VOL. 118, PARTS 1 & 2
31 MAY, 1994.
Contents
Transactions of the
Royal Society of South
Australia
Incorporated
Williams, W. D. Climate change and its implications for South Australia — introductory
remarks - - - - - - - - - - -
Schwerdtfeger, P. A brief overview of climate research - - - - - -
Allan, R. J. Modelling climatic change and variability - = - - - -
Williams, M. A. J. Some implications of past climatic changes in Australia = - -
Fitzpatrick, R. W. & Wright, M. J. Climate change and its implications for South
Australian soils - > - - - - - =:
Bates, B. W., Charles, S. P., Sumner, N. R. & Fleming, P. M. Climate change and its
hydrological implications for South Australia - - - -
Harvey, N. & Belperio, A. P. Implications of climate change for the South Australian
coastline - - - - - - ~ - - -
Tyler, M. J. Climatic change and its implications for the amphibian fauna - -
Stott, P. Climate change and its implications for the terrestrial vertebrate fauna F
Boardman, R. Some possible effects of climate change on vegetation - - -
Burns, M. E. & Walsh, C. The economic implications of climate change - -
McMichael, A. J. & Beers, M. Y. Climate change and human Population health: tne
and South Australian perspectives - -
Thumlert, T. A. & Austin, A. D. Biology of Phylacteophaga Jroggatti Riek (Hymenoptera:
Pergidae) and its parasitoids in South Australia -
Bayliss, D. E. Description of three new barnacles of the genus Elminius (Cirripedia:
Thoracica) from South Australia, with a key to species of the Elminiinae
Lange, R. T., Lay, B. G. & Tynan, R. W. Evaluation of extensive arid Faneclance: the
land condition index (LCI) - - - - - -
Prideaux, G. J. A small sthenurine kangaroo from a Pleistocene cave deposit, Nullarbor
Plain, Western Australia = - - - - - - - -
Zbik, M. The Cook 007 meteorite: a new H4 chondrite from South Australia - -
Brief Communication:
Read, J. L. A major range extension and new ecological data on Oxyuranus microlepidotus
(Reptilia: Elapidae) - - - - - - - - -
Walker, S. J. Growth in the Australian ene. frog, Bink australis (Gray) Anns.
Leptodactylidae) - -
PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS
SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000
owe
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TRANSACTIONS OF THE
ROYAL SOCIETY
OF SOUTH AUSTRALIA
INCORPORATED
VOL. H8, PART 1
Proceedings of a symposium entitled
“CLIMATE CHANGE AND ITS IMPLICATIONS FOR
SOUTH AUSTRALIA”
held on Li November, (993.
TRANSACTIONS OF THE
ROYAL SOCIETY OF SOUTH AUSTRALIA INC.
CONTENTS, VOL. 118, 1994
PARTS | & 2, 31 MAY, 1994
Williams, W. D. Climate change and its implications for South Australia — introductory remarks
Schwerdtfeger, P. A brief overview of climate research - - - - - -
Allan, R. J. Modelling climatic change and variability - - - - - -
Williams, M. A. J. Some implications of past climatic changes in Australia - - -
Fitzpatrick, R. W. & Wright, M. J. Climate wine and its herpligations for South Australian
soils - - - - - - - -
Bates, B. W., Charles, S. P., Sduiner, N.R. & Fleming, P.M. Climate cheng 2 and 1s
hydrological implications for South Australia - -
Harvey, N. & Belperio, A. P. Implications of climate change for the South Australian coastline
Tyler, M. J. Climatic change and its implications for the amphibian fauna — - - -
Stott, P. Climate change and its implications for the terrestrial vertebrate fauna —- -
Boardman, R. Some possible effects of climate change on vegetation - - - -
Burns, M. E. & Walsh, C. The economic implications of climate change = - - -
McMichael, A. J. & Beers, M. ¥. Climate change and human popalaeige health: global and
South Australian perspectives -
Thumlert, T. A. & Austin, A. D. Biology of Phylacteophaga froggatti Riek (ymenopter:
Pergidae) and its parasitoids in South Australia -
Bayliss, D. E. Description of three new barnacles of the genus Elminius (Cirripedia: Thorucica)
from South Australia, with a key to species of the Elminiinae - -
Lange, R. T., Lay, B. G. & Tynan, R. W. Evaluation of extensive arid rangelands: the land
condition index (LCI) - - - - - - - - -
Prideaux, G. J. A small sthenurine kangaroo from a Pleistocene cave deposit, Nullarbor Plain,
Western Australia - - : - - - - - -
Zbik, M. The Cook 007 meteorite: a new H4 chondrite from South Australia - -
Brief Communications:
Read, J. L. A major range extension and new ecological data on OxyaranDs microlepidotus
(Reptilia: Elapidae) - - - e zg . = -
Walker, S. J. Growth in the Australian burrowing frog, Cyclorana australis (Gray) (Anura:
Leptodactylidae) - - - - 7 : . - - =
45
143
147
PARTS 3 & 4, 30 NOVEMBER, 1994
Davies, M. & Watson, G. F. Morphology and reproductive biology of Limnodynastes salmini,
L. convexiusculus and Megistolotis lignarius (Anura: Leptodactylidae:
Limnodynastinae) - - - - - - - - -
Rounsevell, D, E., Ziegeler, D., Brown, P. B., Davies, M. & Littlejohn, M. J. A new genus
and species of frog (Anura: Leptodactylidae: Myobatrachinae) from
southern Tasmania - - 7 - - - - -
Pamment, D., Beveridge, I. & Gasser, R. B. The distribution of nematode parasites within
the stomach of the western grey kangaroo, Macropus fuliginosus -
Ainslie, R. C., Johnston, D. A. & Offler, E. W. Growth of the seagrass Posidonia sinuosa
Cambridge et Kuo at locations near to, and remote from, a power station
thermal outfall in northern Spencer Gulf, South Australia - - -
Molsher, R. L., Geddes, M. C. & Paton, D. C. Population and reproductive ecology of
the small-mouthed hardyhead Atherinosoma microstoma (Giinther)
(Pisces: Atherinidae) along a salinity gradient in the Coorong,
South Australia - ~ - - - = + - - -
Hutchinson, M. N., Milne, T. & Croft, T. Redescription and ecological notes on the pygmy
blubtonstie, Tiliqua adelaidensis (Squamata: Scincidae) —-
Fuller, M. K. & Jenkins, R. J. F. Moorowipora chamberensis, a coral from the early Cambrian
Moorowie Formation, Flinders Ranges, South Australia - - -
Peterson, M., Shea, G. M., Johnston, G. R. & Miller, B. Notes on the morphology and
biology of Ctenophorus mckenziei (Storr, 1981) (Squamata:
Agamidae) - - = ; + % 4 - - 7
McKillup, S. C. & McKillup, R. V. Reproduction and growth of the smooth pebble
crab Philyra laevis (Bell 1855) at two sites in South Australia during
1990-9] - - : E = = =: : - -
Olsen, A. M. The history of the development of the Pacific oyster, Crassostrea @ gigas CTanbee)
industry in South Australia = - “ = -
Bird, A. F. & Yeates, G. W. Studies on Aprutides guidettii (Nematoda: Seinuridae) isolated
from soil at Northfield, South Australia - - - - - -
Brief Communication:
Tyler, M. J., Davies, M., Watson, G. F. The hylid frog Litoria albapentare CGtimhet) in
the Northern Territory - - - 4
Insert to Transactions of the Roval Society of South Australia, Vol. H8, parts 3 & 4. 30 November, 1994
149
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CLIMATE CHANGE AND ITS IMPLICATIONS
FOR SOUTH AUSTRALIA —
INTRODUCTORY REMARKS
By W. D. WILLIAMS*
Summary
It is my privilege, as President of the Royal Society of South Australia, to introduce
this important symposium on climate change and its implications for South Australia.
In doing so, I wish first to welcome all participants and I extend a particular welcome
to Sir Mark Oliphant, an Honorary Fellow of our Society. Our patron, Her Excellency
the Governor, Dame Roma Mitchell, has indicated that Vice-Regal commitments
preclude her attendance. | trust that all participants, students, distinguished scientists,
or mere mortals like me, will find the symposium interesting, useful and a
constructive addition to local debate on this most important matter.
Teiyacnany of the Royal Sacre ofS. Mase (994) TBR Le.
CLIMATE CHANGE AND ITS IMPLICATIONS FOR SOUTH AUSTRALIA —
INTRODUCTORY REMARKS
by W. D. WILLIAMS?
Tis any privilewe, as President af the Royal Society
of South Australia, 09 tntroduee this injportani
symposium on climate chimge and its implications for
South Australia.
Ii deity se, L Wish first iu welcome all participants
und | extend a particular welcome to Sir Mark
Oliphant, an Heaueary Fellow of our Society. Cur
putron. Her Excellency the Governor, Dame Roma
Mitchell, has indicated that Viee-Regal conunitinents
preclude her attendance. I trust that all participants,
dndents, distinguished scientists, or mere mortals like
me. will Fiod che sympesiuny interesting, useful and
¥ Constructive additron te local debate on this most
TMIpOrin Matter.
The Royal Society of South Australia held nis first
meoling i November IR80, shortly afer Queen
Vietoriacassemted ta the use of the word Royal The
Society, however, succeeded the Adelaide
Philosophies! Society tortaed much sarher, and indeed
not lon afler the foundation of the Stace of South
Austauia [rt was formed in 1853. Since those early
yours, the Society has met regularly. published
scientific papers, maintained an extensave Library,
awarded grants fir research oruwards for meritorious
CONTDUTIORS to schence, and its Fellows have docluded
most ot South Australia’s best known scientists,
The overall objective of the Society is the promotion
and ciffunon of saentitic Knowledge, und it is in
support of this objective that the present symposium
hus been sponsored, [need hardly remind this audience
that any change to the climate of South Australia will
have profound, sygarficant, and comprehensive effects
an the South Austrahan environment, economy. social
structure and public health — to name but the most
ohvious features that could be affected, It isto orig
this to the altention of the community in general asa
its decision makers in particular that the present
syinpesiuin us berg held, Such an sum falls squarely
wilhin the overall objective of the Society. To premate
farther the views expressed. the pitpers presented ire
heing published in this part of the Trqnsacrions of the
Riyal Socrerv of South Australia and are alse available
sepairulely ws a bowk,
OD} course, the symposium is only one of many thul
have heen and are plunined to he held to discuss chymnite
chinge At ihe international level. numerous mectings
have been held, several scientific journals specitically
angel rescurch emt climate change, and there 1 an
* Department of Zoulogy, University of Adelaide, South
Austrailia 3005-
iitermational commuter which reports revulurly (POC:
Intergavernmental Panel on Climate Change).
Likewise, at the national level. considerable activity
prevails. Indeed. the Australian Academy of Science
is Currently sponsoring # meciing in Canberra oa the
subject of climate change, The National Greenhouse
Respoase Strategy was finalised in 1992. and we the
Stale level, Various povernment ugencies maintain ut
least a watching brief on the subject, a number of
oatnral and medical scicnlsts, economists and others
ure actively interested in the subject. there is a Climuce
Change Committee (serviced by the Department of
Environment and Natural Resources and including
representatives from # wide variety of government
departments) which reports regularly to Cabinet and
has published several imporiant documents (e.g. South
Australian Climate Change Conimittee 1990, 1991) and,
just a few years ago (1988), a Wide ranging conference
was held in Adelaide on the subject of climate change:
1 refer 10 Greenhouse “88: Planning for Climate
Change, Adelaide Conference (Dendy 1989), Finally,
T note the publication in November 1993 of South
Australian Greenhouse News, Vol. 4, No 1, the first
of what is intended 1 bea regular series of newsletters
with particular eniphasis on South Australia. [tis
published by the South Australian Departament of
Environment and Natural Resources with the support
of the Office of Enersy.
Aguiost this flucry of recent and ongomig activity one
might ask, why should the Royal Society sponsor yet
one move meeting We address the subject? What
possible good can anse from another “ralkfesr”?
The answer 4s simple, First, reseanch and views on
this subject proceeds apace and there isa constant need
Io provide an opportunity for such research ty he
reculasly aired for the benefit af the wider Commutnity,
His indeed halla decade since the last major meeting
took place in Adelaide to discuss this, mater.
Seonnd. the Reval Society is quite independent af
government depanments, research instirudons and
universities, and is therefore in a unique position —
indeed, has # speci responsibility ~ to provide an
opportunity for views to be aired which may not
necessarily conform to the party line, curfent weology
or generally accepted scientific views. As will become
obvious i some of the pupers to be given, there are
views held by some which da nor agree with
widespread views. [Sir Mark Oliphant, ina comment
atthe conclusion of the Symposiuin potuted to az mceat
arhicle in Nerare, hand, which pointed ty the luck at
any evidende af change in the presen climate. |
i]
It is important for me to add that the views of
speakers are not necessarily those of the Royal Society
of South Australia or the Institution to which the
speaker belongs.
And a third reason behind this symposium is the
constant need for the South Australian community, and
through it, government departments and politicians,
to be reminded of the need fo plan ahead tor possible
change, which may, in any event, be more rapid than
predicted. A recent article in the New Sefentist (Nielsen
1993) documents some early results of the analysis of
an ice core from Greenland which records detailed
changes in the earth’s climate over the past 250,000
years. According to scientists who investigated the core.
present models used in predicting climate change may
be too simplistic. In general, such models predict
gradual change. Analyses of the core, however, suggest
that a climate just a few degrees warmer than now may
change very suddenly to become either significantly
colder or warmer. In other words, we may be forcing
W. D. WILLIAMS
the present climate into an unsteady state when large
natural changes in climate could be triggered by
relatively small events. Whether such climatic
instability has already started is a moot point. Bear
in mind, however, that severe storms have cost
insurance companies over $60 billion over the past six
years (Leggett 1993). Meteorologists have already
begun to point io the increasing likelihood of an
increased frequency of natural disasters following
climate change in the next century (Zillman 1993),
1 hope I have indicated sufficiently the need for
and the importance of this symposium. All that remains
for me to do is to thank the speakers for presenting
their papers and the work involved in preparation. to
thank my various colleagues, in particular Dr Margarct
Davies, for her background support for the symposium,
and to wish you all a good afternoon, bon appetit for
the evening meal, and, since this is the last meeting
of the Society for 1993, a Merry Christmas and Happy
New Year.
References
Lraarrr, J. (1993) Who will underwrite the Hurricane? New
Scientist. 7 August 1993, 29-33,
ZILLMAN, J. (1993) Memorial lecture. University of
Melbourne, 7 September 1993. (Unpublished)-
Nievsen. R. H. (1993) Chill warnings from Greenland. New
Scientist. 28 August 1993, 29-33.
Denny, T. (Ed.) (1989) “Greenhouse 88: Planning for
Climatic Change Adelaide Conference Proceedings” (South
Australian Department of Environment & Planning,
Adelaide),
SouTH AUSTRALIAN CLIMATE CHANGE COMMITTEE (1990)
“Implications of Climate Change for South Australia’, First
report of the Climate Change Committee, August 1990,
(Department of Environment é& Planning. Adelaide).
_—_____ (1991) “The Greenhouse Strategy for South Australia’
(Department of Environment & Planning, Adelaide).
A BRIEF OVERVIEW OF CLIMATE RESEARCH
By PETER SCHWERDTFEGER*
Summary
Schwerdtfeger, P. (1994) A brief overview of climate research. Trans. R. Soc. S.
Aust. 118(1), 3-7, 31 May, 1994.
The problems facing those attempting to generate reliable prognostic climate models
is formidable. Good estimates of trends in concentration of all radiating atmospheric
gases are necessary and these must be entered into a tested numerical model
incorporating all of the important feedback processes. Included in these are the
parallel processes occurring in the oceans. The simplest test of the prognostic prowess
of a model is to run it backwards in time — a simple test of veracity that has not
supported any modern long-term prediction model.
Key Words: climate change, models, radiating atmospheric gases, meteorology.
Tremsaeions af te Ravel Neeters af 8 Aust 1994), TEL), 3-7,
A BRIEF OVERVIEW OF CLIMATE RESEARCH
by PETER SCHWERDTFEUER*
Suinmary
SCHWERDTFEGER, P1994) 4 brief overview of cline rescareh, Trans. R, Soe 8. ast USC), 3-731 May, 1994.
The problems taeing those attempting ts generate relinble prognostic climute models is formidable Good estimates
af pends in-coneeatration of all radiating atmospheric gases ure necessary and these must be eniered into a (ested
numerical model incorporating all af the important leedback processes, Included itr these are the parallel processes
besurring in the oceans The siiplest rest of the prognostic prowess of a model is to run it backwards in lime
usimple test of veracity that fag not Supported any modern long-term climate predichan model,
KEY WORDS: climate change, models, rugiitting wimoxpherie pases, meleornlogy,
Introduction
Over a third of a century ago, | found myself as a
freezing. fledgling geo-scientist, upprehensively
prodding the frozen crust of Arctie sea ice beluw, in
order to render advice on our mutual safety to my Red
Indian compunion, who gingerly guided the hungry
dogs that, from meal to meal, dragged the sled which
held our food and yay bare-bones scientific equipment.
We knew thal we were venturing close lo the brink
of chaos when the supporting ice became less than six
inches thick. My yelling out “150 mm” would only have
generated dangerous confusion. The metcorologival
Parameter of greatest concern ta me then was the
profile of temperature from the surface downwards.
und forthe following ten years, [remained interested
in ige-bound temperatures: in the Antarctic as well, and
pondered on how these arose, At the Lime, these were
generally regarded as actvities ambecoming of
tumospheric svientists, but with the growing relevance
attached to the new science of climate, the perceived
significance of the extent of polar ice, and its thermal
as well as physical conten) has been dtamatrcally
changed.
During the last 25 years, I started to raise my head
and examine the way in which the radiant energy
incident on the Earth's surface was transferred not onty
to depths below burt also to the atmosphere above. In
Ihe 1960s, many “real” meteorologists referred
indulgently to the “boundary layer boys” (a phrase that
the well known Princeton University theoretical
Meteorologist Joseph Smagorinsky used ina talk in
Melbourne. when he unveiled the then excitingly new
results of numencal modelling with enormous
computers, leading to the promise of operational long-
range weather forecasts). Smagorinsky was not
sneering, he well knew that the most definitive
iransformalion of energy from the Sun into other
* Institute for Atmospheric and Marine Science, Flinders
University of South Austnilia, GPO Box 2100, Adelaide
SA SOU!
radiative, conducted, convective, and evaporative forms
wikes place at the Rarth’s surface. Apait from the
conducted heat which involves the material below the
surface only, the three remaining fluxes drive that all
important distnbution zone, the diurnal boundary layer,
the thickness of which cun vary from millimetres to
kilometres, depending on the season and Lime of day.
Without the critical knowledge of the flows of energy
into this. boundary layer, reselved for a sufficiently
fincly Spaced grid covering the atea being wodelled,
every numeneal meteorologist would be marooned,
Theretine for some time, | continued to feel happy with
the relevunce of my work, delving im the lowest 10m
of almosphere blanketing the surface of the Earth.
Only ten years ago,as a result of my colleague Jorg
Hacker bringing his combined knowledge of flying and
meteorology to the Flinders. University atmospheric
research vroup. 1 became persuaded that there was
inore to meteorology than could be achieved by
climbing with instruments to the top of a Ladder. The
experience of flying offers meteorologists definitive
perspectives of the almosphere, especially when the
possibility of both making and viewing the results of
actual physical measurements in real time ullows
important physical connections to be grasped, Mare
than ever now, my heart bleeds for those meleonMogists
who spend all of their windowless lives hypnotically
hunched in front of their ceimputes screens.
The science of the weather was, in earlier years,
often conveniently divided into wetearmloxy, which
involves stitements about aunospheric conditions it
vatious given times, including the fulure, when
concepts of forecasting are invulved, and climmelogy,
which is concerned with quintessential summaries of
meteorological conditions over specified places and
periods, Applied sciences ure strongly driven by public
perceptions and wishes, in which mutters, of course,
scientists are increasingly not averse, tf not forced, ta
offer their guiding influence, The last degade has seen
the more geographical science of climearolngy fall into
relative disfayour as the persuasive value of the study
4 FP SCHWERDTIFEGER
At /anan. in being linked with the drama of inexorable
crends aad woystery al extremely loag-ieria lrecasts.
hus been heralded as an-essential fowus oh serious
allention,
Newspapers and television have popularised the
wuiject oP melenrolosy (hrough the regular presenta
av weather forecasts, and Satellite inmages Of tie Barth
hive helped grewly to generuiing an appreciation pl
the global niture ot meteamlogtieal processes reflected
in Visite cloud structures, There is no daubr (hat an
advance koowledge of denils of metearologival
conditions iy rnporimt in many areas ef human
cndewout, Over telatively short terms and for specific
ures suchas landing grounds, aviation safety depends
ui, borceastng. also influenees many other
Commercuil operations, tneluding those connected with
vaviculnure, the manofaentre oF ieeereun ind Urinks,
the saleetion oF routes tor shipping as well as personal
clnices regarding clothing and recreational wenivities,
As iitesalt, the concepts of mereralogy and forecaying
fhe become alist synonymous in the public mind
However, more general climatological summatics
also belp to guide tie public. particularly travellers,
ever (hough a Le caution im required when too bitte
detail is offered. Forexample, the bald stitement thar
Adelkude’s mean January temperature i 23°C vould
easily. prove deceptive toa nuive visiter, when itis
known Ut (he inedn maximum fempenidture is 30°C
und rhe mean minimum 169CL with extreme values ot
these (wo parameters of almost 48°C and 79C hay ing
feed recorded, Equipped with this additional
Information, the intending traveller miht obtaji seine
dew as to what types of clothing te puck An even mure
useless piece ob “stand-alone climatological
(iteration, exgep! pechaps to wind makers needing
ju be usstired (hal (he teniperatures miintuined im leep
cellars aie benefienl to the maturation af their
prodives. ms that Adclades mean unmaal tempensnture
iy 179 Responsible aind useral chmitolegieal
comipilitiuns Usually ain te provide envelopes tor
monges ol vitlues within which porumelers of interest
might be expected ty lie, The choice of these
panimeters as well us the time scule wall depend an
the purpese ail the application. For example. tor
avrivolionil decisions, information about vasefell,
tenperulare, volar rudtton and winediness Gneludine
likely extreties) ts important. [nthe Hight of the present
Knowledge of the Earths regional clinuattes, ub is cisy
ib overlook the aehievernent of colunial planners such
a6 Colonel William Tight, the founderof Adelaide,
who with Sis Weeks of arrival. barely In advance af
the Hirst Puropedn setilers, was able to assure hunselt
as la the general hature of Adelaide's climate and te
realise (Nal ih was the only location near the oust of
what hid been desiznaled us South Austmilia where
a city based an 19th eenfiiny technol ooulh be
estublished.
The need for information on weather in general
terms Js readily accepted for seasonal forecasts for
agricultural and water resources planning, Inevitably,
because Of the: tnstorically observed hok between
protracted extremes in meteorological gonditions and
the economy, the possibility oF rebably uscertiininy
long-term trends. is an auractive propasivon, Therela
he the origins of (he clamour for, and the justification
of, modern cliniate research,
Meteorologioal processes vccur on many Uillergent
limie-seales, The Iwo most fimilar of these are ihe
varhitions in insolation whieh result in the annual cvele
al’ seasons and the daily msing and setting of the sun
Both ane associated With systematic insolation changes
which resull in every olher meieoroloyienl paramelyr
EXporiene ie coMecquent variations. Chanwes mpodher
lime-seales ire fat less predictable. At the breh
frequeney dnd, jurbulenve is best desembed by sing
stalishicul expressions, Low frequency meteurologieal
changes hive beea discovered in geological studies of
the history wl the Earth wath distinet climate events
being separated by frony thousands to millions of years
On this scale, goalogists und glactologists have lony
led meteorologists with the Intensity of their interest
Systematic periagie meteoralogical changes van he
sumimarised by the climatology of the perids under
consideration, Pooe to the development of modern
methods of reconling meteorological valucs, the
deseriptions of historians. including the authors.of the
Old Testment olter investigalurs of past clhimutey a
weulth of iMformaben, hocexwnple, during bis years
in [evpt, by warning of the seven years of famine te
follow years O) plenty, Joseph issued his Pharaoh with
the first recorded suvecesstul lone. range weather
forecast. Phe Lirle lee Age ot the Middle Ages is a
well recorded historical fact, casily verifiable through
s(udies ol the welvance and relreala Alpine glaciers.
Only in ihe 20Ur century, have mine-shulls. lust worked
some 600 vers avo. become visible aus with the
retreat ol these alaciers which begun oy the [9th
centuyy, Modem visitors to the “Belvedere Hotel"
whieh. a hundred years ooo Urinatieally ever looked
the Rhone Glacier in Switzerland. muy well ponder
on how Jong powerful binoculars will suthiee (a offer
a dimpse of the ghicier’s snout following its dramatte
recession, Tt wits this same “Middle Apes” ace ape
which contributed, substantially: to the elimination ot
the Viking colonies from Greenland at about the same
time that Christopher Columbus was busily diverting
Eivepean atention to warmer parts of the New World,
Bryson & Murrey (1977) offer accounts of climaly
changes und the cunsequenves thereel we various
vilisutions over several thutsund years ol documented
history.
There is #bsolutely me coubt that dre clamite oP (te
Fanh is amb has always been changing (Brooks 144)
Haawever whit his wlso been changing during (he lest
A BRIER (VERVIRW OF CLIMATE RESEARCH 4
Iwoilevades iy the pubhe perception of the nature of
climwte chanes and concern thal the contribu bo
tbese by human activines may greatly overshudow
nalural ramlom trends, Quick td respond to such
voncerns. many alinospheric screntists have xoghe te
qoanoly and offer physical explanations for both
primary wbservations dnd vssumed consequential
changes, Chanwes of climate parameters necureine
“quturally’. are un seemingly randori tine scales
because oF the dificrenc rates governing the physical
“teedback” systems which link most, if net sll.
processes in jhe atmosphere, Phe vast tinge of these
pracesves, uod theit intemetions, wher the perfect
regipe Tor chaos,
The subjection of past elimalie data to frequency
analysis in the hope of obtaining eycheally repetitive
iessaies has been in var. My Flinders Universily
colleagues, John Bye. Rolwnd Bynim-Scat and Adtian
Gordon have demonsiraled tis phenomenon of
randommcss in-an unusually ditecl way. They have
translated random levels ol met-insatation. by means
of a simple, physical computer model in whieh
atmospheric and oceanic momentum are plausibly
coupled. inte time series of air amd sea leniperature
which over a period of several simulated centuries
display an uticanny superticig) resemblunce to actual
observations.
Were the Kanth to be 4 simple, heat conducting
hollow shell, simalur perhaps to carly urpfietl
satellites. but not even containing asuffering dag, hen
meteorology would be simple: the surface tempeniture
would be the only reportable factor, and i could be
walculated front a KnoWledge of the sofar cerisreut
(which in fuct is pot quite constant) and the measurable
radiative properties al the satellite surface The resull
iscommonly feterred to as the planetary lemperaiure
and the (wo importint planetary radiative properties
qwhich this temperature is related are the albedo or
surface reflectivity toward the wavelengths of solar
radiation, Which wre predominantly sisihle and the
planet’, emissivity, a factor which determines the
efficiency of theexport of radiative energy, mainly in
the infra-red minge of wavelengths. out wo space
Otherwise, this loss of radianr energy is determined
only by the temperature, a feet discovered by: Newton
three centunes ago and quantified by Stefan und
Bollamann late last century, This relative
meteorological simplicay is further enhanced by rts
instaneinenty: a thin hallow planetary shell cannot store
uny heat enerey and the coupling of tempertare and
maididtional events ogcurs without any delay.
By virtue of its samosphere, not to mention its sulid,
heat sluring mass, out real Earth has made this simple
business far more complicated. For cxample. the
diurnal delay between maximuns insolation at che locstl
geoxraphical noon and the muximiun air lemiperature.
which may Occur 2,5 tours later is uligest entirely
ascribuble to the thermal auimittanee of Conductive-
cupueity of che ground. ‘The Earth's surface usell does
not have & unitorm albedo; this varies (nim 3-4 % over
large parts of the ucean tu 80-90% over freshly fallen
snow, The tevel af heat storage depends in the leat
espacity and thermal conductivity of the material
beneath the Earths surface, but these couductive
processes cannot compete wilh (he efficiency of both
horizontal ued vertical energy transler mechanisins
within the Earth's dynunsie aunmsphere. Viewed from
space, the clouds which result from the presence of
wiler vapour: are the most dominating and reflective
feature. Clouds are larzely responsible lor the Earls
ayerave albedo being about 34%- As in the case of the
hypothetical, hollow, attiosphereless planet discussed
carlier, he 66% of the incident solar enemy which
is accepted by the Earth can alse be usel as the basis
for the galeulanon of a steady state plimecary
cemperiture; in face this turns out to he jhout 229°C.
Although this figure has real meaning i Lerms of the
Earth's temperature when observed fram outer spite,
it js difficult, unless resting ona polar (ce floe, to
appreciate Uy signilicanee hack on Balth rselP where
the wreally integrated mean surfice ten iperalure js about
I6°C The corresponding figure in upper almospherie
levels hich still emit significant amounts of infra-reé
radiation 14 about -f0%C. ft is essentin) t realise whit
the planetary tempermure must be regarded as 4
radiatively weighted nian over the entire thivkness of
the ulmusphere.
Vhe aumosphere, becuusc it contains poly-atonnic
gases which partially absocb and re-emit their
characterising bands of infra-ted radiation which the
Earths surluce ranstorms from absorbed inculent solat
radiation, greatly modifies the simple, 2-dimensiunal
picture of planetary temperature, Sympson (1927, 1928,
1929) was one of the first tu quantity, ia tetms ot
spectrally selective absorption, the Greenhene Lfjecr
which is cssentially responsible lor the mean
temperature ur the surface of the Earth being
approaimately 379C warmer thao tie plinetary
temperature. The over-whelmingly important gas in
this radiative process is water vapour, H,O-
Understanding, how this process contributes tu the
distribution of tenspetuture within the aimosphere is
diffivult enuugh in itself because the concentration af
this vital vapour varies with beth location and ime
Is Cuncentration und temperature in tum decermine
the existence of clouds, Which increase the reflection
of in-cunnog solar radiation, thereby generating the
fundamentally important feedback process of the
Earth's. surface becom shaded below, thereby
reducing its emiperiture and consequently also the level
of evaporation. All of hese linked steps occur neither
instantaneously nor ar ralés which can be uniquely’
specified The physicul clarity of the entire progess
which as most really comprehended in a single,
fi P SCHWERDTFEGER
vertical, dimension, becomes obscured by large seule
horwontal wvective processes, of which (he winds near
the pround and jnovenent of clouds above are the most
obvious manifestations, The variability of HQ i our
altiosphere results from the fact that it readily changes
trom yapour to bquid and even sobd phases within the
range of temperatures encountered cin Earth, all forms
havent strikingly ditferent mobilines.
There are ober gases Which comribuie to the
ainaspherie Greenhouse Effert, which t should be
emphasised, % significantly diferent from the marke
warden (ype. the litter having enclosing impermeable
inembpanes (6 inhibit both convective jind wyter vapour
lusses fram the systens, The aduitioniel gases are alse
paly-atunite Whar is they have three or more atoms per
NOleeole and are present in concentrations which ate
Small compared ta (hat at water vapour, In the absence
of stranaly locilised sources of productionind of sinks,
veiich absonh them. these gases are considered to be
umtermty axed with the other dontimating gayes of
the almosphere. such us oxygen and nitrogen. All of
the early mihalion toodels ol the atmosphere in
Iovussing an the importance of water vapour. deigned
io consider anly carbon dioxide, CO, as being of even
secondary uportanee in their schemes. Other
members af the ulmospheric Emily ol poly -atomic
gases. (luding Methane. oxides of nimogen and ezone
were reparded as nepligihle in influence. The
importunce accorded these eusey churned dramatically
following the first lew years of ahservations, when
these hevume available during che 70s. drum se global
nenwork Ab menor sratiens which detected steady
mereases Th the Mean unnual coneentrarions ot
particularly CO, and CH, in the atmusphere. (ce.
M.LT. 197)) Those are readily explicable in terms oF
the ever inereasimg vombustion of firssil fucts. the
Hling and bumog of the world's remorselessly felled
remnant forests, and the Tatulence of the world’s
burgeoning hents oF tame ruminants. [ots reasonabic
to dewept that with significantly greater concentration
levels. these gases should be ieluded i the weqountings
of Gallelive energy iransfer aid heating. Phe success
wilh which this can be done alse depends on the ability
lo idemly the mature and magnitude of all ot the
lvedbavk processes
With the acknowledged dmercuse mM the “lesser
fHidiatively Thportint gases, urises the ynestion as ta
the Wieans Whereby Une atiiosphere is also able to
continuously shed part af (he crneentration, That this
ich always haye heen so ws clear since CO, wid CH,
huve both been released ti the wir ik leis) since lite
began on Earth. even in tines when their mean
concentrations Were relatively stuble. (da the basis ob
current estimates of the alobul rates of CO,
Tesssta; Moo1 1 f1962)) PHD, Thesis, Metecrolugy
Duparhovet University of Melbourne Unpubl
production and krewledge of tsuctual concentration
in the atmosphere, itis a simple matter to eateylate
the rite al which one af the Earths leedhack systems
is dewling with the “fising” of ever increasing lowds
of this gus by, for example producing carbonates as
uresull Of the dissolving of the CO, tp water, This
has lune been w fertile Geld of tnvestization lor
teulovists. However. it is now apparent thar this
process is nat coping with all af the CQ) currently
being produced. so thut is atmosphere concentration
is CONLINUINE to increase,
in suinmary, the problems facing those wiiiig Wo
generate a reliable prognostic chimate model are quite
lormulable. His necessary to have geod estimates uf
ihe trends jn coneentration of all mUditing atmospherte
guses and (ensure thal (hese ane entered! oto a tested
numerical model whieh incurporales all of the
importum feedback processes made even more comples
hy the upprectation in recent! yours that models of glohil
atmospheric circulation cannot function without
consideration of the parallel processes necurring pn the
ocedns, The simplest test ol the prognostic prowess
ot ainadel is to run it buckwards in time. When Uwe
Radok and Dick Jenssen deyeluped Australia’s lira
numerical short-term weather forecasting model il
Melbourne University in the late 19505. (Jenssen
192"), they used thiy simple and Honest procedure.
‘Yo the best of my knowledge, no modern long-term
climate prediction model his yet withstood this simple
(vst of veraeay:
In spite of the fact that one stmke of a US
presidential pencan huve Qrealer vunsequenees for the
Australian agricultural economy. thin even some of the
mare adverse predichons ollered by the proponents tt
climure models currently im vogue, | believe thal
climate resewreh is importunt. | alse believe that some
of the recommenditions af those whose faith iq curretit
Inodel predictions is clase to absolute, in which 4
radical reduction in the global sesle of combustion ts
urged, deserve substintial endorsement, particularly
where this may lead toa deceleration of the senseless
destruction of the world’s forests. Dihers may applaud
the development of niore vegetarian habus hy the
inereasinaly carnivorous human pace, whether or nor
aw reduction in vlobal catile numbers has any ulimate
significant impaeton climarcce wends. J do, however.
lind it regrettable, that the stranger proponents ol
climate research have dispropertionately triggered i
wide Tuoge of polineul imochurisinys by means of
potentially, alirming information. without proper
substantiation, apparently to generate and maintain high
levels of public attention. So fir the nwin result appears
Ww bave been to generale public confusion bemween thie
concepts underlying the Greenhouse kffect and Une
Ozone Mole T well recall two remarkably different
lectures held withia the space of a sitgle decade
id the seey same Universite of Adelaide. by ai
A BRIEF OVERVIEW OF CLIMATE RESEARCH 7
internationally acclaimed visiting atmospheric scientist.
The first of these chillingly fortold of an impending
Ice Age, the second unabashedly warned of Greenhouse
warming, Of course all scientists need public attention
in order to survive, but for credibility to be maintained.
science normally needs to disclose verifiable results.
even if these may require interpretation in a manner
conducive to greater public appreciation. It is
dangerous for any branch of scientists to test popular
indulgence too far and in the case of atmospheric
scientists, to allow the public to lose sight of the many
other useful purposes their scientific endeavours both
continue to and could be offering, particularly in non-
long-term prognostic ventures. With public and
political attention rivetted on such matters as burping
bulls in Brazil and the saturation of the ever expanding
seven seas with soda water, it is easy to overlook the
multiplicity of climate-related environmental problems
closer to home, where appreciation of diverse themes,
ranging from the security of vital water supply
catchments and the impact of large-scale land
management policies on regional climates (and vice
versa), to detailed investigations of the actual physical
and chemical processes constituting the links of the
complex chain of events loosely described as climate.
are vulnerable to chaotic diversionary forces which
thrive in confused societies.
References
Brooks, C. E. P. (1949) “Climate through the ages: a study
of climatic factors and their variations” (Dover facsimile
edition, New York 1970).
Bryson, R. A. & Murray, T. J. (1977) “Climates of hunger:
Mankind and the World’s changing weather” (University
of Wisconsin Press, Madison).
M.LT. (1971) “Inadvertent climate modification (Report of the
study of man’s impact on climate)”. (Massachusetts Inst.
of Technology Press, Cambridge).
Simpson, G, C, (1927) Mem. Rov. Meteorol, Soc. 2, 16.
(1928) Ibid. 3, 21.
_____ (1929) Ibid, 3, 23.
MODELLING CLIMATIC CHANGE AND VARIABILITY
By ROBERT J. ALLAN*
Summary
Allan, R. J. (1994) Modelling climate change and variability. Trans. R. Soc. S. Aust.
118(1), 9-15, 31 May, 1994.
Recent interest in climatic change has engendered considerable debate about the
enhanced greenhouse effect and its possible global impacts. Numerical computer
models of the climate system are important tools in the scientific assessment of the
enhanced greenhouse effect. This paper briefly reviews the model development and
approaches used to simulate the nature of anthropogenic changes and natural
variability in the climate system. Research in Australia is given particular emphasis.
Key Words: climate change, numerical computer models.
Trmmsaorions ih the Reval Sueiely ofS, dash (994), OBC), 91
MODELLING CLIMATIC CHANGE AND VARIABILITY
by RORERT J. ALLAN*
Abbas Rod (199d) Modelling climate change and variability, Vans, Ry See. So Aust, UBC. Y-15, AL May, 1994
Recent imerest 1 climatic change has engenderec. consideruble debate Whout whe enhanced greenhouse effect
anid ils possible global impacts. Numerical computer models of the chinate svsicu) are tnporuint fools in the
aulctititic wssessinent of the enhanced greenhouse efleet. ‘Thos paper briefly reviews the model development and
approaches used (o simulate the nature nf anthropogenic changes wad matupal variability an the chimate 4ystem.
Research w-Australia is given particulas eniphiois
KEY WORDS; Climate change, numeridal computer jeadels
Tntroduction
he wlobal climate system involves closely linked
interactions betyeen the atnosphere, oceans,
eryosphere and the biosphere, This system is doveu
hy energy derived trom sulur radiation, with the net
ilvoniing energy being balanced by that which ts fost
ta space, The nonlinear and highly dynamic climate
syste is at consequence of the redistribution of this
energy by thermodynamics and. the forces of motion
derived from the planet's rotation. Any Muctuahions or
changes in the climate system can be brought about
hy external foreing. or by provesses and readyustments
intrinsig to such a closely Coupled nuntinear syste.
Since the systein is finely balanced, what might appeir
whe minor changes. inany part of the system can-cause
large chunges in its character.
Until the latter part of dis century, utempts 10
understand the climate systent, and the variahons- and
changes in il, were puryued through observational
studies and the development of physical theories ol the
interactions and motions voverniny it, With the advent
of, und improvements in, computer technalogy, efforts
Jo produce computer models oF the climate syste have
expanded rapidly. Mueb initial modelling work focused
on the need to miprove day to day weather forecasts
usita models capable of resolving, synupuie scale
tealures in (he atmosphere. However, the longer term
envelope of weather events that constitutes the climate
has now become a significant areca in numerical
niodelinog. Concerns about the enhanced greenhiise
effect and a need to understand natural fuctuations in
the eliniuite system have seen u strong focus Ga The
development of models able to capture the progesses
governing interactions inherent ii the climate system.
Such interactions are shown schematically to Fig. 1.
This paper ptovides an overview of the important
characteristics of numerical models that have been used
to improve the scientific understanding of changes and
Nuctuations in the climate system, It foguses on both
Climate Impact Graup. CSIRO Division pf Atmospheric
Research, PMB No. |. Mociialloe 3195. Vietoria,
international efforts and the progress being made in
Australia 10 examine potential climate problems using
numerical computer models.
Low order models
Some models. have been developed lo improve
Understanding of particular features that dominate the
climate over regions of the globe und for various
periods oltime. Such models use what are often culled
“primitive” forms of the basic physical dynamics that
are necessary to encompass the processes underlying
climatic phenomena. These computer models contuin
sets af mathematical equations describing the most
important physical interactions at each point on
horizontal and vertical grids of pots that cover the
region in which the climatic feature oceurs. All the
equations governing, the physical properties of the
phenomenon are solved at time steps indicative of the
observed nature of the pracesses being examined, Ln
thts way, a model of the real system is built up.
However, such models are only as pood as the
mathematical resalution of the dynamics of the feature
being modelled, Some real processes are relauvely
casier to describe mathematically and ther mieractions
are known. Others are difficult to capture fully, or the
present physical understanding of then may be limited.
Tr general, Jow order models dave focused on resolving
the important atmospheric and/or ocewnie character of
a particular climatic phenomenon,
Perhups the mont concentrated emphases in this ares
has been in the construction of low order models aimed
at capturing the essence of EI Nifio-Southern
Oxcillation (ENSO) events, The ENSO phenomenon
is a large-scale ocean-atmosphere interaction. that
occurs uregularly. dnd is centred in the Indo-Pacific
basin. It involves a Cloye coupling of the important
features of the climate system across this regton aml
has significant impacts in the marine and terrestrial!
environments. Some of the maw variables and
feedbucks Known tu operate during ENSO events are
shown schematically in Fig. 2, and a review of the
phenomenon can be found in Allan (1991).
10 R, J. ALLAN
Cr
a
Changes of }=9=-—~ US
solar radiation us “f 4) -
H '\\
Terrestrial
radiation =
H2O, Ny, Os, COz, O4, etc, Clouds (
aerosol Ls sy Y
Atmosphere
Atmosphere-land coupling
Biomass
VW ’
Fey Sea-ice | wy *1 Wind stress
Atmosphere-ice coupling TA pT
: |
A
Ice-sheets, ,
snow Heat Precipitalion |
Fee = Evaporation
Nee ral MN) cl Otel fs exchange Pp
Ice—-ocean A h fi Ocean
Changes of coupling tmosphere-ocean coupling
atmospheric composition . LaF
Changes ol land fealures,
orography, vegetation,
albedo, etc.
Changes of ocean basin
shape, salinity, etc.
COUPLED OCEAN-ATMOSPHERE
ATMOSPHERIC CIRCULATION
aan
siomen Uy
(RAINFALL )
SURFACE WINDS
=— — > ee
CURRENTS => > => SEA-SURFACE
wr ates TEMPERATURE
NESS POSITIVE
7 pYER THICK FEEDBACK
NEGATIVE
FAST RESPONSE FEEDBACK
OCEAN CIRCULATION
SLOW RESPONSE
MODELLING CLIMATIC CHANGE ANI VARIABILITY Il
Most low order tnodels of ENSO have locused on
the Pacific Ocean region botnded by the suimrapics
in_bath Iyemispheres, Such models are very sunple,
with ninimal physical representation of the almesphere
(often only two layers in the vertical) and a constriction
designet! lo resolve (he basic dynamics of the ENSO
phenomenon alone. The building of models in this
context bas required substantial resources and
particular recourse to ahservational studies and
dynamical theories whieh provide a conceptual
lramework of the physical processes likely to underlie
the phenomenon. At present, there is au firm theory
describing all aspects of ENSO events, but rather a
dynamically consistent understunding of seme of the
IMporiant processes Governing its behaviour. Curren
models of ENSO have been built under this constraint.
More recent developments in low order coupled ocean-
almosphere models have scen improvements in the
ibility of these tools ro predict ENSO behaviour The
mist comprehensive review of the current slalus of
ENSO jodelling as given in Neelin er al, (1992). In
Australi, such work ts perhaps best charucterised hy
ihe law order model developments at the Bureau of
Metcorology Research Centre (BMRC) (Kleeman 1991,
1993), Further propress cun be expected as the
underlying dynamics powering ENSO episodes
become better understood. wd the vapabilidies of
models to represent such Complex processes develop
turther,
Gencral Circulation Modelling
More detailed numerical models. for climatic
research are the General Circulation Models (GCMs),
which are three dimensional models of the global
climate system. These models represent interacuions
between features of the climate system through a series
of mathematical equations on a spherical grid. The
grid, representing a latitude by longitude army, is
repeated tn the vertical plane aver the desired number
of levels or layers throuyh the depth of the simusphere.
This structure is shown in Pig 3, and provides: the
Irimework within which the model simulates the
dominant processes of the climate system over cime.
Physical processes and interactions are calculated at
euch grid point simnultancously in the active maul.
In addition, in some models important boeanic links
to the GCM atmosphere are mimicked by including
a“slabyocean” in Lie model. This iy a 50 m deep layer
covering the Eaah’s oceans that performs like a well-
muxed oceame “near surface” layer. However, sea
surface temperatures simulated by this approach most
be continuously corrected to take uccount of the hear
transport by hortvonlal currents and deep ocean
processes. This is termed w Q-flux correction to the
GCM, Using this technique and given the constraints
mentioned, un evolving simulation of the Earth's
climate is produced by the model that captures many
observed aspects of the climate system, Detailed
specifications of the CSIRO Divison of Atmospheric
Research (DAR) and the BMRC GCMs that haye beet
developed in Australia, are-given in MeGireger rf al
(1993b) and Hart er e/, (1990).
Despite the relatively sophisticated nature of GCMs,
they are coarse in their spatial resolgtion with
endpoints spaced al arqund 500 kin apart and vertical
levels representing the atmospheric structure usually
limited to between 10-20 layers. As u result of these
Spatial restrictions nposal by limilalions of computing
resources, many smaller features of the climate system
such as tropical cyclones and deep cyclonic systems
cunnot be resolved by GCMs, Another importunt
problem that stents from ihe spatial limitations is that
some physical processes, such as those relating la
uspects of cloud physics, cannot be fully resolved and
are parameterised in these models. These factors limit
the current capabilities of GCMs to capture the full
inincacies of global climate. At present, for example,
GCMs are unable w produce realistic ENSO events.
Nevertheless, it must be realised that these are early
generation models and that many of the large-scale
aspects of the real climate ere seen in the current GOM
simulations
An important advance in the development of GCMs
is the Coupling of atmospheric and oceanic models
(fortnulated similarly to the-GCMs) in order th) capture
more of the essential elements responsible for observed
climatic patterns. Currently, such coupled madels
require correction for fluxes of energy, miinentum and
moisture exchange berween the atmosphere and the
aceans if they are to resolve full ocean-almosphere
imeractions. Mast simulauons of enhanced preenhuuse.
effect conditions with current coupled ocedn-
almosphere GCMs have used {ux correction,
However, as ned in the section on low order models.
coupled models provide the basis for a more reulixuc
Fiz, | TOP. A simple schematic view of the major interactions in the global climate sysiem.
fig 2. BOTTOM, A schematic diagram of ENSO interactions involving major oceanic and atmospheric yanables in the
Pacifie Basin. Whe lwo responses (fiat and slow) in the ocean refer in differen feedbacks rosulling fran che natire ol
the dynamical waves generated by tropical oceunaumesphere teractions inherent in the phenomenen
12 R J. ALLAN
a. sea
on the surface
Fig. 3. A schematic representation of a GCM as i divides the earth inty a regular three-dimensional grid.
7.0
= 0.8
i
3
® 0.6
=
oO
bd BMRC
s a4 CSIRO
= CSIRO4
o
we GFDLH
ccc
UKMOH
0.0
DJF MAM JJA SON
Fig. 4. Pattern correlation for seasonal yalues of model
simulated and observed mean $ea level pressure over the
Australian region. The abbreviations are for the GCMs and
BMRC, CSIRO (4 and 9-levels in the vertical), United
States. Geophysical Fluid Dynamics Laboratory (GFDL)
(High resolution), Canadian Climate Centre (CCC) and
the United Kingdom Meteorological Office (High
resoluhon).
representation of the real world. Oceanic GCMs have
a similar structure to those described for the
atmosphere, but must represent a more “sluggish” tluid
with spatial and temporal scales of processes and
interactions. that ate slower than in the atmosphere. It
must also be remembered that models of the
atmosphere owe much to a Jong history of scientific
endeavour to improve weather forecasts. Oceanic
model technology has lagged behind and these models
require considerable further development. Much of this
is dué to sparse Observational data coverage for the
aceans, which limits the validation of model output,
and the need to resolve smaller scale processes
involving meso-scale eddies in order to achieve more
realistic simulations. The status of coupled model
development at the Australian Bureau of Meteorology
is given in Power ef al. (1993), Similar development
at CSIRO’s Division of Atmospheric Research is ina
rather more advanced state.
In using GCMs for studying variations or changes
in the global climate, one important prerequisite is that
the models can perform an acceptable simulation of
current climatic elements. A number of
intercomparisons between various GCM simulations
of climatic fields and observed climatic parameters
MODELLING CLIMATIC CHANGE ANT) VARIABILITY 3
have new been perlormed by reseurch groups around
the world (Houghton ef af 1990, 1992), Specific
intemumparisons for GCM simulations aver the
Australian region are detailed in Whetion & Pitoek
(1991, 1993) and Whetton er af. (1994) (Fig. 4). The
most encouraging aspect ot these checks. is (hil they
have objecuvely revealed 4a general improvement ub
GCM simulations as model resolution and physics have
heen refines!
The procedure employed to sei up a model run ta
produce “contra” Ar present climate conditions usually
volves an-experanent where the GCM as run forward
im line under ihe current wlmosphene CO,
concemiration with observed sea surface lemiperatures
und initial ptmospheri¢ input fields, The aim of this
lype of éxperinient is to obtain a sanyple of simulated
climaly lang enaugh for statistically valid analysis. In
practice. this normally requires between 10-30 years
of simulated chmate variables. The various elds
generated ean then be examined using cureful staustical
techniques thir check the pattern of cormlations
between exch of the GCM: control uutpur dite
distributions with the appropriate observed field
patterns. Sivulated fields can also be examined for grid
point and total spatial freld significance, If this proves
lo be satisfactory. then the GCM can be used to address
the possthle change ta climate that could result trom
the enhanced greenhouse effect
Two Lypes of experiments are commonty conducted
to-assess. possible climatic change duc to increasing
CU, The first, and most widely employed technique
mine in which the experiment is periomied by simply
running the GCM with doubled CO, These
equilittrium simulations require about a 10-year time
peciod for the GCM to adjust to the new CO, limit
alter itis instiniunedusly imposed, and then a sample
of the next 20-30 years of data is ken fo represent
the enhanced areenhouse climate stutistics. A doubling
oF COs Is a standard experiment used as a benchmark
ugains! which all (XCMs can be compared. It does nat
refer ly any particular dale in the future and implicitly
\heluides ‘the radiative effect of increases. in other
greenlmuse pases, which are expressed as
CO ,-equivalents, The more physically realistic
approach, which ts being undertaken as coupled necan-
atmosphere modets begome more widcly available and
computing capabrliaes improve. is tw perforant a
tunsient experiment. In this type of simulation the
CO, ithe model atmosphere is gradually increased
Tha manner which loosely appruxemates the proyected
increases in this gas, lo a time when the level has
doubled.
Intemarional evaluations of climiute change research
and GCM siniiitiens and intercomparisons arc
compiled in the reports of the hnfergovernmental Pancl
on Climate Change (POC) (Houghian er af 989(),
W921 Th Australi, GOM research with “on hose”
models 1s concemrated of BARC and CSIRO Division
of Atmospheric Research. Both groups have produced
doubled CO, cquillhniun culis using impreved versions
of their GC Ms, und are dewelopmg GCMs tor transient
experumenis (Power et a/. 1993) and te simulate ENSO
and patueral climate variability, At the CSIRO Division
of Atmospheric Research, resulis of enhanced
ercenluuse effect experiments have been used as part
of the input inte regional climate change scenarios for
Austealia. Tiaportanl contributions io IPCC evaluations
have been made by (SIRO'S Divisiun ul Almospheric
Researeh through studies examining the resoludon of
possible changes in rainfall amaunts ind intensities in
doubled CO, simulations (Gordon et a, 1992), This
anup alsa has tepaned un the nalure ul variahilily
within the climate system sunulatcd hy duuhled CO,
experiments (Gordon & Hunt 1994) Climate variahiliry
studies ai CSIRO include those using the recent
Atmospheric Model lotercompurisan Project (AMIP)
results Io highlight the ampact ol chaotic influences on
the atmosphere (Dix & Huint 1994) and an ongoing
experiment simulating 300 years of climate
Limited Area and nested modelling
The neal po focus on likely changes un regional
climates, together with the poor spatial resolution of
GCMs, trewns thal models that can resolve smaller scale
processes are required, Limited area models (LAMS)
have finer spatial resolutions than the GCMs and can
resolve Jeatures of orography and geography thal arc
smoothed .in coarser global models (Giorgi 1990; Giorgi
et al, 1990, 1992), Typical horizontal grid resolutians
that have been achieved by LAMS in Australia alt as
Jow as 60 km with IS levels in the atmosphere
(McGregor & Walsh 1993; MeGregor er al. 19934).
LAMs are constructed to cover limited regians af the
Earth's surface and to focus on specific feaures of the
chimale system while being computationally economical,
Such models can be used to examine how: a particular
synoptic system, such as a tropical cvelone. may hive
behaved 1 it were in have been wdlueneed hy different
environmental conditions (Evans 1993; Evans eral. in
press}. For enhanoal greenhouse experiments fis aisy
involve increasing, sea surlisce temperatures in mimic
doubled COj conditions snd then tesung ihe sensitivaly
of the tropical cyclone anil the amount of rain i) proaluces
ta uch a change in boundary conditions. These medels
are particularly useful in ussessmy the sensitivity of local
preciptation-producing systems ta @ changed
environment. Juint research by BMRC and DAR has
Concentrated on the simulation of “east-coast’ lows ov
what are often termed "cut-off" lows that can produce
copious rainfall aver southeastern Australia, This cu-
operaive study is examining the effect that increased
Sex Surface Pemperslujes May have on tle precipitalion
regime of these synoptic systerns (MeTrines et al, 1992)
it kod ALLAN
a —
7
ee TEPrr eben,
a ee
pee a)
=
Fig, 9 Schematic showing the nesting-ol the LAM gridowiltin
the CSIRO GCM grid over Australia,
An important new development using both GCMs
and LAMSs is the combination of the two tn what es
termed a “nested” model, The GCM generutes the
information trom a full globsl simulation and provides
the boundary conditions for the GAM that is embedded
or nested within the larger, coarser model. This ts
illustcated in Fig. 5. The nested modelling approach
is capable of resolying features as small as
60 * 60 km. Recent research at the CSIRO Division
of Aunospheric Research has focused on the use of
this approach ta produce finer scale regional resolution
of climatic features and their sensitivity te changed
eavivonmental conditions. ‘So far. LAMs bave anly
been ncstcd in equilibrium boundary conditions
produced by GCMs with a simple Q-flux corrected
ocean model, As such, these LAMs. ure limited in their
performance by the accuracy of the GCM boundary
cundilions,
Other types of models are currently being used or
developed internationally and in Australia. One
interesting, application is the use of tracer transport
tnodels to examine the atmospheric CO, budget,
Studies ut CSIRO DAR using such models are
described in Enting er al, (1993). Such work is
purticularly important in helping to estimate the global
carbon budyet.
Conclusions
Over the last five years, progress in developing
numerical computer models of the climate system has
heen substantial. mainly as a result of the stimulus
provided by concern about the enhanced greenhouse
effect and the need to investigate its possible globat
impacts, Today a suite of numerical models ts being
Weveloped i juckle various problems relating to the
climate system. Specialised low order models focusing
om particular features of the climate systent are best
exeniplitied hy models of the ENSO phenomenon.
Such research has already been responsible for
significant improvements io the torevasting uf ENSO
phases. Larger atmospheric and oceanic GCMs are
used to-simulate the global climate and hkely changes
and fluctuations in climatic pallerms due to the
enhunced greenhouse effecr and nariral variability,
Efforts are also progressing with the full coupling ol
the oceun-atinosphere system und the addition of
realisne und interactive cryospherie and biospheric
elements. Finer-seule resolution. in the numerical
modelling of climate has been attempted using LAMs
that have been nested in the coarser GCMs, or used
as “stind alone” models cupable of resolving specific
synoptic features and their sensitivity to: potentiul
changes in the climate system.
Considerable further research is needed to continue
such developments and to take the current generation
of models to a stage where fully interactive Earth
system representations are possible. Such
improvements then open up the possibility of
simulating the nature of fluctuations and changes in
the global climate system more realistically. However.
these developments must be linked with research to
improve observational data records and understanding
of chmate dynamics, so that model resulis can he
assessed in their proper context.
Acknowledgments
J would like to thank Dr-Chris Mitchelland Messrs
Barrie Hunt and Kevin Hennessy, CSIRO Divison of
Atmospheric Research and De Janette Lindesay.
Department of Geography, Australian National
University for discussions and their constructive
comments on the initial manuscript of this paper. This
work contributes to the CSIRO Climate Change
Research Program «nd is part funded through
Australia’s National Greenhouse Research Program.
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Graneék., H. (1993) Synthesis inversion of ulmospheric
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Melbourne +44
MODELLING CLIMATIC CHANGE AND VARIABILITY 15
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to sea surface temperature. J. Climate 6, 1133-1140,
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J Climate 6, 2012-2033,
McGreoor, J, L. & Wats, K. (1993) Nested simulations
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SOME IMPLICATIONS OF PAST CLIMATIC CHANGES
IN AUSTRALIA
By M. A. J. WILLIAMS*
Summary
Williams, M. A. J. (1994) Some implications of past climatic changes in Australia.
Trans. R. Soc. S. Aust. 118(1), 17-25, 31 May, 1994.
An important question, in the context of possible human impact on global climate, is
whether past climatic history can offer insights into possible future climatic change.
Equally critical is the likely response of the physical environment to any future
climatic changes. Evidence from pollen analysis, lake fluctuations, desert dunes and
coastal plains from Holocene deposits in Queensland, Victoria, New South Wales and
Northern Territory, demonstrates that the response of different elements of the
Australian landscape to geologically-recent changes in temperature and precipitation
was often time-transgressive. Any attempt to use palaeoenvironmental data to predict
possible future change must therefore take due account of the varying response times
of different constituents of the Australian landscape to external disturbance. A
synchronous response to climate change is more likely with relatively simple
biophysical systems such as small closed lake basins or source-bordering dunes than
with more complex systems such as tropical rainforests and tropical coastal plains.
Key Words. Climatic change, Australian landscape, past climates, Queensland
rainforest, Victorian lakes, New South Wales dunes, Northern Territory coastal
plains, response times.
Vrveciins af the Reval Soviery of S. aust (99d). Le 17-25
SOME IMPLICATIONS OF PAST CLIMATIC CHANGES IN AUSTRALIA
by M.A. J. Wittiams*
Sunumary
Wituiams. M, A. J, 994) Some implications of past ulinuitic changes in Australia, Trans, R, See, 5. dusr LBC),
17-25, 31 May; 1994
An important question, in the context of possthle human impact on global climate, is whether past climatic
husiory can offer insigbtyinto possible future clinic change. Equally evitical 1 the likely response of the physical
environment to any furure climatic changes, Evidence from pollen analysis, lake Noctuarons, deseer dunes and
coustal plains from Holocene deposits in Queensland, Victoria, New South Wales and Northern ‘Territory,
demonstrates that the response of different elements of the Australian landscape to pedlogicully-recent changes
in lempecatuce aad precipitation way often Ume-transgressive. Any attempt to use palaeoeayironmental data to
predict possible future change must therefore take due account of the varying response times of different constituents
of the Austruliinn landscape to external disturbance. A synchronous response to climatic change is, more likely
with relatively simple biophysicalsystems sach as small closed luke basis or source-bocdering dunes than with
more complex systems kuch vs tropical ramforests and tropical coastal plums,
KEY WORDS. Climatic chunve, Australiin landscape, past climates. Queensland rainforest. Victorian lakes,
New South Wales dunes, Northern Territory. coasta: plains, response times.
Introduction
There is in¢reasing concern thut the aceclerating
impact of human activities upon our natural resources
of land, air. water, plants and animals is seriously
daimaging many of our more Tragile egosystems,
culminating 10 irreversible losses-of genetic as well as
cultural diversity. Hunan actions may alsa be
contributing. to possible changes in world vlimute
through a combination of burning of fossil fuels und
changing land use (most notably deforestation). both
of which reduced the terrestrial store of carbon by dbint
60 + 0.5 and 1.6 / LO PeC in 1990, respectively
(Houghton e af. 1992).
Although the increase in the atmospheric
concentration of carbon dioxide. methane, nitrous
oxide und other “preenhouse” gases since the start of
the industrial revolution is well documented, the Hkely
impact Upon global climate and ecosystems remains
unclear (Pearman 1988; Houghton ef al. 1990;
Dunnette & O Brien 1992), Notwithstanding the
unavoidable scientific uncertainties over the magnitude
and frequency of future climatic Auctuations, there
appears to be w significam measure of agreement that
the increase in anthropogenic aerosols and trace gases
will enhance the greenhouse effect, culminating it
global warming of the lower atmosphere, particularly
in middle to high Jutitudes (Houghton er al, 1990;
1992). However, there is far less agreement on how
the global patiern of precipitation might respand to
enhanged greenhouse warming, prompting a aumber
of researcher's to look tw the past as a sourec of possible
= Mawson israduate Centre for Environmental Studics,
University of Adelaide, South Australia 5005.
analogues for future global warming (De Deckker er
al, )988; Petit-Maire et.al. 1991; Street-Perrott 1994),
The .aim of this paper is to consider some of the ways
in which wstudy of geologically recent changes inthe
Australian environment can offer insights into how the
various clements Of Our nutoral environment are |ikely
to respond to [ulure climatic change.
Past climatic changes
Will climatic history repear irself!
The search tor past climatic analogues of a warmer
future planet earth is based on a number of explicit
ond implicit assumptions. A major but unproven
assumption is the notion of cyclic change: “as in the
Past, so, lou, in the future”. Linked to this assumption
of recurrent climatic changes is the equally unproven
assumption of similar recurrent boundary conditions.
Boundary conditions in this context means the global
distribution of land. sea and ice; the global (iistribuuion
al major vegetation zones; and the global albedo
pattern. tosofar as global fuctuauons in surtiee
insoladion and in ice volume are linked 16 eyelet
changes in the ocbitul geometry of the carh at time-
scales of 010° years, vertain climauc changes are
indeed cyclic, at least when set against the last two
mullion years of the Quaternary period (Williams v7
al. 1993), But what of time-scales of shorter duration,
more obviously relevant to present human pre-
uccupalious? Here it is important to distinguish
between climatic change ard climatic vartubility.
Climatic change and climatic: variability
Much contusion over climatic variability and
climatic change stems froma failure to specify the time
a) M.A. J, WILLIAMS
scale involved. Fig. | illustrates climatic variability at
different time scales during the last 0.9 million ycars
(0.9 Ma) of the Quaternary. From 20 000 to 10 000
years ago (20-10 ka), the trend in global climate was
from cold to warm, but relative to the preceding
0.9 Ma, this warming was but one peak in a series of
global climatic fluctuations from cold to warm to cold
again. repeated at least eight times in the last 0.9 Ma.
Tt is important to remember that the Little Ice Age
(Fig. lb) affected both hemispheres more or less
synchronously. World temperatures have increased in
the last hundred years, with a brief return to cooler
4 b
Oh 5 ow (ow
1960 1980 (A) Lune fea Age
1700 @
a si Last Glacial
1880
1100)
ho 0. 06 300 (Last Interglacial
are AD, res
(2)
Max. Min
©
02
04
© os
~ 10°C 08
my,
~ 5x107 km
Fig. |. Climatic variability at different scales in time during
the last 0.9 million years (from Williams et a/. 1993, adapted
from Australian Academy of Science 1976).
temperatures between about 1940 and 1960 (Fig. lat.
None of these global temperature changes seem fo be
linked in any obvious way to ihe steady exponential
increase in the concentration of atmospheric carbon
dioxide — # point conveniently forgotten by over
enthusiastic advocates of enhanced greenhouse
warming.
The reconstruction of past climatic events
The reconstruction of past climatic events is based
on many independent sources of proxy data. These
“nalural archives” include terrestrial and marine
deposits and fossils, as well as evidence from
archaeology. isotope geochemistry and archaeology.
The temporal resolution, temporal range and type of
information which may be gleaned from the proxy data
sources are illustrated in Table 1, and are discussed
in detail by Bradley (1985, 1990) and by Williams ef
al, (1993).
Limitations of palaeaclimatie enquiry
Certain elementary precautions are essential when
using any particular type of evidence to reconstruct
past environinents and climates. Each may be useful
for a particular purpose and for a particular spatial and
temporal time-scale. However, as Williams eral. (1993)
note: “Difticulties anse immediately when we use
Proerustean tactics to force the data to yield
palaeoenvironmental information at particular scales
in space or fime for which those data are totally
inappropriate. A related issue is the precision available
in dating the proxy data or samples used in
reconstructing past events” (ap cit., p. 9).
Reconstruction of past climates thus requires good
chronology as well as careful analysis of appropriate
natural archives. Nor must it be forgotten that climate
is a second-order concept. First-order interpretation
TABLE 1. Characteristics of natural archives used in palacuclimatic reconstruction. (After Bradley 1990, Table I.)
* Minimum sampling interval in most cases. T = temperature; H = humidiry or precipitation; C = chemical compasition
af air (C,), water (C,) ar soil (C,)) B = bronuiss and vegetation patterns; ¥ = valeunic eruptions; M = geomagnetic field
variations, L = sea levels; S — salar activity.
Best Temporal
Archive Resolution*
Historical records day/hr
Tree rings season/yr
Lake sedinients tr to 20 yr
Ice cores ye
Pollen 100 ver
Loess TO) yr
Qceon cores 1000 yr
Corals yr
Paléosals JOO yr
Gcomorphic features 1) yr
Sedimentary rocks yr
‘lemporal
Range (yr) Information derived
1 Ta, BV, M,L,S
107 T.H.C,, B, V, M, §
104408 T, H, Cy, B, V, M
10° T.H,C,, B, VM, 5
10° T.H.B
10° H, B.M
0" T, Cy, BM
10" Cy, L
Ww T,.H, C,, ¥
10" T.H.Y.L
in! B.C. V, M,L
SOME IMPLICATIONS OF PAST CLIMATIC CHANGES IN AUSTRALIA 1
of w natural archive usually provides direct information
about some component of the natural environment.
§uch a§ a river, lake or dune. To infer climate (or some
particular attribute of climate, such as temperature or
precipitation) from a particular element of the
landscape is a far more circuitous and difficult
procedure, with far greater scope for interpretative
error.
Consider, for instance a lake. The outstanding work
by Gasse, Fontes, Street-Perrot’ and co-workers on
reconstructing the climatic history of lakes in Africa
has demonstrated the need for calibration ysing presen(-
day chemical, physical und binlogical data, as well as
the need tor taking due account of local hydrological
factory and of exireme events when deducing
palaeohydrology from palaeolimnology (Fig, 2) (Street-
Perrott & Roberts 1983; Fontes et al. 1985; Gasse er
ul, 1987; Street-Perrott 1991). Only then is it possible
to attempt a reconstruction of paldéoclimate. From
reconstruction of local palacoclimates to regional or
even global palaeoclimatic modelling 1s yet another step
removed from the onginal field data, Provided that the
limitations of pulacoclimatic enquiry are clearly
recognised, and the appropriate steps are followed in
interpreting, past climate Irom proxy data. useful
insights are possible when using past environmental
analogues,
Past eavironmental analogues
A decade has clapsed since Pittock & Salinger (1982,
1983) supgested that the early Holocene climate of
Australia towards 9600 — 700U years ago may be a
suitable analogue for the continent in a CO -warmed
carth, Granted that many early Holocene lake and
LAKE AND AIVER FLUCTUATIONS
AS PALAEOCLIMATIC INDICATORS
PALAEOLJMNOLOGY
(lakas and ¢lvars)
calibratian using
yo oo Medern fotereices
PALAEOHYDROLOGY
(walo’, salls and sofules and stable
isotope falaeobudgets)
calrenie evotits —
affects of local
pdralagloat
lachors
PALAEOCLIMATE
(estimation oF palasa-precipilatlon,
-avaporalinn, -temperature, etc)
~_Y
Fins, 2. Lake and river fluctuations as PalacoClimale indicators
(Source: F, Gasse 1986, ynpublished seminar paper to
Department of Geography, Monash Uneversity),
pollen sites in Australia and Papua New Guinea do
seem. t0 indicate a warmer and/or wetter climate at this
time (Williams 1984), just how reliable is the evidence?
Can it be quantified? Is the "Climatic Optimum” or
“Hypsithermal” in Australia indeed a valid analogue
of future warming? Finally, can we glean anything else
of value to future climate prediction from the record
of early Holocene environmental change in Australia?
In an attempt to answer some of these questions. it
is appropriate to considet the evidence from a variety
of different Holocene sites across Australia. In the
ensuing discussion we consider four distinct types of
Holocene site in tour widely separated localities: the
rainforests of northeas! Queensland, “maar” lakes of
western Victoria, the source-bordering dunes of central
western New South Wales, and the coastal plains of
the Alligator Rivers area in Northern Territory.
The Holocene rainforests of northeast Queensland
Kershaw's (1983) palynological studies of the late
Pleistocene and Holocene vegetation history at four
pollen bearing lake or swamp sites in the Atherton
Tableland of northeast Queensland have tevealed
that the period of maximum rainforest expansion
lasted from 6000-8000 to 3600 years B.P. (Fig. 3).
Lake
Euramoo
Quincan| Bromfield Lynch's
Crater Swamp Tater
1
Maxlrrum
2 Rainforast
Expanslorr
3-
a4
m
Ss
a
a)
a &
Oo
=]
- 7
Time of arrival
B of rainforest
9 Period of
precipitation
increases
1490
Present mean annual tainiall (mm)
Fig. 3. Veretahon changes in northeast Queensland deduced
from pollen analysis of Holocene cater lake sediments,
iron De Deckker vr al, 1988, adapted from Kershaw
1983.)
7600. 1800 2000 2200 2400 2601)
20 M.A. WILLIAMS
However. as Figure 3 clearly shows. the response of
the rainforest to increasing warnith and imereasing
effective precipitation was strongly time-transgressive.
Rainforest appeats m the Bromfield Swamp pollen
record towards 8500 years BP, at Lake Quincan
Crater not until afer 7000 years B.P. The demise of
the ramfOrest at all dour sites was roughly synchronous
Go000 years B.P.), indicating that the rainforest
responded slowly to climatic amelioration (warmer and
wetter conditions) but rapidly to climatic deterioration
(cooler and drier conditions).
Application of the CSIRO binclimate prediction
system developed by H, A, Nix to Kershaw’s
Queensiand rainforest pollen dite enabled Kershaw &
Nix (1989) to derive more quantitative estimates af the
Holocene climates. Their analysis revealed that
Holocene temperature maxima were not achieved in
the Atherton Tableland until S000 years B.P., with
mean annual temperatures up to 3.5° higher than today
persisting until abyut 3500 years B.P) Men annual
Precipitation at that time appears to have been at lenst
300 mm higher than today, and most probably
500-800 mm higher (Kershaw & Nix 1989, De
Deckker et al. 1988).
One obvious conclusion to be drawn from the
Queensland pollen record is that the interval 7000-90)
years B.P, was not the ume of “Chmatic Optimum” for
northeast Queensland. The wettest and warmest period
vame considerably later, and on present evidence seems
to have lasted from 5000 to 3500 years B.P. If this is
true for the tropical northeast of Australia with its
monsoonal suramer rainfall regime is it true also. for
the temperate southeast of the continent, where the
rainfall comes mainly from the westerly airmasses
which pass across the southern margins of the continent
most persistently in winter, when the Antarctic
convergence ts Im ts most northerly postion?
The Holocene crater flakes of western Victoria
Lake Keilambete in Victona is perhaps the best
Studied Holocene lake in Australia (Dodson 1974;
Bowler 1981, De Deckker 1982, Chivas ef al. 1985,
1986), Like other volognic explosion-erater or “maar”
lakes in Victoria, 1 occupies a small closed basin anil
is highly sensitive to changes in precipitation and
evaporation over is catchment. Five “maar” lakes on
western Victoria, including Lake Keilambete. have
yielded usefu) information about Holocene changes.in
the balance between precipitation and evaporation (De
P/E Estimates for 5 Victorian Maar lakes for the Holocene
Kellambete Gnotuk
x1000 years before present
wm
19
Bullenmerri
re
ee
changing
[tow | aan [ttn [ cw ecn | ii
West Basin East Basin
siege 2) se
ee a
50 km
Lowest
Fig. 4. Changes in Holocene precipitation/evaporation (P/E) ratios lor five voleune “maar” lakes in western Victoria. (Froipy
De Deckker ef al, WR)
SOME IMPLICATIONS OF PAST CLIMALIC CHANGES IN AUSTRALIA pa]
Deckker 1982; De Deckker et al. 1988). Highest
precipilation/ evaporauon (P/E) ratios, peak lake levels
and lake salinity minima were all within the interval
7000 to SOO) years B.P. From 5000 to 3000 BLP (when
morthcast Queensland was warmer and wetter than
today) Jake levels were lower and salinity valucs wer
hivzher than in the preeeding 2000 years or sa (Fig, 4)-
One possible urference is that winter Tainfall may
have been lower towanis 501)-3000 years BoP. Eauislly
Plausibly, tolal annual raintall (including both summer
and winter precipitation) may have been reduced al that
timc. We cannot, as yel, choose between these vay
possibilities. What does appear certain is that the time
af greatest effective precipitation (7QQ0-SQKX) years
B.P.) was several thousand years earlier than the time
of maximum effective precipitation in northeast
Queensland, It is tempting to speculate thitt we are
secing the etlects of pwo distinctive climatic systems:
one (in northeast Queensland) controlled by the tropical
summer monsoon, the other (in western Victoria)
controlled hy the winter westerlies. Ifso, the response
of both systems io pusiglacial warming was not
synchronous along the eastern. third of Australia. Nor
may jthe synchronous in the face of any furure global
warning. We Cur tow to the semi-arid inland areas
of New South Wales
The Holocene source-hordering dunes of
central western New South Wales
Holocene palaeoclimaue data are-excecdingly scarce
for the semi-arid and arid areas of Australia which
together comprise 75% of our present land areg, ‘The
late Pleistocene fluctuations in the Willandra Lakes
region of western New South Wales so carefully dared
and clucidated by Bowler (1970', 1983) do aut yield
a Holdcene signal and in any event have far more to
do with runoff from the Eastern Highlands vis the
Pleistocene Lachlan-Willandra river system than with
local changes in rainfall and evaporation (Williams, ef
ul, 1986),
One climatically-sensitive area capable ol providing
useful information about /vcal hydrological events is
the desert margin system of central western New South
Wales studicd by Wasson (19757, 1976) und by
Williams er a/, (199}), The study area occupies about
80 000 km! in the semi-arid region bounded by the
Darling River to the north and west, and the Willandra
'BawLer. 7 M_ (i270) Late Quaternary environments: 4
study of Jakes and associ#ied sediments in southeastern
Austratia, Ph.D, thesis, Australian Natuonal University,
Unpubl,
“Wasson. R. J. 0975) Evolution of alluvial fans in two areas
of southeastern Australia Ph.D. thes, Maequane
University, Unpubl.
