SE of Point Hicks, Victoria, 457m; 6
specimens, 128995 33° 2005; 192° to 33°30S; 192707 Ee ott Broken*BayaiNesie
640m; 1 specimen, J12879, 34°18’S, 152°26’E to 34°24’S, 151921 E, NE of Wollongong,
N'S.W,, 497-485m; 1 specimen, J19702, 34°287S, 151°19" Eto 34°347S; (51°17; E; BiotPort
Kembla, N.S.W., 503m; 4 specimens, J12861, 34°39’S, 151915’ Eto 34°32’S, 151°19”’E, E
of Kiama, N.S.W., 412m; 1 specimen, J12694, 33°40’S, 151°53” E to 33°46’S, 151°497 E,
off Broken Bay, N.S.W., 384m; 4 specimens, J12882, 34°24’S, 151°25’E to 34°23/S,
151°25’E, SE of Sydney, N.S.W., 731-768m; 28 specimens, J13203, 33°35’S, 152°01’°E to
3332) 5s lo22034E, eof Broken’ Bay, NsoiW., 823m: 2*specimensn)1 28693 oa:
150°46’E to 35°36’S, 150°43’E, E of Brush Island, N.S.W., 494m; 2 specimens, J12867,
37°48'S, 150°13’E to 33°37’S, 150°16’E, SE of Gabo Island, Victoria, 494m; 1 specimen,
J19698, 38°06’S, 149°58’E to 38°00’S, 150°02’E, SE of Point Hicks, Victoria, 329m; 1
specimen, J17268, E of Nambucca Heads, N.S.W., 274m; 1 specimen, J12870, 35°32’S,
150°46’E to 35°35’S, 150°45’E, E of Brush Island, N.S.W., 503m; 5 specimens, J12881,
33°43S, 151946’ E to 33°41’S, 151943’E, E of Broken Bay, N.S.W., 170m; 1 specimen,
13207, 37°397S; 159719" EF t0372427S)150°18 (ESE ofiGabo Islandy Victonase/oimesl9
specimens, J13193, 33°34 7S, 152°02E to. 33°31 S, 152°047E) E of Broken BayNESiwe
914m; 15 specimens, J13192,33°357S; 152°00) Eto33°330S; 192902, E. off BrokeniBay,
N.S.W., 823m; 8 specimens, J13201(1), J17262(1), J13256(6), 34°22’S, 151923’ E to 34°19’S,
151°25’E, E of Wollongong, N.S.W., 823m; 1 specimen, J12880, 29°45’S, 153°45’E to
29°42’S, 153°46’E, off Sandon Bluffs, N.S.W., 503m; 1 specimen (eight arms), J19703,
35°02"S, 151°06’E to 34°58’S, 151°08’E, off Shoalhaven Bight, N.S.W., 439-420m; 36
specimens, J13332, 33°36’S, 152°06’E to 33°34’S, 152°08’E, E of Broken Bay, N.S.W.,
914m; 3 specimens, 33°32’S, 152°06’E to 33°34’S, 152°05’E, off Broken Bay, N.S.W.,
823m; 9 specimens, J13277(7), J17263(2), 33°39’S, 152°06’E to 33°37’S, 192°07E, off
Long Reef (Collaroy), N.S.W., 1006m; 6 specimens, J17265(2), J17267(4), 33°38'S,
152°02’E to 33°36’S, 152°04’E, off Long Reef (Collaroy), N.S.W., 960-988m; 1 specimen,
J18618, 35°30’S, 150°52’E to 35°28’S, 150°53’E, off Brush Island, N.S.W., 933-988m.
Distribution: Southern Australia (Great Australian Bight) and Tasman Sea, from

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

F. W. E. ROWE ANDE. L. ALBERTSON 189

Fig. 4. Henricia kapalae sp. nov. (holotype; AM J19707). a. actinal surface and marginal plates; b. actinal surface
with oral plate spine detail.

PROC. LINN. SOC. N.S.W,, 109 (3), (1986) 1987

190 THE ECHINODERM GENUS HENRICIA

southeastern Australia (Sandon Bluffs, N.S.W. — Point Hicks, Victoria) to the west coast
of New Zealand, in depths ranging from 146-1150m.

Remarks: H. compacta (Sladen) is based on a juvenile specimen (R=15.5mm) which Sladen
considered ‘sufficiently well marked to Justify its being regarded as a distinct species. He
enumerated its delicate form, large plates with compact groups of numerous spinelets,
small papulae areas, distinct marginals and armature of adambulacral plates as charac-
teristic of the species. Examination of some 170 specimens in the present collection rang-
ing from R=20-85mm has shown that the arms tend to become relatively more elongate
with growth (i.e. the disc does not increase in width beyond R=c.30-40mm), that the mar-
ginal plates become less prominent beyond R=40mm and there are two (often three in
larger specimens) furrow spines on the vertical face of the adambulacral plates, even in the
small holotype. The relationships of this species have been discussed by Fisher (1940).

Henricia obesa (Sladen)

Fig. 2a-b

Cribrella obesa Sladen 1889: 544, pl. XCVI, figs 3-4, pl. XCVIII, figs 5-6.
Cribrella hyades: Perrier, 1891: 100, pl. TX, fig. 1, pl. X, fig. 2.

Henricia hyadest, H. L. Clark, 1916: 60(part); 1946: 148(part).

Henricia obesa, Fisher, 1940: 164; A. M. Clark, 1962: 48, figs 5n, 6a-c.

Diagnosis: R up to 100mm, r up to 14mm, R/r=4.0-5.6, rarely to 7.1; arms slightly inflated
proximally, slender distally, tapering to a narrow, rounded tip; abactinal skeleton reticu-
late, open meshwork, 1-2 small accessory plates sometimes present in papular areas, 1-6
papulae per area; abactinal spinelets stout (0.3-0.4mm long, 0.1-0.15mm wide), rounded
to slightly bulbous at the tip, terminating in many points, up to about 20 spinelets per plate,
in single or double rows; marginal plates prominent in the holotype, less so in any other
material without clearing of spinelets; 16-18 inferomarginal plates per 20 adambulacrals,
marginal plates quadrilobed, superomarginals slightly smaller than inferomarginals; in-
termarginal plates, 1-3 rows proximally, reducing to one row, extending to 1/2-2/3R; two
rows of actinal plates, second row extending only to 1/4R, actinal plates with up to 10 spine-
lets; adambulacral plates with two furrow spinelets on vertical surface; usually a single,
prominent subambulacral spine on the adradial edge of the plate, behind which are 6-8
spines which decrease in size across the plate, these are arranged either 2 or 3 in single
series, the remaining 4-6 spines in pairs, or all are more or less in pairs behind the inner-
most spine; papulae occur between the intermarginal plates and are larger and prominent
between the actinal plates along the arms.

Material examined: Holotype, BM(NH)1890.5.7.831, 51°40’S, 57°50’W, Port William,
Falkland Islands, 22m; 1 specimen, MV75-9, 38°44’S, 141°33’E, 30km S of Cape Nelson,
Victoria, 155.4m; 2 specimens, AM E5933, 58km S of Mt. Cann, Victoria, 205m; 1 speci-
men, E4712, S of Gabo Island, Victoria, 365m; 1 specimen, E5031, off Babel Island, Bass
Strait, 73-110m; 1 specimen, J5855, Bass Strait, trawled; 2 specimens, J8791, off Newcastle,
N.S.W. trawled; 1 specimen, J19697, off Eden, N.S.W., trawled; 3 specimens, J18628, E of
Gabo Island, Victoria, 402-439m; 1 specimen, J12878, 34°09’S, 151916’ E to 34°03’S,
151°21°E, E of Cronulla, N.S.W., 126-132m; 1 specimen, J13287, 34°37’S, 151°16’E to
34°44’S, 151°12’E, E of Kiama, N.S.W., 457m; 1 specimen, J12689, 34°49’S, 151°10’E to
34°56’S, 151°09’E, E of Shoalhaven Bight, 457-475m; 12 specimens, J12691, 34°53’S,
151°08’E to 35°00’S, 151°06’E, E of Shoalhaven Bight, N.S.W., 402-439m; 3 specimens,
J12858, 35°38’S, 150°40’E to 35°32’S, 150°45’E, E of Brush Island, N.S.W., 393-439m;
9 specimens, J12864(6), J12876(1), J12692(2), 37°45’S, 150°12’ Eto 37°38’S, 150°16’E,
SE of Gabo Island, 402-439m; 1 specimen, J12693, 38°10’S, 149°52’E to 38°14’S,
149°43”E, E of Cape Everard, Victoria; 1 specimen, J12690, 34°28’S, 151°19’E to

PROC. LINN. SOC. N.SW., 109 (3), (1986) 1987

F. W. E. ROWE ANDE. L. ALBERTSON 191

% ame x Nyy
Proximal P » eS AY a

Lo 1) Ts Bee
f \ | 4 NS
Bo
/ on
7]
A
Imm

proximal >

& <J distal

Fig. 5. Henricia kapalae sp. nov. (holotype; AM J19707). A. denuded abactinal plates of ray (proximal); B. abactinal
ray spines (proximal); C. denuded marginal plates, from 13th to 17th inferomarginal plates. AB = abactinal
plates, S = superomarginal plates, IT = intermarginal plates, IF = inferomarginal plates, AC = actinal plates,
AD = adambulacral plates; D. adambulacral and actinal plates and spines; E. oral plate pair with spines.

PROG. LINN. SOC. N.S.W., 109 (3), (1986) 1987

192 THE ECHINODERM GENUS HENRICIA

34°34°S, 151°17’E, E of Port Kembla, N.S.W., 402m; 2 specimens, J12877, 35°29’S,
150°47’E to 35°25’S, 150°50’E, SE of Ulladulla, N.SW., 439m; 3 specimens, J19696,
34°39 7S, 151°lb7 Ey to 34.232 7S) ol 19) BE of Kiama, 412m: 4) specimens, sjil2o72-
38°00’S, 150°02’E to 38°06’S, 149°58’E, SE of Point Hicks, Victoria, 330m; 1 specimen,
J10875, 35°32’S, 150°46’E to 35°35’S, 150°44’E, E of Brush Island, N.S.W., 384m; 1
specimen, J12873, 392368; 190°44" Eto 35-318) 190°47 “E) EofBird IslandaiNes.e
604m; 3 specimens, J13195, 35°32’S, 150°46’ E to 35°34’S, 150°45’E, E of Brush Island,
N.S.W., 384m; 3 specimens, J12865, 35°01 ’S, 151906’ E to 34°58’S, 151°07’E, E of Brush
Island, N.S.W., 420m; 1 specimen, J19695, 33°40’S, 151°53 E to 33°46’S, 151°49" E, off
Broken Bay, IN‘SIW)384m- 2) specimens, J12862, 35232) Ss lo0 245) Eu toms oe ome
150°35’E, E of Brush Island, N.S.W., 219-274m; 1 specimen, J13289, 35°00’S, 151°07’E
to 34°59’S, 151°08’E, off Beecroft Peninsula, N.S.W., 420m; 7 specimens; J13288,
35°02’S, 151°06’E to 34°51 °S, 151°08’E, off Shoalhaven Bight, N.S.W. 365m; 3 speci-
NSN, |UAIN/, SIDS) 1S), WOAH) Id, Wo) DH Sisy 1S), NOOO ANI, ALO oan,

Distribution: Widespread in the Southern Ocean, including Tristan da Cunha (Atlantic),
southern and southeastern Australia (off Newcastle, N.S.W., to Bass Strait and westward
to Cape Nelson, Victoria) and Macquarie Island (south of New Zealand) in depths ranging
from 22-604m.

Remarks: Comparison of the present material with the holotype of H. obesa leaves little
doubt that it is representative of that species. Structural variation appears to be relatively
slight, involving less pronounced marginal plates, the occurrence of subambulacral spines
in single to double rows across the adambulacral plates, and similarly, spinelets forming
single to double rows across the abactinal plates.

A. M. Clark noted (1962) that obesa differed from sufflata in its more robust abactinal
skeleton, the arrangement and higher number of abactinal spinelets and in the larger size
of the marginal plates relative to the adambulacral plates. Our examination of the holotype
of H. sufflata has confirmed these differences. However, the extent of the intermarginal
plates in obesa is greater than Clark (1962) suggests, extending in our material to at least
1/2R if not to 2/3R. We do not consider this to be of significance in determining the identity
of our material. In coming to this conclusion, we therefore disagree with Fisher's (1940)
identification of some of H. L. Clark’s (1916) specimens as sufflata. In fact, H. L. Clark
(1916, 1946) was substantially correct in his identification of the ‘Endeavour’ Henricia
material as H. hyadest, since that species 1s considered a Junior synonym of H. obesa (Sladen)
by Fisher (1940) and A. M. Clark (1962).

The identification of obesa in southern and southeastern Australian waters 1s now con-
firmed. The relationships of this species are fully discussed by A. M. Clark (1962).

Henricia kapalae sp. nov.
Figs 3a-b, 4a-b, 5A-E

Diagnosis: R=26-97mm, r=4.2-17.5mm, R/r=4.8-7.2mm, arms slender, with acute tip;
skeletal reticulum relatively compact, abactinal skeletal plates with single (rarely irregu-
larly double), webbed row of slender spinelets; marginal plates more or less distinct, in-
feromarginal plates twice as wide as superomarginal plates; intermarginal plates extend to
1/4R; 2 actinal rows of plates, the innermost extending to 1/4R; 2-3 spines in vertical series
in furrow; 1-4 papulae per area, actinally papulae restricted to disc.

Material examined: Holotype, AM J19707, 33°39’S, 152°06’E to 33°37’S, 152°07’E,
off Broken Bay, N.S.W., 990m; 16 paratypes, J19704(3), 33°35’S, 152°01’E to 33°32’S,
152°03’E, off Broken Bay, N.S.W., 450m; J19711(3), 33°30’S, 152°07’E, off Broken Bay,
N.S.W., 905m; J19710(1), 33°35’S, 152°03E, off Broken Bay, N.S.W., 823m; J19708(3),
33°39°S, 152°06’E, to 33°37’S, 152°07°E, off Broken Bay, N.S.W., 990m; J19705(1),

PROC. LINN. SOC. N.SW., 109 (3), (1986) 1987

F. W. E. ROWE AND E. L. ALBERTSON 193

35-5009, lo2-064Rs oft) Broken Bays) NESW 914m: J19709()) 33°43." 8; 151°46"E to
33241 7S,, 151°43E, off Broken Bay, NiS:W., 170m; J13323(1), 34°54°S, 151°127E to
g4257 25. lololie@k, off Shoalhaven, N:S:\W., 540m; J18617@), 35°297S;,150°52" EF, to
35°26’S, 150°55’E, off Brush Island N.S.W., 1006m; J19679(1), 35°30’S, 150°54’E to
35°27'S, 150°55’E, off Brush Island, N.S.W., 979-1070m; J19706(1), 35°38’S, 150°40’E
to 35°32’S, 150°45’E, off Brush Island, N.S.W., 393-439m.

Distribution: Between Broken Bay and Brush Island, N.S.W., in depths ranging between
170-1070m.

Etymology: Named for FRV ‘Kapala’ N.S.W. State Fisheries Research Vessel from which
all specimens were collected.

Description: Rays 5 slender, tapering to a narrow tip, R=26-97mm, r=4.2-17.5mm,
R/r=4.8-7.2, Br=4.2-17.5 (Holotype, R=56mm, r=8mm, R/r=7.0mm, Br=9mm). The
disc is relatively small. The madreporite which bears small spinelets, occurs interradially,
about 2/3r from the centre of the disc. Abactinal plates are irregularly bar-like to crescentic
in shape, forming an irregular, relatively compact recticulum. The medial region of the
plates is raised into a low ridge, which bears a single, webbed row, occasionally an irregular
double row, of delicate, finely tapered spinelets. The spinelets measure up to 0.5mm in
length.

There are 1-4 papulae per area abactinally. The papulae extend to the actinal surface,
where they occur one per area, restricted to the disc.

Marginal plates distinct due to their regular shape and alignment. The supero-
marginals are much smaller and less prominent than the inferomarginals. The supero-
marginals are quadrilobed, about as long as wide, with a slightly oblique, actinal/abactinal
directed ridge bearing spinelets. The inferomarginal plates are also quadrilobed, about
twice as wide as long, and therefore twice as wide as the superomarginals. The infero-
marginal plates each bear a slightly oblique ridge bearing spinelets up to 0.70mm long. The
spines are usually in a single more or less uniform row but on some specimens, including
the holotype, the spines occur in an irregular double row. Intermarginal plates, similar to
the superomarginal plates, occur in 2-3 rows, the longest series extending to only about
1/4R. There are 25 inferomarginal plates per 20 adambulacral plates. Actinal plates occur
in two rows, the first row extends only to about 1/4R, the second comprising 2-3 plates in
the actinal angle. The plates are centrally ridged, and bear a group of spines, the central
ones of which are the largest.

The adambulacral plates bear 2(3) stout, cylindrical spines in vertical series in the fur-
row. There is a single prominent, cylindrical spine on the adradial edge of the plate behind
which occurs two similar spines. Behind this group of spines is a further group of three
slightly shorter spines followed by up to 20 smaller, slender spines which are similar to those
on the actinal and abactinal plates.

The oral plates bear the usual complement of spines.

Remarks: H. kapalae is immediately distinguished from H. compacta on skeletal morphology
and spine armament, even though they show a similar shape of tapering arms. The stouter
arms, shape of spinelets, fewer inferomarginal plates to adambulacral plates, less promi-
nent marginal plates and extent of actinal papulae all serve to easily distinguish H. obesa
from H. kapalae.

H. kapalae clearly falls into Hayashr’s (1940) B-group of species from Japanese waters.
It differs from each of those species, H. pacifica Hayashi, H. aspera Fisher, H. ohshimai
Hayashi, H. ohshimai acutispina Hayashi and H. pachyderma in: having regularly 2 spines in
vertical series in the furrow; the form of the marginal plates and spines; and spine arrange-
ment. The very delicate, open skeletal reticulum and spine form and arrangement of H.
mutans Koehler, from the Andaman Islands, and H. arcystata Fisher, from Philippine seas,
excludes the identification of the N.S.W. material with either of those species.

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

194 THE ECHINODERM GENUS HENRICIA

ACKNOWLEDGEMENTS

We wish to thank Mr Ken Graham for collecting material on FRV ‘Kapala’ and
N.S.W. State Fisheries for donating the material to the Australian Museum; we thank also
Miss A. M. Clark, British Museum (Natural History), London, U.K., for the loan of type
material, and Dr D. L. Pawson and Miss M. Downey for material borrowed from the
United States National Museum, Washington, D.C., U.S.A. The senior author wishes to
acknowledge support of Marine Sciences and Technologies Grants Scheme for support

(MST: 84/2092) of his study of the echinoderm fauna of N.S.W.

References

CLARK, A. M., 1962. — Asteroidea. B.A.N.Z.A.R.E. Rep. Series B 9: 1-104, 18 text-figs., 14 tables, 5 pls.

CiaRK, H. E. S., 1970. — Sea-stars (Echinodermata: Asteroidea) from ‘Eltanin’ Cruise 26, with a review of the
New Zealand asteroid fauna. Zool. Publs Vict. Univ. Wellington No. 52: 1-31, 3 figs, 3 pls.

CLARK, H. L., 1916. — Report on the sea-lilies, starfishes, brittle-stars and sea-urchins obtained by the F.1.S.
‘Endeavour’ on the coasts of Queensland, N.S.W., Tasmania, Victoria, S. Australia and W. Australia.
Endeavour Res. 4: 1-123, 11 figs, 44 pls.

—., 1946. — The echinoderm fauna of Australia. Publs Carnegie Instn No. 566: 1-567.

FISHER, W. K., 1906. — The starfishes of the Hawaiian Islands. Bull. U.S. Fish. Comm. 23(3): 987-1130, 49 pls.

—— , 1911. — Asteroidea of the North Pacific and adjacent waters. 1. Phanerozonia and Spinulosa. Bull. US.
Natn. Mus. 76: 1-406, 122 pls.

——, 1913. — New starfishes from the Philippine Islands, Celebes and the Moluccas. Proc. U.S. Natn. Mus. 46:
201-224.

—, 1919. — Starfishes of the Philippine Islands and adjacent waters. Bull. U.S. Natn. Mus. 100(3): 1-711, 156 pls.

——, 1940. — Asteroidea. Discovery Rep. 20: 69-306, figs A-M, 23 pls.

Gray, J. E., 1840. — A synopsis of the genera and species of the Class Hypostoma (Asterias Linn.) Ann. Mag. Nat.
Fast. (1)6: 175-184.

HayaSHI, R., 1940. — Contribution to the classification of the Sea-stars of Japan. I. Spinulosa. J. Fac. Scz.,
Hokkaido Imp. Univ. (6)7: 107-204, 63 text-figs, 7 pls.

——.,, 1973. — The sea-stars of Sagami Bay. (Biological Laboratory, Imperial Household) Tokyo: 1-114, 13 text-figs,
18 pls.

KOEHLER, R., 1909. — Deep-sea Asteroidea. Echinoderma of the Indian Museum Calcutta: 1-143, 13 pls.
MorTENSEN, T., 1925. — Echinoderms of New Zealand and the Auckland-Campbell Is. III-IV. Asteroidea,
Holothuroidea and Crinoidea. Vidensk Medd. naturh. Foren., Kbh. 79: 263-420, 70 figs, pls x11-xiv.
PERRIER, E., 1891. — Echinoderma. I. Stéllerides. Mission Scientifique du Cap Horn, 1882-83. 6. Zool., Paris, 198

pp., 13 pls.
SLADEN, W. P., 1889. — Asteroidea. Rep. Scient. Results Voy. ‘Challenger’(Zool.) 30: 893 pp., 117 pls.

PROC. LINN. SOC. N.SW., 109 (3), (1986) 1987

A new Species in the echinasterid Genus
Echinaster Muller and ‘Iroschel, 1840
(Echinodermata: Asteroidea) from southeastern
Australia and Norfolk Island

FRANCIS W. E. ROWE and E. LYNNE ALBERTSON

RowgE, F. W. E., & ALBERTSON, E. L. A new species in the echinasterid genus Echinaster
Muller and Troschel, 1840 (Echinodermata: Asteroidea) from southeastern Australia
and Norfolk Island. Proc. Linn. Soc. N.S.W. 109(3), (1986) 1987: 195-202.

A new species of Echinaster, E. colemanz, is described from the southeastern coast of
Australia between Moreton Bay, Queensland and Ulladulla, N.S.W. possibly as far south
as Bass Strait, and from Norfolk Island. The relationships of E. coleman: with the southern
Australian endemic species E. arcystatus H. L. Clark and E. glomeratus H. L. Clark are dis-
cussed. The new species most likely evolved from E. arcystatus as a result of isolation on the
east coast due to the emergence of Bass Strait during glacial periods. The most recent emer-
gence of Bass Strait occurred 18-20,000 years ago. Distribution of E. coleman: to Norfolk Is-
land can only be explained by trans-Iasman larval transportation.

FW. E. Rowe and E. L. Albertson, Division of Invertebrate Zoology (Echinoderms), Australian
Museum, Sydney, Australia 2000; manuscript received 19 November 1986, accepted for publication 18
February 1987.

INTRODUCTION

During an investigation of the echinoderm fauna of New South Wales, Lord Howe
Island and Norfolk Island, a number of Echinaster specimens in the Australian Museum
(AM) collections were examined. These specimens were without species epithet, or pro-
visionally identified as either E. glomeratus H. L. Clark or E. arcystatus H. L. Clark. More
specimens of Echinaster have been collected from New South Wales waters as a result of this
project, which has been substantially funded by the Marine Sciences and Technologies
Grant Scheme (MST). In addition, specimens from Tasman locations collected by the New
Zealand Oceanographic Institution (NZOI) were examined by the senior author whilst
visiting the National Museum of New Zealand, Wellington. Reappraisal of these speci-
mens showed the occurrence of a new Echinaster species on the southeastern coast of Aus-
tralia and at Norfolk Island. This is the first report of Echinaster from Norfolk Island. Table
1 lists all known species of Echinaster occurring in Australian waters with their distribu-
tions. A key is also provided for the species.

SYSTEMATIC ACCOUNT
Family Echinasteridae
Genus Echinaster Miller & Troschel, 1840

Key to Australian species of Echinaster

1. Abactinal spinelets large, conspicuous, up to 4-5mm long, well spaced, single, on the
primary plates only, arms cylindrical.
E. callosus
1’. Abactinal spinelets smaller, <2mm long, numerous, single or in groups on primary
plates, sometimes spinelets occur also on secondary abactinal plates, arms cylindrical
or widened at base.

oY)
tho

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

196 NEW SPECIES OF ECHINASTER

2. Abactinal spinelets single, on primary plates, skeletal reticulum with relatively small
papular areas, arms cylindrical or widened at the base.
3:
2’. Abactinal spinelets in small (2-4) to large (5-60) groups on primary plates, sometimes
spinelets occur on secondary plates, relatively large papular areas, arms cylindrical.
6.
3. Arms cylindrical, disc small.
Ae.
3’. Arms widened at base, disc relatively large.
E. stereosomus

4. 5-7 arms, autotomus, usually more than one madreporite, subambulacral spines well
developed.
E. luzonicus
4’. 5 arms, not autotomus, single madreporite, subambulacral spines not well developed.
5,
5. Abactinal and actinal spinelets include stout, chisel-shaped or club-shaped forms.
E. superbus

5’. Abactinal and actinal spinelets uniform in size and shape, bluntly rounded at tips or
truncate, some may be pitted at tip.
E. varicolor
6. | Abactinal spinelets in large discrete groups on primary plates (5-60). Papulae extend
to inferomarginal line.
E. glomeratus
6’. Abactinal spinelets in small groups on primary plates (2-4). also singly on secondary
plates, papulae restricted to abactinal surface above the superomarginal line.
Te
7. Papulae 11-40 (up to about 60) per area, papular areas up to about 15.00mm
diameter.
E. arcystatus
7’. Papulae 6-8 (up to about 14) per area, papular areas up to about 6.5mm diameter.
E. colemani

Echinaster colemani n.sp.

Figs la-b, 2

Diagnosis: A species of Echinaster with a well developed reticular abactinal skeleton; spine-
lets occur in groups of 2-4 on primary plates at reticular junctions and singly on secondary
plates between junctions; papular areas of skeletal reticulum up to 6.5mm diameter, con-
taining 3-14, usually 6-8 papulae; papulae occur abactinally as far as the superomarginal
line.

Material examined: Holotype, AM J13076, R/r=110mm/13mm, Bate Bay, off Cronulla,
N.S.W., 24.4m, rocky bottom; (14 paratypes), 1 paratype, J10862, R/r=40/8, off Moreton
Bay, Old, 76.8m; 1 paratype, J15258, R/r=85/13.5, Julian Rocks, Byron Bay, N.S.W.,
24.4m; 1 paratype, J15259, R/r=43/8.4, South Solitary Islands, off Coffs Harbour, N.S.W.,
27.4mm; 2 paratypes, J16541, R/r=71/10, J16534, R/r=100/13, Broughton Island, near Port
Stephens, N.S.W., 18m, reef; 1 paratype, J13000, R/r=65/10.5, Broughton Island, near Port
Stephens, N.S.W., 25m, on rocks; 1 paratype, J13077, R/r=92/15, off Boat Harbour, north
of Cronulla, N.SW., 39.7m; 1 paratype, J13084, R/r=75.5/13, off Cronulla, N.S.W., 30.5m;
1 paratype, J10835, R/r=80/13, Jibbon Point, Bundeena, N.S.W., 25m, sand and rubble;
1 paratype, J590, R/r=106/14.5, Newcastle Bight, N.S.W.; 1 paratype, J9182, R/r=56/12.5,
Bass Point, N.SW., 17m, bottom cover of sponge and coral; 2 paratypes J14137,

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

F. W. E. ROWE ANDE. L. ALBERTSON 197

TABLE 1

Species of the genus Echinaster occurring in Australian waters

(7. L. Clark, 1946, L. M. Marsh, 1976, Rowe (unpublished) )
Species/Author Distribution
E. arcystatus H. L. Clark Shark Bay, W.A. south to Waterloo Bay, Wilsons Promontory, Vict. (Endemic).
*E. callosus von Marenzeller Lizard Island, Qld. (Indo-west Pacific; Clark and Rowe, 1971).

E. glomeratus H. L. Clark Abrolhos Islands, W.A., Dongarra, W.A., south to Kangaroo Island, S.A.
(Endemic).

E. luzonicus (Gray) Exmouth Gulf, W.A. north and east along the coast to Double Island Point, Qld,
south to Solitary Islands, N.S.W.; offshore reefs, Rowley Shoals and Ashmore,
W.A. along the Great Barrier Reef from Murray Islands south to Bunker Group,
Qld. (East Indian region to islands of the south-west Pacific; Clark and Rowe,

1971).

E. stereosomus Fisher Murchison, W.A. north and east to Fraser Island, Old, south to Brunswick

(syn. E. acanthodes H. L. Heads, N.S.W. (Philippines south to Australia; Jangoux, 1978).

Clark; Jangoux, 1978)

E. superbus H. L. Clark Broome south west to Dampier Archipelago (N.W. Cape). W.A. (Endemic).

E. varicolor H. L. Clark Broome south to Esperance, W.A. (Endemic).

E. colemani n.sp. Moreton Bay, Qld, south to Ulladulla, N.S.W., possibly to Bass Strait; Norfolk
Island.

* New record based on 2 specimens held in the Australian Museum collections (AM J9674; 13096).

R/r=137.5/18.5, 115/16.2, Ulladulla, N.S.W., 24.4m, rocky bottom; 1 paratype, ? Bass
Strait, trawled (? 91-110m); 1 specimen NZOI, R/r=137/18.5, Norfolk Island, 5-15m.
Distribution: Moreton Bay, Old, south to Ulladulla, N.S.W., (? Bass Strait), Norfolk
Island (N.E. Tasman Sea), in depths ranging from 17-40m (? possibly to 91-110m).
Description: R=40-137mm, r=8-18.5mm, R/r=4.48-8.4 (av. 6.4). Arms 5, rounded in
cross-section, tapering evenly to a blunt tip, though slightly constricted at the base (Figs la-
b). The body is covered by a thick skin. The disc 1s relatively small with the madreporite,
which bears small spinelets, close to the interradial angle. The abactinal skeleton forms an
open reticulum, the papular areas of which are more or less pentagonal and range from 3-
6.5mm diameter in the specimens examined (Fig. 2). The papular areas contain 3-14
papulae, but more often 6-8 papulae. The papulae extend to the superomarginal line.
Below the superomarginals groups of papula-like patches of skin occur, but these do not ex-
tend to the coelom due to a dense, fibrous tissue occluding the spaces between the plates.
The primary abactinal plates at each of the 5 junctions of the pentagonal papular areas,
bear groups of 2-4 small, bluntly pointed spinelets. One to four spinelets occur singly,
spaced, on secondary plates delimiting the circumference of the papular areas.
Superomarginal plates are irregularly quadrilobed and bear 2-3 spinelets (1 distally
on the arms). Inferomarginal plates are similarly quadrilobed, but larger than the supero-
marginals. They similarly bear 2-3 spinelets (1 distally). An irregular series of inter-
marginal plates extends to 1/5-1/4R. A number of these plates each bear a single spinelet.
Adambulacral plates are rectangular, broader than long. They bear a single, small
furrow spine on the vertical surface. Across the actinal surface of the plate are 3, sometimes
only 2, prominent subambulacral spines. The innermost and outermost are usually stout
and cylindrical, but the middle spine may be flattened chisel-shaped towards the tip, or
widened and scoop-shaped at the tip, the flattening or scooping being parallel to the furrow.
A row of actinal plates extends for about 1/5-1/4R, each plate bearing a single spinelet.
Oral plates bear 1 or 2 small furrow spinelets (where 2 are present they stand adjacent

PROC. LINN. SOC. N.S.W,, 109 (3), (1986) 1987

b. actinal surface (R=110mm

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chinaster colemani sp. nov. (holotype; AM J130

Fig. 1. E

F. W. E. ROWE ANDE. L. ALBERTSON 199

Fig. 2. Echinaster colemani sp. nov. (holotype) abactinal surface showing fine reticulation and papulae.

to one another) and 4-6 subambulacral spines. The subambulcral spines form a single or
irregularly double series behind the first spine at the apex of each oral plate.

Colour: In life this species is velvet brown with purple papulae (Coleman, pers. comm.).
Etymology: Named for Mr Neville Coleman, who has contributed many specimens to the
Australian Museum collections, including specimens of this species.

Remarks: E. colemani is most closely related to E. arcystatus H. L. Clark, 1914, (Fig. 3a, b)
and E. glomeratus H. L. Clark, 1916, (Fig. 4). These latter species are endemic to Australia
and occur sympatrically along most of the west and south coasts. All three species bear
clusters of spinelets on primary abactinal skeletal plates at the reticular junctions between
the papular areas. However, E. glomeratus is immediately distinguished from E. colemani and
E. arcystatus by: its stouter abactinal skeletal spinelets, which are restricted to discrete groups
of 10-20 or more (up to 60) on the primary plates (Fig. 4); the extension of papulae to the
inferomarginal line; and the arrangement of its subambulacral spines. E. colemani and
E. arcystatus share the restriction of papulae to the abactinal surface above the supero-
marginal line of plates, the occlusion of spaces between the plates below the superomarginal
line, the distribution of abactinal spinelets and arrangement of subambulacral spines. £.
colemani (Figs la-b, 2) differs from £. arcystatus (Figs 3a-b) principally in the consistently
smaller size of the abactinal papular areas (up to about 6.5mm, colemanz, up to about 15mm,
arcystatus) and fewer papulae per area (up to 14, usually 6-8, colemam; up to 60, usually 11-40,
arcystatus). The largest specimen of colemani known measures R=137mm; arcystatus

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

NEW SPECIES OF ECHINASTER

3. Echinaster arcystatus (AM J11873) a. abactinal surface (R=189mm); b. detail of reticulation, abactinal surface
of arm

PROC. LINN. SOC. N.SW., 109 (3), (1986) 1987

F. W. E. ROWE ANDE. L. ALBERTSON 201

Fig. 4. Echinaster glomeratus (holotype; AM J1624) abactinal surface (R=135mm).

R=186mm. The velvet brown colour of coleman: contrasts with the yellow/pink background
colour and bright purple papulae of arcystatus (Coleman, pers. comm. ).

It seems likely that E. colemani has evolved from E. arcystatus, or their common pre-
cursor, through isolation in the Tasman region, possibly as recently as 18-20,000 years ago
when Bass Strait was last emergent during a glacial period. E. arcystatus is known to occur
as far east along the south coast as Wilsons Promontory, Victoria (AM J11873).

The distribution of E. colemani south of Ulladulla on the New South Wales coast,
however, requires confirmation. We believe there is an element of doubt regarding the Bass
Strait locality attributed to the specimen (AM J8652) we identify as E. colemant reputedly
collected during the Endeavour (1909-1914) Expedition. Since the specimen was not seen
or reported by H. L. Clark (1916), it is possible that the label attached to the specimen may
have been associated with it by error.

The occurrence of E. colemani at Norfolk Island is interesting. The reproductive
strategy of colemani has not been determined, although its occurrence at Norfolk Island sug-
gests a strategy involving a planktotrophic larval stage. This would facilitate dispersal
across the Tasman from the coast of New South Wales in the known west to east current
tracts (Rowe, 1985). The apparent absence of this shallow-water species from Lord Howe
Island or other locations on the Lord Howe Ridge is perplexing. Considering the few speci-
mens found so far, absence from such intermediate locations may be due to lack of collect-
ing, rather than to any other agency.

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

202 NEW SPECIES OF ECHINASTER

In a preliminary analysis of the distributions of some 440 species which we identify as
occurring on the coast of New South Wales, no fewer than 54 species (12.3%) occur either
at Norfolk Island and/or Kermadec Islands and New Zealand, but not on the Lord Howe
Ridge. Of these only 8 occur in shallow water (< 30m), 13 occur across the continental shelf
(31-200m) and the remaining 33 occur on the slope and deeper (201+ m). Shallow-water
species which share with E. colemani a distribution along the New South Wales coast and at
Norfolk Island are the southern Australian asteroid species Austrofromia polypora and crinoid
species Antedon incommoda. The echinoid species Phyllacanthus parvispinus is distributed along
the New South Wales coast and occurs at the Kermadec Islands, whilst the southern Aus-
tralian asteroid species Allostichaster polyplax, the ophiuroid species Ophiopeza cylindrica and
holothurian species Chirzdota gigas are known also from the coast of New South Wales and
in New Zealand waters.

In discussing the distributions of species of Asterodiscides, Rowe (1985) suggested that
populations of the southern and southeast Australian species A. truncatus (Coleman), which
also occurs only at the Kermadec Islands and New Zealand region, might be self-sustaining
and relatively phenotypically stable in their northeastern Tasman locations. He considered
it is difficult to determine whether larval input was maintaining gene flow across the
Tasman, despite the occurrence of appropriate current tracts. Only further sampling along
the Lord Howe Ridge and in deeper waters of the Tasman Sea, together with detailed
studies of reproductive strategies of these echinoderms, will help to elucidate these
apparently disjunct distributions.

ACKNOWLEDGEMENTS

Mr Neville Coleman is acknowledged for his contribution of specimens to the echino-
derm collections and for bringing the species, described herein, to the attention of the
senior author. The senior author also wishes to thank Vicki Harriott, Lyle Vail and Anne
Hoggett for their collecting effort and earlier association with his project; Dr W. del Main
(New Zealand Oceanographic Institute) for the opportunity to examine the NZOI collec-
tions and Dr A. N. Baker (National Museum of New Zealand) for providing facilities and
discussions on the echinoderm collections in the NMNZ. Mr W. Zeidler, South Australian
Museum, is thanked for the loan of specimens of Echinaster arcystatus. Finally, Marine
Sciences and Technologies Grants Scheme is acknowledged by the senior author for its

generous support of this project (MST: 84/2092).

References

CLARK, A. M., and Rowg, F. W. E., 1971. — Monograph of shallow-water Indo-west Pacific echinoderms. London:
British Museum (Natural History), 238 pp, 100 text-figs, 31 pls.

CLARK, H. L., 1914. — The echinoderms of the Western Australian Museum. Rec. West. Aust. Mus. 1: 132-173,
pls. 17-26.

——.,, 1916. — Report on the sea-lilies, starfishes, brittle-stars and sea-urchins obtained by the F.I.S. ‘Endeavour’
on the coast of Queensland, N.S.W., Tasmania, Victoria, S. Australia and W. Australia. Endeavour Res. 4:
1-123, 11 figs, 44 pls.

——.,, 1938. — Echinoderms from Australia. Mem. Mus. Comp. Zool. Harv. 55: viii + 596, 63 figs, 28 pls.

, 1946. — The echinoderm fauna of Australia. Publs. Carnegie Instn. No. 566: 1-567.

JANGOUX, M., 1978. — Biological results of the Snellius Expedition. X XIX. Echinodermata, Asteroidea. Zool.
Meded. 52(25): 287-300, 2 figs, 3 pls.

MARSH, L. M., 1976. — Western Australian Asteroidea since H. L. Clark. Thalassia Jugoslavica 12(1): 213-225,
2 figs, 1 table.

ROWE, F. W. E., 1985. — Six new species of Asterodiscides A. M. Clark (Echinodermata, Asteroidea). with a dis-
cussion of the origin and distribution of the Asterodiscididae and other ‘amphi-Pacific’ echinoderms. Bull.
Mus. Natn. Hist. Nat., Paris (4) 7 (section A no. 3): 531-577, 1 fig, 3 pls, 3 tables.

PROC. LINN. SOC. N.SW,, 109 (3), (1986) 1987

Two new Species of Delma (Lacertilia:
Pygopodidae) from northeastern Queensland and
a Note on the Status of the Genus Aclys

GLENN M. SHEA

(Communicated by H. G. COGGER)

SHEA, G. M. Two new species of Delma (Lacertilia: Pygopodidae) from northeastern
Queensland and a note on the status of the genus Aclys. Proc. Linn. Soc. N.S.W. 109(3),
(1986) 1987:203-212.

Delma mitella sp. nov. and Delma labialis sp. nov. are described from two specimens each
from northeastern Queensland. The new species are diagnosed on the basis of size,
coloration and rostral scale shape. Records of D. inornata from northeastern Queensland
are rejected. Aclys is diagnosed by two derived character states and, in the absence of a
diagnosis for Delma that adequately demonstrates monophyly, retained as a genus distinct
from Delma. A key to the Delma of Queensland and updated distribution maps are provided.

Glenn M. Shea, Dept of Veterinary Anatomy, University of Sydney, Australia 2006; manuscript received
3 February 1987, accepted for publication 22 April 1987.

Since the publication of Kluge’s (1974) monograph of the Pygopodidae, only three new
pygopodids have been described: Pletholax gracilis edelensis Storr, 1978, Aprasia harold: Storr,
1978 and Aprasia rostrata fusca Storr, 1979!. This would suggest that knowledge of the alpha
taxonomy of the family is nearly complete. The discovery of two spectacular new species of
Delma with restricted known ranges in northeastern Queensland, described below, indicates
that this is not the case.

I have followed the definitions of Kluge (1974) for measurements and body scalation,
with the addition of head length, from tip of snout to rostral margin of ear, and hindlimb
length, from junction of limb flap with body to distal tip of flap. The head length measure-
ment of Kluge (1974) is here given as mouth length. The descriptions give measurement,
in millimetres, followed in parentheses by the value of each measurement as a percentage
of snout-vent length in the case of head length, hindlimb length and tail length, or as a per-
centage of head length, in the case of cephalic measurements. Measurements are linear, to
the nearest 0.5mm for non-cephalic characters and to the nearest 0.05mm for cephalic
characters.

Although the head shield characters defined and employed by Kluge (1974) are
‘homologous quantitative characters’ in terms of the reference points, they are not indepen-
dent, with changes in one scale frequently affecting several characters, and do not fully
describe the variation in individual scale morphology occurring between species. The
stability of many head shields in Delma species allows the more conventional nomenclature
used here. Rostral, rostral supranasal, caudal supranasal, postnasal, prefrontal, frontal,
supraocular, parietal, occipital and upper temporal scales are labelled in Fig. 1.
Supralabial, infralabial, nuchal and gular scale definitions follow Kluge (1974). Loreals are
those scales bordering the dorsal margin of the supralabials, from the caudal margin of the
postnasal up to and including the first scale contacting the enlarged subocular supralabial.
Supraciliaries are those scales bordering the lateral margin of the supraoculars, from the
scale contacting prefrontal to that contacting parietal. Preoculars are those small scales

1 Ofthe three species named by Wells and Wellington (1985), Delma wollemi’ and ‘Pygopus territorianus’ are nomina
nuda while ‘Pygopus kluget’is of uncertain status. P klugei’ was differentiated from P. nigriceps schraderi on the basis
of distinctly keeled body scales and a habitat restriction to black soil plains (vs smooth scales and red sand plains),
and was described from a single specimen. However, Kluge (1974) included numerous specimens from black soil
plains in his redescription of PR. n. schraderi, only one of which had keeled scales.

PROC. LINN. SOC. N.S.W,, 109 (3), (1986) 1987

204 TWO NEW SPECIES OF DELMA

between loreals, prefrontal, supraciliaries and the bony margin of the orbit, while sub-
oculars are those scales bordering the dorsal margin of the subocular supralabial, but not
contacting the preceding or succeeding supralabials.

Head shields are numbered rostrad to caudad, while longitudinal scale rows on body
and tail are numbered from the dorsal midline.