Creek distributary of the Lachlan River to the south
(Fig. 5}. There are po perennial rivers iq the entire
area, The ephemeral sircum channel Known docally as
Crewl Greek of Sandy Creek (Fig. 3) [lows
intermittently during very wel years and the West to
east trending linear dunes are toduy vegetated and
stable.
a
o— Lake Nulohars s
+L
~
atari Range
elvanhce «>
[ns
Fig, 5. Map of central western New South Wales showing
ocation of Sandy Creek. The shaded urcays are isolated
Tanpes over 200 10 in elevation (Saurce: Williams et al:
1991).
The Holvcene climatic history of this vast area ts
poorly understood, but (he pattern of local
environmental changes is now reasonably well
documented (Figs 6, 7). Source-bordering dunes were
actively forming from channel sands Jerried in by
Sandy Creek between about S$00 to roughly 600) years
B.P. Both before and after that mterval the dunes were
inactive, vegetated and stable (Williams et al, 1991),
Souree-bordering dunes are dunes which develop
immediately downwind vf a parent source of sand, such
as the sandy bed of w river, asandy lake beach or a
sandy alluvial fan. There are three prerequisiles tor
the formation of source-bordering dunes:
(1) A regular replenishment of the sand supply (for
instance, from a seasunally-fowing sand-bed
channel),
(2) Strong unidirectional winds for at least part of the
year, und
(3) A sparse or limited vegetation cover adjacent to
the sand source.
Any interpretation of the Holocene climatic histury
of this region Must take into account the requitemnents
for source-borering dune formation, the apparent
absence of carly to middle Holocene dune deposits in
22 M. A. J, WILLIAMS
Netrts csifFS csG Males CSiFS CSG Metres
t+ o7-
1,840 + 50 B.P,
Loamy.
fine-medium
red sands
Soft red
sands
Palaeosol
Low angle planar ay.
cross-bedding
dipping downstream
Crude
horizontal
bedding
Cobbles
Sandy clay
Red clayey sand
+ green mottles
§5B: Bronzewing Valley
Fig. 6, Representative alluvial fan stratigraphic sections exposed in the gullied western piedmont of Belarabon Range (see
Fig. 5). The block diagram shows Sandy Creek, the source-bordering dunes and the wind direction. (Source; Williams
etal, 1991).
Years Before Present
1 o ~~
400-3,600 Aboriginal fires. ()
Alluvial and aeclian ganas
3)
ates BBPP 7,400-7,500 a. Hight lake levels (2)
Pedogenic carbonate (4) ~
13,500-15,500 _ Calcareous dust
Alluvial and aeolian sands F
14,000—1 8.000 <O Gypseous jurette 12)
Fig. 7. Generalised late Quaternary piedmont stratigraphy in
the central western New South Wales study area. The
numbers in brackets denote numbers of samples dated, and
includes both radiocarbon and thermoluminescence dates
(Source: Williams eral, 91, revised to show additional
dates, }
Padagenic tarbonale (6) ~S
24,000-32,000 aie Caleareaus dust
Alluvial anc gealiar sands
a)
SOME IMPLICATIONS OF PAST CLIMATIC CHANGES IN AUSTRALIA 23
this area (Fig. 7). and the fact that the present-day * 600 years BLP. to present:
dunes are vegetated and stable, A tentative Hot summers, cold winters
palaeoclimatic interpretation. based mainly upon the Raintall more uniform
geomorphic evidence and needing to be tested by future Wind vyelocitics low and/or
independent work, is 4 follows; Denser vegetation
¢ 10,000-5500 years BP: If this interpretation is correct, any change to a wetter
Rainfall more uniform climate or to. a more seasonal distribution of rainfall,
Wind velocities low with more runolf during the winter months, could lead
Surface well vegetated to a renewed phase of source-bordering dune formation
Erosion and deposition very slow provided the summers remained dry and windy and
® 5500-600 vears BP: the riparian vegetation relatively sparse. Should the
Higher winter rainfall future climate in this area become both warmer and
Seasonal channel flow wetter, the somewhat paradoxical outcome could be
Summers yery hot, dry and windy a replenished sand supply and reactivation of the linear
Summer deflation suurce-bordering dunes.
Arnhem Land Plateau
(Middle Proterozoic
Sandstones & Dolerites)
Sandstone Outlier / 4
e Je fi aa
Wooded Lowlands
Paperbark Swamps
Shelly Sand Ridge
(Holocene Cheniers)
Uranium bearing
Lower Proterozoic
Metasediments
Strike Ridge
Tidal River Sy 1 2 Ag ae
Late Catnozojc
Coastal Plains 5 Sands & Gravels
Late Quaternary
Marine , Estuarine
& Freshwater Clays
Fig. 8. Block diagram showing the Arnhem Land plateau of Northern Territory and the geologically youthful coastal plains
wo the north. (Source: Williams. 1991.)
24 M.A
The Hivecene coastal plains of the
Alligator Rivers area, Northern ‘Territory
The tropicul eoastal plains of Northern Territory
sti in alinos: diameirie contrast w the ephemeral
streaings and desert dimes of semi-arid western New
South Wales. Their significance stems {ron the Tact
that the Holocene coustal plains which extead tor some
thousinds of kilometres from the far narthwese of
Western Australia to the fac northease of Queenstane
are very dynarme landforms, and highly sensitive ti
even minor chasges in sea devel, sediment tnad, salinity
and wave climate (Williams (991). They ure also hese
toa umque and “abundant fauna and Nora (Haynes e¢
al, W991), In addition, the coastal plains west of Darwin
dbut Ono the Arnhem Land plateau (Bias. 8). with its
uhundant galleries of Aboriginal rock art..and its record
ul over SQ 000 years. of prehistoric Aburipiril
senilement (Roberts er al, 1990),
With the melfiing of the ale Pleistocene ice Sheets
of North America and Europe, world sea levels rose
from their last glaeial maximunt level oF about -[35 m
(at 18 QUU years BOP to their present levels towarils
BOWU-7000 years BOP. (Williams-er al’. 193d
The initial rise was rapid, and vast areas of the
contuvental shell off northern Australia were
submerged at @ rate of over 20 metres.a year or roughly
40 emia week. Quce this rise had slowed down, which
it did in the last few thousand yours, coastal nungroves
hes to colonise the intertidal muds. allowing muddy
sediments to accumulate on the old late Pleistowene
land surface inland of the coastal mangrove fringe. The
process accelerated once the sea altained its present
level. and widespread mangrove swamps developed
across the present area of the coastal plains in mid-
1 WILLTAMS
Holocene times, from about 5000 to 2000 years B.P.
(Woodcotle et al, 1985). ‘These mangrove swamps
proved to be highly efficient sediment tnsps, and Were
eventually buried by estuarine and, ultimately, by
freshwater couds (‘Woodrotle etal, 986; Williams 199);
Wasson 1992), The present-day coastal plains are thus
4 relatively youthtul feature, ind are generally less thin
2000 years old,
How these plains might respond to sca leved change
will depend upon 4 vainet of factors. including the
rate of sca level rise, the impact of any climatic change
upon uno and sediment yield in the litlal rivers; Lhe
magnitude and frequeney of future cyclones. und the
relative duration of the wet und dry seasons. Since none
of these variables is. accuracly predictable, speculation
seems unwarrinted beyond noting that the coustal
plains are likely to remain @ geomorphically dynamic
and ucuively developing feature of (he landscape, just
as in the past SOG) years.
Conclusion
Provided certain commun-sense preciutwins are
observed, an appreciation of Holocene climatic und
olher environmental changes in Austrilia can be i
useful guide to pusmble future landscape responses to
global warming whatever its ultimate causes, The
Austnilian landscape is a palimpsest of landforms, soils
and plant associations, all of which will respond to
lulure climate change in a variety of ways, The way
im which our rainforests, lakes. dunes and coastal plains
have responded to Holocene climatic changes offers
us some guide to their possible future responses, It is
highly unlikely that these will be either simple or
synchronous,
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CLIMATE CHANGE AND ITS IMPLICATIONS FOR
SOUTH AUSTRALIAN SOILS
By R. W. FITZPATRICK* & M. J. WRIGHT*
Summary
Fitzpatrick, R. W. & Wright, M. J. (1994) Climate change and its implications for
South Australian soils. Trans. R. Soc. 8. Aust. 118(1), 27-34, 31 May, 1994.
The nature and distribution of dominant soils in South Australia is briefly reviewed
with particular reference to major issues relating to climate change. For this purpose
the State has been divided into two regions: (i) the southern regions or agriculturally
developed area which lies south of latitude 32°S and is mainly used for dryland
cereal/sheep production and (11) the northern region or semi-arid and arid areas which
mostly lie north of latitude 32°S and are mainly used for low intensity grazing of
natural rangeland. A large proportion of South Australia, including many texture
contrast soils of the high rainfall areas, has dispersive soils subject to sodicity and
which are highly prone to waterlogging and salinity throughout a significant
proportion of the profile.
Key Words: climate change, soils, South Australia, soil moisture, soil temperature,
soil organic content, salinity, sodicity, soil acidification.
Transactions of the Royal Sociery of 3. Aust. (994), TES), 27-34
CLIMATE CHANGE AND ITS IMPLICATIONS FOR SOUTH AUSTRALIAN SOILS
by R. W. Fitzpatrick*t & M. J, Wriout*
Summary
birveatrick, RK. W.& Wear, MJ. (1994) Climate change und its implications for South Austrahan sulle eins
R. Sac. S. Aves, WSO), 27-34, 31 May, 1994
The nature and distribution of dontinant soils in South Ausitalia is briefly reviewed with particular reference
to major issues relacing 1 climate change, For this purpose the Stute hay been divided ints two regions; (1) the
southern eeion or agriculturally developed arca which lies south of latitude 32°S and is mainly used for dryland
ceéreal/sheep production und (i) the northern region or semi-arid and arid areas Which mostly le north of latitude
32°S and are mainly used lor low intensity grazing of naturel sangeland. A large proportion of South Australia.
including, many texture contmst soils of the high rainfall areas, has dispersive soils subject lo sudicily aad whieh
are highly prone to walerlogging and salinity throughout a significant proportion of the profile.
Chis paper attempts to forecast the most likely conscquences of plobal climate change on the doniinant soils
of South Australian, The direet iflugnee of imereasing winter temperatures and decreasing winter rainfall on lhe
wide range of soil types (hat o¢cur in South Austrahy is as yet unclear, However, ihe magnitude and extent of
supposed degradation or enhancement of panicular soil morphological properties, and decline or increase of
auil site properties (e.g. salinity or Crosion) are matters of speculation, In fact, these changes can affect the processes
od global climate change by affecling production of greenhouse gases or causing changes in vegetation, How
swils modify under the influence of changing climate will depend principally on seul type. roposraphy and changes
of vegetation.
South Australia’s soils will respond ot global climate change through changes In soil motsbare, soil temperature
and soil organic maiter content, Decline in winter rainfall in the high raintall regions occurring to (he south of
latitude 32°S will have a beneficial effect by substantially reducing seasonal waterlogging and formation of aquic
soils (including non-dal acid sullate sits), In contrast, dryland salinity may continue to expand. with a corresponding
decline in stream water quality.
Tlis nol yet poxsible to accurately quantify regional sort elagges resulting from climate change given the present
Uncertainty about the amounts und rates of global climate change, and particularly concerning regionul patveras
of lensperature, precipitation und coastal geomorphic changes. The cuprentlysinereasing tate of land and water
exploitation in Sonth Austealia will likely. hayes greater inmpact on scils, adverse or benvticial, than the effects
of limale change.
Key Worws: climate change. setts, South Australia. sed! mersiure, sol temperate, soil organic coment,
salinity. sodicity, seil acidification.
Introduction
The greenhouse effect is the warming of the carth
resulting from increases in the cancentralion of carbon
dioxide (CO,) and other radiatively active gases
including methane (CH;), nitrous oxide (NO), ozone
(O,) and the chlorofluorocarbons (CFC's) (hat reduce
the Joss of outyoing infrared radiation (i.e. limit heat
fosses frony earth into space; Table [). A global
increase in CO, has been reliably documented during
the last 100) years and although this is duc mainly to
the burning of fossil Fuels there isan appreciable Mux
of CO, trom the oxidation of soil organic matter und
the burning of forests (Table 1). CO, levels of twice
the present are forecast by the year 2050 (Pearman
1988; Houghton 1990: van Breemen & Feijtel 1990).
Soil is, not only a source of CO, but also of CH, and
N,O (Table 1), Atmospheric warming of about 0.5°C
* CSIRO Division of Soils, Private Bag No. 2. Glen Osmond,
South Australia. 5064,
+ Cusuperative Research Centre fe Sol and Land
Management
has been estimated over the last 100 years, estirnates
of warming up to 5°C over the next 100 years have
been made (Bouwmun 1990), A corresponding
warming of the oveans-and rise of sea level of up to
0.5 mare also forecast, Changes of this magnitude and
rale represent 4 Significant change in our environment
that would have a profound effect on the Earth's
ecosystems and human activities (Bolin et al. 1986:
Wild 1993).
Much of the emphasis in recent discussions of the
impact of greenhouse induced climate change in
Australia has focussed on the atmosphere and on
agricultural and forestry production (Pearman 1988),
However, the effeets of climate change om Australian
soils is also a fundamental issue. Given the present
state of severe soil degradation in Australia (e.g.
Chartres ef a/, 1992) the pessibility of further
degradation should be of national concern,
South Australia is a region which 1s currently
influenced mainly by winter rainfall but where the
tairrent rainfall varies greatly from north to south
Hence, for discussion purposes, we have divided South
rs
oe
house gases, and there contrthidion to global warming (adapted from Wild
for major green
ie international society af seil science, 1990),
eranimles of th
f
‘centrations, increase, residence time, sources and sinks fe
1993; Bounwaun 1990); Standing committee on international pra
TABLE 1, Atraspheric cone
R. W, FITZPATRICK & M. J. WRIGHT
fa Australia into the following two regions: (i) the
gs southern region — the agriculturally developed urea
z a5 which lies south of latitude 32°S and is mainly used
[=e as e fay
ial 4 ai for dryland -cereal/sheep production and (ii) the
a, _ * ’ - - ‘
Pa as 3 as, northern region — the semi-arid and arid areas which
is
wif 7S I Zé ' H atl Jon a}
ul+.2= & Se mostly lie north of latitude 32°S and are mainly used
SeSAL1 1 a for Jow intensity grazing of natural rangeland. The
> greenhouse effect is likely to influence the soils in each
az region differently, The southern region is considered
3B hkely to suffer a drying climate duc to a reduction in
uv ¥ . 2 < : .
= sit winter rainfall whereas the northern region is more
= aE likely to reveive enhanced summer rainfall, The
ait = BRA ™ . “fee » * a
SIF. § & ZE objectives of this paper are to: (i) summarise existing
Scifaiol | a RS information on soils in South Australia, (i) highlight
the most likely consequences of global climate change.
q . ‘ whe . .
nn on the range of South Australian soils and (ili) identify
z= future priorities for research on global climate change
= cp on South Australian soils.
or af
eS 3 :
Bun a8 Soils
Zz 4S ao
A Ce so F F
ol Ex =. aoe = Ber Soils are complex systems which are strongly
a 25s e¢ ; Sane oy 3
z 2 ae eso Fres z 22 < influenced by processes occurring in the atmosphere,
SSRAG+S3R 7922 2d biosphere, hydrosphere. and lithosphere (Fig. 1).
In general, soils develop slowly over thousands of
S years, the absolute rates depending to a large extent
is on past climates. Their component parts also differ in
= Boe *. 8 their rates of development. For example, soil biotic
; S foi gD tl a f :
a ey = Sra eS gece processes resulting from the interaction of the
< zoe er Em 358 atmosphere and biosphere are much more rapid than
SS = u ee A 7) ' . . 4 ‘
| —- © SUstsety ae 2 = soil weathering processes resulting from the interaction
= 384745323 7 TT i
il 5 = of = a g 228° C2 58 of the hydrosphere and lithosphere. Thus, in response
Die Pongo ee = Eg SEZES=ES to greenhouse imduced climate change. the more
D . = Cc os r a6 r “Tr
eo oma FR ase eS oe Sagat environmentally sensitive soil biotic pracesses would
y lolic p
+4 be expected to respond more rapidly (Bouwman 1990;
al Varallyay 1990; Wild 1993). In coastal landscapes,
os me many sails would also suffer relauvely rapid water
i FS : , Bee :
= = erosion, waterlogging and salinisation as sea levels rise
u Ds vers with global warming.
é G 4&8 ww BS
i Eos me an ‘a oS Zz 2 a A h
G nese 25688 imosphere
Clone sH228 ER FS
Sones Boao
y
a
i
£2
BE Biosphere ———» («ais +— Hydrosphere
a
“3
58
me? Osh
SERFS =
nc eee Bs wz ot
gerbe Er
ue eae a 4 i= } n oO
E BES = 8 z c g Lithosphere
ose D5 a S Et ae . " . ° . . .
dEfuss 3 5 ea = “a| Fig. 1. Simplified diggrammatic representation illustrating (hat
Sake 2 z “4 Lt = € soils are complex systents which are influenced by processes
Bee 2 ze ae 2 2 fram the uttiosphere (climate), biosphere (vegetation and
2258 6fa $ G & 2 fauna), hydrosphere (hydrology) and lithosphere (geology
and topography).
CLIMATE CHANGE AND ITS [MFLICATIONS FOR SOUTH AUSTRALIAN SOTLS 24
Influence of lund clearing
An important factor im assessing greenhouse etfect
is the change already brought about to Australian soils
as @ result of Buropean settlement, Land clearing,
praying un) cullivalion generally bave resolled in a
rapid decline of soil organic niatter and organisms and
consequently a deterioration of soil structure and
increasing suscepuibility ky erosion. A marked
reduction in the diversity of fauna and flora of most
ecosystems bas also accompanied European settlement
and this his made our soils and lundseapes less resilient
ty change
The effects of gecenhouse induced climate change,
therefore, must be assessed for soils and landscapes
already degraded: pristine conditions are virtually non-
existent, Given this scenario, there is a signiticant
environmental question t be answered: If there is
greenhouse induced climate change of the magnitude
forecast, what will be its effects on South Australian
sos and landscapes?
Sotl Carbon and the COs cycle
The amount of carbon stored in the world’s soils as
fresh organic matter, stable humus und charcoal, & two
to three times highee than carbon stored in natural
vegetation and crops (Table |, Bouwman 1990:
Standing committee on international programy of the
International Society af Soil Sctence 1990; Wild 1993).
Distribution and major properties of soils in
South Australia
Ciencially speaking, Sour Australian soil landscapes
are extremely variable and complex due. i part, 10
the great age of much of the continent. A signiticant
proportion (>60%) of soils io South Australia are
saline, or sodie and/or texture contrast (ic. duplex —
have bleached sandy to sandy clay loamy topsoil
hurmens overlying clayey subsoil horizons that may,
or may mol be mutiled) (Table 2), Sodic duplex soils
are falierently subject t waterlogeing because of water
perching on the more impermeable clayey sobsoil
horizon and consequently saturating the topsoil
hocizons, especially where there is.an Increased inpul
of water following removal of native vegetation, A
similar effect cam be caused. in the short term, by
excessive irrigation. Therefore, to proceed from u mure
general understanding of the processes of waterlogging
and dryland salinisation to 4 more definite
understanding. of these processes in an individual
catchment, in order to predict the effects of
management or climale change. 1s extremely difficult.
The centri ynd southern Mt Lofty Ranges area is
further compheated by hwhly variahle geology and
weathering patterns of the soils in those Jandscapes.
Tt 1s apparent that in adjacent landscape positions one
is confronted with deeply weathered soils which
contain ancient stored salt, juxtaposed with very
youthful soils on partly weathered rocks which are
generating sallas a result of contemporary weathering
processes.
The higher rain(all areas (i.e. greater (han 500 mm
average annual rainfall) which he south of littitude 32%
ure restricted to coastal and sub coastal plains and
ranges. A high proportion of both regions: has soils
which are sodic throughout or through a significant
proportion of the profile. Table 3 lists the estimated
areas ol’sodic soils in South Australia, south of latitude
32°S. The names used represent general profile
characteristics that may each include several great soil
groups. ‘This has been done to reduce the table to
manageable proportions suitable for reproduction here.
The semi- and arid regions north of this latitude contain
even more significant areas of saline and sodic soils.
particularly the desert lounts and associated red clays.
The table was prepared by te-interpreting Maps
published by Northcote (1960) und Northcote ef al,
(1948), Sheets | und 10, respectively, of the Adus of
Australian Suis; ihe original scale of which was
12 100 000,
TABLE 2. Uiviribulion of saline ane Sedic sails in relation we rainfall in South Austrilia (after Northern & Skene 19721.
Percentage dred of each map don within anowal rainlall zones
Annual Alkaline strongly sodic Now-alkuline sodic
Rainfall Area Saline Uniform Gradationsl Duplex Neutcal Avid Talal
(mm tke Sails texture texture prafile duplex duplex (%
<)50* 450,215 25 9 6¥ 18.7 - _ 31.35
190-254)** 351.510 | 1.8 470 — — 39.9
250-350"=* 82.570 23 11 54.8 V5 - TOT
3SU-SSD6F* 72,745 - is 224 a4 8 42 _ 62.4
> 5a0t** 25,560 12 | 24.2 a4 28 43.2
Total 983,000
* All within northern region of South Australia.
“+ Largely within northern région hut a significant area of southern region inelodial,
rH AL WITHIN sotiTheen regia cecepl Jor small acca at nortberm region in 250-350 zane:
uw KW. FITZPATRICK & M | WRIGHT
On an area basis two broad groups of sodic sails
are prominent in the southern region, the grey-brown
calcurcous louis (36% of the area) and the duplex suils
with alkaline soil reaction trend (18%), thal is, soils
with pH venerully increasing with depth and alkaline
in the subsoil, These two soil groups inclide the
majority of the cereal and sheep producing, land in the
State, Although of much smaller area, the sadic duplex
soils of southern Eyre Peninsula, Yorke Peninsula, the
Mid-North, eastern Mt Lofty Ranges, Kangaroo Island
and the South-Bust ure critically important beeauge they
have high rainfall and a high production potential. They
are alsey important because Unecy indicate the regions
where waterlogging and secondary salinisatien is 4
current or potential problem,
TABLE 3. Estimated vreas af soits in South Australie seats
af latitinde 32°S (madified ufier Nai et al, 1293),
Area
kunt a
Soils that wre sogie throughyut:
al Cliayéey soils. uniform texture
profiies 3450 if
b) Grey-brown culcareous loans,
unitarm and gradational texture
profiles. 75660 30,1
¢) Alkithine duplex soils 3335 lq
Sub-tutul Rd § 4u4
Soils that fave sodie Bubsolls
i) Alkaline duplex. soils 38685 IBS
ec! Neutral duplex soils W185 24
tf) Acidic duplex soils 2930 14
Sub-total 47800 23.4
Tolal sodi¢ soils 2245 64.2
Now sagdic soils;
) Alkuline duplex soils 947 14
nh) Neutral duplex suils WIG ne
t) Acidic duplex swrls 5682 2.7
i) Clayey soils. uniform textire
profiles 1683 0.8
Kk) Girey-brown caleareous foams,
uniform and gradational bexture
profiles 44212 2uu
lh Sandy soily incudinge podzots WSR 1)
Total not sodic soil 7IATS 36.8
Land area South of latitude 32°8 209720 Tw
Prajections for climate change
Forécasis of greenhouse induced climate change ure
based on global models that incorporate the circulation
of the Earth's almosphere and oceans and the alhedo
effect of its polar ice caps. Present models vary in their
forecasts of global CQ, and temperature changes
although a commonly accepted view is that a doubling
oF CO, will result in a temperature rise of 4°C
(Pearman 1988; Wild 1993), These models also predict.
io varying degrees, a Gonsequent merease In
evaporation and previpitation Le. an intensification of
the hydrological cycle with increased rauntall erosiv ity.
Greater uncertainties exist in the prediction of regtonal
climate change und ils.consequences within continents.
In Australia, a “best guess” scenario tas been
developed by Pittock & Nix (986) based on historic
records and a global model from which a rise at
helween 2 and 4,7°C in meun annual temperature by
the year 2030 is predicted (see also Pearman (988),
[i sumimury, lemperslure increases are predicicd to be
ereatest in southern Australia, particularly in winter;
summer raintall areas will receive um increase in
precipitation of up to 50%, and the monsoonal
influence will extend farther south; the winter rainfall
areay Of southern Australia will receive less
precipitation, particularly in winter,
The climatic models vary even more widely in their
prediction of the moisture status of Australian soils
under doubled atmospheric CO; Only uw broad
generalisahon of increased soul moisture in the north
and a decrease in the south can be made.
The uncertainties in predictions of climate chunge
ints the next century are consideruble, Nevertheless,
it 18 clear thal even a modest greenhouse effect will
significantly change the regional climatic regiines of
Australia, In the southern parts of the continent,
particularly, the climate change forecast will create
Urier soil moisture conditions and impose further
limitations fora range of land uses. When this scenario
is superimposed on the present pattern of soil
Gegradalon in South Australia, it is apparent that large
areas of land could be at risk of further degradation,
Not all the predicted! consequences of the greenhouse
effect, however, are negative. Forecast increased
precipitation in the northern part of South Australia
would have (he potential fo increase soil moisture stores
with 2 positive outcome for soil stability and for the
pustoral and agricultural mdustries. In contrast,
Jecreased precipitation in the higher rainfall areas of
southern Sourh Austraha would reduce seasonal
waterlugging (sce beluw),
Impact of past climate change
Because soils and landscapes develop over long
periods, miny are found to contain information about
past climate change, Among the most significant
examples are the soils and associated sediments of
coastal flood plains wbout Australia (Division of Suils,
CSIRO 1983), At the peak of the workd?’s last glaciation
20,000 years ago, sea levels were about 140. m below
the present and the Australian coastline was
consequently much more cxtensive. AL the cnd of
glaciation, plobal seas rose rapidly ta reach their
present level about 6000 years ago. During this rise,
ocean waters invaded large ureas of coustal land aod
embayed all codstul rivers, The rivers depnsitel
CLIMATE CHANGE AND ITS IMPLICATIONS FOR SOUTH AUSTRALIAN SOILS i]
sediment ia mace and estuarine-enviroamems so thir.
today, their coastal fload plains contain a continuous
record of sedimentation. Around Australia. high water
tables. high salinity, aod sulphates characterise the soils
of these flood. plains.
Vastu 4, Possible negative modifications vf lane use due to
Manes in temperature, rainfall antl windiness in South
Aastralia,
Coastline
Rising sea levely.
= wave vrgsion
* flooding of coastal fond plains
*salinindtion of coastal Hood plains
Northern Region (Arid and semi aril zones which
mostly lie warthoof ketimde 32°98)
Increased rainfall and erasiviiye slighty lugher
LOC LANETEN
* mcreased risk of lindstides
® incrensed pink of water erosian
* fouding of lowlands
Southern Region (High raintall zones which lic south of
latitude 32°S)
Changed rainfall invulence. iicivased vrasnvisy
higher lemperalures
* lower sol moisture
* decreased vegeitive cover
* loss of sbil biotic companenty and otganic matic
® dererioration of soil strucnire
* increased risk ol water crosiay
* increased salinisation
© ditenaranon of strean: water quality
* incredsed wind erosion
® (listicr atmosphere
Tn such coastal landscapes, the ale geological pas;
provides an uccurate guide for predicting the
cumsequences of sea level cise nesulting from the
greenhouse effect, These are increased flooding and
in elevation of water tables, and increased salimisation
und encroachment of marine and estuarine
enV Ironments, accompanied by a build up of sulphidic
sediments in expanded mangrove swampy (‘Tables 4,
S\, This will lead to the development of greater areas
ef avid sulphate soils. The area of coastal Mood plan
actually affected, particularly io the gulf regions oF
Sourh Australia, would depend on the magnitude oF
the sew level rise: a special study would be required
tor relivble estimates.
Rises in sca level also cause widespread wave erosion
of coastal svils. near shorelines, as well as
destabilisation of coastal dune Jandscapes (Table 4)
Much of the evidence for these effects in the late
veolagical past have been. covered by the ocean or
obliterated. Nevertheless, a significant risk ty couxdines
around South Australia is apparent (Harvey & Relperio
Unis volume),
‘Tame 5. Possible positive modificarions in kinduse dup to
changes in tenperature, rainfall and wiadiness in South
Australie.
Cnastline
Ristnse seq levels
® flooding und formation of coastal mangrove swaps
Northern Region (Arid and semi-arid zones whieh
mostly lie north of lautude 32°S)
Increased rainfall: slightly higher temperatures
* increased soil moisture
@ increased vepelalive caver
@ reduced. Wafer erosion
* jncreased soil orwinte matter
Southern Region (High rainfall zones whieh he soutlr ot
fatitude 32°)
Decreased rainfall: higher temperarnres
* lower water lables — ground und perched treduced
incidence of seasonal walerlogping)
@ decreased soll acidification
The usefulness of the geological record for predicting
the effect of greenhouse mduced cliniate change on
other Australian soils and landscapes is much less
certain, This is because much of the stratigraphig and
seomorphie evidence for climaic change during the laxt
20,000 years relates to world-wide (Pleistocene) glacial
climates, In Australia, a large part of Tasmania and
a small arca of the mainland were glaciated. inland
lake levels were high, river discharges were higher than
the present, lurge areas of hill slope land were
crusivnally unstable and eroded sediment filled local
alley floors. In central Australia, the dry phases of
the Jate Pleistocene were characterised by extensive
movement of sand dunes and the mobilisation of large
quantities of dust which wus deposited as a clayey
mantle over Jandscapes in eastern Australia These
unstuble soil and landscape conditions changed to
relative stability as climatic warmed during the fast
{0,000 years (Holocene),
If we use the past ax an analogy, the predicted
greenhouse warning should be seen as leading to soil
formation and. therefore, not expected to result in
widespread instability and degradation of soils and
landscapes um non-coastal Australia, However, two
factors muke the present different fron the late
geological past, Furst, the rate of climate change due
to greenhouse warming is predicted to be much quicker
than climate change of the same magnitude in the
geological past. Second, most of our sotls and
landscapes haye already been modified follawing
European settlement an such a Way as lo make them
more vulnerable to rapid environmental change. In
many parts of South Australia, thresholds of stability
have been exceeded, resulting in accelerated
walctlogging, saliosauon and erosiog (Fiepauick
al. 1992, 1993). If present land use ts not responsive
Ls RW. FITAZPAPRICK & M_ J. WRIGHT
ti ereenhnuse induced climate change, these and other
fanny ol soil degradation are likely to increase,
particularly ip the southern region where decreascd
soil mowiure wuld make present land use marginal
ir sume Areas
In this connovnon, Pitock & Nix (1986) have
modelled changes in plant productivity a5 4 result of
projected greenhouse induced climate change in
Australia. In northern Australia, the positive outeeme
of the greenhouse effect in inereased precipitation and
vegetative cover could mean i shill inwards an anercuse
in the stability of Sauls ly water erosion. As elsewhere,
however. much would depend on Land use practices,
A decrease in plant production is anticipated in the
southem region of South Australia, signalling a
decreuse in soil vegetative cover and soil biotic activity
ft is Suggested that these effects would be relatively
rapid with 4 cotresponding decrease in sei) structural
stability and au) inereased susceptibility 6 erosion Li
southem Australia, lherefure, the year 2050 dould see
w return towards conditions that prevailed m the Tale
Pleistocene given the Opper Qvirst) greenhimse
scenarin,
Projections for Soils and Landscapes
From the foregoing, ts possible 10 oudline some
uf ie faain Consequences of greenhouse induced
climate change in South Australan soils and
landscapes. Although these can only be froadly slated.
they have a potential value in idenefying like areas oF
Positive und negative outcomes. These are summiarisect
in Tables 4 and 5S. Lt may be argued that some of these
projections are alarmist, However, waterloysiig.
sulinisation, acidification, wind and water erosian are
already major problems in South Australia anu there
is no evidence that they are under control (Fitzpatrick
etal. 1092. 1993. 1944: Naidu ep al, 1993). Even the
conservative “niddle scenarios” of greenhouse induced
climate change would exacerbate these problems in
many areas.
Sail erasian
South Australia has a high proportion of sodic and
saline soils (Northecte & Skene 1972: Naidu er al.
(993), The xudie:saline rao of approximately 3-1
mages hetween 4 anil WO times that reported tor other
continents (Szaboles (989) and is conststent with the
high proportion of sodium present in soil solutions and
groundwaters. [nm South Australia, most sodium
affected sails are the result of past inundations by
brackish water supplemented possibly by cyclic sult
Thus, im subsoils, chloride 4s the dominant anion and
exchunveable Mg/Ca ratios are high. The imcidence
of sodicity often coincides with the spatial distribution
iW duplex soils. This associuun nibcates a patern of
environtnemal hazard which appears to at beast broadly
coincide with many areas under agriculture in the
southern region, and substantially limits its
productivity, Saline, sodic and sodic duplex soils are
often predispused te land degradation (e.g. Naidu ez
al, 1993. Isbell e¢ 7/1983) and, signifieunily, cover
> AN ul the South Australian land mass. The effects
of adsorbed sodium on clay dispersien are most
pronounced in denus alkaline subsails which comprise
over 86% of Australian sodic soils (Northcote & Skene
1972),
The impart of sail sodicuy on the eaviranment is
# inost important land degradation issue in Sauth
Australia. Both primary and secondary sodification of
soily cuuses undesirable changes in soil structure,
severe hillslope crosion, waterlogging and erosion of
downstream watercourses. The assucialed increase in
colloid and nutrient loading of streams alsa conte ibutes
ty sedimentution and the consequent loss al reserverr
storuge capacity, Iagether with serous water quality
problems. Heavever. itis the contact of sodium-attected
soils with water in the form of rain. splash, surface
runoff, thoaugh Now, natural ground water flaw, or that
pumped for irmgation frony other catehments, which
provides He expanded scale on Which environmental
problems and harards become must conspicuous
(Fitépatrick ef al. 1994).
Tn South Australia generally, climate change is likely
to increase the tncidence of water erosion (Table 4),
Although lower preetpilauion an the south wall generally
reduce the rate of water erosion, this 1s counterhulunced
by the less intensive sail conservation influence of poor
vegetative growth due tran inadequate moisture supply
for plants (cxucerbited by an increase in temperature,
as well), Lower precipitation and higher temperatures
will alsu lead to increased wind erosion, especially
where sandy soils are dominant (ive i the semi anid
northern portions -of the southern region). In the
northern région, increased summer rainfall is Itkely
to be of higher intensity than at present and i, thus,
expected to present o higher erosion risk here tou,
particularly to the widespread saline and sodic desert
luans and red clays of the stony tablelands,
Waierlogeing und dryland saliniry
Extensive deforestation which followed Europes
settlement im South Australia has conrnbuted to a
detrimental shift in the hydroligic balance of muny
¢utchments, This has resulted in 4 rise of perched and
saline groundwater tables causing waterlogging anil
dryland salinity to be a major land degradation problem
in South Austnilia, Over 220.000 ha oF land are
estunated. to be affected by dryland salinity throughout
the major agricultural districts at an annual cost to this
state of approximately $25 nullion in lost agricultural
production. The problem is worsening. In particulir.
there is # prowimg Concern by pruperty hulders in the
MI JLofiy Ranges over fhe rapal merease in saline,
CLIMATE CHANGE AND ITS IMPLICATIONS FOR SOUTIL AUSTRALIAN SOILS 4s
structureless acid sulphate sails that are waterlogged
and highly prime te water erosion (Firzpauick er al,
1992, 1993)
Bespne an obvious reduction in seasonal
Waterlopgzing due ta déeline in winter raintall in high
rainfall regions south of latitude 32°S, dryland salinity
may continue to expand with x corresponding decline
instream water quality (Tables.4 and 5). According
to Sadler et al, (1988), the processes involved in such
“saline” sodie soul cavironments, due to redueed winter
precipitation, ave further complicated by the balance
hetweern rates of leaching and recharge. Gverall, the
consequent reduction of avater quality and land
capability are, therefore, only products of 4 counples
set of interactions which can occur between. the
following factors in Mediterrancan climates: sadium-
affected soils, ambient levels of soluble salts and the
teniporal Now patterns, spatial distribution. relative
comributions and quality of surface water, and through
Naw and ground water In particular, the leche of
saline sodic soils with good quality water te.
Tatnwalet) also poses a threat iy structural stability in
huildinus, bridves, earth dams and embankments
(Fitzpatrick er ef 1994)
Seil acidification
In the Southern region, a. reduction in winter
Precipijation may decrease downward infiltration and
leaching of water and hence lower the rale at soil
acidification. Similarly, a decrease in winter
preipitation and increase in winter temperatures will
discourage heath vegetation which produces acidifying,
litter (Table 4), In contrast. in those presently high
raintall regivns where climate change will cause an
iNerease Mn Sumer precipitauion, soil acidification may
accelerale due to stronger leaching snd chemical
reactivity (eg carbonate dissolution may occur)
Bidlegieal charges including the rutrient regime
Graetz ef al. (1988) in discussing the implications
of climate change: tor arid zone vegelaton stresses the
importance af the spatial patterns of soil types.
Furthermore, given the importance of this spatial
dimension, there 164 requirement, as yet not met, Lor
models of vegelation response to plant-available
myistire und avaiable nutrients thal.ure specitic to the
major soil types. Three reqhiremients Were identified
tor forecasting vegetation change. The first was that.
becuuse the spatial patterns of suil type are still
expected to determine the distribution of vegetation,
it is essential that the forceasts of future climatic
conditions have a spatial resolution equivalent to that
of the mupped soil landscapes.
We consider that much of the carbon stored in South
Aumralian sous is in the torn of charcoal due to a Jong
history of burning, With climate change in the
directions discussed, it is reasonable to expect that ii
the northern regions, biomass, and hence charcoul will
increase, whereas in the suutbern region reduced
canfall will lead toa generally reduced biomass and
consequent reduction in the amount of charcoal
produced, even though line trequency. could wierease
because of higher temperatures and lowered moisture
Carneclusiins
Changing agricultural practices ({hat lead to erosion,
salinisuhen, sodihealion, acidification and
waterlogging) make jt difficult to monitor the effects
of climate change in individual cutehments, Special
rescarch that takes account of these complications 15
needed on specific South Austritlian soil types lo
quantify regional effects of climate change and ty plan
yinilegies fo cope with te changes because of
implications for the well being of our communities and
the environment. Th would appear thal the diornnmently
pastoral activities of the northern region stand to benefit
by reason of incrensed Summer rainksll ala time when
temperatures are suitable for major growth of herbage:
Salinity of surtiace sails, at least, should alse be reluced
by the increased precipitation (Brinkman & Brammer
1990). In the southem region, on the other hand,
agricultural pursuits could be adversely affected by
reduced rainfall, although increased (emperusures will
improve winter growth of cereal ctops. Agricultural
activities may also be able to expand somewhat in the
northern marginal fringe to take advantage of increased
rainfall there, Finally, a] a time when the demand lor
good quality water js esculating, the limited
untdlersignding of the factors and interrelatinuships
involved in soil and water management may well be
further compheated hy » changing, global climate
induced by the “greenhouse effect”
Acknowledgement
The authors Wish to acknowledge Dr P, H- Walker.
formally. of CSIRO Division of Soils, who contributed
significantly tu dhis paper by providing expert advice
in the form ofa craft review chat formed an original
framework for it
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SzaBocs, [, (1989) “Salt affected soils” (CRC Press, Boca
Raton, Florida).
STANDING COMMITTEE ON INTERNATIONAL PROGRAMMES OF
THt INTERNATIONAL Society OF Sot ScieENCcE (ISSS)
(1990) “At global change . . . Do soils matter?” (c/o ISRIC,
P.O, Box 353, 6700 AJ Wageningen, Netherlands),
Van Breemen, N, & Feurnn, T. C. J. (1990) Soil processes
and properties involved in the production of greenhouse
gases, with special relevance to Soil Taxonomic systems,
In Bouwman, A_ F. (Ed_) “Soils and the Greenhouse Effect”
(Wiley, Chichester).
VARALLYaY, G. Y. (1990) Consequences of climate changes
induced in soil degradation processes. Trans. 14th Int,
Congr. Soil Sci.. Kyoto, 1990. 265-270.
WiLp, A. (1993) Soils and the environment: an introduction.
Chapter 1, (Cambridge University Press, Cambridge).
CLIMATE CHANGE AND ITS HYDROLOGICAL
IMPLICATIONS
FOR SOUTH AUSTRALIA
By B. C. BATES*, S. P. CHARLES*, N. R. SUMNER*, & P.M. FLEMING?
Summary
Bates, B.C., Charles, S. P., Sumner, N. R., & Fleming, P. M. (1994) Climate change
and its hydrologic implications for South Australia. Trans. R. Soc. S. Aust. 118(1),
35-43, 31 May, 1994.
The possible effects of doubled atmospheric concentrations of carbon dioxide on the
North Para River at Penrice catchment are evaluated using results from a general
circulation model, a stochastic weather generator and a conceptual water balance
model. Although results show only a marginal decrease in median monthly runoff
during winter months, large increases in monthly runoff maxima are indicated for
August and September due to large increases in extreme monthly rainfall. Modest
increases in evapotranspiration were also indicated for these months.
Key Words: Climate change, hydrology, South Australia, water yield.
Treva tins vf the Koval Secen of & Ame, US) LET Ly
35-43
au
CLIMATE CHANGE AND f'l'S HYDROLOGICAL IMPLICATIONS
FOR SOUTH AUSTRALIA
by B.C, BatTes* S. P Coaries* N, R, SumNrr*® & PM. FLEMING=
Sumeary
Hates, BL C., Cuaknes, 5. PB, Sumner, N. R., & Fleming. PM (1994) Climate change and its hydrologic implications
for Soot Australia Trans, R. Soe. S. Aust, 80), 35-43, 3) May, 1994,
The possible effects of doubled atmospheric concentrations of carbon diceide othe North Pura River al Penrice calchinent
ure evalualed asm results from) a general circulation model. a stochastic weather generator ynd a conceptual water halance
model, Although results show only a mitrginal decrease in tiedian monthly funoff during winter mondhs, large increases
in Monthly unolt machina are mdicated tor August and Seprember due to large increases in extreme mounttily ruintall, Modest
Increases in evapotranspiration were also indteated for these Maoaths
Kev Woros: Cliniite chunge, hydrology, South Australia. water viel
Introduction
Global climate change caused by rising atmospheric
concentrations ol carbon dioxide (CQ) and other
trace gpuses may have a significant unpuct on regional
Willer resuurecs, Reecot research suggests that plausible
climatic changes will affect the timing and magnitude
of cunoff and soi moisture, change lake levels. and
increase evapotranspiration (Cohen 1986; Lettenmaier
& Gan 1990; Allene al. 1991, Lettenmaier & Sheer
1991. Mimikou er afl 1991; Nash & Gleick 199);
Panagoulia 1992), Such scenarios have important
implications (or future water resources planning and
Management, the cnvironment and the national
economy.
Mest hydrologic scenarios are based on the climatic
predictions of numerical models of the generil
circulation of the atmosphere. General Circulation
Models (GCMs) can produce long-term simulations
of the energy and water Muxes in the atmosphere and
land and ocean surfaces on a global computational grid
uf cells and a number of vertical layers, Predictions
af changes in climatic variables such as precipitation
und teinperature ave considered to be more reliable than
those for runoff and soil moisture (Gleick 1989). The
predictions are provided as sputial averages over areas
of the order of (0% to 10° km? due te ihe limrtations
of present-day: computers. Current GCMs perform
reasonably well in simulating ihe present climate with
respect to annual or seasonal averages at this spatial
scule. However, the direct use of GCM outputs to drive
hydrelugic models is considered to be impnyper due
to the coarse (relative to river basin scale} resolutiog
* Division of Water Resources, CSIRO, Private Bag P.O,
Wembley. Wester Australia 601+
+ Division of Water Resourees. CSIRO, GPO Bow [ber
Canberra, Australian Capital Territory 2601
of the spatial grids used by GCMs and the simplified
GCM represemations of land surface processes, energy
transfer wilhin oceans, wid subgrid-scale atmospheric
processes such as convective storms, Moreover, GCM-
based assessments uf climate change are based on
steady-state simulations of current climate and the
climate associvted with a doubling of current
atinospheri¢ concentrations of CQ,, whereas in
reality the concentration of CO, is. increasing
continuously. Despite these limitations. GCMs offer
the most detailed quantitative information on potential
large-scale climatic changes due to increasing
utrnospherie concentrations of trace gases
Consequently, assessments of the impact of climate
change on hydrologic systems frequently use doubled
CO, scenarios consisting of a spectrum of unitorm
shifts to historical temperature series and scalings of
histurical precipitation scenes based on GCM trends.
The size of the adjustment may vary from month tw
month to reflect seasonality in the assumed changes,
Estimates of potential cvapotranspiration for the
changed climate scenarios are usually oblained by using
simple sculing factors, sometimes varying seasonally,
and by laking qualitative account of precipitation and
temperature trends, The historical series and changed
climate scenurios are usedl as input to mathematical
models of hydrologic processes anu the model outputs
vare evaluated to discern pussible changes in soil
moisture and water yield for a given catchinent.
It is frequently argued that working with hypothetical
scenarios suits the purpose of a sensitivity analysis of
water resources and that results are not intended to be
4 predicuion of changes. However, this approach suffers
from three major limitations. First. ic Ignores any
changes in the distribution and frequency of
precipitation events and any changes in the nature and
variability of temperature series, This may be
regardless of whether such changes wre indieated by
3
GCM predittions Second, the use of hypathetical
soominos based on an arbitrary: number of temperature
wid precipitation perturbations and stationary wind and
relative hunndity series, say, 18 not realistic and may
lack the imerdal consistency of GCM simulations of
fulure climates, Third, concurrent historical climatic
series are relatively: short in length which may
compromise the evaluation of the response of a
hydrologic system to elimaté variability as well vs
climate change,
At alternative to the conventional approach is to vse
4 stochastic model representing daily weather
vuriauons aca iocation, The parameters of such a model
characterise the behaviour of the present day climate,
Changed chute sequenees can be produced by
adjusting the model parameters ina maine consistent
with GCM trends. The model is then used to generate
long-lerm sequences of synthetic dally weather records
which in urn are used to drive a hydrologic model
(Wilks. 1992; Bates et uf, 1993; Charles eral 1993)
Such an approach allows consideration of changes in
the distribution and [requency of precipitation events
und chines in the variability of other efimaue
variahles. It also preserves the mtcrnal coosmtency of
GCM simulations of future climates
In this paper, an attempt made to assess The impact
of a doubling of current atmospheric concentrations
of CO, on the North Puri River ut Penriee catchmenc
in Saoth Austrglia, Historical climatic and hydratogic
series und results from a sinule GCM, a Stochusuc
wauther geucrator, and a conceptual water balance
model have been used. Conclusions are drawn on
possible changes im Water yield ay a cesult of climate
change.
Methads
Creneral circulation nuutel
The GCM used in this study, CS7ROG, has been
develaped by the CSIRO Division ol Atmospheric
Research, The model operates with nine vertical levels
in the atmosphere and a horizontal resolution uf about
UH km ~ 600 km. A computauonal ine step ol St
minutes is used, The simulated climate dala come from
equilibrium (constant CO, concentration) tuus tor
present day (control) and future (dovbled CO.)
climates. A complete descrippon ol the mudel ws given
by McGregor et al (1993).
Charles ev af, (1993) reported distinct differences
between the monthly raintall patterns found in
Australian meteorological station records and those in
the CS/ROY control nuo. The differences were with
respect Ky muinfall amount and its peographicil
distribuuon, Maximuns daily grid cell rainfalls for
CSIRO control runs were also found lo be between
‘wand '4 ofthe maxinam daily rainfalls recorded on
nesoscale walersheds, Nevertheless, the comparison
36 & C BATES. S. P. CILARLES. N. R. SUMNER & PM. FLEMING
of GCM rainfalls. by Charles ef u/.. revealed a general
(rend towards increasing rainfall amounts and changes
in the rainfall occurrence process under doubled CO,
conditions. The latter is consistent with the work ol
Gordon ef al, (1992) who found marked changes jy
(he magnitude and frequency of extreme raintall events
when comparing results from equilibria experiments
with the CS/RO4 GCM,
SMechastic weather generalyr
The stochastic weather generator used in this study
is based on the WGEN generator deseribed by
Richardson & Wright (1984), The daily climatic
variables simulated by WGEN are precipitation
occurrence and amount. maximum and minimum
lemperature, and globitl solar radiation (R,)-
Precipitation occurrence is described by a two-state
(wel or dry day), first-order Markoy chain, the
transition probubililies for a given location are allowed
to vary (hteugh an anaual cycle by defining separate
probabilities for each of the [2 calendar months:
py = Prid, =i | day = 7
where py, = probability that a day Oy state ¢ will be
followed by a day in state 7 (7,7 = 0 denotes u dry day;
i, J =t detwtes a wet day): und /, =indicator
variable denoting the presence or absenee 4
precipitation on day a, The variation of precipitation
ndumly on wel days is characterised using cither a
gamma or mixed exponential distribution,
‘The temperature and solar radiation components are
represented as.
x*(7) = ANF) + Beit)
where x = (3 % 1) veetor oF climatic variables:
© =(3 * 1) random forcing vector consisting of three
independent standard normal variates; A and
B — (3 ™% 3) matrices obtained from the lag-( and
lag-l correlation matrices for the components OF s
(Matthias 1967), and the asterisk Genotes
standardisation:
SAE = ALY — fg) Foyt)
in which kK— 1. 2. 3 and y—O. be yt) = mean
for climatic variable A and state jo and
a0 = corresponding standard deviation. The annual
cycles Of jy, and ay, dire modelled by simgle Fourier
harmonies with fixed phase angles
HYDROLOGICAL IMPLICATIONS OF CLIMATE CRANGE FORK STH. AUSTRALTA x7
Jnvestigation of historical Australian daily max anu
and minimum temperature series conditioned on wet
and dry days has shown thit the use of 4 single
hurmanic to describe annual cycles of means and
standard deviations is often unjustified, Higher
harmontes. ure present if the mean series and the
sland deviation series ure more tealistnatlly
represented by fixed monthly values tsee alsvv
MeCuskill 1992), Thus a mean temperature series (7)
tay be writer as
TA) = Hy) | E Ry coy Jarre (b+ a) / 365] -F F,
where K=t. 2) =O. te + =Jolian date.
4 = constant: ay=3; RK, = amplitude of {he rth
harmonic: w = 3,14 in (his case; = phase anple:
und 5, = tth residual,
Similarly. investigation of Australian solar radiation
series has indicated that the fitting of single harmonics,
Ihe use Of a fixed phase angle and the use of the
generation scheme defined above ts an inadequate
approwch (sve Bates eral. 1993), Our approach tavolves
the calculation of the upper envelope for clear day
conditions, R,. (A documented FORTRAN-T7
compuler program tor calculating the upper envelope
can be ubtained from the authors). This thearetcal
maximum is based on geographical location and
uverage clear day atmospheric conditions Por each
calendar month, a generulised beta distribution is fined
to the daily resuluals (the difference between the
theoretical maximum and recorded data) for wet .and
dry days,
Climate sequences for doubled CO, conditions
Wilks (1992) presented. a method for the adaption
of stochastic daily weather models fitted to current
elinnitic series to the generation of synthetic series for
future climates. “The adjustments to the model
parameters were made in a manher consistent with the
changes in monthly: statistics derived from comparisons
al GCM runs for contrat and doubled CO,
conditions. This approach was based-on the oouon thin
GCM results ure often available in terms of monthly
rather thin daily values or even monthly means and
varianees, The CSTRO9 GCM runs provide daily values
lor 30 climatic variables for 30 year periods, The
variables include; precipitation; humidity at level t;
maximum and minimum screen temperature:
temperature at level |; and.nct solar radiarioo at ground
level (&,,). Consequently, our method for making
parameter adjustments to stochastic duily weather
models uses this information,
Let P = monthly precipitation for a month
comprised of Nuduys, The mean af P and is varinee
ane defined by (Wilks 1992)
wy) — Noreep
wtP) = Naa? ita = abe !(l-d)
where a = uncenditional probability of a wet day
le =poll laa pk a 8 = shape and scale
parameters of the gamma distribution: and
d= pPy-Py WS # Measure OF persistence. For
simulating doubled CO, conditrons, we construct {he
rats (Wilks 1992):
BPD) Cote
wP) rap
(PY) — ae'B [1 bec! (Lay +d)! (ed)
o (PF) pais? [1 + a(l-ryl + d) / ed
where the prime deootes doubled CO, conditions.
The solution of the above equations requires twa
additional constraints, For catchments in the arid and
semi-arid regions of Australia where x < 05, we set
wv! =m, and d' =day/ dy) in which the
subseripts | and 2 denote GCM values fora nearby
GCM wrid cell tor conwol and doubled CO,
conditions, respectively,
For mean doubled CO, daily maximum and
mninimum temperatures, we assume:
Tylt) = TAO + Ty — They
where the prime denotes doubled CO, conditions;
7 = harmonics fited to the observed mean series; and
T, P= harmoniés fitted 1 screen temperatures
from the control and doubled CO, GCM runs,
respectively. Standard deviations are assumed to be
unchanged.
Clear day solar radiation is affected by precipitable
water content and CO, concentration, Here we derive
daily tevel 1 vapour pressures using the temperatures
and relative humidity. at level | and hence upper
envelopes for the control and doubled CO, GCM
runs. The GCM solar radiation is R, = R,, / (a)
where a is jhe albedo assigned to the GCM grid cell.
Generalised beta distributions are fitted to the
corresponding residuals (R,-R,) for each calendar
month and rainfall state. Parameter estimates for
doubled CO, residual distributions are obtained by
the method of moments, Here the historical residual
means and variances tor cactr monthare scaled by the
ratios of the corresponding doubled CO, and control
means and variances, respectively. The upper envelope
for doubled CO, conditions is phtained by scaling
historical daily vapour pressures by the ratio of derived
doubled CO, and contro) GCM vapour pressures.
4S RC BATES, & P. CHARLES, N. R, SUMNER & P.M. FLEMING
Hitter balance model
A modified version of Boughton’s (1984) SFB model
was used for the study (Fig. 1), The madel uses daily
rainfall and potential evapotranspiration data us input
to estimate monthly streamflow. The original model
has three parameters requiring calibration: the surface
slurage capacity (S$). the daily infiltration capacity (F)
cantrolling the movement of water from the surface
store to lower store; and the basellaw factor (8)
determining the portion of the daily depletion of water
in the lower store that appeurs as basefluw (O<B<1),
The faur remaining parameters are fixed: the fraction
of the surtace storage capacity that does not drain to
the lower store (VDC = 0.5): the maximum limiting
rate of evapotranspiration (£,,,, = %.9 mm d!); the
wetee
lower store depletion factor (DPF = 0,005); and a
baseflow threshold for the lower store
(SDR,,,, = 25 mm) defining the depth of water in the
lower stoce al which baseflow will cease.
roll
t Eg
PX 0,
Surface Storage
S
ie
Lower
t
Storage Ob
rt. =8-DPF'SS
| SDRmas
Dp
= (1 B}DPF-SS
Hip. 1 Structure of the SFB model.
Drainage
= (1-NOC}§
Non-drainage
=NDO'S
‘The model operations may he summarised as
follows. Rainfall begins to fill the surlace store and
any water in excess of the non-drainage component of
that store infiltrates into the Jower store at 4 maximum
daily rate of F mm d'. Surface runoff (Q.) oceurs
when the drainage component of the surfave store 1s
full and may be written as
QO, = P — F tanh (P/F)
where P = rainfull excess remaining after the surface
store is filled. The surfiice store contents are depleted
by evapotranspiration which oceurs at the potentlal rate
(E,,,) When the non-drainage component is full,
Otherwise. the actual evapotranspiration rate (&,,) is
given by
E, = min f&,,.5 / (NDCS): E,,,,|
where » = depth of water im the non-drainage
component of the surtace store. The lower store is
depleted by deep percolation (D,,) and baseflow (Q,,)
which are defined by
D, = (!-B). DPF SS
Q;, = BDPE(SS-SDR
wee )
where SS = depth of water in the lower store.
In this study, estimates of daily potential
evapotranspiralion in mm d! were obtained using the
Priestley-Taylor equation (Priestiey & ‘Taylor 1972):
L Ey = aR, ANA + ¥)
where L = latent heat of vaporisation in MJ kg", rz
= Lain this study, A = slope of the vapour pressure
curve in kPaeC', RX, = net radiation in MJ m? d'
and + = psychrometric constant in kKPa?C'.
Although the SFB model parameters have some
physical basis. they cannot be readily determined by
physicul measurement. Thus parameter estimates must
be obtained by fitting computed to observed monthly
streamflow hydtographs, Formal optimisation
techniques can be used to facilitate the estimation
process. These lechniques use a subjectively chosen
colterion (the ‘objective function’) to quantity
discrepancies between the computed and observed
hydrographs for a given set of parametér values. The
estimates of the inodel parameters are those parameter
values which result in the minimum possible value of
the objective function, The accuracy of these estimates
affects the accuracy of streamflow predictions.
In this study, the estimates for the SFB model
parameters were obtained using a simulated annealing
algorithm (Kirkpatrick ef a/, 1983; Press eral, 1992),
Simulated anneulme 15 a stochastic, multivariate
optimisation technique which seeks the global or near
global minimum of 4 user-defined objective function
without geting Wapped in a local minimum. The
function need not be smooth or even continuous. in its
domain. The method can he considered as a biased
random walk that samples the full parameter space and
provides @ solution that is independent of the starting
HY PROLOGICAL, IMPLICATIONS OF CLIMATE. CHANGE FOR SOUTH AUSIRALIA a9
point. I can aecept i ‘move’ thal increases the value
ab the objective function a5 a part of @ full series af
moves for which the general trend is to decrease the
lunctiom value, Dewils of tbe algorithm used will be
published elsewhere,
The objective functron used inthis study may be
Whitten as
a
mus Ls,
fw fF tel
where the model parameters vector 8 = (8. FB, NDC,
Ena DPF. SDR,,,.) in which the prime denotes the
transpose OF a Vector, #4 = number of tonths iy the
observed streamflow hydrograph record; & = number
ol months inthe ‘warm-up’ period which fs excluded
from the calculation of the objective funetion value;
und the disturbance z, ts defined by
MW « Ay
.) (QD, + Ad) > (Q, + Aa) yy &, #0, Ay=U
mowhich @ = observed streamflow. @ = computed
streamflow (Q = QO, + G\; and Ay A =
franstormation constants which can be estimated by
(nial and error (Bates & Walls I988), Generally, the
best ftw are obtarned with Ay < 1 und Ay = 0
Case Study
Deseription of catchment anet cletea
The above methodology was applied tn the Narth
Para River at Penrice catchment in South Australis.
The catchment comprises an weea of (8 kor and 1s
located within the Gawler River basin some 50 kn
north cast of Adelaide. It rises from 300 i to S00 m
AHD and has duplex scils and a mined ground
cover!, Land use includes horticuliure, viticullure,
sawing. and arable farming.
Daily streamflow at the Penrice sauging Station
(34° 28'S, 139° 4B) and daily rainfalls recorded at
Angustun Post Office (34° 30'S, 139° 3’B) and
Keynetun (34° 33S. 139° &'F) for the period January
1978 ta December 1989 were used in the study, Over
this period, the mean annual runoff was 6.200 MI
(30 mm) and the ean annual rainfall (65% of which
fylls mn the period May to September) was 550 inm.
The average daily maximum winter and summer
temperatures were 13°C and 29°C, respectively (Chiew
& McMahan 1993!).
| Crew, FH, S. & McMauon, T, A, (1993) Complete set
of daily minfall, potential evapotanspiration und streamflow
data for 28 unregulittud Australian catchments. Centre fir
Eoviranmenal Applied Hydrology. University of
Melbourne. 34 pp. (unpubl >
Estimates of daily potential evapotrunspiration were
Obtained wusing the guidelines described by Smith
(1991), clear day radiation and day length estimates,
und temperature and sunshine hours data from the
Nurivotpa Vittcuhural climate station (34° 29'S, 139°
O'R) which is located some 10 kim from the eutchment
cuntrotd,
Model calthration
Preliminary investigations reveyled that the
performance of the 3-parameter SFB model was
inadequate. However, satistagtory model fits were
obtained by selfing A, = Of E,.= 89 mm dt,
NBC = 0.5, SDR,,,, = Omm, und allowing the
parumeters $, 68, and DPF to vary dung ealibrarion
runs. The finul parameter estimates obtained wene:
S=l65 mm, F=0.9mm dl; #=HAld, and
DPF = 012, The B and DPF estimates indicate (hat
there is a significant loss of water from tie catchment
lo the regional groundwater,
An important assumption in this study is thal the S&B
model is able to simulate runoff from the North Para
cutchment under climati¢ conditions thit are different
to those for which the model has been calibrated, For
example, the eflect of possible chinges in plint
Lranspinition ralcs und vegeiative cover due tw CO,
doubling on the model parameters is ignored. This was
considered to be a réasoouble assumption given che
current level of uncertainty regarding the nature and
magnitude of these changes.
Results
To simplify the analysis of the results, the behaviours
of three variables under control (Ll « CO.) and
doubled CO, (2 x COs) conditions were examined:
tl) monthly raintall, (2) monthly evapoternspiration:
and (3) monthly total runoff. Synthetic ¢limane
sequences of 1,000 years duration were generated for
control and doubled CO, conditions and used io drive
the S&B model. Exploratory analyses of the data
revealed that the variable distributions were highly
skewed, Thus the median rather than the mean was
adopted as the measure of central tendency
Fig, 2 compares the distributions ol simulated
monthly caintall for control and doubled CO,
conditions, "Three features are worthy of note under
doubled CO, conditions: (1) there is an inerease in
minimum monthly rainfall for the period from June
io August and a decrease for the months of Mary,
September and October: (2) median rainfalls. for the
period from March 10 November ure greater than or
équil to those for present day conditions: and (3) there
ate marked increases in extreme (high) rainfalls for
the month of January. and the period from August lo
October.
Fig, 3. compures the distributions of modelled
monthly evapotranspiation for coairel and doubled
40 B. C, BATES, 8. P. CHARLES, N. R. SUMNER & P. M. FLEMING
TaBLe 1. Seasonality and distribution of monthly runoff for control and doubled CO z conditions.
Percentile Monthly Runoff (mm)*
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Noy Dec
Minimum 0 0 {) 0 0 0 0 QO 0 0 0 0
(QO) (0) (0) (0) (0) (0) (O)} (0) (0) (0) (O) (0)
10th 0 0 0 Q 0 0 0 17 10 0 0 0
(0) (0) (0) (0) (0) (Q) (0) (1.4) (0 4) (0) (0) (0)
30th 0 0 0 0 v0 0 1,3 7.3 5.0 03 0 )
(0) (0) (0) (0) (QO) (0) (1.5) (7.2) (4.3) (0,3) (0) (0)
501h 0 0 0 0 0 0 5.0 10.8 8.7 14 0 0
(0) (0) (0) (QO) (QO) (0) (3.0) (10.6) (8.4) (L,7) (0) (D)
70th 0 0 0 9] 0 Lo 87 7 1.0 4.2 0.1 0
(05 (0) (0) (0) (0) (h4) (8.8) (IL7) (11,0) (4,3) (OU) (0)
90th 0 0 0 0 0 58 IL.6 7.4 N14 8.5 0.5 0
(O) (0) (Q) (0) (()) (6.1) (H.6) (16.0) (13.9) (8,45) (0.6) (0)
99th 1) 0 0 0 3.5 10.9 367 61,5 A().4 1.8 3.9 0.1
(Q) (0) (0) (0.1) (4.7) (108) (44.7) (63.1) (89.5) (16.9) (37) (0,2)
Maximum 0) 0 0.6 4.9 1.0 412 69.5 99,2 53.2 51.9 68 0.4
(20) (2.5) 02) 27) 5) Ol) (966) (145) (103) (82.3) (76) (1.0)
* Values in parentheses are for doubled CO; cOnditions.
1 x CO2 2 x CO2
=) oO
R R
_ Oo oOo
—E & a
£ - _
= oa ~ — fo)
€¢ 2 22280 2 : :
= = | =
oO _ tr 3! : _ = - 5 =
2 =": = ole =: s+
=e Ff). El: z =~) 2) 7 =
= =E-s & = -- =
5 a i AEE: ,
2 °e uy 8 af
a a [. RA
F JF
| core ---
MAMJJASONOD MAMJJASOND
Month Month
Fig. 2, Box plots of modelled monthly rainfall for control (L x CO ) and doubled CO, (2 x COg) conditions. [Edges of
boxes mark upper and lower quartiles, and horizontal blank line within each box depicts the median. Distance between
ihe quartiles is the interquartile range (IQR). End points of whiskers attached to boxes denote either data extremes (no
adjacent horizontal lines) or adjacent values defined by upper quartile plus 1.5 < IQR and lower quartile minus L5 x IQR.
Horizontal lines mark data points that lie beyond adjacent values. |
HYDROLOGICAL. IMPLICATIONS OF CLIMATE CHANGE FOR STH. AUSTRALIA Al
1 x CO2 2 x CO2
150
150
100
I
100
i
+ TM me
-- OR
2 |
mT CMR - - - -
11 EMINC- - 208 --a
50
50
ee)
7m Ces: -- - Sail
Monthly evapotranspiration (mm)
MAMJJASOND J
MAMJJASOND
Month Month
Fig. 3. Box plots of modelled monthly evapotranspiration for control (1 x CO ) and doubled CO, (2 x CO3) conditions.
1 x CO2 2x CO2
Oo oO
sf =
oO Oo
nu ~
E 8 8
t 3 ° ~
°o = ie) ==
Cc — =
2 * ==
© = i=] = => =
~ ive) = _ © ia
xo > = =.
= 9 eas g sg:
Ss ga 5 - zs
2 AaR= ° 7M
m4 eS Ss Se
pag? es
S | a oS pecan.
JFMAMJJASOND JFMAMJJASOND
Month Month
Fig. 4. Box plots of modelled monthly runoff for control (1 x CO ) and doubled CQ, (2 x CO) conditions.
42 GB C, BATES. 8. P CHARLES, N. R. SUMNER & P.M. FLEMING
CO, conditions, There appears us be a veneral
increase in evapotranspiration for the period from
March to November under doubled CO, condifions.
Furthermore, a review of evapotranspitation values
greater than the 75th percentile point revealed un
increase in high evapotranspivation Values oVer the
entire year, The size of the increase is quite large for
the months of January, Februsry and November
Table | lists various percentile points for simulated
monthly runoetf tor control and doubled CO,
conditions, The distributions of the runoff data are also
Ulustrated in Fig. 4. Overall. there is lithe evidence
of changes in the median and upper and lower quartiles
ot monthly runotf. However, a ts evident thai the
catremes (high runotf values) tor the doubled CO,
climate are greater than those for present day conditions
for the period trom July t October. There are also
indications of reductions im the 10th. 30th and 50th
percemile pomts for Seprember.
Fig, 5 shows an empirical quantile-quintile plot of
the annual maxima of monthly runoff. The annual
maxima for both conditions are very similar lar
maxiina below their medians (about 1) mmy. Above
the medians, the maximuy are higher tor the doubled
CO, climate than for the present day and the
difference between the series appears to grow with
invreasing percentile rank, Thus there is a suggestion
of an increase in flood risk A review of the maxima
reveuled that the extreme events were due to the marked
increases in extreme rainfalls and modest inercases in
evapotranspiration for August and September under
doubled CO, conditions.
Percentile ranks. fot 1x CO2
55050
5
ta
99
Percentile ranks tor 2% GO2
Annual maximurn runotl tor 2x CO2 (mm)
t zo 40 60 20 00
Annual maximum runot tor 1x GO2 (mm)
Fiz 5, Empirical quamtile-quintile plot of annual maxima of
monthly runoff tor conrral (| = 05) and sloubled CO,
(2 ¥ COs) conditions
Conclusions
The most prominent effeer of doubled atmospheric
concentrations. of carbon dioxide on the water yield
of the North Para River catchment, South Australia,
is a Sizeuble increase in the annual maxima of monthly
runoff. “This muy indicat an increase in flood risk.
However, there appears to be litthe impacr on the
seasonality and magnitude of the median and upper
and lower quartiles: of monthly runoff.
The wbove results highlight the benefits: of using
stochastic weather generators in assessments of the
impucl ol climate change on hydrologic systems, The
estimation of extreme hydrologieal events based oni
original and perturbed historival records are not
Teliable when the return periods of these events are
of the order of (he original record length, An impasse
arises when the return periods of interest are preuter
than the period af record. Thus the length of the
historical revord Jimits what can be suid ubout the
changes tn the tails of the distributions of hydrologic
variables that may be caused by CO, doubling, In
contrast, our approach is capable of producing climatic
and hydrologic series of arbitrary Jength. This enables
amore detniled investigation of changes in interannval
variability,
The results presented herein were obtained using one
GCM (CS/ROY), a stochastic weather generator and
a water balance model (S78). Thas the quality of the
results rely heavily on the accuracy and relevance of
these models and the implicit assumption that the long-
lerm changes in vegetation that would be associated
with CO, doubling represent a second-order effect.
Nevertheless, it can be argued that the present
generation of GCMs are fundatientally similar and that
they differ principally with respect to their
patameterisuvion of certain processes. such as cloud
formation and rainfall, Thus the use of one rather thar
a suite of GCMs should not seriously: compromise the
results of this stucly.
Acknowledgments
This Work contributes to the CSIRO Cliniate Change
Research Program and is part funded through
Australia’s National Greenhouse Research Prograin.
We are indebted to FHS. Chiew and T.A. McMahon
(Centre for Environmental Applicd Hydrology,
University of Melbourne), the Bureau of Meteorology,
and the Engineering and Water Supply Department of
South Australia for permission to use their data sets,
Special thanks are due ta the CSIRO Division of
Almosphene Research for supplying CSIRO9 GCM
dats.
HYDROLOGICAL IMPLICATIONS OF CLIMATE CHANGE FOR STH. AUSTRALIA 43
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FLANNERY, B, P. (1992) “Numerical Recipes in FORTRAN:
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67-84,
IMPLICATIONS OF CLIMATE CHANGE FOR THE
SOUTH AUSTRALIAN COASTLINE
By NICK HARVEY* & TONY BELPERIOF
Summary
Harvey, N. & Belperio, T. (1993) Implications of climate change for the South
Australian coastline. Trans. R. Soc. S. Aust. (1994) 118(1), 45-52, 31 May, 1994.
Recent attention has been focused on the effects of climatic change on sea level and
the effects of a rising sea level on coastal environments. However, the variation in
physical and geological processes which are responsible for sea level change is often
overlooked or underestimated. This paper presents recent geological and tide gauge
data from South Australia to demonstrate that neotectonic and anthropogenic
influences have resulted in a general overestimation of the current rate of sea level
rise, apart from two sites where the reverse is true. The paper concludes that all tide
gauge sites must be corrected for vertical crustal movements before any conclusions
are drawn regarding local or global sea level change. The implications of this for
South Australian coasts are that adjustments to sea level trends should be made before
any vulnerability assessments are conducted.
Key Words: sea level change, coastal environments, climate change, vertical crustal
movements, South Australia.
Tronsaciions af the Novel Sevvery af S. dead. (1994), ARG), 13°52,
IMPLICATIONS OF CLIMATE CHANGE FOR THE
SOUTLL AUSTRALIAN COASTLINE
by Nick Harvey? & Tony BELPERIOT
Suminacy
Hanvry, N, & Boiehkin, T. (1993) Implications of climate change tor the South Australian coastline, Trans,
KX See. 8. Aas, (1994) LEIB(). 45-52. 31 May, 1994,
Recent attention his been focused on the effecrs of climatic change on sea level and the elfeets of a ral sca
lew! an coastal environments. However, (he variation in physical atid geological processes which ure responsible
Tor sed level change 19 often overlooked or underestimated. This paper presents recent geologic’ vial tide gauge
dati fron South Australis to demonstrate that avotectonic and ynthropapenic influences have resulted in i general
overestimation of the current rate of sea level rise, apart from swo Sites where the reverse is (rie. The paper
voneludes that ull tide gauge sites must be corrected Jor verte crustal movements before any conclusians are
drawn regarding focal or global sca level change, The implications of this tor South Australian cousts are That
adjustinents td sea level trends should he made hetore any vulnerability tasessments are conducted
Key Worps: sea devel change, coastal environments. climate change, vertipal crustal movements, South
Australia.
Introduction
‘The recent Greenhouse debate has focused attention
on the effects of climune change on sea level (Warrick
ef al. 993) and also the effects of a rising seu level
on coastal environments (Bird 1993, Tooley &
Jelversma 1992). Tn addition, the work of the
Intergovernmental Panel on Climate Change IPCC)
has produced scientific assessments of climatic change
(Houghton ef a/. Wl. Houghton er @/, 1992), weether
with the IPCC Common Methodology for assessing
the vwinerability of coastal areas to Sea level mse (IPCC
19M). However, there has been some critigism of the
applicability of this IPCC Common Methodology to
the Australian region (Kay er ul, 1992; Woodrotle &
Mcbeuty 1993),
Belore discussing the implications of this recent
research forthe South Australian coast, itis important
to note that there isa great variation in physical and
geological provesses which are responsible for sex level
change, ‘These variations, which have been categorised
by Pugh (1993) in terms of their spatial and temporal
influence. can vary Considerably from short term wind
waves with periods of about 10 seconds and an extent
af tens Of metres, up to global changes iit sea level
relaicd tw sea floor spreading: with time periods of
hundreds of millions of years. In addition, the pricess
of redistribotion of mass over the earth resulting From
devlaciauon, addition of meltwater to the beeans and
transgression and regression over the continental
shelves, wsclf results in a variety of isostatic fesponses
af the crust to the changing louds, The resultant lack
of uniformity in global sea level change is often
overlooked or underestimated.
* Mawson Graduate Centre tor Environmental Studies,
University of Adelaide, Seuth Australia SOOS_
P South Australian Departmental Mines and Enerey, PC) Baik
15. Enstwood, S$. Aust, 5063
Perhaps the most studied geological period in terms
of sea level Change is the Quaternary (Williams er af.
1993) where the effect of climatic change has resulted
in numerous Muctuations of sea level in response to
the waxing and waning of continental ice shects.
Consequently there has been alternate flooding and
exposure ol-continental margins together willl periods
of erosion and sedimentation
The most recent time when climatic conditions and
sea level were similar to the present was during the
lase interglacial period a around 125,000 years before
present (BP) when sea level in the Australian region
was between 2 and & m higher than today (Chappell
1987). Since then, sea level Nuctuations have always
been lower chan present with evidence from the
Australian region of a law sea level of hetween 130
and 165 m lower (Chappell 1987) at 18,000 years BP,
afer which if rose at.a rate of between 6 and 12 mi
vr! prior to reaching its present level between 6000
and 7000 years BP.
ft is in the record of the last 6000 years that scientists
have focused attention on finding evidence for any long
term trends in sea level cither by direct sea level
indicators in the geological record or by analogous
paleoclimatic change evidence. Careful field studies
from many coastal localities have supported
geophysical models which indicale that subtle
differences in sea level behaviour are the norm even
for the South Australian coast, long regarded as stable
and uniform (Lambeck & Nakada 1990),
There has also been direct measurement of sea level
and detailed analysis of recent tide gauge records te
extripalate relative trends (Gornitz, 1993; Pirazzoli
1991), As noted by Bird (1993) there are humerous
lactors ulfecting rélative sea level change. Apart from
eustatic sea [evel change there iy the tectonic response
of the Jand and the isosialic response of the continental
margins relating to changing volumes of ice. water or
a6 N. HARVEY & T. BELPERIC)
sediment. In addition, human aeniyities such 4s
groundwater br hydrocarbon extructron, land
reclamation, arihicwl coastal sructires, dredging and
pumping of sediment can atteet local sea level change.
Pirazzoli (O89) sugeested that local seculur lide pauge
data are dominated by neotectonic and anthropogenic
effects, resulting in an overestimation of glubal sea
level ise hy 2 lu 3 dames when these factors are ignored
South Australian policy of coast protection
and new develupment
The current South Australian policy on const
protection amd new cuustil development was prepared
by the Coast Protection Board and endorsed by the
South Australian Government in May 91. The policy.
Which is deseribed in detail elsewhere (Coast
Prection Boaid (992). relics in part oo loca records
of coastal erosion. flooding and sea level rise bul mone
importantly has incorporated the IPCC estumiutes ul
wrcenhouse inmluoxd eustatic sca level rise. These
estimates predict a seu level rise to the year 2100 ul
approximately (65 m (range 0.33 m to 1.10 m) for a
“business as usual” scenamo (Houghton eral. WY9l).
Given these extimates the Coast Protection Board used
the “preciuinnary principle” .in preparing ity policy,
The precautionary principle which was adopted by wil
Australian governments states that “where there arc
(reals oF serious or ureversible environmental damage,
lack of full scientific eertainty should not be used as
a reuson lor postponing measures to prevent
emjronmental degradation” (ntergoverninenial
Agreement on the Efivironment 1992, para 3.5.1).
In acgordance with this principle the Coast
Protection Board has adopted the policy that any new
coastal development should be capable al being
reasonahly protected from a bm sea level rise by the
year 2100, The policy estblishes the 100 year average
return interval (ARI) water level as a standard for
coastal development in South Australia. It reconimends
thal sile and building levels should be determined by
adding {4 m to the }00 year ARID water level and
(where appropriate) making an adjustment for locahsed
subsidence or uplift, Floor levels of buildings should
hewn additional 0.25 m above this level. and buildings
should not be approved unless they are capable of being
proweted of raised (o withstand a further 0.7 m of sea
level rise (c.g. by means of a bund wall). In the case
af Mood protected sites. the calculation of the 100 year
ART design flood level must incorporule the extreme
ude (plus surge) and stormwater everts, together with
wave effceis within te development.
The policy also makes a general recommendation
for an crosiog setback distance, This: is to he based
wron 100 years OL erosion at a site, allowing for local
coaslal processes and g sea level rise OF 03 11 to the
yeur 2050, und taking account of slurm erpsion from
a series of severe storms. For majar coastal
development tt ts suggested that calculuejons are birsext
Upon 20) years of erusion.
The policy is less spevitic about the protection of
existing property although a reaffirms an earlier
Government policy not to protect private property,
Although part of the Coast Protection Bouru's duties
are lo protect the coast. most coast protection works
wre carried by local councils. Acvording tothe (991
policy statement, the Coust Protection Bowrd provides
councils with grants of up to 8U% Ob the cust of
approved coast protection works and up to the same
winount tor storm damage repairs. The issue af uns
sharing bewween State and local covernmer os currenny
an issue of debate bue the underlying question for any
protection Works is essentially a devision whether to
protect or reloeate. This is conyplicuted by the level
af public of private involvement and the relative
responsibilities i State and lucal government. bor this
reason the Coast Protection Board and local
2overnment determine these sues an & case hy case
situation,
Global sea fevel rise estiniates
A major problem in identifying the corrent rate ol
eustutic sea level change Irom tide gauge data as the
influence of neotectonie, isostatic and anthropogenic
effects, compounded by a gengraphicul biey in the
dhsteibuvon of reliable tide gauge data (Warrick 1993;
Gornitz 1993; Aubrey & Emery 1999: Woodworth
1993). These problems create uneertuinry 1
extrapolating the custatic companent ofsea level change
and bas Jed authors such as Aubrey & Emery (1995)
ro-expressed caution in attempting lo exlrapolate actual
sea level changes ftom the data. They supecst thar the
apparent post 1930 accelerated sea level cise muy be
relaled to factors other than human induced fichucs such
as a Uclayed response to climatic warming following
the Litle Ice Age, oceanographic factors, or perhaps
my nat even be statstieally significant.
Mrker authors such as Gornitz. (1993) suggest that
after extraction of long-ternt trends and data averaging,
thalatas possible to obtain w true picture of sea level
rise, Gornitz presents evidence based on 16 tide piauge
Ua studies Wy suggest that estimates of global sea level
tise Over the last 100 years has been between 0.5 and
3mm yr’, wath most estimales in the range of 1 fo
2mm yr! (Gernitz 1995),
In addition lo studies alterpting to idenuty eustatic
sea level chunges based on woulysis of tide pauwe dala,
there are alse a timber af studies on sea level rise
projections related co climate change. The kev study
his been the [PCC seu level rise predictions with uw
best estimate of a 165m rise to the year 2100
(Houghton eral. 90) upon which the South
Australian coastal policy has been based, ‘The IPCC
IMPLICATIONS OF CHANGE FOR THE §.A. COASTLINE 47
report provided a significant downwards revision of
earlier sea level rise predichons but more recent
calculations have continued to produce similar best
estimate figures cither by more qualitative expert
analysis. (0.61 m by the year 2087, Woodworth 1993),
or by detailed re-calculation (0,46 m by the year 2100,
Wigley & Raper 1993).
The South Australian coast: Secular sea level rise
Records of sea level in Australia, from the period
1897 to present, have been monitored and analysed by
the National Tidal Facility (NTF) at Flinders
University in South Australia. As for the global
situation, there is significant spatial inhomogeneity in
the secular sea level trends resulting from the myriad
435°
Thevenard
EYRE
PENINSULA
Port
Lincoln
§@ Tide gauge sites
KANGAROO |S,
Victor Harbor
SIR RICHARD
PENINSULA
SOUTH
AUSTRALIA
Fig. 1. Location of tide gauges in South Australia.
Onkaparinga R.
of factors affecting relative sea level behaviour at each
lide gauge site. Although there is a variation in the
quality of tidal records available, the NTF analysis of
lidal data from gauges with an acceptable datum
stability indicate am Australian average of
151 +018 mm ye! at the 95% confidence level
(Mitchell (991). This figure 1s in dgreemtent with global
analyses of 10-20 mm yr! (Gornitz 1993),
notwithstanding the caution expressed about the
validity of these figures (Aubrey & Emery 1993).
South Australian sea level wends based on tide gauge
data (see Fig. 1 tor location of tide gauges) have been
presented by Mitchell (1991). although there are
significant tidal records such as Port Augusta, which
have yet to be analysed by the NTF (Table 1).
139°
Port Augusta
Port Pirie
Port Adelaide A.
@ ADELAIDE
$e
>
4
<)
Lake
Alexandtina
36°
Creek
YOUNGHUSBAND
PENINSULA
|
|
|
|
|
|
|
|
|
|
Mt
e Gambier
Port Macdonell 38°
48 N HARVEY & T. BELPERIO
South Australia — Neotectemic contribution
ta sea level change
Australia is frequently. and incorrectly, quoted as
a stable continent from which absolute sca levels can
be measured. Neovtectonic inovements caused by
structural geoidal. and isostatic processes. together
with factors such as sediment compaction, affect
Australia at various spatial and temporal seales. This
complexity of underlying factors that control relative
sea level change has been demonstrated by the
reconstruction of palaeu-sea level histories from
numerous sites around Australia and South Australia.
‘The sea reached its present level around the South
Australian coast beeween 7000 and 6000 yr BP. South
Australia is in the “far field” in its response to global
devlacttion, and is affected by subtle, onguiny isostatic
adjustment of shelf and coust. This is manifested us
an apparent highstand of the 6000 yr BP shoreline (Fig.
2), the height of which varies systematically and
predictably afound the coast. In particular. the height
Upmar Spencer Guit ¢4_fim)
Por! Wakeltlald (9.0re)
Port Gueter (2 1m
Cottin Bay €O.8m)
aes
Freseal_ Sea.
PALAEO SEA LEVEL RELATIVE TO PRESENT LOW WATER DATUM (mt
4
te] | e 4 4 5 5 7 2 q In
RADIOCARBON YEARS - BP (thousand years)
Tig. 2, Paluco seu levels relative to present sew levels at Various
sites in South Australia
of the highstand mereases up the two galts. with
increasing distance from the contimental margin, Sea
level change over the past few thousand years is
dominated by this regression, which increases in
magnitude from tm or less along Eyre Peninsula to
3.0 m at the head of Gulf St Vincent and 4.5 m in
Upper Spencer Gulf, Isostatic adjustments thus visry
from O.l mm yr? to 0.8 pam yr! averaged over these
ume scules. This has caused slow bul abvinus coastal
regression, particularly at the heads of both gulfs,
Superimposed on this geographically variuble
Holocene isostatic warping are longer term teclome
inevements, Tectonic effects are most noticeable along
the South East coastal plain, between Lake Alexandrina
and Mt Gambier, where Quaternary voleanisin has
resulted mm ongoing uplift and tilting of the coastal plain,
The scale and variability of this upwarp can be
iustrated by the changing elevation of the last
imerglacial shoreline (Fig. 3). This 125,000 year olel
shoreline rises progressively southwards, from 3 i
above present sea level at Salt Creck. fo in excess ol
IK m near Port Macdonnell. Uplift rates in the Port
Macdonnell region are a minimum of 0.2 mo ye! if
averaged our aver this entire time period.
Another, under-rated effect associated with cies 15
sediment compaction and land subsidence associated
with coastal reclamation and withdrawal of
underground fluids (Bird. 1993). Such effects are local.
but are sufficiently frequently associated with harbours
and tide gauge sites as to seriously question the validity.
of global averages obtained from their secular trends
(Davis 1987; Pirazzoli 1989).
The Jocal record from Port Adelaide clearly
illustrates these effects and the inherent danger of using
lide gauge data without adequate neatectonic correction
(Belperio 1989. 1993).. Daw from tide gauges,
muingrove migration patterns and trom dated subsurfice,
strata all indicate a contemporary relutive rise in sti
level within the Port Adelaide estuary. The geographic
restricdion of these effects to the Port Adelaide regio,
together with preliminary geodetic evidence, indicute
that the apparent rise in sea tevel is a local
phenomenon, resulting chiefly. from subsidence of the
lund. Belperto (1993) concluded that up ty 1.0 m ot
surficial compaction and land lowering had occurred
in association with wetland reclamation, acid sulphate
sou development, in¢reasing urban and industrial
development and groundwater withdrawal. Highly
variable rates of land subsidence, between 18 and
10mm yr'. were estimated to be occurring over
different parts of this regiou. More significanily, some
three quarters of the secular rise of sea Jevel indicated
by the Port Adelaide and Quter Harbor tide gauge dita
could be attributed to land subsidence over the last 50
years at this location.
Corrections to tide gauge derived sca level trend
data should be made for these various neotectoniv
IMPLICATIONS GF CHANGE FOR THE S.A, COASTLINE w
contributions. Preliminary corrections have been made
for South Australian tide gauge sites incorporating
known neotectonic variations (Table 1), The tide gauge
data from Port Adelaide and Outer Hatbor have been
used in global and Australian sea level rise averages
without adequate local negtectonic correction. Many
of the world’s tide gauges are similarly biased by land
subsidence effects, indicating the importance of making,
such local neatectonic corrections to all tide gauge data
before inferring local or global sea level changes.
TABLE t. See level trends caleulated from South Australian
tide vuuve data.
preliminary
years of trend* adjusted
record mit yr trend’
min yr!
Port Adelaide (Outer Hb) 48.2 2.82 06
Port Adelaide (inner Hb) 35.1 2.2% 0.5
Port Lincoln 25.4 0.90 07
Port Macdonell 217 0.49— 0.6
Port Pirie 5I- -0,20° 0.3
Thevenard 24 114 10
Victor Harbor 238 117 Vd
*Source: (Mitchell 199), p: 355) except ? Mitchell 1993. pers,
comm.) *
this paper,
Kielnerabilicry of the South Australian coast
io seu level rise
A number of papers presented at the Australian
“Greenhouse 87 Conference” discussed the gencral
coastal impacts of a greenhouse sea level rise around
the Australian coast (sec Pearman 1985). However,
very little work has been conducted on the vulnerability
of the South Australian coast to current erosion
processes, ar on the effects of an accelerated sea level
rise. Harvey: (1993) provides data on the sensitivity of
selected South Austrahan coastal environments to
development, Fothcringham & Caton (1989) give a
broad overview of potential impucts of a greenhouse
sea level rise othe South Australian coast, and Wynne
(1989) examines implications of a sea level rise for
coastal erosion and flooding.
The South Australian coastline is approximately
4000 km long tncluding a variety of coastal landforms,
including clifted coasts, rocky outcrops, mangroyes.
mudflats, extensive sandy beaches, coastal dunes, and
a number of off-shore reefs and islands. These arc
associated with a range of high energy exposed open
ocean coasts through to the protected low energy
shorelines of the upper gulfs.
The immediate impact of any sea level rise will be
to increase the magnitude and frequency of extreme
tides and levels of storm erosion, although the effects
of this will vary greatly around the coast. The least
vulnerable areas will be the resistant rocky coasts of
the Fleurieu Peninsula, Kangaroo Island and Eyre
Peninsula but there is likely to be greater erosion on
the more predominant softer aeolianite and Teruiary
limestone rocky cousts. The actual rate of chill or blutt
retreat on these coasts will vary with factors such as
Tock resistance, structure, the presence or absence of
shore platforms or nearshore reefs, exposure to wave
action, and tidal range (Bird 1993).
On the sandy coasts which represent about half
(1900 km) of the South Australian coast are likely to
have a variable response because of differing rates of
littoral drift, onshore-offshore sediment movernent, and
sediment size variability, The high energy beaches of
.
PORT MACDONNELL
ROBE
ry, + BEACHPORT
1 12 =——* ELEVATION OF THE 125,000 YR BP SHORELINE
Ww
1 10
<
uu
ie] B .
5
7 ir
O65 iva
re .
; :
wg a
6 : ‘
a —*
ii 2 a ee _— X. /
rd 7 -
im 0
=
-3 EYRE PENINSULA SPENCER GULF FLEURIEU SOUTH EAST
GULF ST. PENINSULA
VINCENT
hig. 3, Changing elevation of the last interglacial shoreline (125000 yr BP) m South Australia.
50 N
the Younghusband and Sir Richard peninsulas for
example are barked by an extensive dune coastal barrier
system which would be yulnerable to increiwsed staqn
utlack wilh elevated sea levels. Commline retreat and the
development of dane blowouts is likely to cause a
migration of the barrier towards the Coorg, As this
eccurs underlying calercte ant back harrier muds would
hecome exposed causing variable rates of retreat, In
addition raised water levels will impact on the Coorong.
In contrast. the metropolitan sandy coast lacks the
exrensive backing dune barners of the south-east, Urban
emeraachment across the frontal dune together with
extensive protective works have necessitated a sand
replenishment program to maintain the beaches, Elevated
sea levels are likely to have greatest financial impact
Wi this ared where starm protection will need to be
upgraded tigether with an increased sand replemshnical
programme, if the metropolitan beaches are to be
maintained, To the north of metrupolilan Adelaide, tand
subsilence has already been noted for che Port Adelaide
area, Pleveted sea levels will exuverbite the rate of
relalive sea level rise culsiny inangeoves wo advanes
further inland and changes to ecological zonations of
the intér-tidal wid supra-tidal biota. Jn sume plucus
mangrove advanvé may be restricted hy wrtificial
embankments resulting in mangrove dic back-
Th the gulf retions, similar displacement of ceological
commuities such as seagrasses, mangoes and
sampires would) be pronuiunced along the low ereahent
coasts. This rapid coastal retreat would be ussogiated
with reactivation of ual swamps, localised flooding of
the coastal plains and erosion of beach ridge systents.
In-other parts of the coast. an clevated sea level is
likely to cause flooding of low lying land, enlargement
a) coastal lakes and/or connection of some lakes to the
sea, mised groundwater levels ind alteratim (o estuarine
environments. South Australia has few estuunes
ulthough there could be major implications for urban
development adjacent to estuaries such as the
Onkaparings and the Port River. In the case of the Port
River estuary. upproaimeately 25% of the nearby urban
Ucevelopmenc is currently below high water, Elsewhere,
potential inipacts an the Murny River residual esiuury
(artificially consinéined by the Construction of bartages)
may be less significant for urban areas but could have
major implications for thé operating levels of the
hurrages und iiffect (he management of the Murray
Mouth and lower Murray Lakes region (Harvey 1988),
Assessing oulnerabilirys The IPCC common merhidology
In late IS91 the Coastal Zone Management Sub-group
of the [PCC released its “Common Methouology” for
the asscssmientof vulnerability of coastal areas tn sea
level vise (IPCC W491). An advisory group compresing
the United Nations Environment Progran (UNEP) and
12 nations Initiated a series of case studies to examine
HARVEY & ‘l
HELPERICY
appropriate national response strategies, and
linplementation requirements for cousts vulnerable tu
sea level rise. The Cotmnon Methodology lor assessing
vulnerability Comprises the following basic steps:
1) Delineation of case study ured and specification of
accelerated sea level rise and climate change
boundary condilions,
2 Lnventory of study area characteristics.
3 Projection of relevant development factors,
4 Assessment of physical chinges und naturil
TESPOMscs,
5 Formulation of response strategies and assessment
of their costs und effects.
6 The assessment of the vulnerability profile and
interpretation ol results.
7 Edentifcstion of actions to develop a long tem
coastal Zane management plan_
The assessment process, using !hese steps, 1s
explained in detail together wil tables and checklists
to assist un the compilation of data in 4 consistent. manner
GPEC 1991).
The Common Methodology approwch was fnund bo
be. deficient by Kay ef ail. (1992) in thew Western
Australian case study, They sugges} thar there jire
problems in the biophysical structure of the pssessment
and also in the engineering. dommated approweh bo cost
benelit response mechanisms. They also: indicate: that
there is an essential slep between the assessinent of
impacts and the formulation of a policy response. to
reduce those impacts, In Wester Australia, Ihe policy
formulation includes inter- and jntra-yovernment liaison,
public consultation, and political consultation, eventually
leading to strategic coastal zone management, However,
the lact |hal strategic coastal engineering decysions are
part of wider State und regional planning issues, means
that preference may be given to reactive coast protection
strategies fur short tenn eroston problems rather than
long-term coustal strategies related to sea level rise. This
type of problem highlights some ol the difficulties in
developing a common approach strategy encompassing
biophysical. wdininsteative and legislative factors (Kay
chal, WAR),
Tn South Australia, Ihe PCC Common Methodology
has yet to be tested but tt is likely that butemueratic nnd
politcal problems are curtently compomided by
uncertainty with the review of the 20 year old Cpasr
Protection Act, debale twer cost sharing for coastal
management between Stile and local government,
Discussion
Appropriateness. of current South Australian poitew
Tnitial estimates of the current Tate of sea level reste
obtained by averaging tide gauge dara from around the
world have been malucing., The principle reasui is che
recognition that lind level changes need to he pemenved
IMPLICATIONS OF CHANGE FOR THE S.A. COASTLINE S|
Troum (he tide gauge data before they can be used for
this purpose. Underlying, unrceognised neuieoimic
effects renal the mails reason for the geographic
variubility in secular tide gauge trends A global seo
level cise eannut be cxpecied to be detected untel
udequute Correctivins are ade for these effects at euch
lide gauge ste. With consensus estimates of predicted
sea level rise now down to 4,65 m over the next 100
years, these neotectonic effects will be determining
(actors an overall liscu! sea level behaviour, In the gulfs,
iiny rise of sea level will be mitigated by ongoing
iwostalic upwarp of up ti O83 mm yr'. In the
South-Rash, tectonic uplift will be ua mnitigating
circumstance. In major lowes, particularly where
wetland reclamation has occurred, or excessive
groundwater withdrawal is taking place, any
greenhouse sea level rise will be exacerbated by land
subsidence effects.
In the absence of accurate geadctic or wltimetric data,
the geolpgic record can be used ta obtain fipst order
estimates of land Jevel changes. In South Australis. the
chastal record has produced some useful results that
go some way towards explaining the variability ol sea
level change documented hy tide gauges. However.
there remains & pressing need for locul and global
crustal scule geodetic control of all tide gauge sites
In addition, a geographically suitable spread of tide
fue sites is required to account for the scale and
variability of neotectunic processes. In South Australia.
these fide gauge sites yre required within the gulfs as
well as along the oceanic coastline and offstore islands
The implications of this for South Australia’s coasts!
policy are twofold, First, it bighlights the need for
accurate local sea level data which are then corrected
for neolectonic and anthropogenic factors. The
preliminary corrected sea level trend figures presented
inthis paper indicate that the current rate of sea level
ns¢ in South Australia has generally been overestimated
apart lrom two sites where the reverse is true, Second.
ihis South Australian case study provides support fer
Pirazzoh’s concerns that (he global averages lor current
custatic sea level rise may also be overestiomated. The
compounding effect of uncorrected global sea level rise
trends added to uncorrected local trends will produce
inaccuracies in the sew level trend data upon which the
policies are based.
In addition the uncertainties. Surrounding elimate
change predictions und the associated sea level response
necessitate the adoption of the precautionary principle
allowing safety margins for building levels and erosion
set-backs, Although this: may huve major cost
implications lor copstal development, it is unlikely that
a grealet precisian for climate change models and sea
Jevel response will be reached in the near furure.
However, IL ts possible to reduce stme af the
uncertainties of current sea level measurements by
correcting sea level data for neotectonic and
anthropogenic influences.
Conclusions
The dala presenred in this paper indicate the need
for taking noutectunic und anthrupovenie factors into
atcount for calculating currence rates of seu level rise.
The secular trend obtained from tide gauge data 1s only
a relative sea level trend since jhe tide gauge cannot
distinguish between a real rise of sea level and crustal
subsidence al (he site. ‘This paper indicates thal the
current rate of sea level rise based om South Australia
fide gauge dita has genecally been overestimated apart
From two sites Where the reverse is true. [11s apparent
from this paper that all tide gauge sites must be
corrected for vertical crustal movements betore any
conclusions wre drawn regarding local ur global sea
level change. The implications of tis for South
Australian coasts are that adjustments to) sea level trends
sfiould be made before any Vulnerability assessinents
are conducted. It may also be possible to revise policy:
guidelines baxed on the revised sea Jewel trend data.
References
AuARFY. BD. G & Emery. K. ©. (1993) Recent global sed
levels and land levels pp. 45-56 /a Warrick, Ro A., Barrow,
EM. & Wigley, T M_ 1, (fds) “Climate and sea evel
chunwe, observations, projections and implications”
(Cambridge University Press, Cambridge).
BeLrerto. A. P (1989) The greenhouse debate and rising s¢s
levels: geological and other fictors controlling sea ‘evel
change pp. 77-00 It Dendy, T. (Ed.) “Greenhouse "BR:
planning for climate change” (Department of Pnvironmem
and Planning, South Australia),
__ (1993) Lund subsidence und seu level rise in ihe Port
Adelaide estuary: implicubons Jor moniporing she
greenhouse effect. Australian Journal of arth Scien ic AQ
499-4AK.
Bik. B.C. F 11993) “Submerging cousts: the eects oF a
rising sea level on coastal environments” Hohn Wiley &
Sons. CRichester).
Cuastat, PROTECTION Boaniz (1992) Comsial erssivn, Hooding
and sea level rise standards and protection policy Cyrtst/irte
26.
Tavis. G. H. (1987) Land subsidence and sea level rise on
the ALAC Coust phi Ardeermental Gevliey anil Water
Science 1, 67-80,
POTHERINGHAM, D, & Caros. B (1989) Soulh Australian
coastal landforms response te a greenhnuse sea level rise
pp. 77-80 Jn Dendy. T, (Ed.| “Greenhouse ‘88> planning
tor climate change" (Deparment of Environment and
Planning, South Australias,
"ty
Gorwitz, V. (1993) Mean sea level changes in the récent past
pp. 25-44 In Warrick, R. A,, Barrow, E. M. & Wigley,
T. M. L. (Eds) “Climate and sea level change: observations,
prajections and implications” (Cambridge University Press,
Cambridge).
Haavry, N_ (1988) Coastal management issues for the mouth
of the River Murray South Australia Coastal Managemenr
16(2), 139-149,
(1993) Sensitivity of South Australian environments
la marina construction pp. 327-342, /n Thomas, D. S. G,
& Allison, R, J. (Eds) “Landscape sensitivity” John Wiley
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Houcuron, J. T.. CALtenper, B. A. & Varney, &. K,
(1992) “Climatic Change 1992. The supplementary report
to the [PCC assessment” (Cambridge University Press,
Cambridge) -
Houcuton, J. T., Jnvnins, G: J. & Epriraums, J, J, (991)
“Chmaic Change. The IPCC scientific assessment”
(Cambridge University Press, Cambridge)
IPCC (1991) “Common methodology for assessing
vulnerability to sea-level mse” prepared by the IPCC Coastal
Zone Management Subgroup (Ministry of Transport &
Public Works. The Hague, Netherlands),
Kay, R. C., Eitor. L & Kiem, G. (1992) “Analysis of the
IPCC sea-level rise vulnerability assessment methodology
using Geographe Bay SW Western Australia as a case study:
im coastal risk management” (Report to the Department of
Arts. Sports, Environment and Territories)
MiTcHete, B. (1991) Sea fevel and climate change, pp:
327-342, Proceedings of the Second Australasian
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(Australian Hydrographic Sociely, Sydney).
PEARMAN, G. 1. (1988) “Greenhouse "88: planning for climate
change” (CSIRO Division of Atmospherit Research.
Mordialloc)
Pirazzoit, P. A. (1986) Secular Trends of relative sea level
(rs!) changes indicated by tide-gauge records Journal of
Coastal Research 1, 1-26,
(1989) Present and near-future global sea level
change, Paleogeagraphy, Paleaclimatology, Paleovcolagy
(Global and Planetary Change) 75, 241-258.
2 N. HARVEY & T. BELPERIO
PuGH, D. T. (1993) Improving sea level data pp. S7-71 In
Warrick, R, A.. Barrow, E. M. & Wigley, T, M. L. (Eds)
“Climate and sea level change: observations, projections
and implications” {Cambridge University Press,
Cambridge).
‘Tootry, M. J. & JecGersma, §, (1992) “Impacts of sea-level
rise on European coastal lowlands”, (Blackwell Publishers,
Oxford, }
Warrick, R, A. 1993) Climate and sed level change: a
synthesis pp-3-21 Ja Warrick, R. A.. Barrow, E. M. &
Wigley. T. M. L. (Eds) “Climate and sea level change:
observations, projections and implications” (Cambridge
University Press, Cambndge)
. Barrow, E. M. & WicLer, T, M. L.. (Eds) (1993)
“Climate and sea level change: observations, projections
and implications” (Cambridge University Press,
Cambridge).
Wiciry, TM. L. & Rarer, S.C, B. (1993) Future changes
in global mean ternperature and sea level pp. 1135 In
Warrick. R, A.. Barrow, E, M. & Wigley, T, M, L. (Eds)
“Climate and sea level change: observations, projections
and implications” (Cambridge University Press,
Cambridge).
WiiiamMs, M.A. J., DUNKERLEY, D. L,. Dr Drexker, P.,
Kersiaw. A. Po & Stokes, T. (1993) “Quaternary
environments” (Edward Arnold, London),
Wooororre, C. D. & McLean. R. F (1993) “Cocos
(Keeling) Islands: vulnerability to seaJevel rise”
(Depariment of Environment, Sport and Territories,
Canberra),
Woopworri. P-L. (1993) Sea level changes pp. 379-391 Jn
Warrick, R. A., Barrow, E. M. & Wigley, T. M. L, (Eds)
“Climate and sea level change: observations, projections
and implications” (Cambridge University Press,
Cambridge),
Wane. AR. (1989) tnplications for coastal erosion and
flooding in South Australia pp. 77-80 Jn Dendy, T. (Rd.)
“Greenhouse ‘88: planning for climate change” (Department
of Environment and Planning, South Australia).
CLIMATIC CHANGE AND ITS IMPLICATIONS FOR THE
AMPHIBIAN FAUNA
By MICHAEL J. TYLER*
Summary
Tyler, M. J. (1994) Climatic change and its implications for the amphibian fauna.
Trans. R. Soc. S. Aust. 118(1), 53-57, 31 May, 1994.
The dependence of frogs upon moisture makes them highly sensitive to environmental
conditions. The South Australian frog fauna includes 28 species which collectively
and individually experiences a wide temporal range of temperature and available
moisture. Any climatic change involving warmer and moister conditions is likely to
enhance their distribution and abundance.
Key Words: Frogs, moisture, distribution, South Australia.
‘Transactions at te Reval Spetery of 8, Aust, (1994), WI8(1), 53-57.
CLIMATIC CHANGE AND ITS IMPLICATIONS FOR THE AMPHIBIAN FAUNA
by MICHAEL J. TYLER*
Summary
‘Tyuer, M. J. (1994) Climatic change and its implicationy for the amphibian fauna, Trans. R. Soc. 3 Aisy, HIR(L),
53-57, 31 May, 1994
The dependence of frogs upon moisture makes them highly sensitive to envirenmental conditions. The South
Australian frog fauna includes 28 species which collectively and indivieually experiences a wide temporal range
of (emperature and available moisture: Any climatic change involving warmer and moister conditions is likely
tw-enbance their distribution and abundajice
Kry Wore; Frogs, moisture, distribution, South Australia.
Introduction
‘The relevance of the study of frogs in.any cvaluation
of the impacts of climatic change hinges upon
recognition of the dependence of these animals upon
tnoisture, Nevertheless the survival of frogs throughout
the period that witnessed the entire evolution of all
other terrestrial vertebrates, demonstrates the capacity
ot frogs to survive massive environmental changes.
The complete extent of the diversity of the modern
frog fauna is unknown: Duellman (1993) estimates a
total of 3967 species ut 31.xi),91, and numerous species
have heen described subsequently. In Australia the
current total is 203, but many more await description.
Despite the success and Jongevity of frogs as an
evolutionary lineage they remain dependent upon
moisture because of the relative permeubility of the
skin and (in most species) the need to deposit eggs in
tree bodies Of water.
It follows that frogs are highly sensitive indicators
of environmental pollution and, equally, that they
eomribute an carly warning system in terms of
detecting environmental changes.
Any climatic change in South Australia at a regional
or tobi level is likely fo impact upon the distribution
of species, and upon the number within (his State,
South Australian frog fauna
Currently 28 species of frogs have been reported
from South Austria, The largest number of species
in any area (1) is to be found in the lower southeast
whith is also the area of highest rainfall. However
comparable numbers are found in the arid northwest
(9) and northeast (10) (Table 1).
The northwest and the northeast are also the portions
of South Australia where numerically and
proportionately there are the most species not shared
wilh other areas (Table 1). In each instance these
“unique” species are known within South Australia
“ Department of Zoology, University of Adelaide, South
Austraha S0QS,
from fewer than six Jocalities, and each specics is more
widely distriboted outside the State. Numerous frog
species in South Australia are at the geographic limit
of their distribution, and climatic change may have a
dramatic influence upon their persistence or
abundance.
Tanti ty Geographic characte risies af South Australian frog
fauna.
Spenes = Known from
Nomner of = confined fo ess than six
Gengraphic Areu spevigs That urea localities
Northwest 9 aa 5
Northeast 10 5 5
Flinders. Ranwes 5 | 0
Fyre Peninsula 3 0 0
Yorke Peninsula 2 0 ()
Mt Lofty Ranges 7 0] 0
Murray Valley 9 3 iF
Kangaroo Island 6 i) 0
Lower southeast iy 4 )
Nullarbor Plain 2 0 a
Impact of Rainfall
Although the amount of rainfall. and its reliability
are not the only factors influencing frog distribution,
the generality can be made that the persistence of
moisture at or near ground level is most important.
To predict faunal impact in South Australia it is
pertinent to examine a geographic area in which there
is a progressive increase or decrease in rainfall along
a Jatitudinal gradient. The one area in Australia in
which this requirement is met is the Northern Territory.
where there is 2 progressive reduction in annual raintall
from north to south (Fig. 1),
The Northern Territory model
Based upon the distribution patterns of the 42 species
then known from the Northern Territory plotted by
rytetet ate!
I2°S
“
=, o
t
Newcastle Waters
1Ige—
Tennant Creek
400
e Barrow ade
Creek
Alice 200
prings
i)
2a°— | 50
LS Se es
|
|
|
| |
| |
30°— 455
Fig. | Northern Terrttory showing latitudes, isohyets und major centres,
CLIMATIC CHANGE AND ITS IMPLICATIONS FOR THE AMPHIBIAN FAUNA
Number of Species
19)
1200 1000
Rainfall (mm)
Fig. 2. Number of species plotted against isohyets (r = 2750).
Degrees Latitude
40 42 44 46
TOO = 600 SpQ 400 agg 200 jaQioO
48
55
Tyler & Davies (1986) there is a distinct association
between rainfall and the number of species (Fig. 2)
within rainfall zones where there is 100-500 mm per
annum. At rainfalls higher than 500 mm the trend is
not apparent, but the geographic area invalves Arnhem
Land which has been largely inaccessible to collectors.
I anticipate that the totals for the more northern
latitudes will increase when zoological exploration
becomes possible.
Because of their dependence upon having access to
sources of moisture, the evolutionary capacity of body
size and mass of frogs are closely linked to the
reliability of sources of moisture. Species of small size
have a surface area to mass relationship that is fitted
to reliable sources of moisture. Conversely the capacity
to withstand xeric conditions requires that the surface
area from which water is lost is, by some means,
reduced. Effectively large, bulky frogs are best
equipped for xeric conditions.
Although the length of the body bears no fixed
relationship to body mass, it is the standard expression
of size (S-V = snout to vent length), In Australian
species there is marked sexual dimorphism : males are
always smaller than females (except for the eastern
Australian species Adelotus brevis), To examine any
geographic trends in size | have used a median
eft5
OE dS o =.» anttéi‘“COON,CO~C«CssS;”:C~‘C<‘®SSOSOOSSSSOOOM
50 52 54 56 58 60
Mean of Medians (S-V}
Fig. 3, Number of species occurring within particular latitudes (boundaries at single degree positions) plotted against mean
of medians of snout to vent length (S-V). Two linear regression lines have been plotted: one between 12° and 20°S (r = .948),
the second between 12° and 26°S (r = .975).
Si Moh
measurement, berg the midpoint between the size oF
the smallest adult male and Jatgest adult lemale, The
first of these analyses followed the standard 1°
latitudinal divisions (Fig. 3). The seeond involved
sunilar Iattudinal divisions but separated ut the
intermediate 0.59 divisions (Fig. 4). Essentially the
fosults dre siintlars sinall species are assoeiated with
high levels of quoisture, and in arid areas larger species
predaminate.
\ ar
t
* +h
4
as
“| Lt
Pa
—>—— — — + ——- or
\e Aa ae aa SE ah ~ a aS we
“Wns ot Mudiis CSV)
lig. 4, Nurober of species occurring within particular latitudes
(boundaries at half degree positions) plotled against medians
of snout to yer fength (S-V). Regression line r = -921
The overall lifestyle of frogs also is reflected along
these north-south transects of decreasing available
moisture. Io tecms of the size of the individuals there
is a shift in the predominance of small species to larger
ones associated With rainfall. Equally there ts a distinct
change in the overall Jifestyle of these same species,
such chat the tecrestrial/aquatie mode shifts from more
than S0% in the extreme north of the Nonhern
Terrilory lo zere at 21° latitude south
Exiting geoyraphic trends in South Australia
Within South Australia there is a lineet assocratinn
between the number (and diversity) of species and
geographic area, but without the latitudinal decrease
from the arid north to the moist south, Instead there
are pockets of species, cach of which has ity own
unique features.
For example any gradient ix destroyed by the
presence of a wet refuge like that of the Coongie Lukes
system in the northeast of this State — an intrusion
of anarea wholly atypical within a broad zone of low
and irregular rainfall, The Coongie Lakes frog fiuina
js particularly rich, but not permanent becuuse wt the
period of drought. lt has been suggested that the
northeast af the State presents a dynamic sinmation in
which there is contraction and expansion uf populations
accuTdiog to the availability of moisture and repeated
PYLER
transportation of species. from southern Queensland
Myler 990; Fig, 3),
The River Murray presents a further dimension
because at Constitutes a route for the introduction of
species from the east. Litoria peroni and Crinta
parinsivnifera emer the State by this micans, and are
confined to its vicinily,
The toadlet Preudophryne vectdenalis bos an
extensive distribution in Western Australia. whereas
in South Australia at has been found only at Vietory
Well in the Everard Ranges. in the northwest, The
existence of species within such restricted areas is ot
significance uy palacovlimatic interpretation, Because
of the small gcogruphic area to which it is confined,
the dependance of the species upou morsture. and the
existence of more arid conditions around the area,
can be inferred that since the (ime of dispersal to thut
site, it has not been more arid there than at present.
With the excepuion of Crinta riparia which is wholly
confined to the Flinders Ranges, every species mm South
Australia has a more extensive distribution beyond the
Ste boundaries. In fact the percentage of endemism
in South Australia is less than tn any other Australtin
State.
The implications of climatic change
The climate variables that influence the yougraphic
distribution of Australian species of frogs are very
poorly known. Brattstrom (1970) was {he first tu
demonstrate an dssociaion in muntane species berween
alitudinal location and toleran¢e to temperature
fluctuations. In particular he indicated that the species
accupying the cooler habitats. had the least capacily
to adapt to a change in thermal regime.
Untortunutely the climatic variables that have
parucular influence upon ithe spatial distribution of
frogs aré, at best, inferred rather than demonstrated.
For example, what is the limiting factor of the
distribution of the (ree Frog Literta rubella whieh
extends as far south as Wilpena Creck in the Flinders
Ranges?
This species. ranges across the entire northern hall
ol the continent, principally within the area of summer
rains (Fig. 5), A commensal species. it is clearly
capable of adapting to changing environments, Itcan
he assumed that a climatic change producing warmer
and moister conditions will enhance the geographic
ranye of L. rubella.
Perhaps the most significant influence of climatic
chunge will be the creation of more aquatic breeding
sites and the persistence of these sites for longer
periods. Together these factors will modify habitats in
a manner (hal is advantageous lo frogs. Hence the
Coongie Lakes ure likely lo become more persistent,
and the fauna there nore stable in a temprral sense
(han at present.
CLIMATIC CHANGE AND ITS IMPLICATIONS FOR THE AMPHIBIAN FAUNA 57
Fig. 5. Geographic distribution of Litorta rubella
Conclusion
Any climatic change that results in a moister and
warmer climate in South Australia is likely to enhance
the geographic distribution of the constituent species.
This can be assumed because of the wide range of
temperatures experienced already by South Australian
species, and the fact that a warmer and moist regime
will ameliorate the existing seasonally harsh
environmental conditions.
Acknowledgments
| would like to express my deep gratitude to the
University of Adelaide and the South Australian
Museum, far creating a research environment
supportive of my studies of the South Australian frog
fauna over the past 35 years. I am also indebted to Kelly
Fennel and Veronica Ward for preparing the figures.
References
Bratrstrom, B. H.. (1970) Thermal acclimation in Australian
amphibians. Comp. Biochem. Physiol. 35, 69-103.
DueLLMAN, W. E. (1993) Amphibian species of the world:
additions and corrections. Univ, Kansas Mus. Nat. Hist.
Spec. Publ. (21), 1-372.
Tyzer, M. J. (1990) Biogeography. Chap. 20, pp, 223-226.
In Tyler, M. J., Twidale, C, R., Davies, M. & Wells, C.
B. (Eds) “Natural History of the North East Deserts” (Royal
Society of South Australia, Adelaide).
& DAVIES, M. (1986) “Frogs of the Northern
Territory”, (Conservation Commission of the Northern
Territory, Alice Springs).
CLIMATE CHANGE AND ITS IMPLICATIONS FOR THE
TERRESTRIAL VERTEBRATE FAUNA
By PHILip STOTT
Summary
Stott, P. (1994) Climate change and its implications for the terrestrial vertebrate fauna
(1994). Trans. R. Soc. S. Aust. 118(1), 59-68, 31 May, 1994.
A limited number of studies of the biology of a variety of species of terrestrial
vertebrates is used to speculate about their responses to climate change as predicted
by global circulation models. Dramatic changes in the distribution and abundance of
animals in Australia that has already occurred consequent to European settlement 1s
noted. Speculation about the impact of climate change on the relative abundance of
mammals and reptiles, range changes of kangaroos, rabbits in arid areas, food security
of the Spinifex Hopping Mouse, competition between two species of skinks, and
disease transmission is included. Nest-site selection by tortoises and social structure
of foxes are given as two examples where behavioural plasticity demonstrates some
capacity to cope in situ with the effects of climate change, but the ability of most
terrestrial vertebrates to track rapid climate change across different substrates is
questioned. Reservations are expressed about the knowledge base upon which the
speculations are based. For very few species is a suite of studies available to compare
detailed data on distribution and abundance with climate over a wide geographical
range, backed up with biological information sufficient to explain the mechanisms by
which the species interact with their environments.
Key Words: Climate change, mammals, reptiles, Testudines, abundance, distribution,
rainfall, drought, disease, Macropus giganteus, Macropus fuliginosus, Oryctolagus
cuniculus.
Transquliods af the: Raval Saevery af S. Aust, (19941. UBT S9-t
CLIMATE CHANGE AND ITS [IMPLICATIONS FOR
THE TERRESTRIAL VERTEBRATE FAUNA
by PHILIP S'ror'y
Summary
Sore, P. (1994) Climate change and its implications for the terrestrial vertebrate: tama (1494), Trans. Ro Sac.
So Anst, WBC), 59-68, 31 May, 1994,
A limited number of studies of the biology of a variety of species of lerrestridl vertebrates is. used to speculate
AlbQut the responses.to climate change ws predicted by ylobal circulation madels, Qramatic changes in the distribution
und abundance of animals in Australia that has already oecurred consequent to European settlement ts nord
Speculation about the impact of climite change on the relative abundance of mammals and reptiles, range chames
of kangaroos, ribbits i arid areas, food security of the Spinifex Hopping Mouse, compention between wo species
of skinks, and disease transmission is includvd, Nest-site selection by tortoises and social siructure of loxes ure
piven as (Wr exatnples where behavioural plasticity demonsiraies some cupacity to cope it yily with the effects
of climate change, but the ability of thost terrestrial vertebrates to track rapid climate change uctosy different
substrates 18 questinned. Reservations arc expressed abvut the knowledge base upon which the speculations are
based. Por wry few spewies is asune of studies available to compare detailed data on distrihuden sind abundance
with climate over a wide geographical Tange, backcd up with biological information sufficient w capkan the
mechanisms by which the species imteract with their environments,
KEY WORDS: Climate change, manutiais repoles, Testudines, abundince, distribulion, rainfall, drowght, discise,
Macropus efyanteus, Macropas filiginesus, Orvetolagus cuniculis
(ntroduction
Some 6000 years ago, within the span of 100 years,
ihe vegetation at Cold Water Cave, Towa, changed from
forest to prairie (Dorule ef al. 1992). The changes were
assmcsated with atemperature rise of e 39°C, and with
such a complete change in yeyetauion structure must
ligve come.a profound change in the vertebrate faunis
of the locality. ILimay be, during the next century, that
similar dramatic changes. in biota will result from an
anthropogenically-cohanced “greenhouse” effect, The
terrestrial vertebrate fauna of Austritlit has alrcady
undergone profound changes as a consequence of
European settlement (Recher & Lim 1990), climate
change would compound the impacts of introduced
competitors and predators. destruction and
fragmentation of habitat, altered fire regimes, hunting.
ond diseases.
Realise predictions about the impact of climate
change on the distribution and abundance of terrestrial
verfebrate species require a foundation of information
from several sources. Essential are delinled forecasts
from climatologists; predictions about changes in the
distribution, composition, and productivity of plant
communities and other elements of food webs: and
Uetailed information about the present distribution.
ubundance, physiology, and ecology of vertebrates, ‘The
temporal scale of possible changes should also be
considered.
* Dep of Environmental Scicnee and Ruapeland
Munagement, Roseworthy Campus of the University of
Adelaide, South Australia 5005
The sentiments expressed in this paper are
complementary to those expressed in essays by Arnold
(1988), Busby (1988), Graetz (98%), Muin (1988), and
Possingham (1993). ‘The paper augments. previous
contributions by exploring, some examples of the
mechanisms by which climate change might affect
vertebrates, Tt isa speculative paper, the scenarios are
presented with. the intent of illustrating the level of
complexity, rather than making confident predictions
about the outcome. The predictions are weskened
because a single study of a local population over a short
period of time is not necessarily representative of the
biology of a species during all seasons and over the
whole of its range (see Kemper er al. 1987), und the
range and habitat of a species as observed to date da
hot necessarily represent accurately the reilised
efivironimental niche of the species, nur this in turn its
fundamental niche *Possinghar 1993), Ln addition, the
predictions of global climate models are not considered
10 be reliable at the regional level (Gordon ef al. 1992).
As the number of logical steps increases, sy the crrors
are summed,
The paper reviews those climatic predictions of
particular Importance to animals. uses some exumples
w explore the mechanisms by which climate change
might atfect terrestrial vertebrates, then examines the
capacity of animals to cope wilh the predicted changes,
Predicted climate changes
Evidence from chmute modely jodicates that
emissions of “greenhouse” gases into the atmosphere
60 P. STOTT
will cause global warming of 0.3 (range 0.2-0.5) °C
per decade, and ussociated changes in patterns of
Precipitation (Muughivn eal, 1992), Whilst the general
predictions of the models are broadly aceepted. the
issuc 1s comiplicuted by the lick wl clear observational
evidence of changes attributable with certainty. tdi
enhanced “greentoase” elect, On examining records
of meteorological observations, Nicholls & Livery
(1992) were hot able to identify aay clear treads in
raintall yt reliable meteorological stations in South
Australia up until W988. Although Burruws & Staples
991) note a warming trend in South Australia sinee
WS), they cautioned that the trend was “close te the
bounds of past experience’, Models are not consistent
in their predictions of seasonal changes in temperature
patierns for South Australia (see Boer e7 af, 1992; Gites
ef el, 1992)5 at the present state of refinement of the
general circulation models such uncertainties in timing,
magnitude, and regional palterns are well recoynisert
Mfoughum ev al 1992).
Rainfall has a major influence oo food supply for
serivbrate species, partioularly in arid arcas. Maarsina
et al, (1993) predict a global inerease in tropical
disiurbances, Ab present, some of these disturbances
extend into northern South Australia as tropical-
extratropical cloudbinds (TECBs). und us a result
heavy tain falls on an averuge of nine days per year
(Kulinel 1990), However, the distribution and catent
of the rainfall is erratic. Gordon eral. (4992) cautionsly
report the results of simulations which indicate changes
in daily rainfall intensity across Australia, particularly
in summer. in the form of inereases in the frequency
of heavy (12.8-25.6mm, 7, 31%; > 25.6 min. «, 95%)
nuifall days. ane. 15% decrease in the frequency of
ht (<4.4 nim) rpintall days, and a decrease in the
number of rain days. Whetton eral. (in press) review
the predictions of five glohal circulation models; Hour
predict inereasing summer rainfall over the whole of
South Australia (all five predict invreasing summer
rainfall over much of the far narth), and four predict
a decrease yt winter rainfall over must wt Seuth
Australis.
A weakness Of the models 1s that they do not take
into account all mayor Known influences an climate. The
El Nine Southern Oscillation (ENSO) phenomenon is
nor coupled with the major global circulation models
{including the CSIROY model) despite ms influenee on
variability of rainfall in Australia (Whetton ef al. in
press}. ENSO as ol’ particular importance do the
fectuitment Of many arid zone species [Austin &
Williams 198%; Gractz er a/, 198%). The influecee of
ENSO needs tn be considered m addition la changes
predicted co accompany the cnhanced greenhouse efteet
(Walker ere. 1989). The issue is further complicated
hectiuse ENS# iisell may be affected. HW increased
(O. Works ta equalise temperawres in the gaters vik
the custero aud western Pacific Oeean, ENSO would
he weakened (Rind 1991), thus tending to reduce
Climatic vanability ty custom Australie and countering
ft some extent the predictions outlined above
Direct effects
Direet efteots of climate on vertebrates should be
more readily elucidated than woald be the case with
indirect effects. Two examples are presented al ulirect
cttects of cliimte on reptiles. The first is the process
of lemperature-dependent sea determinition, and the
second is the influence of temperature on che
distribution of testudines (lurtles arid tortoises) Hotls
examples demonstrate that the Knowledge base is
inudequare even inthis more straighrhorward caregory.
Temperature-dependent ser determination
Temperdture-dependent sex determination (TDSR)
has been demonstrated im miny species of testudines
jeg Mrosovsky eral. 1984), croeodilians (c.g. Webb
eral 1983) an aganid (Charnier 1966 vite Bull 1980).
und a gekkonid (Wagner 1980). Slight (<2°C)
departures Tram pivotal incubation temperalures may
result in. entirely male or entirely female sexual
phenntypes, ever in some of the speaes which have
heteromorphic sex chromosomes (Servan eral MR)
Some species have Wo thresholds, Wath mates resulting
from intermediate and females frony extreme incubation
lemiperatures (Yntema 1976: Webb & Smith [984).
TDSD has been demonstrited to occur in an Australian
erocodtlian (Webb er af 983). an Australian
freshwater/estuarine testudine (Webb eral, 1986), and
Australian marine testuclines (Limpus efal/, (985). In
contrast. it does not aceur in several Australian
freshwaner testudines, including species found in South
Austraha (Georges 1938; Thompson [988a),
Preliminary studies suggest that TDSD oceurs in at
least four Australian agamids (G. Johnston pers
cornet).
The influence of environmental temperature on sex
ratios is a mechanism by which climute change can
directly affect the distribution of reptiles. Species with
TDSD would be more susceptible to rapid climate
change than species without TDSD. Wehh & Smith
(1984) noted variauon in sex ratios of Crovvelylus
Jehaston’ hatchlings in the field. and speculated that
it was due to the interuction hetween TDSD jincl
geographic differences in mean air temperatuces. Ln
a-warmer climate. the effect might be ta skew the scx
rao completely. blocking reproduction, dnd thus
leading 1 the local extinclion of the species, Such i
scenario has been advanced by Pieau 1182) as a
pssible reason for the extinction of many Mesovoie
reptiles
IMPLICATIONS UF CLIMATE CHANGE ON TERRESTRIAL VERTEBRATE FAUNA hy
Distribution af wesacdines
Testudines, which require external sources uf fical
for merabalic wetivily, are limited in latitadinal
distribution by remperucure. However, ean anoual
lemiperuture ds far rn crude & meusure te predict limits
tochsunibution. Length of the growing season has been
generally weepted as a limiting factor, since ab the
latitudinal lims ol distnbution adults may aot have
enough tine &) aeeunulate sufficient energy reserves
to Survive the winter (MacCulloch & Seeay 1983).
Alternatively, distribution may be Timited by the abal ty
of hatchlings to survive overwintering in the nest
(Breitenbach ef al. 1984; Congdon eral 1987). Obban
& Brooks (1987) suggest that a critical factur ts
leniperature durlog the matunition of nwa, whieh
requires (he accumulation of heat units wer sponge
surficient for successful reproduction. Csugzest another
possibility, tht distribution is lirmied by the probability
of ambient temperatures being suffiviently high during
The nesting seaseo to permit the energy expenditure
necessary in dieging the nest chamber. un acuity
recouniseil as being eneryetically demanding (Congdon
& Gallen 1984),
To predict the effect |hat-climate change would have
Ona species reqitives detailed Knowledye both of the
nature of the change in climate and of the mechanisms
by which climate change would affect a species Tn
testudines, we need ia know whith of the above
hypotheses is acceptable befure we know whether w
examine number of duys hetween threshold
romperatures. padir winter ternperalures, number uf
spring days above i tinperature threshold, or the
probability of oeeurrence m spring of spot tenipentlares
abiwe a thteshuld.
Indirect effects
Hffewts mediated by other lactors are inherently more
comples than direct effects. Aliempts af predicting
changes in animal distributions based on detiiled
analyses. of complex mechanisms. which include
consideration of matters such as physiology, popalanon
dynamics. interspecific interachuns, behaviours!
changes. und microhabitat conditions are fraught with
potential for error. Ao alternative 15 to Wentify a
smaller suite of influences which drive the system and
determine the end result, Nix (982) saw chmute as
jhe major determinant of the distribution of terrestrial
urgumisms. and sever! authors have used various
climatic indices ti) explain the diversity and abundance
of particular Australian hota. OF particular value are
those fare studies which compare detailed data on
Uistribution and abundance with climate aver a wide
veographical raige Both mechanistic and deterministic
explorarions hallow,
Relative abundance of manmials wane reptiles
And Australia already has a more diverse and
abundant reptile fauna than arid areas in North
Americn and Africa (Pisnka TY85), In parm. the
diversity and ubundatice of the reptiles ts altribuled to
the high sariability of rainfall which is a feature of the
Australian inlanl (Morton & Juines 1988), Proposed
changes in clinmte inight therefore be expected to lead
To un increase in the diversity and abundance of reptiles
relative to mammals.
Not all mammals would necessarily be adyersely
affected, For the large avid arcas of Austrulia, nel
ynnual productivity and hence (he carrying capacity
expressed as Lotal biomass of vertebrites iy related 10
wnnudl ruinfall (Burbidge & MeKenzie 1989).
However, the species compasition of the total biomass
is largely determined by the predictability and
distribution of the tintall, rather than ily amount.
Paichy rainfall favours birds. buts. and mobile large
mammals. such. as kangaroos (Burbidge & MeKensie
1989) which are physically capable of moving long
distances. to environments made favourable by recent
heavy rains. Irregular rainfall favuurs repriles, which
huve very low field metabolic rates relutive tO maninals
(Nagy J9X7) and can survive for long periods without
food (Morton & James 1988). A change in rsinfull
patterns to fewer days with rain, o lower probability
of light rainfall, and an increase in the frequency of
heavy raintall would not have a great impact on mobile
animals, bot would favour tepliles over small
Thammals. Small mummalian cellulose-dependene
herbivores would be particularly disadvamaged; they
ate vulnerable becduse their energy intake is limited
by their gut sive. This represents a stinilar proportion
of the size of the individual as in larger animals, but
the energy eapenditure for mainiainanee of body
lemiperature Must be relatively higher than for larger
owmmals which have a lower surface area i body
mass rato (Mortun 190). ‘Chey ace also limited by
their restricted mobility in ther ability ta explo a
patchy environment; and they ure most vulnerable to
competition from rabbits (Burbulge & MeKenzie 1989,
Morton 1990).
Ib could be argued what the balance bewwcen the
diversity of reptihan and mammalian speeses as at the
tine of European settlement was determined id same
prion more severe period of aridity (such periods ane
known from prefistoric tines- see Sinh 1981), and
therefore the balance would not be affected by a turther
increase in temperature and in the variability of ruintall.
Further, any mammals which might have been affected
are diready extinct as a consequence of European
settlement, The counter argument is, that effective
aridity in the future may be more extreme than in recent
evolutionary time. Climahe aridity (wherein increased
evaponition is in excess of increused ramfally inay be
compounded by “emuluted aridity” becouse of the
2 P STOTT
consequences of the removal of primary productivity
from the arid system in the torm of livestock und
livestock products (Burbidye & McKenzie 1989), and
the sequestration within the arid system of primary
productivity and nutrients in the tissues of livestock
and rabbits. Thus the resources available to native
vertebrates would be significantly diminished,
particularly during the resource “bottlenecks” of
droughts, and in drought refuges (see Morton 1990),
Range changes af kanguroos
The responses of kangaroos can perhaps be predicted
with a little more confidence than those of other
vertebrates. Many studies of Kangaroos have been
undertaken, including thorough studies of their
distributions (Fig. la) as part of the basis for managing
populalions which are commercially harvested.
Cauehley eral. (1987) have demonstrated that the
distribution of three species of kangaroos. is, in the
major part. determined by climate. Whilsc the
distributions are directly determined by land use and
the availability of food, water, and shelter, these
attributes are in turn greatly influenced by climate, The
distributions of the two grey kangaroos, the Eastern
Grey Kangaroo (Macropus piganters) and the Western
Grey Kangaroo (Af. fidiginasys), are closely associated
with the seasonality of rainfall; they overlap in areas
of uniform seasonality of rainfall, but M. g/peeus
occurs inareds where summer ramfall predominates,
and M. fidiginosus oceurs in areas Where winter rainfall
predominates, The Eastern Grey tolentes higher
seasonal lemperalures than the Western Grey provided
(a) Present distributions of grey kangaroos
(Density > 0,1 per square km.)
that there is summer rainfall, Both require a
heterogenous habits! with shelter being an important
component (see Hill 981, Calmes eral. 99h. The
distributional data have been used by Walker (1990)
to develop an integrated modelling and mapping system
which could be used fo predict and map changes in
distribution consequent (o climahic change, Caughley
eral, (\987) sugeest that climate change in the past
has influenced the distribution of mucropods: it is
therefore reasonable to use their conclusions to predict
the distribubonal responses of these (hree species ty
future olimate change.
If. as predicted (ubove), the winter caintall zone
contracts to the south and temperatures rise, the
distribution of the Western Grey Kangaroo would ils
contrac. to the south (Fig. 1b). Perhaps more
remarkable might be changes to the distribution of the
Eastern Grey Kangaroo, At present the species occupies
two small and widely separated areas in Soult
Austatia, but these are minor projections. into this State
oF u distribution whose western boundary runs along
or just to the cast of the State's castern borders
(Caughley ef a/, 1984). M. gigentens could extend a
considerable distance to the west of its present
distribution. and hence across the north of South
Australia (a G¢eupy suitable habitats in the northern
part of the present range of M. fidtinosus, WP verain
conditions are met. They are that summer rain becomes
more common in northern South Australia, that rehable
water is provided by increased frequency of heavy
rauitall and/or divestock water supplies, and Uiat habitat
hetergenenly persias m the face af climate ubange.
MM. /ulquaasus
M gigaritaus <
(b) Possible fulure distribulions of grey kangaroos
Fie. | (4) Present distributions of grey kangaroos, (b) Possible future distributions of grey kKanugutoos. Adapted trom Cire.
efal HY99), Cuughley & Grigg (UXT), Cauehley eral (1983), Coughtey et al (984), Short eral (YK4)-
IMPLICATIONS OF CLIMATE UHANGE ON TERRESTRIAL VERTEHRATE FAL SA fa
Seven af rabies dn eavid arent
Inurid Australia. the Rurmpean Rabbit Oryverohaga.s
Hmioudasy is deercusing the probability af sumival of
small percanial plants during droughts, qd having
profound eflect un the reeruiment of some species,
sufficient th Uite to.eluninare then from the landscape
(Lange & Grahiim 1983: Cooke 1987). The response
af lhe rabbic to climate chinge 4s therefore of particulay
WMperhinee
Historical micords show that rabbits mand ancas have
buen severely reduced i Numbers during past droughts
(Griffin & Friedel 1985), and may, under drought
conditivns, Hecume eatinct over lunge areas (Myers &
Parker 1975). Reevuilment is most unlikely) under
dmauwghl vondilions (King -Cel) W983), ama! prolonged
drwghis such as the '%9a8-64 und 1925-38 droughts
at Alice Springs (Griffin & Friedel JY85) test the
longevity of the species txee e.g. Dunsimore 1974).
Been soca few cabbits s0rvive in refiges, The quitlity
of the refuges is determined by therr ability to harvest
and store water and tutrients flowing from kirger areas
of the landscape (Morton 1990) such that run-on from
Hight ramiilhis sufficient to scoate seme plant growth
during the drought period (Ludwig 1987), At Witehitie.
South Australia, Cooke (982) noted that a sharp Gall
of Hide more than Simm of rain might be sufficent to
yield cun-ofr which, if concenthicd along drainuge
lines, would ensure that succulent fred in the fort of
chenopod stiruby would be available ta rabbits living
U1 warrens along those drainage lines. Ones heavy rin
falls (>20 on near Carnarvon, Western Australi,
Kinw ey u/. 1983). rabbits begin to breed, and ean
expand from the retuges to rmecalonise the hulk of the
landlscupe.
Hoavy roinfalls sre rare in the Australianand zone.
whole years nay pass withour a rainiall event =
12.5 yyy (Stattard Smith & Morton (990). Light falls
ure more common, EVIL IT has been predicted (soe above)
thatthe heavy falls would became more commen, ant
light falls samewhal less cunmmon, Hence, droughts
fre ikely to be shorter in duration, but the refuges
which sustain the residual rabbit population during
droaughis woul! be uy hutle less reliable. With a
comeident mse fn lenaperature exacerbating the severe
physininvied! stress experienced by rabbits under
present summer conditions (Hayward 1961), Incul
extinetion becomes mire likely during drouchts, but.
with decreased return times lor heavy rin, plagues
tight he expected Move frequently in thase areas where
rabbits survive
Vegetation changes consequent lo the inercased
yaciabiliy of preeipitadion might not faviur rabbits.
Rabhiis are found ii chenopod shrublands, but the
majority of the teed is provided by the short grasses
und forbs between the shrubs; the chenopods are eaten
only during draughts (Hall er ad, 1964; Crrittiy &
Predel ¥5), Under condininns nf inereased vlimatic
yariahilily, perennial plants would be favoured iver
ephoment! plants (Sta/tord Smith & Nurtin 90), so
ribbit populations ciuld be expected ta became less
dense unless palatable penennid] grasses such as
Themeda sp repliceal antuul grasses and forbs. Further
expleration.of thin scenaiia weld need to take accent
of a potential southera extension al the dernnanee af
Cymer C. grasses (Henderson ef af, 1h press), the
relative importance of Cy and C, grasses fo rabbits.
und dhe impticutions for rahbity of a change on the
seasonal distribution of rain towards surmmer rainfall
in uric) arews of South Australi
Fond security for Spinifer Hopping Mouse
Nuramys alexis. he Spinifes Hoppin Mouse. ts
widely distributed through sandy aneis ab werthern
South Australia, mainly in association with spanilex
grasses (Walls & Astin 1981), A major component of
ty diet as seed (Pindavson (940). and trom the
carbohydrate in seed it is able to denve sufficient
metabolic waler to Survive indefinitely; one fernale i
known fo have reured wv singhe young without
Supplementary water (Baverstock & Watts 975), Henee
the regularity of seed produchin would iluence
survivability of N. alexis in northern press af the State.
Seed production may be untMuenced by soul olsture,
iemiperature. und CO, levels. and two mechunisnos by
which A. aleviy night be advantaged are explored.
Soil moisture is one of dhe most iiportanl
climatically determined variubles for gnrisshinus
(Pittock (993) and hence lor species such as N. alevis
which depend in large nieugure on grasses (MayeMilliin
& Loe 1969). Walker eral (M89] anticipate that mean
sol moisture 1 likely to diminish in northern South
Australia, wlthough at present the reliability of
prxlictions is questioned (Vinaikov 1991, Piituck 1993).
More important for seed production are episodes of
higher soil moisture following heavier rainfalls, which
are predieted (above) to become more frequent. Thus
pulses of seed supply may become more frequent, and
support denver populations ol N. clean.
The rate of growih and the speed at whieh secu
development occurs following rains inay be
accelerated. Imai ef al, (985) bseryed increased sced
yield per plant for ree grown under enhancesl
precuhouse conditions. Gifford (979, 198K) predicted
thal wheat yields in areas with more strongly seasonal
rainfirit would increase as a result ol the enhanced
“areenhouse” effect, and that some grain growth woukl
become possible under conditions of aridity. which
currently preclude any yield. ‘The factor influencing
yseld was stimulation to plant growth by both increased
CO, and warmer wemperanires, which would resull in
asharter growing season such that the grat was more
likely to he Alling under a favourable soil moisture
regime. Enhanced efficiengy of water tse would also
occur due bo partial stomata closure (Chaves & Percin
hd BO STON'T
1992). Thus dronghts as perevived by NV. ulet/s inay
he yinehorated by (he supply of some seed when pone
would) be mherwise be available.
Australion artd-vene soils are generally intertile
(Morton (990), and nuvlent limitation May counter
growth-stimulaling tiechanisms Atihough seed
production may be hinted by the availability at
phesphorus, nitrogen. is less likely to be lintiting for
Cy plums, bor & Bkely corollary is that the protein
content of their seeds would be lower [see Conroy
W921. NV. alexry wuntld not be disadvantaged. as at is
tore Ukely to sufvive drought on low protein diets
which obviate the need to expend water ty dispose ab
wise nitrogen (MacMillan & Lee 1969).
Competition between Clenotus species
Muny of the 70 speeies in the seincid genus Crenaries
are associated with spinifex (Cogger 1992), und tt is
nol unusual for several species fo be syntepie,
suggesting fine niche separation between them, IP
Climate influences the quiche separation, climate change
may altegt the hilance between the speyies
Creins helenae and C, pantherinus are wo species
pecurring sympatrivally in the far nomh-cast of South
Australia, Pianka (1969) noted that they shared similar
niches. and suggested thar C. panthers would be
excluded by C. helenae bul for its reproductive
capacity. James (1991a) tound a high degree of dietary
rvetlap between the twa species, and noted that dictary
overlap in Crenorus was highest during the driest period
of tis stady, There is evidence to suggest! that the
separatim between these wo species i4 based an theil
thermal responses; Pianka (Y86) found C parthertates
to hyve a lawer tnean body temperature than
helenve, and James (1991b) speculated thag C!
pantherinas (and C. breoksi) can be active at winter
temperatures: which preelude activity bv CL frelenae
(and gthee Credots species), This permits ©
pantherinus and CC breoksi to begin reproduction
carlicr than the nther species. If activiny at different
icmperalures |s critical either for maintaining stable
fiche separation Gr for sustaining a mechanism of
oscillating disequilibrium between. the species, an
increase In temperature during winter may result in
compeliuve exclusion ol C pantherinus by C. helenae
in those areax where they are sympatric
Epidemlology
In-stable ecosystems, there is generally a significant
level oF accommodation between host populating and
disease-uausing agents, particularly if they Tuve cp-
evolved, However, wansmission of infectious disease
16 a dynamic process, and in many cases is dependent
onthe capacity of the infections agent to.survive outside
of the host, Helmunth parasites often have obligatory
larval stages which may survive for long periods un
the ground, and thus be susveptible to elimatic
influences. Should the climate change, the
accomodation between the host and the parasite may:
be disturbed.
Amongst the purisites af livestock there are examples
of species whose Lransmisstbility is known [a be
alteered by climate. The larvae of Haewonchay
comoedns, a gaéstri-intestinal perasile of sheep, require
inean temperatures > 18°C for normal development.
whereas the development of Oytertagia circumcincia
lurvae is suppressed above 15.59C. As both require
moisture, Ihe former isin organism of summer rainfall
sereas, aad the latter of wanter rarnfall areas (Suutheort
et ul, \976),
There is less Knawn about the parasites of Australian
native vertebrates, and most of the published
investigations have been taxnnomie (e.g Beveridge &
Durette-Desset 1992). Arundel e7 al, (1990) undertook
one of the few cpidentulogieal studies, which
demonstrated that helminth parasites can cause
considerable mortality in Eastern Grey Kangaroas, and
concluded that development of free-living larvae os
influenced by climate. In North America, the Morse
Alces alees can exist sympatrically with White-tailed
Deer Qdocoilens virginianus only in Chose areas where
circumstances de not favour persistence of infective
larvac of the meningeal worm Fétre/aphastrongvlus
tenuis (Gilbert 1992). Hence. climate change pay
indirectly influence the distributions of terrestrial
vertebrates through its effect on the probability of
Uisedse transmissien,
Mechanisms for coping with climate change
Possingham (1993) recugrised that there are three
Tiewns by which a species mikhl survive climate change
range change to trick shifting climate zones.
lulerance uf ihe change, and/or microevalutionury
change Examples are presented which demonstrate
thab toleranioe in flee form of behavioural plasticity may
counter climate change. but wacking appears
implausible for many smull terrestrial species
Tealercrive
The Red Fox (hilpes vulpes) has 4 complex sociul
structure which can be modified to cope with
environmental change, Zabel & Taggart (1989) have
demonstrated an effect by the E) Nino phenomenon
on the food supply of 2 population of foxcs.on Round
Island, Alaska, Increased water temperatures in the
Bering Sea were asseciated with widespread nesting
failure dn the seahird species which comprise most of
the summer diet of the foxes. Resorpiton und
preimplantation Joss are known to occur in pregnant
vixens (Ryan 1976), a common cause of which 4s
nutritional stress (see Mousiguard 1969). Hence, tf Ue
available foad was uniformly distrebuted amongst the:
foxes, total reproducrive failure in che fox population
IMPLICATIONS OF CLIMATE CHANGE ON TERRESTRIAL VERTEBRATE FAUNAS 6S
may well have oecurred, However, on Round Island,
diehiry Ghanges th smaller, less common. and tess
accessible seabird species were associated with changes
inthe social structure of the foxes, Polyeyny, the
reproductive mode priee to the dietary change. was
supplamed by monogamy. The male's help ts essential
lov cupliu dig and delivering prey tou lactating female
and ber tiller (Klenmiin 1977); thus the change in the
social SoucTUre meant that assistance provided by the
mile fox was focussed on fewer cubs yl a time when
i would haye beet more difficult for the miles tn
procure food, Individual reproductive success (in terms
of cubs reared to Sexual maturity) of che reduced
number a! breeding females was nim significantly
affected by the Ef Nifo phenomenon. Herve, a
emmporury) climate change which lead wm total
reproductive. failure of the two seabird species mnst
prominent inthe diet al the faxes did notin tum lead
10 fatal reproductive tajlure in the foxes.
The Easter Long-necked tormuse (Clelexdina
longicollis) has simpler behaviount patterns thant the
Rea Fox, but still has seme plasticity, It appears to
idjust Ts selection of nesting sites to lake account of
metcormdlogical pérameiers hkely to affect incubauon
temperatore. At Armidale, New South Wales. the
species digs nesting chambers an unshaded areas. whieh
invreases insolation and hence egg temperature, and
stiriens incubation (Parmenter 1976). The sume
species ut Rosewurthy, South Australia, digs about two-
(hinds of ity nesis in sites shaded for more than half
of the day (Stott 1987, 1988). Nest temperatures were
not recorded at either site, but mean daily temperatures
duting the incubation period are higher at Roseworthy
(39°C higher in January) and cloud vover js less
frequent (07 oktas less in January). Thompson (988b)
has demonstrated that unshaded arests of Benydare
macquarit at Barmera, Seuth Australia cart be 26°C
warmer than Shad mests, and attributed deaths in
some unshaded nests ta excessive heal. Thus it is
reasonable co speculuce that €” /orgicvllis. Tike the
spévies of testudines considered by Bull et al. (1982)
and Schwarzkopf & Brooks (1987), positions its pests
relative fo shade to ohtain optinmint suhsuilace
temperatures (or meubation.
tracking
Toirale ef af, (1992) dated late Hulocene vexctrlion
changes.at two sites in Towa whieh correspond (9 a
rate oF retredt Of prairie of 3041-600m per annum.
However, the anticipated fate of ambhropogenie ¢]i mate
chine is rourh ereater (han in the past (Possirtehaim
1993) Wrh the low relief of the inlind plains of
northern South Australia. mean thermal eradients ane
slight, aid a typical distance between isetberns
vorresponding to the predicted annual rare of
temperature change of 003°C is 2000 t Because
inany bird species of the Australian arid zone are
nomadic (Wiens 199], tricking climate change is
physically and behayiourally possible, However,
sedentaty species may have behavioural difficulties.
Few dati are uviilable on the dispersal capability of
small lerrestrial vertebrates, bul the longest dispersal
movement recorded by James (1991) for any individual
of tive species of Creanras was OStn, indiwating thal
unassisted dispersal 18 most uolikely to be able to track
climate change al the predicted rate,
Possingham (1993) poms out that comparisons of
dispersal cupability wilh (he mile oF climube charge may
be sunplistic. The comparison is useful to identily
species which ure physically incapable of tracking
climate, but cannet by itself determine competence.
There must he subsequent stages in the process ul
identifying spevies at isk, such as consideration of
physical barniers and juter-relutionships berween
species. Even if Crenorus spp. were physically capable
of tracking climate change, there 1s a clase and
presumably obligatory associmtion berween many
species.of Crenatus and Trindia and Plecerechne spp.
(spinitex grasses) and their attendant termites. These
ure primarily distributed in inlertile, sumdy sutls
(Gruetz et al. 1988), which indicates. that spinifex-
dependent species of Creawias which ure less tolenainr
of uvereased temperatures would be unable to track
climate change acnss changes in soil fertiltly and type,
For vertebrates in the north of South Australia. nacking
femperiture changes means iw generally southern
extension in range (With or without @ northern
contraction, a separate issue which would depend on
the upper linus of tolerance}, bur for partherinus
Uiere would be constraints because long distance
dispersal ar even local spread of Uriedia and
Plectrachin: seem ty require considerable ime (acobs
1482). Also, these grasses would not extend ima heavy
clay soils and limestone plains, the lauer being
venetally south of the present distribotion of ©
pantherinus
The steepness uf chinatic gradicits [i MOUNaINOUS
areas ws much prester than on plains, cand thus
allitudinal tracking of climate change ts much more
feasible than latitudinal tracking tor smal! berrestrial
vertebrates. Generally, a shonm chmb in altitude
curresponds to a majur shilt in latijude (Peters &
Darling 1985), Over a distance of abour |S kin in the
Adelaide Hills, a 500i increase in altitude is associated
with a fallin January mean maximum teriperatire of
about 53°C. and arise in mean annual caintall of about
AOO.mm, However, whilst the climate as one
component of a species’ environment may track upa
Mountain, other comipuacnats of the enyiroameit May:
he fixed; for example, substrate structure and fertility
usually change with altitude
To allow tracking requires the linking of areas
managed primarily for conservation dleng lativudinal
and altitudinal gradients (Nertan 1990). The review
oa P STOTT
oF nature reserves in south-eastern New South Wales
undertiken by Marpules & Stem (1989) confirnied thal
u single. long. narrow. tectapgular reserve aligned
along an altitudinal gradient would be the configuration
which would most parsinoniously meet the dominant
environmental requirements of teniperalure, rainfall.
and substrate fiir 2h canopy tree species which o¢cur
inthe region. Mackey ev al. QY88) note the necessity
ind amue the validity of using vegetation dura as
surrugates tor data on fauna habitat in. the present
cireumstnces of paucity of the preferred primary data.
und ailvecate a Iicus of ecological gradients in onder
le provide @ nara of safety in assessing arcas for
conservation value, However becatise of the low tehel
of much of South Austmlia, only latitudinal gradients
wre possthle in Mest ates.