Delma mitella sp. nov.
Figs 1,2

Holotype: Queensland Museum J32597, Herberton area, Old. R. Russel.
Paratype: Australian Museum R65264, Koombooloomba rd, near Ravenshoe, Qld.
T. Bentz, 19.x1.1967.

DIAGNOSIS: D. mitella differs from all other Delma species in its greater size (to 154mm SVL
vs to 133mm, with only D. fraser, D. grayti, D. inornata and D. plebera attaining more than
115mm), almost straight or concave suture between rostral and rostral supranasals (vs ob-
tusely gabled apex of rostral partly projecting between rostral supranasals) and presence
of a dark line along fifth scale row from caudal body to tail, sharply demarcating dark
dorsal and lateral surfaces from light ventral surface.

DESCRIPTION OF HOLOTYPE: Rostral barely projecting between rostral supranasals; rostral
supranasal in broad contact with first supralabial; caudal supranasals present, in point con-
tact with nostril; postnasal single; loreals five, subequal; preoculars nine (left) or ten (right);
suboculars three, third elongate; supraciliaries five, caudalmost large and in line with
preceeding series; supraoculars two, first longer; supralabials six, fourth below centre of
eye, caudalmost low and elongate; infralabials seven, first pair narrowly separated on ven-
tral midline, second pair widely separated; occipital present; upper temporals two; nuchal
scales 13; gular scales 14.

Midbody scales 16; ventral scales 74; ventral body scales transversely enlarged;
preanal scales three; hindlimb scales five.

Snout-vent length 150.5; head length 14.75 (9.8); mouth length 12.30 (83.4); snout
length 6.35 (43.1); eye width 1.90 (12.9); postorbital length 3.85 (26.1); head width 8.60
(58.3); head depth 6.80 (46.1); rostral depth 1.35 (9.2); rostral width 2.95 (20.0); dorsal ros-
tral length 0.90 (6.1); ventral rostral length 1.80 (12.2); hindlimb length 4.5 (3.0).

COLORATION (IN PRESERVATIVE): Dorsally and laterally mid-brown. Head slightly darker,
with four narrow pale bands, irregular edged with black: first across head from third
supralabial and rostral margin of orbit; second across head from fifth supralabial (where
most prominent) and caudal margin of orbit; third from cranioventral margin of ear, along
cranial margin of ear and across nape; fourth (very weakly defined) across nape a little way
caudal to ear. Head markings do not extend ventral to supralabials.

Ventrally yellow-blue to light green, more yellow on throat, more blue ventrolaterally
and on tail. A narrow dark blue-grey stripe sharply differentiating lateral from ventral sur-

faces, composed of small flecks on body, coalescing to a distinct stripe on fifth scale row of
caudal half of body and tail.

VARIATION IN PARATYPE: Loreals four; preoculars seven; an additional subocular caudal
to elongate third on right side; upper temporals fused into single scale on right side; first in-
fralabials in contact; second infralabials moderately separated; nuchal scales 12; gular
scales 15.

PROC. LINN. SOC. N.SW., 109 (3), (1986) 1987

G. M. SHEA 205

Fig. I. Head shields of Delma mitella. Paratype (above) in dorsal view, holotype (below) in lateral view; CF = caudal
frontal, CS = caudal supranasals, FN = frontonasals,O = occipital, P = parietals, PF = prefrontals, PN =
postnasals, R = rostral, RF = rostral frontal, RS = rostral supranasals, SO = supraoculars, T = upper
temporals.

Ventral scales 70; hindlimb scales four (left) or three (right); pair of ventral scales
preceding preanals fused into a single v-shaped scale.

Snout-vent length 154; head length 15.50 (10.1); mouth length 13.90 (89.7); snout
length 6.40 (41.3); eye width 2.10 (13.5); postorbital length 4.45 (28.7); head width 11.30
(72.9); rostral depth 1.45 (9.4); rostral width 3.50 (22.6); dorsal rostral length 0.85 (5.5);
ventral rostral length 1.70 (11.0); hindlimb length 8.0 (5.2).

Coloration as for holotype, but fourth light head band almost absent.

PROG. LINN. SOC. N.S.W., 109 (3), (1986) 1987

206 TWO NEW SPECIES OF DELMA

Fig. 2. Holotype of Delma mitella.

COLORATION (IN LIFE): A colour photograph of D. mitella appears on a Queensland
National Parks and Wildlife Service poster, ‘Reptiles of Queensland’. The specimen illus-
trated has body red-brown dorsally and laterally, the lateral margins of the scales slightly
paler, producing an indistinct series of stripes similar to those reported for D. inornata
(Thompson, 1980). Head dorsum mid-brown, slightly darker than body, with four narrow,
irregularly black edged cream bands. Venter yellow on body and throat, but cream

ventrolaterally on lips and neck, becoming blue-white ventrolaterally on body.
Iris black.

COMPARISON WITH OTHER SPECIES: D. mitella is a member of a group of Delma species
possessing two pairs of supranasal scales, first supralabial distinct from rostral supranasal,
fourth supralabial below centre of eye, a mode of 16 midbody scales, enlarged ventral body

PROC. LINN. SOC. N.SW., 109 (3), (1986) 1987

G. M. SHEA 207

scales and three preanal scales. Other members of this group are D. borea, D. fraseri, D. grayit,
D. inornata and D, nasuta.

In addition to the characters given in the diagnosis, D. mitella may be differentiated
from D. borea by having 4-5 loreals (vs usually three), from D. grayzt and D. inornata by the
caudal supranasal contacting or narrowly separated from nostril (vs broadly separated),
from the nearest populations of D. nasuta by its much blunter snout and lack of dark spots
dorsally and ventrally on the body and from D. fraser: by lacking dark markings on the
throat.

HABITS AND HABITATS: The paratype was found dead on road at 1905hrs.

ETYMOLOGY: The specific epithet is from the Latin mitella, a bandage on the head, some-
times worn to’counteract the effect of wine, in allusion to the characteristic head markings
of this species.

Delma labialis sp. nov.

Figs 3,4,5

Holotype: QM J45563, Paluma turnoff on Bruce Hwy, north of Townsville, Qld, in
18°59’S 146°18’E. G. V. Czechura, S. K. Wilson, 13.iv.1985.
Paratype: OM J30265, Magnetic Island, Old. T. Low, viii.1976.

DIAGNOSIS: D. labialis differs from all other Delma species in having a lip and lateral neck
pattern of alternating mid-brown and cream bars, and a dark brown stripe on the third
scale row on caudal body and tail, separating dorsal from lateral surface.

DESCRIPTION OF HOLOTYPE: Rostral with obtuse apex, distinctly penetrating between ros-
tral supranasals; rostral supranasal in moderate contact with first supralabial; caudal
supranasals present, in broad contact with nostril; postnasal single; loreals four, subequal;
preoculars eight (left) or ten (right); suboculars six (left) or four (right), subequal;
supraciliaries five, caudalmost only slightly larger and lying medially to others; supra-
oculars two, first longer, second (right) divided into lateral and medial scale; supralabials
six, fourth below centre of eye, caudalmost subequal to penultimate; infralabials six, first
pair in broad contact on ventral midline, second pair moderately separated; occipital
present; upper temporals two; nuchal scales 16; gular scales 18.

Midbody scales 16; ventral scales 72; ventral body scales transversely enlarged;
preanal scales three; hindlimb scales three.

Snout-vent length 103.5; tail length 408.0 (394); head length 12.10 (11.7); mouth length
9.35 (77.3); snout length 4.95 (40.9); eye width 1.70 (14.0); postorbital length 2.05 (16.9);
head width 6.65 (55.0); head depth 5.60 (46.3); rostral depth 1.20 (9.9); rostral width 2.35
(19.4); dorsal rostral length 0.70 (5.8); ventral rostral length 1.25 (10.3); hindlimb length 4.5
(G23):

COLORATION (IN PRESERVATIVE): Dorsally and laterally mid-brown. Head slightly more
yellow-brown. A series of alternating cream and mid-brown bars on lips and laterally on
neck from caudal margin of orbit to cranial third of body. A narrow dark grey stripe along
centre of third scale row from caudal third of body to proximal half of tail.

Ventrally immaculate cream.

VARIATION IN PARATYPE: The paratype is very desiccated and the scalation difficult to de-
termine in places, but definitely differs from the holotype in having five infralabials on the
right side (six on left); 18 midbody scales and 71 ventral scales.

PROC. LINN. SOC. N.S.W,, 109 (3), (1986) 1987

208 TWO NEW SPECIES OF DELMA

> eae aS
——

Fig. 3. Head shields of holotype of Delma labialis.

Snout-vent length 115.0; tail length 446.0 (388); head length 12.50 (10.9); mouth length
10.70 (85.6); snout length 4.90 (39.2); eye width 1.85 (14.8); postorbital length 2.95 (23.6);
head width 6.80 (54.4); head depth 5.40 (43.2); rostral depth 1.40 (11.2); rostral width 2.55
(20.4); dorsal rostral length 1.15 (9.2); ventral rostral length 1.15 (9.2); hindlimb length 3.5
(3.0).

Coloration as for holotype.

COLORATION (IN LIFE): Kodachrome transparencies of three individuals taken by S. K.
Wilson, A. Dudley and G. Husband permit the following notes on coloration in life.
Adult body dorsum red-brown becoming grey-brown on tail and cranial third of body.
Slightly paler lateral margins to body scales, producing faint indications of stripes of dorsal
ground. Head dorsum yellow-brown. Lip markings yellow-brown and cream. Iris black.
Juvenile body dorsum grey-brown, becoming yellow-brown on head and tail. Lip
markings yellow-brown and pale yellow. Iris black.

COMPARISON WITH OTHER SPECIES: D. labialis is a member of the same group of Delma
species as D. mitella, and may be differentiated from other members of this group by its

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

G. M. SHEA 209

Fig. 5. Details of head of holotype of Delma labialis in life. (Photo: S. Wilson).

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

210 TWO NEW SPECIES OF DELMA

dorsolateral tail stripe and the nature of the head markings. It may be further differentiated
from D. borea by its more numerous loreals (four vs usually three) and infralabials (5-6 vs 4-
6, x = 4.5), from D. grayzi and D. inornata by the broad contact of caudal supranasal with
nostril (vs broadly separated), from the nearest populations of D. nasuta by lacking dark
spots dorsally and ventrally on the body and from D., frasert by lacking dark markings on
throat.

HABITS AND HABITAT: The holotype was taken in low open forest with a grassy understorey,
while the paratype was taken under corrugated iron near a beach. Low (1978) records this
species, as Delma inornata(?), from ‘under sheets of iron in low open forest on sand adjacent
to beaches; one seen active at midday on dry, very open, rocky hill. A. Dudley and G.
Husband (fers. comm.) observed two specimens on Magnetic Island in February, 1985: an
adult active during day in wet sclerophyll forest on the road to Nellie Beach, and a juvenile
in litter in open woodland on the road to Horseshoe Bay.

ETYMOLOGy: The specific epithet is from the Latin /abzum, a lip, in allusion to the distinc-
tive labial and lateral neck pattern of this species.

SYMPATRIC SPECIES: The only Delma species with a distribution overlapping those of D.
mitella and D. labialis is D. tincta (Fig. 6; Shea, 1987), which is readily differentiated from
both species by its smaller size and very different scalation. Kluge (1974) records collecting
alive D. inornata‘a few miles south of Townsville, but does not list this specimen in his speci-
mens examined lists or on the distribution map, nor does he question three specimens
(British Museum (Natural History) 98.10.19.4-8, D. le Souef; examined) from Cooktown.
These localities are respectively 620km and 1050km north of the nearest D. inornata locality
(Marmor, Qld), itself 430km north of the main body of the species’ range, which reaches
Oakey, Qld (Shea, 1987), while more recent collections from both localities have not in-
cluded this species. The Townsville and Cooktown records must be assumed to be in error.

THE STATUS OF ACLYS: Kluge (1976) reduced the monotypic Aclys to a subgenus of Delma,
largely to resolve the discrepancy between relationships as suggested by external morphol-
ogy (Kluge, 1974) and osteology (Kluge, 1976). This arrangement has received little accept-
ance. Aclys has been retained as a genus by Storr et al. (1983), Storr and Harold (1980a,b),
Cogger (1983) and Cogger et al. (1983) without comment.

Examination of the data matrix provided by Kluge (1976) suggests that the dis-
crepancy between data sets is partly a result of a lack of definition for Delma that adequately
demonstrates monophyly. Of the nine skeletal characters keyed as derived for both Aclys and
Delma, one, the presence of two cloacal bones per side, is shared only with Ophzdtocephalus,
while the other eight (characters 11, 19, 20, 49, 59, 65, 67 and 71 of Kluge, 1976) are all
shared with two or more genera each. In contrast, only three of the thirteen phenetic ex-
ternal diagnostic characters (Kluge, 1974) are shared by Aclys and Delma: large external
auditory meatus (primitive, and therefore unsuitable for inferring relationships), smooth
scales (shared with most other pygopodid genera) and preanal pores absent (shared with
Aprasia, Ophidiocephalus and Pletholax). Aclys has two uniquely derived external character
states among the Pygopodidae: upper temporal scales greatly enlarged, forming a second
pair of ‘parietals’ and rostral separating rostral supranasals on dorsal midline. There is little
evidence for a sister group relationship between Aclys and Delma, and pending a more
adequate diagnosis for Delma, the two genera are here considered distinct.

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

G. M. SHEA 211

140° 145° 150° 140° 145° lS OS

152

20°

30°

40°

140° 145° 150° 155° 140° 145° 150° 155%

Fig. 6. Distribution of Delma species in eastern Australia. Left: Delma borea (A), D. plebeca (B), D. mitella (&), D.
labialis (W). Right: D tincta(@), D. torquata(&), D. nasuta (northern form) (¥). Based on specimens examined

in the Australian Museum and Queensland Museum collections. Open symbols are literature records from Kluge
(1974).

KEY TO THE GENUS DELMA IN QUEENSLAND

ee sheanialScalesutwOvee tac tr cere gui ee Mo wae ae teh ey hc Mg 2
ie amalyscalesathireenyee Aen sowase ikea Gate a) Micke” We eter PTT fe ae 3

2. Head grey, throat white, dark collar variably present on nape, lips barred, venter
(CIN ENON Git. “dois aetieacas eiguces (atoae aes Tata amy eat rees SacueE Nt Ma nen a eRe eR a D. plebeia
Head and throat black with narrow cream or white bands, venter dark grey

o US" 6: ond cl aly yd Hig Oatteerel Ale: 3 ren tony | ate TRC a oe Meteo Op aaena aca tet ea et D. torquata

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

22 TWO NEW SPECIES OF DELMA

3. Single pair of supranasals, third supralabial below eye, midbody scales usually 14

Bud cohen ten AUS Cte EY sae oven ge ane A Oe abe ie Sine nth Keith MN nea et D. tincta

Two pairs of supranasals, fourth supralabial below eye, midbody scales usually 16-18

se as RR Ae! Me ae Rae at: pa Bee ete Ge, OA rd cs Mi AG TTR NC ALE Sao oo 4

4. Caudal supranasal broadly separated from nostril; head pattern absent; south-east
Miurcerislamicley Gs see tee grates Nisin a saeco eis By anhe ec eg aden we nay AM eco D. inornata
Caudal supranasal narrowly separated from nostril or in contact; head pattern present
oOmabsent<morthvandwest OQueenslandiy sn 2s sei se ee 5

Sa Narrowsdarkclongitudinalistripeonitalll basey ee 5 saci ir eee 6
INomarrow darkslongitudinalistripeonitallibase py 5; a) eh eee 7

6. Dark longitudinal tail stripe ventrolateral, sharply differentiating light ventral surface
from dark lateral surface; head with faint narrow cream bands dorsally; rostral
barely projecting between rostral supranasals ................... D. mitella

Dark longitudinal tail stripe dorsolateral, not differentiating colour of dorsal and
lateral surfaces; head without cream bands dorsally, but lips with cream bars;
rostral distinctly penetrates between rostal supranasals ........... D. labialis

7. Head usually with dark transverse bands; venter unmarked; usually three loreals; SVL
UU StOr Ouran any Mess earn ete ade sen tee DOU heraol HREN MAYS ees Ullal Coes oe arene D. borea

Head without dark transverse bands; venter usually with darker markings; 4-5 loreals;
S\/auprtos| Orman es ctu ye ti, es ue sie age eit eoun ergata pear eae D. nasuta

ACKNOWLEDGEMENTS

I thank J. Covacevich and A. Greer for laboratory space; S. Wilson, A. Dudley and G.
Husband for the loan of colour transparencies, and E. N. Arnold (BM(NH) ) for the loan
of specimens. H. Cogger, B. Farrow, A. Greer, G. Ingram, M. Peterson and G. Storr
offered useful criticisms of the manuscript. B. Jantulik prepared the final line drawings.

References

CoccGER, H. G., 1983. — Reptiles and Amphibians of Australia. 3rd edition. Sydney: A. H. & A. W. Reed.
——, CAMERON, E. E., and COGGER, H. M., 1983. — Zoological Catalogue of Australia. Vol. 1. Amphibia and
Reptilia. Canberra: Australian Government Publishing Service.
KLUGE, A. G., 1974. — A taxonomic revision of the lizard family Pygopodidae. Misc. Publ. Mus. Zool. Univ.
Michigan No. 147: 1-221.
—., 1976. — Phylogenetic Relationships in the Lizard Family Pygopodidae: An Evaluation of Theory, Methods
and Data. Misc. Publ. Mus. Zool. Univ. Michigan No. 152: 1-72.
Low, T., 1978. — The reptiles of Magnetic Island, Nth Queensland. Herpetofauna 9(2): 10-14.
SHEA, G. M., 1987. — Delma nasuta (Lacertilia: Pygopodidae), an addition to the herpetofauna of New South
Wales and Victoria, with a note on rapid colour change in this species. Vic. Nat. 104(1): 5-8.
STORR, G. M., 1978. — Taxonomic notes on the reptiles of the Shark Bay region, Western Australia. Rec. West.
Aust. Mus. 6(3): 303-318.
——., 1979. — Five new lizards from Western Australia. Rec. West. Aust. Mus. 8(1): 134-142.
——, HANLON, T. M.S., and DUNLOP, J. N., 1983. — Herpetofauna of the Geraldton Region, Western Aus-
tralia. Rec. West. Aust. Mus. 10(3): 215-234.
, and HAROLD, G., 1980a. — Additions to the Herpetofauna of the Shark Bay Region, Western Australia.
West. Aust. Nat. 14(8): 240.
, and , 1980b. — Herpetofauna of the Zuytdorp coast and hinterland, Western Australia. Rec. West.
Aust. Mus. 8(3): 359-375.
THOMPSON, M. B., 1980. — Delma inornata Kluge (Reptilia, Pygopodidae) in South Australia. S: Aust. Nat. 84:
42-43,
WELLS, R. W., and WELLINGTON, C. R., 1985. — A classification of the Amphibia and Reptilia of Australia.
Aust. J. Herp. Suppl. Ser. No. 1: 1-61.

PROC. LINN. SOC. N.S.W., 109 (3), (1986) 1987

ey
We

PROCEEDINGS
of the

LINNEAN
SOCIETY

NEW SOUTH WALES

VOLUME 109
NUMBER 4

PRESIDENTIAL ADDRESS

Cainozoic History of the Vegetation and Climate
of the Lachlan River Region, New South Wales

HELENE A. MARTIN
University of New South Wales
Box 1, RO. Kensington
New South Wales 2033

(Delivered 31 March, 1982)*

SYNOPSIS

The history of the vegetation based on palynology of the late Eocene to early Pleisto-
cene sediments is presented here for the Lachlan River region. Grey carbonaceous clays
of the alluvium in the valley and in the Hillston region of the eastern edge of the Murray
Basin, are the most fruitful for palynology. Pollen has not been preserved upstream of
Cowra but silicified wood is found in association with basalts. Early Miocene lava flows
dammed the river and the lake which was formed drowned the local vegetation (Bishop
1985a).

The palynological record and interpretations of the vegetation is one of periods of
relative stability with small changes and a series of considerable, step-like change. The
climate of the time is deduced from the parameters controlling generally similar modern
vegetation. The main features are as follows:

1. From late Eocene to late Oligocene — early Miocene, Nothofagus is the most abun-

dant pollen group and most of it is the bvassi type. The vegetation was rainforest with

reasonable diversity. The climate was very humid with a precipitation of about or
above 1800mm. (The levels of precipitation given here are very general with no great
accuracy: the trend is more important.)

2. Inthe late Oligocene — early Miocene, the Nothofagus content declines, particu-

larly the brassi type. The vegetation was still rainforest. There was a decrease in

precipitation, probably to about 1500mm.

3. Inthe early—mid Miocene, the Myrtaceae group was relatively more abundant.

The assemblages are diverse, however, and contain many low frequency pollen types,

which collectively may have accounted for a major portion of the vegetation, which

was rainforest.

4. The mid Miocene was a time of considerable change. The brassz type of Notho-

fagus disappeared and pollen preservation ceases in the Hillston region.

5. Inthe ?mid—late Miocene, Myrtaceae were dominant but rainforest taxa were

still present. The charcoal record, which had been low in the older, rainforest assem-

blages, increased considerably, suggesting that the myrtaceous vegetation was mainly
wet sclerophyll. The precipitation decreased to about 1000-1500mm, with a definite
dry season. Burning had become a part of the environment.

6. In the early Pliocene, Nothofagus, the menziesii and fusca pollen types only, re-

appeared in the Lachlan Valley and gymnosperms were more abundant. Rainforest

had returned, the precipitation increased to more than 1500mm and burning was
infrequent.

7. Inthe mid—late Pliocene, Myrtaceae returned to dominance, precipitation

decreased to the former levels of about 1000-1500mm and burning became more

frequent.

8. Inthe late Pliocene — Pleistocene, the rainforest element disappeared and the

precipitation decreased to about 500-800mm.

9. The forest cover dwindled and in the Pleistocene Gramineae and Compositae

were abundant; indicative of woodland and grassland/herbfields.

These major changes in vegetation and the inferred climatic changes may be related
to changes in sea level and coincide with the major developments of circum-Antarctic
oceanic circulation and ice cap formation on Antarctica.

INTRODUCTION

The Lachlan River has its headwaters in the gently undulating, swampy Breadalbane
Plains at the continental drainage divide of the Central Tablelands. It flows through a

* Copy received for printing 17 December 1986.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

214 PRESIDENTIAL ADDRESS

broad upland valley around Gunning and Dalton. Downstream, the terrain increases in
ruggedness and, at Narrawa, the river flows through a steep-sided granite gorge. Alluvial
flats of significance commence about 13km upstream of Cowra. Valley width increases with
distance downstream and the alluvial flats become extensive. After the river passes through
the gap between the Jemalong and Corridgery Ranges it enters extensive plains. About the
junction with Willandra Billabong, it flows onto the Western Plains, the surface of the
Murray Basin, and eventually joins the Murrumbidgee River (see Fig. 1). For a delightful
pictorial account of the Lachlan River and the lifestyles of the people living in the region,
see Cowan and Beard (1982).

1 HILLSTON REGION, see FIG 5
2 LACHLAN VALLEY, see FIG 2
3 UPPER LACHLAN RIVER, see FIG 3

-——— Edge of Basin
-. Limits of Marine Transgression

Fig. 1. Locality map. See enlargements for further detail.

In 1957, the Department of Water Resources (formerly the Water Resources Com-
mission of New South Wales, and in 1957, the Water Conservation and Irrigation Com- —
mission) began a programme of the investigation into the ground water resources of
alluvium of the Lachlan River valley, between Cowra and Forbes. Prior to this date, most
bores and wells were sunk for stock water and domestic purposes and did not exceed 30m
in depth. There were only a few low-yielding irrigation bores. ‘Test drilling in the valley soon
revealed good quality water at greater depths with much higher yields, suitable for irriga-
tion and town water supplies (Williamson, 1986). At the time of writing, there are 174 high-
yield bores between Cowra and Jemalong Weir, and of these 166 are used for irrigation and
8 for town water supplies. Investigation has continued downstream of Jemalong Weir and
beyond Hillston.

This study of the history of the Lachlan River is based mainly on palynology, geo-
morphology, and geology. Water Resources has provided samples from its bores for paly-
nology and considerable background information as well. A study of the basalts in
headwaters of the Lachlan River (Bishop, 1985a) provides a history of the upper reaches of
the river, beyond the extent of the alluvial flats. In all, three areas have been studied in
detail and these are shown on Fig. 1.

(GEOLOGY AND GEOMORPHOLOGY

The Hillston Region

The Tertiary sediments are some 100m to 170m thick near the edge of the Murray
Basin. Thickness increases downstream, i.e. with distance from the edge of the basin to a
maximum of about 270m in bore 36342 which is situated over the Ivanhoe Trough. When

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

H. A. MARTIN 215

compared with the Murrumbidgee area to the south, the basement of most of the Lachlan
region is shallower and the sedimentary sequence not as thick (Martin, 1984c).

The stratigraphic units used in the non-marine section of the Murray Basin have been
adopted from those used in the Victorian part of the basin, with slight modification
(Woolley and Williams, 1978; Woolley, 1978). Those relevant to the Lachlan area have been
reviewed in Martin, (1984b; 1986a) and are briefly presented below.

The basal Warina Sand of mid —late Eocene age consists of coarse-grained quartz
sands and minor dark grey clay lenses and carbonaceous clays. It is only found in the
deeper parts of the basin.

The overlying Olney Formation is dominated by grey carbonacous clays and extensive
sands. Elsewhere in the basin, thick lignite layers are a feature of the Olney Formation but
they are not extensive in the Lachlan area. The basal part is late Eocene and extends into
the Miocene.

The Calivil Sand, overlying the Olney Formation, consists of coarse sands and fine
gravels with minor bands of carbonacous clay. It is thought to be late Miocene in age. The
uppermost Shepparton Formation consists of polymict sands and variegated clays with
yellow and brown colours dominant.

Most of the samples used for palynology come from the Olney Formation. The upper-
most 60m to 80m do not yield pollen. Upstream from Euabalong to Jemalong Gap, there
are few bores. Knowledge of the age of the sediments is reliant on palynology which is
presented later.

Lachlan River Valley

Near Cowra, the alluvium is 3-5km wide and erosion terraces are a prominent
feature. They become less pronounced further downstream and are not mappable beyond
the junction of Mandagery Creek. Drilling has revealed that the alluvium reaches a maxi-
mum depth of 61m, some 6km downstream of Cowra. However, 3km upstream of Cowra,
the alluvium is only 17m deep. Back Creek, which joins the Lachlan River some 12km
downstream of Cowra, has alluvium extending some 48km upstream. The alluvium in-
creases in depth with distance downstream and the maximum depth in Section 7, south of
Forbes, is 133m. (See Fig. 2 for localities).

Drilling of the valley downstream of Cowra has revealed a buried ‘valley-in-valley’
structure. Remnants of an old valley floor are shown as a shelf which maintains a depth of
27-30m below the present drainage level, and it is present in both Back Creek and the
Lachlan River valleys. However, the depth of the valley carved into the floor of the old
valley, increases markedly with distance downstream (Williamson, 1964). The valley-in-
valley structure is thought by Williamson (1986) to be the result of successive tectonic move-
ments, but this is discussed further, below.

‘Tectonic movements near Cowra have caused a marked change in the upstream sec-
tion of the Lachlan River but they have not affected Back Creek. Evidence from seismic
refraction and bore data indicate that the margin of the area affected is 3km downstream
of Cowra with a north-south trend. This margin is thought to be the western edge of the up-
lifted Eastern Highlands (Williamson, 1986). The valley-in-valley structure is shown with
the palynology in Fig. 16B.

The alluvium of the Lachlan Valley consists of two quite distinct formations, the basal
Lachlan Formation and overlying Cowra Formation. The characteristics of these for-
mations have been described by Williamson (1986) and are summarized here.

The Lachlan Formation consists of a series of interbedded and interlensed sediments
ranging from gravels to clays. The sands and gravels consist almost entirely of quartz of
different kinds and sometimes pebbles of chert. The most significant feature of the sands

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

216 PRESIDENTIAL ADDRESS

FORBES

: :36079

e! px of
JEMALONG GA 12383
ix

eo
O A
‘3 730484 i ~~
:
Big 10/14913 2Ch/.

914 (9) .

ive,
8 21029

2
%G

Fig. 2. Lachlan River valley locality map.

and gravels is that they do not contain the resistant rock types found in the catchment to-
day. The fact that the sands and gravels of the Lachlan Formation consist of almost entirely
quartz is important for ground water. Quartz is stable and water will become less minera-
lized in its passage through aquifers of the Lachlan Formation than through those of the
Cowra Formation (Williamson, 1964).

The clays of the Lachlan Formation may be divided into variegated clays and car-
bonaceous clays. The former are usually thinly bedded and streaky. The most common
colours are pale brown, yellow, grey and off-white but pink and even red are also fairly com-
mon. The carbonaceous clays are the best material for palynology. They are grey to black
in colour and occur in lenses irregularly distributed through the Lachlan Formation. These
lenses are usually limited in extent and often less than 1m thick though sometimes they may
be correlated at the same horizon between two or three bores. These lenses may be 6m in
thickness, and range up to 12m.

Wood is occasionally encountered in the sands and gravels. Podocarpus sp. has been
identified, but most samples are too carbonized for identification (Williamson, 1986).

The Cowra Formation disconformably overlies the Lachlan Formation. The strata
range from gravels to clays, all of which are predominantly brown, sometimes pale brown,
red-brown, or yellow brown and rarely grey. The sands and gravels contain representatives
of the various resistant rock types present in the catchment area today. In this respect, they
differ significantly from the Lachlan Formation. The associated silts and clays in the Cowra
Formation are predominantly brown and occasionally pale grey in thin layers. There may
be greyish mottling near the surface, probably due to leaching. Carbonaceous clays are
extremely rare.

Wood was encountered only once in the Cowra Formation in gravels in Bore 12437,
Section 4, at a depth of 26m. It has been identified as probably Eucalyptus resinifera (R. K.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

H. A. MARTIN 217

Bamber, fers. comm.), but was beyond the range of carbon dating Le. older than 34,000 years
(Williamson, 1986).

There is another formation, the Glen Logan Gravels, consisting of medium to coarse
quartz gravel, usually in a red-brown silty matrix. They occur in elevated positions and
often form hill cappings and are worked for road materials. It is thought that they are rem-
nants of a formerly more extensive formation which is stratigraphically below the Lachlan
Formation. Williamson (1986) postulates that the Glen Logan Gravels were probably the
mayor source of the quartz gravels in the Lachlan Formation, but this is discussed further,
below.

The test drilling in the valley has revealed facets of river history. See Fig. 2 for lo-
calities. Back Creek now joins the Lachlan River 8km downstream of Section 2 but its
former junction was at Section 2. At Section 4, the ancient course of the Lachlan River
went through the southern part of Section 4A instead of its present course, some 6km to the
south. About Section 7, the ancient river turned south-west and passed under Section 8,
some 15km south of its present course near Forbes. The ancient river passed through the
gap between the Jemalong and Corridgery Ranges, as does the present river, for this is the
only feasible gap (Williamson, 1986).

MIOCENE
BASALTS

K-Ar AGE

CROOKWELL
@

Creek

SEE RIG

Fig. 3. The Miocene basalts of the Upper Lachlan. Modified from Bishop (1985a).

The upper Lachlan River

Near the headwaters of the Lachlan River, early Miocene channels have been
preserved by long narrow tongues of basalt which were extruded into valleys and some

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

218 PRESIDENTIAL ADDRESS

tributaries (see Fig. 3). These basalt remnants occur as hilltop residuals as a result of relief
inversion by erosion of the surrounding rock. These remnants have flat tops and are of a
fairly uniform elevation, as would be expected from lava flows which did not completely fill
the valleys (Bishop, Young and McDougall, 1985). The basalt remnants show that the
ancient river ran parallel to the present course of the Lachlan and on the eastern side of
Narrawa Mountain (Bishop, 1985a; 1987).

The lava flowed westwards down a tributary into the main river, damming the chan-
nel. It then flowed northwards for some 15km down the temporary dry river bed. A large
lake, thought to be about 16,500 km? in area was formed (Fig. 4), presumably drowning

NuY
aaah GUNNING

LAKE SHORELINE

| MIOCENE BASALTS

Fig. 4. Probable extent of the Miocene lake formed after the ancient Lachlan River had been damned by lava flows.
Modified from Bishop (1985a).

the vegetation. Continuing lava flows engulfed the trees and preserved the wood. At least
one tree stump has been preserved, apparently zm situ and fragments of silicified wood are
common in the area today (Bishop, 1985a). The stump has been identified as Myrtaceae,
with affinities to Eucalyptus B. Acacia and Nothofagus have been identified amongst other
wood fragments (Bishop and Bamber, 1985).

It is not clear how long the lake persisted or what were the events associated with
breaching the lava dam. It is thought that the present course of the Lachlan, in the gorge

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

H. A. MARTIN 219

west of Narrawa Mountain, may be attributed to the catastrophic events associated with
breaching the dam and draining the lake (Bishop, 1985a).

Pollen has not been preserved in the sediments of these upper reaches of the Lachlan
River (Bishop, 1985a).

PALYNOLOGY

Late Eocene to mid Miocene palynofloras

The palynological zones described for the Gippsland Basin (Stover and Partridge,
1973, 1982; Partridge, 1976) are the most appropriate for this study. The zones are identi-
fied by the presence of diagnostic species in the assemblages and have been dated from in-
dependent evidence of the foraminifera found in the sequence. Extrapolation of the zones
in the Gippsland Basin to the Murray Basin required some minor modifications, par-
ticularly for subdivision of thick sections of the Oligocene — early Miocene P. tuberculatus
Zone. These modifications utilize the variation in abundance of selected species and the
method is presented in Martin (1984a). Fig.5 presents the palynological zones applicable
here.

Million PALYNOLOGICAL

Years
ZONES

0
Pleistocene Floras

SEE FIG 14

MIOCENE T. bellus
Zone

C subdivision
20

P. tuberculatus
Zone

30} OLIGOCENE B subdivision

EARLY

A subdivision

LATE

MIDDLE N. asperus
EOCENE Zone

Fig. 5. Palynological zones applicable to the Hillston region. From Stover and Partridge (1973, 1982); Partridge
(1976), with the modifications of Martin (1984a).

PROG. LINN. SOC. N.S.W., 109 (4), (1986) 1987

220 PRESIDENTIAL ADDRESS

s]
\Ta, A 30415 B’
Wi \ 70, :
Mande. 5 Nea, 30414 36171@

Euabalong

@30410
———@—_ Ot08

30407
@ 30406

HILLSTON \\ \

36342

Fig. 6. Hillston region showing location of bores and sections.

Two sections in the Hillston region (see Figs 6-8) show the distribution of the paly-
nological zones. The oldest zone present is the late Eocene Middle N. asperus Zone and it
is restricted to the Murray Basin. Pollen preservation ceases in the early Miocene and the
upper or C subdivision of the P. tuberculatus Zone is the youngest in the Murray Basin. Up-
stream of the Lachlan River, beyond the eastern edge of the basin, the younger mid
Miocene T° bellus Zone and Pliocene sequence may be present (see Fig. 8).

Fig. 9 presents the counts of the pollen groups in Bore 36342, Tom’s Lake, the deepest
of this study. For a definition of the groups, see Appendix 1. There is relatively little change
between the pollen groups of the Middle N. asperus Zone and the A and B subdivisions of
the P. tuberculatus Zone The C subdivision shows a considerable decrease in Nothofagus, par-
ticularly the bvassa type, and a slight increase in Myrtaceae. (The importance of this change
is discussed further below.) Besides these abundant pollen groups, there are a number of
low percentage angiosperms listed in Appendix 2. There are others which have not been
named and for which botanical affinities are unknown, particularly tricolpate and tri-
colporate pollen types. On the whole, these assemblages are diverse.

The subdivision of the P. tuberculatus Zone into three parts (see Fig. 9) is only possible
in the deep bores of the Lachlan region. Elsewhere, the A and B subdivisions cannot be
identified. The C subdivision, however, is present over the whole area, as shown in Figs 7
and 8 (see also Martin, 1984c). In this respect, the Lachlan area of the Murray Basin differs
from the Murrumbidgee area where the three subdivisions are recognized over the whole
area (Martin, 1984b).

The Middle WN. asperus Zone is distinctive with a variety of proteaceous type pollen.
There may be up to nine species of Proteacidites which may account for 10% of the total
pollen count although 5% is more common. A number of these species, mainly the larger
pollen types, do not extend into the younger zone above. The diversity and abundance of
the proteaceous type pollen decreases in the subsequent, younger assemblages. As well as
Proteacidites spp. there are a number of distinctive angiosperm pollen types which are not
found in younger zones (see Appendix 2).

Nothofagus is the most abundant group in the P. tuberculatus Zone. The brass type
usually accounts for most of this group but some assemblages have a high proportion of N.

flemingu of the fusca pollen type (see Fig. 9). High N. flemingit assemblages may be found

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

H. A. MARTIN 221

36304
30406
30407
30409
30410
30411
30414
30415

36342

ae ee
aca pearl dine SBA CLEVER Shey socal sla Tap are ee Hill a
q oS x
x Ei
4 a
Ses
x
ae
AL No pollen recovered
m is hue
' ves C subdivision, P. tuberculatus Zone
High N. flemingii assemblages
20
B subdivision, P. tuberculatus Zone
10 20 30 40 50
wu It ! \ 1 1 km

5 A subdivision, P. tuberculatus Zone
Approximate scales

Middle N. asperus Zone

Early Cretaceous

Fig. 7. The palynological zones in Section A-A! of the Hillston region.

anywhere in the P. tuberculatus Zone of the Lachlan area (see Fig. 7) of the Murray Basin,
unlike the Murrumbidgee area to the south where such assemblages are restricted to two,
well-defined layers (Martin, 1984b; 1986a; 1986b).

In the C subdivision of the P. tuberculatus Zone, the Nothofagus content declines,
especially the brass type. This decline is moderate in the Lachlan region, as shown in Fig.
9, but in the western part of the basin, the decline is much greater (Truswell et al., 1985).
The Myrtaceae group increases slightly ‘here (Fig. 9), but it becomes the major pollen
group with the decline in abundance of Nothofagus (Martin, 1986a; 1986b). As well, there
is an increase in diversity and abundance of the tricolpate — tricolporate pollen types in
this subdivision. The proteaceous pollen content may increase somewhat, but the pollen
types are relatively small, more like Helicia — Ontes, hence distinct from the late Eocene
proteaceous content.

There are only a few of the 7’ bellus Zone assemblages and they are found upstream
from the edge of the Murray Basin. The identification of this zone relies on several diag-
nostic species, but quantitatively, the assemblages are little different from those of the upper
part of the P tuberculatus Zone. Figs 10 and 11 present the counts of some 7! bellus Zone
assemblages and Appendix 2 shows the low percentage angiosperms. The seemingly poor

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

PRESIDENTIAL ADDRESS

sajyeos ajewlxoiddy INSW3SSVd
wy
@UOZ snsedse ‘N app Hl es DY Oe Oke
au0o pee
Z snjejnosaqn] “gd ‘UOISIAIpqns v is snoeoe1el9
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sebejqwesse mwhuswoajj “N UBIH w ie
0 Nia
8Uu0Z snjejnosaqgn} “g ‘UOISIAIPgns g Es
8UOZ snjejnosaqn} ‘*g ‘UOISIAIPGns 9
euoZ snjjeq ‘1 TEIAEH WES: |be39| = = = ea ae
eouenbes auadolidq
pdjaA00e1 UAajOod ON ——
5 |: 5
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| |
| |
| 7 LZE9E
|
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peeee B80rSe|
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§. The palynological zones in Section B-B! of the Hillston region.

G
2. O
5

Cc
J

109 (4), (1986) 19

N.S.W.,

PROC. LINN. SOC.

H. A. MARTIN DDS,

——____—___——_
40 % of Total Count ZONE

a C subdivision
b f jm

ie)
Oo

pepth SPORE POLLEN COUNTS 0°

Tas
mae es), le

aa
Ml

el h| P, tuberculatus
ee

B subdivision

A subdivision
Middle WN. asperus :

;

i?)

pores Gymnosperms Myrtaceae Nothofagus
Casuarinaceae

Fig. 9. The abundant pollen groups (see Appendix 1) for bore 36342 of the Hillston region. C. Cyathea, the left hand
portion of the spore count. b, brassz type, f, fusca type and m, menziesii type from left to right, respectively. Those
counts of Nothofagus showing only two divisions are the brassz and fusca types only. Low percentage taxa are not
included (see Appendix 2).

representation of pollen types in Appendix 2, in comparison with the older zones 1s a result
of many fewer of T° bellus zone assemblages which are, however, just as diverse as the upper
part of the P. tuberculatus Zone. Unnamed and unknown tricolpate — tricolporate pollen
types are a feature here.

The Relationship of the T: bellus Zone to the ‘Pliocene’ Sequence

The T° bellus Zone and a good Pliocene sequence are found in a bore at Jemalong Gap
(Fig. 10). Both also occur in a bore at Euabalong (Fig. 11) but the Pliocene sequence there
is not as extensive as that at Jemalong Gap. Prior to this study, only one other bore, at
Narrandera (Martin, 1984b), had been found to contain both, hence a detailed comparison
is warranted.

At Jemalong Gap, the 7’ bellus Zone contains two diagnostic species Triporopollenites
bellus and Symplocozpollenites austellus (Stover and Partridge, 1973). The 7. bellus Zone at
Euabalong has only the latter. S. austellus occurs in the Myrtaceae phase immediately above
the 7’ bellus Zone in both bores (and elsewhere, as discussed further below). The assign-
ment of these assemblages to the Myrtaceae phase is based on an increase in Myrtaceae
and minimal Nothofagus, especially the brassii group. The small amount of this pollen type
in the basal two assemblages of the lower Myrtaceae phase at Jemalong Gap, 2% or less
(see Fig. 10), could easily result from reworking of older assemblages.

Stephanocolpites oblatus, commonly present in the Pliocene sequence, is found in the 7
bellus Zone of both bores. It is found occasionally in older zones as well (see Appendix 2).
A number of distinctive early Tertiary angiosperms which range into the 7’ bellus Zone, e.g.
Malvacipollis subtilis, Triporopollenites endobalteus and Polyorificites oblatus, as well as the
nominate species of the zone, 7’ bellus, are usually lacking from the Myrtaceae phase (see
Appendices 2 and 3) but, given the variability of the latter (discussed further below), this
may not be entirely reliable. Thus the main feature used to differentiate the Myrtaceae

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

e ”
224 a" = © 3| ©
oF ° 3 s| 3
< Ss a) | BS] o
a ‘oO ~ ws ~ 3
Bus = i ES = =
{aa he
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Oo ° = J 3
N ° 5 £ °
= > 4
a Oo
©
8 c
5 €
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[o) 2)
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=
iS)
S 2

@
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®
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o
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(dp) =
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o
o £
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x
N
¢ =
o
1S) L,
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£ 4 5
ge -1] - B
Le}
Spe iOpen Ore Me Met iE) Sian

Fig. 10. The abundant pollen groups (see Appendix 1) at Jemalong Gap, bore 36079. C, Cyathea. b, brassi type.
m, menziesii type. Low percentage taxa are not included (see Appendices 2 and 3),

PROC. LINN. SOC. N.S.W,, 109 (4), (1986) 1987

H. A. MARTIN DDS

Depth SPORE POLLEN COUNTS 0 zo 40% of Total Count ZONE OR PHASE
i a
—— SSE iy
el ‘a Myrtaceae
f
ps p r
mb
ireect 1 i. le bellus
Spores Gymnosperms Casuarinaceae Myrtaceae Nothofagus Compositae

Gramineae

Fug. 11. The abundant pollen groups (see Appendix 1) at Euabalong, bore 36171. C, Cyathea. b, brassti type. f, fusca
type. m, menziesii type. Low percentage taxa are not included (see Appendices 1 and 2).

phase from the 7’ bellus Zone is the low proportion of brassiz type Nothofagus.