Conclusion
The paper has tocused on 9 number of studies which
have relevance to the issie of climate change On
reviewing the topre, it is apparent that the voologival
hase from which changes may be predicted 15
imperfect. Reliahle infermaten on the present
distribution, abundance, population dynamics, and
interspecitre relationships of Australian vertebrates 1
lume (Norton 1990), but there 15 sufficient
Information to indicate that climatic influences on the
distribution of many animals Operate through
Mechumsins which are subtle and’ us yet priorly
understood, and salfigient informution to wurrnint i
conclusion that climate change of any mugnituile ts
quite likely (4 affect the distributions of many species
of terrestrial vertebrates,
Conclusiins about the fate of individual spectes an:
ut presem speculative. Deterministic studies such 25
those undertaken on Kangaroo populations ure less
speculative than mechanistic studies because of the
complexity of the means by which climate influences
the biula, dul mevhanistic ckaminutions sre
complementary in thit they muy reves critical aspects
of detail not apparent to deterministic considerations,
Acknowledgments
Thanks are extendeal to Susana Carthew why) provided
constructive eviticisnt of the manuseripy ta Megan
Lewis who prepared lor presentation and publication
the fizures on the distriburion of kangaroos, and to
Kathleen Williams who cared tor the temple of the soul
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SOME POSSIBLE EFFECTS OF CLIMATE CHANGE
ON VEGETATION
By ROBERT BOARDMAN*
Summary
Boardman, R. T. (1994) Some possible effects of climate change on vegetation.
Trans. R. Soc. S. Aust. 118(1), 69-81, 31 May, 1994.
One of the most difficult tasks for managers of land which grows long-lived plant
Species, (species that dominate ecosystems), is how to assess that change in
environmental conditions is actually of significance to the present ecosystems, and
what to do to ensure that the ecosystem, rather than the species themselves can be
maintained. Practical things to do which will make adaptation possible are limited by
the indefinite nature of time frames. Areas of the State north of Kangaroo Island
extending to the northern State boundary, are the parts most sensitive to changes in
vegetation. It is likely that the most strongly threatened species in the long-term are
the perennial species which dominate woodland formations. Low sclerophyll
woodland and mallee formations appear to be particularly at risk.
Key Words: arid zone, vegetation, climate change impact, Goyder’s line, Eucalyptus.
Transactions of the Royal Soviety if 8. Ast, (W994), UBC). 6Y-BL,
SOME POSSIBLE EFFECTS OF CLIMATE CHANGE ON VEGETATION
hy ROBER! BOARDMAN*
Summary
Busknuman, R. T. (984) Some possible effeets of cllivwte change On Vegetation, Prary. AL Se Suse DESC),
69-81, 3] Miiy, 194
One of the most difficult tasks for managers of land which geiws long-lived plant species, (species that dominate
ecosystems), iy how lo assess that charige in enviranimental conditions is actually af signifeinee 10 the presen
ecosystems, and What te do to ensure that the ceasystenn mither than the species themselves can be msintained,
Practical things to do which will make adaptation passible are limited by the indefinite nature of time frames.
Areas of the State north of Kangaroo Istand extending to the northenm Stite boundary. are the pxirts most sensitive
10 changes in vegetation, It is likely that the most stroagly threatened species im the long-term are the perennial
apovics which donate woodland formations. Low scletuphyll woodland and mallee formations appear to be
particalarly at risk. These formations occur m the zone with an average growing season, in the best possible
conditions. of fron 3,5 to 5.5 months, A large proportion of species in the seni arid and arid zones are trees
und shrubs which afe classed as “sensitive” by dendroclimatologists and dendrochronologists, The first visuil
changes to Vegetation will rosult From changes int physiological processes that are mediated (hrouph plait metabolisn.
Physiological processes of plants and microflora are more sensitive to tempeniture than ecological processes,
but this sensitivily to Wimperature change will be observed through the symptoms which will be ceslagical in
natue and reflect the impacts through interactions within and between species. It should be possible to use
dutecotogical information to help track the pathways ta new gynecological combinations of species: There is a
need and an opportunity to identify “indicator species” und particular features for Measurement. Alternative
approaches to the question of minimum water vequiccinent for a woodland without "gaps", und the extent af foliage
cover in the overnwood and whole system, are reported. The optimum cover in the overwood of 3245 % should
be udequate.to sustain low woodland on run-on sites in the 200-300 mim riinfall zone of 8.A. The evulogical
significance of Goyder’s line and its putential usefilness ave discussed. Features which permit adaplabon maliude
planting botjnigally closely-related land-races (pravenunces) und spovies, in five disnnct groups of Eucalypnes
Phat hive been matched ta three climatic mudels. The avn has been te provide a nest-egy of genes. allowine
adaptation lo n¢eur whilst retaining ceosystent structures ussociated wilh particular hind-user. Provision of gene
banks and corridors of fand able fo permil native species to niigtate, are still seen to be the best option in the iniecim
Key Worps: arid zone, vegetation. climate change impact, Goyder's line, Bucalypyits.
Introduction Climate change svenarios for the Australian region
(Climate Impact Group 1992) are now less definite than
South Austrilien vegetation is strongly Sssocisted
wilh climates. which are dominated by coo} wet winters
und waren to hot dry summers, Rainfall vod potential
evapramspiratin occur at levels which iinpart a range
from sub-humid to ard climate types, The vegeuytion
has heen classified by ils form (Specht 1975; Boonisma
& Lewis, 1976; Atlas of South Australia 1986). These.
forms shew marked correlation with isohyets and lines
of equal potential evaporation.
A large proportion of species in the semi-arid and
arid wanes ure trees and shrubs which are classed
as “sensitive” by dendroclimatologists and
dendrochronologists (Fritts 1976). Tree rings represent
the end point of the long-lived perennial plant's
ailocation of carbohydrate resources produced by
photosynthesis. This link emphasises the reliance of
semiarid and und zone vegetation on a tolerable
infrequency of favourable growmg seasons when
conditions enable species to miniain their place in
ecosystems of the region.
* Department of Primary Industries, SA, Forestry, 135
Waymonil) Strect, Adelaide, South Australia 3000
the original single emissions scenario (IPCC 1990)
fram which some of us worked in 1988 (Greenwood
& Boutdiman 1989; Boardman 1989). Global warming
models are less simplistic and now include a tange of
gases, responses in cloudiness. absorption of heat by
the oceans and a degree of climate “Sensitivity”, not
included previously. The CSIRO GCM madel has
provided plausible ranges of local temperature warming
and changes in rainlall per degree of global warming
civerall.
Sub-regions have heen added in the raintall change
scenarios which, so far as S.A. vegetation is
concemed, largely reflegt earlier findings. Arcas of the
State north of Kangaroo Island are placed in sub-region
A by Climate Change Group (1992) and this extends
most of the way to the northern heundary. The
reduction in Winter rain. despite a trend towands &
predter proportion of annual rainfall in summer
together with @ marked risé in potential
cvapotransprration has been modelled, and leads tu
greater droughliness and diminution of growing season
desenhed far this patt of (he Stale (Appendix 1)
(Boardrum 989, 1992),
™ K. BOARDMAN
Arcus of the State south of Fleuneu Peninsula are
placed in sub-region Bi this is a sub-region unlikely
la suffer murked change in gross ramtall and may not
even see a decrease in winter rainfall This fits in with
earlier progniises already deduced for this part al the
State, Linpacts on vegeuttion seem likely to be mininniil
and pissibly will see amore favourable sel of growing
conditions, Dendrometrical data on native forest m the
Lower Sruth-ease (Ruiter 1964) rndicate that the native
Iree species ire better adapted to a summer rainfall
climate than the Mediterranean type which his
persisted since the lust glacial cpoehs. That is, most
of their evolutionary adaplation luok place ina clinute
sunilar ta that now present in the castern part of NSW
(Boardinan 1986),
The impact of ENSO (El Nifto — Southern
Oscillation) Has not been mcluded in GEM models,
This uppears only to affect sub-region A of the Stabe
which Is bisected by it (Allen 1988), The boundary
between ENSO affected areas and those damunuted by
the Antarctic polar circulation systems lies along a NW-
SE direction which crosses the northern Flinders
Ranges. The “settled areas” of the State SW of the
boundary zone are not directly affected by ENSO, and
only indirectly influenced by the alternate system,
called La Nina,
[tis the time-scale of changes rather than their degree
whuch i different, compared with the early niodel. aad
it hay heen extended, A range of ternpenture change
is nOW given rather than an averuge. However the
median temperature change for the 2070 scenarios is
similar to that used in 188. and these will be fised
specifically here, Physiological processes of plants and
nucnoflor are more sensilive in temperature dhan
ecological processes, bul this Sensitivity th Len \perabure
change will be observed through the symptoms Which
will be ecological in nature and reflect ahe impacts
(through anteractions within and between species. They
will be the first changes to attract dtention We expect
to find features which can be meusured and which will
indicate the nature of changes taking place,
What is the nature of changes
that may be expected?
The sequence of reviews adopted by ihe organisers
ol this meeting imples that the most ebviows efieers
wf! climate change indiested tor South Australia will
he noticed in seals (which [think a unlikely) and in
arens We recognise as wetlands: namely, low coastlines
and coustal swamps, swales between sand-dunes, lakes
and lunettes, Most indicytions of the impact uf seasonal
changes in rainfall distribution, both in quantity and
intensity, and a Warmer elinnte incheate that it will be
topographically low-lying areas which will be affected
early during the process of change It seems reasonable
lo suggest (hat it will be the vegetation in areas (0st
sensitive to direct and intense climate change which
shows the visible signs first, rather than the animals
and (he soils. However, if may be lempting to sugeest
thot Wt will be the microflera and microfkunu which
will be must sensitive to climate change until is 1 1s
redlised that both dre more closely linked to the
dominant heher-plant specres uf an ecosystem anal the
weather, than to climate (Howard 1967: Lewin, 1985:
Simpson 1967).
It appears likely that the first yisual changes bs
vegetation will result from changes in physiological
pricesses that are mediated through plant metabolism.
In this respect, the tallest elements jf ecosystems.
especially Irees, are likely to shiw the effects {irst,
Trees are more likely to suffer climate change effects
than Shrubs, the result of thes ability wy pinduce a boke,
or trunk. As the bole lengthens and its diameter
becumes banger a grealer amount of usste feeds
sustenance which is unnecessary forlesser planis.
sheath of new Ussuc m added cach growing scasou
comprising extra phloem, or bust, bark and meristem
tissue, the cambiuin. Ibis a cast dor Keeping the tree
in a lofty posibea in the ceasystem. There ts no
difficulty with maintaining this structure in a stable
climate: allocation of resources is attuned 10 the
situation: wood erowih is laid down ita predictable
fashion including well into old age Waring (1988) and
Franklin er al. (988) have discussed the natute of
changes which may result in tree death, There ts 4
complex series of allernale pathways which lead ko a
single result. The fest of these is a reduction in food
substanee uvailable for new wood productin,
Different environmental stresses alleci the majur
components of the resource-hudgel allocation. There
is a hierarchy far normal allocation of carbohydrate
with a Ifge, Stem growih only occurs unee the resource
demands of new leaves, new roors and an intrinsically-
reguluted reserve ulloewtion into slurage (“reeavery
insunince”) has been agcommedated. Hence, the time
when physiological state (s most suitable for assessment
14 just before a Mush in new leaves becomes wppurent,
Flowers ure not produced annually antl when they de
depends on the amount of stored resources
aocunwulated. At this time carbohydrate and nutrient
reserves in (wigs, alder foliage, large dummeter routs
and the stem self are at their greatest, The urbiter
ofthis is the amount uf new woud produced per unit
of foliage
Fritts (976) bas prodiced model dixgrinis whiet
illustrate the factors which interact und lead by
formation of u narrow growth ring in the trunk. His
mudel has bye parts which can be related to the S.A,
situation, currently and as it may develop over the next
century. Pant A of his model contains relationships
associated with low peecipiration and high temperature
during the grawing seuscn that lead ta the foeniatin
Of a narrow ving in trees on Ury sues. Pin Bois a
SOME POSSIBLE EFFECTS OF CLIMATR CHANGE ON VEGETATIUN 7
possible precursory extension to plirt A. times when
low precipilation and high lemperiliire yecur rir hs
the growing seasert and auld wo the uinount al stress.
The complexity of these reactions is indicated by the
iwo parts which each have 24 possible stages. When
both paris nieer they aci additively and dimecthy on the
cumbiom, bul only combine tor the last three stages
Combination of parts A and Boas not an uncanimon
situution in SA, when dry winter presedes o draughty
growing season, The net result is reduced rates yf ect
division jn the cambium and so fewer wend cells
become differentiated, uy a conseyuence ol law
praluction of growth regulatory, substances (plant
hormones} and lood resource. Severa! other oomurwi
factors affect the cambiumn when a qarrow ring is
formed including years of intense flower and seed
production (mast years) and uttack by pests, mumly
deloliators bul also including bark beetles and sapwood
borers. Dendruchranologists have artempted ta idenmly
the range of causes when a narrow ring is formed and
some models exist for these sittations (Pritts 1976),
Palucoccolggical plan studi¢s have shown cycles of
change have alreauly been expenenced by the vetiera
und species stil] present in Australia. Species may be
adapted ts the seemirio io different ways or may be
tolerant in different degrees. A Sustained trend away
fron) normality will have an effect propertional te any
dormim evolutionary adaptation ulready incorporated
into the penes. We have rio ready way of knowing what
these mivhe be in specific cases, This position 15
agenwvatal by the appurent fag af amny centurles before
consequen| veewtatiin changes vecurred if ihe pust.
The current climate change seenurios preclude this
significant lag period apparent in the past. Ef 4 trend
in the mvc of change in climate becomes apparent ther
the siress impacts are likely ta accumulates tm witys uot
seen before. The impact iy then likely to be sunilar
to one of a range of possible impacts such ds dlustrated.
hy Barkers (1999) looking at different recurrent peyiods
herween fires in Aeacta woodland,
Periods of extended drought are no} particularly
hannful if trees have well-established mot system and
foliage canomes, partly because drought is reckoned
in relative derms 10 the normal climate. Where there
has bees natural selection by drought, trees mirely die
from draught, In the short term, adaptatvon may help
Telain reserves by temporury reduced demund, Trees,
such as those in S.A. are probably more peelimatised
to chrtmic drought than must high forest tree species.
They already have features which reduce the impact
of drought: Jow teal area indices (leaf area per unit
of ground area), smaller, thicker leaves. strap-like
petioles leading to panphotometric (drooping! lest
posture; leal mils and Wax coating, Some species
readily drop older foliage ta reduce leaf urea. The
adjustments can mean Wik woot preductton is
ewinparable with that on mare mellow sles because
respiration load is less A eonsiderable vegetation
inertia oevurs with existing, mature irec-vominated
ecusystems eeaise Irees iifluenee microclimate so
siwiuficantly, especially in the availability of sunlight
andl top-sor] moisture They can also alter the character
of spil chemistry ws they store and retranslocate scarce
NUCreNts in tissues away from recycling,
Consideration of microclimates emphasises thar che
regencration phase. when seedlings hecome
established, 15 passibly the mest eritical phase 1p the
life-cycle of higher plants The gume is one of numbers
and uf tesource reserves in the seed. Must species in
S.A. have adapted so that they produce large numbers
of seeds with small or nevlwible food reserves, 90
regeneration becomes a mater of numbers. Vege
falional change associated with climate change well
depend therefore, an the reproductive vigour of invasive
and endemic specees and whether Microelimates and
provision af soi) conditions are equally-sutted to: the
potential invader.
Persistent drought. however, falls photosynthesis.
Jeads to slepletion of varbohydrate reserves, and
depletion of defensive compounds, Individual species
wilhin a plant community will vary in the pate of
reaction, Air lurbulence and wind shear, especially of
strong. hot, dry winds. as well as chilling winds, will
agoravale drought and damuge loliages caller trees are
more at risk that shart anes. Shallew-rooted trees
growing over stony materials wllocate more resources
to rout erowth than co shoom. Even so, they are
susveptible to infrequedt intense drought anél are liable
( die off in wholé geuups. Drought will inhibit
Microbial activily amd teduee minerahsation of
nutrients from litter: lugher temperatures in tumes of
rain will accelerate mineralisation rates hut also
accelerate leaching losses — both conditions im 4
changed chmate will limit the quantity of nutrients
available. Nearly any stress. if it becomes sustained,
leads to forest und woodland decline: it reduces Icaly
canopy, photosyothelic activity, stored reserves and
delensive factors thoughout trees more so thar on
lesser vegetation. Consequently the balanve between
species und individuals will alter and. in euolopical
terms, a sefal change will be initiated,
Sciemific investigations since 1988 into impacts of
facturs in the climate change siluation on vegetation
mainly have bee conceme@ed upon the effect of elev-
ated atmosphere carbon dioxide concentrations on
physiological and metabolic processes. This hus been.
discussed frequently ds a “fertilizer effect” und evidence
inthe field as well as numerous laboratory studies have:
been made (o agluce is impact, The subject was re-
viewed in Australia by the Ecological Society and
Society of Plant Physielogists 12992, 1993; Gilfid
1993)
The impact on biodiversity has been given much less
allention, It has been reviewed by Possingham (1993)
rF RB. BOARDMAN
at the species level in terrestrial ecosystems. He took
a mechanistic approach to an implied rapidly-changing
physical environment and eoncluded that. in the short -
term, extinctions will oceur through direct interactions
of species with changes in the climate af their
environment. A second set of species face extinction
either beesuse vl uppartunities availahle to disease -
cuUsiNg species uF to loss of mutual support (ee. loxs
of shude) Or Unsustuinable symbiotie relavionships
Parucular “keystume” species ur Une associiqvon may
become extine: und so change the essence of that
system. Direct interspecific competition ts considered
ty be the beast likely cause of species loss frowe a
presenlday community, Passingham (1993) argues that
cach species Will be affected hy une. or any
combinarion ot four hasie pathways, change in
geographical position to follow favourable climatically-
controlled fealures uf (he current range; seleclion of
tolerance to vliniave change within the current 4onation.
Nicto-cvolutionary Ghinge, and extinction. He suggests
\hal ip should be possible ty use autecolumeal
information to help track the pathways to new
synecolimued! Combinations of species. This supports
the ides chat there Is 4 need.andl an oppertunity: te
identify “uidicator species” and particular fealures af
these for measirement
Suwth Anstralian vegelation
in Ube late 20th Century
The foregoing discussion has described! genenil
(rends and effects, Once une Wishes to review the local
Siluation, One has to observe its arregularites. ‘The
Flinders Ranges, for example. cause the seathern
climate patterns to extend northwards but any change
is (odified by 4 reduction in maritime cenditions
(Boardman 1992), This is reflecied, for exanyple, in
the direetion taken by Goyder’s Line between Eudundis
and Crystal Brook. When climate chunge is considered,
however, it hag to be noted that rainfall, temperatures
und particularly marine cunditioas are modified by
the topographically-high region:
Natural yewetalion in rural South Australis has beer
weected wi Jaw allitudes by the direct influence of
changes in sea levels associated with Pleismecne
glaciation. A significant pam has been affected by
marked climate changes associated with the Ice Ages.
Collectively there fuve been both eduphie and
geographic barrlers te plant immigration. Both of chese
barriers have mean| that a considerable, but
unknowable, mumber of native species has the capacity
ty wrow in S.A. 1o colomise sites and cumpete strongly
with locul species. hut these have been unuble to reach
at by Hatural means. On the other hand. the species
which are present haye suffered genetic depletion
through intense environmental selection pressures in
the last we million yeurs in their refigia_ and have
limited intrinsic adaptability io change in directions
not already encountered (Boardmun 1986).
Much of the vegetation outside the settled areas and
influenced by Aboriginal peoples. can be classed as
ancient secondary yegcwon, A une propartoen ol
the remnant vegetation, including conservution reserves
in the settled areas should be regarded as miodert
secondary ur tertiary vegetition (Barker |98Y:
Bourdmiin 1986). These last two categories were lirsi
affected by European migrauon purposefully, and
secondly, but without direction, from escapes of
introduced domestic aniinals and plants which have
become feral. Kloot (1985. 1985, 1987 ) has assessed
plant intraduetions in great detail wand has idenificd
these from plant migrants and sje pative species,
Klon| (1987) also evinced evidence (hal intense hunting,
especially on isolated land, like Kangarmo Island, very
cea Ly an the period uf sentlement by migranes, decunutd
hative grazing specics of kangaroo and Wallaby.
Iransforming the native plant ecosystems mpidly and
dramatically, Fire frequency and species life-span
interact and this affects the stability or vulnerability
of coosystems (Barker 1989), Such information is used
lo manage minge in the pastoral zone Frequent
Cevestating fires in the first half of this century, as the
population increased, and fire protection messunes in
the latter half, alse have had @ strong visible impact
on partially cleared und uncleared nalive ecosystems
io the areas south of the 32nd Paralle) (Cirandison
1983),
The studies of Kloot (1987) are a reminder that weeds
are likely to be the bane of any attempt by native species
al colomisation of new habitat, even ifthe habitat is
an amenable one, Many of the weed species iitretuced
from overseas are much more adaptable (notice Kloor's
method of separating, natural Migration from ganien
escape) and ofien they have a much more aggressive
behaviour, Many sre walural pioneer species, Two
examples of weed species likely to be affceted in the
climate change seenane and able In migrate southwards
and compete with native species needing in do the
same to survive, are Montpellier broom (Genlstr
monspessulana) and Salvation Sane (kchinnw
Plantaginium).
[tis likely that the mast strongly threatened species
in the long-term are the perennial species which
dominate woodland formations Low sclerophyll
woodland and mallee firmations appear ww be
particularly at risk. These farmatins occur in the zone
with an average growing season, in the best possible
conditions, of from 3.5 to 5.5 months. It is changes
in the distribution and remains of the physically-
dominant species in ecosystems which have signalled
past climate changes, whether ie is chrough differciat
levels of pollen deposits, whose abundance speaks fur
lisclh or (hrough macro-fossils, On the other hand,
many ofthe perennint herb and low shnih layer species
SOME POSSIBLE EFFECTS OF CLIMATE CHANGE ON VEGELATION 73
in Jocal ccosystems ane more ubiquitous than the
evosystem-dominants. and frequently less constrained
by particular site factors. This ts partly because of
similaricy i the microclimates they generale, and
punly, because Iree-duminated ecosystems provide
wiclioration of the weather From shelter and shide.
The huge demand for moisture by tees un lpcal
ecosystems means that tolerance of chron drought
hy understorey species is normal. ‘This is the case when
the understorey of the mallee woodlands is conipared
wish that Of forest ceosystens in the Mi Lofty Ranges
shuwing, that many species haye a continuity in
distribution across a wide range in rinkall from near
10250) (nintn the Murray Mallee to around 1000. mim
@ Cleland Conservation Park (McCann 1989; Dashirs
& Fessop 1990),
Investigations af woodland productivity
in the S.A. semi-arid zone in relation to climate
Trees. a3 long-lived perennial pliant spectes.
Frequently dominate all or major pars of the
ceosystems that contain ihem, They accumulate a major
part of ecosystem solid dry matter in their wood and
bark Hssues, Wood and hark ure that part of the
perrental plants which sequester CO, and which helps
to maintain stability of this gas in the aimusphere. The
Research Branch of the former Woods and Forests
Department has investigated the amounts und trends
in woud dicuntulation by wees growing in stands. li
has been rogerded on an unit area basis, either us
vile per hectare (preferred by foresters because i
is bulkiness which is of interest) or as dry weight per
hectare. The overall effect over a long time determines
productivity,
Studies in S.A, were stimulated by apprehension
ubout fossil fuel supplies, future costs of fuel, and the.
need {wv assess tenewable luel resources in the mid-70's
(Kiddle-er al, (985). This work was expanded upon
when the Flectrivity Trust of South Australia (TSA)
wixhed Iu ussess the opportunity to sequester CO,
produved frum their puwer stauons (Boardman e7 al.
1992), Productivity of twenty tree species growing in
pluntations over a range of sales. sails and climate. his
heen measured, mostly as single species, but also ay
mixtures of species. About one-third have been
remeusured Which means we have bec able to establish
trends with increasing age. The use of plantations as
useful as both the age and the density of trees in the
stand, or level of oocupancy of site are known factors,
It also means thal we Were able to extend mecthods trom
existing forestry science and use them fo extrapolate
findings over Lime and space, to other localities, We
have compared aur results with general models od foresc
productivity, to se¢ how local stands compare with their
coullerparts On a global-scale,
World plant life-zone classification (Holdridge 174)
indicates (hil sustainuble tree-duimnated ecosystems
are uilikely co existon land in S.A. receiving less than
250 mam rainfall. Consequently vur research set tui
to include all well-cstablished plantations in a zone with
average rainfall between 250 mm and 6) mim, bh the
ever, nene of the plantitions suitable for Une study
were tound in areas which received rainfall al less than
340 mm. Dryland cereal cropping, whiel) requires
220 mm of winter season taimtall la produce ai
econumic chop of obu/aere [§40L/ha|_ rarely extends
below the 280 mm isohyet (French IY8Y), However,
ETSA asked that petential for tree-growing lithe 200
— 300 mim rainfall zone should be given special
consideration. What we found is of interest in the
preseni coniteat-
Atan ecosystem level. soil moisture replenishment
and capacity lo store water ate critical factors which
determine whether trees will graw or nil, n'S.A, there
ix a surprisingly frequent inverse correlanan whiell
exists between Soil depth and fertility; a large number
of shalluw soils. which have low capacity to store
moisture and physically restrict roor development. anc
fertile and an even larger proportion of deep sulk, atle
to-absorb all the rainfall reaching the round and
provide unrestricted physical capacity for roat
development. are infertile in the topsqil where KS %
Or more of ruots are sonnally found in both cases
productivity is markedly below the level expected when
conditions ane the bese that ca be exposed,
Mallee woodland gives way to ql shrubland with
Acacia specres within the 201-300 min raintall Aue.
‘There are exceptions and an hiatus ts. present as in
western Eyre Peninsula, where it is associated soils
vonditions that are unsuited 10 either life-tormanan.
Average rainfalhas a poor delinuthay factor because it
is the critical periods and intensity of rainfall which
count the mest. The anoual rainfall in drought years
is often as Nitthe ws half that of median years. In Utis
yone, alin, infrequent long periods of drouglit persist.
These considerations have led to Lhe concep) and
definition oF “influential rainfall” The quantity uf
moisture available to prevent willing, expressed as its
equivalent in rainfall, is JOO mm und i has to be present
in the upper root zone, The factors which Jetermime
iiftuertial raanfall have been assessed for dryland cereal
cropping and sites with natural vegetation. potential ty
suitable for vereals. Influentitl rain. uo averawe is
needed for 3-4.5 months in 12. The mininwim amaune
of influential raintall ta produce 2.6 dev t/ha/y of wood
exoceds 200 iin. That os to say, in this zone sites
capable of producing wood have i receive waler im
exoess of thar received from natural rainfall directly.
In uther words, they have to be “run-on” sites whith
receive Water as run-off ar percolated as druinage trum
arcas of higher elevauion, The minimum amount of the
folal Water Which his to be available 1s hiund to be the
equivalent of 300 mint, da months when there is a tall
% RK, BOARDMAN
of tain suflicient to promote prowth (ie a surplus over
respiratory eeds!, u needs ky be supplemented by a
leas! 25% Seustinal vartation is eriti¢al for vegetation
survival if (his zone, lhe frequency distribulion al
sustained Tengths wf rainy scason for a tumber ul
representalrve weather stations has been jnvesiivsted
and is summarised in Table | (Boardiman 1980). It
emphasises that growth figs a Uees way oot be sin
imnval sequence of rings but of adequate rainievents.
Tani |, The feegrency per centiry of rain periods ef a given
rheraniant jor sites fn the 200 mm ta 300 un avertee rainfall
sone of South Australia,
Averige rinfall 300 mm 250 mini 200 mnt
Rainy scason
duration,
Al least Sinonths = (7 years 5 years | year
Atleast 4 months 44) years 17 years 5 years
Atleast 3 months 70 years 35 years 16 yeurs
Al least 2 months. 90 yours 43 years 33 years
Ac lewst J month 96 years 85 years = 45 years
‘Trumble (1937, 1939), considering dryland cereals
Tequirements, showed that 200 mm gross annual
randall will provide armual herbaceous Vegetation with
will-Free conditions for up to 3.6 months. Trees in
woodland, however, intercept-a proportion of the grass
rainfall with their wafy canopies, [f the trees are mature
and spaced so as to absorb all the capacity which is
available for them to lay dawn new wood, ie. the site
ix fully occupied by the smallest number uf trees able
to do so, then throuhfall, rain able to wet the soil.
is reduced to 160 min. This effective aceessian will
only provide for 14 months growing seasun, which
is Quite Insufficient to support woodland, At 300 mm
ratifall. fie throughfall rs likely to be sufficient for
4.5 months, An actual growing season of four months.
the minimum likely to curry a Sustamable stand ot
woodland, calls for a gross rainfall equivalera of
320 inm in Upper Fyre Peninsula and for 310 mm in
the Mutray Mallee. Following Trumble’s argument, the
“run-on" requirement fram these consideranons 1s tor
about 450 mm water, bur it muse be delivered in the
Winter ramfill season to be “influential”.
Altertative approaches to the question of minimum
water requirement fora woodland without “zaps” bas
heen provided by Specht (1973, 1975, 1983) and Walker
eral (J986, 1984}. Both have dealt with natural
*cgetation oF the kind being discussed here. Specht
worked from the direction of evaporative demand and
Uctined the tlerable lnnits to evapotzanspirslion siress.
Evergreen, perennial plant communities. under
conditions which are the best to be expected, need a
minimum net monthly supply of sail morsture
determined by rain that falls and any accession or loss
of! water in the Soil. He called this the “soil availible
wurer” requirement and it ranges fram 37.5 to
62.5 nm. Sites able to sustain woodland are
characterised by a need for additional Water Which 1s
equivalent to 27% of average rainfall for the month.
In turn, this iniplres that there is wdequale storage
available in the soil within reach of foots, Taking
canopy inlerceplinn into consideration indicales thal
Tun-on component needs to be nov less than 47% of
rainfall at a 200 mm site, again showing that abcul
300 mm af rainfall equivalent is needed to sustailp
woodland furmitions.
The stind structure of woodland which will optimise
net dry matter production from the site factors available
can he deduced from the research of Specht and
Walker. Specht has found strong relavonshipy between
the Inliage projected area ('PC) of the tallest plany
structural layer (overwood — FPCo), and of the whole
ecasysten (FPCL), to his evapotranspination coefficient,
and hasedl his findings on a very wide sample of mature
Australian nutive ecosystems. Mature low woodlanil
in the 204-300 mum rainfall zone has an overwood cover
@! 32 to 42% and total ecosystem foliuge cover of
=6%: ic. there will be about 20% bare ground.
Walkers graup took a different view of folie cover
which recognised its fragmentary structure
corresponding to about BOG of FPCo, The opumun
cover hased on their work, and adjusted to FPCo terms.
suggests that cuver uf 324+ 3% shoul be adequate tn
sustain Jaw woodland on pun-on sites in the
21H)-30) mim rainfall zone at S,A- Their models allow
one to assess stocking rates in youg-age and juvenile
Stands an well as mature ones.
Thus, the effectiveness: of water supply can be
associated with net primary production and the length
of the growing season, These have both been mapped
by for present-day climste and the median scenarios
by Greenwood & Boardman, (1989) and Boardaran.
(1992). Nee primary production is likely to change by
13% for cach incremental gain or lass of 10 mm in
average annual rainfall These preliminary findines,
nuw Itkely to be extended further abead in time, ure
sulficient to indicate the level of changes which can
be expected and where the changes are more likely to
uccur. [he maps show the “best that cun be expected”
and until the sub-optimal standing of existing vegetation
cun be assessed, they form a workiag basis Tor taking
adaptive action, Table 2 lists the tree and shrub species
whose survival and distribution Im the zones most
sensitive to change, are the ones likely 10 be influenced
by threats anid uppurtunities presented by the clinusle
change scenaris, This is based on Chippendale & Wolf
(198)): Boomsma (1981), Boomsma & Lewis (1976) axl
Costermatns (1983). Table 9 shows the cucalypt
spécies sorted inte their botanical affinities (Pryor &
Johnston, 1971) in relation to changes if ecological
slatus (hat are possible,
SOME POSSIBLE EFFECTS OF CLIMATE. CHANGE ON VEGETATION
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SOME POSSIBLE EFFECTS OF CLIMATE CHANGE ON VEGETATION
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78 R. BOARDMAN
The ecological significance of Goyder’s Line
\ The practical limits to sustained primary production
for land-use historically have been contentious and are
likely to remain so, G, W, Goyder was put on his mettle
to define such « boundary. which, all things considered,
has been largely vindicated by time. He surveyed his
line of demarcation for sustainable Jand use options
LX oad in 1865 (McGowan 1990), He rode on horseback over
3000. miles (almost 5000 km) to define it. He
associated the boundary with repeatedly observable
facts related to the southernmost extension of drought
as indicated by vegetation and soil features. These were
the “chewed remains of saltbushes and other low
4 4 shrubs" and “light souls which were susceptible (o wind
erosion” when dry, indicating stlty-sandy textures and
low contents of organic matter in the top soi, It has
been alleged that he only surveyed the Upper- and Mid-
north Regions but extrepolated his criteria ro Eyre
Peninsula and the Murray Mallee regions without
Yu’ A VAY \ | Specific attention being given to them. However, the
boundary line in the central Section, (tom near Moonta
in the west to Eudunda in the east, is only 350 miles
long. His extensive distance travelled for the survey
demanded by. the Government suggests he ventured
. <men< wl hf nfie ius i
ra 5 a us <4 y eeaR vee much further afield to justify his boundary,
zee vu 28 £ ft 2 Be7ZzR 4Ax The Line, recognised as a broad limit, has come to
Aa 2 2P27 Rk F 55555 ses] , ¢ : ,
mA an 4 HF FRAADA AAW] be associated with three features, an average Winter
. ~ 7 ‘ 4 ' iB «
raintall of 220 mm, # rainfall/potential evaporation
ratio of 0.26, (French 1989). and the southern extent
of ecosystems dominated by saltbushes species: in
particular Maireana pyramidata and M. sedifolia.
None of these features alone is adequate ty define the
sustainable land-use criteria for the 20th century. OF
particular concern here is whether its original joint-
ermeria will be assessable in time to serve in the latter
x a 2 E half of the 21st century.
ze 5 = C= 2 on ; ge . h
£3 oe: Zz Zve French (1989) showed that if the Winter (April-
= ss mes z
Bx a ° x Ge w= %! October) rainfall/evaporation ratio was to be
oo , eB we KOE FS. Bey) eo ;
$5 6 £53 3g BS Sf eg S 22] maintained, the boundary associated with Goyder's
ssp £ £7 86 5 Se mayer] : y ‘ hab
ei ee £ #§ # AS Ew .| Line would have to be located where the winter rainfall
Fe @ $2 56 £2 2 € See! ! . = E v9 !
33.65 £246 5 2 s o£ 2D; | wsohyer is currently 285 mm: ic. shift some 70 kin
_ | south of its present location on the plains of Eyre
= = =| Peninsula and the Murray Mallee, Greenwood &
Bes = =| Boardman (1989) assessed changes in growing season
a EE = =| and net primary production in native ecosystems. The
& E35 S «| northern limit of low woodland, defined as ecosystems
Ss = > el ;
4 = S$: * © | which increment new growth, under udequate
= & Sf ¢ 3 > ae is Ses contlitions, of less than 2.2 dry t/hit/y would move
PeSse § Boe 2 FES eu eS=| south by about (0Okm. These iwo independent
SEGSZ Fe tSsy, sesso lsezzt : ¥ ; 3 P
= S8Ss0etez Sees £5 SS5 x 35 =| estimates, considering the uncertainty, ave in reasonable
= =a EQ = CZSE*SESSEPRT SER LBES Ke oa aul ;
gut sa5S Ea 2G BE SS BSL S585 SS | agreement. There is likely to be a drastic impact on
ats. 25555 20 25 ges ES=f=S°0 25 =) vegemtion, To the east of the Flinders Ranges the
efsss SQSshsozosys z SSS5SSs| change indicates there would be a much more intense
Besos tert ce epgsagttss rit teywstt : : é ee . a
xR Ssgi< .= = ‘ ere = 5 H B, & fre 7 ‘ e plain: ;
<5 8§$8s5gS232acds VSsstagee gradient of change from the hills to the plains than is
presently the cage,
SOME POSSIBLE EFFECTS OF CLIMATE CHANGE ON VEGETATION cr]
Adlaptations to greenhouse-indsiced
Climate change
One of the mist difficult tisks for managers of land
which grows long-lived plant species, crees and shrubs.
espectally Where these species dommate ecosystems,
is how ty assess that change im environmental
conditions: is actually of significance to the present
ecosystems, and what do do to ensure that the
ecosystem, rather than the species themselves, can be
mamtained, Practical things to do which wilh make
adaplition possible are linmted hy the imlefinite nature
of the time frames. It is difficult to make a positive
contnbulion over such a protracted period without
efforts being conlounded hy normal variation in
weather and climate year by year. One positive
sugvesuion being widely Ginvassed is mainly applicable
to commercial tee species subject tu improvement
through hreeding. This has heen to progeny-test in
areas Which have 4 climate akin ls the changes expected
but currently outside the regions of economic viability:
The soit criteria adopted by Geyer can be
ascenaimed oi the basis Of textural classification and
degree of degradation associated with historic land-
use and the ramfall quanuily and evaporation trends
should be discerhible through use of runiing averages.
Consequently, a fulure equivalent of Goyder’s Line
could be used to highlight the progress of clinute
change. should it eventuate.
Better definition of the boundary could be achieved
with the selection of “indicator” plants, such as Mairued
pyramidifolia, indicative af change in favourable or
undavourable environmental condrions. arl gov
definition of the ecological factors which determine
their limitations or provide for stability is needed.
Ahility of indicator species to colonise sites south and
west of the current distribution should be priority
subjects for coology studies.
Ancther approach which has been adopted by
Primary Industees SA-Rorestry, that recognises forest
and woodland have multiple-uses, has been installed
(in 1992) near Gumeracha in a Demonstration Forest,
This investigation has been designed to integrate both
long term and short-term benefits. Uses included are
witer supply catchment protection, agro-forestry and
potential commercial foresury, Botanically closely-
related land-races (provenances) and species. in tive
disiinct groups of Eucalyptus similar. in fashion to Table
3. have been matched to three climatic models. Trees
are heing grown from seed collected (i) in good quality
lucal populations in $.A4. (in) fram central Victoria
where the rainfall distribution pattern ic like the
postulated change, bur the temperitures do not change,
and (iii) and populations in the central part of eastern
NSW where minfall disteibunen with a tendency to
surinee maxima and warmer temperatures occur. The
Nowering, seed set and natural regeneration phases of
ecosystems, us mentioned above, are likely to be the
MUST sensitive seral Stages to significant climale change
The Gumeracha investigation aims wa cover this
eventuality by providing genes in the dominant species
which contain adaptation to climate in the direcuons
in which it is very likely to change, The triaf is
providing a hank or nest-ege of penes to callow
adaptation to occur whalst sill likely to retain ecosystem
structure associated with land-use
A Forest Reserve al Whyte-Yarcowie. in the zone cast
of the Southern Flinders ranges. which our studies
incicate will suffer the most intense changes and
increases in stress, has been reserved tor scientitic
studies, One option we are considering is to plant a
suite of ‘indicator species, in range from those suited
well at present to others growing in regions akin to
the future climates of the scenario. Planting in a set
design at ten-year interval lor the next century would
be tuimensely valuable in an ccolugical sense. Planting
a bracket of three consecutive years at the decadal mark
would help to reduce the eflects of annual vanaton
in weather. Such a programme calls for dedication,
conscientious adherence to the plan and consistency
to be adopted by managers, and lasting, over several
generations of ecologists. [IL inay be too much toask.
A second question concerns conservation of the
current species composition of ecosystems which are
valued and preserved in Conservation and National
Parks. Greenwood d& Boyrdman (1989) reviewed the
impact of the median climate change scenarto on the
representativencss Of Parks in S.A, There has been no
significant addition to options suggested an 1988,
Provision of gene banks and corriders of land able to
permil species migration, mainly southwards is. still
seen to be the best option in the interim..In the ubsence
of physical evidence to justify the need or conservation
practices, sadly little has been added beyond revision
of the Native Vegetation Clearance Act, to suengthen
its preservation provisions, and to positively protect
roadside native vegetation. There is still a need to
include und emphasise whole-ecosystem conservation,
and to modify current preference and emphasis on rare
und endangered species.
80 R, BOARDMAN
APPENDIX L, Average annual temperature, annual rainfall and growing yeason esfimates Jar recent times and the late 2lst
Century adapting the IPCC 1991 median estimates of change in temperature and rainfall with latitude. Growing season length
has been estimated with the De Martonne Draught Index from mean monthly rainfall and teniperature data and estumates
far recent times and the late 2lst Cennry respectively {see Boardman, 1989),
Region Weathet Mean Average Growing, Season Change
Station Temperature Rainfall months in GS.
Recent Late 21C Recent Late 21C Recent Late 21C 4 of year
Eyre Pen. Cleve 16.5 19,9 404 410 5,75 25 -28
Ceduna 16.9 20,1 321 323 4.25 2.0 19
Polda 16.35 19,7 aq? 431 5.5 425 i
Kyancurla 17.0 20,3 330 340) 4.35 2.0 18
Whyalla 18.0 21.35 273 293 1,25 OS -6
Pt Lincoln 16,35 19,8 486 464 6.0 4.75 “1
Kimba 16.55 19,85 346 352 4.75 15 -28
Minnipa W2 20.45 325 323 4.) 1.0 26
Streaky Bay 17.45 20,75 378 36) 45 3.25 1
Yorke Pen. Price If 6 20,05 332 343 4.25 11 -26
Warooku 16,05 19.55 450) 433 6,0 4.5 -10
Maitland 15,95 19.35 509 50M) 6,75 §.25 -12
Kadina 16,65 20.05 346 495 5.25 2.75 2
Nowth:
Upper Hawker 17.65 20.85 301 $11 3A <0.5 27
Mid Yongala 14.4 {7.7 369. 376 5.5 1.5 -34
Bundaleer 14.8 17.1 554 544 7.0 7.0 +0
Georgetown 16,05 19.35 468 466 6.5 5.0 -12
Snawtows 16,4 19.7 407 405 6.0 3.0 -25
Lower Kapunda 15,75 19.2 496 493 6.75 6.0 -6
Roseworthy Coll, 16.35 19.8 439 443 6.5 4.5 -16
Murrayltands Milang 14.9 18.45 408 4n9 6.25 3.75 21
Nildottie 16.2 19.65 256 277 1.25 0 -10
Wanbi 15.7 19.2 307 325 3.0 <05 “2
Meningie 15.2 18.75 470 465 65 45 -lf
Wajkene 16.7 20.1 31] 342 1,75 0,75 =)
Lameroo 15.55 19.05 493 400 5.5 3.0 -21
Monarto 16.0 19.45 35] 371 4.5 0.5 -33
Mt Lolty Ranges Mt Crawford 13.15 16.6 784 755 R.0 10,0 +16
Myponga 13-6 17.05 763 734 1) 10:0 +20
Upper SE Bordertown 14.7 18,35 541 571 4.25 8.5 42
Keith 15.55 19.15 47] 474 6.75 5.0 -14
Lawer SE Konett: 13.45 17.35 713 716 20 11,75 +23
Naricoorte 14.45 18,05 586 580) 8.0 9.0 +8
Mt Burr 13.1 16.6 782 716 a5 11,0 +13
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THE ECONOMIC IMPLICATIONS OF CLIMATE CHANGE
By MICHAEL E. BURNS* & CLIFF WALSH
Summary
Burns, M. E. & Walsh, C. (1994) The economic implications of climate change.
Trans, R. Soc. S. Aust. 118(1), 83-89, 31 May, 1994.
Feedback effects from the rest of the world on the economic structure and well being
of South Australia are likely to be more significant than direct economic impacts of
climate change in South Australia or even Australia as a whole. Economists can help
to identify climate sensitive sectors of the economy and analyse how change in them
will affect other sectors. Economists can also influence anthropogenic changes by
prescribing appropriate incentive structure to redirect human actions in the common
interest. Because of the nature of the data upon which economic models are built,
their predictive capacity is restricted to short periods of 1-5 years. It is postulated that
areas such as agriculture, forestry, fishing and hunting would be highly sensitive to
climate change whilst electricity, gas and water, construction, recreational and other
personal services together with ownership of dwellings would be only moderately
sensitive. Mining, manufacturing, wholesale and retail trade, transport, storage and
communications, public administration and community services, finance, property
and business services would be negligibly sensitive to predicted climate change. The
problems associated with control of greenhouse gas emissions in an economic
framework are controversial and difficult to resolve in a global context and the
significance of collaborative approaches to these and other problems cannot be
overemphasised.
Key Words: Climate change, economics, greenhouse gas emissions, anthropogenic
influence, economic models.
Transactions af the Raval Saeiety of §. Avy (999), TB), 84-89
THE ECONOMIC IMPLICATIONS OF CLIMATE CHANGE
by MICHAEL E. Burns* & CLIFF WALSHT
Summary
Hrkns, MB, & Watart, C. 1994) The ceonotine diplicatrons of inate change. Trans, R. Soe. S. dust, UB),
83-89, 31 May, 1994
Feedback etfeeta from the rest ef the world on the economic stracture and well being of South Australia ait
likely to be more significant than direct economic impacts of climate change in South Australia or even Australia
us a whole, Economists gun help tt identify climate sensitive sectors of the economy and andlyse how change
inthem will affect other seators. Economists cun also influence anthropogenic changes by prescribing appropriate
incentive structure to redirect hurnan actions in the common interest. Because of the nature of the data upon
which econamic models are built, their predictive capacity is restricted lo short periods of 1-5 years. Irs postilated
that areas such as agriculture, forestry, fishing and hunting would be highly sensitive to climate chunze whilst
electricity, 205 and water, construction, recreational and other personal services together with ownership of dwellings
would be only moderately sensitive. Mining, manufacturing, wholesale and retail trade, transport, storage and
communications. public administwtion and community services, finance, property und business servives would
be neghipibly sensitive 10 predicted climate change. ‘Vhe problems associated with control of greenhouse gas enissiuns
inan economic framework are controversial and diffieule (0 resolve inv global context and the significance of
vollaborative approaches to these and other problems cannot be overemphasised,
KFY WORDS ctimate change, ecnnomlcs, greenhouse gas emissions, anthropogenic miluence, economic models
Introduction
An important feature of the symposiusn tor which
this paper initially was prepared was the bringing
together of at least a litte move information abou the
likely regional and local effects of clitnale change
(beyond very tentative material available from the
Greenhouse "BS Adelaide workshop and elsewhere),
The contribution economists can make doesn't depend
totally on this information, however. Like other
sciences (social us well as physical), economics can
contribute to discussion and debate through increasing
understanding and application of its way of thinking
about issues.and problems. We are not here referring
ta the commion view that economists insist on reducing
everything 10 dollars and cents. Rather, we mean the
more general contribution that economics can make
through its development of frameworks for evaluating
public policy options which:
* emphasise costs as well as benefits of policy action;
* pay particular altentivin W incenlive structures 1
policy desien;
* jecugnise how the actions of one individual,
enterprise. region pr ration impact on others; and
= uotwirlistandine the usual presumption that
econumists think that markets work perfectly,
acknowledge causes Where “the invisible hand” of the
market 1s either arthriic or non-existent, such as
When, there are spilliver effects, poorly defined
property rights. Tigh transactions costs, or the need
*
School of Koonanvics, Flindets University of South
Austrulit, GPO Box 2100, Adeluide, South Austratia 5001
- South Australian Centre for Economic Studies, The
University of Adeliide, South Australia 5005,
to impate “existence values” (eg. of life on earth)
exists, and design policies to moderate their
consequences.
Moreover, like those physical sciemists who are
deeply engaged in uttempting to understand climate
change and the role of greenhouse gases in WW, the
cconomists’ framework is general equilibrium in neture
and, to the greatest extent possible and necessary for
the problem at hand, v1 is global in its recognition of
impacts, To be more specifie, the economists’
modelling includes feedback effects between markets.
including from. world markets.
This observation leads {o our Tirst, simple, and yet
protoundly significant poimt about the economic
implications of climate change, That is, for a small
region like $.A., with # not highly unusual economic
and physical structure, feedback effects on our
economic suructure and well being from the rest of the
world are likely to be more significant than direct
economic impacts of climate change within South
Australia, or even Australia as a whole.
On the basis of modelling of various scenarios
produced in late 1980's, estimates of the loss of World
output and income (often extrapolations from specific
country studies, especially the USA) ranged trom
1%-6% of GDP, for tempeniture increases in the range
2-10 degrees Celsius. These are very large oulput and
income changes in abyolute terms, and their effects on
growth of demand for South Australian products would
be very substantial and likely to ourweigh any direct
adverse effects of climate change in South Australia
oo South Australian production and incomes.
The estimates of climate change, however, remain
highly uncensin and have tended to be progressively
8 M. EF, BURNS & ©, WALSIL
reduced in more recent years, The uncertainties about
Whetlier, when and how climate change will occur.
mhoreover, are compounded by the fact that, they are
bused on a presuniprion that policies will net change
sOon enough, or sufficiently, fo ulier outcomes,
Predicting policy responses is as Important, in the end_
its predicting climate change isclF
Cneertainties in elimate predietian:
the policy feedback issue
Others are beter qualified thin ws la explain the
nalure Gl the models heme used to genenite cline
predictions based on observed jrends in greenhovsy
yas emissions, and (he uncertainties that sucround
them. What we might usefully focus some attention
on. Wowever, 18 the iitet-related iss,ue of patiey
feedbacks,
The ceonomic projections of the consequence of
vlimare vhange that currently exist are based on it
“worst case” policy scemirle of mo significant change
16 policy in response t emerging. or anticipated
climale changes, Thrs degree of pessumisii inty be
uuwartanted, bur climate ehange certainly involves
some characteristios which give pessimistic outcomes
i higher than asoal Weightin probability distributions
delined over Jikely responses.
Policy responses to climate change have
chanicterisnes siomlar to what cconanists, in ather
chmtexts, cull public goods, For example, utthe fanuliar
case Ob a xystemt of national defence, pnee a unit OF
“prowetion” is provided for one person il 1s
Simultancausly provided for all others, and those
unwilling lo contribute to the costs cannot be excluded
from benefits, In large numbers context, individuals
see themselves as having mevliible impact on
quantities produved whether they offerte contribare
te Costs or not, and therefore the rational respoose is
hecuterupt (ofree cide, Even theugh all others muy do
likewise. and nothing is produced ws a result. it sull
Will nat pay te offer fo venerbute because whether you
door aol senchally whl ootaffeet decisions of others
Thisis a classic case of market fullure, and a case
Where government provision, funded through
votipulsary taxes, is Inevitable und desirable.
Tt should he noted, nonetheless, that even mt the
relatively simple case of defence..giving governments
“the power lo coerce” does notensure that un optupul
quantity of defence will be produced. Voting. lobby ding
und other means by which people signal their
preferences in the political market have peculiar
chuructecities. und the inventive structures facing
political wad bureauwerutie decision-makers have a
temleney to encetirage oversupply of publie sect
pounds wld services.
Clearly, polictes to control climare change bave the
same characteristics as defence: effective netion wall
benefit all nations. whether or not they contribute to
the cost (financially and/or their own paliey actions),
They also face the added complennty. of course, that
&@ vovernment does not exist that can coerce mation
stiles Co pursue policy changes, We have to rely ob
IMernational cooperation, a nolariousty rare
phenomenon. pursued through cumplex procedures,
which ts bkely (o see countries and groups holding Out
(free riding) at others” expense — even when
perceptions of the Itkely costs of datlure ty obtain
ugreement ace browdly agreed to be high. The deferred
nature of the likely costs of global warming, even yt
agreed lo be inevitable, add durther difficultics ol
getting comperition
This as not intended to be totally pessimistic — just
to feinforee what common experience (if nOL common
sense) predicts ubout global pohey change. Given
scientific uncertainties, watehful caotion abour global
action ts desirable anyway. The claysic example ul
Thoms Malthus’ (8th Century prediction thatglobil
population would grow exponentially but fond
production only arcthimeveally, and the tailed Club of
Rome predictions tn the W60s, serve as cloquent
WAPNINBs aBATNSE precipitate palicy responses, On ib
more YpuMUsGe pote, ere 1s some evidence thal tree-
rider behaviour gives Way to greater conperation when
the stakes.are ugrecd tobe very high, if only tor tec
(hat failure to participate or contribute will resule im
retaliatory action,
Ta al hkehhood, the less developed world (LDC's)
will be Jess willing to hear the cost ol policy change.
and more likely 10 stay outside any agreements, even
(hough they also are more likely (o use technologies
which are ineffieet from the viewpoint of greenhouse
gas (GG) emissions One consequence of this. ts that
(he “price” the developed world, South Australia and
Australia included, has to pay tuight be not just the
costot reducing emissions oF thew own areas, but alse
substantial resource Trusters to LDCS to buy their
purtipation. Those eosts are likely te be high even
withoul Irinsfers to LDC's: it has been suyeested. fo
ckample. hit paducing areenhouse gas emissions by
the onder proposed under the Commonweulth’s O90
imerim planning. target would be cost 3% ob
Austria's GDP (IC 199).
Despite the lack of uceurate and detailed predictions
ubout Climate chanee, about which elements are due
lu human actions, and in what ways policy-makers
world-wide are likely to respond, there arc still a
number of wis in which economisis can usefully
contribute 1 the debate. For one thing, they can help
tu Wenuly climate sensitive sectors of the economy
und analyse hen clowes in them will alleen wiles
Sectors
THE ECONOMIC IMPLICATIONS OF CLIMATE CHANGT. %
Tanie | Breakdown of Sow Australian wross state product and emplovment by sensuivity to climaie change, 199U-Y!
GSP Employment
Value % of No.
Seclor $m total (000s) %
Stas ‘Toal 25,214 636
High Sensitivity 3 7
Agriculture, torestry, fishing and hunting S34 ' 43 7
Moderate sensitivity 24 i}
Elugtrienty, gas and water ASS A 5.9 !
Construction 208 & 37.1 if
Recreation. persanal and orher scrvives } 00S 4 22.8
Ownership of dwellings 2.546 9 nha
Nepligible sensitivity 73 83
Mining 755 a 3.6 |
Manufacturing 4.746 19 105.6 7
Wholesale and retail trade 3,571 14 135-8 |
Transport. storage and conmmunivation i875 s ASS 0
Public Administration and Cammiuhity Services 5.258 21 1S1.2 28
Finange, property and business services 1972 8 62.1 10
Identifying sensitive areas ancl
relevant Now-an effects
Economists have their own range of “global models”
wimed at capturing, the interactions between markets
within und across counties. As in other areas’ of
niedelling. there are well-understood limitations
associated with the use of models, For one thing, for
operaltonal reasons (iractability of solutions, etc.),
there are trade-offs between the Jevel of disaggregation
und detail, the richness of the dynamic specification
and feedback mechanisms, and the complexity of the
functional forms. use fo describe decision-making
behaviour, For another, not only are data limited, but
laboratory experimentation and repheation of
experiments are not yenerally possible, Moreover,
because of technological, climatic, political and
population chanyes amd so on, many of the parameters
of an ceonomic modg¢l alse will change over time:
depending On the yarjables in question, useful
predictions may only be made for the very short-run
(I-S years ahead).
Subject io the above reservations, however, uw is
possible both to identify qualitatively those parts of
the economy (and of other countries economies) which
ate most sensitive lo climatic change and to discuss
generally how changes in the affected sectors might
flow-o0 through the economy. as a whole, A siniple
approach, illustrated in. ‘Table 1 is lo vonsider a
breakdown of the economic activity of a region such
as South Australia inte the output and employment
contribution of different andustry sectors, noting the
sensiuivily of cach sector to clirnatie change. (We follow
the classification of sectors. adopted by TC 199),
With more detailed climate scenarios for South
Australia. beyond those yet able to be provided, we can
further refine this Classification as well as get ito more
specifics. For example, we could track the impact on
other sectors which supply inputs ta agriculture of a
climate-induced change in agricultural output through
the use of imput-output (or, preferably, computable
general equilibnum) models. It (5 important to note that
the relevant analysis must ke into account the global
economy. To illustrate this, while agriculture may be
hurt. (it may also become mere productive) in Australia,
becuuse it is hurt elsewhere such as China, India or the
USA, Australian exports may become more valuable and
Austrahians beeome better off, even though total
Australian production is reduced. By the same token,
however, if other countnes become poorer, the value
of our exports could diminish. Sorting out these issues
elearly is vital to understanding the consequences. of
climate change for us, and globally (since we assume
South Australians have the well-being of otters as part
of their concerns).
The need to take into account the global international
(ride interactions has already been the subject of analysis
and research by ctonomists in North America and
Europe. At the most recent mectings of the prestigious
American Economic Association, an entire session was
given aver to Greenhouse Warming considerations. At
this session. Reilly & Hohmann (1993), explored the
agricultural effect of climatic change and the worldwide
trade implications of a range of scenarios. Despite the
global nature of the approach adopted, the authors’ work
contributes significantly to the identifying of key impacts
and flow-on effects for South Australia.
The analysis by Reilly & Hohmann, through its
analysis of the interactions of global regions (North
86 M
America. European Conununity, former Soviet Union,
northern Europe. China, Japan, Australia, Argentina,
Brazil and “Rest of World"), gives powertul Ulustration
of the complexity of climatic change elfeets upon an
individual region. Thus. Argentina was predicted to
benefit overall. even though no increase in agricultural
production was expected in that country and their
domestic agricultural prices were expected to increase
(in line with international prices).
Evaluate particular scenarios
As was done by CSIRO (988 and revised 1992)
spewihe (and moderately plausible) chinate change
seenurios can be considered. The mujor characterisucs
of these scenarios for the Australian region aré set oul
in Table 2.
Using the most recent of these. a group of economics
graduates (Hutchins et al. 1993) ina project tor South
Austailin Department of Environment und Land
Management, estimated a loss of 20% in volume and
value of South Australia’s wheat production, The
conclusion regarding value, must be treated with a great
deal of caution, however, bearing in mind our earlier
discussion of the importance of taking into account
global trade interactions.
The results of Such exercises have led to two further
tentative explorations. First, there has been use of
general equilibrium economic models, such as
ORANI, to examine the consequences of teduced
wheat yields, increased forestry and fishery
productivity and so on. Second, there haye been
evaluations of some of the more extreme sources of
costs due to flooding, droughts and cyclones (an
eviluation of costs, of course, opens upon
considerations of costs of “abatement’).
B, BURNS. & C. WALSH
The first af these approaches, embedding the
predicted etfects of climatic change into supply shocks
in the agricultural sector of (national aod global)
economic models, has been widely used. Comparison
of the results GF such different research programs is
difficult. however, not least because the continuing
uncertainty’ about the direction and magnitude of
climatic change has resulted in quite different scenarios
heing considered froni ane study to another.
As an example, the previously cited work of Reilly
& Hohmann draws on the climate change forecasts
summarised in Houghton et al, 1990, The weneral
scenario. which involyes temperature increases of 2°C'
in the ipics and in the range 4°C to 12°C for the polar
regions. has agricultural productivity increasing iathe
North of the (former) Soviet Union, Canada anil
Europe, ‘buf reducing in the United States und most
of the rest of Europe due to drought (see Tobey e7 ul
1992).
In addition 10 incorporating economic feedback
mechanisms and imternational trade linkages, some
attempt has also been made to extend the interface
beiween the economy and the environment hy
embodying possible agricultural sector responses,
Questions of “idaptation potential” and “adaptation
capability” are considered to bear. respectively, upon
crop substitutions that ave poteutially available and
desirable due to climatic change, and upon the
constraints to these substinitions dae to poorly
developed tharkely lor crop inputs and a Tange of
Infrastructure considerations. including the skill level
of the agricultural labour force.
TaRLe 2, CSIRO climate change sconarins [987 anid 1992. Scenaries fiir xear 2030, Sources: Pearman (1988) une Climaiy
Chanue Group (1992).
JORT
1992
TEMPERATURE —210 4°
RAINFALL —50%% in north
-Z0% to south
RAINTALL INTENSITY 20-30% Meteuse
SEA LEVEL. +20 ty 40cm
TROPICAL CYCLONES
Extend 200-400 kia further south
+0).5 to 259C
Suminec O tar «209
Winter SA 0 to 104
St Alist -10 to 110%
General increase
—S ie 35 em
Uncertain
Frequeney increyxe 30-60%
SNOW LINE
WIND SPEEDS
Rise LOU m per L°C warming
Decrease 20% North of 3698
Increase south of 36°S
Rise 100 m per IC warming
Strengthening of monsoon westerlies
in north Aust. & the SE trade winds
in summer.
THE ECONOMIC IMPLICATIONS OF OLIMATE CRANGE Kr
Insights tote the occurrence of
snthrapopene cluanpes
None of the anilyses vonsidened so far distinguishes
between changes ue te human achons andl changes
which have other causes. Ib appears to be true that,
by vecivent Or otherwise, human impacts on climile,
svt ur Gind for some time ahead), may have made things
better tor humans than would otherwise huyve heen the
chse, There is also the face that, to some extent, the
living workd adapts lo new condilions.
‘These possibilities apart, ibis well understoral hy
ceoroviists Hhat malividiiels will sometimes (pack on
their environment in way thal reduces the common
good. through, for esanple:
* externalities (spillover efteces|;
*analeyualely defined’ property rights
* transaction casts
* Gailure lo reciunise “exisience values” (e.g. of ile
on earthy.
Torsuch cases, econdmists alsa ire able to presenhe
the appropriate juventive struwture tu redirect human
yetians in the Cormmon jnterest.
ixternalines occur where the sctiins ob one
decision-muker impose costs (ar bestow benefits) upon
other individuals. Problems aeeur when the relevant
decision-maker is not required to take into acemuint
these external costs fur benelity), sa thal an
inappropriate level of an activity 15 undertaken. As an
cxuniple, if nunulacrurer, in ihe course of generating
income, may Use & process that involves signifieant GG
cussions. Suppose that these emissions lead (oret
lime} to a climatic change which reduces globsl
abriculturel productivity and results in a range-of price
increases of basic commodities. Because the
munulacturer’s decision whether to produce ar not lows
not have to take into account the (external) costs
impased upon others. w situation can arise where the
valle of a tmuiufacturer’s oulpul, over and abowe uie
(inihernal) costs oF produchiod, is less than the oosis
imposed upon the rest of the community through the
impucts upon agricalture A tax on emissions would
eneyurage the manufacturer ty behave in a soceally
preferrecdl way,
Whether the presence ‘of externalities is a major
peoblent in practice Frequently depends upon how well
defined are the property tights of a particular resource,
Ownership of resources such as air, water and other
facets Of the natural enVironinent. for a varlety of
Tewsons, Often is poorly defined, TF itis nol defined
in law who-owns.a particular resouree. then consumers
af the resource are without legal recourse when the
quality and quantity of the resaurce is reduced due tn
the actions of others With regard to the earlier
caample, agricultural producers are Hot UL postion to
sue the Mmadufietuber(s) for a loss ih productieity
brought about by inereaset CMF ernissany.
It might seem that externality problems can be
handled by the sunple aet of cusuring tue che
ownership of resourees (such as air and water, ¢te,)
is tilly detined under the law, Leaving aside tor the
moment obvious Jegal complexities, if, for example,
the agricultural sector was given property rights over
the wimesphere. that sector could (im theery) sue fhe
manufacturer for the costs duc to GG emissions: Even
ifthe manufacturer was granted propercy rights aver
ihe atmosphere. then (in theory) a better wufoome could
be obtained as the aycicultural producers could decade
10 pery the manutacturer to decrease production orw
vest ina cleaner pricess. (This may scent “untyir’,
bur it Would result in more soctally efficient outeomes).
It is. perhaps. useful to remensber here that
economists da not prescribe zero emission of GG's or,
in general, zero levels of any other form of pollution’,
Mast unthropovuenre uclivitres ire “polluting” or have
externalilies (0 some degree and economists see the
problem as hiulancing (an the margin) the full costs
(including externalities: of an activity with the benefits
of thalLactivity, The assignment of property rights ean
fo sume Way by secing that decisions about activity
levels uke some qecount of externilitics,
This question of identifying and juking appropriate
wecuuOt all releviinl Costs ts central co Ure eeumomtist’s
trade. It alse leads directly to a funher complication
in the treatment of esternality producing activilics.
Even if property nghts could be cleurly defined, there
may he substantial transactions casts invelved an
abtaining the desired outcome. With regard to the
emissions of GG's, which manufacturers anc
responsible for which proportions of the emissions.
and how much are individual agneultiral producers
affected (and by whom)? At the very least, there could
be very substintial monitormye costs involved, and these
would be “compounded” by further legal and
administrative costs. Asa result. the costs of identifying
and achieving appropriate levels oMGG emissions tay
be prealer than any benelils oblained, Koonomists must
design policies that take into account the costs of
adnumistering the policy. Ibis for tls reason that a range
of other policy-measures are often considered, such
us “polluhion” taxes or deterrent lines. The proposed
hydro-carbon (ax, considered in more detail below. is
one such example.
Consideration of factors such us these give insight
as to why ordinary decent citizens of planet Earth may
tke decisions that are rational for thete individwal
stlves, but contrary to the global good, Understanding
these factors also is necessary for policy design, but
unlortunately may not he sutfivrent. Poley pardneters
can only be set on the husis of agreed valuation’s of
current and future states of the world. Such estimates
Must include Valaahonis of quite complex items, such
ws environmental resources orever "Ile an earth) The
ui fficullies fiere include the whsence of, on irrelevance
ne MoE
wy a simple market value and the question of how the
future should be valued eclutive to the present time (the
chaive of i discount rate).
The difficulties in evaluating assets such asx the Great
Barrier Reel or Anturedca are well understood The
orivate duction of a conmianal good will always Enere
the wilies held by tidividuals who wil be excluded
Ini use et the assets the imputtion OF value on the
basis ul vurrent use of a resource imores both ie
‘option values” of those who would value the
OPPOTLUTy 1G possibly tse a fesduree in future and
the “existence wilues” fell by those, Jor example. whe
Will never Use a resume Hany Way bur who simply
ville ils presenes und preservation
Analysis of alternative policy measures
Instatis Hibo (he peusors Why HOTA actions zine
likely to tead to undesinthle levels of greenhouse wats
emissary and.assocuted clintatie chune. such as those
discussed above. enable veanoimists to preseribe
inventive stroviures whieh Will redindet huinan detions
toward the common interest, There ure obvious
ivforminonal Obstacles io lhe determination oF such
iecntive strugtures, ws cun be illustrated by brief
consideration Of an ideal” data sinuatio),
Clearly. it would: be useful iit was Kitown exautly:
how che emissions associated with different production
Pricesses und GOMsUMpHON aelivites UAPHeted UPON
future production and consumption. the “tasie™
parameters of the population Thal desembe the
relationships hefween consumer choiwe and prices:
dnd. other faclurs, such as the structures of the diferent
Inarkels nthe ecosomy and the policy abjectives al
damestic und foreign governments, Ef all of this wats
Kaown, thes, 6.01 any other closed equilibrium system
whose descriplive parameters are Kiowa. a optinil
intervention strutegy could be concepluilised und
derived
In prustice. (he Inamework within which we have
Ta work hos al best only some stuble parameters and
wine of shichastic elements, Furthermore, whatever
stuble parnaneters nay exist are nob known wilh
verity but Must be estimated. and the preeisivn of
such estiinates iy itself dependent upon eat
undershinding Of the nature of the framework itself
OF course, no Tess ts (rue regarding the esturiation ot
ihe parameters of sysfenvs in ihe physical setences.
The dmpact oF all ot usis unvertainty Js thar economic
poliey prescription olten best can be thought uf as being
a two stige process, First, largely on the basis of
theoretical considerations, iw ty possible to make
qualindive preseripltons that will be robust with respect
oo ouvber of alternative models and parameters
describing real world behaviour The general
supe, merred to above. that a hydro-curbun Las
BURNS & ¢
WALSH
would reduce the incentive for partieipation in vey hes
which contribute significantly (o GG emissions and
climatic change, is an example of reasonably robust
qualiwive policy prescription,
The second sttae is the more difficult, (his one thing
lo be aware thal a hydro-curbot lax can unpact Open
behaviour ia desituble manner. hut itis quite another
to know what size tax should be used.and on what
particular commodities, 115 to obtain insights rate Ls
lutler question Util much current economic policy
rexcurclt is directed, Ar the regent American
Evonomics Assocation meeting menuoned earlier. two
examples were provided af this kind of researets
activity Not surprisingly, both papers use todels
embodying gross simplification of the ghohal economic
and Gimahie interfive.
to Nordhaus (1993), dynamic integrated chit
economy (DIOR) models are extended ~*~. . by
integrating the ceonomic costs and henelits ol GHG
reductions with uw simple dynamic representation uf the
seientitic hniks of crniissiods, concentrators and elirate
change” (Nordhatts 1993 pp. 314). Conditional upon
an estimated toss in overall National Product ol 1%
for every 3°C warming. Nordhaus’ study suggests that
the optioal carbon-tax policy would involve an mitual
wx of just over $8 per ton, resting steadily aver tune
tw reach abuut $20 per ton (based on currenl prices}
hy the end of the next cemury. Wis estunmdted that the
nel henetits. globally, of such & policy conypared ta
doing ovthing would be of the onder of Slo billion
unnually. More significant is the conclusion that the
populardy recommended inmedine cut ol Gi
anissions (tchitive to 1990) would require an mitial
tax of $56 per ton and overali, compured (o 4 fo-control
situation, would detualy involve a ghabal cost of over
$750 billion annually
Eeonomic policy, whether fo do with GG emissions
or otherwise. is itself a complicated and vuntroversial
issue, There has been much argument that public
bodies wre inelleient in their own eetlvities und thi
lax revenues ure hadly used, Iris also widely necepted
(vat most taxes, including many existing Ones, culse
ineMciencies in the economy as a whole by preventing
ihe price system from signalling the “true” values of
commodities aud resourees. With respeet to tysues such
as these, Nordhaus makes two interesting Observations
with respect ty the imposition of even the “optimal”
rute of lax on appropriate commodities. Fitst, if the
lax revenues are used walt a plausible level of
inefliciency in the public sector all bengltits due to
GHG emission reduction may be totally outweighed,
Second, however, il these x revenues could be usin
to eliminate existing taxes which Were distorting the
price mechanism in a costhy manner, the unnaalisea
global beachite might inciease from S16 billion ta aver
$250 billion,
VHE, ECONOMIC IMPLICATIONS OF CLIMATE CHANGE HY
The above is useful in that it gives a flavour of bow
econaniists approach the mleztated analysis of climate
and the economy over time. It also indicales thal no
Single economic policy can be cousidered 1n isolation
from other government interventions in the economy
What the discussion fails to convey. however, as the:
scale of the uncertainues which economists admit to
in this ares OF investigation,
Guskins & Weyant (1993) offer a vivid, if perhaps
extreme. illustration of this uncertainty, In their
comparisomof 4 "DICE-type” models and 13 climatic
scenariog, estimates of the averige carbon tax required
over the period 2000-2020 to achieve, for example, a
20% reduction in carbon emissions (compared to 1990)
vary frum $350 per ton ty $330 per ton. In fairness.
extreme cstimales tend io flow from extreme
assumption whderlying alternative models and it
Tundamental aspect of model development is to dentify
the potentially sensitive areas and to refine the accunucy
ind level of agreement regarding these ares.
Other policy evaluation exereises, in many ways
simpler are undertaken by economists. including the
previously ented approaches using the ORANL model
Ihis model is a highly dtsageregated veneral
equilibrium imodel of output. employmenr und
Consumplion inthe many different sectors and regions
uf the Ausiralian cconomy. It would not be
Openitionally feasible to construct a DICE model at
this level of delat, but what ORAMNI can do is predict.
for u given well-defined policy initiative, the likely
niero-eflects aeress the regions ol the economy, To
know in ailvance, for example, that # particular section
of the Adelaide inetropolitun region is kely
experience signilicant unemployment hecause of GG
policy measures, can be of significant use wy svcil
planners.
Conclusion
The capacity ol econornisis to provide clear
predictions of the costs and benefits of climate change,
ar of palivies to modify likely future climate changes
1) limited by a number of factors, not least the
uncertainties surrounding, predictions of chimite
change, The work done so far ts shght aod highly
spoculative. but it bas grown — and in collaboration
with our colleagues in the physical and health sciences,
dor example, it could further expand in extent and value
to the policy process — especinily as.the world works
Ws way out of the problems of recession that have
thoroughly captured the attention of policy-makers int
recent years.
The significance of collaborative approgches cane
be over-emphasised. Espectilly in areas where extant
knowledge ts very limited, and subject toa very high
degree of uncertainty, if can ensure thal research und
modelling investinents by different disciplines are
mutually supportive. Because of the nature of their
“business, economists bong not only analytical and
modelling skills to the considerauion of issues such as
vlimute ehanve, but also a sharp foeus on what ts
“policy-relevant” information wad analysis.
Conary to the belief of many, if not most, non-
economisis, economists do not believe that private
seclor markets always work well and provide the rigtnt
signals and ineentives. But their scepticism about
private markets applies equally to political markets
where lobbying bas a powerful role. and where
repulatians can become captured by those they are
supposed to regulate, Ancarly keen eye to these issues
can resull in policy debate und design heme focussed
on achieving better ultimate policy outzomes.at lower
Cost.
Keferences
Cumann CHanar Group 1992) Climate change seenitrios
for the Australian region, ( CSIRO Division of Aimospherte
Rescurch, Mordiallne).
Gaskins, DW. & Weyanr, L P1993) Madel Comparisons
of the Costs af Reducing CO, Emissions, dmentcan
Econcmie Review BAZ), M&32I
Hough, J.T) dawkins, G. J. & Draraums, Jd. (bd)
(1990) "Climate Chane, The IPCC Scientitie Assessment”
(Cumbridge University Press. New York),
Hincions, D., Gross, R., Winsow. B. & Serppaan. P92?)
“The economic impact of climate change on ughenulfune
in Sonth Austrahad. (Department of Environment and Lund
Management. Adeluide)-
IC (tyoesrky Commission) (291) “Costs and benefits of
reducing greenhouse gus Cosine” (AGPS,: Canberra).