The upper part of the P. tuberculatus Zone may have a high content of Myrtaceae (dis-
cussed previously). However, near the eastern edge of the Murray Basin, it also has
appreciably more than a few percent of the brass type of Nothofagus as well as more of the
typical early Tertiary angiosperms, hence is unlikely to be confused with the Myrtaceae
phase. Near the western edge of the basin, the upper part of the P. tuberculatus Zone has
more Myrtaceae and very little of the brassi type (Truswell et al., 1985; Martin, 1986a) but
a Pliocene sequence has not been identified here.

The ‘Pliocene’ Palynofloras

The Pliocene sequence 1s described in Martin (1973b) and has been divided into in-
formal ‘phases’. This description was based almost entirely on one bore, 14747 of Section
6, in which the average of all the assemblages for the phase was presented. With subsequent
work, a total of some forty bores have been examined from the Lachlan River valley, hence
the Pliocene sequence is re-described here with an assessment of the variability. Bore 14747,
which shows the best sequence in the Lachlan River valley, is presented here again (Fig. 12)
to show the variability found in the phases. This present report does not contradict or make
substantial alterations to the original descriptions.

Myrtaceae phase

Myrtaceae is the abundant pollen group and usually constitutes 30-40% of the total
pollen count, but some assemblages may have as much as 70%. Spore content is moderate,
usually 30% or less. The content of gymnosperm pollen is moderate to low, less than 20%
and Podocarpus is usually the most common in this group. Nothofagus is sometimes present
but only in small amounts, 5% or less. Compositae and Gramineae are usually present in
low quantities, 5% or less. Cyperaceae and Restionaceae are occasionally present, also in
low quantities of 5% or less.

Some taxa are only recorded in low percentages that show little variation. These taxa
usually present include Haloragis and Proteaceae (excluding Banksieae) whereas Tasmannia
and Micrantheum are sometimes present. Infrequent occurrences include Anthocerotae,
Acacia, Dodonaea, Banksieae Epacridaceae (tetrad pollen type), Symplocos, Quintinia and
Myriophyllum whereas Monotoca, Coelebogyne, Gyrostemonaceae, Sparganium, Macaranga-
Mallotus and and Goodeniaceae are rare. Unidentifiable tricolpate/tricolporate angiosperm
grains usually account for 10% or less of the total pollen count.

The original description of the Myrtaceae phase included Casuarina, i.e. the
Myrtaceae — Casuarina phase. High percentages of Casuarinaceae are somewhat erratic
and are discussed further, below.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

226 PRESIDENTIAL ADDRESS

PHASE
Upper Myrtaceae
Gymnosperm

Nothofagus
Lower Myrtaceae

Co)

E %

oO =

oO o

vai (eo)

& Q

2 5

5 oO

- 7)

‘ 3

° xs s

SS =

is)

=
°
AN

rs)

° o

Co)

is0j

2 5

> 7)

5 Oo
oO

rr)

= =

4 ®

Ss j

a e

fr =

o oO
(7p)

ce) ©

£ of: a

Awe aga ie

oO oO
+ © 2

Fig. 12. The abundant pollen groups (see Appendix 1) of bore 14747, Section 6, Lachlan River valley. C, Cyathea.
1, fusca type. m, menziesu type. Low percentage taxa are not included (see Appendix 3).

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

H. A. MARTIN 227

Nothofagus phase

Nothofagus is relatively abundant, over 10% with the highest value of 29%. The fusca
pollen type, Nothofagus brachyspinulosa, is the most common with smaller amounts of the men-
ziesu type, Nothofagus aspera. The brassit pollen type which was abundant in the early and mid
‘Tertiary is present as very minor quantities (1-5%), and in this respect the Nothofagus con-
tent of these Pliocene assemblages is quite distinct from that of the older, Nothofagus-
dominated assemblages.

Spore content is moderate (20-30%) with a slightly greater diversity than that of the
Myrtaceae phase. The gymnosperm content 1s usually less than 20% and the composition
of the group is much the same as that for the gymnosperm phase (discussed below).
However, the single occurrence of Lagerostrobus franklinu in the Pliocene is found in the
Nothofagus phase. The Myrtaceae content is relatively low, less than 20%. The low percen-
tage taxa register in much the same way as that described above for the Myrtaceae phase.

Depth
a SPORE POLLEN COUNTS 0 20 40 % of Total Count
m eee eee
60 F
Section 3, Bore 12423 m f
Section 6, Bore 14747 if
¥ f
Section 8
Bore 21019 B 14707
ae b i ore mf
— m
Jemalong Gap,\Bore 36079 Casuarinaceae Myrtaceae Nothofagus | Gramineae
Compositae

|
Spores Gymnosperms

Fig. 13. The Nothofagus phase in the Lachlan River valley, bores arranged sequentially downstream. Jemalong
Gap does not meet the definition of the Nothofagus phase but it is in the stratigraphic position of this phase and
included for comparison. C. Cyathea. m, menziesii type. f, fusca type. Low percentage taxa are not included (see
Appendix 3).

Fig. 13 shows the pollen spectra of the Nothofagus_phase, sequentially downstream. The
percentage of Nothofagus decreases with distance, downstream. Jemalong Gap, with 4.5%
does not meet the definition of the Nothofagus phase, but there is a peak and it occurs at the
expected stratigraphic level of the Nothofagus phase (discussed further below). This low per-
centage could well result from long distance transport, which, however, would coincide with
the Nothofagus phase further upstream. Jemalong Gap 1s included in Fig. 13 for comparison.

The Gymnosperm phase

Gymnosperm pollen exceeds 20% of the total count. Podocarpus is usually the most
abundant of the group, but other taxa may be more abundant, e.g. Cupressaceae,

PROC. LINN. SOC. N.S.W,, 109 (4), (1986) 1987

228 PRESIDENTIAL ADDRESS

Dacrycarpus or Phyllocladus. Dacrydium and Araucariaceae are usually present but the latter
does not exceed 6%. Microcachrys is occasionally present in very low frequencies.

Spores, Myrtaceae and Casuarinaceae are all moderate and values rarely exceed 20%.
Occurrence of the low percentage taxa are much the same as that for the Myrtaceae phase.

Fig. 14. The gymnosperm phase in the Lachlan River valley, bores arranged sequentially downstream. C, Cyathea.
The most abundant gymnosperm of the group is shown thus: K, Cupressaceae, P, Podocarpus. Ph, Phyllocladus.
D, Dacrycarpus. A, Araucariaceae. Low percentage taxa are not included (see Appendix 3).

Fig. 14 shows the pollen spectra of the Gymnosperm phase, arranged sequentially
downstream. The best development is upstream at Section 4, but the decline downstream
is not as marked as that for the Nothofagus phase. One Nothofagus-phase assemblage at Sec-
tion B also meets the definition of the gymnosperm phase. At Jemalong Gap, the highest
percentage of gymnosperms is slightly less than that in the definition, but this assemblage
also contains the highest Nothofagus value of the Pliocene sequence in this bore , thus these
two high values occur in the expected stratigraphic position (discussed further below). The
Jemalong Gap high gymnosperm assemblage 1s included in Fig. 14 for comparison.

High Casuarinaceae Content

A more specific identification of Casuarinaceae pollen is difficult for there is continu-
ous variation through Gymnostoma and Casuarina (Kershaw, 1970a). Some assemblages have
a high content of Casuarinaceae, 50% or more of the total pollen count. These assemblages
are otherwise typically those of the Myrtaceae phase. The distribution of the high percen-

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

H. A. MARTIN 229

tages of Casuarinaceae do not show any clear patterns (discussed further below), hence this
feature is not included in the definition of the phases.

High spore content

Some assemblages have an exceptionally high spore content (50-70%). Cyathea is
usually the most abundant taxon throughout the Pliocene and it may be exceptionally high
in these assemblages. There is usually a greater diversity of spores and most of them are
ferns. Because of the high spore content, the percentages of other groups are depressed and
these assemblages may not fit into any of the phases described above. Some of them,
however, have an appreciable Myrtaceae content and could be included in the Myrtaceae
phase.

Stratigraphic Relationships of the ‘Pliocene’ Sequence

As originally described (Martin, 1973b), the lower Myrtaceae phase 1s the oldest of the
sequence and forms a relatively thin part of the sequence over the valley basement. This 1s
overlain by the Nothofagus and then the gymnosperm phases, both relatively thin. Above
these, the upper Myrtaceae phase forms the uppermost part of the Pliocene sequence. Thus
the Nothofagus and gymnosperm phases divide the Myrtaceae phase into a lower and upper
component, with no means of distinguishing the two from the composition of the assem-
blages (see Appendix 3). The upper Myrtaceae phase accounts for the largest part of the
section and is usually one half to one third of the total Pliocene sequence. Fig. 15 presents
the relationship of the phases.

The distribution of the Pliocene phases in the sections across the valley is shown in Fig.
16 and Fig. 17 shows C-C! section along the length of the valley (see Fig. 2 for localities.).
It can be seen that the whole sequence increases in depth with distance downstream.

The Nothofagus phase has been considered as a marker horizon within the Pliocene.
The evidence from the Lachlan River valley to support this hypothesis is rather sparse as
only five Nothofagus phase assemblages have been recovered. There is no evidence to the
contrary, either. In a broad sense, the Nothofagus phase is still considered a satisfactory
marker horizon and the palaeoecological reasons for this are discussed later. However, if
levels between bores are extrapolated, then a high degree of precision should not be ex-
pected for these are largely fluviatile sediments and cut and fill associated with the changes
in the river system may produce a complex stratigraphy. This is probably the reason for the
Nothofagus phase in Bore 14745 being some 12m below that in Bore 14747, which is only
0.7km distant. See section 6 in Fig. 16E.

The gymnosperm phase occurs above the Nothofagus phase in only one bore. All of the
other gymnosperm phase assemblages occur in bores which do not have the Nothofagus
phase or else coincide with the latter. In the upstream part of the valley, both the Nothofagus
and gymnosperm phases occur close to the floor of the valley (see sections 3 and 4 in Fig.
16). Thus there is doubt whether the gymnosperm phase always occurs stratigraphically
above the Nothofagus phase. Given the distribution of the two phases, an equally valid
hypothesis would be a contemporary mosaic of the two kinds of vegetation (discussed
further under palaeovegetation). Thus it may be more prudent to consider the Nothofagus
and gymnosperm phases as contemporaneous events.

Stratigraphically, the high spore assemblages occur roughly about the same levels or
below those of the Nothofagus and gymnosperm phases (see Fig. 18). Curiously, most of the
high Casuarinaceae assemblages are found about these levels also (discussed under
palaeovegetation).

PROG. LINN. SOC. N.S.W,, 109 (4), (1986) 1987

230 PRESIDENTIAL ADDRESS

PALYNOLOGICAL
SUBDIVISION

Compositae/

PLEISTOCENE Gramineae

Upper Myrtaceae phase

PLIOCENE

Gymnosperm phase

Nothofagus phase

Lower Myrtaceae phase

MIOCENE

Fig. 15. The relationship of the phases in the ?mid Miocene-Pliocene sequence.

Towards the top of the upper Myrtaceae phase, diversity is reduced and the content
of Compositae and Gramineae increases. It may be difficult to distinguish the uppermost
Pliocene from Pleistocene assemblages (discussed further below).

Dating the ‘Pliocene’ Sequence

The lower Myrtaceae phase overlies the 7? bellus Zone, as discussed previously, but
dating the boundary is problematical.

In Section 6A near Eugowra on Mandagery Creek, basalt has been intersected in the
bores. There is 9.4m of alluvium below the basalt (Williamson, 1986) but pollen was not
recovered from these sediments. The texture, mineralogical and chemical composition of
the basalt in the bores is sufficiently similar to the basalt outcrop at Toogong, some 21km
upstream, to allow a common source of both the basalts (Williamson, 1986). The Toogong
basalt has been dated at 12.2 million years (Wellman and McDougall, 1974).

The assemblages above the basalt contain 4-6% of Nothofagus, insufficient to qualify
for the Nothofagus phase. One assemblage fits the Myrtaceae phase and the other has a high
spore content. There is a good representation of rainforest taxa which 1s usually a feature

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

H. A. MARTIN 231

of the older parts of the Pliocene sequence. However, the location of these bores, in the
narrow, steep-sided valley of Mandagery Creek would favour rainforest taxa (discussed
further below), hence its position in the sequence is uncertain. Hence the basalt date in-
dicates that some of the Myrtaceae phase is younger than 12.2 million years.

Basalts dated at 17 million to 14 million years (Wellman and McDougall, 1974) overlie
lake sediments at Chalk Mountain in the Warrumbungle Range. Two pollen assemblages
from these sediments have been described (Holmes et al., 1983). Both are generally similar
to the oldest of the Myrtaceae phase encountered in the Lachlan River System. One assem-
blage has 0.5% of the brassiz type of Nothofagus and there is a variety of rainforest
angiosperms. However, gymnosperms are unusually abundant for the oldest part of the
Myrtaceae phase. It is thought that a montane lake habitat may not be strictly compara-
ble with the alluvial flats of a broad river valley and this may account for the difference.
However, given that there is some uncertainty about the boundary between the 7° bellus
Zone and the Myrtaceae phase, an alternative interpretation is that these Chalk Mountain
assemblages are youngest 7’ bellus Zone. If this alternative interpretation is accepted, then
they are rather different from the 7? bellus Zone assemblages of the Lachlan River System.
Thus evidence from the Chalk Mountain assemblages suggests that the base of the lower
Myrtaceae phase, i.e. the ‘Pliocene’ sequence, may be as old as 17 million to 14 million
years, or middle Miocene.

There is no evidence to date the Nothofagus/gymnosperm phase. However, inference
from interpretations of the palaeovegetation and changing sea levels (both topics discussed
further below) suggest that it coincided with the early Pliocene highstand sea level.

‘Pleistocene’ Palynofloras

Only a few Pleistocene assemblages have been recovered from the Lachlan River sys-
tem. They are presented here (see Fig. 18) for comparison with the Pliocene sequence.

Myrtaceae is still relatively common with some high values, about 40%. Other assem-
blages may have fairly low values, less than 20%. Casuarinaceae is usually quite low, less
than 5% although there are a few assemblages with moderately high values of about 30%.

Spore content is low, usually less than 8% and of restricted diversity. Anthocerotae is
usually present and the most abundant spore. Cyathea is absent from all except one of the
assemblages and other ferns are minimal.

Gymnosperm content is low, usually less than 5%. Cupressaceae, Araucariaceae and
Podocarpus are the only taxa found here.

Compositae is always present, usually with more than 20%. There are some very high
values, the highest being 64%. Two pollen forms are almost entirely restricted to the Pleisto-
cene, Cichorieae and Tubulifloridites pleistocenicus. Gramineae is usually present in values
greater than 10%, sometimes up to 30%. Chenopod/amaranth is also usually present but
in low quantities, less than 7%. Polyporina granulata is restricted to the Pleistocene.
Cyperaceae and Haloragis are usually present whereas Restionaceae and Sparganium are
rare.

The shrubby element, viz. Acacia, Banksiae, other Proteaceae, Epacridaceae, Monotoca
and Micrantheum are occasionally present. Unidentifiable tricolpate/tricolporates values are
usually very low. The rainforest element is absent or rare. The high pollen producers, viz.
Cyathea, Nothofagus and most gymnosperms are absent or present in such low percentages
that they may represent long distance transport or reworking. Other rainforest taxa such
as Cupaneae, Tasmannia, Quintinia and Symplocos are entirely absent.

The deepest of these assemblages are intermediate between those of the Pliocene and
the Pleistocene. They have a good representation of Myrtaceae and Casuarinaceae,
virtually no rainforest element and relatively low Compositae.

PROC. LINN. SOC. N.S.W,, 109 (4), (1986) 1987

DED PRESIDENTIAL ADDRESS

ep)

A N Section 3
©
o =
i a © S
re) al st roo)
© ri P
COWRA FM

LACHLAN FM

M Cy
mg § PALYNOLOGICAL
| SAMPLE

M 10

M N

S 20 {0} 0.5 1.0

Lachlan R

__.. CARBONACEOUS
LACHLAN E> CLAYS

> Wood, probably
Eucalyptus resinifera

e
| PALYNOLOGICAL
SAMPLE
m
0

ie)
20 u

SECTION 4A

COWRA FM
p

CARBONACEOUS CLAYS

e
| PALYNOLOGICAL SAMPLE

LACHLAN

SECTION 4B

0.5 1.0
2 2: 1 km

°
6K
+
nu
-

SECTION 5

12468

YA
Lachlan R

12464

—— ae ~N

CARBONACEOUS
CLAYS

LACHLAN FM

-
[PALYNOLOGICAL SAMPLE

PROC. LINN. SOC. N.S.W,, 109 (4), (1986) 1987

H. A. MARTIN W8)3)

14804
14805

/
ee

COWRA FM

SECTION 6A

LACHLAN FM

e
a I PALYNOLOGICAL SAMPLE
M ()

10 © 2 CARBONACEOUS CLAY LENS

20 0 0.5 1.0
r 1 1 km

F “Ee SECTION7 | SECTION 8 sw
© +
vt a a
: 3 : ells eas S
o x ays Ss nN
M M
Cc M
M
PALYNOLOGICAL
SAMPLE N+G
: N M
(3) M
10)
10
B
35 B/T 9 3 0 1.0
km km

Fig. 16. The distribution of the palynological phases in sections across the valley. For location of the sections, see
Fig. 2. Cross sections of valley, the distribution of carbonaceous clays and the boundary between the Lachlan and
Cowra Formations from maps supplied by the Department of Water Resources. Only those bores yielding pollen
are included. Legend: P — Pleistocene; PP — Pliocene-Pleistocene transition; M — Myrtaceae phase; G —
Gymnosperm phase; N — Nothofagus phase; S — High spore assemblage; C — High Casuarinaceae assemblage;
B — T. bellus Zone; T — P. tuberculatus Zone.

In summary, the most conspicuous difference of the ‘Pleistocene’ assemblages when
compared with those of the mid Miocene-Pliocene is the virtual absence of the rainforest
element, the high Compositae content, moderate Gramineae content and lower diversity.
The changeover to ‘Pleistocene’ assemblages is recognized in other river valleys and may be
used for stratigraphy (Martin, 1979, 1980, 1981) but there is no direct evidence available for
dating. Wood has been retrieved once from the Cowra Formation, but it is beyond the
range of radio-carbon dating, i.e. older than 35,000 to 40,000 years (Williamson, 1986).

PALAEOVEGETATION

Interpretations of the Tertiary palaeovegetation from pollen assemblages rely on
general principles rather than on direct comparison with some analogous, living vegeta-

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

234 PRESIDENTIAL ADDRESS

wo a
°o N
i) +
© t a
~ = bd r
z+ o
Do vt
bd r-
Jemalong Gap SS | ol |
se)
@ Si pe
C KR o
o
o
vT
© o Pliocene —
a o Pleistocene
\<>| Upper Myrtaceae phase
K Gymnosperm phase
P. tuberculatus |_| Nothofagus phase
x g Zone i
vd m >\j Lower Myrtaceae phase
LY ze SEAVLEVEU is inniyane 0
—- “8 ELEVATED Joom T. bellus Zone
0 5 10 20 25 20

Approximate scales

Fig. 17. Section C-C' running lengthwise of the Lachlan River valley.

tion. A good fit with some extant vegetation does not exist. For example, the three pollen
types of Nothofagus are found together in the early— mid Tertiary assemblages but the living
plants do not grow together anywhere in the world today. The brassi type only is found in
New Guinea, New Caledonia and the New Hebrides, whereas the other two types may
grow alone or together in southeastern Australia, Tasmania, New Zealand and South
America. As a second example, Dacrydium and Dacrycarpus are found in New Guinea, New
Caledonia and New Zealand today. In the early — mid Tertiary, the pollen is found with
Lagerostoma franklini which 1s restricted to Tasmania. Other examples could be given, but
if a general, rather than a specific approach is adopted, floristically, comparable vegetation
is found on the Australian mainland, Tasmania, New Zealand, New Caledonia and New
Guinea.

Experiments with surface samples, in which the pollen assemblage on the ground is
compared with the composition of the vegetation, show that changes in abundance of
pollen may be interpreted as changes in abundance of the parent plant in the vegetation.
For high pollen producing plants, deposition of pollen decreases exponentially from the
source (Birks and Birks, 1980). An empirical value may be determined, values above which
indicate local abundance of the parent plant (e.g. MacPhail, 1979). Low values may be in-
terpreted as pollen transported in from a distance or a low occurrence of the parent plant
in the vegetation. Little may be deduced about the abundance or rarity in the vegetation
of low (percentage) pollen producers.

Assemblages which consist largely of tree pollen may be interpreted as closed forest
(=rainforest). With a closed canopy, insufficient light reaches the ground to support a good
cover of low growing, ground covering, herbaceous plants. With a good representation of
the low growing plants, viz. Gramineae, Restionaceae, Cyperaceae and Compositae, the
forest cover would have been more open. If values for these latter taxa are high, they may
indicate woodland, savannah, grasslands or herbfields.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

239

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PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

Fig. 18. Pleistocene and Pliocene — Pleistocene transitional assemblages. A, Anthocerotae.

236 PRESIDENTIAL ADDRESS

Late Eocene to mid Miocene palaeovegetation

The vegetation throughout the whole of this period of the Tertiary must have been
closed forest, judging by the extremely low content of the light-demanding, low-growing
plants (see Fig. 9 and the 7? bellus Zone in Figs 10 and 11). The drassi type of Nothofagus was
prominent from the late Eocene until the late Oligocene — early Miocene when it declined
and Myrtaceae became the more abundant group. This changeover in the most abundant
pollen group is thought to indicate a climatic change (discussed further below).

In the Murray Basin, N. flemingii was more abundant in some levels of the sequence,
e.g. the 230m to 240m level in bore 36342 (Fig. 9). This greater abundance may be found
anywhere in the Oligocene P. tuberculatus Zone, A and B subdivisions, as shown in Figs 7
and 8. It is thought that N. flemingi required a well-drained habitat (Martin, 1984a; 1986a;
1986b) and the Lachlan region of the basin, with its slightly shallower and rather irregular
basement (see Fig. 7) provided more of this habitat than the Murrumbidgee area (Martin,
1984b). Thus there was a forest mosaic, with N. flemingii common on well-drained sites and
the brassit type of Nothofagus prominent elsewhere. This mosaic would change with time,
according to changes in the course of the rivers and sedimentation. Deposition of sands
with the resulting good drainage would favour WN. flemingit.

The Myrtaceae which became prominent in the late Oligocene-early Miocene, is a
heterogeneous group. The ‘eucalypt type’, Myrtaceidites eucalyptoides, which conforms with
the Angophora — bloodwood eucalypt morphology is present but not common. Recent work
shows that some species of eucalypts have pollen which is much smaller and generally un-
like that of the Angophora — bloodwood eucalypt type (Chalson and Martin, manuscript
submitted). Most of the myrtaceous pollen is generally similar to that of Acmena, Baeckea,
Backhousia, Syzygium, Tristania and probably others as well (Martin, 1978). Decaspermum, Aus-
tromyrtus and Rhodamnia may be present during this time also (Truswell et al., 1985).
Whatever the affinities of this pollen, the very low content of ground covering plants
indicates that the myrtaceous-dominated vegetation was closed forest.

The proteaceous component of the vegetation in the late Eocene may have been sub-
stantial. A number of these proteaceous types (Proteacidites spp.) became extinct about the
end of the Eocene and during the Oligocene, but other types, e.g. Helicia — Orites are found
throughout the whole period (see Appendix 2). Proteaceae pollen is under-represented in
surface pollen spectra. In Eucalyptus forest with s sclerophyll shrub layer in which species of
Proteaceae may constitute the major part, the pollen registers in low percentages only
(Martin, 1978). Tree species of Proteaceae may be widespread in northeastern Queensland
rainforests but their pollen is found only in low percentages in surface samples and Quater-
nary deposits (Kershaw, 1970b). It is thought that the ancient proteaceous types may have
been higher pollen producers with more efficient pollen dispersal (Martin, 1978). A change
in the pollination mechanism of the family Proteaceae 1s proposed by A. R. H. Martin
(1981).

Diversity increases in the early Miocene C subdivision of the PR. tuberculatus Zone, par-
ticularly amongst the tricolpate-tricolporate group. Quantitatively these account for an in-
crease of only a few percent, from 7-9% in the Oligocene to 14-18% in the latter. Given that
the parent plants would have been low pollen producers, they could have been quite signifi-
cant in the vegetation. The proteaceous group, low pollen producers also, increase some-
what in the early Miocene as well (discussed previously). Collectively, these low pollen
producers may have formed a substantial portion of the vegetation.

As discussed previously, early Miocene silicified woods in the upper Lachlan have
been identified as Myrtaceae with affinities to Eucalyptus B, Acacia and Nothofagus (Bishop
and Bamber, 1985). This wood assemblage cannot be placed in any palynological zone, but
it further illustrates the mixed nature of the vegetation.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

H. A. MARTIN DIST

The mid Miocene T° bellus Zone vegetation would have been generally similar to that
of the C subdivision, with a slight change in species composition.

? Mid Miocene-Pliocene palaeovegetation

The botanical affinities of the Myrtaceae pollen group of the Pliocene sequence is not
necessarily the same as that of the group in the late Oligocene-early Miocene. Both the
Angophora — bloodwood eucalypt type and the small-grained eucalypt type (discussed pre-
viously) are present. Other types similar to Austromyrtus, Baeckea, Backhousia, Rhodamnia,
Syzygium, Tristania (Martin, 1973a) and possibly others as well are present. Whatever the
identity of the Myrtaceae pollen, its association with some rainforest element (see Appen-
dix 3) and low Gramineae/Compositae counts (see Figs 10, 11, 12) indicate a good forest
cover.

The Myrtaceae phase may have been closed forest (Martin, 1978) and this interpre-
tation would accommodate the rainforest element present in the Myrtaceae phase.
However, tall open forest (= wet sclerophyll), in which species of eucalypts are dominant
and some rainforest taxa are present as small trees (Ashton, 1981) is an alternative interpre-
tation. The rainforest element present in wet sclerophyll includes myrtaceous taxa.
Moreover, tree ferns are a conspicuous feature of the east coast wet sclerophyll (Ashton,
1981) and these assemblages have a considerable Cyathea spore content. The nature of the
myrtaceous forests is considered further in the discussion.

The gymnosperm and Nothofagus phases would have been closed forests. It is likely that
they had a patchy distribution, occupying the well-watered sites in the valley. Myrtaceous
forests were present also, although probably relegated mainly to drier sites, the slopes and
ridges. Nothofagus was more extensive upstream near Cowra, with a noticeable decline
downstream, with very little at Jemalong Gap (see Fig. 13). This distribution suggests that
Nothofagus migrated downstream from the highlands where it was probably considerably
more abundant during the late Miocene. Gymnosperms were most abundant upstream
also, but they were relatively common downstream as well, to beyond Jemalong Gap (see
Fig. 17). Podocarpus was the most common gymnosperm but Cupressaceae, Phyllocladus and
Dacrycarpus were sometimes abundant (see Fig. 14). Today, Callitris of the Cupressaceae is
well known in the inland, semi-arid regions and may grow as forests in the Lachlan Valley,
but there is one rainforest margin species, Callztris macleayana (Boland et al., 1984). There
are other possibilities within the family Cupressaceae. Whatever the identification of the
fossil Cupressaceae pollen, its association with other rainforest taxa and a carbonaceous
clay lens (indicative of a swampy environment) suggests that it was a rainforest taxon also.

Other rainforest angiosperms (see Appendix 3) were usually present. The pollen
registers in low percentages, but the parent plants may have been relatively common. They
may have occupied favourable habitats such as stream sides and sheltered gullies, or may
have been present in the understorey layers of wet sclerophyll.

As discussed previously, a more specific identification of Casuarinaceae pollen is
difficult. Gymnostoma may be found in rainforests. Casuarina is not a normal constituent of
rainforest but it may be found in open forest bordering rainforest. Species of Casuarina may
be found in most kinds of vegetation, from coastal open forests to arid shrublands.
However, the high Pliocene Casuarinaceae assemblages occur stratigraphically close to the
Nothofagus and gymnosperm phases, hence they may represent rainforest taxa or taxa which
border rainforest. Riparian species may have been involved. The group may have consisted
of different taxa at different times.

A shrub or small tree element may be present also (see Appendix 3). Most of the taxa
in this element are represented by only a few percent and little may be deduced about their
abundance. In a study of surface samples, Ladd (1979) did not recover Acacza pollen, yet

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

238 PRESIDENTIAL ADDRESS

Acacia was co-dominant in the vegetation beneath which the samples were taken. Thus taxa
in this element may have been common in the vegetation.

Fern spores are subject to water transport (Ladd, 1978; Birks and Birks, 1980). Con-
sequently, the high spore assemblages may have been produced by runoff from a nearby
gully containing abundant ferns. Such gullies would be particularly favourable for Cyathea.

Towards the top of the upper Myrtaceae phase, Compositae and Gramineae increase
and the gymnosperms and other rainforest taxa decrease. These assemblages are thought
to be transitional between the Pliocene and Pleistocene. The closed forest canopy was
decreasing and open forest, woodland and grasslands were expanding.

In summary, the ?mid Miocene — Pliocene sequence is a record of gradual decline of
the rainforest element. It was moderately common in the oldest part of the sequence. The
vegetation was probably a mosaic of rainforest in the most favourable habitats, wet
sclerophyll (further evidence for wet sclerophyll is presented in the section on fire history)
covering considerable areas and with dry sclerophyll in the driest habitat. For a period,
probably in the early Pliocene, rainforest expanded and would have occupied substantial
areas of the valley during the Nothofagus and gymnosperm phases. Subsequently in the
upper Myrtaceae phase, the rainforest element declined and eventually disappeared from
the Lachlan Valley and its tributaries.

Pleistocene palaeovegetation

The abundance of Compositae and Gramineae (see Fig. 18) indicates open vegeta-
tion, probably woodland and grasslands/herbfields. Most of the trees would have been
myrtaceous or casuarinaceous. The Casuarinaceae group, which is sometimes abundant,
probably consisted of different species to those in the group during the late Miocene —
Pliocene. Gymnosperms were rare and rainforest angiosperms (see Appendix 3) had
disappeared from the valley and its tributaries.

The spore content 1s low and Anthocerotae are the most common of the group.
Anthocerotae are commonly found along stream banks in open vegetation.

The Pleistocene sequence is very disjointed and nothing may be deduced about the
vegetation of glacial and interglacial times. The stratigraphic position of the assemblages
in the Cowra Formation (see Fig. 16) suggests most of them may be contemporaneous. It
is thought these assemblages are older Pleistocene in age. It may be argued that the inter-
glacials were more humid, being times of high sea level, and swamps necessary for preser-
vation of pollen would be more likely at these times, hence all of the pollen assemblages
presented here probably represent interglacial period(s). However, there is no direct
evidence to support this argument. These Pleistocene assemblages are presented here for
comparison with those of the Pliocene and to illustrate the considerable difference between
the two.

Wood has been encountered in bore 12437, Section 4 at a depth of 26m (see Fig 16).
It has been identified as probably Eucalyptus resinifera by H. K. Bamber, Forestry Com-
mission (Williamson, 1986). E. resinifera is found in dry sclerophyll and wet sclerophyll
(Boland et al., 1984). Unfortunately, pollen was not recovered from this level in bore 12437.
The identification of E. resinifera suggests that some relatively denser forest vegetation may
have been present in the Pleistocene, probably confined to the more favourable habitats.

FIRE HISTORY

Black carbonized particles are readily recognized in palynological preparations. Such
particles are usually regarded as charcoal fragments and variations in abundance of the
particles are used to reconstruct Quaternary fire history (e.g. Singh et al., 1981). Carbonized
particles, or fusinites of the coal petrologist, may result from charring, oxidation, moulder-

PROC. LINN. SOC. N.SW., 109 (4), (1986) 1987

H. A. MARTIN 239

ing or fungal attack before deposition, or on the peat surface. Carbonization may also result
from coalification after deposition (Teichmuller, 1982). Thus care must be exercised in the
interpretation of carbonized particles, particularly for the Tertiary and older geological
periods.

Fire 1s not essential for carbonization of plant material. It is thought that dehydration
and oxidation on the swamp surface may produce fusinite. Fungal attack may cause effects
similar to carbonization. For example, dry rot alters the unused part of the wood into
carbon-rich, humic substances. Other fungi and other plants may produce dark material,
which looks similar to carbonization. These and other factors are reviewed by Teichmuller
(1982). By and large, these processes which produce carbonization without burning are not
well understood, but there seems little doubt that a portion of this material may be
attributed to burning (Kemp, 1981).

The re-working of carbonized particles in older sediments, particularly if they contain
coal seams, may cause serious error in the deduction of a fire history. However, re-working
may be detected by older palynomorphs included in a younger assemblage. A very small
amount of early Tertiary re-working has been detected in the late Miocene — Pliocene
sequence of the Lachlan River Valley, but no Cretaceous or older palynomorphs have been
found. In particular, there is no evidence at all of re-worked Permian coals, the most likely
cause of serious error. There may be a very small amount of Early Cretaceous re-working
in the base of the Eocene — Oligocene sequence of the Murray Basin.

When fire is the cause of carbonization, a high degree of charring may result in very
brittle fusinite which readily disintegrates into fine fragments. Less strongly charred wood
may preserve the cellular structure, although probably with distortions (Teichmuller, 1982).
Much of the fuel in forests is bulky and rarely, if ever, burns completely (Luke and
McArthur, 1978). Fires may sweep over herbaceous swamps, burning the plant cover to
water level. Fusinites of such peats shatter easily and are deposited as fine splinters. The
surface of a peat swamp may occasionally dry out and ground burning may occur with the
formation of a great deal of ash. This ash, however, is easily blown away or dissolved in
swamp waters which are rich in carbon dioxide. Some charcoal remains on the ground, but
the fine fragments are readily blown away and deposited elsewhere (Teichmuller, 1982).
These are a few of the possible effects in the deposition of carbonized particles that may
result from fires.

Clark (1984) reviews the effects of different pollen preparation procedures on car-
bonized particles and the problems of identification. The most important for this study,
which presents counts of carbonized particles, is that larger particles may be broken into
small pieces. Burning destroys the stratification of the cell wall (middle lamella, primary
wall and secondary wall) and this feature may be used to identify charcoal from dark-
coloured, unburned plant tissue (Cope and Chaloner, 1980). However, different chemical
treatments may destroy stratification or produce a similar effect in cell walls which have had
stratification destroyed (D. R. Selkirk, fers. comm.). In this study, carbonized particles were
counted in preparations made for palynomorphs where treatment included a moderate
oxidation process. Identification of particles was made with reference to burnt plant
material which was then treated by the same chemical procedure as that used on the sedi-
ments. Particles within the size range of pollen were counted. Estimations of the area of car-
bonized particles (Clark, 1982) which allow for the different sizes of the particles were not
possible as the preparations were made long before the decision to count carbonized
particles and the methods used are unsuitable for area estimations. Counts are presented
as the ratio of carbonized particles to total pollen count, in Fig. 19.

The interpretation of the abundance of carbonized particles is problematical (see
review by Clark, 1983), even with the assumption that most of the particles result from
burning. Charcoal particles may be transported by air currents, particularly at the time of

PROG. LINN. SOC. N.S.W., 109 (4), (1986) 1987

240 PRESIDENTIAL ADDRESS

fire, or washed from bare ground after fires. Larger particles may be broken into smaller
fragments before deposition. A greater abundance of charcoal does not necessarily mean
more fires, it may result from bigger fires. More charcoal probably indicates that more fuel
was burnt, but even this interpretation assumes that the fuel was charred to the same
degree.

The prerequisites for wild fires are

1. a fuel supply;

2. the fuel must dry out sufficiently so that it can burn; and

3. asource of ignition.

Lightning would have been the main agent (Kemp, 1981), and probably the sole
source of ignition for the Tertiary and the portion of the Pleistocene under consideration
here, which predates the arrival of man.

The palaeovegetation was almost entirely forests, with woodlands and grasslands be-
coming prominent only in the Pleistocene. Fuel is not limiting in forests (Luke, 1961).
Closed forests rarely burn, except in drought periods when they are subjected to excep-
tional drying (Webb, 1970; Luke and McArthur, 1978). However, the pollen record is
biased towards those plants growing close to the site of deposition. It may be that closed
forest grew around the site of deposition and was rarely burnt, whereas a drier, more open
kind of vegetation on the slopes and ridges was burnt more frequently, the charcoal being
transported eventually to the site of deposition.

The interpretation adopted in this study is that an increase 1n carbonized particles,
most of which are probably charcoal, indicates a greater frequency and/or intensity of dry
periods which would allow burning, given adequate fuel. The major control of wild fires is
thought to be climatic (discussed further below). Charcoal from fires anywhere in the
catchment would eventually become incorporated in sediments at the site of deposition.

Counts of carbonized particles are presented in Fig. 19. In the late Eocene — early
Miocene of Bore 38342 (Fig. 19A), the ratios of carbonized particles are low but increase
somewhat towards the top of the sequence. In this sequence, there is a variety of dark-
coloured bodies, some of which may result from the coalification process, but only particles
similar to the reference burnt plant material and to the carbonized particles seen in the late
Miocene — Plocene sequence were counted.

The abundance of carbonized particles in the mid Miocene T° bellus Zone at
Euabalong and Jemalong Gap is comparable to the early Miocene in the top of Bore 36342.
In the Myrtaceae phase of the late Miocene — Pliocene sequence, the abundance is much
higher. The ?Nothofagus-gymnosperm phase of Jemalong Gap has lower ratios of car-
bonized particles which are comparable with those of the 7’ bellus Zone. In Bore 14747, the
Nothofagus phase has low ratios and the ratios of the gymnosperm phase are intermediate
between the Nothofagus and Myrtaceae phases.

In summary, burning levels were extremely low in the late Eocene and most of the
Oligocene. In the late Oligocene to mid Miocene, burning increased somewhat but it was
still relatively low. Burning increased considerably in the lower Myrtaceae phase (which
may be as old as mid Miocene). Burning decreased in the Nothofagus and gymnosperm
phases (probably early Pliocene) to levels similar to those of the Oligocene — mid Miocene
and increased in the upper Myrtaceae phase to the highest levels of the sequence.

THE EFFECTS OF CHANGING SEA LEVEL
Changes in sea level have an important influence on sedimentation and the environ-
ment, especially close to the shoreline. At times of high sea level, drainage is sluggish and
the low-lying areas in the landscape are swampy. Sediments accumulate at these times.
Evaporation is higher from the shallow seas over the continental shelves and flooded low-

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

H. A. MARTIN 241

D SECTION 6, BORE 14747
Depth

PHASE CARBONIZED PARTICLES
(m)
40

Upper Myrtaceae

60
Gymnosperm

Nothofagus
80
Lower Myrtaceae

B JEMALONG GAP, BORE 36079

Depth PALYNOLOGICAL |
DIVISION

40 -

Pleistocene

CARBONIZED PARTICLES

60 4

Upper Myrtaceae

80 5

100 ————

Nothofagus —
Gymnosperm

120 -| Lower Myrtaceae

140 | T. bellus

°
zl
o
ny
a

A TOM'S LAKE, BORE 36342
Depth ZONE CARBONIZED
(m) PARTICLES
1004 fe ee

C EUABALONG, BORE 36171
Depth PALYNOLOGICAL |

al — (m) DIVISION CARBONIZED PARTICLES
eee 604.
140-| \secanovenl
P. tuberculatus Myrtaceae [oo
B E 70-
1807 Do nn)
— [as SSe ea
=| = =
T. bellus
{ess 80-7
220-4 ooo
egal ene aP fe) 4 8
| i C RATIO Carbonized particles

Total spore-pollen count

260-| Middle N. asperus
+

RATIO Carbonized particles

Total spore-pollen count

Fig. 19. Carbonized particle counts.

PROC. LINN. SOC. N.S.W,, 109 (4), (1986) 1987

242 PRESIDENTIAL ADDRESS

lying areas. As a consequence, precipitation 1s greater and through this effect, changing sea
levels may have considerable influence along way from the shoreline. At times of low sea
level, with the shoreline close to the edge of the continental shelf, drainage is more efficient
and there may be erosion and/or lack of sedimentation. Evaporation from the colder,
deeper seas at the edge of the continental shelf is less, consequently precipitation is lower.
At these times, the climate is more continental.

Fig. 20 presents the late Eocene to Oligocene changes in sea level, based on the
sedimentary cycles recognized on the southern margin of Australia (from Loutit and
Kennett, 1981). This curve is the most appropriate, for although the changes in sea level
may be correlated on a worldwide basis, local tectonics may modify their expression.
Sedimentary cycles have not been reported for the late Pliocene to Pleistocene on the
southern margin of Australia so the global changes in sea level (from Vail et al., 1977) are
presented for this time range. The changes became more frequent in response to glacial-
interglacial oscillations of the last 2-3 million years.

In Bore 30407 of section A-A! (see Fig. 7), the younger P. tuberculatus Zone extends
almost 60m below the boundary of the older Middle N. asperus Zone. This depth of 60m is
considerable and more than would be expected with a simple change of the river course. It
is thought to illustrate downcutting at a time of low sea level and subsequent fill with
younger sediments.

In the eastern edge of the Murray Basin, the Oligocene low sea level (Fig. 20) is ex-
pected to be a time of erosion. However, the effect of this low sea level is not detectable in
the non-marine sediments (R. M. Williams, pers. comm.). It is thought that with the almost

AUSTRALIA
SOUTHERN MARGIN

10 4
Zz
Ww
(6)
2)
2047
uw
z
Ww
(6)
304°
Oo
=
re}
40-269

B GLOGAL
SEA LEVELS

Fig. 20. Changes in sea level. The southern margin of the Australian continent from Loutit and Kennett (1981)
and global sea levels from Vail et al. (1977).

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

H. A. MARTIN 243

flat terrain and a rainfall sufficiently high to maintain the forest cover, erosion would be
minimal (Bishop, 1985b; Martin, 1986a; 1986b). The late Miocene low sea level, however,
is evident in the sediments (R. M. Williams, pers. comm.). At this time, the climate was be-
coming drier and the cover of vegetation would be more difficult to maintain, thus allowing
erosion (Martin, 1986a; 1986b). The relative scarcity of the mid Miocene 7? bellus Zone
which was deposited at a time of high sea level, may be attributed to subsequent erosion
during the late Miocene low sea level.

The lower Myrtaceae phase 1s not as thick as would be expected if, as discussed pre-
viously, it begins in the mid Miocene. It is thought that the late Miocene low sea level,
accompanied by a somewhat drier climate (discussed further below) resulted in erosion or
lack of deposition and this may account for the relatively thin sections of lower Myrtaceae
phase seen in Figs 10, 16 and 17. Other factors may be involved as well. Tectonics may have
had some influence in the Lachlan River Valley where lower Myrtaceae phase forms a rela-
tively thin section over the basement. As discussed previously, there is also uncertainty
about the age of the transition of the 7’ bellus Zone and lower Myrtaceae phase (see Fig. 5).

The early Pliocene high sea level is expected to be a time of higher precipitation which,
it is thought, allowed the Nothofagus and gymnosperm phases in the Lachlan River Valley.
With the subsequent fall in sea level in late Pliocene, the Myrtaceae phase is evident once
more.