Nokbitaus. W. D. (1993) Optinial greenhouse-gas reductians
and tax policy inthe “DICE” model Diericun Bconeomtie
Review 83(2)_ 313-317,
Pranwan, Go, (¥és) “Greenhouse — plupning dor elimine
change’ (CSIRO Division of Almospheric Research, E.J,
Brill. Melbourne & Leiden),
Remy, J & Hommans. N, (1993) Climate vbange and
agriculture: The rote of mnternational (nade American
Erononne Review 83 (2), 306-312
Yosey. J. Retry, Eo Kase. §. (1992) Economie
implications of global change for world ugriculture, Journal
aw) Auceultural und Resources. Fevicnttcs U7, W5-205-
CLIMATE CHANGE AND HUMAN POPULATION HEALTH:
GLOBAL AND SOUTH AUSTRALIAN PERSPECTIVES
By A. J. MCMICHAEL*? & M. Y. BEERS*#
Summary
McMichael, A. J. & Beers, M. Y. (1994) Climate change and human population
health: global and South Australian perspectives. Trans. R. Soc. S. Aust. 118(1), 91-
98, 31 May, 1994,
Public health scientists now need to think within an ecological framework because of
planetary overload by the human species. Climate change must impact on human
health. Predicted health hazards include increases in thermal extremes which impact
particularly on the very old and the very young. Natural disasters such as cyclones,
floods and ocean storm surges will increase rates of mjury and death. Insect and other
vectors for infectious diseases will change geographic distribution resulting in
changes in the demography of diseases such as malaria, yellow fever, dengue,
encephalitis and cholera. Changes to agricultural productivity will alter food
availability in many regions. Rising seas would, amongst problems, disrupt sewage
disposal causing diarrhoeal diseases. Climate-induced environmental disruptions
would create “ecological refugees” leading to spread of infectious diseases and social
disorganisation. Ozone depletion with increased exposure to ultraviolet radiation will
lead to an increase in skin cancers, eye disorders and immune suppression. The
significant environmental degradation in South Australia, together with climate
change could lead to significant increases in water pollution, outbreaks of Ross River
virus, Murray Valley encephalitis, dengue fever, tick-borne diseases and possibly
Hanta viruses together with gastrointestinal diseases.
Key Words: global environmental change, human ecology, human population health.
Trnsnrtivas Gf the Bavel Seeterm pf So dust (999), TB), SE BR,
CLIMATE CHANGE AND ILUMAN POPULATION HEALTH;
GLOBAL AND SOUTH AUSTRALIAN PERSPECTIVES
by AL). McMicHaAtL*| & M, Y. BEERS*i
Summary
MemMicnarn, A, t & Buers, M. ¥, (994) Climate change and human population health: global and South
Australian perspeetives, Trans, Ro Soe. 8. Auyt. DRL). 91-98, 31) May, 1994,
Publie health seteotists now need ie think within an ecalpgical framework because of planelary overload by
(he human species, Climate change must impact onchumnan health, Predicted health hazards melucde mereages
ii Thertyal extremes Which impact particularly on the very old and the very young, Natural disasters such its
iyelones, floods and ocean storm surges will increase cates of injury and death, Indect and-otber veetors fee infectious
discascs will change geographic distribution resubing in. changes in the demography of diseases such as muliria,
yellow: fever, dengue, encephalitis and cholera, Changes to agricultural productivity will alter food availability
“nmMaty tepions. Rising seas would, amongst problems, disrupt sewage disposal causing diarrhoval diseases.
Climate-induced environmental disruptions would creat: "ecological refugees” Ieading lo spread of infectious diseases
and social disorganisation. Ozone depletion with inercasedt exposure 19 ultraviolet radiation will lead to an increase
ski Cancers, eye disorders. and immune suppression. The significant environmental degradation in South Australia,
toxether with climate change could lead 1 significant increancs i water pollution, outbreaks of Ross River virus,
Murray Valley encephalitis, dengue fever, tick-borne diseases and possibly Alana viruses Woyether with gastrointestinisl
diseases.
Kiy Woris: glabal environmental change, human coology, human popubatiean tealtl
Jntroduction
During ils quarter-million years of existence, Mame
sapien has adjusted to several shifts in the prevailing,
profile of covironmental health hazards. Each shift has
coincided with Gulturally-driven changes in the social
organisation and ecological relationships of the human
species: the advent of agriculture, urhan semlement.
industrialisation, and the spread of “ulMuent’ living.
Today, another category of environinental health hazard
is emerging that is qualitatively dishincl from earlier
hazards and may well have wider-ringing health
consequences, This hazard arises From the plantetary
overload resulting from the cumulative impact ol the
human species, via population growth, land pressures,
energy-intensive technology and 2 high output of waste
vuses — including changing the gaseous composition
of the lower almosphere and, hence, its heat-trapping
propecues (McMichael 1993),
Appraisal of the health hazards of these incipient
wlobal environmental changes requires us ta think
within gn ecological framework, Public health
scientists have aot heeded this framework previously
because, until recently. humans were (apparently)
* Departmen of Community Medicine, University of
Adelaide, Sauth Australia 5005.
+ Curent address: Deparment of Epidennology and
Population Sciences. London School of Hygiene and
prepical Medicine, Keppel Street, London, WCIE 7HT,
+ Curren address Communicable Diseases Comnal Ui,
Epidemiology Branch, South Australivn Health
Commission, Adelaide. SUOT,
living Within the biosphere’s carrying capacity. Now,
however, there ure signs that our aggregate impact 1s
reducing the stability and productivity of various of
Harth's natural life-supporting systems. Each of these
individual problem areas is already a familiar subject
of envicunimental research and debate; greenhouse gas
accumulation, stratospheric ozone Joss, land
devradation, aquifer drawdown, depletion of occan
fisheries. Juss of biodiversity, and the ecological and
social consequences of rapid urbanisation. But their
wider, collective. eco/ogical significance for luinan
population health has been rarely considered
There is much that is uncertain — und controversial
about the causation, course and consequences of
global environmental changes, This has been well
demonstrated by the divergent views of scientists about
the net impact of grecnhouse gax accumulation upon
the world's climate, The consequent difficulty for
health researchers x knowing whether and how 16 reap
further uncertainties about health outcomes upon the
underlying uncertitntics about global-change
processes. Nevertheless, since sustained averload of
the biosphere must, logically and eventually, entail a
reduction of its life-supporting ability, it 1s a reasonable,
and prudent, inference that this poses a fundamental
hazard to human health. Esumations of health inpact
must therefore be attempted
The best-documenmed and most discussed aspects oF
global environmental change ure Those pertaining to
climate change. “Climate chunge” bas been widely
taken to include both greenhouse enhancement and
tvame depletion. Purists may argue that “climate” is
4 lropospheric pheneinenon, and has to do with
2 AJ J McMICHALL & M. ¥. BEERS
temperilure, previpilanon, humidity and winds; since
uzone depletion i occurniig, ii the stratosphere, ald
its main anticipated consequence is an increase in
surface-level ultravioler irradiuean. eis wot pur of the
eimate change, However. the demureation is net
cleurcul — there wre various intersenve and other
relations between these wo: processes, For example.
vhiurofluorocarbons (CFCs) contribuie to hoth
processes; tropospheric warming may cnbisnce
sirtlospherie cooling; and ozone depletion may affect
the nidiative forcing properties of (he almosphere
overall, Lv this pauper we use the inclusive apprough.
Two other qualifying comments are needed here,
First. specifying the adverse health elfects of ¢linusle
chawe rests largely on extrapolation and. in some
eases, ressonuble conjecture (MeMichuel 1993:
Maskell 1993}, Unlike the directly toxien|ogical
hazards posed by localised environmental pollutants,
tins cutewory of health hazard does not readily permit
the empirical observation of health impact im one
Population ws a bass: for esumating the visk ia other
populations, As more information about these cimplex
systems accrues, the modelling of populatien health
impacts will improve.
Sceond, itis generally diMicull (o make predictions
of localised etfects ol climate change upon the health
of South Australians, bor certain. simpler, processes
— such as skin-cancer risks duet increased ultraviolet
irradiation (UVR) and the change in mortality due to
increased frequency ol heatwaves — local estimates
un be made, but for many other processes (c.g. citeets
upon regional and global food production) the health
of South Australians will reflect their sharing af the
experiences ol wider populations. Therefure. this paper
concentrates initially upon the general possibilities ot
population health impact, ater which some more
spocifie comments ane made about the South Australian
UUMMERL
Greenhouse gases and climate change
Various maternal “ercenhouse” gases in the Inwer
atfhosphere thserh tmich of the heat that 4s re- radiated
fram the Earth’s surface, Fossil fuel combustion and
accelerated forest clearance since 1800 has increased
the concentration of carbon dinside trom 275 ppm to
34) ppm. Coneentrations of orher anthropogenic
treenhouse gases. especially methane and the
chlorofluorocurhons (CECS)_ are also increasing (IPCC
(992: Epstein 1992).
The United Nations Intergovermucotal Panel on
Climate Change (IPCC) estimates thal, in consequence
vt the increase in radiative forcing. the average global
temperature will rise by around 2-3°C by the your 2100
(IPCC 1992), While this estimate is acknowledged to
he uncertain, the evidence is firming Urab human
activity 1s how Warming the Exrth’s surface (Lacis &
Carlson 1992). A rapid global temperature merease
al several degrees would be without precedent in all
A) huinaan history. and would pose a range of risks to
human health, both by direct and (probably more
nnponanth indirect mechunisnis (Last (991). The 4-59C
Inercase ih temperature after the more regent ie age
which (several smaller Nuchutliiins aside} occurred yer
several thousand years. caused major ecological
chunpes — including the spread of forests wnd. together
with intensified human predatin, the extinction ol
many large mianiials: Halse presumably contributed
to (he onsel of agneullure
The most likely impacts of greenhouse-induced
climate change are summarised in Tuble 1.
TAHLE 1. Mata potentiol effects of glabul climate change un
population healihs
Increased Mrequency af leatwaves — deaths.
illness. injury,
Respiratory cheers of moisture, dusts, pollens,
Climate instability — disasters (eyelones,
floods. fires),
Alicrald hubstat and (ayasinissian et vecty
horne infectious diseases (uiher contagious,
Uisesses Oso).
Impaired crop production (soil temperiuire,
wiler, Pesto),
Soa-level cise — inundation, sanitation,
salinity: Cle.
Demographic disruplon. environmental
tetugecs
Direct
Indirect
The main direct hazard to health Froni a temperature
increase comes [rom thermal extremes. It is predicted
that the frequency of heatwaves in teniperale and suh-
tropical climates will inerease (Brasseur 1991) — for
example, a doubling in the annual number ol days over
38°C in Washington, DC, is predicted over the next
half-century (Leal 1989). los not unusual op tempenite
vones, such as South Australia. tor three to four days
cach summer to record temperatures greater than 38°C.
However, with an avenigs aanual temperature inerease
of 2-3°C, the number of diys in excess af 38°C 45
predicted to rise (Ewan er af. 1991),
Intesponse, death rates aniong the most suscepaible,
particularly the very old und very young will rise
(McFarlane 1978; Kilbourne 1992). Ciiuses of death
wo rise Would inelude heat-steoke. exhaustion,
cerebrovascular stroke and, probably, accidents. and
acts of violence. On the other hand, winter deaths trom
influenza and trom hypothermia — particularly among
the homeless. poor fram Delhi to London to New York
— vould be expected lo decrease.
Weuther patterns are likely tu be destabilised, with
more frequent cyclones, Hoods and ocean storm surges.
These “natural” disasters would increase rates of injury:
and death. They would alse disrupe agriculture. lon!
transport and snniiry engineering. Fringe dwellers i
CLIMATE CHANGE AND HI MANS POPULATION HEALTH 93
flimsy housing or exposed co rreers and oceans, as in
coastal Bangladesh. wil] be the most vulnerable
(Houghton er al, 1990). In Australia, it is predicted
that cyclonic aclivity would extend down the eastern
coast from a latiiide bound.o 27°S 16 31S (Australian
Bureau of Statistics 1992), However, there gre unlikely
in he consequences lor Australia’s southern coastline.
Variols insects, vertehrates and other veeturs for
infections disease wall change their geographic
disuibution (Cook 1992; Shope 1892). As higher-
lutiiude locations become warmer und wetter
mosquitoes will tend to spread) there, As well as
extending (hei spread und seasonality in tropical
countries, mosquito-borne infections (tmalena. yellow
lever, dengue. and various forms of encephulitis) may
ret t currently Unaffected lempeniie areas (Ancient
Rone and early Eurype were malarious; so, earlier
this century. were puts of the United States, England.
lly and northern Australie.) Vector-borne diseases
spread by (lics, water-snails and vertebrate animals are
also. likely ta change their distributions. Recent
evidence that cholera is “veetor-barne™ by taking reluge
under the mucins puller coat of aquatic slgae
fincluding spectes found in both fresh and vost
Walers), suggests [hal ils spread may also be assisted
by warmer oceans (Epstein 1992)
Lamy-term increases in backe round temperature mury
reduce the yield of same of the world's mest productive
AKIN-growing temperate regions the American
primes, western Euiope, the Ukrdine and coastal
Australias while inereasing the agriculiural
productivity of other regiins (norfhem Europe and
Canuda) (IPCC 1990) Crops. that hive been
selectively bred for specific environments. my ful wo
germinate, Climate change may also alter habitats tor
those insects. fungi and micro-organisms thal cause
“diseases” of grains, fruits and vegciables. While
increased levels of carbon dioxide in the atmosphere
may enbance plant growth (especially the evolutionary
older “C3” plants), most current evidenge indicates
thal. on balaace, this would not be a significant effect,
Further, while the overall net effect of climate chunge
upon world agriculture is uncertain (Parry 1990)
Crosson 1989) ata regional level food security may
be sevivusly threatened — ocspecially im poorer
counties inthe semi-arid end humid tropics. Climatic
vhange would alse perturb the growth and prialuctivity
of livestock and fish, local fuelwood supplies and the
inailability of freshwater - all inspartant determinants
Ot healt.
A rise in sea levels is predicted from thermal
expansion of the ocean. The IPCC estimares a rise of
upproatiiilely 0.65 pietres by the year 2100, with
serious effects upan low-lying coustal commyntics.
The mist vulnerable settlements are those in ccastal
Jowlands such us in Bangladesh, China und Evypt, and
in wall island mativgs. Inundation and storm surges
could lead to significant displacement of peuple
(Maskell ev a/, 1993: Worres| & Cirant 1989), Risine
sens woulll also disrupt sewage disposal (thus causing
diarthoeal diseases), cuuse invreased salinity of Goustisl
freshwater estuaries and turmlands, aid han worlands
und the breeding grounds of many ooean Tish (Haines
& Fuchs (991)
Many of these climate-based environmental
Uistuphims would create “ecological refugees”
(dacohson 1989) Dispassessed people from areas
damaged by Moods. cyclones. risiag seas or droughts
wruld migrate to the relative salely of cines, Shanty
towns would proliferate and so would infectious
diseases and socal disorganisation, It is cancetvable
if climatic and agricultural and clenwyraphie circu
suiftice’s Ueterinrate significantly fiche coming century.
that Australia will have an influx — yvoluniarilyor even
uncontrollably. — of refugecs from Seutheast Asia.
Ovene layer depletion
Stratospheri¢ azone. whieh resides at 1-20 km
allitule, wocunulated over several billion years
principally as a resall ob the evolution of uxygen-
releasing photosynthesis, This ozone “hiyer” provides
life on Furth with substantial protection from
bighowically-damaging UVR (partigulirly the higher-
energy UV-C which js entirely absurbed and the
medium-energy UV-B which is largely ubsorhed),
loday, the stratospheric ovine ts being damaged by
human-niade chlorotluoreearbons (CFCs) and other
industrial pusenus emissions.
AL miid-oorthern latitudes (30-SO°N) the ozone layer
thinned progressively by around 0.5% per year during
the 1980s snd carly 1990s (Stolarski er al 1991;
Brasscur 1991; UNEP 1991). Overall. during that period
(here was an accumulated 8% loss in Winter and a 2%
lass in summer Meanwhile. in the southern
hemisphere, siynificam cumulative azone lass ocurred
during the 1980s from the pole “down” to Lutaude 3U°S,
which. includes the lower parts oF Australia and of
South America, There now appear to be well-
established, strengthening trends in ozone depletion
in both hemispheres. The anticipated increases in UV
irradiance at ground-level would be greatest in the
southern parts-of Africa, Australia and South Anrericn
and inthe mid-latitudes (30-60°N) in Europe. Asia and.
North America (Madyorich 1992),
The predicted direct clects of inercased homan
exposure to UVR include inecrenses. in skin cancers,
eye disonlers, and suppressive cffests upon the MMyn¢e
system. The LN Environment Programme esiimutes
that Jor every 1% decrease in azone, there is iin
Approximately 15% increase in exposure to ullaviolet-
B (UV-B) radiation at the Eanth’s surtace; in tum. each
sustained 1% anerease in UV-B would case un
oF
estimated 2-3% increase in the incidence of skin
cancers, predgminuntly non-melanoma cancers.
Increused UV-B exposure ts also predicted to increase
the occurrence of caturucts und of pterygia (tissuc
overgrowths on the cornea) (Taylor 1989), Less
certainly, it may depress the body's immune system.
thus reducing protection against infectious and fungal
diseases (Morison 1989). Alihough there has been
trequent reference to these various predicted health
outcomes. the estimated incteases in incidence are
based on rather sparse data (Armstrong in press).
During 1993, un attempt was made in Austraha to
predict the number of extra LVR-induced cases of skin
cancer — basal and squamous cell curcmomas and
malignant melanoma — and of ocular cataracts and
plerygia that would occur over the next four decades
(Fraser et al. 1993). The calculation éntailed, first,
estimating the dose-response relahonship between level
of UVR exposure und the incidence from existing
epidemiological data. Then, from knowledge of trends
ja ozone depletion and grognd-level UVR during the
Biological
Aol McMICHABL & M Y. BEERS
preceding decade, and from estimates of future trends
in ozone damage, predictions were made of changes
in ground-level UVR over connng decades. From these
estimates, and appropriate amplification factors, the
numbers of extra incident cyses were predicted, The
predictions for South Australia are Shown in Fig. I.
(Note, however, that the techniques for making such
predictions are still crude, and, im this particular case,
there wus inadequate allowance for mmimual duration
of exposure ahd for cancer latency periods, These
predictions are therefore illustrative, not definitive!),
Increased UV-B exposure would also have
deleterious effects on the world’s biota, including
impairmg crop yields (Worrest & Grant 1989). The
estimated danger to marine organisms could be
sizmificunt, since increased UV-B irradiation of the
ocean's surface could damage the photosynthetic
phytoplankton population ~ the basis of the aquatic
food chain (Smith e7 al. 1992). (A quarter of the protein
supply in the human dict comes from the sea, as does
imich of the protein fed to livestock.)
cance! ar - t
5 CON SA Cancel ome earac
cquat US neat CON a melan prego ea tical &*
: :
E
2.2 17 14 Tz 0.8
amplification factor
10,000
1,000
100
10
Annual incidence of extra cases
(Note: Log scale)
2005
Fi, 1, Estunuted annual extra numbers of cases of skin cancer und ocular lesions due to increased ground-level exposure
tultraviolet radiation, consequent upon stratospheric ozone depledan. Numbers are expressed relative to the (background)
incidence of these conditians in 1990, (Prom: Fraseres a. 1993, Note also the caveats in the accompanying text.)
CLIMATE CHANGE AND HUMAN POPLLATION HEALTH 95
South Australia
Heathcote (2990) his proposed chat climate change
should not be considered in isolatyon from <limate
aritbilvy as Che fatler may privide valuable
informauon on which to base estimates of change. An
examination ol the elimate and ecology of South
Austrailia provides particularly interesting perspectives
of Such variation with which to consider the possible
human health effects of climate change,
Australias a whole und South Australia in particular
has one Of the most variable chinates om each, Phas
is no new phenomenon but has been the case for
hundreds of thousands of years (Nicholls 1992) Cuok
(992), This predictable but irregular variability has
resulted jn the adaptation of the flora and feuna of the
Siate to suit such changeable conditions,
Covering one eighth of the Australian continent,
South Australia lies wholly io the temperate zone. ‘The
land is low, with the inland areas being largely covered
by plains, sandoand gibber deserts. 80% of the State
is less than 300 metres above sea level, The climatic
consequence ty that apart From the coastal areas, there
are ne signilicant mountain ranges ty encourape
precipitation evenly throughout the Stave. Much of SA,
therefore, is arid or Senmearid (Gardner 1993)_
Weather never replicates itsell exactly from year bo
year (Noice er al. 1993). Australia has been found to
hive the highest interseasonal rainfall variability and
rum! an the world (McMahon ef af. 1987) oath
drought being a frequent occurrence when defined as
annual tainfall in the lowest (0% of records (Gibbs
& Maher 1967). SA is ibe driest State with fout-fittlesy
of the Stote normally receiving less than 250 mn of
rain annually = while only one third of Aastralia as
4 whole receives less than 250 mm, The basic fentires
of the SA climate are hot, dry summers wath relatively
mild nights and cool but not severe winters, where most
of the rain falls helween the months of May to August
(Gardner 1993),
Vt is clear that changes to the rainfall pattern alone
could dramatically alter the ecology of this State,
Not only is the geography of the fand and natural
fluctuitions influential in climate variability in SA, hot
4 phenumenun known as ENSO, of the EL Nino
Southern Oscillation plays a significant role in the
climate of Seuth Austritia. The Southern Oseillition
isa major shift in wir pressure between Asiy and easter
Pacilic regions which elects changes to trade winds.
cloud amounts and rainfall over the tropical Pacific
through to Eastern Australia (Voice er al 1993.
Nicholls 1992), extending ity influeuce ay far Westas
the River Nile (Quinn 1992), The Southern Oxcillavion
is measured by an index of the difference in air pressure
between ‘Tahiti and Darwm, A posmive index (low
pressure at Darwin) sieans that (ade winds blowing
strongly aeross the Pacific feed moisture into the
monsoons of Astiand Australia, A negative index (high
pressure al Durwin) means thal the trade winds are
weak ar even reversed.
A significant Warming of the sea surface in the
castern equatorial Pacific occurring at the same: tine
a8 newalive index measurements of the Souther
Oscillation, 16 known as El Nino, The reverse effect,
known ay La Nifia, takes place when the Southern
Osertlation index ts positive and the waters of the
castem Pacific are cold. The combination of the
Southern Oscillation and El Nino is known as the Ti)
Nina-Southern Oscillation or ENSO.
This is nota recent effect. Paleoelimatic records have
Uncovered evidence of the FNSO as far bagk as (000
years (Cook 1992), Thos predicted that ENSO ty
unlikely to change dramatically or vanish onder
conditions brought about by the level of climate change
currently predicted (Voice et al, 1993), However, it
is not yet known whether ENSO may become variable
or more frequent, desiabilising the presen! (2 month
cycle.
AL present, ENSO accounts tor about 90% of major
droughts in Eastern Australia, Further, this same
phenomenon ts responsible for approximately 20% af
all rainfall variations in eastern Australia including
flooding, wet seasons lasting six to alae months as well
us less dramatic events (Yoice ef al, 1993), Tt is
therefore a significant contributor to the extreme
variability in the Australian, and South Australian,
climate,
Over the past 200 years of white settlement, human
activity in adapting und manipulating environments to
provide for hetter living standards, may have incretised
harnan vulnerability to the effects of sysnificant climatic
and environmental variation (Heathcote 1990), In other
words, Hugin adaptation hus been in the form of
modifying the external environment as opposed to the
longer evoludion of native Nore and fauna which modify
their own physical characteristics and systems.
Hustian atichipts at chylroniental modification 1
Souttr Australia have included extensive land clearance
resulting in desertification und increased salinity of soil
and water. Delorestation of catchment areas has led
10 flooding and erosion, and agricultural practices have
contaminated the land with pesticides and heavy
metals. South Australia’s only major river, the Murray,
is heavily used as a water supply. and irrigation base
in the three States (New South Wales, Victoria and
South Australia) through which it Nows, In recent
years, siznificantly lower water levels and reduced flow
in this Stare, combined with pollution with phosphates
und nitrates, lave resulted in extensive algal blooms
(Soong 1993), The Murray bas 4 fall of only 20 metres
across South Australia (Gardner 1993) and is therefore
nota fast Howing Ayer, nembering il susceplatle lu ulaal
blooms,
96 A. . McMICHAEL & M. ¥ BEERS
Within the context of such significant environmental
Degradation, abienpis must be made to predict and
prepeue for humuin health needs in a climate change
scenarin where average global surface temperatures are
predicted © rise by around 27-39C hy the year 2100
(PC 1992).
Over thousands of years of climatic variability,
Australia’s flora and fauna have evolved to accommodate
the climate, and indeed to flourish. Nicholls (1992)
describes adaptations in the behaviour and breeding
panerns of the Red Kangaroo, the long-haired rat and
other Australian birds, insects and plants, All have
Jeyeloped survival pauerns of breeding in plague
Proportions in wet seasons when food is plentiful and
a virtual shut dowa in tinses oF extended drought.
More permanent increases in summer mainfall and
Higher minimutn tempersiures in SA as predicted by
climate models (Heathcote 1990) could lead to plagues
unhampered by the natural controls of drought and cold.
In the past (2 months, excessive rainfall and milder than
usual minnmium lermperatures have led to plagues ol
mice, locusts, various bird species, wasps and
mosquitoes. Floods have created disuster and death i
human populations, Water supplies have been canta
nated wath huntarr and animal sewage (Walters 193!).
The human health consequences of such
meteorological variation have included aw South
Australian outbreak of Ross River Vinas (RRV) of
epiderec proportions. There were in excess of B00 cases
of FURY in the period mid-September 1902 t Mid-May
1993 (Camenin 1995%). By comparison, previous lesser
cpidemacs were in HS4, 136 cases and IORY, 115 cases
(Weinstein 1). The underlying annual mean muinber
of cases ower the last ten years (excluding epidentic
years} 16 less than 10 cases per year (Bell ev al. 1991).
The 99293 epidemic occurred as & result of high
rainfall amd malel temperstures Cron August 1992 tw
February 1993 as reported by the South Australian
Regional Office ot the Bureau of Meteorology (1993)
which provided perfect breeding conditions tor mosquito
vectors dedey vigiler aml Cidex annvliroseriy.
“Watrers, R. 0999) Persunal communication, Slate Water
__labersiory, South Australia,
“Camenos, S$. (1993) Unpublished data, Communicable
Diseases Control Unit, Epidemiology Branch, Public and
Enmiroamentl Health Service, South Australian Heulth
Commission.
‘SuTHERST, R, W. (1992) The Likely Impact of Climate
Chonige on Medical Vectors. Paper presented al the Australian
Tromeal Health and Nutrition Conference, Brisbane.
"Barry, R, D., Hupson, BJ, Saareun, DOR. & Wins,
MC. (1993) Evidence tor an indigenous form af Lyme
Borreliosis in Australia. sr presented ut the Australian
Troptea! Health and Nutrition Conference. Brisbane.
*DowNWAKo, RK. (1993) Disaster management. Report to the
Tth Sanding Coriutice on Industry, Science, ‘Techinolngy,
Transport, Communication and Infrasinuctaun: Wuslinnd, SA
Wed. th GQetober_
Discuses predicted to neresxe include Murray Valley
Encephalitis (MVE), also known as Australian
encephalitis, Caused by & mosquilorbome Maviviris,
MVE was last recorded in South Australia in 1974.
Excessive rains as a result of an ENSO ever had lel
to an extended breeding season for the heren population,
the natural reservoir of infection lor MVE, The virus
is transmitted via the mosquito vector dedes aegyplae
and has a case fatality rate from. 0.39% 0.60% (Benenstin
1990) and is often around 30% (Manson-Bahr 1987).
Nicholls (1986) subsequently devised a method, basexl
on historical research, of predicting outbreaks of MVE
using the Darwin mean air pressure of the Southern
Oscillation.
Other predicted inypaces uf clamate Change en South
Australia include the move south of denpye fever, tick
borne diseases such as Lyme distase, and even malaria
(Sulherst 492°; Ewan ev al, 1993), In Petruary 93,
local wansmission of falciparum malaria is reported 3
have occurred in Queesislind, south of the nineteenth
parallel (Murtay-Snuth & Weinstein 1993). Although
Australia was vertified {Tee of endemie Malaria by the
World Health Organisation on ist, there are over 70)
cases of imported Malaria annually, The common
vectors for malaria, the Anopheles Jarauti 5.1, and An,
punciulatus mosquito are considered to exist only above
the 19th parallel, known as Australia’s malaria receptive
zone. However. An. farauti in known to range south of
the 2ist parallel, while vn. antrilipes, previously im-
plicated in local transmission of malana in Victoria ancl
New South Wales (Ford 1950), ranges as far south as
Tasmania (Russel 1990),
The implication for South Australia is that with
climatic change, at least 4n. annulipes is capable of
establishing a tuche here, becoming infected by biting
cases Which afe at the gametocytaemic stage of the
disease and transmitting the disease locally,
The spread of Lyme disease could occur because tl
increased vegetation due to higher rainfall in South
Australia, Although no vector for the anaerobic
spuochaetes of the Borteliae parasites. he causauve
agent for Lyme disease, bas yet been identified m
Australia (Russel er a/, 1993), there is suggestive
evidence from serology reports from patients with Lyne
type symptoms, thal indigenous Lyme borreliosis (LB)
mewidespread in Austrailia aod that a nurnber of strains
of the disease are present (Barry ef al, 19934),
Perhaps more frequent mouse plagues would bring
with ther new and serious diseases such as the Banta
Virus. which emerged in the US this year (Leduc et at
1993, Mestel 1993). Methods of controlling such plagues
are ud hoc at present, expensive ($1.6rn.) and largely
inappropnate for long-term use, The use of poisons,
such as strychnine in the [993 South Australian mouse
plague (Downward 1993") is not a vinble long-term
response to Problems that have their origins fs coulogival
and climatie disruption,
CLIMATE CHANGE AND HUMAN POPULATION HEALTH uy
Contamination of water supplies has led to anecdotal
reports of inereased incidence of diarrhoeal and
gastrointestinal diseases. There have beer 1250 cases
of Hepatitis A or infectious Hepatitis in South Australia
since 1983, with 510 of these ogcurring, in 1986 (Bell
eral. W9L). Hepatitis A isa disegse of public health
importance in flood situations or other disasters which
affect infrastructure, Waterborne diseases such as
siardiasis (caused by the protozoan Giardia lamblia),
Campylobacter jejuni (found in milk. waler, meat) ond
amoebiasis (Entamoeba histolytica) could also pose
problems ina wetter SA, while increased temperatures
may lead to more inlection with Leyionel/e and food
borne disease such as Salmonella and Yersinniz, the
latter being already on the increase and ol significant
public health concern (Bell et af, 1991), Legionella
bacteria survives better m waler aerosol which conan
blue-green algae (O'Brien & Bhopal 1993), this having
significant implications for the transmission of the
disease ina warmer climate scenario.
Th is predicted that with global surface warning of
2-3°C the main direct hazard to health from uw
temperature increase Comes from thermal extremes,
South Australia endured temperatures in excess of Aec
in February 1993, provoking media headlines ay far
away as Brisbane of “Heatwave kills 1. hundreds in
hospital” (The Courier Mail 1993), while in Adelaide
it was “Heatwave drama as toll rises” (The Advertiser
1993), Children, the elderly and the unwell were the
main groups affected,
Finally, SA currently enjoys up to 3,500: hours of
sunshine annually, out of the approximutely 4,380)
hours possible (Gardner 1993). Increasing levels of
UVR post a worst case scenario of mot only
significantly increasing skin cancer rates, bul also of
initiating Immunosuppression, rendering the population
more vulnerable (0 various infectious agents. Evidence
troro transplant patients, the immuncgcompromised and
AIDS patients illustrate well the range of opportunistic
infections which may: result from sweeping population-
based apumunosuppression. The efficacy of
immunisation may be reduced. and diseases which have
heen “controlled” in the past may therefore re-emeige.
Conéhusion
One result of the increasing impact of lon sapiens
on the world’y natural systems is thal our foeus of
environmental health concern must move beyond the
realm of polluted local environments ancl rite the reali
of disrupted natural systenin. Anticipating effects on
Population health is made difficult by the combination
of unfamiliarity, unceriainty and timing (the
unacceptability of an empirical “wait-and-see"
approach). The risks of cancer and cataracts from
ozone depletion can be easily apprecuited; likewise the
healch hazards. frou heatwaves. More difficult to
conceptualise — but of potentially much greater impact
are the anticipated indirect health consequences of
climatic effects on food production and on the spread
of infections, und, perhaps, immune suppression hy
ultraviolet radiation,
Some extreme, and extremely costly, technological
adaptations may be possible — such us erecting
ultraviolet barriers aver living spaces or investing
massively in seawater desalination, However, im
general, human population health cannot be sustained
if the Earth’s natural systems are not maintained, For
the first time in human history, there is evidence that
various of those natural systems dre becoming over
loaded at a global Jevel. This portends a Jrameshilt
in how we must think about “environmental health”,
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TRANSACTIONS OF THE
ROYAL SOCIETY
OF SOUTH AUSTRALIA
INCORPORATED
VOL, 118, PART 2
BIOLOGY OF PHYLACTEOPHAGA FROGGATTI RIEK
(HYMENOPTERA: PERGIDAE)
AND ITS PARASITOIDS IN SOUTH AUSTRALIA
By T. A. THUMLERT & A. D. AUSTIN*
Summary
Thumlert, T. A. & Austin, A. D. (1994) Biology of Phylacteophaga froggatti Riek
(Hymenoptera: Pergidae) and its parasitoids in South Australia. Trans. R. Soc. S.
Aust. 118(2), 99-113, 31 May, 1994.
Information is presented on the biology of Phylacteophaga froggatti Riek (the leaf-
blister sawfly), an emerging pest of plantation and ornamental eucalypts in south-
eastern Australia. The range of eucalypt species attacked, the extent of tree damage,
seasonality, fecundity and longevity of P. froggatti are investigated for populations in
the Adelaide region. The hymenopteran parasitoids associated with P. froggatti are
reviewed. An illustrated key to the 17 species encountered in the Adelaide region is
presented, along with notes on their biology, relationship with their host, and species
recorded from other parts of Australia.
Key Words: Phylacteophaga froggatti, Pergidae, parasitoids, hyperparasitoids,
Braconidae, Ichneumonidae, Chalcididac, Elasmidae, Encyrtidae, Eulophidae,
Eupelmidae, Pteromalidae.
Transactions of the Reyal Soctery ef 8, Aust, (9949), TIN), GU-ITs
BIOLOGY OF PHYLACTEOPHAGA FROGGATTI RIEK (HYMENOPTERA: PERGIDAE)
AND TTS PARASITOIDS IN SOUTH AUSTRALIA
by I. A. THUMLERT & A. D. AusTIN®
Sommary
THumberi, T, A. & Absyik, A, D, (1994) Biology of Pavlucteaphava froygartl Riek (Hymenoptent: Pergidie)
and its parasitoids in South Australia. Trans. A. Soe. S. Aust. 11B(2), 99-113, 31 May, 1994.
Informution iy presented on the biology of Plhylacreophaga frogeari Riek (the leat-blister sawfly), an emerging,
pest of plantanon and ornamental eucalypts in south-eastern Australia. The range of eucalypt species allacked ,
the extent of (ree damaye, seasonality, fecundity and longevily of FP fraggeit ure investigated for popnlations
ithe Adelaide revlon. The hymenopteran parasitmids associated with PB froggadti are reviewed, An illustrated
key to the 17 species encountered in the Adelaide region ty presented .alogy with potes on ther hialogy, celationship
with then host. and speees recorded frour other parts of Australia,
Koy Worps: Phylacreaphage frogear!, Pergidae, parasitoids. hyperpurasitoids. Braconidae, lehacumonidite,
Chalvididae, Ehumidae, Encyrtidae, Bulophidae. Bupelmidac, Pieronialidac
Introduction
Phylaciwzophaga frogeatti Riek, the leal-blister
sawlly, is 4 native leal-mining species that is generally
restricted in distribution to the south-eastern and south-
western parts of the continent (Riek 1955; Benson
1963; Farrell & New 1980; Curry 1981), Larvae teed
ong ninge of eucalypt species by mining the mesophyll
layer between the upper and lower epidermis of large
leaves. This pattern of feeding results in characteristic
oval sor elongate. brown blisters, which are most
common on the Jower branches of young trees (Farrell
& New 1980; Nuttall 1985). Heavy infestations of P
froggari{ cause substantial loss of photosynthetic area,
Tesulling, i stunted growth or death af trees. This
damage is of most concern where large nombers of
suplings exist in a confined area, such as cucalypt
plantations, ornamental eucaly:pts in parks and gardens,
and potted trees in qutside nurseries.
In 1985 PF frogeanti was accidentally introduced into
New Zealand where m has become a serious pest of
eucalypt forests and ornamental trees (Nuttall 1985;
Kay 1986). This has led to the importation and release
of two species of braconid wasp from Australia in an
attempt to control this sawfly biologically (Austin &
Fauldy 1989; Faulds 1990),
Given the pest status of Fh froegarty, relatively tittle
work has been undertaken on its biology or that af its
parasitoids, Other thun the work of Farrell & New
(1980), who provide a detailed account of some aspects
of the biology of P freggarti in the Melbourne area,
the species has received only anecdotal attention
(Froggatt 1899; anon. 1950; Riek 1955, 1970; Moore
1966; Curry 1981; Naumann 1983; Bungey [986).
Information on ifs panisitoids is even more limited.
* Deparunent ol Crop Protection, Waite Campus, University
of Adelaide. Glen Osmond, 5S. Aust. S064
Farrell & New (1980) record some observations on the
development and mide of parasitism for two species,
Bracon sp, and Cirrospilus sp. in the Melboume area.
while other workers have simply listed the species
reared (Moore 1966: Curry W981; Boutek [988). In
these studies no more than six species have been
recorded and, with the excephion of Boutek (1988),
the accuracy of species identifications 1s questionable.
More recently, Austin & Faulds (1989) have described
two species of Bracen that have been introduced inty
New Zealand, while Faulds (1990) has reported the
release and establishment of these species and
discussed their potential for controlling
Phylacteophaga.
The aim of the present study is to document more
fully the biology of PF fregeatti, focusing, on aspects
that have been largely neglected by previous workers.
Atthe same time, detailed information is presented on
the large parasitoid complex assocjated with P frogeatz
in the Adelaide region. An illustrated kev to their
indentification, as well as notes on their taxonomy.
biology as either primary or hyperparasitoids,
immature stages and seasonality are presented as a
prelude to future studies on the ecology und possible
control measures of this emerging eucalypt forest pest,
Materials and Methods
bighl sites
Rucalypt trees infested with larvae and pupac of B
Jreggatti were monilored al several sites in the Adelaide
region, viz., from the western coastal suburbs (Dover
Gardens), city parklands and adjacent suburbs (Scurt),
and Lobethal-Woodside area (Adelaide Hills), between
August 1990 and July 199), The first two sites comprise
mostly ornamental trees (immatures of numerous
species) in parks and along roadsides, while the latter
site is semi-rural and comprises both ornamental trees
KK) T A, THUMLERT & A, D AUSTIN
und naturally growing trees in. paddocks and along
roadside verges (mostly éucalyptuy camaldulensis, £-
eladecalva and &. leucoxylon). The chimate ot he first
two sites is very similar with average monthly
maximiom and minima temperatures of 23°C and
13°C, while the Lobethal-Woodside site has average
monthly maxins anil mini of WC and BC. The
average annual rainfall for these two areas is
subsiintially different (Adelaide Sk&Sonn; Lobethal
ARR).
Laboralary rearing:
Infested branches. mostly of £. canialdulensis. were
collected trom the field, placed in plastic bags and
relurned to the laboratory. They were placed in
conlainers OF waler al room tempeniture or constant
20°C und P froevart allowed to develop. Leayes with
larvae, pupye and/or developing parasitoids were back-
lit with an opheal libre light to determine the exact
stages present and observe their development, Most
parasitised and unparasitised pupae were separated into
5 crn dia, plastic containers and allowed to complete:
development. Once emerge, adult PB froggaiti and
parasitoids were kept alive by supplying them with &
synear of honey and water via a saturated cotton roll
pushed through the lid of a small glass vial. Virgin
fernale P froggatti were sometimes isolated trom mates
as soon as they emerged lo provide individuals: for
laboratory experiments.
Fecundity and longevity
Virgin female P froegar which had emerged over
a 24 hour period were held at constant 209°C with honey
and water supplied. Ten individuals, niudomly selected
cach day, were measured for body length. killed in 70%
EtOH, squashed on a microscope slide and the number
of lilly developed exes counted.
To determine the longevity of Po fregeari, adult
females were kept in 20 em diameter plasuc capes and
their survival monitored twice daily until al) wasps hud
died. Longevity was compared for J) virgin or mated
females, 2) with or without water and honey, ancl 34
at five lemperatures (10, 15, 20, 25, 30°C"), Ten femates
Unt bad emerged within a 12 hour peried were
rindoniy selected from stock cages and Lransterted int
experimental containers. Three replicates were run tor
each of the abave 20 treatments, Honey or water were
not supplied during mating to those which wen tate
the “no fod” ireatments. All cages were held at 12.12 fy
photoperiod in temperature cabinets with -F0.5°C
temperature ranges. The mean survival time for each
treatment Was calculated and plotted agaiast
temperiture, and the regressions analysed using, it
2-tailed F-test,
Taxonomy and scamming electron microscopy
‘Terminology for morphological structures generally
follow Gauld (1984) and Gauld & Bolton (1988) for
ichneumonid and braconii wasps and Bouéek (!088)
for chalcidoids. Where necessary, morphological
structlres referred to in the key are indicated on the
figures. Voucher material of all species have been
lodged in the Waite Cumpus inseet collection,
University of Adelaide,
Specimens for SEM examination were killed and
washed in a strong detergent solution before being
dehydrated in 100% EvOH and critical point dried in
an Emacape CPD 250. They were coated twice with
carbon and gold/palladium before being cxarnined
undera Phillips 505 Scanning Electron Microscopes
Results and Observations
Tatonemy of Phylacteophaga
The taxonomy ol Piylacteophaga species 15 confused
and requires urgent study, Nominally, only one species,
encalypti Froggatt, 14 recoynised which has becn
divided into four subspecies (Pe, eucalypi, Pe
Jrogearti Rick. P ¢. tlasmanica Riek and Pe, oevideny
Benson) based on colout and size (Benson 1963). At
least two of these subspecies (Pe, froggatti and Pe
accidens) oecur sympatrically and may therefore
represent distinct biological species (Naumann 1983).
However, of the several hundred specimens reared from
leaf nines by us during 1990-91 all belong to the one
colour form, that of Fe, froggarti Riek (ic. antennal
scape and pedicel anc scutellum ot female pale yellow
to wrange-yellow). Furthermore, mating of field-
collected material in the laboratory was always
successful. supporting the premise chat cither oaly one
species is present or one is dominant in the Adelaide
region. Because all recent! authors (e.g. Barrell & New
1980; Curry [¥@). Faulds 1990) have adopted species
level status for the subspecies 2 ¢, froggarti, this
upproach will he used here to save confusion,
Matiny
In the field, males were often observed perched on
the doryal surface of leaves with their body held at 45°
ubove the surface, in the vicinity of ovipositing or
resting females, or flying in small swarms of 5-10
individuals, close ld leaves where mating pairs were
perched, Copulation is strophandous, i-¢. male and
female are joined end-to-end with the mate genitalia
twisted through 180° (Gauld & Bolton 1988), In the
laboratory Mating took 2-3 minutes. During copulation
the femule stands with her wings tolded at rest, while
mules have their wings spread at about 45° and
penodically fan them. After uncoupling, the female
immediately moves away dnd the male remains in
place, tapping its-abdomen severg] times on the leaf
surface before fying off,
PUYLACTORGPHAGA FROGGY ANDITS #ARASITOLDS nl
Fecundity and avipasitieon
The number of exys carried by virgin females ranged
from S57 tw 87, while ege number within these lirnits
Was positively correlated will both age and size of
termales (t « O76and ¢ $0.66, P< 005) On landing,
female P frogeaiti move sideways over the surface of
a leat perpendicular to the oid vein, with this
movement continuing until a avipasiiion site is
chosen. The female saws into the leaf tissue tar 3-10
sec. then remains stationary for 3-4 nun while
oviposiang. Abts ime the body and anteanae are held
at an angle of about 45° tothe leat and to the mid vei,
possibly t@ position the cee so that the larva hatches
in the direetion cowards the apex of the leaf. Once
completed, the female rubs her wbdymen over the
pasition of (he egg spot three or four times, possibly
io seal the hole, and then moves away. either staying
on the same leaf or moving to-another leaf to oviposil
Egps.are penerally laid close ty the mid vei and along
the length of the leaf, but got in the distal 20-30 mm
The small egg spots, about 0.8. min ta digmigter. byron
raised globulous structures (¢ge-galls) on the surface
of the leaf, The number of egy Spots per leal ranied
front 7-170, with numbers over about 90 represeneny:
the ovipositions of more than one female,
Life-history stages
The morphology, colow aad size of the egg, larval
instars and pupa of P froggatti dre virtually identical
fo those described! by Farrell & New (980). Briefly,
the eae is fkattened and oval in shape, measuring
0.75 &.0.5 mim. There are five larval instars. in the
female and four in males, Tor the Adelaide population
females hid mean head capsule widths of 0.49 (1). 0.62
(1), 0.80 (11H, LOL dV) and £25 mm (V) (mn & 25
in cach cause), with males being virtually wdemtical to
Jemate instars I-IV. These sizes are almost identical
to those measured for the Melbourne population by
Farrell & New (1980), with the exception of female
inslar Vo which averaged 1.08 mun ut the latter study,
‘This diserepaney may be due to our selection of larvae
from non-overlapping mines, where there was lo
compelition for food between individuals. At room
temperature in the laboratory during: January ane
February (daily average about 23°C) larval
developyoent to cocoon Wnitation (ic. including the
prepupal period) look 15-18 days and the complcie lffe-
eyele about 30-35 days.
Pupalion is preceded by a prepupal stage.
distinguished from the fast larval instar by a liphter-
coloured head capsule. The prepupa moves away from
the tight perimeters of the mine imty a more central
position where it begins to spin a thin ovoid coenen,
Silk 14 fad down from mandibular glands as it swings
its head from side to side. The cocoon ay attached to
the mine floor and the leaf surface abowe by silk. ond
forms an annulus of creases ot the blister ossue. The
period trom cecoon initiation to adult emergence took
about eight days m the laboratory for both sexes, Two
to three days afler cocoon formation the prepupal
cuticle was shed revealing the cVe spots and appendages
of the developing popa. Ry the filth day the pupit
beeame pigmented, al which stage the scx of
indiwiduals could be determined by the coloyr of the
thor and presence of the female avipositar, the thorax
af males being black and that of females orange, The
pupa moulted on day sever ants the preadule stage.
whi¢h is characterised by having all the features of uw
fully formed adult. The body is completely pigmented
and the wings. which begin lo untold, are frequently
futined wrthin the mine to help them dry, On the eighth
day an oval-shaped hole ys cul in (he lop ob the mune,
und the wasp voids a yreen-erey meconium just belore
exiting or inmmediately atterwards. Ln the laboratory
upproximately 25% of adults died after failing 10
emerge from their mine, This mortality was hyshest
for leaves containing multiple individuals:
Longevity and overwintering
‘To determine the optumal holding conditions for P
frogeari in the laboratory und determine the potentual
for the species to overwinter as adults, longevity of
Wasps was Tieusured ab yarious temperatutes aid
combinations of mated or unmated females, with fond
and water supplied or not. ‘The regressions of mean
survival time against temperature were signifteant tor
the four combinations of virgin and mailed females,
witli und without food and water supplied (Fiz. 1). The
slope of the regression for lemales with food was
significantly different fram those without foow and
water for both virgin (F = 685; d., = 2.4, P<00s
Fig. Ta) und mated females (Fo © 474, df. = 2, 5:
P<005 — Fig, Ib), but mating did not affect the
survival of the fermales for either af the Lreatents with
(Rom 4.5; df. © 2.5; P>0.05) or without food und
water (F = | 6d, df) = 2, 6; PS 0.05). In Puzure Ib
Ihe data lor mitted firnales. with food and water supplied
al 10°C are excluded because of the breakdown of the
temperature cubinet part way through (he experiment,
The lungevity of Jemules was inercased when held at
lower temperatures and when water and honey were
supplied, but was not affected by malting. Ib rs mot
known whether both water and honey were (esponsible
for the increase in the survival rate of the females, as
these factors were not tested separately, Possibly only
water is required to prevent females trom desiccating,
and food ts ununportant for longevity. chough it may
he necessary for increased feeundity.
The longest that adult femules survived in the
Jaboralory at Lemperatupes below LO%C was 28 days
Hence, il seems unlikely that 2 frogearte overwinters
only as adults. A small number oFoceupied mines was
found on trees in coastal suburbs of Adelaide during
tune and July. but at no offer sites, Ht seers likely.
We T. A. THUMLERT & 4. D. AUSTIN
therefore, thal some pupae continue developing in
refuge areas until about mid winter and the subsequent
adults remain quiescent for the rest of winter before
ovipasiting into leaves in late August! or early
September (see Fig. 2).
Mine developinent
First-and 2nd instar larvae first mine towards the
distal end of the leaf following {he direction of the
venation, butas they grow with each subsequent moult,
the mine becomes progressively larger and circular or
oval m shape. As observed by previous authors (Farrell
& New 1980: Nuttall 985; Kay 1986). mines generally
follow along the mid-vein of the leaf and, when more
than one larva is present, their mines often merge so
(hat almost the whole leaf can beconie blotched. The
largest number of larvae observed to complete
development on a single large leaf was 21, so thal.any
remaining eges fron) the large numbers of ovipositions
(see ahiove) either died beforehand or were ealen by
the first emerged larvae. Leaves supporting, more than
about 10 larvae often resulted in Smaller adult wasps
compared with leaves supporting fewer larvae. Other
authors (Farrell & New 1980; Curry 198]) have
reported Phvlacteophaga as confining its development
only to the op surface of a leaf. This was confirmed
here lor cucalypt species that have differently textured
500
400
Mean Surval Time (bh)
‘5 10 15
20 25 ao 35
Temperature (°C)
Add (b)
200:
MWéan Survival Time (ht
5 10 15 20 25 30 35
Tampesrature (°C)
Fig. |. Longevity of @ PAvlucteophaga Jrogeani measured
as fhe mean survival Lime for three replicates of [0 waspy
(4S, E.) ut different temperatures) (a) far virgin 2 2 with
food and water supplied (IM) and without food and water
(J, andth) formated 2 2 with food and water supplied
(@) and without food and water (1).
dorsal and ventral surfaces, but for species with
jsolateral leaves (ie. identical surfaces on leaves
hanging verucally), mining was carried aul on eilher
side, sometimes resulting in individual larvae being
separated only by the internal venation of the leaf.
Damage 1 trees
The levels of damage varied between infested trees
und between individual sites, poxsibly because of
differences in the Lree species present, their size, age
and health Quantifying damage for these variables was
beyand the scope of this study but casual observations
in the field allow for some useful generalisations to
be made: Trees which supported large numbers of P
froggarti mines were sitnilar in several respects. they
were usually small (<4 m tall), were offen situated
on roadsides, carparks, parklands or isolated groups
in paddocks, and were usually sheltered to some degree
frou westerly winds. Possibly sheltered localities
prevent adult wasps from being blown away from their
host plant, since they wre not strony fiers, Trees that
became heavily damaged appeared healthy in the carly
stages ol aliack, in that they hid dense lush foliaye and
did not appear to be stressed. As reported hy Farrell
& New (1980), damage is more severe towards the byse
of trees. particularly on low hanging branches, The
upex region is usually not attacked, especially for trees
above 4-5 m in height. In some suburban areas, where
(here were many young frees of the same species (1-2 m
in heivht), damage often reached very high levels,
causing leaf shedding and sometimes death of saplings.
Well-cstablished trees (>8m high), were far less
frequently altached and, if so, were usually in Close
proximity to heavily infested young trees. Mines on
such trees were venerally confined ta low hanging
branches with semnu-imature leaves.
Host range
The range of tee species attacked by Phylacteaphaga
spp. wus compiled by reviewing previous studies. uy
well as surveying a wide range of trees in the Adelunle
revion during 1990-9|. Trees in open. situations,
nurseries and gardens were examined, with those in
(he laller two areas being used a contirnl apecies
indentification. The data presented in Table | shaw
that at least 27 eucalypt species arc susceptible to attack
both in Australia and New @ealand. with most of these
records being attributable to PR: frogeami. in Australia
the most widely infested species ure Enculyptuy
borryatdes, E. camaldilensis, EB, citriedora, &
cladacalyx, E. ficifolia, BE. wlobilus and EB. grandis
Also, several non-eucalypt species have heen reported
as bosts, but there is some doubt as lo the accuricy
of at least some of these accounts as they have mostly
not been confirmed since the initial reports.
PHYLACTOEOPHAGA FROGGATTI AND ITS PARASITOIDS 13
Parasiroid species
Over the period of this study 17 species of
hymenopteran parasitoids were associated with PR
Froggatt. A list of these species and summary of their
biology is presented in Table 2, and a key to their
identification given in the Appendix, Eleven species
were primary parasitoids of either the larval or pupal
stages. Three were confirmed as facultative
hyperpurasitoids (Elasmus australiensis Girault,
Cirraspilus margiscutellum (Girault) and Eupelmus
sp.), while three have not had their biology conformed
(Lartophagus sp., Pediobius sp, and Chrysonotemyia
sp. 2). The species in this latter group were not
positively reared from P. froggarti, but rather emerged
into rearing, containers with eucalypt leaves infested
with host larvae. There is a possibility that these species
had come from other leaf mining hosts, though none
of the latter was seen after careful inspection of the
leaves. lt is likely, therefore, that these (hree species
are parasitoids of P froggarti, but further study is
required to confirm this.
Seasonality of Phylacteophaga froggatti and its
parasitoids
Fortnightly field excursions from August 1990 to July
199) to collect P froggatti and parasitoids were used
to determine the approximate seasonality of the species
involved in the Adelaide region. Egg spots and early
mine initiation were first evident on trees in the
Adelaide Plains in late August 1990 and pupae and
preadult stages in mines were present up until late May
1991, A few occupied mines were found in June and
July on several trees, only in the western (coastal)
suburban areas, Although relative abundance of species
was not determined, it was clear from the number of
infested leaves and trees that P. froggatti is most
TABLE 1. List of tree species which have been recorded in the literanire and during this study as susceptible to damage
by Phylacteophaga spp. (sources of information are as follows: 1, Anon, [1950] and 2. Moore [1966], coastal N.S.W: 3,
Riek [1955] and 4, Farrell & New [1980], Melbourne region, Vie.) 5, this project [1990-91], Adelaide region, S.A.) 6, Curry
{1981}, Perth region, WA.; 7, Nutal [1985] and 8, Kay [1986], New Zealand; sources 3-3, 7.and § are for Phylacteophaga
froggattr)-
N.S.W.
Host Plant |
bo
Eucalyprus borryoides 4.) 26.5 + tyekepee
. camaldulensis .. JJ eet 2.
ee ae Te oe
. chtriadord . 6. - - eee. ce ey eee ile
» AMICI o:5- 6a section stejerejens tdhoicboabetete
. diversicolor oo. cra dp ewtetetete cotcick pete
Jivifolitt -a000.5 222524 -00acecuaes ae
e021) a
- gomphocephala 20.2 22 eee ee
QYANAIS oo cc eee teen eee
lehmamnit 2.02 a ee eee ee
leucoxylon
leucuxvlon MOS 6 ee ee
~ longifolia occ. 5s parpeletlecirettsa ule
macula. cu coe 0d wtataretent-¢ de paetete fod
_ macarthurti
marginala 0. . eee eee iit ines
LMONIOMANE 66 oe cece ene eeenes
PEW oe eee
%
AN SN ih
VA
. perrinidnd
- punctala -..---..-+-------
TOBUSID ole cena a tet hooeentastTe Male
TUAIS oy vee ee CHEE De ereee ot pike nite:
~ WOEEOMED Cece nese et eee cee 00 bejere
~ yideroxylon.......
WET ERLE 00.0 os OOS baa ls
PPP SMEARS MMe
AgOnIS flOXUOSA. 0. eee
Betula (Birch) -. . Vyigeie oe aot ee
Lophosiemon confertus (Tristania) 2. ~~ -
Quercus palustris oo... ccc us teesenvel
CF FQBUP Cojajeitee ett a wsteteteten GOO Odes
Recorded Phylacteophaga damage
Vic. S.A. W.A. NZ.
3 4 5 6 7 8
- i“ “ -
“ o ad
i - -
” - al
” i”
i
“ -
” ae -
-
“ -
-
fea
-
-
-
-—
“
-
-
- -
a
-
- -
-
-
-
- ’
-
-
104 T. A, THUMLERT & A. D. AUSTIN
TABLE 2. Summary of the relationship between Phylactcophaga froggatti and its parasitoids in the Adelaide region
(— = information not available; pa = preadult).
Stage of Solitary (S) Endoparasitoid (N) Primary (P)
Species of P. froggatti or or or
Parasitoid Family attacked Gregarious (G) Ectoparasitoid (C) Hyperparasitond (H)
Bracon confusus Braconidae larva; pupa S fe, P
Bracon
phylacteophagus Braconidae larva; pupa S Cc P
Paraphylax sp. Ichneumonidae pupa S Cc P
Brachymeria sp. Chalcididae pupa S N P
Elasmus australiensis Elasmidae pupa S Cc PH
Apleurotropis sp. Eulophidae larva G 2N P
Chrysonotomyia sp. | Eulophidae larva G Cc P
Chrysonotomyia sp. 2 Eulophidae - — — —
Cirrospilus
margiscutellum Eulophidae larva; pupa; pa S Cc P; H
Cirrospilus occipitis Eulophidae pupa s Cc P
Cirrospilus sp. 3 Eulophidae larva 8; G c P
Cirrospilus sp. 4 Eulophidae — Ss Cc P
Cirrospilus sp, 5 Eulophidae pupa s Cc P
Diaulomorpha sp. Eulophidae larva; pupa G Cc P
Pediobius sp. Eulophidae — — — -
Eupelmus sp. Eupelmidae pupa S Cc P; H
Lariophagus sp. Pteromalidae — = — —
1990 1991
A s Oo N D F M M J J
Phylacteophaga froggatti
Adelaide Plains eke -_- _ 4
Adelaide Hills
Parasitoid
Bracon confusus _
Bracon phylacteophagus —_—_OO
Paraphylax sp.
Brachymeria sp. _
Elasmus australiensis -_
Apleurotropis sp. —
Chrysonotomyia sp. 1 .
Chrysonotomyia sp. 2 .
Cirrospilus margiscutellum al
Cirrospilus occipitis te
Cirrospilus sp. 3 —
Cirrospilus sp. 4 —=—=—=—_—_—_——_———_
Cirrospilus sp. 5 °
Diaulomorpha sp. —_—___
Pediobius sp, *
Eupelmus sp.
Lariophagus sp. ad
Fig. 2. Seasonality of Phylacteophaga froggatti and its parasitoids in the Adelaide region for 1990-91, determined by the
presence of larvae and pupae in leafmines.
PUYLACTOLOPHAGA FROGGATT AND ITS PARASITOIDS KA
numergus at sites on the Adelaide Plains frorty about
early December to Jare March (Fig. 2). and drat there
are three, possibly four, peaks in numbers over the
pertod September to March. In the Lobethal-Woodside
ured ( Adelaide Hills), largest numbers occurred slightly
later, from about carly January to late Mareh, with egg
spots and mine initiajion not appearme unul later
Novernber. presumably because of the cooler average
lecpperatures there compared with the Adelaide Phuns.
Of the 17 species of parasitoids, six were encountered
over a period of two months or more (.c. Braco
piylacreophagus Austin, Cirraspilas margiscureltin,
C. ovcipitis Girault and Cirrespilus sp. 4, Kapelinis
sp. and Diaulomorpha sp.) (Pig. 2), and of these (it
descending order) ©. mareixentellum, B
plylacteaphagaus, Eupelmus sp. and Diavlemarphe sp
were reared in the largest numbers, No parasitised
hosts or parasitoid pupae were Mund in maiies during
July and August, though two adult female B
phylacteaphagus ank-one CL margivcarelhan were
located i old piiies during this tine, indicating that
these and possibly all parasitoids in the 2 fregeatti
comples overwitter quiescently as adults.
Levels of parasitism
Collections of mined larvae during January and
February 1991 enabled ussessment to be nade of the
levels of parasitism in the field at three sepurate sites,
viz,, Lobethal (Adelaide Hills), Sturt and Dover
Gardeos (Adelaide Plains) (Table 3), These data show
4 large range in the level of parasitism of P /reasyarti
(15.6 to 60.3%), and in all cases C marviseutellian was
by far the most important parasitoid, Other parasitoids
contributed O to 115% of total parasitism, of which
B phyvlacteophagus was most common, These date.
however, represent a minima estimate of percentage
parikwalisin, particularly for the Start sample. where F
frageaim were mostly represented by carly like-history
stages when collected, so that those sampled could have
beech parasitised if left in the field. Because ©
margiscutellum ws cupable of paralysing and leouling
on all stages of PB frovgain trom carly instars to the
preudult stage, un estimate of the maxinium potential
parasinsm was obtained by adding the measured level
ol parasitism with the proportion of hosts sull viable,
Biology of parasitotd species
Faouly lehnewumonidae
Paraphylax sp.
FIG. 5
There aire more thin SO recognised species ol
Puruphylax from Australia, the majority of which are
undescribed (Guuld #4). Where thew biology ts
known, they have heen recorded mostly as primary
ectoparasitojds and hyperparasitoids of lepidopteran
hosts, The sinvle male specimen reared here in March
199] ts an ectoparsinoid of P frogeary larvae. It belongs
to the covax species-group, and is probably the same
species as thal recorded by Gauld (1984) from FP
frogsatit iv south-eastem Australia. Being the only
ichncumonid in this purasitoid complex, Paraphylax
sp. cum be easily identified by is characteristic wing
vention,
Family Braconidae
Bracon confusus Austin & B plylacteophagus Austin
FIGS 3, 4,6
Both these specres ure solitary primacy
ecloparasitoids of third instar larvae to early stage
pupae of 2 fraysaut (Yable 2). B. pliylacteaphagus was
the More commonly encountered species of the two,
and was found assoviated with ils host for the same
period chat occupied mines were present in the field
(September to May), B confustis was collected for only
a short period during Deceinber-January, The biology
of these speeies has been discussed by Austin & Faulds
(1989) and Faulds (1990) and only observations
complementary to those given by these authors are
presented here. Pnor ty N89, fcherences to Bracen
associated with Phylacteophaga did not reengnise the
presence of two sympatrically occurring species.
B. confusus and B. phylactepphayus oviposit one egg,
rarely two, ina mine, either next to or onto the surface
oi the host. The host larva is paralysed at oviposition,
thereby allowing the Secure attachment of a newly
emerged Brucon larva in the absence of an active host
which normally whips its abdomen about when
disturbed. Late stage Braco: laryae are easily
distinguished from other parasitoids by their lange size.
dorsal ampullae and covering of long hairs, but they
wre identical io each other, Contrary (o the lindings
of Farrell & New (1980), most larvae which we gently
dislodged trom a host with a fine brush did not dic,
but rather successfully reattached themselves. When
feeding is completed, Bracen larvae normally move
away from their host before commencing cocoon
construction but occasionally pupate inside their host's
pupal cocoon, When spun away torr a host pupa,
Bracon cocoons often have loose Trass and moulted
cuticles imcOrporited into their silk matrix. They are
white in colour, ovoid in shape and measure about
IW * 13mm, The confined prepupa deposits a
meconium at one end of its cocoon and then bunehes
Lalo Ue Opposite end so that it eeccupies only alent
half of the internal space, Adults emerged in the
laboratory. 1-12 days atter the comniencernent of
cocoon construcvon, Adult B phivlacteophasuy Weld
at constant 15°C were still alive afler 68 days and
individuals were observed to feed on the honey uni
water provided
Wty TA
Along with Puraphylax sp., B confusus and B.
phylacteophagus can be easily recognised by their
relatively complete venation and large size (<3 thm),
und the species separated by their colour and
differences in pilosity.
Farnily Chaleididae
Brachymeria sp.
FIG. 7
This genus is 4 moderately large genus in Australia
comprising primary parasitowls and oblgatery
hyperparasitoids of the pupae of a greal range of
holometabolous insects, but particularly |epiduptera
(Boucek 1988). The species recorded here wus
reared from several pupae of P frogeatli during one
week in December 1990, Brachymeria sp. is casily
identified by its small robust form, enlarged hind
femur, elongate postmargimal vein, and densely
punctate dorsal mesosoma,
Family Eupelimidae
Eupelmus sp.
FIG, 8
This species Was generally reared as a solitary
primary parasitoid from the pupal stage of its host, but
on several occasions was also found fo be
hyperparasitic on smaller conspecific larvae and those
of several other undentified parasitoids. Mature larvac
can be identified from the other common parasitoids
in mines by the presence of dorsal ampullac and dense,
long hairs on the thoracic segments. Expelpris sp. was
reared from PF. froggattl trom mid summer to mid
autumn. Adults can he separated trom the other species
in (he complex by the presence of a greatly enlarged
mesopleuron, exposed ovipositor, and form of the wing
venation.
-
PTHUMLERT & A
1D AUSTIN
Family Elasmidae
Elasmuys australiensis Giraull
FIG, 9
This species was reared either as a solitary primary
cvluparasitoid oF hyperparasitoid of & froggattd during
December and January, Previous reports (Riek 1967;
Austin & Allen 1989; Allen 1990) have recorded this
species as beitg, gregariously hyperparasitic on
ichncumonid pupae assoerated with the Jepidopteran
defoliator Uraba lugens Walker. [1 can be readily
identified by its Jarge dise-shaped hind coxa, cream
voloured legs with black hairs and spines, and
distinctive criss-cross pattern of black hairs on the hind
tibia, See Rick (1967) for additional taxonomic
information and list of synonyms.
Family Ptreromatidae
Lariaphigus sp.
FIGS 10, U1
According to Boudek (1988), this small genus ts
represented jn Australia only by L. distinguendus
(Foerster}, a cosmopolitan parasitoid of beetles
associated with stored grain. The single specimen
reared here was removed in early February 1991 from
a P froegani jilne that did not contain any other
parasitoids orother potential hosts, Given (he paucity
of biological information available on-this genus ‘we
are unable to speculate further on this species
However, given that 4 dead P fragyatti larva was the
only other occupant of the mine, we have assumed that
Lariophagus sp. is either a primary or hyperparasitoid
on it. This small nondescript species is the only
pteromalid in the parasitoid Complex and it can be best
identified by the presence of five tarsal segments and
six funicle segments of the antenna.
4 , “ a,
Figs 3. 4+. Dorsal yiew of the head of 9 Bracen spps 3. Brecon phvlecteophayuy Austin, 4, Bracen cenfiisus Austity
Seales = 250 jan.
PHYLACTOEOPHAGA FROGGATT! AND ITS PAR ASITOIBS 107
Family Eulophidae
Peiltobius sp.
FIG. 12
Mernbers of (his genus are known (0 be primary and
hyperparasituius of eges. pupae and sometioes larvae
of various Insects, mainty of Lepidoptera, Coleoptera,
Diptera and other Hymenoptera, With several species
being reported as attacking leafminers (Boudcek
1988). An unidentified FPediabins species is
hyperparasitic on UL /ugens (Austin & Allen 1989:
Allen 1990), the latter occasionally appearing on the
same host plants as 2 fraggaii. However variation
in sculpturmg on the scutellum indicates that the
Pediobius associated with these two hosts are different
species, Previously, Po bruchicida (Rondani), a
gregariuus species (Boucek 1988), has been found as
a hyperpurusitoid on Be ophylacteuphagus in New
Zealand (Paulds 1990), and his species resembles
closely the single specimen recorded here, Pediobius
sp. Was reared in a comainer froma nuinber of leaves
mined in late Decemher 1990 Inspection of these leaves
revealed (he presence of no other potential hosts. Liki
Apleurorrapis sp. and Diaulomeanahe sp.. this species
can be identified from others in the camplex by the
presence of several long hairs on the scutum and
seutellui, afd the posterior expansion of (he
seutellum. dn addition to the characters in the key. it
can be separated from Aplenrotrapiy sp. by the absence
of a pusféro-mediul suleas on the scutum and trom
Diaulomerpha sp, by the very distinctive colour ol (he
latter species.
Apleurciropiy sp.
FIG. 13
This genus fas previously been revurded us emerginy:
from Fo frugeatd mines (Riek 1955, Farrell & New
1980), with Boucek (I988) listing 4. panetiperiniy
(Girault) as parasitising P encalypi: in (he Melbourme
area. Material reared over a ‘three week period in
December 1990 here confirms that this species is
gregarious, with the larvae probably being
endoparasilic, as holes were found in the body af the
host next to the parasitoid pupae. A single oie of a
late stage PP fregett? larva was found to contain 12
parasitord larvae and pupae, of which most completent
development in the Jaberatory. Compared wiih
Pediobius sp.. this species can be easily recagnised
by the presence of 4 large postero-medial sulcus on
Whe scubim (sce above),
Diaulomorpha sp-
FIGS 14, 15
This gregarious primary ecloparasitoid allacks the
larvae and pupae of FL froggani. with up to 10
parasitoids beg recorded froma single mine. Artoum
lemiperature most individuulsy i a Single mine emerged
within 13 days after the commencement of pupation,
Mature larvae ean be easily recognised from the other
common parasitoids in the complex. beeduse of their
pregarious behaviour and absence of dorsal ampullae.
Adults can be recognised by their metallic green-yold
colour, white legs afd fight brown antennae (see
contents under Pediohius sp.) An apparently
uadeseribed species belonging ia this genus has been
recorded fron Phvlacteaphaga mines in. Western
Australia and elsewhere in Australia (Boucek 1988),
and this may be the same species as reared here.
Chrysonoronivie sp, land sp. 2
FIGS 16, 17
According to Boudek (988), members of this
meusuii-sized werius of about 40 deseribed Australian
species, develop in the eggs ar young larvae of leaf-
inimng and gall-forming Diptera and Lepidoptera,
Virtually no biological (nforjation is available for any
Australian species, excep! lor one reared [rom Perthida
glyphopa Common (jarrah lea miner) in the warmer
regions of the Darling Ranges and coastal regions of
south-west Western Australia (Mazanee 1988), It is a
solitary endoparasitoid most frequently of the 2nd and
ard instr larvae,
Two species belonging to this genus have been
associated with Bo frogeatie im the Adelaide region.
Chrysunotomyla sp, | was reared as a gregarious
promary ecloparasitoid Gf a single late stave larva o
P fragvatti collected at Lobethal in March 1991, while
(wo specimens ol Chrywenotomyia sp. 2 were reared
only from mined leaves in February 1991 collected al
te Dover Gardens site, and therefore are only
tentatively assumed 10 be patasive on this host,
Although very small in size (about | mm in length),
these Iwo species ean be separated from the: other
culophid genera in the complex by the antennal funicle
being Only 2-segmented and the scutellum lacking a
Submedial groove, They can be sepurated from each
other by the form of the wing venation and their colour,
Cirrospailus margiscutellum (Girauiy)
FIGS 18, 19, 22
Cirraspilus ts a large and taxonamically difficult
genus of sinall often brightly patterned wasps which
are well-known primary and hyperparasitaids of
eucalypt leal-mining insects. Two described species
hive previolisly been peared fram Phylacreophaga spy,
vid, Co maruscurellam (Girwuly) trom Western
Australia. and 2 oecipiis Girault from NSW,
(Boucek 1988), while undelermined species haye been
reared from J! frogeum in Melbourne by Riek (1955)
and Barrell & New (1980) and im Western Australia by
T. A. THUMLERT & A. D. AUSTIN
108
PHYLACTOEOPHAGA FROGGATTI AND ITS: PARASITOIDS
OY
TAGLL 3, Morraliny of Phytacteophaga froggattt ay revealed by collections made during Jannary and February 1991, from
Loaleial where trees were in relatively early stages of ultack, and from Sturt and Dover Gurdens where trees Nad beet
keayily attacked during the preceeding weeky (— = intormation unavailable),
Lobethal
Total number of mines examined»... 4555 434
% already enierged = es on 45.0
Viuble Po froggarti
% larvae, pupae & preadulis. --.-- - 2. 13.8
Cause of Death Unknown
Se larvae und pupae... 6.50... 10,1
% preadulis... 222220 2222-2 ----- 10.6
% Total 20,7
Parasilinnt
%e parasitized by
Cirruspilus marvivcutellamt 5. - 13.3
% parasitized by Bravan spp. — 16
% parasitized by others . 0.2
% Total parasitism... 2... ; [5.6
% mix. potential parasitism ,..-..... 29.4
Curry (1981), Clearly, at least some Cirrespilus spp.
are widespread in Australia. Here we record C.
margixculellum trom the Adelaide region and. C.
ocetpitis from Perth (material in ANIC) for the first
time, indicating that these species are distributed across
the southern part of the continent, at least.
C. margiscutellum was by far the most commonly
reared parasitoid associated With PB frogeatti in ihe
Adelaide region (Table 3), and was present in mines
for nearly the whole time that host larvae and pupae
were available (late ‘September to late May). As shown
by Farrell & New (1980), laboratory observations
confirm i to develop as a solitary primary parasitoid
of third instar larvae to late stage pupae, us well being
hyperparasitic, in the Adelaide region, on the larvae
of Bracon spp., Eupelmus sp., Dieatlomarpha sp. and
small conspecific larvae occurring im the host mine.
When multiple ? freggatri were found in mines, with
some parasitised by Aracan spp., the latter appeared
to be preferentially parasitised by C. mrargiscutel/um
rather than unparasitised primary hosts.
Usually only one Cirrespilas larva was found feeding.
on a host and generally i did not confine iis feeding
to one feeding site, as indicated by numerous dark
melanised spots found over the host's integument. In
the laboratory, ©. margiscutellam held at roam
temperature took 16-17 days to complete development,
once emerged from the egg, Mature Cirrospilus larvae
Collection Sites
Sturt Dover Gardens
899 410
0 12
79.6 63
3.8 21)
4.4 60).7
0 it}
11 i
16.6 60.7
96.2 67.0
could be recognised from those of the other common
parasiloids in the complex by the absence of long setae
over the body and the presence of ampullae on both
the dorsal and ventral surfaces. The larvae of different
species of Cirraspilus could not be separated. Adults
of Cirrospiluy spp. can be distinguished from other
eulophids in the complex by the presence of a
Submedial groove on the scutellum, submarginal vein
with at Jeast two brisiles. and their metallic and/or
striking colour patterns. C. margiscutellum is the only
member of the genus in this Complex to have the face
with transverse black bands.
C. aceipitis Girault & Cirrospilus sp. 3-5
FIGS 20, 21, 23-26
Four other species of Cirraspilus were reared from
P. froggatti mines, all of which are solitary primary
ectoparasitoids, with the exception. af Cirraspilus sp.
3 which was found to be gregarious. C. veeipitis and
sp. 5 were observed to feed on host pupae, and sp.
3 and sp, 4 on the larval stages. C. occipitis and sp.
4 were found in mines over the suromer months, early
December to late March, and early January ta late
March, respectively, while sp. 3 and sp. 5 were
recorded from only a few samples in one week in early
March and late March, respectively. The live species
encountered here can be separated relatively easily by
their distinctive Colour patterns.