The relatively rapid rise and fall of sea level in the Pleistocene cannot be detected in
the palynological record which, as discussed previously, is extremely fragmentary.
Moreover, the sea was denied access to the Murray Basin by the closure of the entrance,
probably in the late Pliocene (Brown et al., 1968; Abele et al., 1976). As a consequence, the
drainage route to the sea became much longer and this would have made a difference to the
erosion/sedimentation associated with low/high sea levels (respectively).

PALAEOCLIMATE

The palaeoclimate may be deduced from the climatic requirements of comparable,
present-day vegetation, but this can only be attempted at a very general level. Interpreta-
tions of the palaeovegetation are very general and as discussed previously, a good fit with
some extant vegetation does not exist. However, some general climatic parameters may be
deduced from the extant vegetation and applied to the palaeovegetation to illustrate
climatic trends.

Brassit species of Nothofagus may be dominant in the mid montane zone of the New
Guinea Highlands (Johns, 1982). Nothofagus is generally associated with high precipitation
of 1500-1800mm and considerable cloudiness which reduces light intensity and maintains
high humidities. It is generally absent from areas which suffer a regular and sustained
water deficit (Ash, 1982). In New South Wales, one species of the menzzeszz type, Nothofagus
mooret is present in the Eastern Highlands and may be dominant where precipitation
exceeds 1800mm. It is usually restricted to sites that are commonly fog bound (Baur, 1957).
Boland et al. (1984) give the annual rainfall of N. moore, not necessarily dominant in the
vegetation, as 1500mm, with the driest months receiving 60mm, augmented by mountain
mists. In general, Nothofagus requires a high precipitation and maintenance of relatively
high humidities throughout the year.

As discussed previously, the myrtaceous vegetation may have been closed forest. In
New South Wales, Myrtaceae are common in rainforests which require a precipitation of
1500mm for widespread development (Baur, 1957). Alternatively, the myrtaceous vegeta-
tion may have been wet sclerophyll (or mosaic of rainforest and wet sclerophyll) and the fire
history favours this interpretation. Wet sclerophyll may be found over large tracts receiving
between 1000mm and 1500mm (Ashton, 1981). Thus the Myrtaceae phase probably

PROG. LINN. SOG. N.S.W.,, 109 (4), (1986) 1987

244 PRESIDENTIAL ADDRESS

indicates a precipitation of somewhat less than 1500mm, probably between 1000mm and
1500mm.

The disappearance of the rainforest element from the landscape is another important
parameter. The lower precipitation levels of the major eucalypt species of the drier end of
the range of wet sclerophyll (in Ashton, 1981) is about 500-700mm (precipitation require-
ments from Boland et al., 1984). The limit of subcoastal rainforest pockets across northern
and northeastern Australia falls between the 600mm and 800mm isohyet (Webb and
Tracey, 1981). Whichever interpretation of the Myrtaceae phase 1s favoured, it makes little
difference to the climatic parameter at the drier end of the range.

Using these parameters and the trends shown by the carbonized particles, changes in
climate may be reconstructed. It should be emphasised that there is no great precision in
this reconstruction: the parameters are used more to illustrate a climatic trend.

In the Hillston region, the precipitation of the late Eocene and Oligocene was high,
probably above 1800mm. High humidities were maintained throughout the year and the
vegetation rarely dried out sufficient to allow burning. Fires would have been very limited
in extent. There may have been a gradual, slight decrease in precipitation during the upper
part of the Oligocene. In the late Oligocene — early Miocene, precipitation was probably
less than 1800mm but above 1500mm. There was no definite seasonal dry period and fires
were only slightly more frequent than in the preceding period. This level was maintained
into the mid Miocene 7, bellus Zone.

In the Hillston region, pollen preservation ceased in the early Miocene. Pollen preser-
vation requires permanently wet sites which remain wet long enough for burial to a depth
below the influence of the fluctuating water table. The disappearance of permanently wet
sites from the landscape would have been the result, in part, of climatic change over the
region as well as of a decrease of water transported into the region by the river (discussed
further below). Subsequently, the sites of pollen preservation were located upstream from
the edge of the Murray Basin, particularly around Jemalong Gap and in the Lachlan River
Valley to Cowra.

In the ?mid Miocene, 1.e. the start of the lower Myrtaceae phase, there was a further
drop in precipitation probably to somewhat below 1500mm, but not less than 1000mm. A
well-defined seasonal dry period became established at this time, and burning became a
regular event in the landscape. The drier slopes and ridges were probably most subject to
burning.

In the early Pliocene, the time of the Nothofagus and gymnosperm phases, precipita-
tion increased, probably to more than 1500mm but not more than 1800mm. The level of
burning decreased. In the mid — late Pliocene, the precipitation decreased once again to
about or below 1500mm and the level of burning increased as well. Precipitation continued
to decrease so that at the end of the Pliocene, it was probably about 500mm-800mm.

Wood recovered from the base of the Pleistocene Cowra Formation, identified as prob-
ably Eucalyptus resinifera, indicates a precipitation of at least 800mm, the lower limit for this
species. This evidence does not contradict the above deductions, which are at best very
general, and given the lack of any evidence to date the ‘Pliocene — Pleistocene’ boundary.

The changes in precipitation are shown diagrammatically together with the major
palynological events in Fig. 21.

‘Today, the mean annual precipitation for Hillston is about 140mm, Forbes 200m to
210mm and Cowra, 240mm to 250mm, respectively.

The climatic parameters discussed above have been deduced from plant growth and
in this context, ‘precipitation’ is more precisely ‘effective moisture’. Effective moisture in-
cludes both rainfall and water transported by the river systems (Martin, 1986b) and the
latter was probably important in the maintenance of the permanently wet sites required for
pollen preservation. The climatic change described above could not occur over the Lachlan

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

<«— Bulseaiou

+— Bulseesouy

+ Bulseasou
0 0 «Bulseasou; 0

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+ Bujseeiou

EOCENE

MILLION Y

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fo)
1

OLIGOCENE

snsedse “N SIppIW

snjejndseqn} ‘gq

H. A. MARTIN 245

EARS

-O2

Bn
= vu
1 | r
m
MIOCENE PLIOCENE |a
1 1 im
(@)
fe} U
| 8 § lg|_ 2
= 8 2 8/2 Ss
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® = =z 'o/qG O
> < < = ie
S| = = (S\2
© o '8|/= @
> Sens ?)
ee one lee
oO Ira

snBejoyjon
JO edA} JIssesg:

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SdNOYS N3AM10d YOrviNn

KXXKKK EKER Y
lietonnnnicnianen eaten

LEER

=||5
aie
alle
Oj}|>
Ze
D

Ble
ey m
Zila

ROO Cy AX RERRKXINNY

SAA SOS AAAS Od RR |
ARRAN RRRY, ST OT RORY =
| RAK RAXRAXX RAK RRR | oe

RAXKKXRXX XNA RRR |
ER KRY ”
Cooeeuaaaaay eee =

OXEKX NR

| ER? Oo
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<

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Fig. 21. Summary of major palynological events and the inferred changes in precipitation (Not to scale.)

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

246 PRESIDENTIAL ADDRESS

Valley in isolation from the surrounding areas. Parallel changes would have occurred in the
Eastern Highlands over the headwaters of the river and these would have been very impor-
tant for the volume of water transported by the river.

Effective moisture also includes the influence of temperature through its effect on
evaporation. Temperature 1s also very important for plant growth but it is not considered
here because effective moisture is thought to be the major control. Oligocene — early
Miocene temperatures were probably somewhat higher than those of today. Surface sea
temperatures during the mid Miocene were about 5°C higher than those of today (Savin
et al., 1975; Shackleton and Kennett, 1975), hence land temperatures were probably some-
what higher also (Martin, 1986b).

DISCUSSION

The nature of the late Miocene — Pliocene myrtaceous forests with its component of
rainforest taxa has long been problematical. It was thought that they were closed forests,
perhaps akin to the ‘heath forests’ of Borneo (Martin, 1982). This interpretation was based
on an inadequate appreciation of the diversity of pollen types within the genus Eucalyptus.
With a greater eucalypt content, an alternative interpretation of wet sclerophyll forest is
possible. The record of carbonized particles 1s extremely important in this context. Rain-
forest rarely burns (Webb, 1970; Luke and McArthur, 1978) whereas ‘there can be little
doubt that fire is an integral part of the environment’ of wet sclerophyll forests (Ashton,
1981). The great increase in carbonized particles in both the lower and upper Myrtaceae
phases supports the interpretation of wet sclerophyll forest. As discussed previously, the late
Miocene — Pliocene palaeovegetation was probably a mosaic of rainforest, wet sclerophyll
and dry sclerophyll, each type of vegetation occupying the appropriate habitat in the
landscape.

A more precise identification of myrtaceous pollen is possible. The grain may be
scored for a number of characters and a combination of several character states may be dis-
tinctive. It is found that the character states of the fine detail are the most promising for this
purpose (Chalson and Martin, manuscript submitted). Taxonomically closely related
species may not be separable (Martin and Gadek, manuscript submitted), but pollen of the
family should be divisible into a large number of groups. A more precise identification of
the pollen awaits the compilation of a reference set scored for the distinctive character
states. For the late Miocene — Pliocene assemblages, this set should include all species com-
mon in eastern Australian rainforests, wet sclerophyll and dry sclerophyll — a formidable
task. For older assemblages, the reference set should include species in New Guinea, New
Caledonia and New Zealand.

Late Oligocene — early Miocene assemblages from the western Murray Basin have a
considerable myrtaceous content (Truswell et a/., 1985) but it should not be assumed this
content is the same as that of the late Miocene-Pliocene assemblages in the Lachlan valley.
A study of the carbonized particles may shed some light on this problem, but more precise
identification of the myrtaceous pollen is required for the western Murray Basin also.

The major changes of palaeoclimate, as inferred from the vegetation, are thought to
be linked with the development of circum-Antarctic oceanic circulation and the extent of
glaciation on Antarctica (Martin, 1986b). Antarctica has had, and still has, a profound in-
fluence on world climate (Flohn, 1978), particularly on Australia because of its close prox-
imity. About late Oligocene, circum-Antarctic circulation was established (Kennett, 1977;
1978). This factor probably reduced the efficiency of the heat transfer from the equator to
the pole, thus increasing the temperature gradient between these regions. The extent of ice
cover on Antarctica is uncertain but all the evidence indicates high latitude cooling and a
northwards shift of westerly winds which would have influenced much of southern

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

H. A. MARTIN 247

Australia (Kemp, 1978; Flohn, 1978). It is thought that these events initiated the decrease
in precipitation which resulted in the decline of the brassz type of Nothofagus (Martin,
1986b).

The early Miocene was a time of increasing temperatures (Savin et al., 1975;
Shackleton and Kennett, 1975). At the beginning of the mid Miocene, a major global cli-
matic threshold was reached with the development of the East Antarctic ice cap (Kennett,
1977; 1978). The cause of this event remains unknown, but Kennett (1978) notes that this
development occurred at a time of warmer temperatures which would have increased
precipitation over Antarctica. (The mid Miocene was a time of high sea level, as discussed
previously.) By the late middle Miocene, temperatures began to fall again. In the late
Miocene, the ice cap retreated somewhat. It is difficult to match up the complex, poorly-
dated events in the Lachlan River region with the complex developments in Antarctica.
However, the mid Miocene was a time of profound change in both, and in this respect, they
are in agreement. The late Miocene was a time of major cooling (Savin et al., 1975;
Shackleton and Kennett, 1975). Cooler oceanic temperatures result in lowered precipita-
tion and this is in accord with the lower Myrtaceae phase of this time and the inference of
reduced precipitation.

The charcoal record suggests that a well marked dry season, which allowed fires to be-
come an integral part of the environment, dates from the ?mid—late Miocene.

A major extension of the Antarctic ice cap occurred in the late Miocene — early Plio-
cene (Kennett, 1977), a time of somewhat higher temperatures (Savin et al., 1975) and high
sea levels. This is thought to coincide with the Nothofagus/gymnosperm phases. Cooling
continued from the mid Pliocene (Kennett, 1977; Savin et al., 1975) which would have
resulted in lower precipitation and the return of the Myrtaceae phase. A further global cli-
matic threshold was passed in the late Pliocene when the glacial/interglacial oscillations
commenced (Kennett, 1977). It is thought that this event coincided with the elimination of
the rainforest taxa from the Lachlan River valley.

Climatic changes such as these could not .have occurred over the Lachlan valley in iso-
lation from the surrounding areas which must have experienced similar or parallel changes.
There is evidence of a climatic gradient across the Murray Basin, parallel to that of today,
during the late Oligocene — mid Miocene (Martin, 1986b).

Decreasing precipitation over the Eastern Highlands would have had extremely 1m-
portant consequences on the activity of the rivers. The mid Miocene decrease in precipi-
tation would have reduced the volume of water carried by the river such that it was unable
to maintain the permanently wet sites in the Hillston region, but it was sufficient to main-
tain some permanently wet sites upstream in the Lachlan valley.

Weathering subsequent to deposition may destroy pollen and this may alternatively
account for lack of pollen preservation. However, it is unlikely to be the sole cause. In the
Hillston region, pollen preservation ceases in the early Miocene and the upper 80-100m of
sediment are barren. In the Cowra district, however, grey carbonaceous clays containing
Pleistocene assemblages may be found at depths of less than 20m. Thus the observed
pattern of the cessation of pollen preservations being younger, further upstream best fits an
hypothesis of a climatic gradient and a progressively diminishing volume of water carried
by the river.

Changing sea levels and climate have probably played a part in shaping the Lachlan
valley itself. As discussed previously, the late Miocene low sea level was a time of erosion
and this would have removed most of the older sediments in the valley. Early and mid
Miocene sediments are only found south of the present river near Forbes (see Figs 2 and
16F). The river once turned southwest about Section 7 and passed under Section 8, some
15km southwest of Forbes (discussed previously). When base levels were lowered and down-

PROC. LINN. SOC. N.SW,, 109 (4), (1986) 1987

248 PRESIDENTIAL ADDRESS

cutting commenced, it is likely that the river adopted its present, shorter course to
Jemalong Gap, thus bypassing the older sediments to the south.

The late Miocene was also a time of reduced precipitation hence the river carried a
reduced volume of water which would have cut a narrower valley within the existing wide
valley to produce the observed valley-in-valley structure. Williamson (1964; 1986) argues
that the valley-in-valley structure was produced as a result of uplift of the highlands but
Bishop et al. (1985) note that there does not seem any need to invoke dynamic (active) tec-
tonism to account for the geomorphology of the Lachlan Valley. However, isostatic rebound
(1.e. passive tectonism) probably occurred in response to erosional unloading (Bishop,
1985b). Moreover, signs of minor isostatic uplift may be expected mostly at the edges of the
highlands (Bishop, 1987). Earthquakes in the region today show that adjustment to stresses
in the earth’s crust is still in progress (Denham et al., 1985). Thus passive tectonism, com-
bined with a reduced precipitation probably produced the valley-in-valley structure.

Climate has probably had an influence on the nature of the sediments themselves.
With a high precipitation, various rock types would be decomposed leaving only the most
resistant quartz and chert, as seen in the Lachlan Formation. The Cowra Formation, which
contains an assortment of the rock types found in the catchment area, was deposited under
much lower precipitation: insufficient for their decomposition. As discussed previously,
Williamson (1986) postulates a formerly widespread quartz gravel formation as the source
of the quartz in the Lachlan Formation. No doubt, older sediments have been reworked
and they were probably mainly quartz. An hypothesis about the source of the quartz sands
and gravels in the Lachlan Formation would have to be suitable for a very wide application,
for all the Pliocene and older sediments of the river valleys of the western slopes of New
South Wales have similar quartz gravels and sands. Experience with sediments of Creta-
ceous to early Oligocene age in the Gippsland, Bass and Otway Basins (Martin, unpubl.)
has revealed similar quartz sands and the almost complete absence of other resistant rock
types. Precipitation was also higher throughout this time span. Thus it is thought that
climate has been the major factor in the production of these quartz rich sediments.

ACKNOWLEDGEMENTS

I am indebted to the Water Resources Commission of New South Wales for samples
from bores and for financial support. Comments and criticisms from Dr P. Bishop, Dr
A. P. Kershaw and Dr A. R. H. Martin have been invaluable but the opinions expressed
here are my own.

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VaIL, P. R., MITCHUM, R. M., Jr., and THOMPSON, S., III, 1977. — Seismic stratigraphy and global changes
of sea level, Part 4: Global cycles of relative changes of sea level. Amer. Assoc. Petrol. Geol., Mem 26: 83-97.

Wess, L. J., 1970. — Eastern Australian environments in relation to fire. Jn: ‘Environments to order?’ Proceed-

ings of asymposium, Australian Museum, 8th Aug. 1970 (unpubl.): 5-8.

,and TRACEY, J. G., 1981. — Australian rainforests: patterns and change. Jn: A. KEAST, (ed.), Ecological

si0gcography of Australia; 605-694, The Hague: Junk,

WELLMAN, P., and MCDOUGALL, I., 1974. — Potassium-argon ages in the Cainozoic volcanic rocks of New
South Wales. J. Geol. Soc. Aust. 21: 247-272.
WILLIAMSON, W. H., 1964. — The development of ground-water resources of alluvial formations. Paper B 11

in Water resources use and management. Proceedings of a symposium. Canberra 9-13th Sept. 1963; 195-211.
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PROC. LINN. SOC. N.SW,, 109 (4), (1986) 1987

H. A. MARTIN 251

——,, 1986. — Investigation of the groundwater resources of the Lachlan Valley alluvium. Part 1: Cowra to
Jemalong Weir. Water Resources Commission of N.S.W. Hydrological Report No. 1986/12.
WOOLLEY, D. R., 1978. — Cainozoic sedimentation in the Murray Drainage Basin, New South Wales. Proc.
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Basin, N.S.W. Aust. Soc. Soil. Sci. Symposium, Griffith 1977: 45-65.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

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PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

Post-Fire Demography in the Resprouting Shrub
Angophora hispida (Sm.) Blaxell: Flowering, Seed
Production, Dispersal, Seedling Establishment

and Survival

TONY D. AULD

(Communicated by P. MYERSCOUGH)

AULD, Tony D. Post-fire demography in the resprouting shrub Angophora hispida (Sm.)
Blaxell: Flowering, seed production, dispersal, seedling establishment and
survival. Proc. Linn. Soc. N.S.W. 109(4), (1986) 1987: 259-269.

After fire, the resprouting shrub Angophora hispida (Sm.) Blaxell rapidly forms an
extensive canopy from epicormic and lignotuberous shoots, culminating in most plants
in extensive flowering and fruiting. Flowering was virtually absent in populations which
had not been recently burnt. There are virtually no predispersal seed predators in
mature fruits.

Seed dispersal was minimal and confined to the immediate vicinity of the parent
plant. In this study, released seeds were all viable and germinated following the first
heavy rains. Seedling mortality was highest in the establishment phase (1.e., before the
first leaves were produced) and declined markedly afterwards, remaining at a much
lower rate for the following eight years. Seedling growth and lignotuber development
were slow and after eight years the young plants may not be fire resistant. The impor-
tance of the length of inter-fire period for the survival of populations of this species is
discussed.

Tony D. Auld, National Parks and Wildlufe Service N.S.W., P.O. Box N189, Grosvenor St, Sydney,
Australia 2000; manuscript received 21 October 1986, accepted for publication 18 February 1987.

KEY WORDS: Demography, Angophora hispida, fire, resprouter.

INTRODUCTION

Two modes of post-fire regeneration have been documented. These are regenera-
tion solely from seed following the death of all adult plants during a fire (obligate
seeders), and regeneration from protected vegetative buds on surviving plants
(resprouters) (see Gill, 1981). Whilst several workers have categorized plants according
to their mode of recovery after fire (Purdie and Slayter, 1976; Purdie, 1977; Benson,
1985), information relating to the long-term survival and reproduction of species is
sparse.

Keeley and Zedler (1978) developed a model to explain the evolutionary develop-
ment of these two major modes of post-fire recovery in relation to fire frequency, in fire-
prone Californian vegetation. Based on the demography of several representative
species, they predicted that high fire frequencies favoured the evolution of resprouters
over that of obligate seeders. One aspect of the life cycle which they did not consider was
the importance of the primary juvenile period in resprouters. Bradstock (1985), who
worked on two resprouting and two obligate seeding species, predicted that repeated
fires of short frequencies may eliminate both resprouters and seeders. In resprouters,
this was because of the slow growth rate in plants originating from seed (juveniles), and
their delay in gaining fire resistance. Hence, frequent fire may prevent recruitment of
plants. Abbott (1985) found a similar result for the resprouting Banksia grandis. Auld
(1987) has shown that short fire frequencies (with <10 year intervals) are likely to reduce
the abundance of the obligate seeder Acacia suaveolens by a decline in the buried seed
bank. Consequently, any model trying to predict long-term changes in plant popu-

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

260 POST-FIRE DEMOGRAPHY IN ANGOPHORA HISPIDA

lations in relation to the fire regime should consider vegetative survival, growth, sexual
reproduction, establishment and the subsequent length of the primary Juvenile period,
as well as the longevity of adult plants.

Angophora hispida is a common understorey shrub found in heaths and woodlands on
Hawkesbury Sandstone around Sydney (Leach, 1986). It grows to 2-3m in height and 3-
4m in width and flowers from November to January (Price, 1963). Plants are usually
able to survive fire (Beadle, 1940), and regeneration occurs by a combination of ligno-
tuberous and epicormic shoots from protected bud strands, as is typical of many species
in the closely related Eucalyptus (McArthur, 1968).

Several components of the life cycle of the resprouter Angophora hispida (Sm.) Blaxell
were examined using demographic methods in an effort to link post-fire regeneration to
survival and reproduction. These components included seed production, dispersal,
seedling recruitment and seedling survival. Predictions were made of the long-term
behaviour of A. hispida populations under different fire regimes.

METHODS

Study sites were chosen in Ku-ring-gai Chase National Park and Manly Dam
Reserve, some 10-30km north of Sydney. Annual precipitation for the region is around
1300mm with the wettest period being, on average, from January to June. The average
maximum temperatures are 27°C in summer and 17°C in winter, whilst the average
minimum temperatures are 17°C and 4°C, respectively.

Eleven study sites were chosen, each with a different time since fire. A range of
times from one to 13 years was available, with replication for 1, 6 and 13 years. The sites
are briefly described in Table 1. The soils at each site are derived from Hawkesbury
Sandstone. Rainfall levels are roughly uniform across all sites throughout the study
period. Rainfall data during this study were taken from the station at Manly Dam,
approximately 1km from Site 1. Plant names follow Beadle et al. (1982).

TABLE 1
Location of study sites with time since burnt and characteristic vegetation. Vegetation type after Specht (1970)

Years since
Site fire Vegetation and notes

1 1 Open heath with occasional emergent Eucalyptus gummifera and E. haemastoma.
Understorey dominated by A. hispida, Banksia ericifolia and Casuarina distyla.

2 1 or >10 Vegetation as for site 1. This site was divided into two neighbouring populations of A.

hispida. In both populations the fire which had burnt the site had stopped in the

middle of the population, once through natural causes and once because of a small

fire trail. This left one half of each population burnt and one half unburnt for at least

10 years.

Low woodland of Angophora costata, E. gummifera, B. serrata and Xylomelum pyriforme.

Low (1-2m) shrub understorey of A. hispida, Eriostemon australasius and Dillwynia

floribunda.

Open scrub of A. hispida and B. oblongifolia.

5 5 Open heath with occasional emergent E. haemastoma. Heath dominated by A. hispida
and B. oblongifolia.

6 6 Open heath with A. hispida and B. oblongifolia.

7 6 Open heath with A. hispida, B. marginata and Petrophile pulchella.

3 7 Closed scrub of A. hispida, Leptospermum attenuatum and P. pulchella. Occasional
emergent E. haemastoma.

) 10 Low woodland of E. gummifera and E. haemastoma. Understorey dominated by A.
hispida and D. retorta.

10 13 Closed scrub with A. hispida and P. pulchella.

1] 13 Closed scrub with A. hispida and L. attenuatum.

Sw)
1SN)

PROC. LINN. SOC. N.SW., 109 (4), (1986) 1987

T. D. AULD 261

Fruit production

At each site, 20 individuals were randomly selected using a transect/baseline
method. From randomly selected points along a baseline placed adjacent to the popu-
lation to be sampled, 1m-wide transects were run into the population. A 1m interval was
randomly chosen along these transects and any individuals touching the transect in this
interval were sampled. The number of ripe fruits on each individual at the time of fruit
maturation (February 1978) was counted. Where no fruiting individuals were en-
countered, the population was searched for any flowering and fruiting plants. Where
possible, a sample of ripe fruits was harvested.

A further 10 individuals were similarly chosen and the number of living leaves on
each counted, as an indicator of growth since the last fire, at sites 1,3,4,5,6,8,9 and 10.

For each fruiting individual that was sampled three size components were
measured:

i) plant height (m);
11) girth of the main stem (cm) at 15cm above ground; and
iii) total girth of all aerial stems (cm) at 15cm above ground.

Diameter at breast height (DBH) is not a useful measure in A. hispida because of the
species shrubby, frequently multistemmed habit. Two girth measurements were in-
cluded because of the heterogeneous nature of regeneration after fire, 1.e. epicormic
and/or lignotuberous regeneration. The data were analysed using a multiple regression.

An additional 21 individuals were randomly selected at Site 1 and fruit production
on both types of regrowth i.e. epicormic and lignotuberous shoots recorded. This gave a
better estimate of the variation in fruit production and allowed a comparison of fruiting
success in relation to type of regrowth.

Predispersal seed predation

The components of each fruit harvested at Site 1 were examined to estimate the
proportion of seeds that were intact, eaten by a seed predator, or undeveloped.

Seed dispersal

Initial seed fall on the soil surface was mapped along four, 50cm-wide, transects
radiating from the centre of a cluster of A. hispida individuals.

Potential secondary dispersal by ants was examined:

1) Using 20 permanent quadrats (50 x 50cm) at Site 1 with 10 seeds of A. hispida
and 10 seeds each of Acacia linifolia and Acacia suaveolens, both known myrmecochores
(Berg, 1975; Auld, 1986). Movement of seeds by ants was observed intensively for five
days. This trial was set up during seed fall for the acacias (December) and not when
abundant A. hispida seeds were on the ground.

ii) During seed fall of A. hispida at Site 1 (March 1978), a permanent 1m? quadrat
was set up near eight individuals. Thirty A. hispida seeds, each marked with a small
white paint dot for ease of recovery, were placed in each quadrat. The fate of these seeds
was followed for a month.

Seed dormancy and viability

Samples of 32 seeds from the seed lot collected from ripe fruits at Site 1 were tested
for dormancy and viability monthly for six months. Seed collected from the soil surface
some two weeks after seed release was also tested. Seeds were placed in petri dishes on
Whatman’s Seed Test Thick filter paper with distilled water. These were placed on a
laboratory bench at room temperature. Germination was determined by the emergence
of a radicle.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

262 POST-FIRE DEMOGRAPHY IN ANGOPHORA HISPIDA

Seven replicates of 50 seeds in small hessian bags were randomly buried at a depth
of between 0-5cm in Site 1. Mesh sides on the bags allowed the penetration of water, soil
air and small soil animals, whilst preventing the movement of the large A. hispida seeds.
One bag was to be recovered each month for seven months, with the component seeds
being extracted and examined for dormancy, viability and signs of predation.

Seedling survivorship

The transect/baseline sampling method was used to locate 10 permanent quadrats
at Site 1. Within these, the locations of 143 germinating seeds were mapped. The fate of
these seeds was followed at weekly intervals for six months and then sporadically for a
further seven years. Where possible, causes of mortality were identified.

At Site 1, 30 eight-year-old plants were unearthed in August 1986 and the depth of
burial and size of the developing lignotuber were measured with a vernier caliper. The
volume of the lignotuber was estimated by assuming it was an ellipsoid. The height and
the number of live aerial stems and leaves of these plants was also measured.

RESULTS

Fruit production

Flowering and fruit set were confined to those plants which had been burnt in the
past 12 months ie. plants at Sites 1 and 2. This was most evident at Site 2, where only
half the plants had been burnt. Here burnt plants regrew extensive canopies and some
flowered, whilst unburnt plants remained largely inactive and failed to produce any
flowers or fruits. Whilst all burnt plants showed vigorous regrowth (Table 2) not all
flowered (75% of plants at site 1 and 65% of burnt plants at site 2). No extensive,
regrowth was evident at those sites not recently burnt (Table 2).

TABLE 2

Mean number of living leaves per individual

Time since

last fire mean number of leaves

Site (years) per plant (+S.E.)

1 1 2452.8 (373.1)

5 3 399.5 (107.8)

4 4 284.5 (49.5)

5 5 465.1 (173.1)

6 6 219.7(101.1)

8 7 68.1 (22.3)

9 10 49.1 (24.2)

10 13 104.6 (27.0)

The immediate post-fire growth and reproduction were evident at other sites as in-
dicated by either the presence of fruits from previous seasons (Sites 3 and 4), or typical
inflorescence structures, 1.e. terminal cymes (Sites 5-11), from past flowering.

At Site 1, significantly more fruits were produced on epicormic shoots than on
lignotuberous shoots (1 — Factor Analysis of Variance (ANOVA), P <0.005). This, in
part, reflects the greater abundance of epicormic shoots produced after the fire.
However, even where individuals produced both epicormic and lignotuberous shoots,
more fruits were found on the epicormic shoots (paired t-test 0.01 > P > 0.001). Where
the main stem had a sufficient bark thickness to survive the fire and produce epicormic

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

T. D. AULD 263

shoots, successful fruit production was virtually assured (94% of such plants set fruit).
However, where the main stem was killed and regrowth was entirely lignotuberous, the
chance of an individual successfully fruiting was greatly reduced (35%), and was con-
fined to those plants which produced a new shoot of at least 50cm in height. Most of
these plants had small fruit-crops (<100 fruits), and only one plant had a large fruit-
crop which was totally lignotuberous in origin (871 fruits).

The number of fruits produced varied greatly between individuals. At Site 1,
around half the fruiting plants sampled produced less than 100 fruits (Fig. 1), though
some plants produced up to 1500 fruits. This variation could be partially explained by
size of individuals. The linear regression of the number of fruits per fruiting plant on
girth of the main stem (Fig. 2) was significant (P < 0.001) and accounted for some 44%
of the variation in fruit-crop size. Adding the height and total girth components did not
significantly improve the regression. This is expected, as all three variables were signifi-
cantly correlated. A large amount of the variation in fruit production remained
unexplained.

Frequency (%)

co)

=
'

ep)
(2)
(=)

00s} -Lovl]]

OOlL-LOOL
OOCL-LOLL
OOElL-LOcl
OOvl-LOEL

Number of frui
Fig. 1. Number of A. hispida fruits produced after fire at Site 1.

Predispersal seed predation

At Site 1, some 463 ripe fruits were harvested, with a maximum of 10 fruits being
taken from any one plant. The majority of locules in a fruit contain an intact seed (Table
3). Seed predation in fruits was negligible (0.4% seeds lost), with occasional unidenti-
fied hymenopteran wasps reared from seeds.

TABLE 3
Extent of predispersal seed predation in A. hispida

mean (+ S.E.) % of available locules
locules/fruit 2.996 (0.013) 100
intact seeds/fruit 2.361 (0.039) 78.8
seeds/fruit lost to predators 0.011 (0.005) 0.4

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

264 POST-FIRE DEMOGRAPHY IN ANGOPHORA HISPIDA
1600

1200

800

Number of fruits/fruiting plant

400

=
20 40 60 80
Girth of main stem (cm)
Fig. 2. Relationship between fruit production and girth of main stem for A. hispida at Site 1. y = -43.9 +
12.1x.
Seed dispersal

Initial seed dispersal was restricted to less than 5m around the parent plant (Fig. 3),
with most seeds falling below the parent’s canopy. Seed movement by ants was minimal
(Table 4), indicating that A. Azspida is not myrmecochorous. Where some ant dispersal of
seeds occurred, it was only over a small distance (<50cm) and seeds may have been
moved because of their close proximity to the two known Acacia myrmecochores.

TABLE 4

Evidence for seed movement by ants

Test species % seeds remaining after 5 days(+ S.E.)
Angophora hispida 79 (5.6)
Acacia linifolia 6 (4.7)
Acacia suaveolens 9 (5.5)

For the marked seeds, some im situ seed predation by an unknown seed predator oc-
curred (20% +7%), along with seed germination (18% +6%) following heavy falls of
rain in March 1978 (300mm). A large component of the seeds could not be traced (62%
+9%). These seeds were either: 1) buried by moving sand, ii) washed away from the
sampling area during heavy thunderstorms during the sampling period or 111) removed
by seed predators or over small distances by ants. In all cases, the seeds would have ger-
minated and established following sufficient rainfall in March 1978 (see below), unless
they were destroyed by predators or buried too deeply for successful establishment. Seed
movement during storms was small (<2m) as this was impeded by vegetation and litter.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

T. D. AULD 265
350

300

250

Rainfall (mm)

200

150

100

Frequency of seeds (%)

Dee aEeO 2.0 3.0 4.0 5.0
Distance from centre of parent (m)

Fig. 3. Initial seed dispersal around A. hispida parents.

Seed dormancy and viability
All seeds showed 100% viability for up to six months after collection. There was no
indication of any dormancy mechanism.

Seeds buried in hessian bags germinated within the first month of burial presum-
ably following good rainfall in March (300mm). The remaining seed bags were
retrieved two months after burial and all seeds had germinated.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

266 POST-FIRE DEMOGRAPHY IN ANGOPHORA HISPIDA

TABLE 5
Size of eight year old plants, n = 30

Dimension x S.e. range
number of live aerial stems 1.8 0.3 1-8
number of live leaves* 8.1 0.9 2-27
plant height (mm) 140.2 8.0 40.5-220.2
depth to top of lignotuber (mm) Boll 2.1 -25-29
lignotuber length (long axis) (mm) 20.3 1.6 5.9-36.1
lignotuber width (widest point) (mm) 16.4 1.0 6.3-26.7
lignotuber width (orthog.) (mm) 13.8 1.0 4.7-24.7
lignotuber volume (mm_~) 4736 780 213-16297

@ None of the leaves sampled was of adult leaf size.

@ 2.0
S
iS
~16
e)
o
19) 4,2
E
=)
Ss
20.8
a

0.4

@) 12 24 36 48 60 Me 84
Months

Fig. 4. The survival of A. hispida seedlings from germination to 8 years of age at Site 1.

Seedling survivorship

In the field, germination was confined to a 3-week period. Mortality was highest in
the first few weeks after germination when the radicle was emerging and penetrating the
soil surface and the cotyledons were unfolded. Once the first leaves were produced sur-
vival increased and remained relatively higher for the remainder of the study. The sur-
vival of seedlings from germination until eight years of age is shown in Fig. 4. During
the first six months of seedling growth some 60% of deaths were due to desiccation, 16%
to seedling predators; 5% to burial by moving sand; and 19% to unknown agents. The
8-year-old plants were small with small lignotubers close to the soil surface (Table 5).

DISCUSSION

A. hispida is a typical resprouter (Gill, 1981) with regrowth after fire from a com-
bination of epicormic and lignotuberous shoots. It is among the fastest resprouting
species in those communities in which it occurs, and where the main stem is not killed an
extensive new canopy is formed some 6-8 months after fire. These mature plants have a
very short secondary juvenile period and are potentially able to flower in the first flower-
ing season (Nov-Feb) following a summer fire. Plants burnt in autumn to spring will

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

T. D. AULD 267

probably not flower until the second summer as regrowth will still be incomplete in the
first summer. Other resprouting species showing this same pulse of flowering in the im-
mediate post-fire period include; geophytic orchids, lilies and herbs in South Africa
(Levyns, 1966; Martin, 1966), coastal sage shrubs in California (Keeley and Keeley,
1984), Xanthorrhoea spp. and Kingia spp. in Australia (Gill and Ingwersen, 1976; Baird,
1977; Lamont and Downes, 1979). The change from a suppressed shrubby individual in
long unburnt communities, to vigorous regrowth after fire, as shown by A. hispida, has
not been recorded before in a resprouting species in Australia. The berries Vaccinium
spp. show a similar response in the U.S.A. (Gill and Groves, 1981). Some non-sprouting
plants (obligate seeders) can also grow rapidly and flower extensively in the first 18
months after fire, e.g. Acacia suaveolens (Auld, 1987).

In A. hispida, fruit production was clearly enhanced on plants resprouting via
epicormic shoots. After the 1977 fire at Site 1, some 65% of plants which resprouted by
lignotuberous growth only, failed to flower and may be juveniles. Most of these in-
dividuals were small. For these plants to reach maturity a sufficiently long inter-fire
period is needed to allow development of a stem large enough to survive fire. However, it
is important to note that one plant was able to produce a large seed-crop from a 1.5m
aerial stem produced solely from the lignotuber. This is in contrast to Banksva serrata
(Bradstock, 1985) where a minimum of 6-10 years is required for a resprouting Juvenile
plant to develop a stem of sufficient size to be fire resistant. Only plants of this size were
capable of flowering and even then the subsequent secondary juvenile period was
around two years. In B. serrata, repeated fires at less than 10 year intervals may produce,
in a proportion of the population, continually suppressed plants which never flower. In
A. hispida, the proportion of plants flowering will be higher.

Jacobs (1951) and Majer (1980) suggest that insect abundance 1s reduced at least in
the short term after a fire. Although there are no studies on how fire affects the popula-
tions of predispersal seed predators, heavy flowering and fruiting after a fire may lead to
reduced levels of seed predation through satiation of available seed predators e.g. Acacia
suaveolens (Auld and Myerscough, 1986). A species which only flowers after a fire is likely
to be both spatially and temporally irregular because of the irregular nature of fires.
This is comparable with mast seeding species (Janzen, 1976; 1978). In contrast, other
resprouting species after a post-fire secondary juvenile period, either maintain their
reproductive output throughout their life span (e.g. Banksza serrata, Bradstock, 1985;
Arctostaphylos glandulosa, Keeley and Keeley, 1977; and Protea nitida, Kruger, 1983) or show
a peak after fire with a decline after approximately 2 years (Xanthorrhoea spp., Gill and
Ingwersen, 1976; Telopea speciosissima and Lambertia formosa, Pyke, 1983) or around 10
years (Isopogon anemonifolius, Bradstock, 1985; and Protea cynaroides and P. speciosa, Kruger,
1983).

Dispersal of A. hispida seeds is minimal and seeds will remain in the boundaries of
the existing population, except perhaps near its edges. Like the closely related A. bakeri
(Auld, 1986), ant dispersal of seeds was not extensive. As well as the possibility of
decreased seed predation (Majer, 1982) and increased nutrient levels (Siddiqi e al.,
1976) the immediate post-fire environment has ample light for developing seedlings. For
A. hispida, with no seed dormancy, successful establishment and growth is largely depen-
dent upon lack of discovery by post-dispersal seed predators and the amount of available
moisture. Specht (1981), Bradstock and Myerscough (1981) and Bradstock (1985) have
shown that the seedling establishment of several proteaceous shrubs was directly depen-
dent upon soil moisture immediately following a fire. With high moisture availability,
establishment should be high, although unless moisture levels are maintained sub-
sequent mortality will be high. As seed of A. Azspida maintains its viability for up to six

PROG. LINN. SOC. N.S.W., 109 (4), (1986) 1987

268 POST-FIRE DEMOGRAPHY IN ANGOPHORA HISPIDA

months it is likely that sufficient rainfall will occur for germination. During drought
periods, post-dispersal seed predation may severely reduce the available seed.

For recruitment in A. hispida to be effective, the inter-fire period must be sufficient
to allow seedlings to reach a stage where they are fire tolerant. This will vary depending
on the intensity of the fire and the depth of burial of the lignotuber. After 8 years, seed-
lings (or juveniles) were still distinct in size from all individuals which survived the 1977
fire at Site 1. Whilst the development of the lignotuber had commenced, its size was
small and may not ensure survival in the next fire. Abbott (1985) found that 3%-year-old
plants of the resprouter Banksia grandis were small in height (11cm), whilst the length of
the long axis of the lignotuber of these plants was also small (1.04cm). He predicted that
it takes 35 years for this species to reach maturity from seed, although no estimate was
made of how large the lignotuber must be to survive fires of differing intensities. To en-
sure the continued survival of populations of A. hispida via seedling recruitment, a fire-
free period of greater than eight, and possibly many more, years may be required if the
small juveniles described here are not fire resistant. Clearly, for the long-term manage-
ment of populations of A. hispida, an investigation of the minimum fire-free period that
is required under varying fire intensities is required. Kruger (1983) has suggested that in
fynbos the primary juvenile period is usually less than eight years, whilst for chaparral it
is 8-10 years. Data from Abbott (1985), Bradstock (1985) and this study indicate that in
fire-prone Australian plant communities the primary juvenile period may be much
longer for resprouting species. This emphasises the importance of considering the mini-
mum length of the fire-free interval required to maintain populations of resprouting
species through seedling recruitment. Whilst many resprouters are vegetatively
vigorous and floriforous after fire, this cannot be directly interpreted to mean that
resprouters are capable of maintaining population levels under short fire frequencies.
The length of time required for seedlings to become fire tolerant dictates the minimum
fire interval, irrespective of any post-fire flushes in vegetative growth or flowering.

ACKNOWLEDGEMENTS

I wish to thank Warringah and Manly councils for permission to work on their
reserves. Drs R. Bradstock, K. Mullette, P,- Myerscough and N. Shepherd made helpful
comments on the manuscript. M. Ellis drew the figures.

References

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J. Bot. 33: 381-391.

AULD T. D., 1986. — Population dynamics of the shrub Acacia suaveolens (Sm.) Willd.: Dispersal and the
dynamics of the soil seed bank. Aust. J. Ecol. 11: 235-254.

——.,, 1987. — Population dynamics of the shrub Acacia suaveolens (Sm.) Willd.: Survivorship throughout the

life cycle, a synthesis. Aust. J. Ecol. 12: 139-151.
, and MYERSCOUGH, P. J., 1986. — Population dynamics of the shrub Acacia suaveolens (Sm.) Willd.:
Seed production and predispersal seed predation. Aust. J. Ecol. 11: 219-234.

BAIRD, A. M., 1977. — Regeneration after fire in King’s Park, Perth, Western Australia. /. Roy. Soc. WA. 60:
1-22.

BEADLE, N. C. W., 1940. — Soil temperatures during forest fires and their effect on the survival of vegetation.
J. Ecol. 28: 180-192.

——, Evans, O. D., and CAROLIN, R. C., 1982. — Flora of the Sydney Region. 2nd Ed. Sydney: A. H. & A. W.
Reed.

BENSON, D. H., 1985. — Maturation periods for fire-sensitive shrub species in Hawkesbury Sandstone vege-
tation. Cunninghamia 1: 339-349.

BerG, R. Y., 1975. — Myrmecochorous plants in Australia and their dispersal by ants. Aust. J. Bot. 23;
475-508.

BRADSTOCK, R. A., 1985. — Plant population dynamics under varying fire regimes. Sydney: University of
Sydney, Ph.D thesis, unpubl.

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, and MYERSCOUGH, P. J., 1981. — Fire effects on seed release and the emergence and establishment of
seedlings in Banksza ertcifolia L.f. Aust. J. Bot. 29: 521-32.

GILL, A. M., 1981. — Adaptive responses of Australian vascular plant species to fire. Jn: GILL, A. M.,
Groves, R. H., and NOBLE, I. R., (eds), Frre and the Australian Biota: 243-272. Canberra: Australian
Academy of Science.