Figs 3-9, 5-6, Fore and hind wings; 5. Paraphylax sp.. 6. Bracvon confiusus Austin, 9; 7-9. Lateral view of whole body:
7, Brachymeria sp.: 8, Eupelmus sp; 9, Elasmus australiensis Girault, Scales: Figs 5.6 = 1.0 mim; Figs 7-9 = 0.5 mm.
Abbreviations: bu = bullae; he = hind coxa; hf = hind temur, mp = mesopleuron; pp = prepectus, pv. = postmarginal
vein; &V¥ = stigmal vein.
HO
T. A, THUMLERT & A. D. AUSTIN
Figs 10-17. 10-11, Lariophagus sp.: 10, antenna; I, hind leg; 12, Pediobius sp., dorsal view of mesosoma and metasoma;
13, Apleurotropis sp., dorsal view of scutellum, propodeum and metasoma; 14-15, Diaulomorpha sp,: 14, antenna; 15,
hind leg; 16-17, Fore wings: 16, Chrysonotomyia sp. 1, 17, Chrysonotomyia sp. 2. Scales: Figs 10, 14 = 0.5 mm; Figs
ll, 15 = 0.5 mm; Fig. 12 = 0.5 mm; Fig. 13 = 250 wm, Fig. 16 = 200 wm; Fig. 17 = 0.5 mm. Abbreviations: cl = clava;
cv = cubital vein; f = funicular segments; pe = pedicel; sc = scape; smy = submarginal vein.
PHYLACTOEOPHAGA FROGGATT! AND ITS PARASITOIDS il
Figs 18-26. 18-19, Cirrespilus margiscuiellum (Girault): 18, fore wing; 19, anterior view of head showing transverse banding;
20-21, antennae: 20, Cirrospilus occipitis Girault; 21, Cirrospilus sp. 4; 22-25, Cirraspilus spp., dorsal view of mesosoma
showing colour pattern: 22, Cirrospilus margiscutellum (Girault); 23, Cirrespilus vecipitis Giraull; 24, Cirrospilus sp.
3: 25, Cirrospilus sp. 4; 26, Cirrospilus sp. 5, dorsal view of mesosoma and metasoma showing colour pattern. Scales:
Figs 18 19 = 0.5 mm; Figs 20, 21 = 200 jm: Figs 22, 23 = 0.5 mm: Figs 24, 25 = 250 pm; Fig. 26 = 0.5 mm.
Abbreviation; sg = submedial groove on scutellum,
12
T A. THUMLERT & A. 1. AUSTIN
Key fo the parasitoids of Phylacteophaga froepath
2
uo
in the Adelaide region (based on females}
Fore wing wihomore than one enclosed vel) (Figs 5, fy:
Prepertus absent... - 2... 2
Fore wing with no more than one 2 enclosed cell (Fign
7, 10-18); prepectus usually present (Figs & 9) seniotines
absent (Pig. 7) (Chaleidoidew) ~~. 2... 4
Fore wing with vein 2meu present; ‘hind wing
with vein em niweting Rs after Rs diverging from
Se + R (Fig. 5) (Lohneumenidae) | fore wing with twe
bullae if) 2ry-cu (Fig, 5), mesosculum smooth ane
UNSCHIPLUTBt] pyc ee eee rey Faraphviive spt
Fore wing with vein 2meu absenl: hind wing with vein
rm meeting Ks before Rs diverging Inm Sc +R
(Fig. 4) (Braeonidaey : 4
Vertex, uccipur and temples. ivtostly hairless except
lur row of short hairs around margin of eyes und
postenor vertex (Fig. 3), propadeum black
Bravvin phylucteuphoens Austin
Vertex veciput and temples sparsely covered wath
long hairs (Fic. 4h propadeum ormnge-yellow
» Braver vanfasuw Austin
Find leraur Colirpea and with (eetls on-yentnl edge (Fig,
7); prepectus very small, virtually absent (Chalewdidue)
|malar suture diétinet: postmarginal vein longer thin
stigenal yer (Pig. 7). propoeduim with rough alveolate
se dipruring| _ _Brochymeria sp,
Hind formar not enlarged a and without teeth (Figs 8, WN),
prepectus distiner tFigs 8, 9).
. Mesopleuron greatly enlarged, conver, ¢, developed into
a large Undivided shield (Fig 4) [postmarginal vein
slightly shorter (han sligmal vein; ovipositer protruding
from posterior ineaoma, shealhs black with addle
third white (Pig. Bp... .. —Eupelmus sp.
Mesopleuron not greatly enlarged (Fig. Wy... of
. Hind coxa developed as a large flat dise (Fig. 9)) bind
ubin wilh seac tanning distinct criss: cross panier, fore
Wing nao, pustmanginal vein mach lanper thant stygrntl
vein (Fig. 9) (Elesmidae) [body dark, tezula and legs
pale except for bind coxae|Flasiitks australicriviy Giraull
Mind cond mot as large flat dise (Fig. 1) hind tibial
hiirs not forming eriss-cnoss pattem; fore wing relatively
brovid (Pigs (IB) eee 7
. Hind tarsi 5-seginented (Fig. ty, antes without dstinet
funeulur segments and clava (Big. 10) (Pterormalidae)
head round im anterior view; basal tind of fore wing
virtually deverd of hairs]. --.- , _. Larivphaguy sp.
Hind tars} 4-seamented (Fig, 15); antenna wath distiner
funicular seyrneaty and clave (Figs 44, 20, 21
(Hulophidae) a i LR
. Dorsal mesosoma with long ‘stout hatirs (Fig. ‘W);
metisoma distinevy petiolate (Figs 12, 13), body black
and shiay- ..- oe ed
If dorsal mesasoma with hairs then most ly fine and short;
Melasoina sessile, or pet(ole less Conspicuous (Big. 26);
fody colour -vartable .. 6... yp eee ery 10
Seutellum lengittidinally strigose with excepunn of
longitudinal medial line which is smooth (Fig, 12); Tt
of metasoma with anterior Mange (Fig 12); legs all black
except proximal 4 wiikn segments of md and bind lees
which are while 2-0... yy Pedinbiies sp
Scuteilum with coriaceous sculpluriag (Fi. 13); TL of
mélasoma without (hinge (Pig, 13); all legs distal to coxae
pate in colour eee yy Aplewrorrmpis sp
Funicle of antennae J-segoented (Fig. 14) [body metallic
Breen-gold ip colour, cll legs distal (a coxae white; becky
about 2 men mm length} . Biaulamerpha sp,
Funicle ot dnteomie 2 segmented (Miga 20, 21), 1
H, Scutelliu withour submedial grooves: submmargimal vein
swith 2 dorsal bristles (Figs 16, 17): body less than 1 (it
in length (Chryvenotongia Ashmead)... 6.00.) 12
Seurellun) with submedial grooves (Figs 22, 73):
submarginal vein with more than 2 dorsal bristles (Fig
18): body greater than | mm in Jength elses
Wexlwood) 260. cs yi y cee ees bene
12 Cuil vein of fore wing Indicated by 4 ra
tow Of hairs (Fig. 16); stigmal vein distine( and infuseate
around distal emch sep ba mesosomea.and head metallic:
ereen . 6.64) Chrysencaronvia sp. 2
Cubital vein of w ing fen indicated (Fig. 17); stigmal yein
short und not infuscale (Fig. 17); mesosome and head
non-metallic and dark in colourChrysenoramvid sp |
2 Clava of anlenna continuous wath funicular segments
(Fiz. 20) 0.00.0... potty tN
Clava separated fron FaniClifar ‘segments by distinct
constriction between segments (Fig, 21). lt
14 Stigmal vein infuscate around distal end, Sornesinnes onl
faintly {Fie. (8): vertex with short stow) black hairs
(Fig. 19)... OA
Sugmal vein without infuscation ‘distally’ Vertex with only
fine pale hairs [Mesosonta with yellow and metallic green
markings Fig. 23)] ,,. -Cirrospiluy oecipitis Girault
(8 Darsal mesosoma metallic green-blue with slight yellow
markings in posterior half (Pig. 22), surface with
relicolare sculpluring; face with lransverse durk
stripes (Fig. 19); pronotum shorter than seu
(Fig 22)... Croraypilus margiscatellam (Giraulo
Pronatum yellow wilh J longitudinal blaek bands, rest
of mesosoma oringe wilh black markings (Pig. 24):
surface with coreacious sculpturing: pronatum nearly
as long as seulum....-- e Cirraspilus sp. 4
16 Metagama rounded i in dorsal View. pronotum yellow-
orange. fest of dorsal mesosema orange with black
markings (Pig, 26)... ——- _Cirrnspillis sp, A
Metasoma — brentely. ‘elongate, pointed POSteTiOr]y,
dorsal mesosonm yellow with black markings (Pip.
45) aoe ee, _ Cirrespilus sp. 4
Discussion
This study confirms a number of aspeets of the
hiulogy al 2 fraygarni first reported by Farrell & New
(J980), as Well as doeurienting the fecundily ans
longevity of adults, and details of its parasitoid complex
for the first time. Previously, this insect had been
recorded only as a sporadic pest of eucalypts in the
Adelaide region and then mostly on ornamental trees
However, the status of P fruggarti as an emerging
serious pest both in Australia and New Zealand has
increased dramarically over the last lew years and this
is correlated with a corresponding expansion in native
woodlor plantings in south-eastern Australia. Indeed.
the Engineering & Water Supply Department of Soath
Australia has rated P froggatr as one of the most
important pests at its Bolivar woodlot (pers, comm,
S. Shaw).
Dunny this study we recorded newrly three times
ihe number of parastinid species associated with
frogeani as any reported by previous authors (Riek
1955; Farrell & New (980; Curry 198). The size of
the parasitoid complex in the Adelaide region, however,
is Unlikely to be significantly larger than elsewhere in
PHYLACTUBUPHAGA FROGUATTS AND ITS PARASTTOIDS "a
Australia. Rather, the concentration of this study on
rearing parasitoids and its longer duration has meant
that several sarer species have been colleeted, Also,
the tixonomy of the groups involved is now better
Known (see in particular Boucek 1988 and Austin &
Faulds 1989). so that species can be more easily ond
reliahly identified. However, the makeup of the
parasitoid complex and relative abundance of species
does seem to differ belween regions. For instince. in
the Adelaide region Cirrospilay sp. 1 was far more
abundant than 8. phylacteophagus, while in Melbourne
the Iailer species was more abundant (Farrell & New
980), In Western Australia Curry (O81) reared six
parasitoids froay Phylacteaphuga (three Cirrosytlas
spp., Elasmus sp. Eupelmus sp., an unnamed culophid
und Apanieles sp.) but no Bracon, Austin & Paulas
(1989) bave pointed out thar the record af Aparteles
ap. must be erroneous. as members of this penus anil
all mictogastrine braconids ure endoparasnoids of
lepidopteran larvae. The only major parasitoid nexan
not recorded in this study is Jsephuoides westralianis
Girault, a pleromalid, which has been reared from F
frogvamt only in Western Austratia (Boudek 1988).
Farrel) & New (1980) suspected the Cirraspilus sp
reared by them 1 Melboume to be polyphagous and
to use alternative hosts at different times of the year
() breed continuously. Although we did mol pear
parasitoids from other eucalypt leal minings in the
Adelaide region, the Ract the taulliple Crrrespilus
species have often been recorded from other hosts dacs
provide some indircet evidence that menibers of Hus
paTasitoid genus are polyphagous (pers, comm. N,
Gough, [. D. Naumann).
This study should provide a solid basis for more
detailed work on the ecology of FB froegain,
particularly the factors that influence its abundance,
medns of overwintering, and the susceptibility of its
¢ucalypt species. These and other factors will be
iportant in understanding how A frogger develops
as a pest and how best it can be controlled. Programs
simed at limuing the danvage caused by this sawlly
Will need to take ite account the mortality caused by
the above parasitoids. In this respect future research
might profitably examine both seasonal and regiunal
differences in rates Of parasitism and the species
involved.
Acknowledgments
We thank Dr Jolin LaSalle, Internatyonal Institute ol
Entomology. London tor cantirming the identity of the
chaleidoid parasitoids, and Mr Paul Dangerfield for
the line drawings, electron micrographs, and his.
comments on the tanusery pt,
References
Ason (950) The Jeaf blister sawlly — Phylacieuphaya
encalyp. The Avricultiral Gazette 470 471.
ALLIEN, GR. (1990) Uraba lugeas Walker (Lepidoptera:
Nogtuidae) larval survival and parasitoid biology in the
field in South Australta. J Ause ear Soe, 29 401-312,
Ausiin, A.D. & AtLais, G. R. (1989) Porasitoids of L'vaba
fagens Walker (Lepidoptera, Nectiktae) in South Australia,
wilh deseriplion of two new species of Braconidae. Trans:
R. Sue & dase V3, 169-184
& Pautos W. (1989) Two new Australian species
of Bravon & (Ay menopten Braconidae) parasitic or
Phylacteuphaga spp, (Hymenoptera: Pergidae). 0 Ais ent,
Soc. 28. 207-213,
Benson, R. B. (1963) Some new Western Australian saw !Ties
ot the Huryinae and Phylacteophaginae (Hy menoprers’
Pergidite). J. Rey Soe. Wo Aust, 46, 81-84,
BuNGey, R. (1986) Insect pests, pp. 233-24). fn Wallace.
WR CRE. "The Bealogy of the Porcats and Woodlands
of Sour Austrailia.” (Govt. Printer, Adelaide).
Bouter. 2% (J988) “Australasian ~~ Chaleiduidea
Clymenbptert). A Biosystemutic Revision of Gener ot
FPoarteen Families, with a Reclassification of Species” (CAB
International, Williford),
Curry, J. §. (81) Native wasps hit introduved tree post. /
Agric W Auyt 22, 61-62.
Fagkrett, GS. & New, TR. (1980) Some aspects at the
hiology of the Gucalyptmining sawlly Phylacteapteapa
Jrongani Rick (Hy menoplera: Perpidae). lust, J Zee! 28,
83-00),
Fabios, W. (1990) Introduction into New Zealand of Breceoe
phylacieophagus, a tocontrol agent of Phylacteaphape
JSroggaui, Fucalyprus leat muning sawtly N42 Fioress
Sei 2), 54-04
Froeoatr. W. W. (1899) 4 new genus und species of sawlly,
Prac. Linn, See. NSW 34, 130-\34.
Gavip. [, D, (984) “An introduction to the Ichneumonidae
of Australia” (British Museum [Natural History} Landon).
& Boiron, B. (Eds) (1988) “The Hymenoptera’.
(British Muscuin [Natural History|, London and Oxtord
University Press, Oxtord).
Kay. MK (1986) Phylapteophava froeear Rick
(Itymenoptera: Pergidae) Buealyprics leat mining sawfly
Forest and Timber Insects in New Zealand. No 4.
MAdANEC, Z, (1988) Tairnature stages and dite history ot
Chrysenviomyia sp. (Hymenoptera: Eulophidae), a
parasioul of the jarrah leafminer, 2 ausr enn See 27,
279 292.
Moors, K, M, (1966) Observatians on some Australian
forest insects 22. Notes on some Australian leaf-miners.
Aust. Zowl. 1, 303-349,
NAUMANN, TF, D. (1983) A neW genus ot Phylacleophaginae
from. northern Australiu with a key to the penera of the
subfamily (Hymenoptera! Tenthredinoidea! Pergidae). J.
Aust. ent. Sue, 22, 237-242.
NurraLy, M. J. (1985) New inseet pest atracks cucalypts
Phylactevphaga jroggalti (Hymenoptera; Pergidue), The
New Zealutnd Harmer 106, 124-5,
Riek, E. F (1955) Australian lovt-niining sawities of the
genus Phylacteophaga (Hymenoptera: Tenthredinoidea).
Aus I foal 3, 95-98.
(1967) Australian Hymenoptera Chalcidoiles farnily
Eulophidae, subfamily Elasmidac. Jbid, 1S, 145-99.
(1970) A redefinition of the subfamily
Phylaecteophaginge will deseriphon of a sew penus and
species (Hymenopters: Symplivta. Pergidae), J. aust. enti,
Soe, % 25-218.
DESCRIPTION OF THREE NEW BARNACLES OF THE GENUS
ELMINIUS (CIRRIPEDIA : THORACICA) FROM
SOUTH AUSTRALIA, WITH A KEY TO SPECIES OF
THE ELMINIINAE
By D. E. BAy.iss*
Summary
Bayliss, D. E. (1994) Description of three new barnacles of the genus Elminius
(Cirripedia : Thoracica) from South Australia, with a key to the Elminiinae. Trans. R.
Soc. 8. Aust. 118(2), 115-124, 31 May, 1994,
Three new species of intertidal barnacles from South Australia are described.
Elminius flindersi sp. nov. is a comparatively large species, grey to white in colour
which favours habitats with strong water movement although not direct wave action.
It has a long hatchet shaped tergum with a centrally localised fold. In comparison with
congeners the shell is strong and the cirri are broad and robust. E. placidus sp. nov. is
a smaller and more fragile species found in very sheltered localities, including
mangroves in the Spencer Gulf. It can be distinguished by its banded shell and wedge-
shaped tergum. The body and cirri are very elongated. E. erubescens sp. nov. is
common in the high intertidal zone in the Adelaide region. It is easily distinguished
by the reddish coloration of its translucent shell which gives a dark purple appearance
on rock.
It is suggested that South Australia has Eliminiinae distinct from those of the Eastern
States of Australia.
Key Words: Cirripedia, Elminius flindersi sp. nov., Elminius placidus sp. nov.,
Elminius erubescens sp. nov., Elminius modestus, Elminius adelaidae, Elminius
covertus, intertidal, South Australia, taxonomy.
Transactions of the Royal Society of 8. Aust. (1994), W8(2), 115-124.
DESCRIPTION OF THREE NEW BARNACLES OF THE GENUS ELMINIUS
(CIRRIPEDIA : THORACICA) FROM SOUTH AUSTRALIA, WITH A KEY TO
SPECIES OF THE ELMINIINAE
by D. E. BAYLIss*
Summary
Bayiss, D. E. (1994) Description of three new barnacles of the genus Elminius (Cirripedia : Thoracica) from
South Australia, with a key to the Elminiinae. Trans. R. Soc. S. Aust. 118(2), 15-124, 31 May, 1994.
Three new species of intertidal barnacles from South Australia are described. Elminius flindersi sp. nov. is
a comparatively large species, grey to white in colour which favours habitats with strong water movement although
not direct wave action. It has a long hatchet shaped tergum with a centrally localised fold. In comparison with
congeners the shell is strong and the cirri are broad and robust. E. placidus sp. nov. is. a smaller and more fragile
species found in very sheltered localities, including mangroves in the Spencer Gulf. It can be distinguished by
its banded shell and wedge-shaped tergum. The body and cirri are very elongated. E. erubescens sp. nov. is common
in the high intertidal zone in the Adelaide region, It is easily distinguished by the reddish coloration of its translucent
shell which gives a dark purple appearance on rock.
It is suggested that South Australia has Elminiinae distinct from those of the Eastern States of Australia.
Key Worns: Cirripedia, Elminius flindersi sp. nov., Elminius placidus sp. nov., Elminius erubescens sp. nov.,
Elminius modestus, Elminius adelaidae, Elminius covertus, intertidal, South Australia, taxonomy.
Introduction
In South Australia barnacles belonging to the genus
Elminius are highly abundant (Womersley & Edmonds
1958; Hutchings & Recher 1982; Bayliss 1982). Until
recently, they were classified as Elminius modestus
Darwin, as were those found in eastern Australia,
Western Australia and New Zealand. This situation
arose from the very broad definition of this species
obtained from a combined examination of Darwin's
(1854) original description and illustrations with those
of Pope’s (1945) paper.
Foster (1980) suggested that Pope's description
confused a new species with E. modestus as known
in New Zealand (Moore 1944; Morton & Miller 1968;
Foster 1978) and that Darwin's original material may
also have been a combination of these two species. The
new species, £. covertus, was subsequently described
by Foster (1982) together with a six-plated barnacle,
Hexaminius popeiana, A new subfamily, Elminiinae,
was proposed to accommodate them. Studies of larvae
by Egan & Anderson (1985) support the establishment
of this subfamily.
Foster (1982) furthermore suggested that E. modestus
was a New Zealand species which had been introduced
into Australia by fouling on shipping, probably in the
nineteenth century; E. covertus was, therefore, the only
known endemic Australian species. Bayliss (1988)
described another species, E. adelaidae, which is
abundant in the Adelaide region, and suggested that
* Kathleen Lumley College, 51 North
Adelaide, South Australia 5006
Current address: 7 Thomas Street, Kingsgrove, N.S.W. 2208
Finniss Street,
E. covertus was not found in South Australia although
E. modestus was present. Jones (1990) has identified
both E. covertus and E. modestus from southern
Western Australia.
There is little fossil material, but Buckeridge (1982)
has described E. chapronierei from the lower Miocene
in Victoria and E. pomahakensis from the upper
Oligocene in New Zealand (Buckeridge 1984) and
suggested the subfamily originated in south-east
Australia, He proposed a two-phased migration from
Australia to New Zealand and South America, firstly
in the Oligocene for Elminius with a spur on the tergum
and subsequently, in the Cenozoic, for E/minius lacking
a spur.
This paper describes three new species of Elmunius
which, with E. modestus and E. adelaidae, brings to
tive the number of species of this genus found in South
Australia.
Systematics
Suborder Balanomorpha Pilsbry, 1916
Superfamily Balanoidea Leach, 1817
Family Archaeobalanidae Newman and Ross, 1976
Subfamily Elminiinae Foster, 1982
Genus Elminius Leach, 1825
Type species Elminius kingii Gray, 1831
Elminius flindersi sp. nov.
FIGS 1, 3
Holotype: SAM C4242, on iron pilings of ferry jetty,
Penneshaw, Kangaroo Island, South Australia;
35°44'S, 137°57'E; D. Bayliss, 21.vii.1992; dissected
(partially).
16 D. E. BAYLISS
Fig. 1. Elminius jlindersi sp. noy. holotype. A. External view of shell; B. inner view of shell; C, scutum, internal view:
D. tergum, internal view; E. scutum, external view; F. tergum, external view; G, body; H. labrum; I, mandible; J. maxillule;
K-M. cirri L-HII; N. middle segment, posterior ramus, cirrus VI.
THREE NEW BARNACLES FROM SOUTH AUSTRALIA U7
Paratypes: SAM C4243, WAM 727-92, D. Bay iss,
21Lvit.1992, same locality as holotype.
Description of holotype: Shell (Figs 1A, 1B): conical,
white lo grey, opaque. Purielies folded. Orifice
pentagonal in outline, width 4 length, carimal sides
almost twice as long as lateral sides. rostral side
straight, Basal outline sinuous, deeply undulating. Alae
wider than radii with less oblique summits, Radii with
oblique summits, narrow, Shell 10mm in basal
diameter, 9.5mm in width, 4mm in height
Opercula (Figs 1C, 1D, TE, 1B); Solid white to grey
in colour. Seutum longer than high, basal margin
slightly convex with slight upward turning al tergal
corner, crests for depressor muscles absent, articular
ridge, articular furrow moderately developed. adductor
ridge not apparent, adduetor pit faint. externally growth
ridges present.
Tergum hatchet shaped, vertical articular ridge
folding inward to. form centrally limited fold, articular
murgin gently curving in apical region From carina!
margin, becoming parallel with basal margin to form
long garrow handle, basal margin curying sharply,
almost at right angles, to form projection with carimal
maczin, projection wilh prominent, deeply curved
crests tor tergal depressor muscles, spur Confluent with
basiscutal angle,
Body (Pig. 1G): prosoma white, broad. rounded
Cirri light reddish brown in living specimens, cirri 1,
Iland I slightly darker than ci LV, V and VI, colour
lost on préservation.
Mouthparts (Figs TH-IJ): Labrum with three teeth
and setulae, on each side of central notch, Mandible
with live teeth, lower short pectinate edge terminating
in short curved spine. Maxillule with two large spines
above notch, five smaller spines in notch, four large
spines. below notch, smaller spines at lower angle.
Cirri (Figs IK-IN): Cirrus T with anterior ramus
* 1.5 length of posterior ramus, Segments broad,
slightly protuberant, Anterior ramus of cirrus IT slightly
longer than posterior ramus, segments slightly
proluberant anteriorly. Cirrus 17 with anterior ramus
slightly longer than posterior ramus, segments slightly
protuberant anteriorly; setae on both rumi extremely
long, stout pectinate setae on six distal segments of
posterior ramus. Cirri TV to VJ all subequal in length.
segments with three large parrs, two small pairs of setae
on anterior lace, small proximal pair of setae between
segments. Number of segments in rami of cirri -VI
shown in Table |.
Penis: Penis.as long as cirrus VI, setose, basidersal
point absent,
ariation: Shell can be tubular, comical or Aattened,
Parieties can be smooth or undulating with variable
number of longitudinal folds. Maximum basal diameter
Vim. Colour varies from white to light grey,
Etymology: The species name refers to the
Flindersian biogeographic region.
Comparison with other species: Elminius flindersr
is larger and more robust than other members of the
genus found in Australia. The shell is thicker and
comparatively strong. The opercular plates are opaque
rather than translucent, The body is broad and rounded
with broad cirri.
E. flindersi cannot be reliably distinguished from. £.
modesius on the basis of external shell appearance. The
opercular plales are. however, quite different, The
fergum (Fig. LD, IF) is hatchet shaped with a long
handle and an articular furrow which is. restricted to
the central portion,
E. modestus (Fig. 2B,.2D) basa deep articular fold
“running from the apical end to almost the basiscutal
angle. The articular margin, Unless Worn, is straight,
The scutum (Fig. 2C) has a grey band, but this feature
is not always apparent.
TABLE 1. Cirral counts, Elininius (indersi ay. ate fanterior
ramus first),
Basal diameter
(mm) Cirrus
‘ I i lv Vv Vi
10.0 (holotype) 11,7 10,9 12.11 22.20 24.25 26.27
5.0) 12,6 LOL 12.01 1817 2L,20-23,22
80 12,7 V1,0P 13.73 21,20 23,24 26,25
80) 12,7) 10,9 13,13 23,23 27,27 27,28
10.0 17 11,10 13,11 22,19 24,24 23,23
lines
Fig. 2. Elminius meidesiuy Darwin. A, sculum, internal view;
B. tergum. internal view; C, scutum, external view; D,
Jergum, external view,
HA DB, bh BAYLISS
EL mmrlesins also can be distinguished from other
embers of the genus im lacking pectinate sete on the
pnsteriur ramus of cirrus Of, The apercutar Mapes are
pure white wilh a grey band at che rostral end und a
small orange dot centrally. &. flindery? Has a dark
brown band wt the rostral end and another brown band
ventrally on flaps which are dull write
&. adelaidge (Bayliss 988 Tig 2B) has a very small
articular furrow, which imiay not be visible if the tengun
rs viewed directly from ebove. The shell is much
thinoer_translucent, and is light brown. The opercutir
fips are, however, ino close tw &. ffinderst in
appeataice to be liseful for identification.
Habitat, Flinius flindersi js found tm the intertidal
yore in waters sheltered from direet wave action, ft
1s found to habitats with stronger water Haw snd rere
turbulence than other members of the germus in Seuth
Australie, Inthe gull regiogis, where wave impact is
diminished. ft occurs On exposed rocks. di grews in
. flindersi sp. nov-
. placidus sp_ nov. :
. efubescens sp. nov. |.
large -sives on jetlies although i 1s nul found an surfaces
faving directly inty waves at more exposed Inculities..
Outside the gulf regions it is found in habitats protected
from oceanic waves such as in boal enclosures behind
breakwaters.
Senlemenr occurs on a wide variety of Surlaces
including roeks. cement, wood, nietal and -plastie. I
worurcly found jn janproves Where Water HoW Is
gentle. AY Port Pirie ik eau be found on rocks in the
strongly- flowing sections uf the Hdal nven but nal im
nearhy mungraves,
Distrikunon: Elminins flinderst is widespread uw
South Austpalia frony Kangaroo Island to Ceduna and
the western Eyre Petinsula, and occurs in both gulls
(Hig. 3). ln the Spencer Gulf it can be found at Port
Augusta indicating that it ean tolerare the wide salinicy
ond temperiture range found m South Australnin
walers.
Pt Augusta’ 2 :
Os:
Penneshaw
oom
Fig, 3, Distribution of Elinjntay madesius Darwen, Ciminiay udetvidae Bay lias, Elrnintins flinderse sp tov, , Elelintias placidiss
ap nov, and Efi cbesceyiy sp, poy. in South Australie,
YHREE NEW BARNACLES PROM SOUTH AUSTRALIA Wo
Elminins placidus sp. nov,
FIGS 3, 4
Eliiinus Modestis: Womersiey & Edmonds 158:
243. Thomas & Edmonds 1979; lol, Hutchings &
Recher (982: 95.
Holotype: SAM C4244, on branches of Arcevia
marina at Cowell, South Austnilin. 33°8'S. 14694 "EB:
1), Bayliss. 6.4).1992: dissected (partially).
Paratypes: SAM ©4245, WAM 728-92, D. Bayliss,
6.111992. Same lacalily as holotype.
Description of holotype: Shell (Fig. 4A, 4B): uprieht
conical. grey ta white with thin brow) and dark erey
bands parallel to base. Pariecties gently folded, Shell
thin and nearly transluvent, Orifice large, pentagunal
in outline, rostral side coneave and broad, carinul sides
longer than lateral sides, width 4 length. Basal outline
sinuous. Alae wider than radii with less obhque
summits. Radii with oblique sammils, narrow: ead
and alae with prominent growth Hines. Shell 7mm ito
basal diameter. 5mm in width, Strom in height,
Opercula (Figs 4B-E): thin, translucent, white in
colour, grey margin near articilating margins of
internal Surlaces Of terguifi and scutum, Seuting longer
than high, basal margin slightly convex with prominent
depression for depressor muscles, fo eresis upparent,
articular ridge, articular furrow moderately developed.
adductor ridge, adductor pit not apparent, externally
growth ridges present.
Tereum wedge shaped, articular margin and basal
margin gently curving from apical end, coriverging on
basiscutal angle, articular furrow shallow, apical portion
large with small. feeble crests for tergal depressor
muscles; spur confluent with basiscutal angle.
Body (Fiz, 4G); Progomia white, narrow, clongated ,
palps and cirri | to U1 very dark brown, cirri LY to
V1 light brown wath dark brown pigment along anterior
face of segments, cirrus | covering Oral cone.
Mouthparts (Figs 4A-J); Labrum wath three teeth
and setulae on each side of central notch, Mamlible
with five teeth, lower pectinate edge temminating 19
short curved spine. Maxillule with two large spines
above natch, live smaller spines in notch, four large
spines below notch. smaller spines at lower angle.
Cirri (Pigs 4K-Ni: Cicris I with anterior ramus 61,7
length of posterior ramus, unterior ramus wilh broad
proximal segments, narrow distal segments. with long
sclac, segments slightly protuberant. Amterior Tamus
of cirrus Uo slightly longer than posterior rannus,
segments slightly protuberant anteriorly, Cirmis I with
amerior ramus *J.25 length of postenor rans,
segments slightly protuberant anteriorly, setae on pam
very Jon. Stout pectinate setae on six distal segments
of posterior ramus. Cirri IV to VI all subequal in
Jength. segment with four large pairs, two small pairs
of setae on anterior face, smal! proximal puir of seme
belween segments, anterior fee with dark brown
pigment. Number of segments 19 rai of efrr) 1 taVI
shown in Table 2.
Tama: 2, Cirral cunts: Elmigins plactlis sy gov farilerior
reams first),
Basal diameter
ammi Crus
| I i ly yvoo£«#r;
7.0 Lhilotype) bhA LO Pa44 27.24 28.30 32,30
40 $6 98M (212 Wz 26.29 3b,.29
40 12,6 TOO Vp,1) 24,23 26.24 5026
5,0 14,7 00,9 [4,83 23,23 27,27 28,27
6 ek TOM 14.94 25.24 28.23 24,27
Penis: Penis a long as cirros V1, setose, basidarsal
point absent.
hiviation: Shell is usually upright conical orcubular
and rarely Alartened. Shell way have visible banding
but = 10 iiagnitication is offen required to see the dark
and light bands. On mangroves the Darnacles nay
appear solidly ercy, Some speeunens colleeved from
Tocks have dark grey bands and the shell may appear
almost bluish, Thomas & Edmonds (1979) deserbed
the shell as being “bluish-areen" Some specimens huve
a slight pink tinge towards the top of the panetivs.
The tergum ts cXtremely thin near the basal margin
and wear may alter the shape, in some specrmens the
tergum 18 virtually triangular with the basal margin
yery gently curving from the tergal resis to the
basisculal angle, Qa unworn specimens. the external
surface of the terguim is grey with.a white area ar the
apical end.
The sewtui in matny specimens has a nearly straight
basal margin and (he arlicular inacgin 1s. almost at right
angles forming w right angle iriangle shape, The
depression for the depressiur musele is usually well
formed with the shell being extremely thin in this area
FErymology; The species name is derived Irom the
Latin plverdus meaning calm. crmquil with reference
to the habitat of ihis species.
Comparixon with other spectes: The shell is thin and
translucent wilh narrow dark bands. Other species arc
uniform in colour and lack the alternating Ifght and
dark banding. The opercular plates are thin with grey
margins igternally along the articulating margins.
The tergum ts distinctive in shape. The apical region
1§ large with very leeble crests. The articular margin
and the basal margin curve gently to the basiscutal
angle forming a inangular or wedge shape.
The prosoma js narrow and elongated as are the cirri.
The dark colour of the cine) contrasts with the white
prosoma. The coloration survives preservation,
The opercular flaps are cream with a dark black band
at Lhe rostral end and anothes black band centrally, This
enables it to be distinguished from £, medesius, bul
it cunnot be readily distinguished from other South
Australian species which have the scane pattenn of dark
bands on lighter coloured Naps.
120 D. E. BAYLISS
Fig. 4, Elminius placidus sp. nov. holotype. A. External view of shell; B. inner view of shell; C. scutum, internal view;
D. tergum, internal view; E. scutum, external view; F. tergum, external view; G. body; H. labrum; I. mandible; J. maxillule;
K-M. cirri I-IIf; N, middle segment, posterior ramus, cirrus VI.
THREE NEW BARNACLES FROM SOUTH AUSTRALIA 121
Fig. 5. Elminius erubescens sp. nov. holotype. A. External view of shell; B. inner view of shell; C. scutum, internal view;
D. tergum, internal view; E. scutum, external view; F. tergum, external view; G. body; H. labrum; I. mandible; J. maxillule;
K-M, cirri I-IIl; N. middle segment, posterior ramus, cirrus VI.
122 Db. L. BAYLISS
Habhar Elias places is.an intertidal species
which is found jm sheltered hatitats ft 1s highly
abundant in mangroves in Spencer Gulf Ie settles on
branches and pneulnatophores, but is rarely found on
leaves, His not restricted to jrangroves bul alsiy selthes
un rovks, cement, wood, metal and orber surfaces.
Distribution: Ebninies plactdys ts widespread, being
found on Kangaroo Island, Yorke Penigsala, Spencer
Gulf and Kyre Peninsula (Fig, 3). Pers nol foural tn
The mungroves north of Adelaide which are vecupied
by £) adelaidue.
Elminius erubescens sp. nov.
FLGS 3, 5
Liminins covertus Faster 1982; 26,
Holotype: SAM C4246, collected on rubber tyres
at Glenely bowt ramp, Adelitide, South Australia,
44°54'S, 138°36’E, D, Bayliss, 19,Vi.1991, dissected
(partially).
Paraty pes: SAM (4247, WAM 729-97, D. Bayliss,
1Y.vi 199), fran the saine locality as: holotype.
Deseription of the halotype: Shell (Pigs SA, SBy
flattened conical. translucent wilh red coloration.
appearing dark purple before the body nemoved, colour
ereyish purple when preserved, Paricties gently folded
(rilice small, pentagonal in outline, width %4 length,
rostral side short and suinght. other sides almost equal
in length, Basal outline sinuous, Alae wider than radii,
with Jess oblique summits. Radii with oblique summits,
narrow, Shell Simm in basal drarmater, 7mun in width,
3mm in height.
Opercula (Figs SC-F). Thin translueent. reddish.
greyish purple with white areas when preserved
Scutum longer than high, basal margin slightly convex
with upward turning al tergal corner, articular ridge.
articular furrow moderately developed, apex retlected
outwards, externally growth ridges apparent.
Tergum wilhurticular pidge folding inwards to form
well developed lurrow, articular margin concave, basal
margin curvine sharply to form projection with carina
Mafeln, Well developed crests for tergal derpessor
muscles, spur confluen! with basiscutal angle.
Body (Fig. 5G): Prosoma light brown, palps and
cirri Pand UE with some dark brown pigment, creri TL
to Vi clear, partially light brown, preserved material
with body and cirri almost uniformly light brown,
prosoma wnd eicn elongated. cirrus Loverlapping oral
came. :
Mouthparis (Pigs SH-J)) Labriim with three teeih
and setulae on. each side of central fete, Mandible
will five teeth, lower short pectinate edge tenminating
in short curved spine Muaxillule with two large spines
helow notch, smaller spities at lower angle.
Cir (Figs SK-N)}) Cirrus] with anterior ramus <5
length of posterior ramus, segments slightly
prowbenint, Anterior ramus af cirrus TE slightly longer
(han posterior ramus, segments slightly protuberant
anteniarly Cirrus fl wilh anterior ramus slightly longer
Than posterior ramus, scpments shvhtly protuberant
anteriorly, $ctae on both int tong, stout pectinate setae
on six distal segments uF posterior camus. Cirrus [VY
to Vi subequal in length. segments with three Jarge
pairs, (wo small pairs of setae on anterior fice. sniall
proximal pair ot setae between segments. Number ot
segments io rami of cinri | to Vi shown in Table 3,
Vault 3, Cirnal counts: Elminits erubescens sp. rn
(amierter rams first,
Basal diameter
(rim) Cirrus.
I I iW IV Vv VI
3.0) (holoryywy) = 2,7 99 LTP 21,24 25,24 28,30
TA 4.6 99 [242 25.28 29.27-°52.3)
a) 15H 18 Lill 23.20 25.24 25,24
75 13,6 Vb,00 1201 20.19 23,23 25.25
oO (2,6 9.9 13,72 23,23 25,26 30,27
Penis: Penis as long as cirrus VI, setose, busidorsal
point absent.
barialion: Elminius erubescens is usually Mattened
although tubular and upright conical forms can vccur.
‘The she!) often has broad longitudinal folds whieh vary
in murmber between specimens, Maximum basal
diameter iim.
Arymology: The specific name is derived fram the
Lutin ervheycere, to blush with modesty. in reference
to its redness.
Comparison with other species; Elminuts erubescens
can be easily distingished [rom other species in South
Australia by the red coloration of the transiucent shell
which, ithe eld, appears dark purple. The shell is
btherwise vlose to E jhodestus. It lacks the ridges seen
in & coverius, allhough worn specimens of the two
species are similar in appearance. The tergoscutal Haps
are straw lo) pale yellow with two dark brown to black
bands. A sinall portion of the flaps between the postral
end and the tirst dark band are white. £. cevertus his
six puirs uf dark spots on white tergoseutal flaps.
Habuar. Elminius erdbescens 1s comimon in sheltered
waters in the Adelaide region although it is nor found
in mangroves. 1 Setfles On 4 wide variety of surfaces
including rock, cement, Wood. metal and rubber. [r
is the highest barnacle in the intertidal zone, There js
litle overlap with EL madestus whieh forms a zone
below ft On cocks it is usually found on surfaces which
are overlapping or de oot free directly into the
atiermoon sun, although it can stand exposure to
sunlight for part of the day. E. udeleidae avoids sunlight
und is found vader rocks Which are lower in the
intertidal zone,
Distribution: This species 1s very common in the
Adeluide region, but was not dound elsewhere in South
Australia (Fig. 3). tts distribution outside of South
Australia is unknown,
THREE NEW BARNACLES FROM SOUTH AUSTRALIA
Key Lo species of the Elminiinae
Shell with tour
compurimental plates... 2
Sheil with six
compurtmental plates. 4
2 Tergum with spur
confluent wilh bitsiseutal
angle 3
Tergum) with distinet spur
Elmunits kingi: Grity
(South Anierival
Bt
3 Shell white or grey.
Shell not while or grey 5
4 Shell white, tergum with
straight urtcukin margin,
Jeep urticular furrow. Elminvay modestus
Darwin (Australia, New
Zedlund, Europe, South
Africa)
Shell grevish- white,
tergum hatchet shaped,
concave ariicular Margin,
centrally localised furrow Eliminivs flindersi sp nov
(South Australi)
Shell grey, narrow dark
bands alternaung with
white, lergum wedge
shaped, weak crests,
shallow furrow Elminius placidlas sp. ov
(South Australia)
5 Shell light brown, terguem
with small furrow
restricted ( apical end, Elmunius adelatdac
Bayliss (South Austral)
Shell translucent with ted
coloration, purple in
field, tergum with
concaye articular margin,
deep Tirrow . ...-- . Elminius eribeseens sp
nov, (South Australia)
Sheth butf red With ereany
rulges, lencun hatchet
shaped... , 0c Elminius coverius Fosver
(NSW. Western Australia)
6 Shell pale brown with
reddish brown bands
between low ribs, lerguin
with spur longer than
basiscutal angle ..
Shell preyish vrearn with
darker radial bands,
terguin with Spur not
longer than basisentul
angle - - 4
Hexaminius folioram
Anderson, Anderson a
Egan (NSW)
Hexaminiuy poupeiana
Foster (NSW)
Discussion
The diversity of extant species, as well as fossil
evidence, suggests that the Elmiminae originated it
south-eastern Australia (Buckeridge 1982, 1984). The
species present | South Australia are, with the
exception of Elminius modestus, not found in NSW,
The genus Heyanninius is not represented in Syuth
Australia and Elainins caverns is also ubsent.
33
South Australian barnacle populations are
geugraphically isolated from the eastern States by a
long expanse of coastline. from Robe to Cape Orwway
in Victoria, in Which intertidal species are virtually
absent (Womersley & Edynends 1958). In South
Australia there are extensive areas of coastline which
(ire protected from oceanic waves in the gulfs which
contain ideal habitats for Elminjus. Speciation may have
vceurred ia the variety of sheltered habitats which are
available.
It is also possible that species from elsewhere along
ihe southern coustline of Australia have been
introduced. At present the distribution of Elamnius
species in other States has not been determined. The
presence of E. covertus in Western Australia (Foster
1982) has been confirmed by Jones (1990), Tl is possible
that ic was introduced from eastern Austraha. £,
modestus may have been introduced from New Zealand
(Foster 1982; Plowerdew 1984), but an electrophoretic
comparison involving South Australian forms hits yet
to be done.
At present there is @ large scule attempt to establish
an oyster industry in South Australia using spat from
Tasmania. This could lead to introductions of species
not endemic to South Australia
Two species of Elminius are extremely abundant in
mangroves in South Australia, buc their distributions
do not overlap, &. udelaidae 1s found in the mangroves
north of Adelaide whereas AL placuluy is tound ii
mangroves in the Spencer Gulf The two gulfs have
considerable differences in thetr marine invertebrate
fauna (Shepherd 1983)
Only &. adelaidae utilises the leaves as well-as the
branches and pneumutophores, Anderson e7 al. (1988)
have deseribed a species Hexaminius foliorum, which
is specialised for living on leaves. The adaptations they
list as important for this species, whieh include thin
shell, thick basal membrane. long elongated cirri and
rapid cirral beating, are also found in £. adelaiduc.
Nevertheless, £. adelaidae grows to ainuch larger size.
is found in other habitats and has a larger variety of
cirral beating patterns, including the ability to hald the
cirral fan fully extended. & placidus has a more
restricted range of cirral acuivity and lacks the ability
1a beat ramidly. he basal membrane ts also thinner.
Tis thin shell und elongated body suggest a species
adapled for very calm habitats, bur not necessarily
mangroves.
In New Zealand E nrodesius ys reported to live in
mangroves (Moore 944; Morton & Miller 1968), The
present author has observed several spattalls in
mangroves near Adelaide. hut they failed to persist.
ft is uncommon for &. flindersi to be found in
mangroves and no FE. rrubeseens were found in
mangroves despite its abundance on rocks in the
Adelaide region.
124 D, FE. BAYLISS
Acknowledgments
{ would like to thank Dr A. J. Butler for providing
lise Of microscopes for examining specimens and for
reading a draft of the manuscript.
References
Anberson, D. T.. ANbERSON, J, T, & Ecan, B.A, (1988)
Balanoid barnacles of the genus Hexaminius
(Archaeobalanidae: Elminiinae) from mangroves of New
South Wales, including a description of a new species. Ree.
Aust. Mus. 40, 205-223.
Bayuss, D. E. (1982) Switching by Lepsiella vinosa
(Gastropoda) in South Australian mangroves. Oecologia
(Berl) 54, 212-226.
______ (1988) A new intertidal barnacle of the genus Elminius
(Cirripedia; Thoracica) from South Australia. Trans. R.
Soc. S. Aust, 112(2), 75-79,
BucKERIDGE, J. S. (1982) The barnacle subfamily
Elminiinae- Two new subgenera and a new Miocene species
from Victoria. J. R. Soc. N.Z. 12(4), 353-357,
(1984) A new species of Elminius from Pomahaka
River. Southland, New Zealand, N.Z. j. geol. geophys. 27,
217-219,
Darwin, C. (1854) “A monograph on the subclass Cirripedia,
with figures of all the species. The Balanidae, the
Verrucidae, etc.” (Ray Society, London),
Ecan, E. A. & Anperson, D. T. (1985) Larval development
of Elminius covertus Foster and Hexaminius popeiana
Foster (Cirripedia; Archaeobalanidae: Elminiinae) reared
in the laboratory. Aust, J. Mar, Freshw. Res. 36, 383-404,
Fiowrrpew, M, W, (1984) Electrophoretic comparison of
the anlipodean cirripede, Elminius modestus, with
immigrant European populations. /. mar. biol. Ass. ULK.
64, 625-635,
Foster, B. A. (1978) The marine fauna of New Zealand:
Barnacles (Cirripedia: Thoracica). Mem. N.Z. Oceanogr.
Inst. 69, 1-160.
(1980) Biogeographic implications of re-examination
of some common shore barnacles of Australia and New
Zealand. Prac, Int. Symp. Mar. Biogeography & Evolution
in the Southern Hemisphere, 613-623. N.Z. DSIR
Information Ser. 137.
(1982) Two new intertidal balanoid barnacles from
eastern Australia. Proc. Linn. Sac. N_SAo 106(1), 21-32,
Hutcuinas, P. A. & Recher, H. F (1982) The fauna of
Australian mangroves, /bid. 106(1), 83-121,
Jones, D. S. (1990) The shallow-water barnacles (Cirripedia:
Lepadomorpha, Balanomorpha) of southern Western
Australia, Jn Wells F. E., Walker D. 1., Kirkman H, &
Lethbridge R. (Eds) “Proceedings of the Third International
Marine Biological Workshop: The Marine Flora and Fauna
of Albany, Western Australia, 1988" Vol. 1, 333-437
(Western Australian Museum, Perth),
Moore, L. B. (1944) Some intertidal sessile barnacles of
New Zealand. Trans. R. Sac. N.Z. 73, 315-334.
Morton, J. E. & Miter, M. C. (1968) “The New Zealand
Sea Shore.” (Collins, London, Auckland).
Pore, E. C. (1945) A simplified key to the sessile barnacles
found on the rocks, boats, wharf piles and other installations
in Port Jackson and adjacent waters. Rec. Aust. Mus. 21,
351-372.
SHEPHERD, S. A. (1983) Benthic communities of upper
Spencer Gulf, South Australia. Trans. Ro Soc. S. Aust.
107(2), 69-85.
THomMas, L. M. & Epmonps, S. J. (1979) Intertidal
invertebrates. pp. 155-166, /n Tyler, M. J., Twidale. C. R.,
and Ling, J, K. (Eds) “Natural History of Kangaroo Island”
(Royal Saciety of 8, Aust., Adelaide),
Womers._ey. H. B. 8, & Epmonps, S. J, (1958) A general
account of the intertidal ecology of South Australian coasts,
Aust. J. Mar. Freshw. Res, 9, 217-260.
EVALUATION OF EXTENSIVE ARID RANGELANDS:
THE LAND CONDITION INDEX (LCD
By ROBERT T. LANGE*, BRENDAN G. LAY? & RODGER W. TYNANT
Summary
Lange, R. T., Lay, B. G. & Tynan, R. W. (1994) Evaluation of extensive arid
rangelands: the land condition index (LCI). Trans. R. Soc. S. Aust. 118(2), 125-131,
31 May, 1994.
Social and historical circumstances that have given rise to rigorous new rangeland
management legislation in South Australia are outlined. A program specified by the
legislation is explained and described which will determine the condition of the 40
million hectares of arid rangeland in South Australia relative to criteria about
sustainability and prevention and rectification of degradation. An objective
assessment of average rangeland condition, the Land Condition Index (LCI) is
introduced and results obtained by its use are illustrated and appraised.
Key Words: Arid zone, rangelands, Australia, vegetation, assessment, technique.
Transeidnions of the Rawil Society of S. Awyt. (994). M803), 125-13)
EVALUATION OF EXTENSIVE ARID RANGELANDS:
THE LAND CONDITION INDEX (LCT)
by ROBERT T. LANGE, BRENDAN G. LAY? & RODGER W, TYNAN?
Sumunary
(anGe, RT), baw B, G. & Trvan RW, (1994) Evaluation of extensive arid rangelands: the laid condition
index (LCT). Trans. Ro Soe 8. duasy 1802), 125-43L 3) May, 1994,
Social and historical circumstances (hat haye given rise to gorous new rangeland janagernent legislation in
South Aastra a are- outlined. A prowran) specified by the legistation is explainéd and desenbed which will determine
the condition of the 40 mallion hevtires of urid ranjclund in South Australia reluuve to crite about sustarability
dnd prevention und recification uf degradation An objective assessment of average rangeland condition, the Land
Conduinn trdex (10) is introduved and results oblained by its use are jllustrated and appraised
Kiee Wows, Arid zony, tangelinds, Australi, vegetation, assessment, technique
luteoduction
The Pastoral Land Management and Conservation
Ace (1989) came into operation in South Australia in
March, 1990, after years of public contention. Lt
challenged the then Outback. Manapertent Brancly of
the South Australian Lands Department (us. the
responsible agency) with an immense, mandatory “ind
assessment task within an 8-year deadline. In tts
combination of difficult features und its emphasis on
sustainability, this task may have parallels in other arid
parts of the world, where our procedures might,
therelore, be applicable:
The particular combination of challenging features
isa vast rangeland area (the size ot Tray or Somalta
for exainple) poorly documented and difficult of access,
lo be assessed for Lhe integrity of its vegetation and
soils in detail, acvording to characteristics nol
uccessible by remote sensing, using minimal time, staff
and money,
We faced (he further difficully (given the linjited
arount of resources relative to the immense size of
the task) that aur assessments even of small tracts of
Jand (a (ew hundred ba or so) would have to be capable
of withstanding challenge and close-scrutiny in a court
of law,
This paper outlines (a) the histoneal and social
context of the Act, (b) an approach (inelbding a new
index of fand conditian) which we have devised to help
uctueve Whal the Act reqilires and (c) some initial
resulls abtained by application of the index,
+
National Key Centre for Teaching and Research m Dryland
Apricullure and Lund Use Systems, The Universi(y of
Adelaide Roseworthy Campus, Roseyorthy, South Australia
ae |
Current address: Box 330, Whyalla. Seuth Australia 5600,
~ Pastoral Management Branch, Resource Conservation
Division, Department of Environment and Natural
Resources, GPO Box 1047, Adelaide. South Australia S001
=
Australian Background
Whether native inland Australian vegetauion can
Ultimately withstand the Impact of Ongulate Mocks and
herds is yet to be determined, Since it evolved in total
isolation from) ungulates. it may prove unayoidably
susceptible to the effects of Jarge numbers of them.
From prehistoric ime until the mid-I9th century, the
inland and regions of S.A. sustained only sparse
Aboriginal nomadic hunter-gatheters. Due to lack of
surface waters most vegetation probably experienced
only very light grazing pressure, om average, by
kangaroos alone. That regime changed “vata-
clysmically” (Adamson & Fox 1982) when Europeans
colonised much of the country. lapped ground waters
and introduced millions of uagulate stock and various
feral animals. creating aw sedentary prazing industry
which now vecupies 40 m ha (Fig. 1) and is mostly
i the early part of its sevond century of operation,
The saying “Our of sight, our of mind” aptly
summarises public and governmental attitudes to
pastoral zone landcare throughout much of the
indusiry’s history (Lange 1983), Nearly all of South
Australia’s | m people dwell in the arible caastal
regions and of these the vast majority live in and about
Adelaide. Only a low hundred al any one time have
had extensive pastoral zone experience. Cily-dwellers
have only recently become better aware and more voeul
about the arid zone, as tourism, wildlife protection,
recreation and mining have increased.
Sheep stocking enterprises. typical of the more
southerly parts of the zone are protected from predatory
dingoes by a special fence (the Dag Fence). Features
include the subdivision of the rangeland into wire-
fenced paddocks with water available tor stock at fixed
points, approximately fixed flock size year in, year out
anda system of vehicular access tracks. This leads. to
the development of repealed drink-centred patterns of
unequal flocktime distnibuiion (piosphere pattern,
Lange 1969, 1985: Andrew 1988) in which flocktime
126 R. T, LANGE, B, G. LAY & R. W. TYNAN
Soil Conservation Districts in the pastoral zone.
Fig. |) The pastoral lease lands of South Australia, showing also their subdivision into Soil Conservation Districts,
concentrations can rapidly cause damage to the
vegetation and soil, particularly during drought
periods. The northerly extensions, exclusive to cattle,
show the same patterns ona broader scale, with most
pastoral properties there without internal fences.
Seientinc and general Intersture, government
inspectors’ reports and the like show that from its
inception onwards, this industry has caused great
damage to the land, in some cases denuding the soil
in just 2-3 decades (Dixon 1892: Waite 1896; Ratclille
1936, Pick 1944; Lay 1979). This was of public concern
particularly in the 1930s when the issue of soil erosion
was topical worldwide (Jacks & Whyte 1939) and has
subsequently remained a problem, Half a century later
in the 1980s, government administration again publicly
conceded that some pastoralists were still failing to
meet landcare obligations and that land degradation
from overstocking and feral animal pests was
extensive!. In an attempt ta accommodate all
interested parties, the Pastoral Land Management und
Conservation (PLMC) Act was finally passed in 1989.
This Act is profoundly different from earlier South
Australian pastoral laws in its great emphasis on
resource protection. It is, in fact, unequivocal landcare
legislation (in some ways setting searcely-attamable
ideals of landcare perfection) but it also provides
provisions for grazing industry protection and appeul,
How these opposites will be reconciled in practice is
yet to be shown,
Of special relevance to this paper, the new Act gave
the Minister of Lands greatly increased scientific and
technical responsibilities. The administration of the Act
is delegated to the Pastoral Board (whose membership
reflects major interests in pastoralism and conservation)
but the technical tasks miust be carried out by
the Pastoral Management Branch. Of these tasks,
this paper refers to the requirement for vegetation
assessment. This immense task. which the Act requires
"South Australian Government. Lands Department (1981)
The administration, management and tenure of South
Australias Pastoral Lands. Interdeparimental Report
(Chairman: J, Vickery, Lands Department).
THE LC} (27,
ti be carried Gut within witht years and at id-year
imervals thereafter, réfates lo 40 tn ha of the rangelands
und is to specify the condition ot the land geeording
To entena derived fram the Objects of the new Act.
These effeciively require that the whole
area he divecily inspected by tears of Lmined assessors
on the ground. As is usual where a lange and
complicated lask must be weconiplished piecemeal by
Uifferent (euims, Ohyective, stundardised procedures
need io he adlopled io ensure thal the findings ol any
assessment are independent af the particular team
enyployed ‘The development ol the index arid training
plocediires deseribed below |§ an allewpt at suen
slimdardisatien
The Lesyl Stiptilations. about Assessinenr
Ii Soul, Austala’s nonarahle inland regions a
pastoral Iease is the only form of tenure than con be
gramed over Crown (government) land that is to be
Used lor long-term grazing purposes, a pastoral tease
connot be granted orextended without an assessinent
if the conditian of the Land first having been made
(PLMC Act, 1989, See, 20, 25), This is why the
assessient pragian is urgent aad eructal. The pastarul
Industry ts awaiting the issuing of new leases.
specilying new Stocking fintits and ciher condinons
consequent Upon these assessments.
Section @ of the PLMC Act slates (hat suc
assessment ot the condition of the Jand must be
thoraugh, must tnelude an assessment oF the capacity
of tie land tq carry stock, musi be conducted jn
mccnrdance with recognised scienhtic principles and
must be carried out by persons who are qualified and
experienced in land assessment techniques.
The further meaning of assessment derives directly
from relevant Objects of the PLMC Act (Sec. 4) which
are: (4) lo ensure that all pastoral land in the State is
well-managed and utilised prudently so thar its
renewuble resources ure maintained and its yield
sustained and (b) to provide for (1) the effective
inanmitoring of the eandition of pastoral land, (2) the
prevention of degradation of the land and its indigenous
plant and animal Tite-and (3) the rehabilitalion af the
land in cases of damage. The Minister is expresshy
charged (PLMC Act. See. S) with the duty of acting
consistently wilh and an furtherance ot those Objects,
Assessivient pursuant to the Act thas is the process af
obtaining scientifically based answers to the Gbjcets
posed as questions, viz, Is this pastoral land well-
managed and utilised prudently so that ils renewable
resources dre being maintained and its yield sistined”
Is degradation of the tand and of its indigenous plant
and animal life being prevented? Is there degradation
which requires rehabilitation? Has eflective monitoring
of the condition of the land been proylded? What is
the capacity of the land to carry stock consistently with
answers fo the foregoing questions?
Since “degradauon” is defined (PLMC Act, Sec. 3)
as d decline jm the quality of the natural resaurves of
the land resulting From human aetivitves, same ol the
inipliestions unsiny from the Objects are imotediately
lipparent. For instance, the Capventionsl rangelands
dogma thal the desirable condition of fhe vogetation
is thar Which feeds most stock is potentially denied by
this legislation. The optitnal candition instead 14 that
where indigenous plant and animal life retain their
prepastoral integrity while as far as possible Suslaniny
au Viable pastoral industry Appreciation of this
difference is. crucial in developing the condition index
outlined below and | Will be further examined jf (he
Discussion.
That the unswers to Wiese assessment questions ure
meant to be taken seriously is made plain in many parts
of the PLMC Act including See, 43 for example, a
Section excluded from the appeal provisions (See 54_
55). Section 43 makes it clear that the Pastoral Board
should require the rennoval of stock fren an area not
only ifits hehieves that the land has been danniged but
ulso if the Bound considers thar the land is likely to
suffer damage.
Essential Preliminaries to Assessment
fy should be noted here that the assessment program
as develaped cunsists of two pans viz — the Iease-based
Land Condition Index determinution (the subject of this
paper) and a petwark of paddock-based photo-
monitoring sites, These latter sites, with vegetation and
soils data collected, form the base line fur the
asbessment of future trends ip cach paddock,
Access and Sampliriz
Any thoroligh assessment of a station or lease in
accordance with revagnised scientific principles (Sec.
6 ol the PLMC Act) should ensploy many mdependent
samples in an effort jo achieve a petite which is as
balanced and equitable as ts practicable. In practice,
sampling falls somewhere between systematic and
random, as in the way the industry itself informs
potential buyers about the contents of a bale of wool,
for example, where the sumple is (rom an arbitrary
grab. In sainpling station condinon, rehunee ts ustially
placed upon the pastoralist’s systemt of vehicle access
tracks to spread sumples thraughout the overall area.
Then, to avid possible observer bius, samples are
draWn by strict pestricted-randoin processes from
whal i8 track-uccessible. There are some obvious
consequences of such sampling but they are more
academic than practical and there ts na evidence that
they sigmficantly uffeer comparisems. All well-
developed pasioral land has adequate watering points
and a network of access tracks linking them, Thus there
js a higher chan random likelihood of sarmpling closer
(a avitter, and a currespondingly reduced likelihood al
intermedia ranges, than would result Go randem
128 R. T. LANGE, B.
Kity fram water
G LAY & R, W, TYNAN
Km from water
45° -& 375-4
' | '
te 6
or the LC
sampling route
o 4S oo 8 6 TH 4 105 o
| \
&;
ro)
pas |
=
&
= 2 x
5
at Ly 40 =
SE corner, McDouall Peak Station
Fig 2
same length af
route relocated
without reference
to water
SE corner, Millers Creek Station
Histograms showing the percentage of sampling points at various distances from water paints on the access tracks
(above) and when the sume trick-yrid 1s offyet from witer-points (below).
tracks. This effect is consistent from station to station
(Fig. 2) and can always be taken into account.
Recognition af vegennian Type and Component
The next essential of the assessment procedure is that
the assessment team member, at any sample point, must
recognise Which of the many different vegetation types
js represented and, of that type, which of the various
sub-lypes called components. In practice Uns ability
comes from instruction manuals. and field training
provided by us, as experienced team members, who have
a collective 60 years of botanical work experience in
the region to draw upon. Learning is simplified by
restricting consideration to manuals drawn up for one
Soil Conservation District at 4 time, thus reducing the
immediate diversity to be handled, The assessment
feams then attack the task by completing assessments
of one District before re-calibration and assessment of
the next District and so on,
We write keys for the identification and separation
of the Vegetation types and components in each district,
with sets of enteria to distinguish condition classes (see
below) and we illustrate the resulting manuals with
extensive colour photo-guides, All teams are then
trained en bloc (in extended field calibration exercises)
to achieve a uniform, consistent interpretation of all
erieria,
All main diagnostic criteria refer to perennial plants,
rather than to ephemerals which appear only alter
significant rainfall. The main reason for reference 10
perennials is that (hey embody the drought fodder
reserves of the rangeland as well as being the reliable
buffer against wind erosion, There are a few places,
particularly in northern cattle rangelands, that lack
perennial vegetation sufficient for present purposes:
these will be treated in separate ways,
TARLE 1. Fevewtion type ane component used in LCL
Type Component
Chenopod shrublands Atriplex vesicaria-Maireana
astrotricha \reeless plains
Acacia aneuray A, ramulosa-
frass sp. on deep sands.
Woodlands with grassy
understorey
The level at which types and components in general
are distinguished is indicated by the examples giving
the names of predominant species in Table 1,
Condition Classes
Arguments about non-linear “state and transition”
models of rangelands dynamics notwithstanding
(Westoby et al. 1989), experience of S.A. rangelands
shows that in general any component can be placed
into one or other of three condition classes viz,, Class
3, with the orginal stock-palatable perennial species
suill present and reproducing, Class 2. with the original
palatable perennial species being eliminated (if grasses
or low shrub species) or lacking all juveniles (if large
long-lived perennials) and with replacement by much
less-palatable perennial species including weeds or by
ephemerals and Class J, with all palatable perennial
species eliminated except old trees, unpalatable species
heavily pruned, many weeds present or, in some cases,
the integrity of plant cover totally destroyed. It has of
course been demonstrated for nearly 60 years thal
deterioration to Class. 1 goes hand in hand with the
onset and rapid acceleration of sail erosion (Ratcliffe
1936; Jacks & Whyte 1939; Pick 1944; Lay 1979).
The advantage of distinguishing only three classes
is that only two sets of separation-criteria are then
required Le, separating Class 3 from Class 2 and Class
2 from Class 1. Obviously, it 1s easier to train all
THE 101 129
jssexsment tearus ly absolute cangisteney when they have
oapply only Wwe sets of sepanition criteria ty deseribe
the condition of some given vegetational compo ent
instead of three, four ot more sets. Contrary to what
al first sight might seem to be the case, this restricuion
{o just three condition classes im no way limits the
yencral sensitivity of inter-levse comparisons. When the
condition of two leases is to be compared. the sensitryity
of the index of comparison depends instead on a
sufficient number ol saniples plus the use of weighted
averages. as will be demonstrated. So, in specitying
arbitrary but consistent criteria for the separation of
condition classes. counts cat be avoided in favour of
absolute differences. Regarding the 4. vesicarig-M,
astrotricha Component of the chenopod shrublands. for
example, the main criterion for separaiing condition
Class 3 from condition Class 2 is the elimination of 4,
veyivaria (a grazing-susceptible species of the orizinal
palatible perennial component). Condition Class | is
distinguished from Class 2 by loss of M. asrratricka,
a palatable bul more grazing-resistant perennial. “aking
the exatnple of the Acacia areure-A. ramulosa-2riss spy
component of (he woodlands with grassy understorey,
condition Class 2 involves. Joss of all of, vtetre
Tegeneriion, appearance of a distinct browse line on
old wees, elimination of the most palatable perennial
grasses (&g. Monechather puradoxa) and severe
pruning, at teust, of palatable shrubs (e.g. Ereynophile
letrobet), Similar sets of criteria haye been adopter (o
distinguish condition classes of all yegetatinnal
components of all types. in the rangelands, supported
by photogmphic examples of these classes,
Training
An essential for condition assessment condovted
piecemeal by different teams. using hus method. ts a
toll scale combined field-training exercise, The Soil
Conservation District to be assessed it [raverseat ty
intersect all vegetation types and components, At stop
after stop on this traverse, samples are assessed by the
party as a Whole, according to the manual, untrl wall
questions have been resulved on the spot, guaranteciys
anilormity of interpretation. Only then are the ditterent
teams allocated their particular shares of the district
workload and the actual assessments conducted
Predetermination of Sampling Points
Using available maps of slation tracks. the
assessment sampling route is pre-selected for best
syslemutic coverage and the total length involved is
calculated using, an opisometer. That length is then
divided into JO0 equal parts within each of which a
precise Stopping-point is drawn at random. The tesults
are then programmed into a computerised trip-meter
(Halda Instrument Co.) mounted in the access vehicle,
Which then signals the stops. This guarantees restniceed-
random sampling of the route,
Assessirent and LC! Calculation
Field teams comprising a scientitic officer and a
echnical officer operate self-contained 4-wheel drive
expeditionary vehicles equipped far remote res
operations, including radia communications.
Nuvigation is by reference to available maps und
satellite imagery using a vehicle-mounted GPS unit.
At each sampte site (of which the trip meter sounds
prior warning and displays a. count down to 10 ny), the
reference is made Lo the SU pa-square plot located
immediately beyond a line parallel to the direction of
vehicular travel, 10 m offset from the Jett side of the
vehicle. The sample within this plot is rated according
to vegetation type, component and condition, is
photographed and the data are recorded. At the
conclusion of sampling, (he station land condition
(LCD is calculated as in the hypothetical example in
Table 2,
YABLE 2, Bromple of caleniation of LOL qstarion lithe
cendition),
Condition Percentase Mollipher Totals
Class of ssaraples
3 50 wR = 150
z 30 *2 = 60
I 20 =] = 20
100 2340
Lel = 2.30
Note that this index has fimits of 1.00 (the whole
stalion in “degraded” condition) and 3,00 (the whole
station in “undegraded” condition), Actual stations have
values between these limits. Note also that the index.
once calculated, is not to be interpreted mechanistically
bul instead is used for first-order sorting of stations
according ta condiboen, Other evidence and dala
collected independently about cach staved are then
laken into account, Stations can be compared directly
by the LCT only where they involve srmifar pasture
types and components jin similar proportions:
The Nature of Initial Results
Assessment leams obtained first resujts during 1994
from the Kingoonya Soil Conservation District of South
Australia using J9 stanons, big. 3 makes a companson
between the LCI scores of hinds making up the stations
it this district and the maximum stocking permitted
on them by the provisions of existing lease documents,
As can be seen, no station attains the ideal of the Act
(all samples in Class 3), Instead, stalions range very
widely in condition, from as low as 160 up to 270, with
an average of about 200). This low average score would
imply that maximum stacking allowances will have to
be reduced, on average, 6 enable some progress with
130 R. T. LANGE. B. G. LAY & R. W. TYNAN
the rehabilitative process prescribed by the PLMC Act.
As can also be seen, there is no consistent relationship
between the present condition of stations und the
maximum stock that they are currently permitted to
carry. This is in contradiction to the Objects of the
legislation which imply clearly that stocking. if it is
to be intense, must go hand in hand with the
maintenance of a high LCI score.
From the descriptions above if can be seen that the
LCI provides a comparative assessment of stations on
standardised criteria derived specifically from the
Objects of the Pastoral Land Management and
Conservation Act, synthesismg an overall picture from
data provided by means of teams of which none has
seen more than a few of the total stations, [1 thus fulfils
iis design purpose and can deliver defensible
comparative assessments to the Pastoral Board, which
has the task of finalising judgments affecting stocking.
Fig. 3 shows the crucial difference betwecn station
country as perceived according to the Objects of the
Act on the one hand and as perceived through “pastoral
production” eyes on the other. For example, station land
occupied by salt flats with inedible samphires
(Chenopodiaceae tribe Salicorneae) vegetation might
score a perfect 3.00 on the LCI scale. for completely
satistying the landcare Objects of the Act, while being
intrinsically useless for pastoral production whereas
overgrazed and denuded Jands (e.g. chenopod
shrublands reduced to ephemeral grasslands) of very
low LCI score might earn the pastoralist substantial
profits as a result of high animal production. The aim
of the Act where LCI scores are law is to arrest decline
and then reverse it into a long-term trend of ascending
LCI scores as a first pnority. Any stocking of the lands
has to be commensurate with that and be a secondary
consideration.
Two major advances in ecological data-collection
arise from the LCI sampling program. Since the
distribution of the sampling sites is known, and is as
close to an even scatter a8 can be achieved, the data
yield estimates of the proportions of the district that
are occupied by the various types and components of
the rangelands vegetation as well as distribution details.
Further, since the condition of the component at each
site is recorded, the condition-profile of each
rangelands vegetational component can be specified ,
Land Condition Index (LCI)
100 160
200
270 300
'
0 6 12
15. = 12
21 28
Stock maximum sheep/km’
Mig, 3. Uhe prevailing mismatching between stocking allowances amongst leases of the Kingoonya Soil District, South Australia,
THE 104 yy
reveuling the relative degree to which (he different Lypes
have been degraded. This is imporant management
information, Tis upon the basis of data such as these
thal the Pastoral Land Management and Conservation
Act must operate and without which its administranon
cannot address its legally-stipulated imperatives.
Discussion
‘Well-reasoned technical preseriptions for landcare
in the State’s pastoral zone haye been ayaileble [01
ulinost.a century, Waile (1896) aimed for a bulanee
hetween Giitake and sustainability via u well-reasoned
ecnlugical argument jivolving land and flock
subdivision. waterpoint multipheation, nutrition of
lactating ewes, the distances sheep walk, drought
strategy, deferred grazing, spelling and economics,
Waites: Key principles have been firmly linked with
good landcare by Lay (1979) on the hasis of long-term
studies. The mest recent managerial prescription
(Statford-Sith & Morton (990) is lithe different in
ponciple from the first (Waite 1896) su i is mol
deficiency of management adviee that explains a
century of ongeing degradation,
From initial experience with the LCE it now seers
probable thar most stations in South Australia will fall
short of the land vendition Weal of the new Act. This
means (tel rechamatory, Pehabiliiatory action will
predictably he required which involves
recommendations for reduced stocking levels. We
believe that those stations mangged in closest accord
with the Act (highest LCT) should retain for the present
Vheir existing stocking wllowunces and act as
benchmarks for other stations with similar vegetation
types but lower indices. These other suitions should
be given lower stocking limits while rehabilitation
programs are inttiated. Qne reason for nor
recommending stocking allowances higher than ut
preseol in uny imstance (even on stations with highest
LCV is due lo a feature of the index itself, In order
I wtrard against likely accusations that the assessment
scoring procedyres are too severe, (hey Were structured
in favour of high LCT scores rather than otherwise. For
example, downgrading asaltbush (4friplex vestear fa)
shrubland from condition Class 3 to 2 is forestallod
While just a scattering of saltbush remain, even thoagh
the ungrazed density wus tnany thousand/ha ‘Thus
Where even the best-preserved vegetation in an area
scores less than the ideal, there is no full comphance
with the sustamabilily motion central lo the Act.
In ils inititl considerations of the first Ieases assessed
by this inethod in the Kingoonya Soil Conservation
Distrier, the Pastoral Board has been reluctant to
adjust leases with low LCI scores unless these scores
can be dipeetly attributable fo current management
Rather, it has indicated it would await air indication
oF trend From permanent photu-muniloring sites set ap
ut the time the LCL is determined.
Of the many further points that might be made, the
ane warranting most emphasis here ws that the LCL and
associated procedures bave to be taken against the
backdrop of the strinvencies, pressures and urgencies
that allend il, Those who right regard is as hasty
{compared with whal has been allempled in some other
Australian States) should recall the timeframe, The
whole rangelands (40 m ha) must be by law assessed
in eight years: this is a formidable and costly task.
Ultimately, however, landcare in the zone will
depend not on the assessments but on the determination
with which executive government lises {he assessment
information to ensure that more sustainable tand
mangement practices are adopted in our arid
rungelands.
Referenves
Avamson, BD. & Pox, M. 982) Change in Australian
veseration since BLnopean setlioment pp 109-146 Jn Savith,
J, (Ed) "A History of Australasian Settlement” (MeGraw
Hill, Sydney)
Anpelw, M2 H (1988) Grazing inipact in relation ta
livesiock walering pols, Trends rr Beal, and’ Eval. 3,
436-434.
Dixon, S. S92) The efleels oF settlement aad pastacsl
pecupation im Australis upon the ddigenous yegelstion.
Triny. Ro Sue S Aust 13. 145-206.
Jacks, G. Vo & Wryre, Ro OF (1999) “The Rape of the
Barth. A World Survey Wt Soil Erosion”, (Faber && Fiber,
Lundon)
LANGE, Ro T (1964) The plosphere. sheep track and dune
paterns, J Range. Manag, TZ, 396-100,
(1983) Arid zone lindeare: Educanon and Polines.
pp. 12-129 Jn *Whaut Future for Australia’s Arid Lands?”
(Aust. Comsery. Mound., Vie, Aust.)
—.... IRS) Spatial distributions of stacking intensity
priduced by sheep flovks grazing Australian chenopod
shriblands. trans. Ro Suv §. Ase, 9, 167-174.
Lay, BG. (1979) Shrub population dynamics under grazing
— a lone term study, pp. 1074129 fy Graera, R. & Howes,
K. (Rds) “Studies al the Australian Arid Zone lV, Chenopod
Shrublunds.” (CSIRO, Aust.).
Prek, J) Ho (1944) “Australia’s Dying Heart” (Melbourne
University Press, Oxtord University Press)..
Bareiiren, FON. (1996) Soil Drifi in the Arid Pastoral Areas
of Soorh Australia, CSTR (Aust) Pamiphler No. 64.
South AUSTRALIAN GOVERNMENT (1989) Parliament, Pastoral
Land Management una Conservation Aer (Govi, Printer,
Adelaide).
Srarrorn-Svire, Do M. & Moron, § RB, (990) A
framework for the ecology of arid Australia, J, of Arid,
Env, (8, 255-278,
Warn, P (1896) Pastoral development — past and present.
“Adelaide Observer” Sept, 26. 1896 (S, Aust.)