, and GROVES, R. H., 1981. — Fire regimes in heathlands and their plant ecological effects. In: SPECHT,
R. L., (ed.), Ecosystems of the World 9B Heathlands and Related Shrublands: 61-84. Amsterdam: Elsevier
Scientific Publ. Co. :

, and INGWERSEN, F., 1976. — Growth of Xanthorrhoea australis R.Br. in relation to fire. J. Appl. Ecol. 13:
195-203.

Jacoss, M.R., 1951. — The growth and regeneration of eucalypts. J. Aust. Inst. Ag. Sct. 17: 174-183.

JANZEN, D. H., 1976. — Why bamboos wait so long to flower. Ann. Rev. Ecol. Syst. 7: 347-391.

——., 1978. — Seeding patterns of tropical trees. Jn’ TOMLINSON, P. B., and ZIMMERMAN, M. H., (eds),

Tropical Trees as Living Systems: 83-128. New York: Cambridge University Press.
KEELEY, J. E., and KEELEY, S. C., 1977. — Energy allocation patterns of a sprouting and a nonsprouting
species of Arctostaphylos in the California chaparral. Am. Midl. Nat. 98: 1-10.
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and seedling strategies. Am. Midl. Nat. 99: 142-161.
KRUGER, K. J., 1983. — Plant community diversity and dynamics in relation to fire. Jn: KRUGER, F. J.,
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LAMONT, B. B., and DOWNES, S., 1979. — The longevity, flowering and fire history of the grasstrees Xanthor-
rhoea preissti and Kingia australis. J. Appl. Ecol. 16: 893-899.

LEACH, G. J., 1986. — A revision of the genus Angophora (Myrtaceae). Telopea 2: 747-779.

LEvYNS, M. R., 1966. — Haemanthus canaliculatus, a new fire-lily from the western Cape Province. Sth. Afr. J.
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McCARTHUR, A. G., 1968. — The fire resistance of eucalypts. Proc. Ecol. Soc. Aust. 3: 83-90.

Majer, J. D., 1980. — Report on a study of invertebrates in relation to the Kojonup fire management plan.
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——, 1982. — Ant plant interactions in the Darling Botanical District of Western Australia. In: BUCKLEY,
R. C., (ed.), Ant-Plant Interactions in Australia: 45-62. The Hague: Dr. W. Junk Publishers.

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PRICE, D., 1963. — Calender of flowering times of some plants of the Sydney district, possibly associated with
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PyKE, G. H., 1983. — Relationship between time since the last fire and flowering in Télopea speciosissima R.Br.
and Lambertia formosa Sm. Aust. J. Bot. 31: 293-296.

Sippiqi, M. Y., CAROLIN, R. C., and MYERSCOUGH, P. J., 1976. — Studies in the ecology of coastal heath in
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SPECHT, R. L., 1970. — Vegetation. Jn: LEEPER, G. W., (ed.), The Australian Environment: 44-67. 4th Ed.
C.S.1.R.O.

——.,, 1981. — Responses to fires of heathlands and related shrublands. Jn: GILL, A. M., GROVES, R. H., and

NOBLE, I. R., (eds), Fire and the Australian Biota: 395-415. Canberra: Australian Academy of Science.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

ty ies

)

A Review of the Biology of Acacia suaveolens (Smith)
Willd. (Mimosaceae)

DAVID A. MORRISON

(Communicated by P. J. MYERSCOUGH)

Morrison, D. A. A review of the biology of Acacia suaveolens (Smith) Willd.
(Mimosaceae). Proc. Linn. Soc. N.S.W. 109(4), (1986) 1987: 271-292.

Published and original data on a number of aspects of the biology of Acacia suaveolens
are presented, including: taxonomy, geographical distribution, climatic, topographic, and
altitudinal limitations, substratum, communities, gregariousness; response to biotic fac-
tors; performance in various habitats, effect of frost, drought, and waterlogging, mor-
phology, chromosomes, physiology, biochemistry, perennation and reproduction,
phenology, flowering and pollination, seed production and dispersal, viability of seeds and
germination, seedling morphology, mycorrhiza, animal feeders and parasites, plant
diseases and parasites, and history and conservation status.

D, A. Morrison, Department of Applied Biology, New South Wales Institute of Technology, P.O. Box
123, Broadway, Australia 2007, manuscript received 9 December 1986, accepted for publication 22
April 1987.

INTRODUCTION

In 1941 the British Ecological Society began publishing a series of papers on the ‘Bio-
logical Flora of the British Isles’ (Anon., 1941), a series that is still being actively published.
This series was intended to be an introductory reference source of published and unpub-
lished data on various aspects of the biology of the plant species occurring in Great Britain.

No similar series has been undertaken for Australian plants, partly, at least, because
very few data exist for most of the species. However, extensive data do exist for a number
of the more common species, particularly of the genera Eucalyptus and Acacia. This paper
is an attempt to summarize the published data on one particular species, Acacia suaveolens
(Smith) Willd., that is common in south-eastern Australia and for which considerable data
do exist. The data are arranged in the format used by the British Ecological Society in pub-
lishing their Biological Flora, and I have attempted to collate whatever data exist concern-
ing A. suaveolens for each of the aspects covered in that series (see Anon., 1941), and to
contribute original data for aspects for which published data are not available. The refer-
ences cited are mainly a subset of the bibliography of 186 sources of data on A. suaveolens
listed in Morrison (1986).

DESCRIPTION AND TAXONOMY

Acacia suaveolens (Smith) Willd., Sp. Pl. 4:1050 (1806)

[Mimosa suaveolens Smith, Trans. Linn. Soc. Lond. 1:253 (1791) non Salisb. (1796); Mimosa
obliqua Lam., J. Hist. Nat. Paris 1:89 (1792) non H. H. Wendl. (1798); Mimosa ambigua Salisb.,
Prodr. Stirp. 325 (1796); Mimosa angustifolia Jacq., Pl. Hort. Schoenbr. 3:74 (1798); Acacia
angustifolia (Jacq.) H. H. Wendl., Comm. Acac. 34 (1820); Acacia suaveolens var. platycarpa DC,
Prodr. 2:453 (1825); Phyllodoce suaveolens (Smith) Link, Handbuch 2:133 (1831); Phyllodoce
angustifolia (Jacq.) Link, Handbuch 2:133 (1831); Hecatandra suaveolens (Smith) Raf., Sylva Tellur.
120 (1838) ]

Subgenus Phyllodineae (DC) Ser.; Section Phyllodineae DC; Subsection Racemosae (Benth.)
Maiden

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

Die. ACACIA SUAVEOLENS

Usually erect, slender, glabrous, little-branched, open shrub to 2(-3)m high; bark smooth,
(bluish-)green. Branchlets terete below but acutely triquetrous above due to decurrencies,
sometimes almost flattened, glabrous, brownish-green or sometimes glaucous, new growth
often pinkish. Phyllodes alternate, erect, glaucous, coriaceous, glabrous, flat, straight or
rarely slightly falcate, narrow-oblong to linear-lanceolate, acute or obtuse, acuminate or
mucronate, narrowed towards the base, (5-)7-12(-20)cm long, (2-)3-7(-10)mm wide, (9-)13-
27(-35) times as long as broad, one-nerved more or less central, margins thickened,
yellowish-brown; pulvinus 1-2mm long; small, flat, elongated, non-porate gland (1-)2-3
(-4)mm from base of phyllode, sometimes another gland at tip. Jnflorescence of globular heads
5-8mm in diameter, each of (3-)4-7(-12) flowers, in glabrous, axillary, often crowded, (5-)6-
8(-12)-branched racemes; rhachis 1-2(-3)cm long, slender; peduncles 2-5mm long, slender;
flower heads andromonoecious, usually twice as many male as hermaphrodite flowers; be-
fore development, racemes enclosed by imbricate, scarious, fimbriate, ovate, obtuse, pale
yellow-brown, pink-tipped bracts to 2.5mm long, crowded at base of axis, with larger ones
subtending individual peduncles, all deciduous before anthesis. Flowers actinomorphic,
(4-)5(-6)-merous, creamy to pale lemon-yellow, protogynous, sweetly-scented. Sepals free,
thin, linear-spathulate, acuminate, glabrous with a few hairs at the tip, 1.2-1.3mm long.
Petals thin, free, ovate-oblong, glabrous, 1.7-3mm long, 0.7-1mm wide, less than twice as
long as the calyx. Stamens (35-)40-55(-80), 3-5mm long; anthers small, almost round, bi-
lobed with 4 loculi per lobe. Ovary unilocular, central, superior, yellow-brown, oblong, later-
ally compressed, acuminate, glabrous, with (4-)5-7(-8) anatropous basally-attached ovules.
Style yellow-brown, filiform, bent, up to twice as long as the stamens; stigma simple, ter-
minal, acute. Pollen grains yellow, non-reticulate, 4-porate, pores placed towards the angles
of the grains, 4-furrowed; grains aggregated into polyads of 16, long equatorial diameter
of the polyad 52-64um, 1 polyad per locule. Legume stalked, glaucous, purplish-red at
fertilization, turning bluish-green, often reddish-brown over the seeds, brown when open
at maturity, glabrous, coriaceous, pruinose, (elliptic-)oblong, obtuse, apiculate, laterally
compressed but slightly raised over the seeds, 2-4(-5)cm long, (10-)12-20mm wide, twice as
long as wide, margins thickened. Seeds smooth, shiny, dark brown to brownish-black (rarely
maroon), transverse, (elliptic-)oblong, 5-8mm long, 2.5-4.5mm wide, 2-3 times as long as
wide; areole closed, 3-4.5mm long, 0.6 times the length of the seeds; funicle 1.5-2mm long,
filiform till nearly mature then thickened into a slightly oblique, fleshy, 1-3(-4)-folded aril
covering the hilar end, same colour as the seeds; no albumen present.

Variable in erectness, height, phyllode axillary angle, phyllode shape and size, and
number of flowers per inflorescence, but most of this has no apparent genetic basis. In the
Myall Lakes area, populations of plants with very narrow phyllodes (< 2mm wide;
Armitage, 1977) are common in the sclerophyllous forest community on the Holocene
sand, and this may have a strong genetic component. In the Grampians, populations with
phyllodes held conspicuously close to the stem and a small rootstock can be found in the
deeper sands on the west-facing slopes. Along the New South Wales coast north of Sydney,
plants with a prostrate, spreading habit and much broader phyllodes are often found on
thin soils on exposed headlands. Elliot and Jones (1982) also report that a form with a cream
band on each side of the phyllode midvein and flowers with a deeper yellow is often culti-
vated, but that it ‘must be propagated from cuttings to retain the variegation.

GEOGRAPHICAL DISTRIBUTION
A. suaveolens is endemic to the southeastern coast of the Australian mainland, around
the coast of Tasmania, and on the larger off-shore islands (Fig. 1). It is generally restricted
to the coast, although it does occur inland, notably in the Sydney Basin, in the Grampians,
and at the South Australia-Victoria border.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

D. A. MORRISON 273

Fig. I. Known distribution of Acacia suaveolens. Each outline represents at least one record on the national 0.5° x
0.5° grid. Data from dried specimens held at AD, BRI, CANB, CBG, HO, MEL, NSW, PERTH, and SYD (codes after
Holmgren et al., 1981).

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

974 ACACIA SUAVEOLENS

In the literature, populations have also been recorded on the Victorian coast east of
the South Australian border (Churchill and de Corona, 1972; Costermans, 1981), east of
the Grampians (Churchill and de Corona, 1972; Beauglehole, 1980b; Maslin and Pedley,
1982), in the Albury district (Jacobs and Pickard, 1981; Maslin and Pedley, 1982), and in
the northern tablelands of New South Wales (Simmons, 1981; Maslin and Pedley, 1982),
but these records have not been confirmed and are not supported by herbarium voucher
specimens (Beauglehole, 1980a; Morrison e¢ al., 1983).

CLIMATIC, TOPOGRAPICAL AND ALTITUDINAL LIMITATIONS

As a result of its large latitudinal (24°26’S to 43°15’S) and altitudinal (0-950m)
range, A. suaveolens experiences a wide range of climatic extremes (Table 1), and does not
appear to show any particular climatic preferences. It also does not exhibit any climatically-
determined slope preferences. Burrough et al. (1977) have found plants growing at both ex-
tremes of a strong gradient of sites varying from a strongly positive to a balanced precipi-
tation/evaporation budget over a short distance on the New South Wales central coast.

TABLE 1

Long-term temperature and precipitation extremes within the geographical distribution of Acacia suaveolens
Data from Australian Bureau of Meteorology (1975a-e) for a total of 163 meteorological stations

Climatic parameter Value Meteorological station
Mean annual maximum temp.:— max. 28.5°C Tabbimoble S.F., N.S.W.
min. 13.4°C Tasman Is. Lighthouse, Tas.
Highest mean monthly maximum temp.: — max. 31.6°C Tabbimoble S.F., N.S.W.
min. 10.2°C Tasman Is. Lighthouse, Tas.
Mean annual minimum temp.:— max. WAC; Tabbimoble S.F., N.S.W.
min. Bo) AC Geeveston (Forestry), Tas.
Lowest mean monthly minimum temp.: — max. 22.8°C Tabbimoble S.F., N.S.W.
min. 0.0°C Geeveston (Forestry), Tas.
Mean annual rainfall: — max. 1814 mm Alstonville Res. Stn., N.S.W.
min. 538 mm Geelong (S.E.C.), Vic.
Mean annual no. raindays: — max. 235 Strahan (Vivian St), Tas.
min. 57 Waterloo, Old.

Populations are most commonly found within a few kilometres of the coast, and below
an altitude of 300m. The only places where this species is recorded above 500m (to 600m
in the McPherson Range, to 950m in the Blue Mountains, to 650m in the Budawang
Range) are where the upland sandstone soils of the Great Dividing Range are contiguous
with the sandy coastal plain.

SUBSTRATUM

On the mainland, A. suaveolens is most commonly found on the coastal Quaternary
sands. The northern boundary of the distributional range is near the northern boundary
of the lime-free sandy and sandy-loam soils that form the dominant soil type along the
eastern Australian coast. However, coastal populations are occasionally reported from the
perimeter of clay soils over sandstone (e.g. Webb, 1981); and on Wilsons Promontory the
species is also found as a rare occurrence on the perimeter of sandy soils over Devonian
granite (Gillham, 1960; Ashton and Webb, 1977).

Inland along the eastern coast, the species occurs on sandy soils over Jurassic sand-
stone in the McPherson Range, over Triassic sandstone in the Sydney Basin, over De-
vonian sandstone in the Budawang Range, and over Ordovician sandstone near Orbost.
It is not found on the inland sandstone areas which are isolated from the coastal sandy soils.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

D. A. MORRISON DHS

It has also been recorded from acid ‘Tertiary rhyolite on the Lamington Plateau (McDonald
and Elsol, 1984) and on Mt Coolum (Sharpe and Batianoff, 1985).

Between Lorne and Warrnambool, populations are found on inland sandy soils over
Tertiary sandstone, rather than on the uplifted Cretaceous shale of the Otway Range
adjacent to the coast. The species does not occur on the coastal soils derived from the Ter-
tiary limestone and Quaternary volcanics west of Warrnambool, and the isolated popu-
lations in the Grampians occur on Quaternary sand and on sandy soils over Devonian
sandstone. Near the Victoria-South Australia border, populations are found on the inland
siliceous Quaternary sands and sandy soils over ‘Tertiary sandstone, but do not occur on the
calcareous Quaternary sands nearer to the coastline.

In Tasmania, this species is common on sandy soils over the many sandstones (e.g.
Silurian, Permian, Triassic, Tertiary) which form most of the northern and eastern coast-
line, but it only occurs sporadically on the ‘Tertiary and Middle Proterozoic sandstones
along the northwestern coastline. On the Tasman Peninsula, it is also found on thin sandy
soils over Jurassic dolerite (Kirkpatrick, 1977b). In the Furneaux Group, plants occur on
sandy soils over both Silurian sandstone and Devonian granite, as well as on granitic col-
luvium (Kirkpatrick, 1977b); and on King Island, plants are found on Middle Proterozoic
sandstone soils.

A. suaveolens has been recorded from the following soil groups: — lithosols (Ashton and
Webb, 1977), siliceous sands (Parsons, 1966; Firth, 1969; Durrington, 1977; Kirkpatrick,
1977a, 1977b; Benson and Fallding, 1981; Forbes et al., 1982; Opie et al., 1984), earthy sand
(Buchanan and Humphreys, 1980), yellow earths (Benson and Fallding, 1981), humus
podzols (Myerscough and Carolin, 1986), and peaty podzols (Parsons, 1966). However, it
is most commonly found on siliceous sand podzols, of various colours and stages of
differentiation: e.g. poorly-developed (Kirkpatrick, 1973; Myerscough and Carolin, 1986),
moderately-developed (Clark, 1975; Thatcher and Westman, 1975), well-developed
(Groves and Specht, 1965; Kirkpatrick, 1975; Ingwersen, 1976; Ashton and Webb, 1977;
Burrough et al., 1977; Clifford and Specht, 1979; Buchanan and Humphreys, 1980).

These soils may be very shallow (e.g. Cambage, 1923; Petrie, 1925; Hannon and
Evans, 1963; Kirkpatrick, 1977b; Auld and Myerscough, 1986) but are usually quite deep
(e.g. Groves and Specht, 1965; Ingwersen, 1976; Durrington, 1977; Kirkpatrick, 1977b;
Clifford and Specht, 1979; Opie et al., 1984; Myerscough and Carolin, 1986). The soils are
predominantly freely-draining, but are occasionally seasonally waterlogged or per-
manently moist (see below). The surface pH has been recorded from 4.3-6.9 (Davis, 1941a;
1941b; Gillham, 1960; Parsons, 1966; Siddiqi et al., 1972; Burrough et al., 1977; Kirkpatrick,
1977b; Buchanan and Humphreys, 1980). The soils are extremely infertile, being relatively
more fertile in the wetter habitats. The surface loss-on-ignition varies from 0.9-30.0%
(Pidgeon, 1938; Davis, 1941a; 1941b; Ingwersen, 1976; Burrough et al., 1977; Myerscough
and Carolin, 1986); and total nitrogen ranges from 0.02-0.18% (Hannon, 1956; Siddiqi et
al., 1972; 1976; Myerscough and Carolin, 1986). ‘Total phosphorus varies from 0.001-0.01%
(Beadle, 1962; Parsons, 1966; Myerscough and Carolin, 1986), with ‘available’ water-
soluble phosphorus ranging from 0.0002-0.0013 % (Siddiqi et al., 1972; 1976; Ingwersen,
1976). The exchangeable calcium 1s reported to be in the range 0.52-4.0 meq.%, the ex-
changeable potassium from 0.06-0.95 meq.%, the exchangeable magnesium from 0.2-1.77
meq.%, and the exchangeable sodium from 0.05-0.95 meq.% (Siddiqi et al., 1972; 1976;
Ingwersen, 1976; Burrough et al., 1977; Myerscough and Carolin, 1986).

A. suaveolens grows well in the laboratory in a range of soil types with higher fertility
levels than those on which it is normally found in the wild (Beadle, 1962); and the absence
of plants from higher fertility soils has been ascribed to an inability to compete with faster-
growing species under these conditions (Beadle, 1962).

PROC. LINN. SOG. N.S.W., 109 (4), (1986) 1987

276 ACACIA SUAVEOLENS

COMMUNITIES

A. suaveolens can be regarded as a short-lived pioneer species (Clemens and Franklin,
1980), and it often occurs in early successional communities (but not in the early exposed
stages) (Pidgeon, 1938; 1940) or in early phases of regeneration cycles. As a result, the
presence of the species in a community 1s very dependent on the past history of disturbance,
especially in relation to fire. The species has been recorded from a wide variety of com-

munities, including forests, woodlands, shrublands, heaths, and sedgelands (Table 2).

TABLE 2

Plant communities in which Acacia suaveolens has been recorded

Alliance* Suballiance* Structural form ~* Area Referencef
Eucalyptus pilularis a) Eucalyptus pilularis woodland Bulli, N.S.W. (9)
b) Eucalyptus pilularis- open-forest Nth Stradbroke Is., Old. (31)
Eucalyptus intermedia- —_ open-forest Moreton Is., Qld. (10)
Eucalyptus stderophylla
c) Eucalyptus pilularis- open-forest Myall Lakes, N.S.W. (2), (21)
Angophora costata open-forest Sydney, N.S.W. (15)
Eucalyptus botryoides low open-forest Sydney, N.S.W. (5), (15)
woodland Sydney, N.S.W. (26)
low woodland Sydney, N.S.W. (4)
Eucalyptus gummifera- a) Eucalyptus gummifera- _ open-forest Sydney, N.S.W. (23)
Eucalyptus racemosa- Eucalyptus racemosa- low open-forest Sydney, N.S.W. (5), (23)
Eucalyptus sieberi Angophora costata low open-forest Barren Grounds, N.S.W. (6)
woodland Sydney, N.S.W. (2), (5), (15)
(23), (26)
woodland Barren Grounds, N.S.W. (6)
woodland Jervis Bay, N.S.W. (17)
b) Eucalyptus sieberi- open-forest Sydney, N.S.W. (3), (15)
Eucalyptus piperita- open-forest Fitzroy Falls, N.S.W. (6)
Eucalyptus racemosa low open-forest Bulli, N.S.W. (8)
low open-forest Macquarie Pass, N.S.W. (11)
c) Eucalyptus eximia- woodland Sydney, N.S.W. (3)
Eucalyptus punctata woodland Bulli, N.S.W. (9)
Eucalyptus globoidea open-forest East Gippsland, Vic. (13)
woodland East Gippsland, Vic. (13)
Eucalyptus baxtert open-forest South Australia (29)
low open-forest South Australia (32)
woodland Wilsons Promontory, Vic. (1)
low woodland Wilsons Promontory, Vic. (24)
Eucalyptus amygdalina open-forest Schouten Is., Tas. (16)
woodland Tasman Peninsula, Tas. (18)
Banksia aemula low open-forest Nth Stradbroke Is., Old. (31)
woodland Sydney, N.S.W. (26)
open-woodland Nth Stradbroke Is., Qld. (30)
tall shrubland Moreton Is., Old. (10)
tall open-shrubland Nth Stradbroke Is., Old. (7)
tall open-shrubland Moreton Is., Qld. (10)
closed-heath Moreton Is., Qld. (10)
closed-heath Myall Lakes, N.S.W. (21)
closed-heath Sydney, N.S.W. (27), (28)
Banksia ercifolia open-scrub Sydney, N.S.W. (5)
closed-heath Sydney, N.S.W. (28)
open-heath Macquarie Pass, N.S.W. (11)
Banksia oblongtfolia closed-heath Sydney, N.S.W. (23)
open-heath Myall Lakes, N.S.W. (21)
open-heath Sydney, N.S.W. (27)
Banksia robur sedgeland Sydney, N.S.W. (5), (15)
Banksia marginata closed-heath Wilsons Promontory, Vic. (14)
closed-heath Rocky Cape, ‘Tas. (12)
closed-heath Ocean Beach, Tas. (19)
open-heath Schouten Is., Tas. (16)
Leptospermum myrsinoides closed-heath Wilsons Promontory, Vic. (1)

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

D. A. MORRISON Dae,

TABLE 2 (Contd.)

Alliance* Suballiance* Structural form * Area Referencet

Leptospermum scoparium low open-shrubland Western Port, Vic. (22)
closed-heath Flinders Is., Tas. (20)

Casuarina littoralis- closed-heath Northern coastal N.S.W. (2)

Banksia oblongifolia

Casuarina distyla closed-heath Sydney, N.S.W. (2)
closed-heath Mt. Wilson, N.S.W. (25)
closed-heath Bowen Is., N.S.W. (17)
open-heath Sydney, N.S.W. (23)

Melaleuca squarrosa open-scrub Wilsons Promontory, Vic. (24)
open-heath Wilsons Promontory, Vic. (24)

* after Beadle (1981)

+ after Specht (1970)

} Sources of data: — (1) Ashton and Webb (1977); (2) Beadle (1981); (3) Benson (1981a); (4) Benson (1981b); (5)
Benson and Fallding (1981); (6) Burrough et al. (1977); (7) Clifford and Specht (1979); (8) Davis (1941a); (9)
Davis (1941b); (10) Durrington (1977); (11) Fallding and Benson (1985); (12) Firth (1969); (13) Forbes et al. (1982);
(14) Gillham (1960); (15) Hannon (1956); (16) Harris and Kirkpatrick (1982); (17) Ingwersen (1976); (18) Kirk-
patrick (1973); (19) Kirkpatrick (1977a); (20) Kirkpatrick (1977b); (21) Myerscough and Carolin (1986); (22)
Opie et al. (1984); (23) Outhred et al. (1985); (24) Parsons (1966); (25) Petrie (1925); (26) Pidgeon (1940); (27)
Siddiqi et al. (1972); (28) Siddiqi et al. (1976); (29) Specht (1972); (30) Specht et al. (1977); (31) Thatcher and
Westman (1975); (32) Whibley (1980)

The forest communities are usually found only on deep coastal sand masses, and have
an open, heathy understorey. The woodlands also have a heath rather than a scrub under-
storey, and usually occur on the shallower soils. The heath communities vary from open to
closed, and include both dry and wet heaths. Plants also occur occasionally in ground-water
heaths, and sporadically around the edges of swamps in the northern areas (e.g. White,
1945; Hannon, 1956). The species is not found in closed coastal communities such as the
exposed Leptospermum laevigatum and Acacia sophorae closed-scrubs which front many of the
beaches, occurring instead in the more open woodlands or heaths slightly further inland
(e.g. Giliham, 1960). Plants do, however, occasionally occur in exposed headland heaths on
thin sandy soils over sandstone along the New South Wales coast (Beadle, 1981) and over
dolerite on the Tasman Peninsula (Kirkpatrick, 1977b), and also on leached sands in ex-
posed situations within the salt spray zone along the northern and eastern coasts of Tas-
mania (Kirkpatrick, 1977b). The species also does not appear in the coastal
closed-grasslands, but plants are occasionally found in understoreys with clumped grasses.

The community dominants vary with the geographical area (Table 2) and substratum.
In Queensland and northern New South Wales on low fertility soils, A. suaveolens is found
in Eucalyptus intermedia open-forests, and also in the Banksia aemula low open-forests and tall
open-shrublands on the deep sand coastal islands. On more fertile soils, but not usually in
valleys, it occurs in Eucalyptus pilularis open-forests.

On the New South Wales central coast, it is found in Eucalyptus botryoides low open-
forests or woodlands in the higher fertility areas on the hind-dunes, and in FE. pilularis open-
forests and low open-forests on the deep coastal sand masses and higher-fertility inland
soils. More commonly, it occurs on less fertile soils in low open-forests or woodlands domi-
nated by Eucalyptus gummifera (in wetter areas), Eucalyptus siebert (further south), Eucalyptus
racemosa (at higher altitudes), Eucalyptus piperita (in more fertile spots), and Angophora costata
(ubiquitous). Itis also common in B. aemula low open-forests, heaths, and shrublands on
sand throughout the New South Wales coast. It is also common on sand and sandstone in
dry heaths of Banksia erncifolia and Casuarina distyla, and with Casuarina littoralis on exposed
sandy headlands. In wetter heaths, it occurs with Banksza oblongifolia, and occasionally with
Banksia robur.

PROC. LINN. SOC. N.S.W,, 109 (4), (1986) 1987

278 ACACIA SUAVEOLENS

In Victoria, the open-forests and woodlands are dominated in the east by Eucalyptus
globoidea (on more fertile soils) and E. szeberz (on less fertile soils), and in the west (and also
in South Australia) by Eucalyptus baxter: (on the less fertile soils) and Eucalyptus obliqua (on
the more fertile soils). In eastern Victoria, the species also occurs in Leptospermum myrsinoides
heaths, and, in wetter more saline areas, in heaths and thickets dominated by Melaleuca
squarrosa. On soils derived from granite, plants are occasionally found around the edges of
Banksia marginata heaths.

In Tasmania, the species is very common in B. marginata heaths, especially on sand. It
is also found in woodlands and open-forests of Eucalyptus amygdalina, often with Eucalyptus
viminalis and Eucalyptus tenuiramis in more fertile areas. On the west coast, it is also a rare
occurrence in open-scrub formations of Eucalyptus nitida and Eucalyptus ovata in low-fertility
sand deposits, often in isolated patches among low heaths on exposed headlands.

Within the communities, A. suaveolens may be associated with a wide range of species.
For example, in eucalypt forests along the southern coast of New South Wales and east
coast of Victoria its occurrence has been found to correlate positively with 27 angiosperm
species, notably Eucalyptus gummifera, Leptospermum attenuatum, Caustis flexuosa, and

Ricinocarpos pinifolius (Table 3).

PAB ICES

Percentage frequency of occurrence with (+) and without (—) Acacia suaveolens of some angiosperm plants associated with A.
suaveolens at three sites covering two eucalypt alliances in which A. suaveolens is commonly found
Data from Miller (1972), Ingwersen (1976), and Forbes et al. (1982) respectively

E. gummifera — E. racemosa — E. sieberi Alliance E. globoidea Alliance
Royal National Park Jervis Bay Territory East Gippsland
Species ai ee ibe CG) hae (+) A aaa

[No. quadrats] [ 9] [80] i [23] [52] [34] ae pal]
Acacia oxycedrus —_ — a — — 29 7 940 **
Actinotus minor 0 1 35 12 DAY = =
Banksia ericifolia 22 10 39 15 Dells) c = =
Banksia serrata 56 84 65 42 4.96 * 79 69
Bossiaea ensata = = 48 aL} = =
Cassytha glabella 33 16 26 21 74 49 Bl +
Caustis flexuosa 78 31 (Aouad, rh) 13 oe aot 24 28
Correa reflexa = = 26 8 65 38 O87 *
Dampiera stricta = = 61 S/o ee OS 83
Darwinia taxtfolia — = 30 10 RA) — =
Empodisma minus — _ = — 29 8 1:82, Ses
Epacris microphylla 22 1 39 Ge eS 2 ee eels — =
Eucalyptus gummifera 78 Bee SS) ® 74 38 Sh 2a 21 14
Eucalyptus racemosa = = 43 21 Be) — —
Gompholobium huegelu — = — — 32 10 Gil)
Hardenbergia violacea 44 13 is tO 23} Te 17 25 = =
Hibbertia fasciculata — = _ ~ 29 8 1h °°
Hypolaena fastigata = = 4 6 32 SiG 74.9 icra
Isopogon anemonifolius 11 10 35 8 Bolen ees = =
Leptospermum attenuatum 67 MS ORS) 2 BY OO SAG asa eee 23 O08
Leucopogon collinus = = = _ 62 13} IS). (8333,
Lycopodium fastigatum _ = 30 12 Sel — _
Monotoca scoparia = = 13 6 53 hy SO) FX0) ">
Persoonta caleyi = = 43 Spy, VTS KAO leas sie = =
Platysace lineartfolia 89 68 39 10 MS) Na sarees _ =
Ricinocarpos pinifolius 22 40 35 12 Qo 59 24271272 Ones

Themeda australis — - 13 44 26 11 SEO Dk aks

+ Yalues indicate statistical association between the species and A. suaveolens at that site. Lack of any details in-
dicates that the association was NS at P<0.05, or that it was not tested because one of the four values in the
2 x 2 contingency table was <1 or more than one was <5. * 0.01 <<P<0.05, ** 0.001 <P<0.01, *** P<0.001.
4 Missing data indicate that the species was not recorded at that site.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

D. A. MORRISON 279

GREGARIOUSNESS

Population size and density vary markedly with the past history of disturbance by fire.
Monospecific stands of up to 250 plants/m? have been recorded in mature recently-burnt
populations, although population density is usually in the range 0.8-6 plants/m? (Auld,
1984). In older less-disturbed habitats, plants are usually widespread but not abundant. As
the plants are relatively short-lived and do not normally regenerate without fire (Auld and
Myerscough, 1986; Auld, 1987), density decreases with age of the population.

RESPONSE TO BIOTIC FACTORS

Plants without rootstocks are destroyed by even the mildest fires. However, these fires
render the seed coat permeable to water, and the seeds in the soil then imbibe and ger-
minate if suitable rains fall (Bradstock, 1981; Auld, 1986c). Thus, if plants are old enough
to have produced seed, the species will readily regenerate strongly immediately following
a fire (e.g. Specht, 1975; Siddiqi et al., 1976; Harrold, 1979); and A. suaveolens is often the
major understorey species (commonly with Pterzdium esculentum and Imperata cylindrica) in
communities which have been frequently burnt.

This species does not respond well to disturbances other than fire. It does not regener-
ate following disturbance by sand mining without being treated with fertilizer (Clark, 1975;
Thatcher and Westman, 1975), but it does compete well with exotic grass cover after this
disturbance (Thatcher and Westman, 1975). Gillham (1960) has noted that this species is
characteristic of the unaffected areas of sea-bird rookeries but does not occur in areas which
have been dug or trampled by the birds. Similarly, Yates (1976) found that it does not
regenerate well in easements under electrical transmission lines, attributing this to con-
tinued disturbance from maintenance, a higher water table, and the use of herbicides.

PERFORMANCEIN VARIOUS HABITATS

The size which plants attain varies greatly between habitats, as does the size at which
they begin to flower and the number of inflorescences and fruits produced per year. Death
rates vary between sites, apparently depending on the rate at which the soil dries out during
summer (Auld, 1987), being greater in thin sandy soils compared with deep sands. Auld
(1984) has also reported between-site variability in seed weight and seed viability.

Seed production is reduced through predation by the weevil Melanterius corosus, and the
extent of this predation varies markedly between sites. Auld (1983) and Auld and Myer-
scough (1986) found losses to vary from 10-61%, with a mean of 47%, at six sites during
several fruiting seasons. Populations which suffer from this weevil predation have only been
found north of Jervis Bay in New South Wales; and the predation is usually more prevalent
in larger populations, with scattered populations of only a few individuals commonly being
free from predation.

Leaf area does not vary (range 8.0-8.3 cm*) when grown in the laboratory in a range
of soil types (Beadle, 1962), but total dry weight of the plants (as well as root nodulation) is
reduced when plants are grown in swamp soils as opposed to the lower-nutrient eucalypt
forest soils (Hannon, 1956).

In cultivation, plants will grow in light from filtered sun to full sun, but not in semi
shade or full shade (Elliot and Jones, 1982). Response to salt winds and exposure to full salt
spray is reported to be variable (Allen and Allen, 1981; Elliot and Jones, 1982), and plants
vary in habitat from sheltered leeward dune slopes to fully exposed headlands.

EFFECT OF FROST, DROUGHT, AND WATERLOGGING

The altitudinal and large latitudinal range of A. suaveolens suggests that this species can
tolerate considerable exposure to frost, and Simmons (1966) lists 18-month-old plants as

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

280 ACACIA SUAVEOLENS

surviving temperatures as low as -7°C in late June in Tasmania without frost damage.

A. suaveolens is particularly common in dry soils, which indicates that this species also
has considerable tolerance to drought. However, plants show a much higher death rate
during summer than at other times of the year, and Auld (1987) ascribes this to the con-
siderably lower moisture levels of the soil during extended periods of high temperature and
low rainfall.

Populations usually occur on freely-draining soils, but can be found on seasonally-
waterlogged sites (e.g. Parsons, 1966; Siddiqi et al., 1972; Myerscough and Carolin, 1986)
and occasionally in permanently moist areas (e.g. Parsons, 1966; Siddiqi et al., 1976;
Benson and Fallding, 1981). However, plants were not recorded as regenerating after fire
in ground-water heaths (Siddiqiet al., 1976); and plants have not been recorded from sites
where a standing water table reaches the shoot system.

MORPHOLOGY

The flowers are borne in axillary racemes of heads, these racemes being produced only
by buds in the axils of phyllodes on those vegetative shoots that were produced during the
previous December to January. Shoots produced in previous years do not usually flower.
The axillary primordia may or may not expand and differentiate into floral buds. These
floral buds expand lengthwise, the bracts fall off, and the flowers open. These flowers are
andromonoecious, with about 13-50% of the flowers being bisexual (Morrison, 1986).

As each individual flower forms, it follows a spiral developmental sequence from the
outside to the inside, described in detail by Newman (1936). Each flower is protogynous,
the style first being exserted well beyond the petals, which still enclose the contorted un-
dehisced anthers. The style is folded in bud, and pushes its way out between the over-
lapping petals before the bud opens, straightening progressively as the flower begins to
open. The cup-shaped stigma 1s of the wet non-papillate type, and is sited terminally on the
long narrow style. Several days after the exsertion of the style, the stamen filaments unfold
and lengthen; although the style still projects well beyond them. The anthers then dehisce.
Each flower head in a raceme flowers synchronously, so that each head is also protogynous.
However, the racemes open subacropetally and subsynchronously, while the racemes
usually open basipetally along the vegetative shoot. Therefore, each raceme and each shoot
usually contains flowers at all stages of anthesis at some time.

When the ovary has been fertilized it changes colour from dull brown to reddish-
maroon. The unfertilized flowers drop off early, leaving the small fruits on the bare stalks.
The fruits then expand very slowly in both length and breadth for 5-8 weeks, after which
they rapidly expand in length only for a further 3-5 weeks, before breadth again begins to
increase. The fruits reach their final size rapidly after this, and they begin to thicken the
fruit walls. The fruits then dry out and open along the ventral and then the dorsal sutures,
releasing the seeds. The fruits open about 15-22 weeks after they are formed, with seed
release being fairly synchronous in any one population.

The buds that form the next season’s vegetative shoots are in the same axils of the phyl-
lodes as the floral buds, and adjacent to them. Thus, the new shoots of one season are the
sites of both the ensuing flowering and fruiting and also of the following season’s shoot
production. If no new vegetative shoots survive on a particular branch of a plant in a par-
ticular season, then that branch dies back to a branch that is supporting a surviving shoot.
If no new shoots survive on a plant, then that plant dies back to the main stem, where it
may or may not produce new shoots from buds that are several years old. Such plants do
not usually flower again, and they die the following summer. Therefore, the successful
production of new vegetative shoots is the key to the long-term survival of a plant.

If the growing points of the plants are destroyed in any way, reversion foliage consist-

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

D. A. MORRISON 281

ing of pinnate or bipinnate leaves may appear on the proximal parts of branches or along
the stem (Cambage, 1915; 1917; Fletcher, 1920). Similar foliage may be produced at the
base of the stem if the upper shoots die back during summer, although this usually consists
of small phyllodes with a number of leaflets at the tips. Plants with reversion foliage rarely
produce floral buds, and they usually die within the next 3-12 months.

The leaflet buds are often found along the margins of the phyllodes as well as at the tip,
resulting in the appearance at different places along the phyllode margins of pairs of
reduced pinnae, pairs of leaflets, and single leaflets (Fletcher, 1920).

The phyllodes are xeromorphic microphyllous leaves. The epidermis is narrow, with
a thick cuticle (Lemesle, 1965). Most of the mesophyll is composed of chlorenchymatous
palisade cells, but it passes gradually into a smaller central zone of spongy cells less rich in
chloroplasts (Lsmesle, 1965; Boughton, 1986). The primary and secondary veins are
covered by a crescent of sclereid fibres, often with tracheids, and the surrounding tissue
sometimes has calcium oxalate inclusions (Lemesle, 1965). The stomata are mesogenous,
paracytic and bicytic (Grosso, 1987), fairly large (25.0 + 2.5um long; Connor and Doley,
1981), and occur on both surfaces of the phyllodes at a density of 249 + 21mm (Connor
and Doley, 1981).

The woody stems have a perennial vascular cambium, which produces very little
secondary phloem. The secondary xylem consists of diffuse porous vessels with
paratracheal axial parenchyma. There is no ray parenchyma, and no heartwood is formed.
Periderm is present only on older stems very close to the base of the stem.

The contribution to the community biomass (i.e. wood component) in any one area
is usually very small (e.g. < 0.05%, Clark, 1975; 0.06-0.09%, ‘Thatcher and Westman,
1975; 0.1%, Specht, 1979), and the percentage cover (i.e. leaf component) is also low (e.g.
0.02- 0.32%, Russell and Parsons, 1978; > 0.17%, Posamentier et al., 1981; 0.7-1.4%, Weste,
1981; 0.2-0.8%, Fox and Fox, 1986). The percentage cover does not apparently show any
particular pattern of change with population age (Russell and Parsons, 1978).

CHROMOSOMES
2n = 26 in material from New South Wales (Hamant et al., 1975).

PHYSIOLOGICAL DATA

Specht and Groves (1966) and Groves and Keriatis (1976) have investigated the phos-
phorus relations of A. suaveolens in water and sand cultures respectively. The plants showed
little growth below 0.1ppm P, but there was a significant increase in dry weight accumula-
tion between 0.1 and 1.0ppm P. No further dry weight increase occurred at 5.0-10.0ppm P,
and all plants died at 50ppm P and above. At the higher P levels, root weight showed a
greater decrease than did shoot dry weight; and the plants showed toxicity symptoms above
P levels of about 4% of the shoot dry weight, with irregular necrotic areas appearing on the
phyllodes before they began to die from the tips down. At levels of 50ppm P or more, the
cotyledons senesced and the juvenile bipinnate leaves became red-purple, with the pinnae
and petioles dying before the formation of phyllodes. At low P levels, < 30% of the dead
leaves were shed and 85-90% of the P was translocated from the dead to the living tissues.
The plants did not show any deficiency symptoms at low P levels. The P content of the phyl-
lodes tended to increase with increased P levels in the substrate, and the maximum P
uptake from the substrate was 57% (in water culture).

The nitrogen relations of A. suaveolens in sand culture have been investigated by Groves
and Keriatis (1976). The plants showed no response to changing N levels from 0-250ppm
without the addition of P. The most favourable combination for growth was low P concen-
tration and high N concentration (see also Hannon, 1956). At low N levels the plants

PROG. LINN. SOC. N.S.W., 109 (4), (1986) 1987

282 ACACIA SUAVEOLENS

showed deficiency symptoms, with a reddening of the leaflet tips; and the N content of the
phyllodes did not vary with the N concentration in the substrate. The root/shoot ratio was
highest at intermediate N concentrations with no added P.

Beadle (1962) grew A. suaveolens plants in a range of soils ina glasshouse, and found
that plants grew larger (ranging from 12 to 170cm tall after 5 months) and produced more
leaves on soils with higher nutrient levels (ranging from 23 to 230ppm P). Leaf area did not
differ between these treatments. Plants grown on full nutrient solution in the laboratory
absorb P far in excess of field requirements (Beadle, 1968); and plants given insoluble phos-
phate in the form of ground fossil laterite concretions could not absorb this ‘unavailable’ P
(Beadle, 1968).

Clark (1975) and Thatcher and Westman (1975) have shown that the addition of
fertilizer to A. suaveolens seedlings following sand mining increases their contribution to the
community biomass in the following 2-3 years (i.e. bigger plants are produced), but that by
the fourth year the species has returned to its pre-mining level. Specht (1975) and Specht
et al. (1977) have concluded that an increase in the nutrient level in the soil increases the
growth rate and speeds-up the life cycle of these plants, resulting in their earlier death; and
Specht et al. (1977) recorded the disappearance of an unusually high number of A. suaveo-
lens plants during the 8 years of their fertilized treatment.

The seeds contain about 0.2% total P, 4.0% total N, 1.0% K, 1.3% Ca, and 0.4% Mg
(Beadle, 1968; Groves and Keriatis, 1976; P.J. Myerscough pers. comm.). Phyllodes from
field plants contain 0.01-0.05% P (Beadle, 1968; Lambert and ‘Turner, 1987) and 1.8-2.1%
N (Hannon, 1956), both increasing with increased nutrient status of the soil, and 0.01% Al,
0.40% Ca, 0.32% Meg, 0.86% K and 1.92% Cl (Lambert and Turner, 1987).

BIOCHEMICAL DATA

Seneviratne and Fowden (1968) and Evans e¢ al. (1977) have found the following free
amino acids in the seeds: S-carboxyethylcysteine (the predominant amino acid),
S-carboxyethylcysteine sulphoxide, S-carboxyisopropylcysteine, 6-acetyl- &,@-diamino-
propionic acid, «-amino-@-ureido-propionic acid, pipecolic acid, 4-hydroxy-pipecolic
acid, 5-hydroxypipecolic acid, djenkolic acid, djenkolic acid sulphoxide, y-glutamy]l-
djenkolic acid. Conn et al. (1985) did not find any cyanogenic glucosides in either fresh
leaves or herbarium material.

PERENNATION AND REPRODUCTION

A. suaveolens is normally a nanophanerophyte, or occasionally a microphanerophyte.
Plants without rootstocks have a half-life of 3.7 years (Auld, 1987), with most populations
lasting a maximum of 15-25 years (cf. Siddiqi et al., 1976; Specht et al., 1977; Russell and
Parsons, 1978; Clemens and Franklin, 1980; Bradstock, 1981; Auld, 1987). The lifespan of
the form with rootstocks is unknown. The annual death rate is about 22% up to 8 years of
age, and about 12% after this (Auld, 1987).

Vegetative growth continues throughout the life of the plant, with 1-6 shoots being
produced each year (Morrison, 1986). Shoot production is low for old plants, and very
variable for younger plants (Morrison, 1986), with the younger plants producing much
longer shoots (Morrison, 1986).

Reproduction is entirely by seed, although this species can be cultivated from cuttings
(Elliot and Jones, 1982). Seedlings are rarely found in the field unless recently stimulated
to germinate by a fire (Auld, 1987), with fire-free periods of 10-30 years being the most
appropriate for the long-term maintenance of viable populations of this species (Auld,
1987).

PHENOLOGY

Flowering is strictly seasonal, with floral buds usually initiated in autumn, but with

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

D. A. MORRISON 283

a distinct geographical sequence from north to south (Table 4). Flowers are most profuse
in mid winter, with the fruits ripening and releasing the seeds from late spring to early
summer (Table 4).

TABLE 4

Times at which flowers and mature fruits of Acacia suaveolens have been recorded
Data from 216 dried specimens at CBG, MEL, NSW, PERTH, and SYD which had sufficiently detailed data on collection-locality and
date, plus Rodway (1903), Ewart (1930), Groves and Specht (1965), Court (1972), Beadle (1976), Armitage (1977), Rogers
(1978), Clifford and Specht (1979), Pedley (1979), Whibley (1980), and Beadle et al. (1982)

Area Flowers Mature fruits
Queensland early March — mid August early June — end October
N.S.W. north coast* mid March — end August early June — end October
N.S.W. central coast

and tablelands mid March — early September early June — end November
N.S.W. south coast + early April — end October end June — early December
Victoria end April — end October early July — mid January
South Australia end May — end September early September — mid January
Tasmania mid May — mid September early September — end January

* north of Newcastle; + southof Nowra

Flowering time 1s very population-specific in any one area (Morrison, 1986), but there
is considerable variation from year to year, with low rainfall at the beginning of the season
delaying the onset of flowering (Morrison, 1986). However, Blakely (1941) suggests that
early onset of flowering is a!so related to low rainfall. Individual plants flower for about 4-7
weeks, but this duration decreases with plant age (Morrison, 1986). Any individual floral
bud on a plant flowers for about 3-5 weeks (Morrison, 1986), with most buds opening fairly
synchronously.

Fruiting phenology closely follows the flowering phenology (Morrison, 1986), except
that fruits ripen and release their seeds over the same 2-3 week period each year, irrespec-
tive of when flowering was initiated (Morrison, 1986). Consequently, only early-opening
flowers ever ripen fruits, as ripe fruits are usually 15-20 weeks old when they release their
seeds (Morrison, 1986).

Vegetative buds are usually initiated immediately after the seeds are released. Vegeta-
tive growth continues for 8-10 weeks, when the floral buds are initiated on the new shoots.

Very small numbers of seeds germinate without stimulation from fire, and these may
be found at any time of the year (Auld, 1987). Only 14 seedlings emerged at seven sites over
three years, and only one of those survived longer than six months (Auld, 1984).

FLOWERING AND POLLINATION

Plants can flower within 1.5 years if germination occurs in summer, but not until the
second year if germination occurs in later seasons (cf. Clemens and Franklin, 1980;
Benson, 1985); however, individual plants may take up to 4-5 years to flower (Auld, 1984;
Benson, 1985). 80- 93% of the plants in a population flower each year, and this is consistent
from year to year for the life of the population (Morrison, 1986).

Flower production varies greatly from year to year, but there is a close inverse relation-
ship with plant age, the first 1-4 years being the most prolific (Morrison, 1986). This pattern
seems to be related to a reduced number of floral buds being produced per vegetative shoot
as the plants age (Morrison, 1986), while the large inter-year variation in flower production
is related to inter-year variation in the number of vegetative shoots produced (Morrison,

PROG. LINN. SOC. N.S.W., 109 (4), (1986) 1987

284 ACACIA SUAVEOLENS

1986). Some plants also appear consistently to produce more flowers each year than do
others (Morrison, 1986).

Superimposed on the age pattern is a relationship with rainfall, increased rainfall
early in the flowering season being correlated with increased flower production (Morrison,
1986). However, floral buds produced in the middle of the season produce more flowers and
ovaries than do floral buds produced early or late in the season (Morrison, 1986), and the
ovaries contain more ovules.

Ovule number per ovary is not very variable, being reported as 5-6 (Newman, 1936),
5-7 (Kenrick and Knox, 1982; Knox and Kenrick, 1983), and 6-9 (Morrison, 1986). Anther
number per flower is also not very variable, at 60-80 (Newman, 1936), and 44-52
(Morrison,1986). Pollen fertility is fairly high, at 85-95%, but this decreases with plant age
(Morrison, 1986).

About 15-31% of the ovaries produced per plant are fertilized, with about 1-12%
aborting, and 2-48% being eaten by insects (Morrison, 1986). Of the ovules produced per
plant, about 3% are aborted, 12% are eaten, 1% are not fertilized in ovaries that are
fertilized, 23% are fertilized, and 61% are dropped from the plant (Morrison, 1986).

Pollination of flowers is very consistent in this species at about 15-31% (Morrison,
1986), and so fruit production closely follows the pattern of flowering discussed above
(Morrison, 1986). However, pollination rate is markedly decreased in floral buds opening
late in the flowering season (Morrison, 1986).

The pollinators are apparently a range of non-specific insects (Morrison, 1986), in-
cluding beetles (Coleoptera: Chrysomelidae, Cerambycidae, Apionidae), bees (Hymeno-
ptera: Apidae, Halictidae), flies (Diptera: Syrphidae), ants (Hymenoptera: Formicidae),
and hemipterans. However, the introduced honey bee, Apzs mellifera (Hymenoptera:
Apidae), seems to be the most effective pollinator in some areas (Morrison, 1986). Pollen
is the only reward, as no nectar is secreted by the phyllode gland (Hardy, 1912; Boughton,
1981; but see also Carne, 1913b). Wind pollination is unlikely, as the 16-grain pollen polyad
(see Cookson, 1954; Guinet, 1969; Kenrick and Knox, 1982; Knox and Kenrick, 1983) is
not easily windborne.

SEED PRODUCTION AND DISPERSAL

Of the fruits formed on a plant, about 38-85% abort during the first 5-10 weeks after
they are formed, 5-25% abort later on, 2-41% are eaten by insects, and 1-13% mature and
release seeds (Morrison, 1986). The abortion of the young fruits apparently allows the
plants to regulate tbe number of seeds released per plant quite closely (Morrison, 1986).
About 48% of the plants in a population mature fruits in any one year (Auld and
Myerscough, 1986), and there is considerable inter-year variability in the number of fruits
per plant (Auld and Myerscough, 1986; Morrison, 1986). The number of fruits matured
per plant follows the same inverse relationship with plant age as does flower production
(Auld and Myerscough, 1986; Morrison, 1986), and there is the same increased production
with increased rainfall (Auld and Myerscough, 1986).

Of the seeds formed per plant, about 60% abort while small, 12% abort later, 16% are
eaten by insects while small, 7% are eaten by insects later, 3% are consumed by weevil
larvae (Coleoptera: Curculionidae), and 2% are matured and released (Morrison, 1986).

The dispersal unit is the seed with its aril. Average seed weights (with aril included)
per population of 23-41mg have been reported from a wide geographical range (Specht and
Groves, 1966; Beadle, 1968; Groves and Keriatis, 1976; Drake, 1981; Westoby et al., 1982;
Auld, 1983; Morrison, 1986), and average weights from 27-41mg have been reported from
populations within a few kilometres of each other (Auld, 1983; Morrison, 1986).

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

D. A. MORRISON 285

The seeds are released passively from the fruits. Auld (1986b) reports that the average
distance of fallen seed from the parent plant is 45cm, with 90% of the seed within 1m of the
parent. This distance is partly dependent on the height of the parent plant (Auld, 1986b).

Ants have been observed to move seeds to their nests in the Sydney region (Rice and
Westoby, 1981; Auld, 1986b), on North Stradbroke Island (Drake, 1981), and on Wilsons
Promontory (Andersen and Ashton, 1985). Drake (1981) found seeds to be removed at the
rate of 10 seeds in 55 min, with Rhytedoponera metallica removing 9 seeds and Aphaenogaster
longiceps removing the other. Auld (1986b) reports two unidentified species of Pheidole and
one of Iridomyrmex to move seeds, although only Pheidole sp.A actually took them into their
nests. Pheidole sp.A moved seeds an average of about 220cm, while the other two species only
moved them about 10-15cm (Auld, 1986b). Removal rates vary from 93-100% (Drake, 1981;
Auld, 1986b; Andersen and Ashton, 1985), although Auld (1986b) reported that only 38%
of the seed removed by Pheidole sp.A was incorporated into the nests, and none of the seed
removed by the other two species were. Seeds are found in the top 5cm of the soil outside
ant nests, but in the top 2-15cm inside the nests (Auld, 1986b). About 65% of the seeds end
up in sites that are unsuitable for germination (Auld, 1987).

All ant species drag the seeds by the tip of the aril. The elaiosome 1s the folded aril,
which in A. suaveolens is unusual in being dark brown instead of the more usual whitish
colour of other acacias (Vassal, 1971; 1972; Drake, 1981). This elaiosome comprises about
5% of the weight of the dispersal unit (Westoby et a/., 1982; Auld, 1984).

The released seeds are incorporated into the soil seed bank, at a density of 6-23
seeds/m? outside ant nests (Auld, 1986b). The annual decay rate of seeds in the soil is
6.5%, with a seed half-life of 10.7 years (Auld, 1986b). The peak size of the soil seed bank
occurs at a population age of about 6 years, and the population self-replacement point (ie.
where the number of seeds in the soil equals the initial mature population size) is about 60
years (Auld, 1987).

Dispersal of seeds onto bare rock has also been reported. Five years after an 8-acre
[3.2ha] area of Hawkesbury sandstone was cleared of both plant and soil cover to expose the
underlying rock, A. suaveolens was one of the few species recorded to have become estab-
lished (Cambage, 1923); and 44 years later plants were reported to be scattered in-
frequently over the still extremely thin sandy soil (Hannon and Evans, 1963).

VIABILITY OF SEEDS AND GERMINATION

Staining with 2,3,5-triphenyl-tetrazolium chloride reveals that mature seeds heavier
than 20mg are more than 98% viable, independent of site or age of the parent plant (Auld,
1986a). Seeds less than this weight have significantly reduced viability (down to 40%). The
seeds have a hard impermeable seed coat 180um thick (Cavanagh, 1980), which means that
the seeds are in induced dormancy (sensu Harper, 1977). Less than 1% of the mature seeds
will germinate spontaneously on release from the fruit, and the remainder enter the soil
seed bank (Auld, 1986a). Most of the induced dormancy is acquired during the first 2 weeks
after seed release (Auld, 1986a).

The seeds can remain viable in storage for many years. P. J. Myerscough (pers.
comm.) found that after 83 months storage 16/23 seeds were still capable of imbibing after
scarification and treatment with boiling water, and Ewart (1908) reported that 1/25 seeds
stored for 51 years was still capable of imbibing after sulphuric acid treatment.

The seed coat impermeability can be overcome by heating (either by adding boiling
water, heating in an oven, or exposure to microwaves), treatment with sulphuric acid and
dilute ammonia or lime water, or mechanical chipping or abrading (Ewart, 1908; Clemens
et al., 1977; Cavanagh, 1980; Auld, 1986c). Clemens et al. (1977) achieved maximum
germination of 77% by chipping the seeds, while Auld (1986a) achieved 98% germination

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

286 ACACIA SUAVEOLENS

by abrading with sandpaper. For mechanical chipping, seeds germinate more rapidly, with
a maximum rate of 48.5 seeds/day as opposed to only 2.2 seeds/day for the boiling water
treatment (Clemens et al., 1977). Seeds imbibe within 15 hours when mechanically chipped
(Clemens et al., 1977). The optimum temperatures for breaking seed dormancy are 60-
80°C, for any length of time (Clemens et al., 1977; Auld, 1986c). Below this temperature,
germination percentage is low, even if heated for long periods of time, and above this the
seeds are killed even if exposed for short periods.

Cavanagh (1980) considers the strophiole to be responsible for overcoming the hard-
seededness. He found that, after heating the seeds to 100°C by microwave exposure, the
shortened palisade cells covering the vascular bundle at the strophiole broke down, allowing
water penetration in this area. The seeds then swelled from this end. Heat-treated seeds
that had the strophiole covered with petroleum jelly did not imbibe, and so there is no
general water permeability after treatment.

Heat-treated seeds have been observed to germinate and emerge from depths of up to
10-15cm in laboratory trials (Drake, 1981; Auld, 1986c); but in the field, emergence has only
been recorded from a maximum depth of 6cm (mean depth 2.4cm) (Auld, 1986c).

SEEDLING MORPHOLOGY

Seedling growth in A. suaveolens has been monitored and described in detail by
Cambage (1915) (see his fig. 1, fig. 3, plate IX nos 8-10) and more recently by Vassal (1970;
1972)

ee germination, the curved upper portion of the hypocotyl appears above the soil
first, the cotyledons free themselves from the testa, and then the hypocotyl elongates and
becomes erect, pulling the cotyledons out of the soil, where they open out. The hypocotyl
is erect, terete, pale reddish-violet, 5-40mm long, thicker than the epicotyl, with their
boundary marked by an annular crest. The cotyledons are opposite, shortly (but distinctly)
petiolate, oblong, distinctly lobed, sagittate, 6-9mm long, 3-4mm wide, erect at first but be-
coming horizontal in a few days, reddish-violet on the lower surface, greenish-brown be-
coming greenish-red then green on the upper surface. They are persistent at least until the
production of the 15-16th leaves.

The first leaf produced is pinnate, followed by a succession of alternate, bipinnate
leaves, phyllodinization beginning at the 5-8th leaf stage and ending between the 7-12th
leaves. The first two leaves are produced at right angles to the plane of the cotyledons, and
subsequent leaves appear in two helices. This pattern does not change with the transition
to phyllodes. The first leaf is usually 5-7mm long, and is a similar colour to the cotyledons.
Subsequent leaves sequentially increase from 1-2cm to 3-4cm long, and are pale green. The
early leaves are lyrate, with the number of leaflets sequentially increasing from (2-)3-4
(-5) to 4-6(-7), before phyllodinization. Up to the 4th leaf stage, mixed leaves are sometimes
produced, where the proximal pair of leaflets may be replaced by a pair of pinnae which are
as large as the distal part of the leaf. The phyllodes appear gradually as the petioles in
subsequent leaves become more dilated vertically, with reducing numbers of leaflets at the
tips.

MYCORRHIZA

Rhizobial root nodules have been reported on this species in the field in the Sydney
region (Carne, 1913a; Benjamin, 1915; Hannon, 1956; Norris, 1959; Barnet et al., 1985) and
in Queensland (Bowen, 1956). Thess rhizobia are of the slow-growing Bradyrhizobium type
(Norris, 1959; Barnet et al., 1985) as well as the fast-growing RAyzobium type (Barnet et al.,
1985). The nodules can produce a urea-splitting enzyme (Benjamin, 1915). In cultivation,
the bacteria show increased growth with increased Mg in the medium (Norris, 1959); and

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

D. A. MORRISON 287

root nodulation decreases with increasing N levels in the substrate (Groves and Keriatis,
1976), but increases with added P at low N levels (Hannon, 1956; Beadle, 1962).

ANIMAL FEEDERS AND PARASITES

The fruits of A. suaveolens are reported to be an important food source for lorikeets,
parrots, cockatoos, and native pigeons (Adams, 1980), although evidence has been
presented only for crimson rosellas [Platycercus elegans (Gmelin) ], which cut the edge from
the unripe pod to extract the half-ripe seed.

The only invertebrates reported to be associated with A. suaveolens are insects. Froggatt
(1902) reports that larvae of Rhinotia hoemoptera Kirby (Coleoptera: Bruchidae) live inside
the branches, while the adults feed in the foliage. Also, Sextzus virescens (Faimaire) (Homo-
ptera: Membracidae) lay their eggs in slits cut through the bark of young branches (Cook-
son and New, 1980). New (1983) reports 11 unidentified species of arthropods inhabiting
18-month-old seedlings, 3 of these species being Araneae, 2 Coleoptera, and 1 Lepidoptera.

Morrison (1986) reports unidentified beetles (Coleoptera: Chrysomelidae) and grass-
hoppers (Orthoptera: Acrididae) consuming flowers.

Auld (1983), Auld (1986d) and Auld and Myerscough (1986) report that larvae of
Melanterius corosus (Boisduval) (Coleoptera: Curculionidae) feed in developing seeds in the
field, as well as consuming whole fruits. Auld (1986d) also reports that adults of Melanterius
maculatus Lea will oviposit in fruits in the laboratory. Auld (1983) and Auld and Myerscough
(1986) report unidentified lepidopterans, hemipterans, and grasshoppers feeding on all or
part of developing fruits, while Morrison (1986) lists unidentified species of adult grass-
hoppers (Orthoptera: Acrididae), sap-suckers (Hemiptera: Psyllidae), and beetles (Coleo-
ptera: Chrysomelidae) doing the same. Morrison (1986) also notes several unidentified
species of coleopteran larvae (Coleoptera: Lagriidae, Pythidae, Tenebrionidae, Nitidulae),
and five unidentified species of lepidopteran larvae, all eating developing fruits.

Drake (1981) reports Aphaenogaster longiceps F. Smith and Rhytidoponera metallica F. Smith
(Hymenoptera: Formicidae) eating the aril of mature seeds, while Auld (1986b) reports two
species of Pheidole and one of Iridomyrmex to do the same.

PLANT DISEASES AND PARASITES

Weste and Law (1973) and Weste (1981) consider A. suaveolens to be a species which is
tolerant of Phytophthora cinnamom Rands rather than resistant to it. In their study plots on
Wilsons Promontory, the species did not show early signs of becoming affected by the root
rot, unlike the majority of the surrounding woodland (Weste and Law, 1973), but did
eventually develop mild but fluctuating symptoms (Weste, 1981). While the surrounding
community changed from a low shrub woodland to an open sedge woodland, with a reduc-
tion in tree density and a loss of susceptible species, A. suaveolens was the only species to
actually increase in percentage frequency. On the diseased plots, A. suaveolens plants showed
severe and permanent die-back during periods of high evaporation and low rainfall, while
on the control plots no die-back occurred. This behaviour may merely be a response to the
opening of the tree canopy.

Fletcher (1920) records that A. suaveolens plants are ‘particularly liable to fungoid
attacks, which sometimes interfere with, or even kill the growing point, but he does not
specify the fungi involved. Similarly, Cambage (1917) comments on ‘pathological trouble’
at the growing points but is not specific.

HISTORY AND CONSERVATION

A. suaveolens was first collected by Joseph Banks and Daniel Solander at Botany Bay in
1770 (Britten, 1905), and it was among the first species to have seeds sent back to Europe

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

288 ACACIA SUAVEOLENS

from Australia (Lebler, 1980), so that by the end of that century it had been established in
a number of the botanical gardens of Britain (Loudon, 1830) and Europe (e.g. Jacquin,
1798). The only report of the direct human exploitation of A. suaveolens is the use by the
early European settlers of the aromatic leaves in infusions as teas (Nakao, 1976). The pre-
history is unknown, as no pollen directly referrable to A. suaveolens has been found in the
fossil record.

A. suaveolens was apparently common throughout its geographical range when it was
first collected by Europeans, as it was readily collected by many of the early exploration
parties. However, it is now much more restricted in occurrence in Queensland, Victoria,
South Australia, and Tasmania due to human destruction of suitable habitats (Morrison
Ci Ale 983)):

In Queensland, populations are now rare, principally because of the many new coastal
housing developments that are occupying the available habitats (Morrison et al., 1983). The
only areas where this species appears not to be at risk are the less-disturbed parts of the
large sand islands.

In Victoria, there are apparently no longer any large coastal populations west of
Wilsons Promontory, although this species was frequently collected between there and
Melbourne at the turn of the century (Morrison et al., 1983). This appears to be a result of
pastures and settlements encroaching on the somewhat restricted habitats.

In South Australia, this species has only been recorded since the early 1960's, and the
majority of the known populations are in vacant lots in plantations (Morrison et al., 1983).
More recently, the western-most known population was apparently destroyed by roadside
vegetation clearing.

In Tasmania, this species is also rarely encountered, principally due to destruction of
coastal heathlands. In particular, the species’ distribution along the northern and eastern
coastlines is now very patchy.

The species is most widespread and common in New South Wales, and it is very easily
located along most of the coastal areas. In particular, even in the disturbed urban areas A.
suaveolens is commonly encountered in the coastal parts, unlike the other states.

The two geographically-restricted morphological forms are each present in conser-
vation areas, the narrow-leaved form in the Myall Lakes National Park, and the Grampians
form in the Grampians National Park.

ACKNOWLEDGEMENTS

I wish to thank: Tony Auld for stimulating my interest in this species in the first place,
and for helping in numerous ways over the years; Peter Myerscough for encouraging me
to expand and complete what started out as a minor review, and for providing unpublished
data; Kerri Gallagher for help with many things; Roger Carolin for the electron microscope
work on the wood anatomy; Leon Costermans and Brian Mitchell for many helpful letters;
and Cliff Beauglehole, Estelle Canning, Bruce Maslin, and David Whibley for help with
the distribution data.

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PROC. LINN. SOC. N.S.W,, 109 (4), (1986) 1987

Multiple Folding of the Ordovician Sequence,
‘Tambo River, eastern Victoria

CHRISTOPHER L. FERGUSSON

FERGUSSON, C. L. Multiple folding of the Ordovician sequence, Tambo River, eastern
Victoria. Proc. Linn. Soc. N.S.W. 109(4), (1986) 1987: 293-309.

Ordovician quartz-nich clastics of the Tambo River region in the Tabberabbera Belt
of eastern Victoria show the effects of four folding episodes. A strong to weak bedding-
parallel slaty cleavage is developed throughout the region and predates the earliest folds
so far identified. F, folds are tight to close with steep easterly trending axial surfaces and
large amplitude-to-wavelength ratios. The F, folding is thick-skinned and is responsible
for major crustal thickening in the area. F, folding is only locally developed and is
associated with a pervasive stripy cleavage in sandstones. F3 folds are widespread upright
northerly-trending structures with close to open interlimb angles. Superimposition of F,
on F, has caused bending of F, fold trends and formed an eye-type interference fold
pattern in one instance. F, folds consist of warps and kink-like folds.

The east-west F, folds extend into the Omeo Metamorphic Complex and predate
the Late Silurian volcanic and sedimentary sequence near Benambra. F3 folds have been
traced westwards to Tabberabbera where they are of the Middle Devonian age and are
related to east-west compression.

C. L. Fergusson, Department of Geology, University of Wollongong, PO. Box 1144, Wollongong, Aus-
tralia 2500, formerly Department of Earth Sciences, Monash University, Clayton, Australia 3168;
manuscript received 9 December 1986, accepted for publication 22 April 1987.

INTRODUCTION

The Palaeozoic tectonic history of the Lachlan Fold Belt in eastern Victoria is charac-
terized by several intermittent compressional events (VandenBerg and Wilkinson, 1982).
The styles and orientations of structures produced by these deformations are known only
from some restricted areas (e.g. Beavis, 1967; Fagan, 1979; Wilson e¢ al., 1982). The aim of
this paper is to describe the structure of the Ordovician rocks in the Tambo River region
of eastern Victoria (Fig. 1) and to relate the two main deformations to the Early-Middle
Silurian Benambran and Middle Devonian Tabberabberan deformations respectively.

REGIONAL SETTING

The Ordovician sequence of the Tambo River region lies partly in the southern Omeo
Metamorphic Complex and mainly in a belt of very low-grade metamorphosed rocks called
the Tabberabbera Sub-zone by VandenBerg (1978) and the Tabberabbera Belt by Fer-
gusson (1985). The eastern boundary of the Tabberabbera Belt is the Kiewa Fault which
has a steeply dipping mylonite zone up to 2km in width (Scott, 1985). East of the Kiewa
Fault is the Omeo Metamorphic Complex which consists of lower greenschist to upper
amphibolite facies metamorphics derived from Ordovician quartzose clastics and intruded
by S- and I-type ‘granitoids’ (Fagan, 1979).

Fagan (1979) has shown that major folds in the metamorphics are upright and easterly
trending. This deformation extends throughout the Tabberabbera Belt and is the major
folding in the Tambo River region (Fergusson, 1985). North and east of Benambra the Late
Silurian volcanic and sedimentary sequence postdates the major metamorphism and defor-
mation in the Omeo Metamorphic Complex (Fig. 1; Bolger, 1982; Bolger e¢ al., 1983). The
Silurian sequence was strongly deformed prior to the deposition of the unconformably-
overlying Lower Devonian Snowy River Volcanics (VandenBerg and Wilkinson, 1982).

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

294 MULTIPLE FOLDING

Upper Devonian —-Carboniferous Avon
River Group and equivalents

Cambrian-Devonian stratified rocks of
the Melbourne Synclinorial Zone

Omeo Metamorphic Complex

Ordovician sedimentary rocks and
Siluro—Devonian granitoids

NEW SOUTH WALES

T.|N.

Granitoids

Ordovician
sequence

0 F, syncline
La

4 F, anticline
7

e F, syncline
d
7

Fig. 1. a. — Geological sketch map of eastern Victoria with the location of the Tambo River region shown by the
shaded inset.

b. — Geological map of the Tambo River region showing the locations of Figs 2 and 5 and domains 1 to 12. Note
that the Kiewa Fault is not shown as it is cut by the ‘granitoid’ between domains 1 and 2.

At Tabberabbera the Ordovician sequence is overlain with a high angular uncon-
formity by the Emsian Wentworth Group and both are, in turn, overlain with a high
angular unconformity by the Upper Devonian-Lower Carboniferous Avon River Group
(Talent, 1963). The latter surface was called the Tabberabberan Unconformity by Harring-
ton et al. (1974) and VandenBerg (1977). This was caused by uplift and erosion associated
with the deformation that formed a tight northerly-trending syncline in the Wentworth
Group. The deformation is commonly referred to as the Tabberabberan Orogeny.

West of the Tabberabbera Belt lies the Melbourne terrane of Fergusson et al. (1986).
The Melbourne terrane consists of Cambrian mafic volcanics and a conformable Ordo-
vician to Middle Devonian quartzose clastic succession. Structurally the Melbourne
terrane is dominated by northerly-trending upright folds with a major fault zone along its
eastern margin. The Cambrian rocks occur as slices within the fault zone which is uncon-

PROC. LINN. SOC. N.SW., 109 (4), (1986) 1987

C. L. FERGUSSON 295

formably overlain by Upper Devonian— Lower Carboniferous fluviatile sediments and
volcanics. These relations establish that the deformation that affected the Melbourne
terrane was the Tabberabberan Orogeny (VandenBerg and Wilkinson, 1982).

STRUCTURE

The Ordovician strata of the Tambo River region consist of a monotonous quartz tur-
bidite succession (Stewart and Fergusson, in preparation). The structure of the region is
dominated by two main deformations with two additional deformations of lesser extent and
significance. Each deformation is characterized by folds and accordingly each event 1s
labelled F,, F,, F3 and F, in order from oldest to youngest. These fold episodes are
mapped on the basis of orientation and fold style groups whose age relations have been
determined by overprinting criteria. The nature of axial plane foliations associated with
each fold generation has been the least useful guide in structural mapping (cf. Williams,
1985). Macroscopic structure is indicated by many sections of homoclinal strata and the
abundance of younging criteria (Figs 2, 3, 4, 5 and 6).

The earliest structure developed in the Ordovician sequence is a bedding-parallel slaty
cleavage that predates all folding (Fig. 7d). A similar fabric has been described from the Or-
dovician at Mallacoota (Wilson and de Hedouville, 1985). The significance of this fabric
will be discussed elsewhere (C. L. Fergusson and D. R. Gray, unpub. data).

F, FOLDING

The F, folding is the most pervasive structural event developed throughout the
Tambo River region. F, folds are upright isoclinal to open structures with northeast to
southeast trends that were initially easterly trending (Figs 8 and 9). Hinges are narrow and
angular with long planar limbs (Fig. 7a). Sandstone layers have class 1B and 1C shapes
whereas mudstones have class 3 shapes (Fig. 7a-c). Many of the F, folds are flattened
chevron folds, and probably formed by flexural slip along bedding planes with accompany-
ing flexural flow in mudstones, and fold flattening.

An §, axial surface cleavage is associated with the F, folds (Fig. 7c). In mudstones the
cleavage has formed from microfolding, accompanied by dissolution, of the early bedding-
parallel slaty cleavage and forms a zonal crenulation cleavage (Fig. 7d). On the limbs of
some F, folds the angle between S, and the early cleavage is low and the two fabrics are in-
distinguishable. Throughout domains 9-11 (Fig. 1) a pervasive S, stripy cleavage is de-
veloped in sandstones. The stripy cleavage consists of planar domains of aligned micas and
other pressure solution residue up to 0.5cm in width separated by microlithon domains of
quartz sandstone up to 2cm in width. The microlithons have abundant chlorite over-
growths (especially on quartz grains) indicating extension within the cleavage plane.

On the map-scale the main F;, structures are an anticlinorium in domains 1 and 2
and a synclinorium, with a refolded southern limb, in domains 3 to 12 (Fig. 1). These
structures are described separately below.

Domains 1 and 2: Domains 1 and 2 lie to the north and south of the Kiewa Fault respec-
tively but as there are no major differences in structure these domains are described
together. Domain 1 contains the hinge region and the northern limb of the F, anticli-
norium (Figs 5 and 6) whereas domain 2 has the relatively planar south-younging limb of
the anticlinorium (Figs 2 and 4).

Four orders of folding are developed within the anticlinorium. Fourth order folds are
slightly overturned Z-shaped structures, with wavelengths up to 5m, found on the upright
limbs of third order S-shaped folds on the steeply dipping to overturned northern limb of
the anticlinorium (Fig. 6). Third order folds have wavelengths of 10-20m. The core of the

PROC. LINN. SOC. N.S.W,, 109 (4), (1986) 1987

Lower Ordovician

MULTIPLE FOLDING

Lower to Middle Ordovician

296

“

aac

Cae

“N

a
°

Nos

Bedding

Upright beds

Overturned beds

Vertical beds

S; cleavage

Fy, syncline
with plunge

F, anticline

F, z-shaped
fold couple

F, s-shaped
fold coupje

n
°

\ vA

F, m-shaped
folds

F, downward

facing synform
F, axial trace
Fo downward
facing antiform

Sg cleavage

Fg, axial surface

Fg anticline

Fg m-shaped
folds

Fg axial trace

So cleavage

109 (4), (1986) 1987

Fig. 2. Geological map of the Ordovician sequence in the southern Tambo River region showing the main struc-
N.S.W.,

tural features and locations of cross sections and Figs 10 and 12. Note the small amount of overlap across the double

pa ge.
PROC. LINN. SOC.

C. L. FERGUSSON

P
<o) 8 B2 84 86
( \) + + +
ry? s@O \+ + + 0 1 2 3km
Oe Sam ee a
aot Peete:
‘ay cq ec fg os
36
| ay + + ys = Peeters
78 v9 tee o s @ ee , 6 Fey,
AVY nears Z| Pa? + 44
2A era Me” ye? Gu i Rs Bare”
85 ry 42
** SA ee : 2) 5 w., 4p 24 aur a2 ae & as ot R-
AN 5 af => © y) [\fea 24
60
Se

Fig. 2. For explanation see opposite.

109 (4), (1986) 1987

PROC. LINN. SOC. N.S.W.,

298 MULTIPLE FOLDING

Pa Younging direction

_-— Cleavage
— Bedding

Fig. 3. Regional cross section across the major F, syncline in the southern Tambo River region. See Fig. 2 for
location.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

C. L. FERGUSSON 299

pe SOAS SS Ze
Se ANN ce

3 De LO oO

7 SSM i A AN

t+) 200 400 SPEEA SpE GCE alae a aah 800 1000m
HS = VS.

Fig. 4. Cross sections for the southern Tambo River region. See Fig. 2 for location.

anticlinorium consists of three second order folds up to 600m apart. The northernmost
hinge contains several third and fourth order mesoscopic folds (Fig. 6).

One stretch of the southern limb in domain 2 consists of approximately 1km of south-
ward dipping overturned beds with some parasitic F,; downward facing folds (Figs 2, 4 and
7b). Along this section the 8; cleavage dips steeper than overturned bedding and is axial
planar to the F, folds (Fig. 7b). These overturned beds are fault-bounded and projections
to depth indicate that they are pinched out at about 1km below the surface (Fig. 4, cross
section NOP). This section is unique within the Tambo River region and reflects local
deformation prior to the F, folding.

The lack of significant younger deformation in domains 1 and 2 provides a clear cross
section of the F; structures. From cross section QRS (Fig. 6) these folds have narrow
hinges and long planar limbs and a fold style characteristic of flattened chevron folds.
These folds have a large amplitude-to-wavelength ratio which coupled with the 4-5km of
succession indicates a relatively thick-skinned style of deformation.

Domains 3 to 12: The southern Tambo River region is dominated by the north-younging
limb of the major F, synclinorium (Figs 2, 3, 4, 8, 9 and 10). The axis of the synclinorium
occurs along the Tambo River in domain 3 (Figs 1 and 2). The hinge is angular with a
planar and locally overturned northern limb (Fig. 4, cross section KLM). Throughout
domains 6 and 7 the southern limb of the F,; synclinorium is shallowly dipping and

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

300 MULTIPLE FOLDING

Bedding

Upright
—younging
determined

Overturned

S, cleavage

F, fold
with plunge
(s-folds)

o
o

F, z-folds

F, syncline

F, anticline

F4 fold
with. plunge

Granitoids

Foliated
granitoid

at
av

Fig. 5. Structural map of the Ordovician sequence in the northern Tambo River region.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

C. L. FERGUSSON

7
77 Sicleavage O, Bedding
a Younging

Fig. 7a. — Tight F, syncline from domain 1, Omeo Highway (D. W. Durney for scale). Location — GR 734534

(Grid references from Omeo and Bairnsdale 1:100 000 Topographic Sheets).

b. — Downward facing F, synformal anticline from domain 2 (see text), Omeo Highway. Note that the southern
limb of the fold is truncated by a steeply dipping fault. Width of view is 6m. Location — GR 764432.

c. — Angular upright F, syncline from domain 1. Hammer 33cm in length for scale. Location — GR 734538.
d. — A well-developed S, crenulation cleavage frem the core of an upright F, anticline. Note the S, stripy
cleavage in the sandstone bed and the cleavage refraction. Hammer is 33cm in length. Location — GR 809333.

PROG. LINN. SOC. N.S.W., 109 (4), (1986) 1987

302 MULTIPLE FOLDING

(1) (2)

+ F, (5)

Fig. 6. Lower hemisphere equal-area stereographic projections from the Tambo River region (for domains see
Fig. 1). Heavy dot is the calculated fold axis in each stereographic projection. Stereographic projections have
bedding poles contoured at: 0, 3 and 6% per 1% area (nets 3, 4, 6 and 7); 0, 3,6 and9% per 1% area (nets 5, 9,
11 and 12); 0, 3,6, 9and 12% per 1% area (net 2); 0, 3, 6, 9, 12 and 15 per 1% area(nets 8 and 10); and 0, 3, 6,
9, 12, 15, 18 and 21% per 1% area (net 1).

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

C. L. FERGUSSON 303

So poles
AS2 poles
Fo axes

iS Fy axes
e AS3 poles ° areanioe:

Fig. 9. Summary lower hemisphere equal-area stereographic projections for structural elements in the Tambo
River region. Points are contoured at 3% intervals per 1% area. Structural elements include: F, fold axes (100);
S, cleavage (212), F, axial surfaces (AS1); F, fold axes, Sy cleavage, F, axial surfaces (AS2); F3 fold axes (137), S3
cleavage (347), F3 axial surfaces (AS3); F, fold axes, S, cleavage and Fy axial surfaces (AS4).

dominated by abundant F, parasitic folds and refolded by F3. In domains 8 to 10 the
southern limb 1s steeply dipping and becomes overturned farther south (Fig. 3, cross sec-
tion ABCDEFG and Fig. 10). F, parasitic folds are largely restricted to two major Z-
shaped fold pairs along cross section BCD (Fig. 3). The northern fold pair consists of F,
minor folds plunging steeply to moderately to the east (Fig. 8, domain 8) with a faulted
axial surface along the major syncline hinge. The southern fold pair (cross section BQ, Fig.
3) contains a central limb with abundant upright F, folds in contrast to the steep limbs
which only have rare Z-shaped parasitic folds. The overturned beds to the south of this fold
pair in domains 11 and 12 are strongly refolded by Fy and F3 (Fig. 1).

F, FOLDING

In domain 10 this deformation formed a discrete hair-like crenulation cleavage that
overprints S; and is in turn overprinted by S3 (Figs 10 and Ila). F, folding is restricted to
domains 11 and 12 (Figs 1 and 10). Throughout domain 11 the overturned limb of the major
F, synclinorium is refolded by close downward facing F» folds with axial surfaces dipping
moderately to the east (Fig. 10). The Fy folds have relatively narrow hinge zones and are
extensively refolded by the F3 folds (Fig. 10). The inset in Fig. 10 shows the shape of the F>
folds prior to folding by F3.

Across the boundary between domains 11 and 12 (Fig. 1) the Sy axial surface cleavage
rapidly intensifies and bedding is transposed (with the exception of exposures along the
Omeo Highway in domain 12). Throughout domain 12 S, is developed as a pervasive
stripy cleavage in sandstones (Fig. 11g) and as a strong continuous cleavage in mudstones.

PROG. LINN. SOC. N.S.W., 109 (4), (1986) 1987

304 MULTIPLE FOLDING

Zone of strong
~ So cleavage

Fig. 10. Map and cross-sections of downward facing F, and F3 folds in domain 11 (see Fig. 1 and Fig. 2 for
location). See text for discussion. Inset shows a reconstruction of the F, folds in cross section prior to the F3
folding.

The Sz stripy cleavage is similar to S, cleavage in sandstones except that the former is
much more pervasive. F» folds have not been found in domain 12 and the Sy, cleavage itself
is refolded by younger structures (see below).

F; FOLDING

F; structures occur throughout the southern Tambo River region (Fig. 2). They are
northerly-trending folds with steep axial surfaces and variable plunges (Figs 2, 8, 9 and
llb-e). F; folds are close to open with slightly rounded hinges and characteristic small
amplitude-to-wavelength ratios (<0.5, Fig. 11b-e). An axial surface crenulation cleavage
is developed in mudstones and occasionally a weak stripy S; cleavage is developed in sand-
stones (Fig. lla-d).

Throughout most of the southern Tambo River region the F; folds are upward facing
(Figs 2, 3 and 4). In domain 11, however, the F3 folds are developed on the overturned
limbs of the downward facing F, folds and are therefore downward facing (Fig. 10).
Locally the combination of F, and F; folding has formed box-fold structures with the
western hinge formed by an F, fold and the eastern hinge by an F; fold (e.g. cross section
XY, Fig. 10). F; folds are abundant in domain 11 in contrast to the general scarcity of Fy
hinges.

On the map-scale the most obvious F; fold is the large open syncline shown in cross
section DEFGH (Fig. 3). Along the Tambo River this structure has a hinge zone 800m
across with an open conjugate fold style (cross section EF, Fig. 3). The hinges in the con-
jugate pair plunge shallowly to the north-northwest (Figs 2, 8 and 9) which indicates that
they must have developed along a shallowly dipping part of the north-younging limb of the

PROC. LINN. SOC. N.SW., 109 (4), (1986) 1987

C. L. FERGUSSON 305

Fig. 11 a. — An 83 crenulation cleavage is shown locally cross-cutting a less steeply dipping S. crenulation
cleavage. Bedding dips steeply to the east and is overturned. Hammer for scale. Location — GR 793315.

b. — Broad F; fold couple with an axial surface S; crenulation cleavage. Note an early S, crenulation cleavage
is folded in the core of the fold. Match stick for scale. Location — GR 808336.

c. — Close F; folds with a well-developed axial surface crenulation cleavage. Lens capis 5.5cm across. Location
— same as (b).

d. — Broad to open F; folds with S,; crenulation cleavage oblique to bedding on the limbs and in the fold cores.
Match stick for scale. Location — same as (b).

e. — Upright close Fs fold with a box-shaped hinge. Hammer is 33cm in length. Location — GR 819362.

f. — Steeply plunging close F; fold with an axial surface crenulation cleavage. Pen cap is 6.5cm in length.
Location — GR 809331.

g. — Sp stripy cleavage in sandstone in the core of an upright F; fold with a steep S3 axial surface cleavage (in
domain 12). Location — GR 783308.

PROC. LINN. SOC. N.S.W,, 109 (4), (1986) 1987

306 MULTIPLE FOLDING

RIVER

Fig. 12. Schematic block diagram of the F, and F3 folds along the Tambo River in domain 5 (see Fig. 2 for
location). Note that the dip of the limb of an F fold has been locally decreased by post F3 warping.

F, synclinorium. On the east limb of the F; syncline bedding dips moderately to the west

whereas on the western limb it dips steeply to the northeast (Figs 2, 3 and 8, domain 7).
AS1 (syncline)

AS1 (anticline)

<— Faulted
AS1 (syncline)

Fig. 13. Schematic block diagram of the structures in the southern Tambo River region (domains 3-10), Note the
easterly trending F, folds and the large F, syncline developed on the shallow limb of the F; syncline.

Map-scale F, anticlines occur to the east and west of the major F3 syncline (Fig. 2).
The eastern F; anticline is partly shown on cross section GH (Fig. 3) and contains abun-
dant M-shaped shallowly plunging F; folds (Figs 2 and lle). Farther north the limbs of F3
folds are extensively refolded by F, structures (Fig. 12). At GR 850 417 (Fig. 2) the F3
anticline is superimposed on an F;, anticline producing a prominent dome with bedding
dipping outward from a central focus (see also Fig. 13). F,; minor folds show significant
plunge variations in this area (domain 4, Fig. 8). North of the dome the F3 anticline causes

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

TAMBO ae

C. L. FERGUSSON 307

the bending of F, trends from east-northeast to east-west either side of the ‘granitoid’ at J
(Fig. 2). The western F; anticline caused a change in structural trends from southeasterly
to easterly from domains 2 to 7 respectively (Figs 1, 2, 8 and 13). Thus the main effect of the
F; folding on the map-scale is the bending of F, structures with only local development of
dome and basin type interference fold patterns.