Wesrusy, Mo. Wacker. B. & Noy-Mein, 1, 11980)
Opportunistic management for rangelands mot at
equilibviiiny. .f, Ranke Manak. 42. 26h%-274
A SMALL STHENURINE KANGAROO FROM A PLEISTOCENE
CAVE DEPOSIT, NULLARBOR PLAIN, WESTERN AUSTRALIA
By G. J. PRIDEAUX*
Summary
Prideaux, G. J. (1994) A small sthenurine kangaroo from a Pleistocene cave deposit,
Nullarbor Plain, Western Australia. Trans. R. Soc. S. Aust. 118(2), 133-138, 31 May,
1994,
Simosthenurus maddocki nullarborensis ssp. nov. is described from Lindsay Hall
Cave, northwest of Madura Pass on the Nullarbor Plain, Western Australia. Although
the upper molars are very close in morphology to S. maddocki maddocki Wells &
Murray, 1979 from eastern Australia, they are notably smaller. The P’ is small,
narrow and inflated posteriorly. Geographic dwarfing due to resource limitations is a
possible explanation for the small size of this Nullarbor subspecies.
Key Words: sthenurine kangaroo, Simosthenurus maddocki nullarborensis ssp. nov.,
Simosthenurus maddocki maddocki, Lindsay Hall Cave, Nullarbor Plain, Pleistocene,
geographic dwarfing.
Transactions of the Royal Society of 8, Aust, (1994), 18(2). 133-138,
A SMALL STHENURINE KANGAROO FROM A PLEISTOCENE CAVE DEPOSIT,
NULLARBOR PLAIN, WESTERN AUSTRALIA
by G. J. PRIDEAUX*
Summary
Pripraux, G. J. (1994) A small sthenurine kangaroo from a Pleistocene cave deposit, Nullarbor Plain, Western
Australia. Trans. R, Soe. §. Aust. 118(2), 133-138. 31 May, 1994,
Simosthenurus maddocki nullarborensis ssp. nov. is described from Lindsay Hall Cave, northwest of Madura
Pass on the Nullarbor Plain, Western Australia. Although the upper molars are very close in morphology to 5,
madidocki maddocki Wells & Murray, 1979 from eastern Australia, they are notably smaller. The P? is small,
narrow and inflated posteriorly. Geographic dwarting due to resource limitations is a possible explanation for
the small sive of this Nullarbor subspecies.
Key Worps: sthenurine kangaroo, Simosthenurus maddocki nullarhorensis ssp. noy., Simosthenurus maddocki
maddocki, Lindsay: Hall Cave, Nullarbor Plain, Pleistocene, geographic dwarfing.
Introduction
In April 1991, a recently collapsed opening to a
limestone cave was discovered by members of the
Western Australian Plane Caving Group on Madura
Station, northwest of Madura Pass on the Nullarbor
* School of Biological Sciences, Flinders University of South
Australia, GPO Box 2100, Adelaide, S. Aust. 5001
MADURA
A
NARACOORTE
TANTANOOLA
Plain (Fig. Ll). Fossil material was removed from the
cave, named Lindsay Hall Cave, during exploration in
September 1991 and April 1992. Several cranial
fragments belonging to a medium to large size
sthenurine, and a very small partial left maxilla were
recovered. This latter specimen represents a new form
smaller than any sthenurine previously described from
the Pleistocene. It most closely resembles
Simosthenurus maddocki Wells & Murray, 1979 from
eastern Australia with which it is compared, This paper
describes the new sthenurine.
BINGARA A
WOMBEYAN CAVES &
MT FAIRY &
A
A & LANCEFIELD
Fig. |, Deposits yielding Simosthenurus maddocki in southeastern Australia.
134 G. J, PRIDEAUX
Material and Methods
The material is housed in the Western Australian
Museum, Perth (WAM). Mensuration, dental
terminology and nomenclature follow Tedford (1966).
As the homology of premolar cusps is currently being
re-examined by Prof. David Ride (pers. comm.), they
are referred to here an anterior labial cusp, posterior
lingual cusp, etc. All measurements are in millimetres.
Abbreviations: L = length; AW = anterior width
protoloph(id); PW = posterior width metaloph,
hypolophid; AH = anterior crown height, labial side;
PH = posterior height; n = sample size, Note crown
height measurements are heavily dependent on degree
of enamel wear.
Systematics
Order: DIPROTODONTIA Owen, 1866
Suborder: PHALANGERIDA Aplin & Archer, 1987
Superfamily; MACROPODOIDEA Gray, 1821
Family: MACROPODIDAE Gray, 182]
Subfamily: STHENURINAE (Glauert, 1926)
Genus: SIMOSTHENURUS Tedford, 1966
Simosthenurus maddocki maddocki
Wells & Murray, 1979
FIGS 1, 5
Holotype: SAM P16999, a near complete juvenile
skull collected from Victoria Fossil Cave, Naracoorte,
South Australia. Diagnosis, description and
comparison of nominotypic form of S. maddocki is
provided by Wells & Murray (1979), This subspecies
is also recognised from Greenwater Hole Cave, near
Tantanoola in South Australia (Pledge 1980), Bingara,
Wombeyan Caves and Mt Fairy in New South Wales
(Flannery & Hope 1983), and Lancefield in Victoria
(pers. observation). Age of type locality is late
Pleistocene (Wells et al, 1984).
Simosthenurus maddocki nullarborensis ssp. nov,
FIGS 1-5
Holotype: WAM 92.9.8, a partial left juvenile maxilla
collected from Lindsay Hall Cave, near Madura,
Nullarbor Plain, Western Australia (31935/’S,
126°40'E). Collected in September 1991 by Wendy
Binks and Katherine Crisp, Western Australian Plane
Caving Group. Age of type locality is ?late Pleistocene,
Diagnosis: Maxilla smaller than Simosthenurus
maddocki maddocki Wells & Murray, 1979 and
Sthenurus gilli Merrilees, 1965; molars low crowned,
very similar in morphology to S. m. maddocki, but
smaller. Molars possess very fine enamel crenulations
with lophs notably convex anteriorly. P* very small,
narrow to tapered anteriorly, but inflated posteriorly.
Fig. 2. Stereopair of Simosthenurus maddocki nullarborensis ssp. nov. left maxilla (WAM 92.9.8, holotype) in occlusal view.
SMALL STHENURINE KANGAROO 55
Hig. 3. Stereapaits of Siosthenurus wiaddocki hillabomevisis ssp. Ao (etl Pe, top, and P! bottom, (WAM 92.48, holotype)
in occlusal view,
Description of holetype; Maxilla (Fig, 2),
Fragmentary nature of specimen has resulted) in
preservation of yery few non-dental characters suitable
for description or comparison. Palatine vacuities
appear to have extended anteriorly to level of dP’
anterior loph, Only base of masseteric process
preserved in holotype making an estimation of size
difficult. Posterior aspect of buecinator musele scar
is laterally wide
Dentition (Figs 2-5, Table 1), Includes P*®, dP4,
M!3, excavated P*, with teeth exhibiting. only
beginning, wear.
P*®, Very small with relatively high labial crest
containing prominent anterior labial cusp and small
cuspules posteriorly. Transverse ridge joining labial and
lingual posterior cusps encloses posterior basin. Fine
iransverse ridgelets Occupy longitudinal basin, with
anteriormost slightly larger and dividing off sivall
anterior basin.
dP’: Completely molariform, Wider seross
posterior loph than anterior loph, and smaller (han
succeeding, molars,
P*: Posterior third of P’ inflated both Jabially and
fingually. Posterolubial stylar cusp is well developed
but does not attain height of posterior labial cusp,
Labial crest divided into three cuspules bordered at
both extremes by prominent anterior and posterior
cusps, ‘Transverse ridge leading, halfway into
longitucinal basin from labial crest slightly overlaps
with small ridge descending across from Ituapural crest
Fig, 4, Driwing of the holotype P* of Simosthenarus
maddock nullarbarenyis asp nov AL anterier basin: P.
posterior basin; Ly loangitucinal basin. Lit. labial crest; Li,
lingual crest; PLSC. posterolabial stylar Cusp. (Length =
13.3mm).
tA G.I
tu form anterior border uf pusterror basi, Srvall
inlerior basin separated by transverse ridge descending
Jingually from prominent antéerdlabial cusp to lingual
cnunterpart, Ridge appears i have formed foi
unifieation of two sinaller ndges descending
transversely front each antenor cusp:
MW) Molars.inerease in length und width from M1!
lo M4. (M"* not preserved), Respective widths acrass
peotoloph and metaloph in M! are identical. In M2,
protoloph wider than metafoph. and jn M’ wider
apain, Molars low crowned with lophs tiatably convex
anteriorly, Very fine crenulations on molars ape
extensive, slightly coarser on posterior side ot
menoph. Anterior cingulum well developed, hut nel
capecially broad. It extends anteriorly from up of
paracone, then labrally across almost entire width of
protoloph. Low, weak postprotocrisia extends
posterolabially from protocone and meels with very
small erest Originating from metaloph. forming acrista
obliqua, Lingual to this -structore, U deep fossette 15
presemt in median valley. Labially, @ notch ys tormed
between Well developed postpara—and prentetucristae
Posterior cingulum broad and formed by fine
posthypoerista descending posterolatnally, then lwbiutlly
verass back of molar and overtapping with less
PONTE! PostMetuerista
Farmitype: WAM 9212-7. «singular, unassouated P!
PRIDLAUX
fron Lindsay Hall Cave. Collected in April 1992 by
Lindsay Hatcher, Western Australtan Plane Caving
Group
Variatinn: 1018. Only possible to get an impressfou
of yarmition within 5. on. rullarbofensis by comparing
the P? of WAM 92,9.8 und WAM 92.12.7. They differ
Slihily in size. namely length and posterior width, bul
more noticedbly in morphology, Whereas the postertor
third of the P? In WAM 92.98 is inflated both labially
and lingually, (here is only'a lingual inflation in WAM
9212.7 (Fiy. 41. This is due largely to (he reduced
development of the posterolabial stylar cusp in WAM.
92,127 compared (6 the holotype. This is.an especially
variable character in other sthenurines, including S
m. maddocki and 8. occidentalis, and probably alse
represents intrasubspecifie variation in S$. nt.
aullarborensis, Other notable differences between the
premolars are the lateral tapering present in WAM
92.98 where the tooth becomes more nartyw
anteriorly than WAM 92.12-72 and the incipient
development of the anterior cingulum in WAM 9212.7
compared to WAM 92.9.8: Intraspecific variation in
preinalar morphology is common in all sthenurine
species tor which numerous individuals are knawn
(pers, Observation).
Comparison wilt other taxa: Clearly, the skal ot
Som. nullarbarensis was very small even considering
Fip S. Stereopair of Sumeasttenaruy madi k, nullarburehsis ssp nay. P’, mp | (WAM 9212.7, puritype), and typacal
cw.
Simastherurny maddockt enaddackt PP potton (SAM 027752),
im ocelisal v
SMALL STHENURINE KANGAROO
the difficulty involved with interpreting skull size from
fragmentary juvenile specimens. ‘The preserved
maxillary region is smaller in every respect than
similarly-aged individuals of §. gilli, previously
recognised as the most diminutive extinct sthenurine,
IL is much smaller than similarly-aged S$. yn. meddocki,
Anterior extension of the palatine vacuities and the
morphology of the buccinator muscle sear in S. rm.
nullarborensis are both similar to S. am. maiddocki.
The P? and P* of S. m. nullarborensis are
considerably smaller than S$. m. maddocki (Table 1).
Although only two P* specimens were available to
conduct one-tailed t-tests, both length (t=3.3, P=0.09)
and anterior width (t=30, P=0.0l) for Soom.
nullarborensiy fell significantly outside of the Known
range of §. mm. maddocki. As with the maxilla, they
are smaller in size than those of any sthenurine. Bused
on the comparison of mean length and width
1x7
tneasurements, they are 24% smaller than S$. m.
maddacki,
Som, nullarborensis and §. m. maddocki differ in
several morphological features of the P*. The
posterolabial stylar cusp is less prominent in S. mm.
nullarborensis, especially in WAM 92.12,7, Neither
Specimen possesses u stylar cusp which allains the
height of the posterior labial cusp, contrasting §. m7
maddovki in which the height of the stylar cusp
consistently exceeds that of the posterior labial cusp.
Tn S. nt. maddocki the labial crest is divided into either
two or three cuspules between the larger anterior and
posterior labial cusps. Both S. om. nullarberensiy
specimens possess three cuspules but a similar
variation in number could also occur in this subspecies,
In the holotype P* the anterior basin is well formed
and separated from the longitudimal basin by a
transverse ridge (Fig, 4), This structure is not present
TABLE 1. Cheek teeth dimensions of Simosthenurus maddock: nullarborensis ssp, nev, S. mi, maddeeckt and S. gilli
[mean jstandard deyaitiun) jubserved rangel]
Tooth Species L AW
Pp Sm. nullarborensis WW 49
Soom. maddocki 9,2 (h15) 6.2 0.53)
190-4. [5871]
S. gilli 10.0 (ay 7410.20
[90M 5] f60-75]
dP Sm. nullarborensis 8.5 15
Soom. maddoackt OR wih 8.7 (24)
(4.6-46] (8.4.94)
S. pili 9.2 wrt 9.2 (0.26)
(90-46) (89.96)
pi S. om. nullarborensis 12,5 11.13) 5.8 1014)
(713.3) {8.7-5.9]
Sm. maddacki 16.0 (00.59) 8.1 105i
(15.2110) (75-90)
S. gilli 16.1 (0.63) 9.2 4.54)
(52-1711 [MOR]
§. andersoni 15.5 (0/53) 1.7 3m)
14.8-46.5] [72-82]
M! Soom. nullarborensts 94 8.7
S. om. madédocki 10.8 (29) 10.) (24)
(W.341,2) 97 WS]
S. gilli 10,3 Wwos0 TO.2 (hay
(9341 [Yee 07]
M*— S. m. nullarborensis 10.1 9.2
Soom. maddovki H.2 (0.21) TOK (0.98)
JW9ALS] [ML aelt.of
S. villi 11,0 W49) LOS (aay
(10.9.9) 194 141
M3 Soom. nullarborensis 10.2 96
Som. maddocks (1.4 (2) Tl.) (45)
(NOFT] HOS-th9)
S. pillt 11.6 (0.46) 10.7 (0.34)
1W8-12,2 [IOU- 11,6]
PW AH PH n
57 55 55 I
7.3 5) 6.3 way 6.2 59) 5
J6.7-8.1| 1576.9} [57-04]
8.9 (O17) 6.5 W239) 6,7 (0,46) 7
[86-91] [od-o.9] {3-7 5)
8.2 4.9 49 I
QA) (tv 30) 50 wat 4Y way 7
[8.0.4.0] [4.65.9] 14.6-5.2]
9.6 (0.42) 4.9 (035) 4.2) IN
{9:2-10,2] {43-53} [4.95.5]
8.0 (ar) 6.5 W711 5,9 (21 2
17.5-8,5] [6,0-7.0) [S760]
1.1 (0.50) 8.3 (oa WT Wszh §
CORBI AT (77-86) {7-8}
1b (724 9.8 (ory 9.8 kx il
[W212] {MAP HR] PRAM]
9.5 (aK) 9.4 (40) Od ion) 8
j9.0-10 3] 14.9 100] [43-05]
8.7 48 4.9 |
10.0 (0.45 5.3 109) 5.2 (0.49) 9
19419) 48571 14.3 G0)
10.0 (0,43) 5,5 (0.5% 5,9 (say iH
[44-15] 14.3-6.2] [4k0.4)
8.9 5.5 6.0 l
10,3 (a4) 5.5 (045) 5.5 (that) 7
[9.9107] 4.90.2 14464)
10,2 (0 36) 6.1 (v.79) 0.6 (043) il
(9.6 10M) [56.47] fa27n
BA 47 4.5 1
10.1 (4K) ST a7) 5.5 (aK) 7
(95-107) [5-62 {50-f0)
10.3 (0.50) 6,2 (0.66) 6.5 168) i]
{9.6-11.2) {S578} 1557.9
{48 6.1 PRIDEAUX
in§. mn meddecki where mward curving ol (he anterior
exuernes uf the lubial and lingual erests necurs (Pig
5), These often do not meet and result if ait anterior
basin which is very poorly designated or completely
ubsent. A more intermediate condition ts observed in
WAM 92.127.
Complete molarisation of the dP? is eharnctenstic
of all sthenurines, Likewise, the ineregse in molar
length and relative changes in widths aloe the tooth
row in S. vn nudlarherensis are typical oF rnost species,
Molar torphology is vary siotilac tod. a. eneedelaeks
Making them readily separable from the sinilarly-sized
S. gilli, which possesses a larger crista Obliqua und
less extensive fine enamel erenulations. Therefore,
\ipper molars of the subspecies are separable on size
only, Wath Measurements of the & an nnllirborensiv
molars approximately 13% smaller than mean values
for Sn meidackl,
Discussion
Although some doubt existed mitially over which
laxonumic tank should apply to the Nullurbor form,
morphological similarily to nominotypic 8, ¢iadddoeks
precluded w specific separation. Designation of fussil
subspecies has previously been made in re(crenee wy
dwarfing macropod lineages (Marshall & Cofeuceiny
1978, Dawson & Flannery 1985). They ure aneffective
form of recognising clear-cut temporal of geagraphical
variants within a species. Size reduction m PF? and
molar dimensions (24% aid 13% respectively) from
Som maddacki to 8S. in nullarborensix 1s comparable
to that observed by Marshall & Corrnecini for other
jnediuin to large-sized macropods fron the Pleitacene
io Holocene, They concluded that resource limitations
probably uecounted for dwarfing im these macn ipods
and Sareophilus herrivit, wart ts commanty
observed in island populations conlempurancous wah
large manmnal populations on the mainland (Lenina
1985), A similar ctfect between mainland regions
caused by climatic unpredictability, whieh today
typifies he Nullarbor Plain, may also have been
siumiicant in the Pleistocene environment, perhaps
louding to size reduction in §, yi. nullarborensis
Inierestingly; analysis of size variation in /iylecinas
eynovepludus fram the Nullarbor Plain (.owry 1972)
did not statisticully separate fas u subspecies) the
population sample from larger 2 eyrocephalas, even
though several simall individuals were present
Unforunately, the fick ofS, ot nellarberensix material
ul thts stage prevents a more thorough statistical
analysis thin that conduewd for the permanent
preniolar. Flowever, dimensions of most meusurements
fall well outside the Known range ofS. m, maddoacki
SUppOTtIN, Hts current designation as a separate
subspecies
A probably tate Pleistocene age is attributed to the
Lindsay Hall Cave deposi, hased on the sinnlarily of
Som aullarborensis Ww Som. mneddockr, and the
co-occurrence of another widespread, but undeseribed
sthenuring, The hone-bearing strata of Lindsay Hall
Cuve muy equate with unit 2 in Madura Cave
(Londelivs 1963; Lundclius & Turohull 1989) which
has also Vielded this undescribed species. Support lor
this would rely on a detailed straugraphie investigation
of cave deposits in the area
Acknowledgments
Thanks to Dr Rod Wells (Flinders University), Dr
John Long (W.A. Museum) and an anonymous
reviewer for offering comments on the manuseript, Dr
Long ulso provided Fig, 4, and with Dr Alex Baynes,
wos responsible for loun ol the fossil material, bam
wyratctul to the Western Australian Plane Caving Group
for (heir efforts, particularly Wenuly Binks, Katherine
Crisp and Lindsay Hatcher who discovered the deposit
and collected fossil material from the suc.
References
‘
Dawson. Lo & Foannery. J) F OY8S) ‘lixononmic and
phylogenetic status of living aml fossil kungarins and
wallabies of the genus Macripuy Shaw (Macrapodidae
Marsupiala) will) @ few subgenerie name for the large
wallabies, elusi, J, Zeal, 33, 473-498
Fiannery, Tb & Hove, 3. A, (1983) Oecurreness at the
extinet macroppdid marsupial Semesthenuras eaddock! in
New South Wales. dase Masui, 6, 27-39.
Lomouso, M. V_ (1985) Body size of maninnuls an tslunds:
the land Role roxamined, Amer Net $25, 3D Sif,
Lowry. J. W. J. (1972) The taxonomie statues tf sinall fossil
Ibylacies (Marsupiili, Thylicinidae) from Westerti
Australia, bo Sue We aust, $5, 19-29.
Lewprenios, BE. b., Sr, 1963) Vertebrate remains from the
Nullarior (wes, Western Australia, (if, 46, 75-80)
& ‘Toraneta, WB, (889) The mammalian fives
o! Madurs Cave. Western Atarralia Part Vib:
Moenopodidae: Sdienurinad, Macropochinae willl review
of (he marsupial portion of (he fauna, Fleldtana, Geol.
Wins), 7h
MarsHatin L. 6) & Corruccini, Ry S. (1978) Variability.
evolubanury nes, und ullometry in ulwarfing lineages,
Falvebtology 4, (OWL,
Prener, NS (980) Macrapodid skeletons, including:
Simostheaurns Tedford, fron. an unusual “drowned cave”
deposit jn the southeast of South Australia. Ree: 5, Aueyi.
Mus, 18, (el,
Tronarp. R.H, (1966) A review of the macropodid genus
Sthenurus. Univ Calif, Pabl Geel. Sei. 87, 1-72,
Wrais RT & Morray, Bb. (1979) A new sthenurine
kangarou (Mansupiala, Maenspodidae) from southeastern
Sout Australia. Trans. R. Sec. 8. Aust 103, 213-219.
Mirkiariy, Ko & Wirttams, DB. LG. (1984) The
fassil vertebrate deposits af Vietoria Fossil Cave Naracoorte:
an introduction to the gealoey wid tune, tas) fool 2b,
US $53,
THE COOK 007 METEORITE:
A NEW H4 CHONDRITE FROM SOUTH AUSTRALIA
By M. ZBIK*
Summary
Zbik, M. (1994) The Cook 007 meteorite: a new H4 chondrite from South Australia.
Trans. R. Soc. S. Aust. 118(2), 139-142, 31 May, 1994.
The Cook 007 meteorite, a single stone of over 100 kg in weight, was found at Cook
in 1989. It has been classified as an H4 chondrite of shock facies S5-6 and contains
olivine (Fajo5,.93 n = 30), orthopyroxene (Fs,,5,,. n = 15), clinopyroxene maskelynite,
nickel-iron and troilite. Mineral compositions and textures indicate that Cook 007 was
a metamorphosed part of the H-meteorite parent body and was very strongly shocked
before reaching the Earth.
Key Words: Cook, meteorite, chondrite.
Transactions of the Royal Socrety of S. dust. W994), LBD), 139-142
THE COOK 007 METEORITE: A NEW H4 CHONDRITE FROM SOUTH AUSTRALIA
by M. ZBIK*
Summary
Amik, M, (1994) The Cook 007 meteorite: anew Ha chondrite from South Australia, Vea, Ro Soe S dur
H8(2), 139-142, 31 May, 1904
The Cook 007 meteorite, a single stone@of over 100 ky in weight, was found al Cook 1 1989, If has been elussified
as an H4 chondrite of shock facies 85-6 and contains olivine (Hayy sg) 0 = 30), orthopyroxene (Psj754) 70 = 15).
clinopyroxene maskelynite, mckel-iron and troilite, Mineral Compositions and textures indicate that Cook 007
Was a metamorphosed part of the H-meteorite parent body and Was very strongly shocked before reaching the Earth,
Key Woros: Cook, meteorite, chondrite.
Introduction
A single mass of the Cook 007 meteorite weighing
over 100 kg, was found at Cook on the South Australian
part of the Nullarbor Plain (Fig. 1), in 1989. The
approximate co-ordinates of the locality are 30°37'S.
130°25'E. The meteorite was collected illegally and
exported to the United States of America. The South
Australian Museum obtained a piece from an American
Ineteorite dealer, Mr Allan Lang in Apmil ot 1991,
In recent years the Nullarbor Plain has proved ta be
a productive area for the recovery of meteorites (Bevan
NULLARBOR PLAIN
. *Ooldea
® Cook
Nullarbor
*
Gulf St. Vincent
KANGAROG
ISLAND
Fig. |. Map of South Australia showing the approximate
location of the Cook O07 meteorite.
* Polish Academy of Sciences, Space Research Centre,
Warsaw, Poland
1992; Bevan & Binns 1989a, 1989b) and in the Last few
years has altracted the attention of illegal meteorite
collectors, Under legislation enacted by the
Government of South Australia, all meteorites found
in the State are the property of the South Australian
Museum, An unfortunate consequence of the iflegal
trade in meteorites is the loss of important information
on the exact date and location of the find. In accordance
with the guidelines on the nomenclature of meteorites
rom the South Australian Nullarbor (Bevan & Pring
1993), the meteorite has been named Cuok (07, being
the seventh meteorite to be recorded from the Cook
wren,
Physical description
The piece of meteorite obtained from the United
States weighs 26.2 kg. It is irregular in shape and is
about 35 em in length, 50 crm in width, 4-1 em thick
and polished on one side, The specimen is currently
on display in the meteorite exhibition in the South
Australian Museum. The picee has a dark brown | inm
thick Weathering Crust covering (he outer surlave. ‘The
interior of the stone shows fio sign of weathering, but
the heavy weathered crusty covering the surface
indicate that the meteorite had been exposed to the
elements for many years. A number of quartz grains,
probably of aeolian origin, are incorporated into the
weathered crust, The interior of the meteorite is black
in colour and medium to fine grained. In thin section
the meteorite is generally dark coloured (Fig. 2). The
chondrules and chondrule fragments are partly
recrystallised and well defined boundaries are
recognisable even without using crossed polars. They
are typically less than 0.5 mm in diameter but some
chondrules measure up to | em in diameter. Metal and
troilite occur us finely disseminated grains throughout
the matrix. A slice was cut and a polished thin section
was prepared and used for petrographic examination
and electron microprobe analyses.
140
Meteonte (length of rhe section 5 cm) showing hirge dark
(staining) area and granular structure full of chondrules
and chondrule fragments.
Mineralogy
Compositions of the silicate minerals were
determined with a JEOL electron microprobe at the
University of Adelaide Centre for Electron Microscopy
and Microbeam Analysis. Analyses were made using
an accelerating voltage of 15 kV, a sample current of
3 nA, and a beam width of 5 xm. Mineral anilyses
are presented in Table |.
Well-defined chondrules and chondrule fragments
are composed predominanily of olivine and
orthopyroxene. “Barred” chondrules are composed of
olivine or olivine and orthopyroxene and contain thin
lamellae of feldspar glass. Fig. 3 shows a fragment of
a large barred chondrule which contains bars of olivine
and turbid lamellae of glass. The glass has a bytownite
composition, and is probably maskelynite. Some large
and strongly shocked olivine grains that occur as bright
Fig. 3. Photomicrograph of the Cook 007 meteorite in thin
section, showing frayment of barred olivine chondrule
(about | mm in diameter).
purple crystals under crossed nicols are unusual and
need further investigation. Chandrules composed of
radial pyroxene crystals. are also present (Fig. 4),
Several of these display a cryptocrystalline structure
(Fig. 5) with strong wavy extinction. A number of
granular olivine-pyroxene chondrules and porphyritic
pyroxene chondrules are present. They contain some
coarse, euhedral olivine grains and have a poikilitic
texture. These olivine grains contain a network of
planar fractures (Fig. 6), which are filled by troilite
and metallic Fe, Ni. The occurrence of strong mosaic
extinction, solid state recrystallised areas and abundant
TABLE |. Average chemical compositions ef majer minerals in the Cook 007 meteorite.
olivine orthopyroxene wt% clinopyroxene = maskelynite
oxide WL eore rim WL Wh
THs 4 2.8. 30 15 ie) 30 Il
SiO, 37.8 53.5 53.6 $1.3 47.1
9] 9 Tn 0.1 0.1 0.2 0.7 0.1
AlsO4, = 0.3 v4 1.3 3U,7
FeO . 13.1 7 9.9 | 0.6
Mno 5 V5 05 03 0.1
MgO 41.9 30.2 27.0 189 is)
CaO 0.03 0.5 4.8 17.0 13.2
NaQ, - - - 0.6 2.8
K,0 - = i)
C0, - 04 06 0,9 —
1 {+)"| ene poe es Oe ' 98.3 UTA 96,7 96.1 96.4
Fajo.s Fsjy5 FSj5 5 Fay 4 Abo) 7
Engg 7 Evra 4 Enss,6 AN.
Woro Wop 5 Wo36.0)
THE COOK 007 METEORITE 14]
Fig. 4. Photomicrograph of the Cook 007 meteorite in thin
section, showing a radial pyroxene chondrule (about
0.5 mm in diameter).
Fig. 5. Photomicrograph of the Cook 007 meteorite in thin
section, showing a cryptocrystalline pyroxene chondrule
(about | mm in diameter).
= ’ :
Fig. 6. Photomicrograph of the Cook 007 meteorite in thin
section, showing planar fractures in olivine crystal (about
0.2 mm in diameter).
opaque “mixed” melting which occurs as veins and
pockets within the matrix indicates that the meteorite
has undergone a high degree of shock metamorphism.
Distinct darkening referred to as “shock blackening”
(Heymann 1967) is a common feature of this meteorite.
Polycrystalline “mixed melt” material surrounds
“islands” of unmelted but partly recrystallised crystals.
Several porphyritic olivine chondrules display a zoned
mineralogy. Euhedral olivine crystals are in contact
with low-Ca pyroxenes, with calcium-rich pyroxenes
present as thin rims on these and also as fine needles
set in the glass of plagioclase composition. The matrix
consists of melts, melt pockets and veins, forming a
network of complex branches surrounding the
unmelted chondrules and chondrite matrix. This
indicates in-situ melting of the host material and shows
shock blackening. Such a structure has low porosity.
The absence of a network of joined intergranular pores
protects the interior of the meteorite from being
weathered (Zbik 1982). All feldspar present has been
changed to maskelynite which is abundant as small
turbid patches throughout the matrix. Nickel-iron metal
and troilite occur as accessory minerals.
Microprobe analyses show that the olivine in the
Cook 007 meteorite is equilibrated with a mean fayalite
content of Fay 5,030 = 30, The orthopyroxene shows
only a small variation in chemical composition with
a mean ferrosilite content of Fs,75,;. 1 = 15 and a
wollastonite content of 1 mol%. Some pyroxene
phenocrysts contain calcium-rich rims. These rims
have a composition of Fs,;, with a wollastonite
content of Woy; mol%, The composition of the
clinopyroxene needles within the glass is Wo369 Enss¢6
Fsg 4 (30 analyses) (Fig. 7). The maskelynite glass has
yielded a composition similar to bytownite (ll
analyses). The poor quality of microanalyses is a direct
result of the high degree of shock metamorphism.
The pyroxene geothermometers of Wells (1977) and
Lindsley (1983) suggest that the Cook 007 meteorite
was heated to temperatures of between 600°C and
700°C during metamorphism while the meteorite was
CLINOPYROXENE
30 analyses
ORTHOPYROXENE
15 analyses
Fig. 7. Silicate mineral chemistry in Cook 007 meteorite.
Isotherms show temperature calculations based on the
calcium content of orthopyroxenes and clinopyroxenes
(Lindsley 1983).
142 M, 4BIK
sull part of the large H type of the meteorite parcnt
body. After this the meteorite was probably remelted
in a major part by impact and heated to a temperature
between 00°C and 1200°C, as indicated by the
chemical zoning of pyroxene grains.
Classification
The Cook 007 meteorite has been classified as an
H4 chondrite. The olivine composition (Fajg 5,4) 4) is
within the range of the H chondrites (Keil & Frediksson
1964). The highly equilibrated mineral compositions,
crystalline matrix, and a well defined chondrule
boundaries, suggest that Cook 007 belongs to the type
4 classification of Van Schmus & Wood (1967), The
wollastonite content of the orthopyroxene is similar to
that found in other H4 chondrites (Scott ef al. 1986),
The presence of undulatory extinction, the strong
planar fracture formation in olivine, plagioclase
feldspar altered to maskelynite, occurrence of
mechanical polysynthetic twinning in orthopyroxene
and the degree of crystal fracture all indicate that the
meteorite was strongly shocked after metamorphism.
The shock facies is estimated to be S5-6; strongly to
very strongly shocked according to the classification
scheme of Stoffler er al. (1991),
Acknowledgments
The author wishes to thank Mr Huw Rosser of
CEMMSA, University of Adelaide, for assistance with
the electron microprobe analyses. Mr B. McHenry is
thanked for helpful suggestions to the manuscript. This
manuscript was also improved by the helpful comments
of the reviewers Dr Margaret Wallace and Dr Alex
Bevan. The financial support of the Friends of the
South Australian Museum is gratefully acknowledged.
References
Bevan, A. W. R. (1992) Australian Meteorites. Rec. dust.
Mus, Supplement 5, \-27.
—__ & Binns, R. A. (1989a) Meteorites from the
Nullarbor Region, Western Australia; 1. A review of past
recoveries and a procedure for naming new finds.
Meteuritics 24, 127-133.
& __ (1989b) Meteorites from the Nullarbor
Region, Western Australia: II. Recovery and classification
of 34 new meteorite finds from the Mundrabilla, Forrest,
Reid and Deakin areas. hid. 24, 135-140.
& Princ, A. (1993) Guidelines for the nomenclature
of meteorites from the South Australian Nullarbor. /brd.
28, 600-602.
HrYMANN. D. (1967) On the origin of hypersthene
chondrites: Ages and shock effects of black chondrites.
Tearus 6, 189-221.
KeiL, K. & Freprixsson, K. (1964) The iron, magnesium
and calcium distributions in coexisting olivines and rhombic
pyroxenes of chondrites, J. Geaph, Res. 69, 3487-3515.
LinnsLey, D. H. (1983) Pyroxene thermometry. Am, Mun.
68, 477-493,
Scorr, E. R. D.. Taytor, G. J. & Kem, K. (1986)
Accretion, metamorphism, and breeciation of ordinary
chondrites: evidence from petrologic studics of meteorites
from Roosevelt County, New Mexico. Prac. Lun, Plan.
Se. Conf. 17, EW5-E123.
Srorrver, D., Kei, K. & Scorr, E. R. D. (1991) Shock
metamorphism of ordinary chondrites. Geach. Cosm. Act
55, 3845-3867.
Vas Scumus, W. R. & Woop, J. A. (1967) A chemical-
petrologic classification for the chondrite micteorites,
Geach. Cosm, Acta 31, 747-765.
Wetxs, P. R. A. (1977) Pyroxene thermometry in simple and
complex systems. Conrr, Min, Perr, 62, (29-139,
Zpik, M. (1982) Pore Spaces in Stony Meteorites. Bull,
Acad. Pol. Se. Terre 30, 59-65,
A MAJOR RANGE EXTENSION AND NEW ECOLOGICAL
DATA ON OXYURANUS MICROLEPIDOTUS (REPTILIA:
ELAPIDAE)
BRIEF COMMUNICATION
Summary
The inland Taipan (Oxyuranus microlepidotus) is a large elapid snake endemic to
Australia. Considering both venom toxicity and average venom yield per bite, O.
microlepidotus is the world’s most dangerous snake’. Despite its size and medical and
scientific significance, the status and distribution of this snake, have been difficult to
ascertain. After being described in 1879, O. microlepidotus was not found again until
1974°, Following its rediscovery, it has been recorded from the channel country of
Cooper Creek and Georgina and Diamantina Rivers of south-western Queensland and
north-eastern South Australia’. O. microlepidotus is a rarely seen snake because most
of its life is spent in rat burrows”.
Transacnions of dhe Revel Sacre of §
BRIEF COMMUNICATION
desi, (W994), TIBE2), 145 145.
A MAJOR RANGE EXTENSION AND NEW ECOLOGICAL DATA ON
OXYURANUS MICROLEPIDOTUS (REPTILIA: ELAPIDAE)
The Inland Taipan (Otyereenis mfervlepidomes) is a large
elapid snake endeiie to Australia. Considering both venom
toxicity und average venom yield per bite, Qo riervlepuidors
iy the world’s most dangerous snake!. Despite its size and
ynedical and scientific significance, the stalus and distribution
of this snake, have been cifficull to-ascertain. After being,
described in 1879, OL mierolepidaras was not found again until
\U74-_ Following its rediscovery, it has been recorded Irom
the channel country of the Cuaper Creek and Georgins and
Diumantiog Rivers of south-western Queensland and north-
eastern South Australia®. Q. micralepidimuy is a rarely seen
snuke because most of its life is-spent In rat burrows
In April 1992, a large elapidl was collected on the Coober
Pedy to William Creek Road (2993'S, 135°03'E) (Fig. 1)
hy local contractor Jef? Boland. He recogrused that the siuake
was different from the Western Brown (Preudonaje nuchatis)
and Mulga Snake (Peeudechiy caestratis) with which he was
very funnhiar Jeff Boland subsequently collected a sloughed
skin from the Moon Plarn (28°52'S, 134°50'E) (Pig. L) and
sent both the specimen und the slough to the author,
Mt Barry vA
Lora Cre a
aigepulloultt
— _—
Coober Pedy
*& 0. miéraiepidotus iocalities
Fig, | New records of Grywruis mierelepadetuy in (he
Coober Pedy region.
The dorsal scales of the specimen were uniformly dark
brown with the head nearly black. ‘The xpecitien and the
slouyt had 23 mid-body scale rows and a single anal seale,
These characteristics enabled the snake und (he slough lo be
confidently Wenufied us O. micwlepideoms. The sealation of
the Coober Pedy Specimen closely matched that or Q
micralepidotus speaamens fram Moomba excepr that naher
than the lower primary lemporal scale extending to the lip
between the 5th and 6th labials, this scale bad fused with the
6th labial in the Coaber Pedy specimen. laspeetion of other
specimens in the South Australian Museum indicates that the
status of the lower primary temporal scale is considerably
plastic in CO. anicralepidars
O. microlepidoms can be further distinguished fron the
sympatric. und highly variable P nuchal’ in having a longer
head, srmiller and more numerous sucha) scales, a pronounced
canthus and plain ventral seales as distinct fram spotted vr
dark edged ventral scales in P nuechafis, Inaddition, most
P ntichalis trom within the range of the QO) microlepidotus
eshibit irregular black spots or black crossbands. wheres
these bands were absent from the Coober Pedy and Moomba
specimens of the Q. mucrilepitonis.
Ii the winter of 1992, two noad-killed O. mlcrolepideties
were found on (he Coober Pedy (o Mt Barry Road (28°34'S,
199°54'E, 28933'S, 134°95'E [SAM R40464)). a live
specimen Was sighted in the Dingo Ferce four kilomenes
south ofthe original locality (29°905'S, 125°03"E) and a further
specimen was killed at Torn Cat Hill (29°00'S, 134945'E)
in Coober Pedy (Pig. 1). A further vad killed specimen way
Jocated on the Coober Pedy to Mt Barry Road (28°57'S,
134°47'E [SAM R42484)) 19 September 1993. South
Australian Muscui records are denoted by the prefix SAM,
On September 6, 993g live C2 wrerodemidotus was captured
by Jeff Boland on the Moon Plaia, adjacent to the Breakaway
Reserve. The following week a road killed specimen (29°00'S,
135° 0'F) was collected approx taiately 40. kin east of Coober
Pody,
The live specimen was captured in the rain wath a stron
wind blowing und a recirded temperature at Coober Pedy
ol only 18°C. This snake was uniforidy black on the dorsal
suclace with & white, unmarked ventral surface. One month
uller ifs vaplure the snake slouvhed. revealing a yellowish
belly and a dark brown dorsal surface with black cdges to
some svales producmy a siucht herring -bone pattern. The head
remained a glosyy black colour following the slough
The © microlepidotus from Coaber Pedy region were about
500 kin from the nearest known O. ¢icrolepidonts locality
at Goyder Lagoon, in north-eastern South Australia. The
Coober Pedy population of Q. micrelepidotus is separated
from that of the channel country by the huge salinaof Luke
Eyre and the dunefields of the Simpson Desert. The discovery
of O. microlepidotus near Coober Pedy is highly significant
as tt represents the first Known qeourrence of the species near
aconsiderable populavion centre afd also raises the possibility
ofa much more expanded range than previvusly recognised
lor (his imponant snake
Jee
The Coober Pedy specimens of 0, micralondoins wore
found from April September, Interestingly, the Goyder Lagoor)
popufation of © qurerufepidviay fas been recorded
predominantly in Maretvand Apel? whereas Covacevich*
reports thar O, aerelepidoms wily emerges lar 2-3 weeks
in late-winter-early spring, Althoutr Covacevich? indicates
that 2 mierolepideras are predominantly active oa still days,
Mirtschin® collected several mdividoals an windy days. The
capriye of the specunen near ihe Breakyways Reserve on a
cold, wel, windy dity indicates (hat these saakes may be
encountered at any Time throughout auntie, winter and
spring. CL microlepidotus are possibly ulse achive in cracks
and manna) betes daring the wartier mraychs but aire et
recorded because Ihey de not need ty come lo the surface
to bask.
GO omniertepidetas pecords fond the Coober Pedy region
were found in two diffeccnthabmaws, Kive-ol rhe records were
from cricking eypseous sails. Fourof these reeerds were from
a line areal wy pseois soils knows as tite Moon Plan. These
sails Suppert little or na perennial vegetiuion. Cover and
species diversity ob andital vegelaiien’ valy dninatieally
depending on the season. The dominant, albeit sparse,
vegotnon din October 1992) consisted of Alniples
spongiosa, Sulsdle kali, Helinterum florihindim, ialacna
lepialegis, Aralidella Nasturtium and the wrasses Faneapogurt
polyplioitus. Peernicuny deceompositum, Aristide
anthoxanthoides, Astrebla peciinaiat, Enigrastts verifolte: and
& dielyiz. Vegetanion cover Was preatost ithe small puneen
areas in the undulating plain, These low lying areas were also
mre most Heavily eracked.
Two other records were tony gibber country darpireted
hy the Oodnadatta Saltbush Aripker renuilarta, Onher
commen plants meluded Selerofuena inidvuta. Atriples
spongiosa, Searcostema and Salsola ele, Small gilgar
SeprensiOns within this habitar contained cracking ebay’ and
were Vegetdled with several griss and daisy species.
Che plains country around Coober Pedy is very
heterogeneous and the Wwo habitats described above oflea forts
ammosate with the gibber more common on high around and
the gypseous sails usually in low tying country, Therefore
snakes. this region may depend apan, or preter, either hatirat
and yet be tecorded crossing between areus of favured Habitat.
The cracking gypseous soit habitat i very: similar to the
prelerrec habitar described for ©. micrelepidenis at Clifton
Hills, South Austria’ and ig Queensland” although the
have also heen récorded front gibber plains und sand dunes”,
These cracking plains therefore appear lo be the key habilal
for the OL microfepidorus. The cracking gypsebus plains are
quite widespread near the south-west margin of the Lake Lyre
RBasiiand are offen associated wilh cun off vones tron break-
away country. These plains are (ften inter-connected by creeks
which possibly also provide appropriate habit
Interestingly the Moon Plain and surrounding pypseous
rewions were oot identified as potential sites tor ©
microlepideins based oa climatic indices’. Habmar and
biological mndices, rather than climatic variables may (heretone
be more iraportant than climite in deterntining the range of
(2h miprolepidones,
The eastern population of QO: microlepidamy feeds
extensively on the Long-huired Rat (Rartuy wllosistrnies) wrth
which its ecology and distribution have beon inuxtrivably
linked’, The evolution ofa large size, a rapid Snap-release
bite and extremely potent venom in Qo micrelepidetics is
believed to bed response 10 Ther predation on mkontials whies
ca defend themselves by bring savagely! Uherefiny.
particular ermphasiy was placed on searching lor &,
villastsitnes Of ON sanitlar saree mearimals: (yp tie Coles
Poly twpion, lo predit the potential range of @
microlepidonss,
Tield surveys were condicted during 1992-1993 in cracking
Chay regiotis adjacent wi the OL mierolepidéons records uti ML
Barry, Anna Creek, Billo Kalina and Stuart Creek Startoms
fouscertain whyther O. microlepidotuy, or their prey species,
occupred a more extensive range, The serikimg feature af the
cracked pypseaus sil localities was the crverstty of manimals
and paucity of reptiles. OF particular note was the presence
of Planns Rats (Prenderis austealls). Although A utllisisirenes
reached the Coober Pedy region m (97374 (1. Boland. Kk.
Greenfield pers, camny,) they do not uppear to ynhabir rhe
ego normally and were nol recorded ia this supvev /
australis, Corresr’s mice (Lexpardita forrest), Desert Mine
(Prendvnes desertors, House Mice (Mis domesrrinss,
Poueiert Phonwates (Planikele viltesi) and Buna
(Sminthopsic macroura, 8. crassicaidara) were wweuled over
a wide range, Maynval duasities were higher in Ure cracking
gypseous Sous than in other habitats in the region®., While
The ado) OL wicratepidetys probably teed predominantly op
P wtestralls, L. forrest’ and the oveasional bind, juvenile snakes
vould teed on P oyvilesi and MM dumesticis
Beoalise eight Confirmed fecords now exist lon dierent
Jovalities, initia) suspicicns thal the specimens represented
snakes that were wtificially tanslocated from rheir orginal
nie van probably be discounted, However, itis not known
whether this apparent tinge expansion is a recent
Phenomenon, facilimued by mobilicy of The nomh-easien
population, or whether QO, yicrofepidoras tive occupied Ihe
plains anuAad Coober Pedy for a long period, Considering
the dependence an the &. villoyisemuy in the channel country,
4 porenhal scenario ts hair the G2 mferolepidoms tollowed
the nels from north-eastern South Austnuta thigh te Coober
Pedy during the nit plaphe of WG. (his vinlikely, however,
that GO. ieralepidetas could have expanded its nine by over
400k dure the cate of a single rat plague. A note
feasible explanabon 1S that Qo micratepidetay have alway
inhubited! the gypsecus pluins around the southwestern margin
of te Luke Ewer bast But the sparseness of the huni
pepulation combined with the predominantly uaderonyund
abit of the snake fave accourted lor tne paucity of records,
These sume factors probably explain why OQ. microlepidotus
eluded discovery inthe Birdsville repion forse tong, Increased
4nuke numbers ds dcresult of linge mimmal pypulations which
have responded to good seasons, combined wilt mercased
Vigilance and awareness of local residents probably explain
why the O. mtreralepidatus Wis cuscovered in the repicn mn
1992, A continued interest in this species is predicted (a result
In the discovery of OL micrealepidetis over a broader range
inoneky Pypseous coomery to the south and west of Lake
Eyre.
This paper is dedicated to Jef Boland whose mtvrest ated
invereniess resulted inthe discovery of O, qicrolepidoruy new
Coober Pedy. Thinks also yao to Miek and Ben Evins, Gree
Stazbouer, Peter Paisley, Bee Mupsared, Katherine Moseby,
Steve and ‘Tania Green, John Fewster and Zoe Rowen tor
axsistanice With field work and io Mark Hutehingen. leanete
Covacevich and Peter Mirtschin fur help in the preparation
at this paper, Frank Badman assisted with plant identification.
'Broad, A. J., Sutherland, S. K., & Coulter, A. R. (1979)
Toxicon. 17, 661-664.
2Covacevich, J., Peam, J. & White, J. (1988) Venoms
and Victims. Amphion Press and Queensland Museum:
Brisbane.
3Covacevich, J. (1993) Journal of the International
Herpetological Symposium.
J. L. READ, 3D Eyre Court, Roxby Downs, S. Aust. 5725
145
4Mirtschin, P. J. & Reid, R. B. (1982) Trans. R. Soc. S.
Aust. 106, 213-214.
SMirtschin, P. (1981) Herpetofauna 13, 20-23.
Shine, R. & Covacevich, J. (1983) J. Herpetol. 17, 60-69.
7Longmore, R. (1986) Australian flora and fauna series 7.
Bureau of flora and fauna, Canberra.
8Read, J. L. (1992) Rangel. J. 14, 145-156.
GROWTH IN THE AUSTRALIAN BURROWING FROG,
CYCLORANA AUSTRALIS (GRAY)
(ANURA: LEPTODACTYLIDAE)
BRIEF COMMUNICATION
Summary
The majority of known Australian fossil frogs have been identified by the
characteristics of the ilium, a distinctive bone in which the morphology varies greatly
between family, genus, and species'*. Hence it is now used extensively as a
diagnostic tool*®.
The physical characteristics of the frog can be identified from the size and points for
muscular attachment on the ilium. These features represent adaptations to the
environment. For example a short ilial shaft is characteristic of a burrowing frog
which does not make long jumps'’**.
Transactions of the Reval Sectery of S. Aust, (1994), 1IR(2), 147-148,
BRIEF COMMUNICATION
GROWTH IN THE AUSTRALIAN BURROWING FROG, CYCLORANA AUSTRALIS (GRAY)
(ANURA: LEPTODACTYLIDAE)
The majority of known Australian fossil frogs have been
identified by the characteristics of the ilium, a distinctive bone
in which the morphology varies greatly between family, genus,
und species! *, Hence il is now used extensively us it
Uing nostic tool
The physical characteristics of the frog can be identified
from the size and ports for muscular atlachnient on the ilu,
These features represent adaptations to the environment, For
example a short ital shaft is characteristic of a burrowing
frog which does not.make long jumps)?"
‘To permit extrapolation of the size of a frog fromthe length
of the ium requires un understanding of the relationship
between the ilium length (IL) and snout to vent length (S-V).
I is generally considered that the itu grows linearly 0
relation to the S-V of the donor animal’,
However, the dala from which these assumptions ure mude
have been based on small “ng sizes and little is Known
of carly ontogenetic changes! 7. [is also (portant to note
that it is commen in the Animal Kingdom for a change in
sive lo result ia change of the proportions of the body. A
simple example of such allometric growth can be seen during
the development of human beings, in which a large change
in the proportions of (he limbs and the head, in relation tu
(trunk size, cun be seen when children are compared witht
adults”,
This study was un attempt to determine the quture of growth,
from juvenile to adult form, of the burrowing frog Cyclorartr
australis (Gray, 1842) via the relationship between ilial length
and snout-vent length.
The jlia exumined were dissected from 24 preserved
specimens in (he collections held ul (he Department of
Zoology, University of Adelaide. Before dissection the S-V
of eavh specimen was measured witha pair of NSK eleetronic
digital calipers. The pelvis was (hen removed, and soukeu
in bleach to loosen musele, betore being transferred to a 47°C
oven lo dehydrate for 24 hours. The UL. for each iliam was
measured using electronic digital callipers, with the aid of
D prem
Lt pfotut \ Daeer exp
Wiel stant ™
\
Nestea
ee a eee 5
— a
Proven sy
‘Tubes
~ AcbINhuluny
ay
Fig | Leti lateral aspect of Cyclorana australis pelvic girdle
(LL = 306 mm, $-¥ = 82.1 mm),
Abbreviations: D, acet, exp__dorsal acetabular expansion,
D prom, dorsal prominence; D. protub._darsal
protuberance, pre-acel, zone__presacetabular zone,
fechinity
a Wild M3 dissecting microscope. Iial length was expressed
as the distance between the tip of the dorsal acetabular
expansion, and the end of the ilial shaft!
The characteristics of the iliurm of the upecitnens examined
did nol differ significantly from those described! (see Fig,
Ll). The $-¥ ranged 20.0 mm to 83.1 mm, and IL. 6.1 1m to
32,0 mm
A linear regression comparing IL with S-V showed a direct
relationship between the body length and the length of the
iim, ie. the ilium grows linearly in relation to the body
length throughout ontogeny (see Mig. 2),
The lineur growth of C. australis permits a very accurate
method for estimating the size of an individual from a
disarticulated ilium.
1am greatly indebled to Prof. M. J. Tyler lor the
opportunity (6 undertuke this research and lor constructive
criticism of the early drafts, and Dam extremely grateful for
the assistance from K- Maurice-Jones in the production of
the figures,
|
| 2910 = Dnbda4 OHa72 72 logan X |
1.90 r Pliers es easy na24 ~
—E 7.80 |
aS
= > .
is) .
5 1.70 - /s *
e
/ *
Te pf
— -60 -
ie) .
c. °
ep) .
Qa 7 50 “| / *
> /
i) fp
| la
*
1.40 4 ° f
/
| /
1:30 +—+, 1
T T ~ ’
0,7 0.9 14 1.3 1.5
(09; Ilial length (mm)
Fig. 2. Regression line of ilium length of Cycloratia australis
With snout-vent length. For = = 1830 mm, y « 51.25 mm
(95% confidence limits = 49,36 53,23),
148
'Tyler, M. J. (1976) Trans. R. Soc. S. Aust. 100(1), 3-14. *Trueb, L. (1973) pp. 107-108 Im Vial, J. L. (Ed.)
“Evolutionary Biology of the Anurans. Contemporary
Tyler, M. J. (1982) pp. 159-168 Jn Rich, P. V. & Thompson, _ Research on Major Problems” (University of Missouri Press,
E. M. (Eds) “The Fossil Vertebrate Record of Australasia” | Columbia).
(Monash Univ. Press, Clayton). Tyler, M. J. (1993) Pers. comm.
Bogin, B. (1988) “Patterns of human growth” (Cambridge
5Tyler, M. J. (1990) Mem. Qld. Mus. 28(2), 779-784. University Press, Cambridge).
STEVEN J. WALKER, 12 Preston St., Blair Athol, S. Aust. 5084
VOL. 118, PARTS 3 & 4
30 NOVEMBER, 1994
Contents
Transactions of the
Royal Society of South
Australia
Incorporated
Davies, M. & Watson, G. F. Morphology and reproductive biology of Limnodynastes
salmini, L. convexiusculus and Megistolotis lignarius (Anura:
Leptodactylidae: Limnodynastinae) - - - - - -
Rounsevell, D. E., Ziegeler, D., Brown, P. B., Davies, M. & Littlejohn, M. J. A new
genus and species of frog Peary Spee ee te oe
from southern Tasmania - -
Pamment, D., Beveridge, I. & Gasser, R. B. The distribution of nematode parasites within
the stomach of the western grey kangaroo, Macropus fuliginosus -
Ainslie, R. C., Johnston, D. A. & Offler, E. W. Growth of the seagrass Posidonia sinuosa
Cambridge et Kuo at locations near to, and remote from, a power station
thermal outfall in northern Spencer Gulf, South Australia - -
Molsher, R. L., Geddes, M. C. & Paton, D. C. Population and reproductive ecology
of the small-mouthed hardyhead Atherinosoma microstoma (Giinther)
(Pisces: Atherinidae) along a salinity gradient in the Coorong,
South Australia - - - - - - - - -
Hutchinson, M. N., Milne, T. & Croft, T. Redescription and ecological notes on the
pygmy bluetongue, Tiligua adelaidensis (Squamata: Scincidae) -
Fuller, M. K. & Jenkins, R. J. F. Moorowipora chamberensis, a coral from the early
Cambrian Moorowie Formation, Flinders Ranges, South Australia -
Peterson, M., Shea, G. M., Johnston, G. R. & Miller, B. Notes on the morphology
and biology of Ctenophorus mckenziei (Storr, 1981) (Squamata:
Agamidae) = - - - - - - - - -
McKillup, S. C. & McKillup, R. V. Reproduction and growth of the smooth pebble
crab Philyra laevis (Bell 1855) at two sites in South Australia during
1990-91 - - - - - - - - - - -
Olsen, A. M. The history of the development of the Pacific oyster, Crassostrea gigas
(Thunberg) industry in South Australia - - - - -
Bird, A. F. & Yeates, G. W. Studies on Aprutides guidettii (Nematoda: Seinuridae) isolated
from soil at Northfield, South Australia - - - - -
Brief Communication:
Tyler, M. J., Davies, M., Watson, G. F. The hylid frog J Litoria se seas Lis saint
in the Northern Territory - - : 2
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TRANSACTIONS OF THE
ROYAL SOCIETY
OF SOUTH AUSTRALIA
INCORPORATED
VOL. 18, PART 3
MORPHOLOGY AND REPRODUCTIVE BIOLOGY OF
LIMNODYNASTES SALMINI, L. CONVEXIUSCULUS AND
MEGISTOLOTIS LIGNARIUS
(ANURA: LEPTODACTYLIDAE: LIMNODYNASTINAE)
By MARGARET DAVIES* & GRAEME F. WATSON?
Summary
Davies, M. & Watson, G. F. (1994) Morphology and reproductive biology of
Limnodynastes salmini, L. convexiusculus and Megistolotis lignarius (Anura:
Leptodactylidae: Limnodynastinae). Trans. R. Soc. S. Aust. 118(3), 149-169, 30
November, 1994.
External morphology, osteology, structure of the larval chondrocranium, development
and call characteristics of the limnodynastine frogs Limnodynastes salmini and L.
convexiusculus are described and compared with similar data derived from
Megistolotis lignarius. These data are presented to contribute to our understanding of
the biology of the anuran fauna as well as to a wider analysis of the genera
Limnodynastes and Megistolotis being undertaken to examine the hypothesis that M.
lignarius is the sister taxon to L. convexiusculus and that L. salmini is the closest
relative to this species pair.
Key Words: Limnodynastes convexiusculus, Limnodynastes salmini, Megistolotis
lignarius, frogs, calls, larvae, life history, osteology, morphology, larval
chondrocrania.
Transnertons ef the Beyal Sectety wh So Aust, (994), TIBCS), 149-169,
MORPHOLOGY AND REPRODUCTIVE BIOLOGY OF LIMNODYNASTES SALMINI,
L. CONVEXIUSCULUS AND MEGISTOLOTIS LIGNARIUS
(ANURA: LEPTODACTYLIDAE: LIMNODYNASTINAE)
by MARGARET Davits* & GRAEME F. WaTSONT
Summary
Davirs. M, & Warsos, GT, (1994) Morphology and reproductive bielogy of Livmrvdynastes safaring 2
conve iusculus and Megistolotis fienarius (Anura, Leploductyliduc, Limnodyoastinae), Trans, R, Soc 8) das,
HIKER). 149 lo¥, 80 November, 199dq_
External morphology, osteolowy, structure of the larval chondrocraniuim, development yn call chanetetpsuies
ofthe Timmnodvnastine Fors Lorirroedynrastes selmi anh, convertusculas ave described and compared with simitar
duu derived from Megistoleris lignaruis. These data are presented to contribute ty our understinding of the biology
ofthe anuran faunas wellas toa wider analysis ofthe genera Limnacedynayies and Megisdous bemy undertaken
lo cxumine the hypothesis that Ad, lienartay is the sister taxon lo £. convedtuseulis unet that 2. selena is the
closest relative (o (his species pair,
REY WORDS; Lamodynasies convesiusculus, Lamnedynastes salmuini, Mesistolotis lignarins, (rows. calls, harvae,
lite history, Ostealbgy, morphology, larval Chondrocrania
Introduction
Limnodyrastes Fitainger comprises 13 species (Tyler
1992), generally accepted to fall into three species
groups — the L. kesmeniensis group, the L. dorsalis
group und the ZL. ernarus group (Tyler er al 1979;
Roberts & Maxson 1986; Muhony & Robinson 19k6)
Phe composition of these three groups: and. the
complement of the penus has been challenged by
Hutchinson & Maxson (1987) who proposed, on the
basis of data derived from comparisons of albumin
using the immunological techniqhe of micro-
complement fixation, that Lomnvdyniastes is
paraphyletic and thal Megixteloris Menarius is. the
closest relative of L. comvectusculus. Limnodyviuistes
yeni 1s proposed as the closest relative of these Iw
species, all three falling in the L. fasmeniensis species
group.
This comention has nor been widely avcepted
(Cogger 1992: Tyler 1992. Littlejohn er al, 1993;
Roberts & Watson 1993) and the question will not be
resolved until dat are provided for a broader dinalysis
of the genus, Such data should include morpholoyy,
ostedlovy and biological data such as calls und
developmental information. Although the reeognition
of (hese three Laxu is not in dispute, availability of these
data varies: between them.
* Department of Zoology, University of Adelaide, South
Australia S005,
; Department pt Zoolowy,
Parkville. Victoria aQS2-
University of Melbourne.
Parker (1940) provided morphological descriptions
of most species of Limmedviastes and included sume
information on the osteology of (he carpalia as well
us the nature of the hyoid. In many cases he bad limited
mulerit available to him. Moore (1961) provided sorne
further morphological data for selected species together
with distributional dita whilst Lynch (1971) diagnosed
the genus on the basis of ostedlogieal characters derived
from four species, not the taxa under consideration
here. Megisnioris lignariny was not available, of
course. lo any of these warkers. That genus and species
were Uescribed very comprehensively by Tyler et al.
(1979), However, the deserption did not include a
formal description of the skull or of post-cranial
elements. although the skull was illustrited. None of
these wuthors examined the chundrocrania of larval
forms of the (xa,
As part ofa wider study examining the phylogenetic
relanonships of Larneedveustey and Megistolotis and
of onyoing Studies of the binlogy of Australopapuart
frogs, we provide data on the morphology (including
data from the chondractaniuin) aad biplagy of L.
converiuscnluy, £, salming and M- lignarius und
consider these data in the light of published literature.
Although Parker (1940) provided relatively
comprehensive desenpuians or the external morphology
of L. sa/miné and L. converuiseculas, his sample size
was necessarily stall, so we provide: further
descriptions including measurement) in a standard
format to allow tor direct comparison between taxa.
Bor the samme reason, we include a further jlustratior
of the skull of ML Iignarias uy a composite figure ol
the skulls of all taxa,
150 M. DAVIES & G. F. WATSON
Materials and Methods
Material cited here is lodged in the collection of the
Dept of Zoology University of Adelaide (UAZ), South
Australian Museum, Adelaide (SAM), Queensland
Museum, Brisbane (QM), Australian Museum, Sydney
(AM), and Queensland National Parks and Wildlife
Service (Dept of Environment and Heritage),
Townsville (QNPWS).
Measurements of adult specimens were recorded to
the nearest 0.05 mm by means of dial calipers. The
following data were obtained according the method of
Tyler et al. (1979): snout-vent length (S-V), head length
(HL), head width (HW), eye to naris distance (E-N),
internarial span (IN), eye diameter (E), tympanum
diameter (T).
Initially embryos were reared under fluctuating field
conditions; on return to the laboratory, they were reared
at 30°+1°C in glass aquaria (25 x 25 * 8 cm)
containing aerated dechlorinated tap water. Larvae
were fed boiled, organically-grown mignonette-lettuce
leaves supplemented with Sera Bioflakes pond fish
food. Samples of embryos and larvae were preserved
in Tyler’s (1962) fixative. Measurements of
developmental stages were made with dial calipers
reading to 0.05 mm or a stereoscopic microscope and
ocular micrometer.
The developmental staging system used is that of
Gosner (1960). Descriptions of chondrocrania follow
the nomenclature of Sokol (1981).
Osteological preparations were made after the
methods of Davis & Gore (1947) (single stained) and
Dingerkus & Uhler (1977) (double stained).
Illustrations were made using a Wild M8 stereo-
dissecting microscope and camera lucida.
Calls were recorded in the field using either a Uher
4000 (L. salmini) or a Tandberg S11 (L. convexiusculus
and M. lignarius) portable tape recorder and Beyer
M-69 dynamic microphone, at a tape speed of 19 cm/s.
The effective temperature of each recorded frog (either
water temperature or wet-bulb air temperature
depending on where the male was calling) was
measured at the calling site. All recordings were
analysed on a DSP 5500 digital Sona-Graph (Kay
Elemetrics Corp.) using the in-built set-up #10, with
playback on a Revox B 7711 reel to reel tape recorder,
For each call, two attributes were determined: (i)
duration as the interval from the beginning to the end
of the note; (ii) dominant frequency (Hz) as the
maximum value of the spectrum of power between the
cursors for the whole note. Levels of resolution were
less than | ms for temporal aspects, and less than
40 Hz for dominant frequency. Three calls of each
individual were analysed. Calls of M. lignarius are
those of the holotype and paratype described by Tyler
et al. (1979) and reanalysed here using more modern
equipment to facilitate comparisons between the three
species.
Limnodynastes salmini Steindachner, 1867
FIGS 1-14
Limnodynastes salmini Steindachner, F. (1867) Amphibien
in “Reise der ostereichischen Frigate Novara yon die Erde
in den Jahren 1857, 1858, 1859". Zoologie 1(4), 1-70. (State
Printer, Vienna).
Type
NHMW 14849 (two syntypes) from Cape York Qld
(as Australia) (Cogger et a/. 1983).
7
Fig. 1. A. Limnodynastes salmini, B. L. convexiusculus and
C. Mevistolotis lignarius in life.
MORPHOLOGY AND REPRODUCTIVE BIOLOGY 51
Definition
A large speeies (iialés 43-76 inm, lemales 61 im)
characterised by yellowish dorsolateral skin folds. a
relatively distinet: tympanum, glandular nuptial
exerescences inthe male, poorly flanged fingers in the
Jemale, first finger longer than the second. male cull
a resonant “unk’, deposition of eggs in a foam nest.
Description (based on SAM R 41969)
Head depressed: slightly longer than broad (HL/HW
1.07), slightly more than V4 of snout-vent length (HL/S-
V 0.35), Snout prominent, ovoid when viewed from
above and rounded in profile (Pig. 1), Nares dorsal in
position, thew distance from end of snout less than that
from eye. Eye to naris distance less than internarial
span (E-N/TN 0.92). Canthus rostralis well defined and
straight. Loreal region straight, sloping ventrolatera lly.
Eye moderate in diameter, Tympanum distinet, 0.6 of
eye diameter (Fig. 1), Vomerine teeth m Jony horizontal
rows, meeting in midline, posterior ts small lateral
choanac, but not contacting thera, Tongue broad, Vocal
sac slits lateral to tongue.
Fingers cylindrical. unfringed, basally webbed (Pigs
2. 3); subarticular and palmar tubercles well
developed. Supernumerary tubercles at junction of first
and second fingers and second and third fingers, Inner
palmar tubercles Jarge. outer divided. Nuptial
everescences not detectable. Fingers in order of length
43>] >2 =4.
Hind legs moderately short (TL/S-V 0.40), ‘Toes
long, cylindrical with narrow lateral fringes and basal
webbing, (except between toes 4 and 5) (Pig, 3),
Subarticular tubercles large, conical. Large prominent
inner metatuursal tubercle; rounded thickening of skin,
. \ M ) ss ‘a aN f \ H
‘. ae f i /
aan ( Tt
~ A th :
ee | rd i ?
4 -
Fig. 2. Palmar view of rhe hand af A_ ferale Linmodynastes convertusculas (SAM R41983) BL female /
2
but no outer metatarsal tubercle. Toes in order of length
4>3>5>2>1.
Dorsum with well-developed, elevated, pigmented
glands. Supralabial glands elongate and prominent
(Fig. 1), Skin not covered by spines. Ventral skin
smooth, pigmented, Canthal stripe through eye and
tympanum to axilla. Two pale (cream!) stipes laterally
Jrom scapula to groin. Well-defined black spots on
dorsum along each side of stripes and medial to them.
Groin and back of thighs dark chocolate with white
spots,
kariation
The second finger and the thumb on their medial
sides are ftinged im females (Fig. 2) and nuptial
excrescences are glandular in males. The species varies
little in its external morphology. The tympanum is
more indistinct in some specimens. Hind legs are
uniformly moderately short (TL/S-V mean = 0.405,
range 0,38-0.42). The head is as long or longer than
itis wide (HL/AW, mean = 1.09. range [.00-1.18),
Head length is about 4 snout-vent length (HL/S-V,
mean = 0.36, range 0.31-0.39). In all but one case, eye
to naris distance was less than internarial span (E-
N/IN, mean = 0.87, range 0.70-1.07).
Material examined
Qld: QNPWS NI17095, Bundaberg, nr Mon Repos,
N282090, N28228, N2825]. Lake Nugea Nugega, A851,
Gladstone, A6, Southwood N.P. (nr Moonie). SAM
R41969-70, UAZ BI730. Al729. BI726 or Ban Ban
Springs.
Osfeology (based on AUZ Al6l) (Fig. 4).
Skull moderately well ossitied, Sphenethmoid poorly
ossified, not in bony conlact with nasals, extending
iV
vali (SAM
R469) and Clone Megistolativ lenarins (SAM RAI986), Scule bar = 3 nim
152 M. DAVIES & G, FE. WATSON
Fig. 3. Palmar view of hand and plantar view of foot of A. Limnodynastes convexiusculus (SAM R41983) and B. L. salmini
(SAM R41969). Scale bars = 5 mm.
MORPHOLOGY AND REPRODUCTIVE BIOLOGY 153
Fig. 4. A. Dorsal and B. ventral views of the skull of Lirnnadynastes convexiusculus (UAZ A 233); C. Dorsal and D. ventral
view of the skull of L. se/ming (UAZ Al6l); E. dorsal and F, ventral yiews of the skull of Megisrolotis lignarius (UAZ.
Al742). Seale bar = 5 mm.
184
’ length of orbit in ventral view, Prootic and
exoccipital nut fused, Exoccipital not ossified dorso-
or ventro-medially, Crista parotica short and stocky,
not articulating laterally with short expanded otic ramus
of squamosal. Frontoparietal fontanelle poorly
exposed, Frontopurietals well ossified; antertor
extremities extending 44 length of orbil. Orbital edges
of frontoparictals straight, not angled posterolaterally.
Anterior margin of frontopartetal fontanelle formed by
sphenetlimoid ata level about 6 anteriorly along length
of orbit. Posterior margin undefined because ot
juxtaposion of frontoparietals medially and Jack of
medial ossilication of exoecipitals. Nasals poorly
ossified, maxillary process elongate and moderately
broad, in contact wilh well-developed preorbital
process of pars facilis of maxilla. Palatines broad.
expanded laterally, running under dentigerous
processes of vomer to overlie edges of ventral
sphencthmoid medially, Parasphenoid robust. Cultri-
form process elongate, broad almost reaching medial
extremities of pulatines. Alae broad. moderately long.
expanded laterally, at right angles to cultriform process.
Pterygoid robust. Anterior ramus in short contact
with poorly developed pterygoid process of palatal shelf
of maxilla, Medial ramus slender overlying alae of
parasphenoid. Posterior ramus very robust and
clongate, Junction of three rami very robust.
Quadratojugal robust and entire. Squamosal robust
with moderately long zygomatic ramus and short
expanded olic ramus, Maxilla and premaxillit dentate.
wT
ee,
Fig. 5, Dorsal view of the carpus of A, Limnedynastes convevtusculus, BLL. sadtmini and C. Megtstoloris lignarius, Scale
bars = 5 mm
M. DAVIES & G. Ff. WATSON
Pars facialis of maxilla deep with well-developed
preorbital process.
Alary processes of premaxilla browd, slightly
bifurcated dorsally, directed posteriorly, Pterygoid
processes of palatal shelf poorly developed. Vomers
reduced medially with extremely elongate horizontal
dentigerous processes. Columella bony, sigmoid in
shape.
Pectoral girdle arciferal and robust. Slender
omosternum, broad xiphisternum. Sternum
cartilaginous, Clavicles slender, closely apphed
medially, Coracoids very robust, widely separated
medially. Bicupitate scapula robust, Suprascapula about
' assified. Anteroproximal crest of humerus
moderately well developed.
Carpus of five elements (Fig. 5). Lateral process on
medial surface of first metacarpal. Sesumoids absent
at junctions of metacarpals and/or phalanges,
Seven prococlous non-imbricate presacral yericbrac.
Vertebrae | and I fused, Relative width of transverse
processes:
{i>IV>U = SD>V>Vi>VU> VIL
Sacral diapophyses poorly expanded. IHia extending
very slightly anteriorly to sacral diapophyses. Urostyle
crest approx. %4 length of urostyle.
No ihal crest. Dorsal prominence prominent (Fig.
6). Dorsal protuberance ovoid and lateral, Pubis
calcified) Three tarsal elements in foot. Prehallux larpe.
hastate, cartilaginous dorsally with bony base.
MORPHOLOGY AND REPRODUCTIVE BIOLOGY 5
ay
£ } 4
7 PP \
a A \
Fig. 6. Lateral views of the ilia of A. Liminodynastes salmini, Fig. 7. Ventral views of the hyoid in A. Linmodynastes salmint,
B. L. convexiusculus and C. Megistolotis lignarius. Scale B. L. oh asia iid and C. Meg/stolotis lignarius. Scale
bars = 5 mm. bars = 5 mm.
oh ie
FREQUENCY aye TIME (ms)
FREQUENCY (kHz)
25 50 75 100
TIME (ms)
Fig. 8. Power spectrum, waveform and sonagram (sampling frequency 59 Hz) of the advertisement call of Limnodynastes
salmini. Note that the ordinate for the waveform display is not labelled because it depicts a relative linear scale in volts.
156 M. DAVIES & G. F WATSON
Hyoid plate wider than long; posterior processes
slender, asymmetrical in length, Anterior processes
short, blunt perforated. Short, broadly based
anteromedial processes on anterior hyale. Posterior
cornua ossified (Fig, 7).
Variation
Variation in ossification is restricted to the
sphenethmoid, nasals and crista parotica, Maximum
ossification of the sphenethmoid occurs when the bone
extends between the nasals to a point alinost midway
between them. The condition in Fig. 4 is the minimal
condition. However, such ossification does not appear
to be size related.
The posteromedial processes of the nasals can show
greater development than illustrated while the
maxillary process is slightly separated from the
preorbital process of the pars factalis of the maxilla
in the very large female UAZ A1729_
In two of the specimens examined, the exoccipital
and prootic are confluent dorsally, but not ventrally.
but again such ossification does not appear to be size
related.
The other features of the cranium do not appear to
be yariable.
Material examined
UAZ A1729, female, Ban Ban Springs, BI730, male,
same locality, B1726, male, same locality, Al6l, sex
not known, Brisbane, B1736, Stage 33 larvae, B 1737,
Stage 35 larvae, QNPWS 17095, ?male, Mon Repos
nr Bundaberg,
Habitat
Limnodynastes salmini calls from flooded tall
grassland, culverts and swamps. Frogs are found
beneath dense vegetation and are difficult to locate.
Cogger (1992) records that the species burrows,
Structure of the metatarsal tubercle indicates this to
be a possibility, but we have no direct evidence of this
habit.
Advertisement call
The advertisement call of L, salmini is a well-tuned,
single note (mean duration 67.4 ms, Table 1) repeated
regularly in long calling sequences. Call repetition rate
) ~~ wer : a
ce sate A 6 \
_._ 9) Xu \ “4%
Fig, 9. Distribution of Linmnedynastes salmini (open squares) and L. convexiusculus closed circles. Symbols may represent
more than one close locality.
MORPHOLOGY AND REPRODUCTIVE BIOLOGY al
Tasee |. Durations (ns) af tue cally of Limnodynastes
salmini, L., convexiusculus ame Megistolotis lizmarius. Mecrs
and ranges (showin in parentheses) based en analysis of three
cally fram each individual listed
Species No. recorded Call duration
ty 674
Linmitynastes salmini 3 (50-83.2)
Limnedynastes convexiusculis 6 Lag
" pes wre : (67.6-85.6)
» 26.6
Mevistoloris lignartus 2 (21.5-32.4)
for the three jndividuals reported here was
approximately 0.91 calls/s (at an effective temperature
of 25.2°C). Waveform and spectographie displiys of
4 single note and a power speetrum are shown in Fig. 8.
Distribution
Limnodynastes salminé fas an eastern Australian
distribution, the most northerly record being St
Lawrence (Moore 1961). The most southerly record
is near West Wyalong, N SW. (Fig. 9). These records
cast doubt upon (he interpretation of Cogger er al.
(1983) that the type locality is on Cape York Peninsula,
Natural history
The species is predatory on other species of trogs.