F, FOLDING

Local post-F3 folds, warps and kinks occur throughout the area and these are
grouped together as F; structures. Some of these structures may belong to different fold
generations but these are not distinguished due to the lack of overprinting criteria. In
domain 1 there are several moderately-plunging east-southeast trending late-stage
mesoscopic folds with rounded open hinges. These late-stage structures are grouped with
F, for convenience, as their orientations differ from the Fy and F3 structures.

Late-stage F, folds thoughout Fig. 2 are mainly easterly trending broad to open folds
and/or kinks (Fig. 12). Several of these are map-scale F folds (Fig. 12). Rare kink-like
folds with rotated limbs up to 50m across occur at GR 806 322 (Fig. 2). In addition to the
late-stage folds a number of weak crenulation cleavages and crenulation lineations are de-
veloped in the Tambo River region. These structures are not related to other deformations
and they reflect only low values of shortening.

DISCUSSION

The F; east-west folding event extends throughout the Tabberabbera Belt (Fergus-
son, 1985) and the Omeo Metamorphic Belt where it is constrained between the Ordo-
vician and Late Silurian. Bolger (1982) has emphasized that the timing of deformation and
metamorphism in the Metamorphic Belt is only broadly delimited and is not confined to
the Early Silurian as is sometimes implied (e.g. Crook et al., 1973; Powell 1984).

Powell (1983, 1984) has proposed a tectonic model for southeastern Australia relevant
to the formation of east —west folds in the Omeo Metamorphic Complex and the Tab-
berabbera Belt. The basis of his model 1s that in the Ordovician a back-arc basin (Wagga
Marginal Sea) was bordered to the east by a volcanic island arc associated with a westward
dipping subduction zone. In the Early and Middle Silurian Chilean-style locking of plates
on the subduction zone was accompanied by dextral oblique-slip plate movement and
east — west upright folds were formed at several localities in the Lachlan Fold Belt (see Cas
et al., 1980). The east —west folds of the Tabberabbera Belt and Omeo Metamorphic
Complex are the best examples of this deformation.

The F, folding of the Tambo River region caused shortening of 50% or more (calcu-
lated from regional cross sections, Fig. 14). Estimates of the thickness of the Ordovician
succession obtained from cross sections indicate a minimum of 4-5km. The stratigraphic
thickness, fold style and shortening indicate that at least 12km and as much as 20km of the
present crustal thickness consists of the folded Ordovician succession. Thus the F, folding
represents a major episode of crustal thickening in this part of the Lachlan Fold Belt, and
was probably associated with closure of the Wagga Marginal Sea.

The F, folding is areally restricted and is only constrained in time as post-F, and pre-
F;. This folding event reflects localized east-west shortening of unknown tectonic
significance.

The F; folding is extensive throughout the southern Tambo River region but no con-
straints on the timing of this deformation exist within this area. Farther west at Tabber-
abbera, however, similar northerly trending upright folds with low amplitude-
to-wavelength ratios and an axial surface crenulation cleavage affect the Ordovician suc-
cession (Fergusson, 1985). These structures have been mapped in reconnaissance between

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

308 MULTIPLE FOLDING

Fig. 14. Regional cross section of the main F, structures in the Tambo River region. Note the relatively thick-
skinned style of folding.

the two areas. At Tabberabbera they are related to the deformation that strongly affected
the overlying Lower Devonian Wentworth Group (Fergusson, unpub. data). Thus the F3
folding is regarded as a consequence of the Middle Devonian Tabberabberan Orogeny.
This event reflects major east —west compression and overthrusting of the Melbourne
terrane over the western part of the Tabberabbera Belt (Fergusson et al., 1986).

The F, folds must have postdated F3 and reflect several mild compressions of
differing orientations.

CONCLUSIONS

(1) F;, folding in the Ordovician sequence of the Tambo River region involved the for-
mation of east —west upright folds with a shortening of at least 50%. The major F,
structures are a anticlinorium — synclinorium pair with a thick-skinned style of defor-
mation. They may have formed due to an episode of regional dextral oblique-slip
shear associated with a westward dipping subduction zone in the Early to Middle
Silurian. This event was responsible for closure of the Wagga Marginal Sea and
caused significant crustal thickening in the area.

(2) The F; folding formed due to east-west compression in the Tabberabberan Orogeny.
The largest F3 structure is a box-shaped F3 syncline that developed on the flat-lying
undeformed part of the southern limb of an F, synclinorium. Shortening associated
with F3 is variable and this event, though of regional significance, did not result in
extensive crustal thickening in this region.

(3) The Fo and F, events consist of locally developed and variably oriented folds that
formed during relatively mild compressional events.

ACKNOWLEDGEMENTS
Funded by grants from A.R.G.S. (E83 315675; principal investigators Drs R. A. F. Cas
and D. R. Gray) and from Monash University. The work was carried out in the Depart-
ment of Earth Sciences at Monash University. The diagrams were drafted by Draga Gelt.
The final manuscript typed by Therese Carmody of the Department of Geology at the
University of Wollongong. David Gray kindly reviewed the draft.

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PROC. LINN. SOC. N.S.W,, 109 (4), (1986) 1987

ees

c
J
;

:
Litet
maeye A = |

Geomorphic and physicochemical Features of

floodplain Waterbodies of the lower Hunter
Valley, N.S.W.

B. V. TIMMS

TimMs, B. V. Geomorphic and physicochemical features of floodplain waterbodies of
the lower Hunter Valley, N.S.W. Proc. Linn. Soc. N.S\W. 109(4), (1986) 1987:
311-324.

The lower ends of 42 tributary valleys of the lower Hunter and Paterson Rivers
contain floodplain lakes because the tributary outlet has been blocked by alluvial
deposits of the main stream. On average these waterbodies are elongated to dendritic in
shape, 4.3ha in area, 2m deep and have a Shoreline Development index of 2.13. Typi-
cally they fill from local run-off and subsequently their levels are largely determined by
fluctuations in the water table.

In the 5 waterbodies studied in detail, mean values for Total Dissolved Solids
varied between 215 and 468mg 1', pH between 7 and 8, turbidity between 26 and 318
FTU’s, Secchi disc ae between 30 and 100cm, and nitrates were c. 1.4mg 1! and
phosphates c. 0.4mg 1°. These parameters fluctuated widely as the waterbodies filled
and dried according to variable rainfall and evaporation. Water temperatures ranged
from 12-32°C with no persistent stratification. Waters were of the sodium chloro-
carbonate or sodium chloride types.

Almost all waterbodies are adversely affected by man, mainly via drainage,
nutrient accessions and cattle usage.

B. V. Timms, Sciences Department, Avondale C.A.E., P.O. Box 19, Cooranbong, Australia 2265;
manuscript received 11 November 1986, accepted for publication 22 April 1987.

KEY WORDS: floodplain lakes, morphometry, water chemistry, turbidity, nutrients,
water temperatures, conservation.

INTRODUCTION

Floodplains typically contain areas of ponded water. Such wetlands encompass a
wide range in sizes, depths, geomorphic origins, degree of permanence, physicochemi-
cal features and aquatic macrophyte communities, so that even reconnaissance classifi-
cation is difficult (e.g. see Cowardin et al., 1979, for USA and Riley et al., 1984, for
NSW). Detailed and long term data sets are needed for each perceived type before such
schemes can be much improved. A contribution towards this is made here for a small
area of the Hunter Valley in which wetlands are genetically similar.

According to most textbooks, ponded water on floodplains typically lies in oxbow
lakes (i.e. cut-off meanders) and in broad depressions (swamps) behind levees. These
certainly occur on the Hunter floodplain from about Maitland downstream and on the
floodplain of the lower Paterson River (Pressey, 1981). However in the section of the
Hunter R. between Singleton and Maitland and also adjacent to the Paterson R. near
Paterson, most ponded waters lie in depressions where small tributaries meet the main
valley. These form when the main stream ‘by deposition of levees and of sediment else-
where on its flood bed, aggrades its course faster than aggradation can occur in lateral
tributary valleys’ (Hutchinson, 1957: 115). Hence streams in side valleys tend to become
obstructed and water accumulates so they become partly drowned. In many cases the
main river, when in flood, flows into the lateral valley and so lengthens the obstruction
by reverse delta formation. Hutchinson (1957) uses the term ‘lateral lakes’ for these
waterbodies, but this is unfortunate as it implies relationships to floodplain processes

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

312 FLOODPLAIN WATERBODIES

such as lateral migration. The more appropriate term of ‘blocked valley lake’ (Blake and
Ollier, 1971) is used here.

All of the present waterbodies are wetlands in the broad sense (Cowardin ¢¢ al.,
1979) but called ‘lagoons’ in the local vernacular. Overseas this term is usually used
genetically for coastal marine waterbodies, but in Australia it generally refers to shallow,
often small, inland waterbodies in any geomorphic situation. This general descriptor is
appropriate here though technically those lacking emergent macrophytes are lakes or
ponds according to size and depth and those covered with emergent macrophytes are
swamps (Bayly and Williams, 1973; Riley et al., 1984). A few lagoons in downstream
parts of the study area were included by Pressey (1981) in his inventory, but otherwise
little is known of those investigated here.

METHODS

Locations of possible lagoons were ascertained from 1:25,000 topographical maps
and aerial photographs and checked first from a light plane and then on the ground. The
depth (when full) and siting with respect to local geology and landforms were ascer-
tained in the field for each lagoon. The area and shore length of all except the smallest/
shallowest lagoons were determined in the laboratory by planimetry and measurement
from enlarged vertical aerial photographs.

Three lagoons, ‘Birds’ (4), ‘Bootlands’ (19) and ‘Murphys’ (21) were sounded at 5m
intervals along numerous lines (30, 42 and 26 lines respectively in each lake) stretched
between known places on opposite shores. From the resultant bathymetric maps, direct
and derived parameters were determined using formulae given in Bayly and Williams
(1973) and Hakanson (1981). Altitudes were estimated to + 2m from topographic maps.

Five representative lagoons in the Gosforth series with a wide range in size and
degree of permanence were chosen for a study of the physicochemical features of the
waters of the 42 floodplain lagoons. A causeway separated ‘Bootlands’ (19) into two parts
on almost all visits, so both parts were sampled separately. Visits were made at monthly
intervals for 63 consecutive months commencing in October 1979 and information was
collected on water depth, temperature (by a resistance thermometer), light penetration
(by a standard 20cm Secchi disc), pH (by a Selbys 800 pH Meter), Total Dissolved
Solids (by gravimetry), and on turbidity, phosphate and nitrate (determined on a
HACH Environmental Laboratory DR/EL 1). Samples were always taken in the morn-
ing between 0800 and 1200hrs and in the same sequence (Lagoons 19 to 23). From water
samples collected in February 1981 the major ions were measured — Na and K by flame
photometry, Ca and Mg by titration with EDTA, Cl by titration against AgNO3, HCO;
by titration with 0.01N HCl to an end point of pH 4.5, and SO, by the turbidimetric
BaSO, method (Anon, 1975). Accuracy for all physicochemical methods was + 2.5% or
better.

Where possible, landowners of each lagoon were interviewed in an attempt to
establish the influence of river floods, local heavy rain and droughts on water level fluc-
tuations. Their opinion of the lagoons (e.g. water resource value, nuisance value) and
the extent of their (or their predecessor’s) modifications, if any, of the lagoons was also
canvassed.

RESULTS

(a) GEOMORPHOLOGY

Of the 42 floodplain lagoons in the study area, 31 occur along the Hunter R. and 11
along the Paterson R., giving densities of 0.7km and 1.2km respectively. The lagoons

PROC. LINN. SOC. N.SW., 109 (4), (1986) 1987

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PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

FLOODPLAIN WATERBODIES

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PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

B. V. TIMMS Bly)

range in size from <0.25ha toc 26ha (Table 1), with the average being 4.3ha. However
few approximate the average as size distribution is negatively skewed and somewhat bi-
modal. There is one peak in the 1.1-1.5ha class and another lesser one in the 5.1-7.5ha
class (Fig. 2a). All lagoons stand many metres (c. 2-10) above the normal water level of
the river. Limited data on maximum depths suggest the lagoons are relatively shallow,
with the majority c. 2m deep and the deepest only 7m. The latter (No. 28), though, is
artificially dammed so that the deepest natural lagoon is only 4m.

Shoreline Development (i.e. the ratio of the length of the shoreline to the length of
the circumference of a circle of the same area) lies between 1.3 and 3.3, with an average
of 2.13 for the 21 lagoons measured. This indicates these lagoons are either branched or
have indented shorelines (see Figs 1,3 and 5). Actually the average for all 42 lagoons is
probably a little less than 2.1, because the unmeasured lagoons generally had smoother
shorelines than measured ones.

ELDERSLIE

Pacific
Ocean

Fig. 1. Map showing location of the 42 lateral lakes studied in the lower Hunter. Three inserts give details of
the shape of 3 quite different lagoons — No. 17, alagoon in an essentially unbranched valley; No. 26, alagoon
in a branched valley and No. 38, a composite lagoon with one part a blocked valley lake and the other a lateral
levee lake.

Water regimes vary widely but are accommodated within the four categories of the
classification of Cowardin et al. (1979). Just four (9.5%) lagoons are permanently flooded
and even two of these are artifically dammed. A natural example is No. 19 (‘Bootlands’)
which contained water for all 63 months it was studied, even during the 1980 and 1983
droughts (Fig. 4). In a few lagoons (19%) the bottom 1s intermittently exposed, 1.e. water 1s
present except during extreme droughts. Examples are No. 21 ((Murphys’) which only
dried for 5 months during the 1980 drought (Fig. 4) and No. 25 (‘Greens’) which has
dried 3 times during the last 26 years during the final stages of an extended drought (J.
Green, personal communication). Many lagoons (35.7%) are semi-permanently flooded

PROG. LINN. SOC. N.S.W,, 109 (4), (1986) 1987

316 FLOODPLAIN WATERBODIES

containing water for 1/3 to 2/3rds of the time. Lagoons 20, 22 and 23 are examples (Fig.
4). There are also many ephemeral lagoons (35.7%) that are seasonally flooded for <3
months in most years.

O 25 5 GS 10 W519 >20
Lagoon Area(ha)

Number

O (a NSO 225)" COO), 8751450) 52560082675
Catchment Area (ha)

Fig. 2. Histogram showing the distribution of (a) lagoon sizes and (b) catchment sizes. In the latter case the
proportion of catchments containing lagoons in each size class 1s indicated by the solid part of each block.

The relative number in each group of the above classification, especially in the
semipermanently flooded and seasonaly flooded groups, has been influenced by
drainage programs (Table 1) so that in the past more were in the intermittently exposed
and permanently flooded classes. In fact most of the lagoons have been partly or com-
pletely drained, so that only a third have natural hydrological regimes.

The three lagoons that were mapped are similar morphometrically (Table 2, Fig.
3). ‘Birds’ Lagoon with its three distinct arms is the most branched, but ‘Bootlands’ has a
higher S.D. because of its irregular shoreline. Volume development (i.e. ratio of the

TABLE 2
Morphometric Parameters of three lateral lakes of the lower Hunter River

Parameter ‘Birds’ L. ‘Bootlands’ L. ‘Murphys’ L.

Area (ha) 6.79 8.04 7.40
Volume (m? x10*) 9.10 14.80 8.44
Max Depth (m) Dol 3.9 2.4
Mean Depth (m) ASA: 1.84 1.14
Shoreline length (m) 2350 2890 2170
Shoreline Development 2ESD 2.87 2.20
Volume Development 1.59 1.42 1.42
Length (m) 460 625 655
Width (m) 305 170 205
Altitude (m) cal8 cal6 cal5

All measurements made at full lake level. See text and Bayly and Williams (1973) for explanation of the
parameters Shoreline Development and Volume Development.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

B. V. TIMMS 317

volume of a lake to that of a cone of basal area equal to the area of the lake and height
equal to the maximum depth of the lake) in each is relatively high in keeping with the
steep littoral area and flat floor. In ‘Birds’ (Fig. 3a) and ‘Bootlands’ the deepest point is
well removed from the levee dam, but in ‘Murphys it is close by (Fig. 3b).

(b) PHYSICOCHEMICAL PARAMETERS

All physicochemical data refer to the five lagoons (19-23) in the Gosforth series
which were studied over a 63-month period from October, 1979 to December, 1984.
Rainfall (see Fig. 4) was well below average during 1980 and again in late 1982 — early
1983, so these were drought years. There were some periods of high rainfall (e.g. Febru-
ary 1981, March 1982, Fig. 4) which produced significant local run-off, but there were
no river floods during 1979-84.

Levee

Fig. 3. Bathymetric map of a (above) Lagoon No. 4 ‘Birds’ and b (below) Lagoon No. 21, ‘Murphys. Contours
at 0.5m intervals, with some dashed ones at 0.25m intervals.

Mean TDS for the lagoons varied between 215 to 468mg |! (Table 3, Fig. 4). The
lowest value recorded was 87mg |! in Lagoon 22 and the highest was 2208mg |!" in
Lagoon 21. During the two droughts there were steady increases in TDS to unusually
high values, particularly in Lagoons 19b, 21, and 22. Fluctuations were a function of the
relative input of run-off and loss by evaporation (Timms, 1970a) as expressed by the sig-
nificant correlation between the ratio catchment area: lagoon surface area and annual

PROC. LINN. SOC. N.SW,, 109 (4), (1986) 1987

318 FLOODPLAIN WATERBODIES

1980 1981 1982 1983 1984

Rainfall (mm

a 8

Total Dissolved Solids (mg!)

1980 1981 1982 1983 1984

Fig. 4. Monthly rainfall (and long term average) at Singleton (nearest Meteorological Station) and TDS for
lagoons 19-23 for period October 1979 to December 1984. Gaps in the TDS curves indicate the lagoons were
dry for those periods.

percentage fluctuation in TDS (r=0.8239, n=6, P<0.05). Fluctuations in lagoon 19a
were much less than in 19b, yet both are sequentially located in the same valley (Fig. 5)
and joined during high water. This is explained in part, by lagoon 19b receiving propor-
tionally more run-off than 19a, but it also has salty springs along its western shore (R.
Bootland, personal communication).

Waters in the 5 Gosforth lagoons are of the sodium chlorobicarbonate type.
Cationic dominances were Na>Mg>Ca>K in all lakes, but anionic dominances
varied between HCO; >Cl>SO,, HCO; =Cl >SO, and Cl >HCO; > SO, (Table 4).
Generally all 5 lagoons were slightly alkaline with pH’s between 7 and 8 (Table 4). No
regular pattern of variation was evident. Unusually high pH values of 9.0-9.4 were seen
in Lagoons 21 and 22 in low rainfall summers and equally unusual pH values of 6.2-6.8
were measured in Lagoons 19b, 20 and 23 after large inflows.

Water temperatures varied seasonally from late winter lows of ¢. 12-13°C to sum-
mer highs of c. 27-32°C (Table 3). The extreme range was 10.8°C in Lagoons 19a to
34.4°C in Lagoon 23. All values, especially the maxima, were influenced by the time of
measurements. This explains the steady increase in values from 19 to 23, but even so

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

319

B. V. TIMMS

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PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

320 FLOODPLAIN WATERBODIES

lagoons 20 and 23 tended to be warmer in summer and colder in winter because of their
relative shallowness. During the summer months Lagoons 19a, and to a lesser extent
19b, 21 and 22, were often thermally stratified. In that bottom temperatures were only
occasionally constant or slightly elevated from one month to the next, it is likely stratifi-
cation rarely persisted between visits.

Turbidity also varied between and within the lagoons with mean values between 26
and 318 FTU’s (Table 3) and extremes ranging from 0 in Lagoon 21 to 1600 in Lagoon
23. Values in Lagoon 23 were largely unnatural, as cattle trampled in it particularly
during low water levels. The high values in Lagoon 19b may also be unnatural as Euro-
pean Carp, which are thought to muddy waters by their feeding activities (Tilzey, 1980),
were present. There was little pattern in turbidity variations though, in general, values
were highest following intense run-off. Algal blooms only had a noticeable effect in
Lagoon 19a where Oscillatoria sp. regularly bloomed in late summer.

Light penetration, as measured by Secchi disc depth, varied from a mean value of
30cm to 100cm. The lowest recorded value was Icm in Lagoon 23 and the highest 190cm
in Lagoon 19a. Mean values in Lagoons 20, 21 and 23 should be a little higher as oc-
casionally the disc reached the bottom before becoming obscured. Not surprisingly,
turbidity and Secchi disc values were negatively correlated (r= -0.7756, n=6, P>0.05).

Nutrients in the 5 lagoons were of the same order of magnitude, averaging c. 1.4.mg
1! NO3-N and c. 0.4mg 1! PO,-P (Table 3). Lagoon 23 had the highest values and
Lagoon 21 the lowest, with the former situation easily related to its use by cattle. Values
in Lagoon 20 were probably elevated by intermittent agriculture and use of fertilizers in
its catchment, and in Lagoon 19a the relatively high phosphates could be due to the 50-
100 commercial ducks kept in a partially submerged pen on one bank. Nutrient values
fluctuated erratically, but in lagoons which dried nutrients were elevated soon after
filling and often during low water periods as well.

DISCUSSION

(a) GEOMORPHOLOGY

Almost all of the lagoons studied fit the characteristics of the blocked valley lake
type of Blake and Ollier (1971) (= lateral lake type of Hutchinson, 1957). Typically they
are contained wholly or largely within small side valleys cut into Permian or Carbonifer-
ous sandstones, siltstones and mudstones or into high river terraces (e.g. No. 11) and
with their lower ends blocked by natural levees of the Hunter or Paterson Rivers. Many,
like Nos. 19 and 21 (Fig. 5) end abruptly at the floodplain — country rock junction, but
others project onto the floodplain of the main stream e.g. Nos 6 and 8. There is one, No.
38, that lies partly in a tributary valley and partly between the levee and the scarp de-
fining the edge of the flood plain (Fig. 1). Finally in this continuum, there are a few that
lie almost entirely within the alluvium of the floodplain and are dammed by the natural
levee of the main stream — embankment lakes of Blake and Ollier (1971).

Alluvium is often deposited in the form of an obvious levee, as it is at Nos 9 and 11
(Kaludah Lagoon), but in many cases the levee is complex, wide and of low uneven slope
e.g. along the Hunter River between Nos 19 and 26 (Fig. 5). Reverse delta formation is
apparent in some lagoons e.g. No. 4 (‘Birds’) (Fig. 3a) and in these the deepest point is
well away from the dam. In others no such fan of alluvium is present so that the dam
front is relatively steep and the deepest part is consequently near the dam e.g. No. 21
(‘Murphys’) (Fig. 3b).

Lagoon size is correlated significantly with catchment size (r=0.7305, n=42,
P>0.001), though catchments >600ha rarely contain lagoons and those below 75ha are

PROC. LINN. SOC. N.SW., 109 (4), (1986) 1987

B. V. TIMMS B32

Onze 900m 23 Lee ope

Fig. 5. Position of Lagoons 15-16 and 19-23 with reference to levee alluvium (stippled) of the Hunter River.

less likely too (Fig. 2b). This is probably because large catchments generate sufficient
run-off to keep the channel to the main river open and very small catchments do not
generate enough water to pond or more likely any depression is easily infilled with
sediment from the main river.

The geomorphology of the tributary valley substantially influences the depth,
shape and size of any lagoon in it. Lagoons in wide shallow valleys are typically shallow,
large and with low S.D. values (e.g. Nos 11, 21, and 26), whereas those in narrow steep-
sided valleys are usually smaller, deeper and more dendritic (e.g. Nos 19 and 38). The
contrast in the two extremes is seen in the one lagoon, No. 4, ‘Birds, where the deep
northeast arm is in a steep-sided gully while the shallower southeast arm is surrounded
by more gentle terrain.

Lagoons usually fill from local run-off following heavy rain, but rarely flood waters
from the main stream contribute also. Following such filling, water levels are probably
controlled largely by surface evaporation and by exchange with the regional watertable.
However, given the great salinity increase as some lagoons dry (see later), perhaps some
lagoons are at least partly perched. Although shallower lagoons are the ones most likely
to be ephemeral, sediment types and geomorphic setting are also important. Lagoons

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

Be. FLOODPLAIN WATERBODIES

largely on alluvium of sandy silt are less permanent than those in tributary valleys cut in
rock and abutting alluvium.

Major floods may change the alluvial dam and hence the geomorphic and hydro-
logic features of a lagoon. An example is No. 19 which before the 1955 Flood was a
marsh, but with the deposition of 5 to 10m of alluvium on the levee (telephone poles were
almost buried) it became a permanent lagoon. Similar episodic changes to floodplain
lakes on the Macdonald River near Sydney have been reported by Erskine (1986) and
Henry (1977). Artificial drains silt after floods and many landowners redig their drains
every few years.

(b) PHYSICOCHEMICAL ENVIRONMENT

Probably the most significant aspect of the physicochemical environment in these
lagoons is its variability. Basic to this is the variable hydrologic regime as determined by
their geomorphic position. The mean annual fluctuation in TDS of 177% is higher than
for most freshwaters in southeast Australia (Bayly and Williams, 1966; Williams, 1967),
but is nevertheless typical of small lentic waters such as billabongs of the Murray River
around Albury (Shiel, 1980) and farm dams, including those nearby (Timms, 1970b;
1980). In common with these environments, but in contrast to larger reservoirs and
lakes (Bayly and Williams, 1973; Powling, 1980; Timms, 1976) there are also great vari-
ations in pH, turbidity and nutrient levels. Such variations are influenced more by
irregular inflows from the catchment than by within-lake processes and are exacerbated
by the shallowness of the lagoons.

Other physicochemical characteristics of these lagoons indicate features typical of
Australian freshwaters either countrywide or regionally, or of small shallow lentic waters
such as river billabongs and farm dams. These include:

(a) TDS content in all is higher than usual for the region where only 12% of
waters exceed 225mg |! (Timms, 1970c). This is probably due to their closed
hydrologic regime for most of the time. Certainly, flushing after heavy local rain
reduces TDS levels (Fig. 4) and this happens on average for a few days in wet
years. The influence of rare river floods is unknown, but presumably flushing
occurs then also.

(b) Ionic composition is typical of Australian waters as a whole (Bayly and
Williams, 1973), though HCO; levels are relatively high, as is generally the case in
northeastern NSW (Timms, 1970c).

(d) Annual temperature range of surface waters is c. 15-20°C which is similar to
that reported for other small sites in lowland southern Australia (Shiel, 1980;
Timms, 1970b). Minima of 11-12°C are characteristic of low-altitude waters at this
latitude (33° S) (Shiel, 1980; Timms, 1970b). The perceived lack of persistent
stratification distinguishes these lagoons from many farm dams (Timms, 1980),
typical larger reservoirs and lakes (Powling, 1980; Timms, 1976) but not from
billabongs (Shiel, 1980). Any stratification in these lagoons is short-lived because
of their shallowness and exposure to winds. However, more detailed studies are
needed as there are some indications that at least one stratifies for weeks at a time,
and given its eutrophic status, extensive deoxygenation could occur.

(e) In common with farm dams (Timms, 1980) and most inland waterbodies
(Bayly and Williams, 1973) the waters of these lagoons are turbid. Consequently
Secchi disc depths of <1m, often <0.5m are characteristic.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

B. V. TIMMS Bue

(f) Nutrient levels, particularly phosphate, are higher, than for most freshwaters
in Australia (Bayly and Williams, 1973) but similar to values recorded in farm
dams (Timms, 1980). As discussed earlier at least some of these nutrients are
anthropogenically derived. Despite these high nutrient levels there is no evidence
that the lagoons are hypereutrophic; indeed some appear at most to be meso-
trophic based on evidence on their planktonic and benthic standing crops (author,
in preparation). It seems high turbidity and associated limited light penetration

limit production (Williams and Wan, 1972).

In summary then, while the physicochemical environment in these lagoons is
typical for Australia, it is distinctive by virtue ofits elevated TDS content, high turbidity
and nutrient levels. Most charcteristic of all though is the high variability in all
parameters.

Not one of the lagoons is unaffected by man. Indirectly, they are probably being
flushed less frequently because of the flood mitigation effect of Glenbawn Dam up-
stream on the Hunter R. (Erskine, 1985). More directly, most landowners consider the
lagoons occupy valuable grazing land, so they endeavour to drain them. Two-thirds of
the lagoons are affected in this way (Table 1) with the most drastic changes being seen on
the larger lagoons 1n wide shallow valleys. Other less obvious changes are wrought by (1)
cattle wading which increases turbidity in smaller lagoons and in larger lagoons during
low water periods and (11) by use of fertilizers on upstream catchments.

Nutrient levels are probably elevated in all lagoons, particularly in those with dis-
crete sources, e.g. duck pens. The effects of these changes on their ecosystems are
unknown. Nevertheless, while some eutrophication may have enhanced waterfowl
habitat in many lagoons, their natural value as drought refuges has been further
reduced by drainage. Fortunately, should the opinions of landowners change, the river
can right wrongs by depositing new alluvial dams or silting drains, so tributary valleys
can once more be effectively dammed to become permanent lagoons again.

ACKNOWLEDGEMENT

I wish to thank many of my geomorphology students for help in mapping the
lagoons, and in particular Glen King and the late Edu Nerinckx for assisting in the
initial survey and interviews as well. I am grateful to numerous landowners who allowed
access to their lagoons and spent time talking with us, and to the Director of the Bureau
of Meteorology for supplying rainfall data for Singleton. Dr I. A. E. Bayly of Monash
University and Wayne Erskine of University of N.S.W. are thanked for reviewing the
manuscript.

References

ANON., 1975. — Standard Methods for the Examination of Water and Wastewater 14th Edit. Washington: APHA.
BayLy, I. A. E., and WILLIAMS, W. D., 1966. — Chemical and biological studies on some saline lakes in
south-east Australia. Aust. J. Mar. Freshwat. Res. 17: 177-228.

, and , 1973. — Inland Waters and their Ecology. Melbourne: Longmans.

BLAKE, D. H., and OLLIER, C. D., 1971. — Alluvial plains of the Fly River, Papua. Z. Geomorph. N.F. Suppl.
Bd. 12: 1-17.

CowarRDIN, L. M., CARTER, V., GOLET, F. C., Classification of Wetlands and Deepwater Habitats of the United
States. Washington: U.S. Fish and Wildlife Service, Office of Biological Services.

ERSKINE, W. D., 1985. — Downstream geomorphic impacts of large dams: the case of Glenbawn Dam, NSW.
Applied Geogr. 5: 195-210.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

324 FLOODPLAIN WATERBODIES

——, 1986. — River metamorphosis and environmental change in the Macdonald Valley, New South Wales,
since 1949. Aust. Geogr. Stud. 24: 88-107.

HAKANSON, L., 1981. — A Manual of Lake Morphometry. Berlin: Springer-Verlag.

Henry, H. M., 1977. — Catastrophic channel changes in the Macdonald valley, New South Wales, 1949-
1955. J. Proc. Roy. Soc. N.S.W. 110: 1-16.

HUTCHINSON, G. E., 1957. — A Treatise of Limnology. Vol. 1. New York: John Wiley and Sons.

POWLING, I. J., 1980. — Limnological features of some Victorian reservoirs. Jn: WILLIAMS, W. D., (ed.), An
ecological basis for water resource management. Canberra: ANU Press.

RILEY, S. J., WARNER, R. F,, and ERSKINE, W., 1984. — Classification of Waterbodies in New South Wales. North
Sydney: N.S.W. Water Resources Commission.

SHIEL, R. J., 1980. — Billabongs of the Murray-Darling System. Jn: WILLIAMS, W. D., (ed.), An ecological basis
for water resource management. Canberra: ANU Press.

TILzEY, R. D. J., 1980. — Introduced Fish. Jn: WILLIAMS, W. D., (ed.), An ecological basis for water resource
management. Canberra: ANU Press.

TIMMS, B. V., 1970a. — Variations in the water chemistry of four small lentic localities in the Hunter Valley,
New South Wales, Australia. Aust. Soc. Limnol. Bull. 3: 36-9.

—, 1970b. — Aspects of the limnology of five small reservoirs in New South Wales. Proc. Linn. Soc. N.S.W.
95: 46-59.

—, 1970c. — Chemical and zooplankton studies on lentic habitats of north-eastern New South Wales. Aust.
J. Mar. Freshwat. Res. 21: 11-33.

—., 1976. — A comparative study of the limnology of three maar lakes in Western Victoria. I. Physi-
ography and physiochemical features. Aust. J. Mar. Freshwat. Res. 27: 35-60.
—, 1980. — Farm Dams. Jn: WILLIAMS, W. D., (ed.), An ecological basis for water resource management.

Canberra: ANU Press.
WILLIAMS, W. D., 1967. — The chemical characterisation of lentic surface waters: a review. In: WEATHER-
LEY, A. H., (ed.), Studzes on Australian Inland Waters and their Fauna. Canberra: ANU Press.
, and WAN, H. F., 1972. — Some distinctive features of Australian inland waters. Wat. Res. 6: 829-36.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

Records of Eudendrium (Hydrozoa: Hydroida)
from New Zealand

JEANETTE E. WATSON

WatTSON, J. E. Records of Eudendrium (Hydrozoa: Hydroida) from New Zealand. Proc.
Linn. Soc. N.S.W. 109(4), (1986)1987: 325-330.

Three species of Eudendrium are recorded from New Zealand, all from the North
Island. These are EF. novaezelandiae Marktanner-Iurneretscher, 1890, E. terranovae Watson,
1985 and E. ritcher Millard, 1975. The holotype of E. novaezelandiae has been re-examined
and the description amplified from study of fresh material. The doubtful record of E.
novaezelandiae from North Cape by ‘Totton (1930) is now known to be E. terranovae, while E.
insigne Hincks, 1861, reported from the east coast (Ralph, 1953), is here referred to E. ritchev.
This is the first record of the latter species outside South Africa.

Jeanette E. Watson, Honorary Associate, Museum of Victoria, Melbourne, Australia 3000; manuscript
received 8 April 1986, accepted for publication 20 May 1987.

INTRODUCTION

Records of the genus Eudendrium are comparatively rare in the New Zealand hydroid
literature. The earliest record is of Eudendrium novaezelandiae Marktanner-Iurneretscher,
1890 from Auckland, the holotype of which is lodged in the Naturhistorisches Museum of
Vienna. This species was included by Farquhar (1896) in his checklist of New Zealand
hydroids. There are no other references to Eudendrum in the literature of early authors (e.g.
Hutton, 1873; Coughtrey, 1875; 1876; 1876a; Thompson, 1879; Hilgendorf, 1897;
Hartlaub, 1901; Bale, 1924; Trebilcock, 1928).

Totton (1930), doubtfully assigned to E. novaezelandiae a specimen collected at North
Cape by the Terra Nova Expedition (1910-1913). Comparison of the cnidome of ‘Totton’s
material with that of the holotype (Watson, 1985) showed the North Cape material not to
be E. novaezelandiae but a new species of Eudendrium. This species, named E. terranovae
Watson, 1985, isa common oceanic hydroid of the southeastern Australian coast. The sys-
tematics, ecology and distribution of E. terranovae are fully described by Watson (1985:
189-191).

The remaining reference to Eudendrium in the New Zealand literature is a very brief
description by Ralph (1953) of a hydroid which she assigned to E. insigne Hincks, 1861, from
the east coast of New Zealand.

These records are all from the North Island; the genus has not yet been recorded from
the South Island. Judging by the number of species of Eudendrium now known from com-
parable latitudes and habitats in Australian waters (Watson, 1985), the New Zealand fauna
should, however, include many more species than are known at present.

A small collection of hydroids made by the author, using SCUBA, from the Coroman-
del Peninsula (37°48’S, 175°30’E) in the North Island in February 1983, yielded two
species of Eudendrium. One, identified as Eudendrium novaezelandiae Marktanner-
Turneretscher, 1890, now provides a basis for redescription of that species. The other,
referred by Ralph (1953) to E. znszgne Hincks, 1861, is here identified with E. ritche: Millard,
1975. This is the first record of E. rztchei outside southern Africa.

Material of each species is lodged in the Museum of Victoria (MVF) and held in the
personal collection of the author.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

326 EUDENDRIUM FROM NEW ZEALAND

SYSTEMATIC ACCOUNT

Eudendrium novaezelandiae Marktanner-Iurneretscher, 1890

Eudendrium novaezelandiae Marktanner-Iurneretscher, 1890:201, pl. 3, fig. 21.

non £. novaezelandiae Marktanner-Iurneretscher. Totton, 1930:141.

Material Examined: Holotype, AN12389, Naturhistorisches Museum of Vienna. Other

material, MVF51780, male colony on shell of Atrina zelandica, Coromandel, Haurakai Gulf,

8m deep, bottom gravel and mud, coll: J. E. Watson, 20/2/83.

Description: The holotype material comprises an alcohol preserved distal fragment of a

larger female colony. The stem is about 6cm in height and is broken into two pieces. The

following description of the holotype supplements that of Marktanner-Iurneretscher.

The stem is branched irregularly in various planes, the main branches being fascicled
almost to their tips, perisarc smooth and shining. The distal branches are roughly alternate,
passing upwards at an acute angle, with up to seven indistinct proximal annulations and
additional groups of annulations occurring at intervals along the distal branches; hydranth
pedicels annulated throughout. There are no remaining hydranths. The gonophores are
mature and are borne in groups of up to five scattered along a blastostyle devoid of ten-
tacles. The single egg is enclosed in a thick transparent pellicle.

Nematocysts of three sizes present but poorly preserved; none fully discharged:

(3) —microbasic euryteles (probably from tentacles), capsule 5-6 x 2-3, abundant in
patches in pedicels of hydranths,

(ii) large microbasic euryteles, capsule bean-shaped, 17 x 8u, abundant in pedicels of
gonophores,

(111) microbasic euryteles, smaller than above, 11-13 x 5-6, capsule bean-shaped. Pre-
sent in the pedicel of the female gonophore and possibly on the gonophore itself.

The colony from Coromandel is 30mm in height, comprising two separate stems
arising from the same hydrorhizal plexus. Main stems 0.7mm in proximal width, lightly
fascicled to about half the height of colony. Branches alternate, proximal width 0.1-0.13mm,
rebranching in the same plane with up to eight distinct annulations at origin of branches,
perisarc otherwise smooth and shining. Pedicel of hydranth long, distal width 0.13-0.15mm.

Hydranth large, with about 20 tentacles, a nematocyst ring on the lower body, hypostome

elongate and clavate, maximum width below tentacles (preserved material) 0.3mm. Male

gonophores immature, blastostyle without tentacles.

Nematocysts of two sizes present:

(1) small microbasic euryteles, 6-7 x 3y, shaft 5-6y long, with a few spines, moderately
abundant on tentacles, :

(ii) larger microbasic euryteles, capsule 18-19 x 8-10u, bean-shaped, shaft 11-15 long,
spines not clearly visible, thread with fine bristles. Abundant in nematocyst ring; a
few present on the hypostome and on gonophore.

Colour: Living colony cream coloured, stems dark brown proximally, becoming lighter

distally.

Remarks: The main differences between the Coromandel material and the holotype of £.

novaezelandiae are the flatter and more regular branching, the more regular appearance, and

the absence from the cnidome of the third, intermediate-sized microbasic eurytele. The
larger and smaller microbasic euryteles agree well with those of the holotype. The absence
of the intermediate-sized nematocysts from the present specimen may be due to association
with the female gonophore only, in a similar manner to that noted in E. generale von

Lendenfeld, 1885, by Watson (1985). All other characters agree well with those of the

holotype.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

JEANETTE E. WATSON 327,

Figs 1-4. Eudendrium novaezelandiae. 1. — Distal part of branch of holotype colony (AN12389, Naturhistorisches
Museum of Vienna) showing clusters of female gonophores. 2. — Whole stem of specimen from the Hauraki
Gulf, New Zealand. 3. — Cluster of mature female gonophores drawn from holotype. 4. — Nematocysts: A,
tentacular microbasic euryteles from holotype; B, medium-sized, undischarged microbasic eurytele from female
gonophore of holotype; C, large microbasic eurytele, discharged, from specimen from the Coromandel Peninsula.

Eudendrium ritchei Millard, 1975
Eudendnum ritche: Millard, 1975:87, fig. 30.
non E. insigne Hincks, 1861. Ralph, 1953:63, pl. 1, fig. 2A, 2B.
Material Examined: MVF51781, male colony, MVF51782, female colony; east coast of
Coromandel Peninsula, on brown algae, 2m deep, coll: J. E. Watson, 21/2/83. British
Museum (Natural History) alcohol preserved specimen 1912.12.21.90 labelled £. inszgne
Hincks, 1861.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

328 EUDENDRIUM FROM NEW ZEALAND

Description: Colonies growing luxuriantly on algal holdfast. Stems up to Icm in height
and 0.13-0.18mm in width, unfascicled, irregularly branched, arising from a smooth
reticulating hydrorhiza. Stems completely and closely annulated throughout, perisarc very
thick. Hydranths terminal on branches, with 20-24 tentacles.

Male and female gonophores borne on different but closely associated colonies on
algal substrate. Immature male gonophore with an apical tubercule, mature gonophore 1-2
chambered, distal 5 chamber 0.14-0.2mm in diameter, borne on lower stems of colony in
a tight cluster of 20-30 on a blastostyle devoid of tentacles at all stages. Young female gono-
phore with a strongly bifurcated spadix; gonophores borne in clusters of 4-6 in various
stages of development below a hydranth with a reduced number of tentacles. At maturity,
the tentacles are completely absent, with 2-5 mature gonophores scattered along the
blastostyle. Length of mature female gonophore 0.3mm, width 0.25mm.

Nematocysts of two kinds present:

(i) | microbasic euryteles, capsule 6-7 x 2.5, shaft 54 long, abundant in tentacles, few
discharged,

(ii) larger microbasic euryteles, capsule bean-shaped, 16.5-18 x 7-8y, shaft thick, at least
37 long, spinous, the thread very long and covered with bristles. Abundant on
hydranth, spadix of female, and on apical tubercule of male gonophore.

Colour: Hydranths and gonophores cream coloured, perisarc dark shining brown.

Remarks: Although the material upon which Ralph’s (1953) identification was based has

not been examined, her description, figure and locality notes leave no doubt that specimens

collected by the present author on the east coast of the Coromandel Peninsula are identi-

cal with those recorded by her as E. inszgne Hincks, 1861. Comparison of the cnidome of a

specimen of fresh material from the Hauraki Gulf with the British Museum material of E.

insigne shows that Ralph’s material is not this species.

The New Zealand specimens collected by the author agree well in colony morphology
with Millard’s (1975) description of Eudendnum ntcher, including important structures such
as the bifurcated female spadix and the strongly annulated stems. They differ from the
South African species in the smaller size of the mature colony and in having strictly
monosiphonic stems. Although the tentacular and supplementary nematocysts (Watson,
1985) of the New Zealand specimens are similar to those described and figured by Millard
for E. ritcher, there are, however, certain differences, namely the greater length-width (L/W)
ratio (Kubota, 1976; Watson, 1985) and the greater ratio of length of shaft to capsule (S/C)
in the southern African material. These ratios are compared below:

New Zealand S. Africa
Capsular L/W
Tentacular ratio 2.8:1 Mee /} alk
microbasic
euryteles
S/C ratio 0.8:1 0.8:1
Capsular L/W
Supplementary ratio Peso Poge sll Zn 2eoel
nematocysts
S/C ratio = Pil Mell 3), 2473
(not fully

discharged)

It has been suggested that variations may occur in the relative size and dimensional
ratios of nematocysts across the geographical range of a single species of Eudendrium

PROC. LINN. SOC. N.SW., 109 (4), (1986) 1987

JEANETTE E. WATSON 329

we, ))

SMI,

Ua)

Figs 5-8. Eudendrium ritche: from the east coast of Coromandel Peninsula. 5. — Single stem from female colony.
6. — Cluster of mature male gonophores. 7. — Cluster of female gonophores in various stages of maturity. 8. —
Nematocysts: A, tentacular microbasic eurytele, discharged; B, microbasic eurytele showing distal end of shaft.

(Kubota, 1976; Watson, 1985). The differences in size between the supplementary
nematocysts of the southern African and the New Zealand specimens further support this
view. Thus, until the limitations of variability of nematocysts within a species is established,
the New Zealand specimens are assigned to E. ritchev.

Millard classified the larger (supplementary) nematocysts of E. ritcher as macrobasic
euryteles. However, according to her definition of nematocysts (Millard, 1975: 21), they
should be considered microbasic euryteles since, according to her figure, the shaft is less
than four times the length of the capsule. Although no fully discharged shafts were found
in the New Zealand material, the S/C ratio would clearly be less than four at full eversion.

PROG. LINN. SOC. N.S.W., 109 (4), (1986) 1987

330 EUDENDRIUM FROM NEW ZEALAND

ACKNOWLEDGEMENTS

I wish to thank Dr Paul Cornelius of the British Museum (Natural History) and the
Director of the Naturhistorisches Museum of Vienna for loan of reference and type
material.

References

BALE, W. M., 1924. — Report on some hydroids from the New Zealand coast, with notes on New Zealand
Hydroida generally supplementing Farquhar’s list. Trans N.Z. Inst. 55: 225-268.

COUGHTREY, M., 1875. — Notes on the New Zealand Hydroideae. Trans Proc. N.Z. Inst. 7: 281 -302.

—., 1876. — Critical notes on the New Zealand Hydroida. Trans Proc. N.Z. Inst. 8: 298-302.

——.,, 1876a. — Critical notes on the New Zealand Hydroida. Ann. Mag. nat. Hist. 8 (4): 22-32, pl. 3.

FARQUHAR, H., 1896. — List of the New Zealand Hydroida. Trans Proc. N.Z. Inst. 28: 459-468.

HARTLAUB, C., 1901. — Hydroiden aus dem Stillen Ocean. Zool. Jb. (Syst.) 14: 349-379.

HILGENDORF, F. W., 1898. — On the hydroids of the neighbourhood of Dunedin. Trans Proc. N.Z. Inst. 30: 200-
218, pls 16-22.

HINcKS, T., 1861. — A catalogue of the zoophytes of South Devon and South Cornwall. Ann. Mag. nat. Hist. 8(3)
(1861): 152-161, 251-262, 290-297, 360-366; 9 (1862): 22-30.

Kusota, S., 1976. — Notes on the nematocysts of Japanese hydroids. J. Fac. Sct. Hokkaido Univ. (Zool.) 20(2):
230-243.

HuTTon, F. W., 1873. — On the New Zealand sertularians. Trans Proc. N.Z. Inst. 5: 256-259.

MARKTANNER-I URNERETSCHER, G., 1890. — Die Hydroiden des k.k. naturhistorischen Hofmuseums. Ann.
naturh. Mus. Wien 5: 195-286.

MaARISCAL, R. N., 1974. — Nematocysts. Jn: MUSCATINE, L., and LENHOFF, H. M., (eds), Coelenterate Biology:
129-178. New York: Academic Press.

MILLARD, N. A. H., 1975. — A monograph of the Hydroida of southern Africa. Ann. S. Afr. Mus. 68: 1-513.

RALPH, PATRICIA M., 1953. — A guide to the athecate hydroids and medusae of New Zealand. Tuatara 5: 59-75.

THOmpsoN, D’A. W., 1879. — On some new and rare hydroid zoophytes (Sertulariidae and Thuiariidae) from
Australia and New Zealand. Ann. Mag. nat. Hist. 3(5): 97-114, pls 16-19.

ToTToNn, A. K., 1930. — Coelenterata. Part V. — Hydroida. Nat. Hist. Rep. Br. antarct. Terra Nova Exped. 5:
131-252.

TREBILCOCK, R. E., 1928. — Notes on the New Zealand Hydroida. Proc. Roy. Soc. Vict. 41 (n.s.)(1): 1-31.

WaTSON, J. E., 1985. — The genus Eudendrium (Hydrozoa: Hydroida) from Australia. Proc. Roy. Soc. Vict. 97:
179-221.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

Description of a new Species of freshwater
Hardyhead Craterocephalus kailolae
(Pisces: Atherinidae) from Safia, northeastern
Papua New Guinea

W. IVANTSOFF, L. E. L. M. CROWLEY and G. R. ALLEN

(Communicated by P. SELKIRK)

IVANTSOFF, W., CROWLEY, L. E. L. M., & ALLEN, G. R. Description of a new species of
freshwater hardyhead Craterocephalus kailolae (Pisces: Atherinidae) from Safia, north-
eastern Papua New Guinea. Proc. Linn. Soc. N.S.W. 109(4), (1986) 1987: 331-337.

Craterocephalus kailolae is described from specimens collected in Foasi Creek, Safia,
northeastern Papua New Guinea. The new species is considered to be related to members
of the C. eyreszz species group. C. kazlolae is the only freshwater hardyhead known from the
northeastern drainages of Papua New Guinea as well as being the only representative of
the C. eyresti group known from outside Australia . It can be distinguished from other
species of Craterocephalus by a spatulate maxilla and other osteological characters and a com-
bination of morphological features. The zoogeography of this group in relation to this dis-
tribution 1s discussed briefly. As no literature exists on the species composition of the genus
Craterocephalus, its members and their authors are given.

Walter Ivantsoff and L. E. L. M. Crowley, School of Biological Sciences, Macquarie University, North
Ryde, Australia 2113, and Gerald R. Allen, Department of Ichthyology, Western Australian Museum,
Francts Street, Perth, Australia 6000; manuscript recerved 25 November 1986, accepted for publication
20 May 1987.

INTRODUCTION

The predominantly freshwater genus Craterocephalus, is endemic to Australia (Merrick
and Schmida, 1984) and Papua New Guinea. In the most recent revision, (Ivantsoff, 1978)
divided the genus into two species groups (see also Patten, 1978), ‘C. eyresi’and and C.
stercusmuscarum: Recent work by Crowley and Ivantsoff (unpublished) suggests that a third
group, which includes the marine and estuarine species, can also be recognized. The C.
eyresiv’ group comprises C. eyresz: (Steindachner, 1884). C. cuneiceps Whitley 1944, C. marjoriae
Whitley 1948 as well as two species recently described by Ivantsoff et al. (1987). The C. ster-
cusmuscarum’ group comprises C. stercusmuscarum (Gunther, 1867) which includes spotted and
unspotted subspecies (Ivantsoff et al., 1987). C. nouhuysz (Weber, 1910), C. randi Nichols and
Raven 1934, C. lacustris Trewavas 1940, C. dalhoustensis Ivantsoff and Glover (1974) anda
new species from northern Australia (Ivantsoff et al., 1987; Allen, 1982). The ‘C. honoriae’ or
the marine/estuarine group includes the remaining known species: C. honoriae (Ogilby,
1912), C. pauciradiatus (Gunther, 1861) and C. capreoli Rendahl! 1922 (now regarded as
distinct, see Potter et al., 1986).

Two related monotypic genera Quzrichthys Whitley 1950 and Allanetta Whitley 1943 are
under review and on present evidence (Crowley and Ivantsoff, unpublished) will probably
be included in the synonymy of Craterocephalus.

The new species, C. kazlolae, which is herein described, shows morphological, meristic
and osteological characteristics which align it with the ‘C. eyresi’ group. It is the only
representative of that group to occur in Papua New Guinea.

MATERIALS AND METHODS

The specimens in this study were collected from a quiet backwater using a small seine

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

332 FRESHWATER HARDYHEAD

10mm

Fig. 1. Holotype Craterocephalus kailolae. AMS 1.24640-001.

net of about 20mm mesh size which provided an adequate sample in a short time of
collecting.

The methods of counting and measuring were modified from Munro (1967) and were
as described by Prince et al., (1982) and Patten and Ivantsoff (1983). In Craterocephalus spe-
cies, the second dorsal and anal fins may or may not have an unbranched ray preceding
branched rays. For the sake of uniformity, the first ray following the spine is not considered
as part of the branched ray count throughout this work. The vertebral counts were obtained
by radiography and the osteological studies were made on alizarin specimens prepared
using standard techniques developed by ‘Taylor (1967). Morphometric measurements and
meristic counts were recorded for 31 specimens, designated holotype and paratypes (Table
1). These specimens are now deposited in The Australian Museum, Sydney. N.S.W.
(AMS); Western Australian Museum, Perth, W.A. (WAM); Kanudi Fisheries Research
Station, Kanudi, Papua New Guinea, (KFRS); Museum of Zoology, Ann Arbor,
Michigan, U.S.A. (UMMZ).

DESCRIPTION

Craterocephalus kailolae sp. nov.
(Fig. 1)

Holotype — AMS 1.24640 - 001. 57.5mm standard length (SL) collected with a small
seine; type locality, 3km west of Safia airstrip in still backwater of Foasi Creek , Papua New
Guinea 9°36’S, 148°37’E, collected by Walter Ivantsoff and John Paska. September 16.
1985.
Paratypes, 30 (25.2-52.3), locality as for holotype, collected by G. R. Allen using small
seine, September 8, 1982.
WAM P27783-001 (11 + 3 alizarin specimens); AMS I1.24640-002 (10); KFRS F.5390.01
(3); UMMZ 213857 (3)
Overall size range 25.2-57.5mm SL. Measurements expressed as proportions and counts
for the holotype and 30 paratypes are presented in Table 1.

DIAGNOSIS

Distinguished from all other species and subspecies of Craterocephalus by the combin-
ation of the following characters: small, moderately robust fish with seven rows of trans-
verse scales with two rows above, one covering, and four below midlateral band. Midlateral
scales 31-34, interdorsal scales 5-7. Mouth small, gape restricted by labial ligament. Gill
rakers in first lower gill arch 8-10, those in angle of first arch (1-2) slightly elongated, others

PROC. LINN. SOC. N.SW., 109 (4), (1986) 1987

W. IVANTSOFF, L. E. L.M. CROWLEY AND G.R. ALLEN 333

TABLE 1

Measurements (expressed as proportions) and counts of the holotype and 30 paratypes of Craterocephalus kailolae from Safa,
Papua New Guinea

Holotype 30 Paratypes SD
SL 57.5mm 25.2-52.3mm
In SL

mean range

Head 3.8 3.6 (3.3-4.0) oll
Recle 5.0 49) (4.6-6.0) 35
H max 4.3 4.0 (3.6-4.5) 21
H min 9.8 9.5) (8.8-10.2) 31
Sn-ODi1 2.0 2.0 @9=271) .06
Sn-OD2 1.4 1.4 (1.3-1.5) .04
Sn-OV 223) Dee (Cae;S))) .08
Sn-TV 1.8 1.7 (1.6-1.8) 05
SN-OA 1.5 9) (1.4-1.5) 05
Sn-TA 1.3 1.3 @251°3) .05
In Head
Eye B45 3k (3.0-3.8) 18
Interorbital ed 2.6 (2.4-2.8) 12
Postorbital 2.3 2.3 (1.9-2.5) a2
In Eye
Snout 1.0 1.1 (0.9-1.5) 13
Premaxilla 1.0 1.0 (0.9-1.2) 07
Dorsal process of premaxilla 1.1 1.2 (1.0-1.5) a2
Scale counts
Midlateral 34 S3e2 (31-34) 82
Transverse 7 7.0 — =
Predorsal 12 12.5 (11-14) 90
Interdorsal 6 6.0 (5-7) .26
Fin elements
First dorsal spines 6 5.6 (4-7) .63
Second dorsal branched rays 6 6.2 (5-7) 48
Anal branched rays 8 8.2 (7-9) 56
Pectoral branched rays 11 12.0 (11-13) .68
Position of fins
OD1 to TV F4.0 193.83 (285-455) 50
OD!1 to TPec B1.5 B1.4 (0-B2.5) 57
OV to TPec F1.0 F1.7 (0-F3) .68
Other values
Gill rakes in first lower gill
arch 8 8.8 (8-10) .55
Position of anus to TV B1.0 B1.0 (0-B3) 78

Vertebrae 34 SA (33-35) .69
* 17 specimens.

Abbreviations used in table: SL, standard length; Pec L, length of longest pectoral ray; H max, greatest body depth;
H min, least body depth at caudal peduncle; Sn, snout; OD1, origin of first doral fin; OD2, origin of second dorsal
fin; OV, origin of ventral fins; TPec, tips of pectoral fins; TV, tips of ventral fins; OA, origin of anal fin; TA, point
of last ray insertion of anal fin. Position of fins and anus is expressed as a number of scales in front (F) or behind
(B) point of reference. SD, standard deviation.

short, rounded and with spinules. Dorsal process of premaxilla long, extending into in-
terorbital space. Anterior process of maxilla broad, spatulate, almost meeting its opposite
at midline (Fig. 2). Palatine cylindrical, without pointed dorsal end. Branchiostegals 5.

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

334 FRESHWATER HARDYHEAD

Canals tubular and closed on nasals, anterior infraorbitals and postemporals; those on
frontals and temporals partially closed.

DESCRIPTION

Meristic counts and morphometric proportions for the holotype and paratypes are
presented in Table 1.

Small, moderately robust fish, largest specimen known 57.5mm SL. Dorsal profile
somewhat rounded, continuing in unbroken curve from origin of first dorsal to snout. Lips
thick, mouthparts protrusible. Premaxilla almost never reaching vertical through anterior
margin of orbit. Dorsal process of premaxilla long, reaching into interorbital space. Upper
jaw with two rows of teeth pointing posteriorly. Ramus of dentary highly elevated pos-
teriorly. Anteriorly, dentary expanded, forming wide edentulous plate. Other elements of
mouth edentulous also. Pharyngeal teeth sharp and fine, never molariform. Body scales
moderately large, scalloped and prominent, dorsoventrally elongated with circuli complete.
Single large interorbital scale with one smaller scale on either side always present. Pre-
opercle scaled.

Intercalars large. Anterior infraorbitals reduced, sometimes fused (Figs 3, 4). Large
dorsal and ventral postcleithra present. Urohyal with well developed dorsal plate, reduced
ventral plates, ventral pocket absent. Basihyal bone and cartilage about equal. Other
osteological features similar to other members of the ‘C. eyveszz’ group as described later.

COLOUR

Preserved specimens yellow brown above silvery midlaterlal band and pale yellow
below. Scales on dorsal surface and sides outlined with melanophores . Scales in row im-
mediately above midlateral band pigmented only to end of first dorsal fin along side of
body. Body below midlateral band unpigmented. Spine and rays of first and second dor-
sal and caudal fins with rows of melanophores, other fins unpigmented. Dorsum of head,
snout and lips peppered with melanophores. Posterior border of orbit outlined with
melanophores. Live specimens yellow brown with silvery band, not distinctly different
from those preserved.

ETYMOLOGY

kailolae. Named after Mrs Patricia Kailola, a major contributor to the knowledge of
ichthyology of Papua New Guinea. Without her help much of the work on Papua New
Guinea species of Craterocephalus would have been very difficult.

RELATIONSHIPS WITH OTHER SPECIES OF CRATEROCEPHALUS

Craterocephalus kailolae is a member of the ‘C. eyresi’ group (Ivantsoff, 1978; Patten,
1978). The group can be identified by the following characters: gut elongate, longer than
body length; small finger-like epiotic crest; unbranched posterior myodome extending into
basioccipital but without posterior opening; urohyal with ventral plates reduced; lower
pharyngeals close but not fused and with sharp non-molariform teeth; mesopterygoid
small; posterior edge of coracoid rounded, medial shelf reduced; scapular foramen large:
interdorsal pterygiophores usually weakly developed or absent; anal plate small with no
anterior elongation. Colour of midlateral band usually silvery in contrast to members of
stercusmuscarum group where band darkly pigmented.

DISTRIBUTION AND ZOOGEOGRAPHY

Craterocephalus kailolae is at present known only from the type locality in the north-
eastern highlands of Papua New Guinea. It appears to be present in relatively large

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

W. IVANTSOFF, L. E. L.M. CROWLEY AND G.R. ALLEN 359

Figs 2-4. Craterocephalus kailolae — Paratype WAM P27783-001. 2) upper jaw elements; 3) first and second infra-
orbitals — right side; 4) first and second infraorbitals fused — left side. Scale line represents 1.00mm.

numbers in slower flowing branches of the main stream, in very shallow water, often only
several centimetres deep. It also occurs in backwaters which may be shallow or deep, murky
or clear.

PROC. LINN. SOC. N.S.W.,, 109 (4), (1986) 1987

336 FRESHWATER HARDY HEAD

The location of this species is of great interest since all other hardyhead species from
Papua New Guinea occur in southern drainages. These are C. randi, extending from the Fly
River to the Port Moresby area, C. lacustris, at present known only from Lake Kutubu, but
possibly also occurring in the Kiori and Wago Rivers draining the lake to the south, and C.
nouhuyst, occurring in the Lorenz River, Irian Jaya, near the border of Papua New Guinea
and possibly in the upper tributaries of the Fly River near Tabubil. These species are all
members of the ‘C. stercusmuscarum’ group. Their closely related counterparts, C. stercusmus-
carum stercusmuscarum and C. species B (Allen, 1982) occur on the other side of the Arafura Sea
and Torres Strait. C. marjoriae, C. marianae Ivantsoff et al., 1987 and C. species A (Allen, 1982)
also occur in the northern coastal areas of Australia but there are no known south coast
representatives of the eyzesiz’ group in Papua New Guinea.

It appears that speciation of the genus Craterocephalus may have been slow, unlike the
Melanotaeniidae, another speciose group of freshwater fishes common to both Papua New
Guinea and Australia (Allen and Cross, 1982). C. kazlolae must have been separated from
its north Australian relatives 2-5 million years ago, that is, since the uplifting of the high-
lands in Papua New Guinea during Plio/Pleistocene (Veevers, 1984). The members of the
‘C. stercusmuscarum group, on the other hand, could still have been sympatric 7-10,000 years
ago, prior to the last transgression on ‘Torres Strait, thus allowing for gene flow to continue
and maintain similarity. C. randi and C. s. stercusmuscarum for example are morphologically
and osteologically very close. The fact that C. kazlolae is readily identifiable as a member of
the C. eyresw’ group (although morphologically distinct from other members) supports the
proposal that members of the genus Craterocephalus do not speciate rapidly.

ACKNOWLEDGEMENTS

We thank John Paska of Port Moresby who helped us to collect the fish, and Mr
Patrick O’Connor, the manager of Bulmacau Station in Safia who gave us shelter and
allowed us to collect in Foasi Creek. We also thank Miss Betty Thorn for her drawing of the
holotype and Basim Said and John Patten for their help with osteology and comments on
the relationships within the genus Craterocephalus. Dr D. Hales and Dr J. Bassett are thanked
for reading and commenting on the manuscript.

References

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, and Cross, N. J., 1982. — Raznbowfishes of Australia and Papua New Guinea. London: Sydney: Angus and
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GUNTHER, A., 1861. — Catalogue of the acanthotperygian fishes in the British Museum. Vol. 3. London: British
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, 1867. Additions to the knowledge of Australian reptiles and fishes. Ann. Mag. Nat. Hist., 20: 45-68.

IVANTSOFF, W., 1978. — Taxonomic and systematic review of the Australian fish species of the family Ather-

inidae with references to related species of the Old World. North Ryde: Macquarie University, Ph.D.
thesis, unpubl.
——., CROWLEY, L. E. L. M., and ALLEN, G. R., 1987. — Description of three new species and one subspecies
of freshwater hardyheads (Pisces: Atherinidae: Craterocephalus) from Australia. Rec. West. Aust. Mus., 13(2):
171-188.
, and GLOVER, C. J. M., 1974. — Craterocephalus dalhousiensis new species, a sexually dimorphic freshwater
teleost (Atherinidae) from South Australia. Aust. Zool. 18(2): 88-98.
MERRICK, J. R., and SCHMIDA, G. E., 1984. — Australian Freshwater Fishes. Biology and Management: 143-152.
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Munro,1.S.R., 1967. — The Fishes of New Guinea. Port Moresby: Dept. Agr., Stock and Fisheries, 650 pp.

NICHOLS, J. T., and RAVEN, H. C., 1934. — Two new freshwater fishes (Percesoces) from New Guinea. Amer.
Mus. Novit. 755: 1-4.

OGILBy, J. D., 1912. — On some Queensland fishes. Mem. Qld Mus. 1: 26-65.

PATTEN, J. M., 1978. — Osteology, relationships and classification of hardyheads of the subfamily Atherininae
(Pisces: Atherinidae). North Ryde: Macquarie University, M.Sc. thesis, unpubl.

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, and IvANTSOFF, I., 1983. — A new genus and species of atherinid fish Dentatherina mercert from the western
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PRINCE, J. D., IVANTSOFF, W., and POTTER, I. C., 1982. — Atherinosoma wallaceii a new species of estuarine and
inland silverside (Teleostei: Atherinidae) from the Swan-Avon and Murray Rivers, Western Australia.
Aust. Zool. 21(1): 63-74.

RENDAHL, H., 1922. — A contribution to ichthyology of north-west Australia. Meddr Zool. Mus. Krist., 5: 163-197.

STEINDACHNER, F., 1884. — Beitrage zur Kenntniss der Fische Australiens. Sttzungsb. K. Akad. Wiss. Wien 88(1):
1065-1108.

TayLor, W. R., 1967. — An enzyme method of clearing and staining small vertebrates. Proc. U.S. Nat. Mus. 122:
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TReEwavas, E., 1940. — On new Papuan fishes. Ann. Mag. Nat. Hist. 6(33): 284-287.

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WHITLEY, G. P., 1948. — Ichthyological descriptions and notes. Proc. Linn. Soc. N.S.W. 68(3&4): 114-144.

—., 1944. — New sharks and fishes from Western Australia. Aust. Zool. 10(3): 252-273.

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PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

“am 2
; ws

Acacia suaveolens, biology of.............
AGUS, SAWS Oils so ag b0cnve sg odo abou 0006
INGENTEOSES soo conc ooo oem ooUdD HOD OOD NRS
Albertson, E. L., see Rowe, F. W. E.
Allen, G. R., see Ivantsoff, W.
Amycterinae, adult and immature.......
Anderson, D. T., The circumtropical bar-
nacle ‘etraclitella divisa (Nilsson-
Cantell), (Balanomorpha, Tetracliti-
dae): cirral activity and larval
Gevelopmentee rs cry aie: eens
Angophora hispida, post-fire demography... .
Aquatic angiosperms in coastal saline
lagoons of New South Wales. I. The
vegetation of Lake Macquarie, by R. J.
Irina peices eae eases hs Seis Mads ea
Aquatic angiosperms in coastal saline
lagoons of New South Wales. II. The
vegetation of Tuggerah Lakes, with
specific comments on the growth of
Zostera capricorn Ascherson, by R. J.
King & V. M. Holland.............
Aquatic angiosperms in coastal saline
lagoons of New South Wales. III.
Quantitative assessment of Zostera
capricorni, by R. J. King & J. B. Barclay
Aquatic angiosperms in coastal saline
lagoons of New South Wales. IV. Long-
term changes, by R. J. King & B. R.
IModgsomey ctor eetid ala a hysyekiete ee

Index

VOLUME 109
Page
271 Bulahdelah, N.S.W., dinoflagellate cysts. . 175
210
129

107

ENS EEMOIGE A fc tees Seiki ist alae tee eens 183, 195

Atlas of seeds and fruits from Macquarie
Island, by D. M. Bergstrom.........
Auld, T. D., Post-fire demography in the
resprouting shrub Angophora hispida
(Sm.) Blaxell: flowering, seed produc-
tion, dispersal, seedling establishment
ATAU SUGVAV all oeeesteiays ks sa ee ete staestleys)
Australian megafauna, extinction of.....

Balanomorpha, ‘Tetraclitidae...........

Barclay, J. B., see King, R. J.

Barlow, C. G., McLoughlin, R., & Bock,
K., Complementary feeding habits of
golden perch Macquaria ambigua
(Richardson) (Percichthyidae) and sil-
ver perch Bidyanus bidyanus (Mitchell)
(Teraponidae) in farm dams.........

Batswlone-canecam ey ir ieaca eee

Bergstrom, D. M., An atlas of seeds and
fruits from Macquarie Island........

Bidyanusipidyanusti tree ey ae

Bock, K., see Barlow, C. G.

Brayton, N.S.W., isotopic dating of granites

143
153

69
143

Cainozoic history of the vegetation and
climate of the Lachlan River region,
New South Wales, by H. A. Martin... 213
Carr, P FEF, & Jones, B. G, Non-

contemporaneity in the Marulan

Batholith sys seamen eaters a vats nt 63
Carter, J. J., Metagenesis as a possible key

toxanimialdfornmpmm nee reine 117
Chapmans Creek Granodiorite......... 65
Chiroptera: Vespertilionidae........... 153

Circumtropical barnacle Yeétraclitella divisa
(Nilsson-Cantell) (Balanomorpha,
Tetraclitidae), cirral activity and larval

development, by D. T. Anderson..... 107
Coastal saline lagoons, vegetation of 11, 25, 41, 51
Coleoptera: Curculionidae............. 91

Complementary feeding habits of golden
perch Macquaria ambigua (Richardson)
(Percichthyidae) and silver perch Bid-
yanus bidyanus (Mitchell) (Teraponidae)
in farm dams, by C. G. Barlow, R.
McLoughlin & K. Bock............. 143
Craterocephalus kailolae sp. nov............ 332
Crowley, L. E. L. M., see Ivantsoff, W.

Delma in Queensland, key to............ 211
Delmaglabialisispanovere nie rire chat 207
Delmarmitellarsp anova acura 204
Description of a new species of freshwater
hardyhead Crateroceohalus _—_ karlolae

(Pisces: Atherinidae) from Safia, Papua

New Guinea, by W. Ivantsoff, L. E. L.

M. Crowley & G. R. Allen.......... 331
Dinoflagellate cysts, Late Pleistocene..... 175
Distribution and taxonomy of the long-

eared bats Nyctophilus gould: Tomes,

1858 and Nyctophilus bifax Thomas, 1915

(Chiroptera: -Vespertilionidae) in

eastern Australia, by H. E. Parnaby... 153
Dracaena-ty pe @row thar alerts rte 129
Echinaster colemani sp. nov.......-....+-. 196
Echinaster in Australian waters........... 195

Echinoderm genus Henricia Gray, 1840
(Asteroidea: Echinasteridae) in south-
ern .and_ southeastern Australian
waters, with the description of a new
species, by F. W. E. Rowe & E. L.

(ANID ENtSOMM peahity nasa ewseisce eee eos aie es 183
Eudendrium novaezelandiae............... 326
IOOGORCTHR. TUBIB ss o00'bs 0060 don doe oo oR 327
Extinction of Australian megafauna...... i

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

Fergusson, C. L., Multiple folding of the
Ordovician sequence, Tambo River,
CAGISIIN WG yo vc osscodsndonaue

Floodplain lakes, Hunter Valley.........

Geomorphic and physicochemical features
of floodplain waterbodies of the lower
Hunter Valley, N.S.W., by B. V. Timms

Golden perch, feeding habits...........

Hardyhead, new species of freshwater... .
JEG RCER COMMA ooo 000 n8040 8005085"
JU FORYEHD 127 ONT 9), WON, poo 06 560 be 905
ELCNTICIs ODES ARR en Ae eee
Hodgson, B. R., see King, R. J.

Holland, V. M., see King, R. J.

Howden, A. T., Notes on the biology of
adult and immature Amycterinae
(Coleoptera: Curculionidae).........

Hunter Valley, N.S.W., floodplain lakes. . .

liv. drozoasbly droidameerenien ice er eae

Isotopic dating, Marulan Batholith......
Ivantsoff, W., Crowley, L. E. L. M., &
Allen, G. R., Description of a new
species of freshwater hardyhead
Craterocephalus kailolae (Pisces: Atherini-
dae) from Safia, Papua New Guinea. .

Jones, B. G., see Carr, P. F.

King, R. J., Aquatic angiosperms in coastal
saline lagoons of New South Wales. I.
The vegetation of Lake Macquarie....

King, R. J., & Barclay, J. B., Aquatic
angiosperms in coastal saline lagoons of
New South Wales. III. Quantitative
assessment of Zostera capricormt........

King, R. J., & Hodgson, B. R., Aquatic
angiosperms in coastal saline lagoons of
New South Wales. IV. Long-term
Chan gesey comtiat jeiy alte daninesu enue easels:

King, R. J., & Holland, V. M., Aquatic
angiosperms in coastal saline lagoons of
New South Wales. II. The vegetation of
Tuggerah Lakes, with specific com-
ments on the growth of Zostera capricorn
Aschersongynianti sci mire akucrn iad

acextilias sbyeopodidde! yaar rere:
Lachlan River region, N.S.W., Cainozoic
Vegetation and climaterry cee a
Lake Macquarie, aquatic vegetation.....
Late Pleistocene dinoflagellate cysts from
Bulahdelah, northern New South
Wales ibyeA Vicinity mara terete:
Linnean Society of New South Wales.
Record of annual general meeting 1985.
Reports and balance sheets. . annexure
Lockyersleigh Adamellite..............

Macleay Memorial Lecture 1984........

293
311

oli
143

331
186

192
190

91
311
325

25
203
213

17/3)

PROC. LINN. SOC. N.S.W., 109 (4), (1986) 1987

McLoughlin, R., see Barlow, C. G.
McMinn, A., Late Pleistocene dinoflagel-
late cysts from Bulahdelah, northern
INewsS Oultha Vall ese ee ee
Miacquaniaambvoaniyriets: see eee
Macquarie Island, seeds and fruits.......
Martin, H. A., Presidential Address 1982.
Cainozoic history of the vegetation and
climate of the Lachlan River region,
News Souths Valesm ees eee
Marulan Batholith, isotopic dating......
MemonialeSeniess:nowZ/e nao eee
Metagenesis as a possible key to animal
form byA)en|s Canter as = -i ese eeeerae

Mim osaceaci ees nigh eee ee ice

Morrison, D. A., A review of the biology of
Acacia suaveolens (Smith) Willd.
(Mimosaceae) serie ears

Multiple folding of the Ordovician

sequence, Tambo River, eastern Vic-
toma, by Grae hergussonty. cee ee

New light on the extinction of the Aus-
tralian megafauna, by R. V. S. Wright.
New species in the echinasterid genus
Echinaster Muller and Troschel, 1840
(Echinodermata: Asteroidea) from
southeastern Australia and Norfolk
Island, by F. W. E. Rowe & E. L.
Albertson) ij. jenstnieienaha ie Meee
New Zealand, records of Eudendrium.....
Notes on the biology of adult and immature
Amycterinae (Coleoptera: Curculioni-
dace) by Atlee owcentes seers
INyctophilusmbujaxn eee eee

Np atop nolusy could cen mascara site ee teenie
Ordovician sequence, ‘Tambo River,

WACO Tilia bie key ate Se ae
Palynology sis edict cuospspecieay cee AONE

Papua New Guinea, freshwater hardyhead
Parnaby, H. E., Distribution and taxon-
omy of the long-eared bats Nyctophilus
gould: Tomes, 1858 and Nyctophilus bifax
Thomas, 1915 (Chiroptera: Vesper-
tilhonidae) in eastern Australia..... e
Rercichthyidde mere tr era ene
Pexchateedinguhabitstese nn aera
Phylogenetic significance of Dracaena-type
growth, by J. I. Waterhouse.........
lies AINEOIMCEOs 65 ¢0¢0r0000000000%
Pleistocene, dinoflagellate cysts.........
Post-fire demography in the resprouting
shrub Angophora hispida (Sm.) Blaxell:
flowering, seed production, dispersal,
seedling establishment and survival, by
TD) Auldtin oan cvanae tes ae eee enn
Presidential Address 1979, see Waterhouse,
a
Presidential Address 1982, see Martin,
H.A.

175

195
B28)

91
155
154

293

219
331

153
143
143

129

331
175

259

By copodicaennnn seria ers ole ioe ce ts

Quinn, C. J., John Teast Waterhouse 1924-
1983. Memorial Series no. 27........

Records of Eudendrium (Hydrozoa:
Hydroida) from New Zealand, by J. E.
WVALSO Merete aeastines iols ote hsmcean ly Se

Review of the biology of Acacia suaveolens
(Smith) Willd. (Mimosaceae), by D. A.
IVMOGTISO TM ap ctoecepete Sein erkve tel

Rowe, F. W. E., & Albertson, E. L., A new
species in the echinasterid genus
Echinaster Miller and Troschel, 1840
(Echinodermata: Asteroidea) from
southeastern Australia and Norfolk

Rowe, F. W. E., & Albertson, E. L., The
echinoderm genus Henricia Gray, 1840
(Asteroidea: Echinasteridae) in
southern and southeastern Australian
waters, with the description of a new
SC CLES RSI Neicahmiyar as chue senichninan stat

Safia, Papua New Guinea..............
Seed production, etc, Angophora hispida... .
Seeds and fruits, Macquarie Island......
Shea, G. M., Two new species of Delma
(Lacertilia: Pygopodidae) from north-
eastern Queensland and a note on the
status of the genus Aclys.............
Silver perch, feeding habits.............

‘Wermmloya Rungere, WAVGtO NaI. 5 6 da sb bein Goode c

203

139

325

271

195

183
Sil

259
69

203
143

293

dleraponiclacaemeeet ari an eraclerenstee:
Tetraclitella divisa, first Australian
OCCURKENCEN siete nner eee
Timms, B. V., Geomorphic and physico-
chemical features of floodplain water-
bodies of the lower Hunter Valley,

Tuggerah Lakes, aquatic vegetation......
Two new species of Delma (Lacertilia:
Pygopodidae) from northeastern
Queensland and a note on the status of
the genus Aclys, by G. M. Shea.......

Wespertilionidachee tines
Wilctonlayelam borer ments ane

Water chemistry of floodplain lakes......
Waterhouse, J. T., Presidential Address
1979. The phylogenetic significance of
Dracaena-ty pe erow the =.- ce soe
Waterhouse, John Teast. Memorial Series
iO), 227/,; I Cr, Ifo @Qlewnain, ocevesocvss-
Watson, J. E., Records of Eudendrium
(Hydrozoa: Hydroida) from New
Leealandinrs arige city stec ese tayenaene en omits
Wright, R. V. S., Sir William Macleay
Memorial Lecture 1984. New light on
the extinction of the Australian
Ne vakauniays A eoiras aces enone:

Xanthonmhocaccaceew eee ene

ZO SLELAICADTICOTN Ee ee Ene

143

107

317

129

139

325

pier ieaae

Je aes:
a AP Garena
es toh

259

271

293

311

325

331

339

T. D. AULD
Post-fire Demography in the resprouting Shrub Angophora hispida (Sm.) Blaxell:
Flowering, seed Production, Dispersal, seedling Establishment and Survival

D. A. MORRISON
A Review of the Biology of Acacia suaveolens (Smith) Willd. (Mimosaceae)

C. L. FERGUSSON
Multiple Folding of the Ordovician Sequence, Tambo River, eastern Victoria

B. V. TIMMS
Geomorphic and cae cect nical Features of floodplain Waterbodies of the lower
Hunter Valley, N.S.W.

J. E. WATSON
Records of Eudendrium (Hydrozoa: Hydroida) from New Zealand

W. IVANTSOFF, L. E. L. M. CROWLEY and G. R. ALLEN
Description of a new Species of freshwater Hardyhead Craterocephalus kailolae
(Pisces: Atherinidae) from Safia, Papua New Guinea

INDEX to Volume 109

PROCEEDINGS of LINNEAN SOCIETY OF NEW SOUTH WALES
VOLUME 109 |

Issued 22nd December, 1987 |
i a ne
CONTENTS:

NUMBER 3

129. J. T. WATERHOUSE
Presidential Address 1979. The Oe aenctic Significance of Draceana-type Growth

139 C. J. QUINN
-_ Memorial Series no. 27. John Teast Waterhouse 1924- 1983

143. C. G. BARLOW, R. MCLOUGHLIN and K. BOCK
Complementary feeding Habits of Golden Perch Macquaria ambigua (Richardson)
peareieiionidee) and Silver Perch ee bidyanus (Mitchell) (Teraponidae) in-
arm Dams

153 H. E. PARNABY
_ Distribution and Taxonomy of the long-eared Bats Nyctophilus. spud Tomes, 1858
_-and Nyctophilus bifax Thomas, 1915 (Chiroptera: Vespertilionidae) in eastern
Australia

175 A. MCMINN
Late Pleistocene Dinoflagellate Cysts from Bulahdelah, northern New South Wales

183 F. W. E. ROWE and E. L. ALBERTSON
The echinoderm Genus Henricia Gray, 1840 (Asteroidea: Echinasteridae) in
southern and southeastern Australian Waters, with the Description of a new Species

195 F. W. E. ROWE and E. L. ALBERTSON
A new Species in the echinasterid Genus Echinaster Muller and Troschel, 1840
(Echinodermata: Asteroidea) from southeastern Australia and Norfolk Island

203 G. M. SHEA
Two new Species of Delma (Lacertilia: Pygopodidae) from northeastern Queensland
and a Note on the Status of the Genus Aclys

NUMBER 4

213 H. A. MARTIN
Presidential Address 1982. Cainozoic History of the Vegetation and Climate of the
Lachlan River Region, New South Wales

Contents continued inside
a a a ne a aS Te TE

Printed by Southwood Press Pty Limited,
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PROCEEDINGS

OF THE

LINNEAN SOCIETY

NEW SOUTH WALES

VOLUME
110
(Nos 481-484; for 1987-88)

Sydney
The Linnean Society of New South Wales
1989

Contents of Proceedings
Volume 110

NUMBER 1 (No. 481)
(Issued 12th July, 1988)

STEFFE, A. S., and PEASE, B. C. Diurnal survey of ichthyoplankton abundance,
distribution and seasonality in Botany Bay, New South Wales...........
CAMPBELL, K. G., and MOORE, K. M. Two new species of Glycaspis (Homo-
ptera: Spondyliaspididae) from potentially endangered Eucalyptus species
EMNGIEOMCMKORMY Loon SUI CUT aoa tents sence Mime ponents ey Gate NERO eu ell ae
Moore, K. M. Associations of some Glycaspis species (Homoptera: Spondyli-
aspididae) swith) thei, Zucalypius species NOStsi ass ee
MOORE, K. M. A new species of Glycaspis (Homoptera: Spondyliaspididae) and
SOIMMEeRME WAN OSEIRECONGS | carnewe yt tat ohare eaten A pn ararn Ed Ma Nenu
WRIGHT, A. J. First report of Late Devonian trilobites from eastern Australia .
MCALPINE, D. K. Studies in upside-down flies (Diptera: Neurochaetidae). Part
lee Systematics and phylocenyeyae ine cir eee: ee ce: ee Sue
MCALPINE, D. K. Studies in upside-down flies (Diptera: Neurochaetidae) Part
II. Biology, adaptations, and specific mating mechanisms .............
ROWE, F. W. E., and ALBERTSON, E. L. A new genus and four new species in the
family Echinasteridae (Echinodermata: Asteroidea)..................

NUMBER 2 (No. 482)
(Issued 12th July, 1988)

CARR, S.G. M., and CARR, D. J. Sir William Macleay Memorial Lecture 1987.
The elastic-sided gumleaf, or: The rubber cuticle and other studies of the
(CC) MO OSACM esas. Soe oie meted Ae Wee eR tin es ae Penn ee taster a sheer an:

GLASBY, P., SELKIRK, P. M., ADAMSON, D., DOWNING, A. J., and SELKIRK,
D. R. Blue Mountains Ash (Eucalyptus oreades R. T. Baker) in the western
Bey Milo uta ita Sot thai tid ra nee a te ag ee ee AEE te, Keg nN iia eat

YASSINI, I., and WRIGHT, A. J. Distribution and ecology of Recent ostracodes
(Grustacea)iromsPort Elackinie,. New Souths Walesiy.s-5 7 e see oe

HOTCHKISS, A. T., and IMAHORI, K., A new species of Nitella (Characeae)
belonging to the pluricellulate species group in Australia ..............

HOTCHKISS, A. T., and IMAHORI, K. Additional observations on Nitella verti-
cillata (Characeae) from a new locality in New South Wales ............

SOUTHCOTT, R. V. A new Australian larval callidosomatine mite (Acarina:
Erythraeidae) parasitic on flies, with notes on subfamily and tribe classifi-
CAT OTN ys eM I Vaile Sad ya PO Ni. hada ck cast MAR Nn RG UE Near Se eNO

ARCHER, A. W. The lichen genus Cladonia section Cocciferae in Australia ......

BREWER, D. T., and WARBURTON, K., A dietary study of Sillago analis and its
VAIallOneInEthire ew AUStralane| Ocalilties mam spear ere ete) eee ee

Annexure to Numbers I and 2. The Linnean Society of New South Wales.

Record of the Annual General Meeting 1986. Reports and balance sheets

101

141

NUMBER 3 (No. 483)
(Issued 13th January, 1989)

JOHNSTONE, R., Koop, K., and LARKUM, A. W. D. Fluxes of inorganic nitro-
gen between sediments and water in a coral reeflagoon...............
YASSINI, I., and JONES, B. G. Estuarine foraminiferal communities in Lake
MlawarraINe SW cece eae al ea ee lech ah aie toh gr
VAIL, L. L., and ROWE, F. W. E. Status of the genera Ophiopeza and Ophio-
psammus (Echinodermata: Ophiuroidea) in Australian waters, with the
GEScrIPlOMOltasmewiSPE GIES ee peels cit gaia Orca Ciera aoe
KENNEDY, C. M. A. Pycnodithella harveyi, a new Australian species of the Tri-
denchthoniidae (Pseudoscorpionida: Arachnida) ....................

NUMBER 4 (No. 484)
(Issued 13th January, 1989)

TIMMS, B. V. A study of the crustacean zooplankton of six floodplain water-
bodies of thellower hunter Vallley,.NESWae. 2 2 a) ne ee eee
SYEDA SALEHA TAHIR and CAROLIN, R. Seed type and seed surface patterns in
Galandrinia'sens Mats (Rortulacaceae)isy 2s 5a eee ee
KODELA, P. G., and DODSON, J. R. A late Holocene vegetation and fire record
from Ku-ring-gai Chase National Park, New South Wales .............
ZEIDLER, W. A new species of Melita (Crustacea: Amphipoda: Melitidae) from
northern New South Wales with a note on the genus Abludomelita Karaman,
IO STR sae An. te Rts Mire Me BN Tet aCe ee ae A
HUTCHINGS, P. A., VAN DER VELDE, J. T., and KEABLE, S. J. Baseline survey of
the benthic macrofauna of Twofold Bay, N.S.W., with a discussion of the
INakine:species imtvoduceduntothe!lbayes-4 eee ee oer a eee
FARRANT, P. A., and KING, R. J. The Dictyotales (Algae: Phaeophyta) of New
NOuth Wales: Ai ee Seen ey A FN? APM Bean Vet POTTS ta
SMITHERS, C. N. A distinctive new species and new distribution records of
Stilbopteryginae (Insecta: Neuroptera: Myrmeleontidae) .............
INDEX tosProceedings VOLO: gyi0 0 eal e elsniotngngm ecole a on Wee Ce eee

ZS)

229

267

289

297

307

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