One was collected with aL, entanes head first down
its throat in the process of being swallowed,
Life history
Eggs are laid in a large, foamy egg mass. Two such
spawn clumps contained approximately 1630 and 2430
eggs respectively. These clumps were collected soun
after deposition at 120 on 211.1991 near Ban Ban
Springs, SE Qld. A sample of ten eggs bad a mean
capsular diameter of (55 mm (range |.44-1,68 mm)
und aw mean ege diameter of 1,27 mm (range
118-134 mm). There were two jelly layers surrounding
pigmented eggs which hatched at stage 19 about 24 h
after collection (Fig. 10)-
Just prior to hatching (still al stage 19), the
stomodaeum was a pit anteriorly and a pair of well
developed pigmented udhesive glands projected
ventrally. Two pairs of external gills were apparent,
the first with two und the second with six short, broad
filaments,
At hatching, a depression in the presumptive cye
region was appurent.
At O8S0 on 24.1199], embryos were at stage 2) but
the external gills had disappeared, The cornea was
transparent, The mouth had perforated and
keratinization had commenced upon the upper beak.
The adhesive organs remained extremely prominent.
elevated und slightly pigmented (Fig, 10), The nares
were perforated.as was (he unus as a median aperture,
Twenty-four hours later on 25,1, 1991 embryos were
al stage 24, The operculuny was not fully formed. The
horny beak was keratinized and some of the tooth rows
were partially detectable, All of (hese had keratinized
hy stage 26,
Stage 27 was reached by 271.1991, six days after
hatching. The spiracle Was fully formed. The beak was
keratinized as were two upper and three lower labial
tooth rows. The labial papillae were fully formed at
this stage, Larvae were at stage 29 on 317.1991. Stage
35 was reached on 2.111991.
A larva at stage 34 is illustrated in Fig. I, The body
is ovoid and widest at the level oF the eyes, The snout
is evenly rounded in dorsal view and slightly truncated
A
SS, in
( Vo fae iat
UO se he Sate
Sa 5 = —
a? \ .
Fig. 10. A, Embryo of Linneevnestes seliqind at stage 20,
Boat stage 2T and C. at stage (9. Seale bars = | mm.
B
Pia JL A, Lateral and B, dorsal views of a larva of Limnodynastey satmin( al stage 34. Seale bar = Womm,
ISS M,.
in dateral view with an anteriorly projecting oral dise.
The nares are dorsolateral and not raised. Moderately
lurge cyes are dorsolateral. The spiracle is sinistral and
attiehed to the bady wall along its medial edge. It is
Short, opening slightly dorsally and visible when
viewed from above. Its diameter is constant along its
length.
The anal tube is broad and opens medially. The tail
fins are gently arched and rounded terminally. The
dorsal fin commences posteriorly to the body und is
deepest about halfway along its length. The anal tube
is broad und opens medially. The ventral fin
commences postenorly to the body and is deepest
about half way along its length. Tail musculature is
thick tapering to a point posteriorly. Neuromast cells
of the lateral line are well differentiated (Fig. 12) and
extend from behind the eye, along the canthus rostralis
and over the end of the snout (one sequence) and along
the loreal region and under the eye (second sequence),
The mouth is anteroyentral. The oral dise comprises
a horny beak of maderate proportion und there are four
DAVIES & G. TI) WATSON
to five upper and three lower rows of labial teeth (Fig.
13). The most anterior upper row and the two most
posterior lower rows are undivided. Labial papillae
are interrupted anteromedially but extend laterally and
posteriorly around the oral dise,
The dorsal surface of the body
musculature und fins are heavily
melanophores.
Larvae reached stage 36 by 7.11199] and stage 39
by 10.11.1991, Stage 41 was reached by 12,11-199] and
the species had fully metamorphosed by 5.111. 1991, 43
days after spawning,
Measurements of developmental stages are given in
Table 2,
Chondroacranium (Stage 34 larva
B1736) (fig. 14)
Neurocranium approximately T-shaped box
compnsing anterior braincase and posterolateral
spherical otic capsules. Large ovoid frontoparictal
fontanelle exposes braincase dorsally bounded by
orbital cartilages laterally and by tectum synoticum
and the tail
suffused with
based on UAL
> aa
_
: ie Se
Fig. 12. A. Lateral and Bo dorsal views of a larva ef Limo verisies sulin? at stage 34 showing location af the Heuramast
cells of the lateral fine system. Seale bar — 5 min
ee meee
jul
lin Nib
=
¥ Ne, ‘ey evra
Sf “ony, (ul
Meng i “nmin uel!”
Y
spp Vl
i # yl oe at .
ti iki,
a
——- ; ae
uy OT HNN ivan
SAA Agee
AG
rt)
‘ae
pes,
Pe
3
—
iy wil ni MA gu
Witaye, withow lita tH igang
vy tet STA \
“ \\ ip {i cacao
fg alicuet
et
Th Nem rae
Pig. 0 A, Oral dise of larva of Lintodyaastes vonverisemus at stage 36 aed Boot £ selminé at stage 34. Scale bur = fmm.
MORPHOLOGY AND REPRODUCTIVE BIOLOGY
posteriorly. Narrow ethmoid plate forming anterior
margin of frontopanetal fontanelle, Ethmoid plate
projecting anteriorly from) neurocranium as two
anterolaterally diverging moderately broad finger-like
cornua trabeculae. Floor of neurocranium comprising
basis cranii and planum basale. Basis cranii perforated
by paired foramina caroticum primarum. Posteriorly
planum becoming confluent with otoccipital arch and
perforated by notochord.
Massive approximately L-shaped palatoquadrate.
Processus descendens (attachment to neurocranium)
widening laterally to form arcus subocularis separated
from basis cranit by oval subocular fenestra. Medial
processus ascendens overlain dorsally to about half its
width by anterior extremities of otic capsules. Arcus
subocularis curving laterally while sloping ventrally,
Anterolateral edges bearing low crest, becoming
progressively higher and confluent with broad
Tanie 2. Merve data on developing Linnodynustes salmim. ip = awmber of individuals, + =
Age, days und
19
processus muscularis quadrati. Arcus subocularis
attached anteriorly to neurocranium by commussura
quadratocranialis anterior, Pinger-like projections
extending ventromedially into subocular foramen.
Processus muscularis quadrati inclined medially,
bound to laminorbitonasalis by non-chondrified
ligamentum (cartilago) tectum forming tunnel with
commissura quadratocranialis anterior through which
pass M. levator mandibulae posterior and M. 1, m,
anterior.
Processus articularis. quadrati: of palatoquadrate
extending anteriorly from processus muscularis
quadrati and articulating yentrolaterally with slightly
curved Meckel’s cartilage.
Ligamentum quadrati ethmoidale attaching laterally
about half-way along cornua trabeculae, Dorsomediil
edges of Meckel’s cartilages attaching with small
shallow infrarostral cartilages which form shallow
PCA,
Body length x,
range in
Tot length &,
range in
dates Stage parentheses parentheses ai
4, 24 3,35 8,59 iL
25.i.199] (3,2-3.44) (8-8.72)
7, 35 4.64 92 !
28.1199]
6-8, 26 5.24 13.42 23
T9499} (4.6-6.1) (1L.6-16.4)
6-7 27 5.67 13.01 in
27-28.1.1991 (4.96-7.04) (12.16-16.96)
15-30 2k 717 18,00 10
5-20.11, 199] (6.2-8,16) (12.3-19.14)
41-46 29 793 20.41 \l
SLi -5.1.199] (6.5-8.5) (18.0-22,2)
44-56 3) 8.16 21.46 5
3-15.i1.1991 (7.3-8,2) (19.1-23.3}
43-56 a] 8.45 22,43 IS
2-15.11.199] (71-109) (20-27.5)
43-61 32 B.KY 24.16 10
2-20,11,199] (8.0-10,7) (21,3-28.7)
43-59 Aa 9.54 25.05 x
2-18.11, 1991 (8.6-10.8) (23.3-27.6)
48-61 a4 10.8 29,24 8
7-20.41, 199] (4.2-14.2) (23.9-40.3)
43-61 AS 10.76 28.66 R
2-18.11, 199] (9412.1) (25,7-32.4)
db-6| 36 9 32.2 hy
720,11, 1991 (11.0-13,6) (28.0-41.0)
56-61 37 13.5% 35.48 12
15-20.41.1991 (1.9-15.6) (30.8-40.0)
54 3k 4 45.6 |
12.41, 199]
53-61 34 16.2 42,92 x
12-20,11,199] (147-174) (39.9-4.5)
55-61 40 18.03 48.43 4
14-2011 199] (16.3-19.9) (45,0-S1.5)
53-61 4l IS.84 43.54 9
12-20,11.199] (13.9-18,5) (39.4-47.4)
45 16,52 25
(13, 119.9)
46 20.6 |
1a) M, DAVIES & G. F WATSON
MORPHOLOGY AND REPRODUCTIVE BIOLOGS \6}
inverted U in ventral sew with broad non-chondrified
symphysis. elements joming fo form lower jew of
tiilpole.
Suiprarostral cartilages forming upper juw of tadpale
proyevling antenorly ventral to corqua trabeculse and
io both Meckel’s cartilages. and intrarostritls dorsally,
Central corpus of suprarosta lying anteriorly 10
extremmiues of cornua trabeculae with ron-chondrified
medial symphysis, Alae extend to level ol Meckel's
cartilages haterally.
Himnodynasles convexiusculus
(Macleay. 1877)
FIGS 17, 9, 1345
Karasrer eonveamascalias Macleay, WRT TE Prove, Lowy Soe
N APP 2 135-138 (1361
Limmodynesies Glivacens Der Vin (84h) Prov
NSE 9, 65°68 (66).
Phaneritas aevacguineue van Kurypea, PN. 909% Nove
Cinined 9, Ado (Abs.
I\pe
Holotype, presumed lost, frome Mawatta, Binuturi
River wear Darou. Papua New Guinea (as Katow)
(Copgrer er al, 1953),
Befiniton
A moderately large species (males 44-54 mm $-V,
females 44-64 mm S-V) characterised by a prominent
sacral hump, a mid-vertebral stripe. at ragose dersum
with Jongitadinal skin folds, an indistinct tympanum,
nuplial excrescences in the formol line spines ta the
male, females with well-developed flanges on first and
secund Fingers, first finger shorier or equal to second.
ii fength, male call a resonant “unk” deposition of eggs.
inv loam nest,
Desrriplien (bused on SAM R4AI983}
Head high, slightly longer than broad (HL/HW 114),
fess than hall snout to vent length (EL/S-V 0.41. Snout
not prominent, rounded when viewed from above and
im profile (hig. 1). Nares dorsal in position, their
distance from end oF snout less than that [rom eye.
Internanal span approximately equal lo eye to nuris
distance (R-N/IN 0.96). Cunthus rostralis poorly
defined and straight, Eye moderate, diameter greater
han eye to naris distance. Pupil shape vertical.
Tympanum relatively small, not clearly delined, about
(8 cye diameter (Fig. U1). Vomerine teeth i two
elongate sertes, Corved aeross the palate pustevior to
vhounde; not meeting medially: and extending lalerully
to extremities of small chounde-
‘Longue broad, Fingers cylindrical Second finger
slightly fnoaged medial to thurnb (Figs 2. 3), unwebbed.
Suburticular tubercles moderalely developed, rounded.
Inner palmar tubercle very prominent, raised: onrer
divided. Nupuial cacrescences m form of large ovoid
group of line spines on medial surface of thamb with
small group on dorsal surface of base of second finger.
Interdigital webbing absent,
Lin Nae
Fingers in order of length 3221-4. Hind legs
nioderately shom (TL/SV fide). ‘Toes long and
cylindrical with very marrow lateral fringes,
Subarucular tubercles conigal apd proaunent; elongate
ovoid inner ametitursal tuberele: outer metatarsal
tubercle absent, Toes unwebbed other than basal
lringing (Fig, 3), In order of length Fea >5>2>1
Dorum rugose with pale mid-vertebral stnpe. Pelvic
reioo humped (Fig. 1). Chocolare-coloured, well
detined markings of elongate or smull circular shapes
trending as dorsolateral anteroposterior patterns,
Body and limbs wilh numerous very small conical
hlack spines: not prominent oo lateral side of bead and
on evelids; on mandible. posterior poction af body,
Jaleral surface of forearms, dorsal and fateral surhices
of legs and plantar surkice ut loo,
Ventral surface smooth, cream, ihroal speckled wath
pigment, Well-developed supralabial gland (Fig. 1)
kariativn
The hind legs ure moderately short, TL/S-V mieun
0.43, range 0.38-0,47, The head is longer than broad
iL mose specimens (HL'HW mean = lll range
U.98-1.23) and is Icss than hall the snout vent length
(HL/S-V mean = 0.38) range 0.33041). Eye to nuris
distance ty usually less than inlernanal span (E-N/IN
mean = 0,93, range 077-114)
The small-group of fine spines on the dorsul stirfuce
of the base of the segond finger assaciated with the
nuptial exerescence in the desenbed specimen was aot
observed inany other material ihe quptial excreseences
being confined to the medial surface ol the thumb,
Fenmles have well-developed fanges on opposite
surfaces of the thunib and second fingers (Fig. 2),
‘The dorsum varies in tuberculatity and can he more
or tess tubercular than deseribed. A depree of
regularity i back pattem between specimens 1s
evident, The eonical spines un the dorsum are less
developed in some specimens and whe inner palnuat
tuberele js extremely well developed in some male
Frogs.
Marcrial examined
Northern Territory, UAZ BI738-9 Jabiru, SAM
R41979 Baral] Ck oe Jabiru, SAM RAIY73 Darws
(northern suburbs). UA BI773 Radon Ck Rd (av Mt
Brockman}. SAM R41974 17 kin E Jen Roper River
Rd/ Stuart Hwy. Darwin, SAM R41971-2. UAZ BIT28.
Snake Bay Swamp. Melville Istand.
Western Australia! SAM R41Q80-83, Mitchell Platewu
(upstream from Crusher site), SAM R4l984-S, Michell
Plateau campsite. SAM R41978, Kununurra, SAM
R41977 Parry Ck Rd Kununurrit.
Queensland; SAM R41975-6 Edward River
Township, ONPWS A393 Supur Cane Creek, Tully
Mission Beavh Rd, A395-6 Kurrumine. Murdering
Point, N2641-2. N2659 Barmage (Cupe York), N2633
hip. 14. Dorsal und ventral views of the chondroermant of Ay be Mevistolatis dionaries, sue 2 0, 1 Lieieretlvnesoes
wonventiacilay Stan > urd BS PL. \uledni steps 34 Seate bar — ) qin
lo? M
Somerset (rip Cape York), UAZ. AITTA, NISMUR-Y,
N28030 Townsville Town Commen, N33523 Pine
River Bay np Weipa,
Ostevlagy (based on UAZ AloS, BIT2S8) (Fig, 4)
Skull well ossified. Sphewethmoid rinvderately
oseified, in bony coptagh with nasals execyding half
length af orbit in ventral view, Proohe fused with
exoecipilal. Exocerpilal not ossified dorso- or yentro-
nedially. Ossificuuien of oc region reduced ventrally.
Crista purolicu short and stocky, not articulutinu laterally
with eXpahded oe ramus of squamosal, Frontoparictil
fontinelle very poorly exposed. Fronlypartetal well
ossified: anterior extremities extending 74 leagih of orbit
(about half length ossitied portion dorsal sphenetimoid)
Orbital cdges of Trontaparictals straight, nor angled
posterolaterally, Apterior margins. of frontepatietal
lonianelle formed by sphenethmoid uboul halfway along
fenpth of orbi. Posterior nrgin masked by juxtaposilon
Of rnedial margins of frontopurictals. Niasals moderately
Well ossified, crescentic anteriorly, overlying
sphenethmoid wong posterumedial extremities wand with
well-developed. browd maxillary process indeng contact
with well-developed preorhital process of pars etalis
of maxill. Pulatines broad, complete, panning hepeath
dentigerous processes of yomers and overlying ventral
spheneihinoid medially. Parasphenoid robust, Cultnitorm
process urn Shaped, moderately broad extending about
4 length of orbit, Alae broad. expanded slightly
lalerally and angled slightly posterolaterally.
Prerygoid robust. Anterior ramus long. in shart
contact with moderatcly well-developed pierygoid
mrocess of pulutal shelf of maxilla, Meditl ramus
moderately Jong, subactniinate, not overlying aluc oF
parasphenoid, Posterior ramus extremely robwst,
Juncuan of three rami extremely rabust,
(uadratojugal robost and entire. Squamosals
Modertely pabust with moderucly long zygomatic
Tamus aud short, expanded olie ramus, Muxilla anc
premunilla dentate, Pars facilis of maxilla deep with
well-developed prearbital process.
Alary process uf premaaxitta broad. bifureaw
Jursally. direeled postenorly. Posterior process al
premaxilla moderately long, straight aling
anieromedial edges, then curved postepamedially.
Mulital shell well developed, with moderiely well
developed pterygoid process. Yomers reduced medially
willhextremely long dentigerous processes. Columellit
curved and bony.
Pectoral gardle arcilerai and rohust. Slender bony
mosternum with stilked kKaobbed cartilaginous
uilenior extension, Broad Agplosternum, Sternum
cartiliginnus, Clayrcles slender, curved, closely applied
medially Coraceids robust, moderately separaned
medially, Seapula bicapitate. robust, Suprascapula
about 24 ossificd. Antensprexiial crest oF lurmerus
miuderatcly well developed.
DAVIES & G, F
WATSON
Curpus uf five elements, Puirs of scsamoids ut
junctions of metacarpals and proximal phalanges and
other junctions between phalanges: except lerminal ones,
Lateral flange on first metacarpal (Pig. 5).
Seven provuelaus non-imbrieate presacral vertebrae,
Vertcbrac | and I fused, Relative widths of transverse
processes:
UL>IV>SD>U>V>VI=VU=VIL
Sacral diapophyses poorly expanded. Via extend
slightly anteriorly to sacral diapophyses, Urostyle crest
upproxmiuately hall length of urostyle.
Slender aial erest. Dorsal prominence prominent ( Fip.
6). Dorsal protuberance lateral. Three tarsal elements
In foot, Prehallax large, wedge-shaped with bony base
und narrow dorsal curtilaginous edge,
Hyeid plate wider than Jong. Posterior processes
hroad and trunvated. Anterior processes slender,
expanded and truncated distally. Short anteromedial
processes of unlerior hyale. Posterior comma ossified
(Fig. 7).
bariation
Variation exists in the degree of ossification of che
sphenethmoid which can be slightly greater or less than
lustrated. tn one specimen (UAZ BI728). a large male,
The posteromedial process of the masul is more extensive,
overlying the sphenethmoid and almost reaching the
anterior extremes ob the Hroatyparnetals.
Che length of the demuigerons processes of the yomers
may be slightly longer than those illustrated.
Material examined
UAZ Al65 no data. A233 sex unknown Kununurra,
BI728 male Melville tsland. HI738 male Jabiru. BI739
nae Jabiru. BI230-stage 34 larva, BISO4 stage 30 larva,
61773 Radon Creek Rd nr Mt Brovkman, Al774
Townsville Town Conumnen,
Aalbirar
Limnadvnayies Convexcusculus calls trom sinall
hollows, crab holes, among long grass and dense
yerctation in geassland, culverts and swamps (Tyler
al W983: Davies. Martin. Tyler und Watson unpubl,
observuljons).
Advertisement call
The advertisement call of L. cartveatuseculis iy a well-
tuncd, single note (mean duration 74.7 ms, Table |)
repeated regularly in long culling-sequences. Cull
repetition rate tor the six mdividuals reported here was
2.14 (ange |,59-2.57) calls/s (at an effective lermperature
nf 280°C). Waveform and speetragraphic displays of
@ single note and 4 power spectrum are shown in Piz. 15
Distribuient
Livihodynastes convextusculus. has a distribution
across the north oF Australia ia the wet/dry tropics. and
including southern New Guinea (Pig. 9). The species
is confined tothe Kimberley region in Western Australia
and oeeurs as far south as 34km S Delamere
Highway/ Victoria Highway Jen in the Northern Territory
and Eurimbuluh N.P. near Mirfamvale in Queensland.
MORPHOLOGY AND REPRODUCTIVE BIOLOGY 163
Life history
Eges are pigmented and laid in a foamy egy mass.
Tyler et al. (1983) described the tadpoles of L.
convexiusculus as being an intense black with long,
deep tail fins. There are four or five Upper and three
lower rows of labial teeth. The first upper and the first
two lower rows are divided. The oral dise is
surrounded by a single row of labial papillae which
are interrupted anteriorly (Fig. 13), The full life history
of this species has not been described.
Chondrocranium (stage 33 larva, based on UAZ B1531)
(Fig. t4)
Neurocranium approximates T-shaped box
comprising anterior braincase and posterolateral
spherical otic capsules. Large ovoid frontoparietal
fontanelle exposes braincase dorsally, bounded by
orbital cartilages laterally and by tectum synoticum
posteriorly. Narrow ethmoid plate forming anterior
margin of frontoparietal fontanelle, Ethmoid plate
projecting anteriorly from neurocranium as two
anterolaterally diverging, moderately broad, finger-like
FREQUENCY (kHz)
FREQUENCY (kHz)
P=)
25 50
cornua trabeculae. Floor of neurocramium comprising
basis cranu and planum basale, Basis cranii perforated
by paired foramina caroticum primarum, Posteriorly
planum becoming confluent with otoccipital arch and
perforated by notochord,
Massive approximately L-shaped palatoquadrate.
Processus descendens (attachment.1o neuroecranium)
widening laterally to form arcus subocularis, separated
from basis cranil by oval subocular fenestra. Medial
processus ascendens slightly overlain dorsally by
anterior extremities of otic capsules. Arcus subocularis
curving laterally and sloping ventrally. Anterolateral
edges bearing low crest, becoming progressively higher
and confluent with broad processus muscularis
quadrati. Arcus subocularis atlached anteriorly jo
neurocranium by commissura quadratocranialis
anterior, Finger-like projections extending
ventromedially into subocular foramen,
Processus muscularis quadrati inelined medially,
hound to lamina orbitonasalis by nonchondrified
ligamentum (cartilago) tectum, forming tunnel with
i
25 50 75
TIME (ms)
75 100
TIME (ms)
Fig, 15, Power spectrum, waveform ind sonagram (sampling frequency 59 Hz) of the advertisement call of Linnadynastes
convexiusctlus, Note that the ordinate for the wavelorm display 1s not labelled because it depicts it relative linear scale in volts,
14 MM DAVIES & G. [, WATSON
commiussuta quadrate Cranialis anterior through whieh
pass M. levatoy mandibulae posterior and M_ bn.
amerior.
Processus drucularis quadrati of pulaloquadrate
extending aneniorly from processus muscularis quadrati
and articulating ventrolaterally with slightly curved
Meckel’s cartilage.
Ligamentumt quadrat: ethmoidale attaching laterally
injdway along cornua trabeculae. Dorsontedial edges
of Meckel’s cartilages attaching with infrarostyal
curblages. Infrarostral cartilages small, forming shallow
mverted U in ventral view with broad non-chandritied
symphysis, elements joining to form lower jaw ol
tadpole,
Suprurostral cartilages forming upper jaw of tadpole.
projecting anteriorly ventral to cornu Wabeculac and
dorsal to both Meckel’s. cartilage and intrarostrals
Central corpus of suprarostral lyimg ulterior to
extremities of cormua trabeculae giving appearance of
paired Structure with non-chondrified medial symphysis.
Lateral wlue extending to level of Meckel's cartilage
fakerally.
Megistolotis. lignarius
Tyler, Martin & Davies, 1979
FIGS 1-7, 9, 14, 16
Megestlonls livnarms Tylor, Mod Marin, A A. & Davies,
M 11979) Ausa Zoe 27 125-150.
ype
WAM R58299 Lake Argyle-Kunuourra Rd. 6.5 km
N Lake Aneyle Tounst Village, Lake Argyle. Kimberley
Division, W.A-
Definition
A moderule-sized species (males 43-62 mm 5-Y-
females 47-61 nny S-V) characterised by a large and
prominent Qmpanurn (approximately the size of the eye)
(rig. 1), male nuptial exerescenees Consisting of smal!
clusters of large conical Blick spines, lenmales with
moderately developed Mange on the media) side of the
svcond finger, frst finger longer than second,
advertisement call consisting of a soft regularly-repeuted
“ap. eges deposited ina foai nest.
Dexseripiion
We have little to udd to the comprehensive deseriphon
of byler eval (979). However, in five females reared
fron) the original spawn clump we recorded moderately
developed Manges on the medial sarface of the sevund
finger. a character absent from the paratype material
(Fig, 3),
Osivoluyy (based on UAZ AI742) (Fig. 4)
Skull moderately well ossified, in bony eontiwt with
nasels and extending hall length of orbit in ventral views,
Praoric fused with exoceipinil Exovepital net tused
Jorsoy or ventrostaterally, Ossification of prootic reduced
venirolaterally. Crista parolica moderlely long antl
slender, nol wticulating laterally with short. very poorly-
expanded offic ramus of squamosal. Fromtoparictal
fonutoelle moderately to well exposed. extending about
hall length of orbil unlerrorly, Orbital edges at trontor
parictals straight, not angled posterolatetally. Antener
margins of frontoparietal tontanelle formed by spheneth
mow abot halbyay along length of orbit, Posterior
inargin undefined. Nasals slender. poorly ussified,
Maxillary process ucuminute, widely separated front
moderately-developed preorbital process of pars facial ts
of maxilia. Palutines broad, nol expanded farenally, over
tain about halfway along fength by lateral ‘4 ot
denligerous process of vomer, Purasphenoil robust.
Cultrifonm process moderately broad. alae now quite
horizonl, not expanded. laterally.
Plerygoisl robust, notin contact with poorly-developed
pterygoid process of maxilla. Medial ramus long,
moderarely slender, ucuntinate: pasterior ramus
inoderatcly stocky; banes expanded at junction of three
Tani,
Oyadrutojugal robust and cormplele. Squirnosal
moderately robust with short slender zygomatic ramus
and short slightly expanded ote ramus. Maxilla and pre
maxila dentate, teeth extending past plerygoid provess
of palatal shelf, Pars tacialis of maxilla moderately deep
with moderately well-developed preorbiaal process
Alary processes of preniaixillit broud at base, slender
Iuterally. directed posteriorly. Paluline processes Widely
separaled, slightly angled posteromedially. Palatal shell
poorly developed. Vorners entire, widely separated
medially. Dentiverous process elongate, crescentic,
behind choapawe, Coluinella bony, curved.
Pectoral gindle arciteral and robust. Well-developed
omosteraum and xiphisternum. Clavieles poorly
separated medially, Coracoids widely separated
mediully. Saprasedpula 25 vissified. Well-developed
humeral crest,
Phulangeal formula of hand 2.2.3.3. Carpalio appear
tu be fused with Os centrale postuxiale. Very buny pre-
pollex With spine antermedially, Metacarpal T elongate
with medioliteral Qunge (Fig. 59, Pairs of sesamneids
at junctions of metacarpals and proximal phalanges und
proximal phalanges and next phalanges.
Seven procoeelous nean-imbricate presacral vertebric.
Relative widths of transverse processes:
HWI>SD>1V >> V-VI-VURHVII
Urostyle crest extending #4 lengih of element}, [iui
with short slender crest, Prominentdorsal prominence
(Fig. 6), posterolateral protuberance. Phalangeal fortula
of foot 2.2,3.4.3. Two tarsal elements. Tervinial
phaliunges Kyobbed.
Hyoid phile wider than long. Posterior processes
Slender und rounded distally. Anterior processes slender
slightly expanded distally. Ameroncdial processes of
antcnior hyale expanded and broadly truncated distally
(Fig. 7) Posterior curnua short and Ossified.
hurtation
Varullion occurs im the ossificalion of tre
sphenerhmoid and erist puronca in the material
examined Munim) ossification of both these elements
MORPHOLOGY AND REPRODUCTIVE BIOLOGY 165
is as illustrated although slight ossification of the nasal
capsules was noted in UAZ Al741. In this specimen
ulso, the gap between the extremities of the dorsal crista
parotica and the otic ramus of the squamosal was
slightly less than shown.
Material examined
Northern Territory: UAZ A228 Cannon Hill, A588
male, BI236 larvae stage 27, B1237 larvae stage 37
Birndu.
Western Australia: UAZ A248, Al768 sub adult,
Al769, A398-400 males, B428 BI7771-72 females.
laboralory reared trom spawn from type locality.
BI7410, B1742 males no data.
Habitat
We have encountered a number of M. lignerius since
the species was described and all have been true to
the habntat description of Tyler et al. (1979).
Distribution
The species is confined to the escarpment country
of the Kimberley and northern Northern Territory as
detined by Tyler ef al. (1979).
Advertisement call
The advertisement call of M. /ignirius is a relatively
poorly-tuned, single note (mean duration 26.6 ms,
Table 1) repeated regularly in long calling sequences.
Call repetition rate for the two individuals reported here
was approximately 1.56 calls/s. Waveform and
spectrographic displays of a single note and a power
spectrum are shown in Fig. 16.
Life histery
Tyler er al, (1979) described the complete life history
of this species. Tadpoles are Jotic with mouthparts
adapted to fast flowing water and are characteristically
heavily pigmented. They actively select areas of
streams in which water is fast-flowing over riffles and
avoid the calmer pond areas (Davies, Watson and
Williams unpubl. data). There are five upper and three
lower rows of labial teeth. The two most anterior upper
rows and the three lower rows are undivided. Labial
papillae are arranged in a double row around the oral
dise and are interrupted anteromedially.
80
60
dB
40
20
Ny
2 4 6 8 25 50 75
FREQUENCY (kHz) TIME (ms)
6
N
x
=
> 4
oO Spasth
z
i
=) ‘ “
ge Sem
cc :
= i Tipe
ae
Tt. rar of —Ty cl, = 1
25 50 75 100
TIME (ms)
Fig. 16. Power spectrum, wavelorm and sonagram (sampling frequency 59 Hz) of the advertisement call of Megistolotis
lignarius. Nove that the ordinate for the waveform display is not labelled because it depicts a relative linear scale in volts.
lb
Chondrocranium (Stage 27 larva. based on UAZ
Bi236) (Fig. 14)
Neurocranium approximates T-shaped box
comprising anterior braincase and posterolateral
spherical otic capsules. Large ovoid frontoparietal
fontanelle exposes braincase dorsally. bounded laterally
by orbital carulages and posteriorly by tectum
synoticum. Narrow ethmoid plate forming anterior
border of frontoparietal fontanelle. Ethmoid plate
projecting anteriorly from neurocranium as lwo
elongate anterolaterally diverging, finger-like, cornua
trabeculae,
Floor of neurocranium comprises basis cranii and
planum buasale, Basis cranii perforated by paired
foramina caroticum primarum, Planum becomes
confluent posteriorly with otoccipital arch und is
perforated by notochord.
Massive approximately L-shaped palatoquadrate.
Slender processus ascendens (attaching pulatoquadrate
to neurocranium) widening laterally lo form arcus
suboculuris, separated from basis cranii by oval
subocularis fenestra. Medial processus ascendens
M. DAVIES & G. F WATSON
extensively overlain dorsally by anterior extremities of
otic capsules. Arcus subocularis curving laterally while
sloping ventrally. Anterolateral edge bearing low erest.
hecoming progressively higher and confluent with
broad processus muscularis quadrati. Arcus
subocularis ultached anteriorly to neurocranium by
commissura quadratocranialis anterior, No finger-like
projections of element extending ventromedially into
subocular foramen.
Processus muscularis quadraui inclined medially.
bound to lamina orbitonasalis by nonchondrified
ligamentum (cartilago) tectum, lorming tunnel with
commissura quadratocranialis anterior through which
pass M. levator mandibulae posterior and M.t.m.
anterior.
Processus articularis quadrati of palatoquadrate
extends anteriorly from processus muscularis quadrati
and articulates ventrolaterally with slightly curved
Meckel’s cartilage. Anteroluterally. ligamentum
quadratoethmoidale attaches laterally to cornua
trabeculae about one third along its” length.
Pane 3. bariation.in morphalogical, osteclogical, developmental and behavioural characteristics of Limnodynastes salmini,
L., convexiusculus and Megistolotis lignarius. Karyetypre det are from Mahany & Robinson (1986).
Character L sabnini L. convexiusculus M. lignarits
Morphology
Nuptial pad, spiny - fine large
Spines on skin - t +
Glandular dorsum + + -
Tympanuiti mod. distinct indistinct very distinet
Flanged fingers in female present well developed present
Relative finger lengths A> >2=4 A>leS |=4 I>) >2>4
Karyotype (2n) 22 24 24
NOR (chromosome number) 2 1} 2
Osteology
Nasals approx. iriangular “+ + -
Frontoparietal fontanclle exposure poor poor moderute
Double tooth row on vomers = = +
Rel, of pars fucialis with maxillary
process of nasals G + ~
bust jen of pterygoid rami | { -
Rel anterior ramus of pterveoid
and pax illa + + =
Mod. slender cultriform process = — +
Hastare prehallay + - +
Carpalia r + =
Double sesamoids on fingers — + +
Developmental
Highly melanic larvae — + +
Heavily pigmented eggs ai + -
Larval type lentic lentic Jolie
Larval mouth La/dH/12 1,3/3:V12 2 Ald
Cornua trabeculve extending over
mouth cartilages - - +
Behaviour
Calling site swamp swarrip) Rocky strean
a+ + —
Welltuned call
Riffle-seeking: larvae =
= -
MORPHOLOGY AND REPRODUCTIVE BIOLOGY [67
Dersomedtal edge of cach Meckel’s cartilage attaches
with infrarostral cartilages Which form shallow U in
ventral view with conjoined broad, non-chondofied
symphysis. Curtilages lorm lower jaws of tadpoles.
Suprarostral cartilage form upper juw and project
ventrally to level of anterior extremities of cornua
irtbeculue; posteriorly do not reach Tevel of Meckel’s
cartilages dorsally. Central corpus of suprarostra) lies
between extremities of cornuy trabeculae: appears to
be painad, nacrow medial non-chondrified symphysis.
Litera) alae just extending to level of Meckel’s
curtilage laterally,
Discussion
Table 3 consolidates thase chatacters showing
vanitlion between the three taxa. However, this 1s ut
phenelic comparison and ay such does not clarify the
relationships between the taxa, which await a.clacistic
stuily OF the genus fr tote with appropriate outgroups
(Davies in prep.). :
Information on call characteristics generally have not
been included in phylogenetic analyses, although these
data are of particular interest. The udvertisement calls
Of all three species described here appear to be of
similay harmonic structure (Figs 8, 15, 16). Ul is not
clear how calling: frogs produce such frequeney speetra.
ulhough most interpretauions suggest that the vocul
cords generate a carrier frequency which is modified
hy secondary pulsing (amplitude modulation) chat
results ina pattern of side hands of {he dominant
frequency (Watkins 1967: Rywuy 988). Afternatively.
imaes may produce a fundamental frequency with
assucuited harmonics, with one of more of (Aese
harmonic frequencies being emphasised by the
resonance characteristics of the body cavities. In either
case. Wwel-tuned cally produce ua distinctive spectral
structure, ani this pattern is particularly clear in the
vals of L. conbextuseulus and L. yelminé, where the
fourth frequency band cyuntains most energy (Table 4,
Tigs 4, 15), Despite the overall structural similarity of
the calls of these Two Species. there is a clear-cut
difference ot 400-500 Hz in the dominant frequencies
(Table 3), In anurans, such differences in frequency
are commonly inversely related to the body sizes of
calling males (Schneider T988) and. io this case, are
consistent with the overall trend in mule body size in
the species with L, satiat being senerally larger than
L. convexiuscilus.
The mede of production of the advertisement cull
of M. lignarins is equivocal. While shuring basic
spectral sinilarities with the calls of the other species,
the cail rm fess well-tuned and the envelope shape is
markedly different, having a rapid rise-tirne uber a4
likely 10 generale a (ransient wide spread of frequencies
(Rossing 1982) and to obscure the harmonic frequency
specuim (Table 4. Fig. 16). There is also a clear
difference in the call characteristics of the two recorded
individuals (Table 4). Individual #1 (SAM R16229) was
calling ina rock crevice in u stream that, based on
many subsequent Observations of this species in (he
field (M. J. Tyler. M. Davies & G. F. Watson unpubl. ),
1s a Eypical calling position. In contrast, the call of
individual #2. the holorype (WAM R58229), was
recorded while the male was moving in the open, a
situation that has never again been encountered by us,
Because of the behaviour of the holotype at |he time
when ils call was recorded it is more likely that the
call of individual #f represents a typical call of this
taxon.
Major developmental differences between the three
axa tie in the Jotic nature of the Larvae of WW. /igtarins
as Opposed to the lentic larvae of ZL. convexiuveulus
and L. salmini. Lone adaptations. in M_ /igharijis
include a flattened body, strong tail musculature with
relatively narrow tal fis and a relatively large yentral
suctorial mouth disc. This is ofa less elaborate nature
than found in seme Jotie species (Davies 1989),
‘Time to metamorphosis in. M. /ignarius js longer than
nL. Safin’ (not known ip L. cunpexiuseuluy) being
about 65 diys as opposed (o 43. Both inhabit seasonally
arid areas and the difference in developmental rate
probably reflects the relative permanence and depth
of the aquatic habitat in which development occurs,
Tanith 4. Derribadon of frequencies (Az) withitt the calls af Linniodtynasies salami, Lo convextusculus end Mepistalotis
ligemtrius, The requertey wit most energy (the dominant frequency) iy underlined.
Le yighmini
L. convextusenties M lixniriys
Individual? i 2 x) 1 2 3 4 5 6 ] Z
360) 360) 3A) 480) 520) 480 520 480 AR) 460 -
720 720 720 920 ooo 96d LOO 6) Yao YO) SU
IOs) Ho 1120 140) {RO 1400 i4k0 1440 tds) (F20 1460
Vi} 1820 1520 LRRO. 2000 1920 2000 1901) 1Y4ot) TREO 1940
TkOW baa 1920 2320 2440 2400 2480) 2440) 2440 2770 ~
2160 280 2280 Zsth) 2961) Zs4u 300) 2960) 2920 268U =
2520 2640 2640 _ — - _ = — 2900 -
2RRO 3000 3080 — ~ — _ - 4160 -
(66 M, DAVIES & 1
None of the taxa appear to have exceptional adapaoons
WW rapt birval developnijent, again reflecting the less
ephemeral naiune of the waler bodies in which they
Spawn.
The Julie nature ol the Mevistoloris larvae ts alse
retleered in the chondrocranium, The leneth of the
vornud iriheculue an their relationshig with the rostral
Curtilages differs in these love and lentie species. The
cornuat trabeculae ane elongate in locie M. lignarts
and relulively shorter in lente L. sefowee and 2
convertuscutus This observation does pot fit: the
generality of Sokal (981) Jor lotic species of other
farmnbhes m whieh shortening of the cornua Lrubecalae
is correlated with shitts of (he meauth away. from a
subrerminal position in lotic liryae,
Descriptions of chondrocrauia of limnodynastine
taxa ave ujtaVailable for comparison, the only published
dala on Australias species being those from the
myobalruchine species Uperafeia lilamoda (Day ies
RU) und two Speciey of Pyendopheyie Gacahsen
964), Hone of Which js. u lotic spectes.
Ossification of the skull is negalrvely correlated wall
ste. The large f.. xedmind has the poorest assifioution
anid the Sniullest species Lo cenverisealus has the
Steatest, Intuitively one would expect larger speeies
to have stronger, more ossifed skullsthan smaller anes
However. the adult skeleton is (he product of botti
emtogerietic and phylogenetic constraints (Lruch &
Aiberch 1985: Davies 1989b; Hunken 1992) and such
J simple correlation rarely holds.
Lynch (1971) provided a diagnosis fat Linnodynastes
based on examination of the skulls of L. dorsalis. b
fleichert, Lo peroni and Lo tashianiensiy Data from
the fhree lax investigated here du moet confer te this
diagnosis ina number of features. The nasals are very
small in. Megistal/eny rather than beiny relatively small
although L, sadmine and Lo converiaveutus Ci the
diagnosis, The nasaly are notin broad contact with the
maxillae in Megisralons and the frontoparetal
fontunelle is poorly exposed in both L. salmine and Lb,
Convexiusentus as Opposed to the ninderate expusure
defined tor the yrenus. Lynch deseribes (he 2ygonvatic
ramus as beige as long us the ollie ramus in
Limnodasies but the former ts longer than the liter
in both /. sadmin? and L. convesiaveulas. The palatines
ube not narrowly separated medially in Mevisroletivy
and the sphenerimord does not extend amenorly to ihe
centre of the nasuls in Lo yalmint. The wae ot the
parasphenoid are not broadly overlapped laterally by
The piedian eae of the ptery@oids. in any of the three
tinal and (he anterior ramus of the plerygoid does not
conform fo the generic description ino either
Mexiyrtotis ov L, sualmini-
tarker (1940) discussed the relative lengths: of
jnctacarpal Ly) Lined yedsres and reported a trend
Towards feduetion in the phalanges of the first finger,
bk WATSON
Internally, the metacarpal bone is dispropertronalely
Jong. The trend was noticed nL, flenheri To perom
and (6 a lesser extent in L. salminé. Parker suggested
thal il ts significant that these species do not have
nuptial excrescenees, He went on to argue thal species
thut spawn in Water such as Lawiedyenistes, need
nuptial cXereseenees.to uid mm Clasping the Jemuale and
that lack of the sume must be compensated tor ja some
way, 7¢.. by changes in the relative length of the
lingers. The same trend is recorded in ML Iienariiy
hut this species has well-developed spinous nuptial
cacrescences, 80 does fol support fhe assuription.
Further. we record bere (he presence of ghindulal
nupuul exerescenees ind. salmini contrary lo the
observations of Parker (940),
Parker (1940) provided X-rays of the hands of L,
rasmantensis, Lo penmi, Le fletcheri and £. salmine
Wo iustrate the nature of the melacarpals and digits,
The material in this stody for which compurable data
arcavaiable are all (nates and all have a lateral Mange
on the medial surface of the first metacarpal,
Examination of Parkers material shows the presence
wl this structure only in male Le pera but rot int
feniale £ salminiand male 2. flewher?. Whether this
is asexually dimorphic character remains io doubt us
jhe X-rays ure extremely granular and may. nob be or
sufficient quality to detect the siructure,
Linmodynasies salinity’ and £. convexiuseulas share
simi habitus and, exeept for the difference in
dominant drequeney, have very simihic calls. The
distributions of these two species have 4 vone of overkip
between Mirtamvate and $( Lawrence i Queenstiand,
(dentilication by wity of call from these areas must
therefore be regarded as dubious when compiling
distribution dita.
The data recorded here coniribule to our knowledye
ol the morphology and reproductive biology at tHiree
Austrilupapiin anurans and also wall contribute to a
wider analysis of the pbylogentic relationships within
Linmodynasley Fitzinger,
Acknowledgments
This study was supported by un Australian Research
Grants Couneid Grant ro the first author and Michael
J. 'Tyler, We have alse dirawe tipom material collected
tovether with our colledgucs Angas A. Martin and
Miehael J Tyler whose support and company we
sritetully ackaowledee, Leeanne Seller and Michael
J. Tyler assisted with tadpole rearing. Keith MeDonald
provided distributional dala for Queeastand. Michael
fyler critically read the manuscript and we (hank him
for his comments. We thank Angus A. Martin for
allowing the analysis of original recordings of the taxi.
MORPHOLOGY AND REPRODUCTIVE BIOLOGY [69
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A NEW GENUS AND SPECIES OF FROG
(ANURA: LEPTODACTYLIDAE: MYOBATRACHINAE)
FROM SOUTHERN TASMANIA
By D. E. ROUNSEVELL*, D. ZIEGELER*, P. B. BROWN*, MARGARET DAVIEST
& M. J. LITTLEIOHNE
Summary
Rounsevell, D. E., Ziegeler, D., Brown, P. B., Davies, M. & Littlejohn, M. J. (1994)
A new genus and species of frog (Anura: Leptodactylidae: Myobatrachinae) from
southern Tasmania. Trans. R. Soc, S. Aust. 118(3), 171-185, 30 November, 1994.
Bryobatrachus nimbus gen. et. sp. nov, is described from moist cool habitat in
southern Tasmania. The genus is distinguishable by its direct development, fusion of
presacral vertebrae VII and VIII with the sacrum, dentate maxillary arch, absence of
vomerine teeth and the presence of a columella. The species is small (males 19-27;
females 25-30 mm S-V) with distinctive dark patterns on the dorsum and an
advertisement call that is a series of “toks” with a pulse repetition rate of 5.3-6.1
pulse/s, Eggs are laid in cavities in moss in groups of 9-14 and hatch as four-legged,
tailed froglets. B. nimbus gen. et. sp. nov. is cryptozoic in vegetation at poorly
drained sites on wet peat, or in edaphic moss in implicate rainforest and subalpine
moorland at altitudes from near sea level to 1,100 m. The genus and species are
endemic to Tasmania and bring the frog fauna of the island to 11 species.
Key Words: Anura, Bryobatrachus nimbus gen. et. sp. nov., Tasmania, new genus,
new species, morphology, osteology, advertisement call, development, habitat,
cryptozoic, distribution.
Transactions of tie Koval Seviery ef S, Aust, (904), TREAD, t
TASS
A NEW GENUS AND SPECIES OF FROG (ANURA:
LEPTODACTYLIDAE:; MYOBATR ACHINAE) FROM SOUTHERN TASMANIA
by D. E, ROUNSEVELL* D. ZIRGELER* P. B. BRoWN* MARGARET DAVIES!
& M. J. LItTLeEIOHNe
Summary
RouNseve.(, 0. B., Zieorer, D., Brown, PB. Dawes, M, & Lirti tot. Mt (1994) A new genus and
species of frog (Anura: Leptoduetytidae: Myohatruchinae) trom southern Tasmania, Tries, Ro Sir Aiese- 183).
IIARS. A) November. 1994,
Bryobarravhus nimbus gen, el, sp, nov. is described from moist cool habitat in southern Tasmania. The genus
jy distinguishable by its direct development. fusiun of presuceal vertebrae VIP und VIL wih the sacrum. dentate
maxillary are, absence of vonerine teeth and the presence of at columella, The species 1s siiall (mates I-27;
females 25-30 mm S-V) wilt distinctive dark patterns on the dorsum and dn advertisement eal (hat i a series
of “loks” wilh a palse repetition rate of 3.3-6.1 pulses/s, Eggs ore kud i cuvities in moss in groups of 9-14 and
fateh as four-legged, tailed froglets, & nimbus gen. et sp. tev. is Ceyptozvie in vegetation al poorly drained stiles
on wel peat. of in edaphic moss in implicate rainforest and subalpine moorland at altitudes from) near sed level
io 1,100 m, ‘The genus and species are endemic to Tasmania dnd bring the froy fauna of the land to 1 species
Key Worps: Anura. Bryubarrayhiesy nimbus gen, et sp, noy,, Tasmania, new genus, new spevies, morphology.
usteology. advertisement call, development. habital. eryplazie, distribution.
Introduction
The frog fauna of Tasmania is an clement ol’ the
Bassian Province of south-eastern Australia (Littlejohn
& Watson 1985), As well as species with wide ranging
distribuvions throughout eustern Australia (eg,
Ranidella sienifera Girard, R58 and Liminoevnetstey
tasmuniensis Giinther, W5% [Leptodactylidag:
Myobatrachinae and Limnodynastinae}), there are two
endemic species, Cituria burrowsae (Seatt, 1942)
(Hylidae) and Ranidella tasmeniensis (Giinther, 1364)
(Leptodactylidae: Myobatrachinae).
Ten taxa are recognised in Tasmamia und aspects vt
their biology are relatively well Known (Martin &
Littlejohn 1982). With a greater appreciation of the
need to document the biodiversity of the continent,
efforts have been direcied (oward surveys und the
mapping of distribution patterns of local fauny, Durmg
one such project aimed at providing an atlas of the [rogs
of Tasmania, one of us (D.Z.) recalled having nerd
at several high altitude sites, 4 distinctive frog call that
could not be attributed to any deseribed ‘Tasmanian
inxon,
In late November 1992 cally were recorded at the
Hartz Mountain National Park and specimens were
collected (Robertson 1993; Rounsevell & Swain 1993)
* Parks and Wildlife Service, Deparment of Environment
and Land Majayement, GPO Box 444, Hobart, Tas. 70411,
+ Department of Zoology, University of Adelaide. S, Aust.
SCS,
} Department. of Zoolugy. University of Melbourne,
Parkville, Vie, 3052.
' ANDERSEN, M, L. (197%) The cimparauve myulogy and
osteology of the carpus and tifsus of selected antirans, PHD
Dissertation, Dept of Systematics and Beotoyy, University
of Kansas (Uppubl,).
Examination of the collection at the Tasmaman
Museum und Art Gallery. Hobart. revealed further
material,
The frogs could not be rélerred to any known penus.
and soa new genus is erected ty accomodate Them
here, We describe the species and provide information
on its higloey and distribution
Materials and Methors
Material reported bere ts deposited in the Tasmanian
Museum and Art Gallery, Hobart (TMAG), South
Australian Museum, Adclaide (SAM), Museum of
Victoria, Melbourne (NMY)}, the Australian Museum,
Sydney (AM) und the Department of Zoology,
University of Adelaide (UAZ).
Measurements were taken using dial calipers reading
to OOS mor or wilh an eye-picee micrometer
Measurements (mim) were; eye diameter (EB): eye-to-
maris distanee (h-N): internarial span {IN}: snoul-vent
length (S-V) and ribia length (TL), und for selected
specimens, head Jength (HL) and head width (HW).
‘The methods of measurement follow Tyler (1968) an
interpretation of data follows Tyler (1978), Daya are
presented as means with ranges in parentheses, The
lympana of muny specimens are mdistinet ar nai
visible, dnd hence wecurace measurements of head
width and head length were aut always possible,
Osteologicul dati were obtained from specimens
cleured and stained with alizarin Red § for bone aftet
the method of Davis & Gore (1947). and with alizarin
Red S for bone and aleian blue for cartilage after the
method of Dingerkus & Uhler (1977). Ostelogical
Jeseriptions follow Troeh (1979), and Andersen
(1978") for the curpus and tirsus.
pl
tL
Db ROUNSEVELL, D ZIEGLLER, Po
Line drawings were made with the aid of a Wild M8
slereoscopie dyssecling Ticroseape with adtached
camera bueidue
Most field observations were made jn the spring and
summes pl (993) atthe oype lovality at Hiv. Mountains
National Park, and during a field survey in Qether
und November 1993 (Ziegeler [994). Air temperature
and relative hamidity were measured with a “Zeal?
whirling psyctrometer (BS 2842/66),
Recordings of the adverisement calls of une
individual were nade with a Sany WM-D Pro-
Walknian cassette (ape recorder umd at Beyer M88
cantiond dynamic microphone by Ci, Fo Watson atthe
type locality on 5 December 1992 ar about 1330 hours
ES.7. The wet-huib wir temperature was 85°C and
a Ury-bulb dirtemperutire was 9.2°C in the vaginity
of (he calling pale.
Recordings oF advertisement calls of several
Weis tkus alse were made at the type lovulity by
PR Brown on 30 Novernber 1992 using o Marantz
Superscope C-205 cassette recorder wod « Sencheiser
MLS) electrel microphone: temperatures. were net
measured
The calls were repliyed on a Nakanicht Dragon
cassele recorder, with the le output direeted toa Kay
lemetries DSP-S500 digital Sonu-Graph, Calls of
sufligvent intensity fhal did nok overlap those af
ncighhours were analysed. ‘Phe number of pulses in
Leal (determined by inspection), Jind measurements
of cal] duration (inst and pulse repetition rate (as
pulses’s; trom the peak of the first pulse to the peak
ol (he last pulse) were determined from the wave-fonn
display of euch clear call
Por the six clear calls obtained by -G, F Waison,
Joy nant frequencies (H4) were determined trom jhe
power Spectrum for the complete call. with the
NIM ahd gext highest peaks being measured.
Pulse duration, and attuck anil déeeay times of the
middle pulses alsa were estiniited to the nearest
milliseeaind (ms) from the wave-form displays, The
repetition rate of the calls (as calls/min) also was
determined from this sequence.
Rivhwen calls from the recording sequeave obtained
by P: BL Brown were analysed, und durition, number
Of plilses and pulse pate detennined, As calls of several
individuals are yoeluded, the values are treated as 9
group,
Systematios
Rout anuran lamilies ure nitive to Australie Hylidae,
Ranidue, Microhviidae and Leptodactylidae
(Myobatrachidac ol many authors). The mew frog wits
7 Paynes. M, (987) Ta sonomy and Systematies a the aerrus
(inerilece Cray (Anura Leptodactylidue), PHD Thesis,
Departrient of Zadhary. University of Adelaide Clnipobt
BROWN, M DAVIES & M, J. LITTLEJOHN
identified as belonging to the faniily Leptodactyliduc,
sub-family Myobatrachinae, on the basis of (1)
terminal phalanges knobbed; (2) inlercalary elements
absent: (4) apical element of ML intérmenedibutaris
ubsenl, (4) pectural girdle areierml, (5) palarines
discrete: (6) prepharyingeal folds absent; (7) alary
processes OF hyoid plitle moderately broad and wing-
like: (8) cricoid cantilage divided ventrally: (9)
Intervertebral discs not fused to centra in adultss (10)
alachment oF Mo fitermandihylaris. pon M.,
submentalis absent
The firsethree charaerers exclude the species from
the Hylidae, and the fourth exeliides a0 from: the
Ranidae, whilst the first. fourth, fifltrand sixth exclude
iL from the Microhylidac. The remaining characters
are definitive of the Jeptodaetylid sublanily
Myuhatruchinae (Parker 1940; Lynch 97, Tyler 1972,
Davies 987%),
Litdejohn ef a/, (1993) report 9-12 genera as being
yariously recognised. within the Myobatrach lane, The
neW species canfot be placed in any of these gener
on the busts of the following combination of characters:
an pparent autapomorphy of the yertebrul column at
fusion of the presacral vertebrae VIL and VIM with the
sacrum, presence of 4 dente maxillary arch, the luck
of Vomerine teeth, presctice af a columella, und direct
develupmeat of the young with a four-legged. tailed
frogler hatching trom the ege membranes.
the [roy is exeluded specificully from the
myobalrachine genera us tollaws: together wah the
vertebral fusion, (I) trom ywe by the absence ot
parental care hy the male, by the absence of voinerine
teeth ahd by the complete phalangeal formula: (2) from
Crinue (sensu Blake 1973) by the absence of vomerine
teeth and the absenve of a free-swimming tadpole: (3)
trom. Geverina by the absence of vomerine teeth and
of a projecting dorsal ange on the-anterior ranius of
the pterygoid, by the projecting snout und by the
granular ventral surfaces (4) from Rendell by the
width of the bases ol the alary processes of the hyoid.
by the absenee of free-swimming tadpoles and by the
nature OF the Gmosternum: (5) Front daudacrylus by the
absence of T-shaped terminal phalanges on (he dizils,
by the absence of yomerine teeth and by the lack ol
free-swirruning: tadpoles: (6) from Lperolera by tie
absence of hypertrophicd dermal glands and of two
raised Compressed metatarsal tubercles, and by the
ubsence of w (ree-Swimming tadpole, (7) trary
Preadophrvie by the presence of a columella and ot
teeth on the maxillary arch, and the absence of a Ipee-
awiinting tadpole, (8) trom Arenephryne by the
presence of wu columella and of teeth om the maxillary
arch, and by the ubsence of modified phalungeu!
formulae on the hands and feet: (9) from Myabuiriedis
by the presenee of teeth on the maxillary arch, by the
absence of a modified phalangeal formula of the hands
and by the absence of reinforcement of dhe anterior
A NEW GENUS OF PROG FROM SOUTHERN TASMANIA VW
portion of the skull by anterior placement of {he nasals,
(10) from Metacrinia by the presence of teeth on the
maxillary arch. by a large omosternum, by the lack
of extensive reduction of the palatines laterally, by the
less extensive nasals, und by the projecting snout and
shape of the head,
Genus Bryobatrachus gen. nov.
Type species: Bryobarrachus nimbus
Diaynostic definition
|. Alary processes of hyoid plate broad and wing-
like, 2, Cricoid cartilage divided ventrally 3,
Intervertebral dises untused jn adults. 4. Attachment
of M. intermandibularis on M. sutmentalis absent. 5.
Hypertrophicd dermal glands absent. 6, Small Mitlened
inner metatarsal tubercle. 7 Tiny, or no outer
metatarsal tubercle. 8. Snout projecting when viewed
from above, 9 Masillary arch toothed. 10. Vomers
reduced to one or two fragments at edge of chounue.
I. Columella present. 12. Palutines slightly reduced
laterally. 13. Nasalsy not located anteriorly on skull, 14.
Omosternum large, mushroom-shaped with narrow
stalk. 15, Phalangeal formula of hand 2,2,3,4. 16.
Phalangeal formula of foot 2,2,3,4,3, 17. Terminal
phalanges knobbed. 18. Vertebrae VIL and VII fused
with sacrum. 20. Development direet — four-legged
lailed froglet hatching from egg menibranes,
Etymolowy
Derived from the Greek bryan (= moss) und
buirachay (= trog), alluding to the hubil of breeding
in moss or moss-like vegetation.
Bryobatrachus nimbus sp. nov.
FIGS 1-10
Holotype VMAG C1012, an adult male, 304) 14 north
of Lake Esperunce (146°46'E_ 43°13"30"S), Harta
Mounting National Park, 920 m, collected hy D. E-
Rounsevell & D, Ziegeler on 8.x, 1993.
Paratypes; 9 oo, 3 9 @ and 2 sa! TMAG
C1009, ov. same data as holotype, except collected
1-x.1993: TMAG CIOIO, or sume data as hulotype;
TMAG CIOIS, s.a.. Mt Sprent (14595 8'E, 424 7'30"S),
eall. D. Ziegeler, 21.4.1993; TMAG C024, oF,
topetype, coll, P. B. Brown, 2%;xi/,1993, TMAG
C1025. 9, same data C1024 and C1025 in amplexus:
TMAG C350, (2 o ory. Mi La Perouse Base Cumip,
ANZSES Expedition Jan. W984. TMAG C345. sa.
same data as C350; TMAG CR69, ot. Mi Lit Perouse
(46°44 30 "TE, 43°30'20"S). tocks above Pigsty Ponds
on Mt La Perouse track, 1.7 kin NW of summit, coll,
M. N. Hutchinson & S. Hudson, 911.1990; SAM
R43671 (cleared and stuined), or, same locality us
holotype, coll. D. E. Rounsevell, 25.x7.1992,; SAM
R43672, or, sume data: AM R143566 (cleared und
stained), 9, same locality as holotype. coll D. E.
Rounsevell, 25,xi.1992: AM RI43565, co, same
locality as holotype, coll. BB. Brown & D. Ziegeler,
29.41.1992; NMV D67310, oF, sume locality as
holotype. coll, A. & J, EB. Wapstra, 29.51.1992.
Definition
A small species (males 19-27 nm S-V. females
25-30 mm S-V) characterised by a toothed maxillary
arch, presence of a columellit, Lick of vomering teeth,
unmodified phalangeal formulae, knobbed terminal
phalanges, widely exposed fromoparietal fontanelle,
presacral vertebrae VIT dnd VIT fused with the sacrum,
untringed fingers and toes, granular Ventral surface.
dorsum consistently marked wilh dark chevron-shaped
muark between cyes, pair of purallel dark lines Tram
shoulder along anterior portion of hack und pair of dark
patches in coccygeal vegion, eggs large and
unpigmented, four-legged tailed froglet hatches. from
egg membranes, advertisement call a series of “toks”
with w pulse repetition rite of 5.3-6.1 pulses/s.
Description of halompe
Maxillary teeth presenr, vormerine teeth absent.
tongue oval, free behinds tympanuny obscure. Snoul
short, projecting and slightly truneuted when viewed
from above, overshot and sloping posteriorly when
viewed in profile (Fig. WA). Eye lo naris distance les»
Fig. 1. Bryebulrachus nimbuy genet sp. nove A, literal view
ot the heads B. palmar view ol hand; and C_ planta’ view
of foot (Holotype, TMAG IOI2), Seale bar = 5 mim.
174 DET, ROUNSEVELL. 9. ZIPGELER, Po
than internarial span (B-N/IN = 0.71), Nares located
laterally on snout, directed dorsally, Canthus rostralis
stralht, loreal region coaneave. fueial shelf
conspicuous. Eye prominent. pupil horizontal when
constricted, Fingers short, unwebbed with flattened
subarticular (abercles. (Fig, 1B), Terminal dises
undilated. Palmar jubercles flattened but conspicuous,
Several supernumerary Lubercles. present: fingers in
order of length 3>4>2>1,
Hindlimbs: short (TL/S-V 0.36), ‘Toes relatively
short, Unfringed and unwebbed (Fig. IC); inorder of
length 4>3>5>2>1. Small flattened inner
metalursal tuberele, and tiny rounded outer metatarsal
tubercle. Subarticwlar tubercles small and not
prominent,
Dorsal surface sparsely tubercular, tubercles linear
wong length. Prominent tubercles on dorsal surface
of legs. Veotral surfice granular with well-developed
Fig. 2. Bryobatrachiy ninins gen. ct sp. now A dorsolateral
and B. ventral views in lite (Paratype TMAG 1009) (Photo,
J. Vows)
BROWN, M, DAVIES & M, J, LITTLEIONN
coarsely granular pelvic patch. Small bifurcated
unfimbriated cloacal flap,
Dorsum dark tat in preservative with cheyron-
shaped black mark between eyes. purred clonguled
hluck markings from seapula region, dnd paired
moderately elongate black patches jn the coccyge!
region. Canthal stripe anterior to narty, through loreal
region and eye to axilla, Pale patch beneath eye (Pigs
I. 2),
Ventral coloration chocolate with cream speckling
(Fig. 2), dark chocolate suffusion on throat, Paired
voowl slits posteriorly al angle of jaw:
Colour in life
Dorsal surfave Shades of dark brown. Darkes! brown
markings distinctive varying in intensity and occurring
bilaterally in pairs in the coceygeal region and in
association with scapulae, Coceygeal pair le within
region of paler brown or “ground” colour, not hidden
when colour intensity changes. Larger patch of dark
brown occurs between scapulue and anteromedially
Which, when at darkest. can conceal upper puir ol
bilateral markings, Chevron-shaped mark of dark
colour between eyes. Small white patch above clouca
Limbs barred with dark brown. Other broad dark
hrowh patches Occur along paler brown flanks. Dersul
surfice of snout and inner thighs unmarked and paler
brown or “ground” colour. Paired canthal stripes from
thirts (0 flank broadest in ryinpanal region, Dark brown
canthal stripe passes (hrough most of the eye except
uppermost part of iris. Tris dark brown below and
iridescent pold above facial stripe. Lower stripe of
white or cream, commencing between eye and paris
and varying in width being narrowest, or with lower
edge hotched, below eye and including neither lowet
eyelid nor upper lip. Ventral surface dark brown
covered with irregulirly-shuped fine white spots of
similar size extending aver the limbs and sometimes
to Jower throat, Chin somenmes paler and less spotted.
Pale lemon-yellow tint on paler parts of throat and
forelimbs,
Dimensions (mm); S-V 22.6: TL 8.2: E-N 1751N 14,
| A
variation
There is little variation in external morphology, other
than colaur pattern. The protruding snour is not as
pronounced jn-all paratypes; in those in which it ts
most developed. it appeurs to be thickened and whitush.
The legs unr uniformly short (TL/S-V = 0,35
[0.31-0.38]), and the Head usually longer than width
(AL/HW = 106) [0.96-1.23, n= 9) ‘Phere 1s
variitbility in the relationship between eye-tu-naris
distamee and internarial span (E-N/IN = 0.89
JO7-L2 |). The tympanum as obscure or indistines
A NEW GENUS OF PROG FROM SOUTHERN VASMANIA mm
(Figs 1, 2). foes are unilormly untringed, but there
is Variability in development of the palmar tubercles;
in TMAG C009, these are particularly pronounced.
When present, the ouler metatarsal (ubercle is tiny and
(he inner metatarsal tubercle usually Mattened, A farsal
fold js present in TMAG C1O1), Rugosity of the dorsum
varies bul is aot conspicuous. Ventral granularity of
the belly 1s variuble,
The ground colour of the dorsum and the extent and
nature of the markings varies. The three pairs of dark
markings on the dorsum are consistently present. The
markings between the eyes are chevron-shaped, whilst
those in the scapular region are ollen lyrate and can
vary in length, occasionally almost coulescing wilh the
voceygeal pair.
Dorsal colour varies from very durk brown to grey-
brown or lan, and, in darker specimens the anterior
pair of markings becomes obscure and often nierges
with the suprascapular pair. The coceygeal pair always
contrasts with the general body colour and qs distinctive
(Pig. 2A),
A pale or tan mid-vertebral stripe is present in several
specimens, und oveasionally a pale medial ventral
Stripe is present in puler specimens. The vental surface
can be dusky grey with white spots, or pale with dark
spots, The throat is always pizmented, but added dark
sulfusions are present in calling males. The throat has
a sulon iridescent hue in fife, and a lemon coloration
often occurs in thé axillae. Reddish patches occur along
the lower Hanks and in the inguinal region, A pale spot
ofien vecurs above the cloaca.
‘The pale stripe beneath the dark canthal stripe of
the head varies mextent. maximal development being
shown in Fig, 2,
Ostedlogy (bused on SAM R43671)
Skoli poorly oassificd. Sphenethinoid ussifted
medially, with ossification extending anteriorly
between the nasuls dorsally and ventrally, Curtilages
of nasal capsules. are considerably caleified veotrally
(Fig, 3). Prootic and exoccipital saperticially fused by
extensive culcification giving short and stocky erista
purotica. Exoccipial calcified dorsomedially and
ventromedially. Crista parotica not articulating with
elongate unexpanded otic ramus of squuinosal. Carotid
canal roofed on frontopirieuils medial to well
developed epiutic eminences. Occipital condyles
widely Separited. Fromtopurictal fontanelle widely
exposed for approxiniate length of orbit. Anterior
extremities. of frontopirietals: extend anteriorly to
anterior margin of frontoparietal tontanelle, Orbjtal
edges of frontopartetals slightly curved and angled
slightly posterolaterally. Nasals moderately well
ossified, approximately triangular and widely separated
medially, Maxillary process short, and wadely
separated trom well-developed preorbital process of
pars facjalis of manila.
Palalines tmoderitely robust, reduced slightly
laterally and curving posteromedially to overlie
sphenethmoid ut anterior extremibes of orbit
Parasphenoid moderately robust with moderately
Hroad, deeply creme culiriform process extending
approximately “4 length of orbit in yentral view, Alite
moderately short. relatively broad. extending slightly
posterolaterally, not overlain by medial ramus of
plerygoid, Pterygoid robust: anterior ramus in long
cantact with misxillit anteriorly, and with base ot
squamosal shaft and with cartilaginous quadrale
posteriorly, Squamosial robust. with shart zygomatic
Pig. 3. Briehatrachus nimbus gen. et sp. ney. A. doe) and Bo ventral views of the shull of S4M R43671. Seale bar = 5 mim
iy DF ROUNSEVELL, D2, ZIRGELER. PRO BROWN. M. DAVIES & Mo LITTLEIOHN
ramus and Jong unexpanded oie vans Hot overlying
erista paroled (Pig. 3).
Maxilla and premanilla dentate, Pulilal shell deep
with well-developed pitlaline processes of premuaxillae
not abutting medially. Well-developed pterygoid
process of maxilla. Alary processes of prethanillac
broad at base with narrower posterodorsal projections,
slightly concave, Vomerine fragments present on edge
of choanae on right hand side, Bony columella present
(Fig. 3),
Pectorul girdle arciferal and robust (Fie, 4A),
Omosternun Jarge. mushroon shaped will an elongate
narrow stalk; xiphisternum alse large and mushroom
shaped, with a short broad stalk. Some calcification
Of Niphisternum and epicoracoid cartilages, Slernum
cartilaginous. Clavicles moderately slender, curved,
poorly separated medially: coracuids robust,
moderately widely separated medially. Seapula
bieupnate, longer than chivicles. Supraseapula about
Vs ossificd,
Eight nor-imbricate prococlous presacral vertebrae.
Vertebri VILL Tused with saerum and with vertebra VIT.
Vertebra VII fused with vertebra WIE and with sacrum
(Fig. 5). Transverse processes of vertebra 1V with
bilateral anomaly (Fig. 5). Sacral diapophyses poorly
expanded, Relative widths of transverse processes:
(N>1VSIL>SD>V>VI>VU> VII
Urostyle bicondylar with dorsal eres! extending
approx. | length. Small rounded dorsal prominence
on ilium (Fig. 5). Dorsal protuberance pot prominent,
Vial crest absent, Pubis calcified,
Huinerus with strongly-developed deltoid crest
unteroproximally. Phalungeal formula of hand 2.2.3.3.
Curpus of six clements exhibiting moderate torsion,
©. rudiale and O. ulnare present, O, radiale larger of
the Qwo, Both elements articulate with O. radioulna
proximally and wilh each other posteromedially.
Distally, both articulate with durge transversely
elongaied ©. centrale postaxiale which articulates
distally with buses of QO. metucarpi Wl, TV und V.
Moderately well-developed Hange extends from
lateroproximal corner. Small calcified palmar sesumoid
proaimally on ventral surface (Fig, 4). O. centrale
preaxiile articulates laterally with O, raciale, distally
with ©. centrale postaxiale and with carpal element
oF ©, distale carpale 2 and O, distde carpale 3 and
luterally with basal prepollical element.
Carpal element of O. distale carpale 2 articulates
with carpal clement of OL distale carpale 3, Distal tips
of terminal phalinges knobbed.
Phalangeal formula of foot 2.2,3.4.3. QO. tibiale and
©. fibulare elongate and fused at either end. Bones of
approximately equal length. Two distal tarsal elements
present. Lateral elements largest, lying at base of O-
metatarsus HL and extending laterally to articulate with
medioproximal side of base of O. metatarsus TT. Second
element appears to be result of fusion of two elements
and lies at base and slightly lateral to ©. metanuirsus
Hand arhculates with base of O, mietatarsus | and ©
centre prehallucis, Distal prehallical element small
wid kKnubbed, calcitied.
Hyoid plate longer than broad (Pig. dC). Base of
ulury processes occupying Vs to 4 of lateral edges ol
hyoid plate; not pedunculate, Anteromedial processes
OF anterior hyale long und slender, Posterolateral
processes of hvoid plate irregularly shaped, moderately
long. Posterior cornu ossified,
\ ~~ J
c ir
i
i
Wl rd
yf ,
Fig, 4. Bryobacrachuy inibas gen, et sp. ney. A dorsil view
Ol the pectoral girdle B. lateral view of the ilu C. ventral
view of the hyaid (SAM RAR671). Seale bars = 1 ii
A NEW GENUS OF FROG FROM SOUTHERN TASMANIA 177
C
——
Fig. 5. Bryobatrachus nimbus gen. et sp. nov.: A. dorsal view of the vertebral column B. ventral view of vertebrae VI.
VIL and VIII and the sacrum (AM R143566) and C. ventral view of the vertebral column (SAM R43671). Scale bar = 5 mm.
ell
Fig. 6. Bryobatrachus nimbus gen. et sp. nov.: A. dorsal and
B. ventral views of the carpus (SAM R43671). Scale
bar = 5 mm.
Variation
Paratype (AM _ RI143566) has been cleared and
stained. In comparing this specimen with the described
male, it is apparent that the latter is probably a
senescent specimen on the basis of the high degree of
calcification lacking in the larger female specimen. For
example, calcification of the nasal cartilages, of the
omosternum, xiphisternum and epicoracoid cartilages,
of the tiny cartilaginous prepollex and the palmar
sesamoid is lacking in this specimen. In addition, there
is less calcification of the crista parotica region between
the exoccipitals and prootics.
Consistent with this view is the lack of ossification
of the sphenethmoid between the nasals dorsally and
ventrally, and the lack of fusion of the medial tarsal
elements. Remnant vomerine fragments occur on the
edges of both choanae in this specimen. The vertebral
anomaly on the transverse processes of presacral
vertebrae IV is not present, but fusion of presacral
vertebrae VII and VIII with the sacrum is consistently
present (Fig. 5).
There is little variability in other skeletal elements,
other than in the palatal shelf of the premaxilla which
is much more extensive in its articulation with the
palatal shelf of the maxilla.
(78 D, EF. ROUNSEVELL, D. ZIEGELER, P, B. BROWN. M. DAVIES & M.S. LITTIAIOHN
Advertisement call
Males call from the ground surface beneath dense
vegetation, and from prepared breeding chambers
within cushions of sphagnum or edaphic lichen. Male
advertisement culls are heard in loud chorus, diurnally,
in spring and carly summer.
The following description of the advertisement call
(Fig, 7) is based on the sequence recorded by G, F.
Watson, The call is a single quasi-periodic pulse train
with a duration of 1009-1281 ms (mean = 1157), and
consists of a series of 7-8 (mean = 7.5) short pulses
(duration; range = 7,2-10.4 ms; mean = 8.74 ms) with
rise times (attack) of about (0-3-5 ms and fall times
(decay) of about 4.9-5.9 my. The pulse rates range from
5,3 to 6.1 pulses/s (mean = 5.74) (Table 1). The calls
ure repeated at 4 rate of 2.65 calls/min, Although there
is a wide spread of spectral energy (as a consequence
of the short rise-times of the pulses), there are two
dominant frequencies of equal energy at 2100-2140 and
2680 Hz in three of the calls, there is only one peak
at 2140 Hz in two calls, and in the remaining call there
are two peaks al 2120 and 2740 Hz (with the latter
being lower by 2 dB).
0 200 600
The values obtained from each of the 18 calls
recorded by P. B, Brown are presented in the recording
sequence in ‘Table 2. From an inspection of the
numbers of pulses and dominant frequencies, it is
suggested that calls of three or four individuals may
be included in the sequence. The values tor call
duration, number of pulses and pulse rates, although
of greater range, include those of the individual
recorded by G. F. Watson. The dominant frequencies,
Where measured, however, are lower, ranging from
1540 to 1960) Hz.
Calling period
The calling period in alpine habitat is seasonal, Culls
in chorus were heard at the type locality from carly
October to late December. Earliest culling heard was
in the Hartz Mts on 1.x.1993, and the latest at Mt
Norold (146°IS'40"E, 43°15'30"S), on 2.111.1994,
Chorusing oceurred in rain and whilst snow fell,
In early spring, calling was interrupted frequently
hy heavy snow falls, and recommenced when thaw set
in. During summer, calls ceased during the warmest
part of the day at temperatures above 15°C and as the
800 1000 1200 1400
milliseconds
Rig 7 A wave lorm display of an advertisement call of a male of Bryehiurrachus nimbus. gen. et sp. noy, recorded at a
wet bulb air temperature of 8.5°C, in the Hartz Mountains. The depicted call is the first of the series recorded by
G. 1. Watson. (See Table | for more information. )
Tania |, Malues for si. advernyement cally af a male af Bryobatrachus nimbus gen. et sp. naw at the type locality ala
Wweebulh air temperature ef 85°C on 5.xth 1992.
Duration Pulse rate Pulse duration Dominant frequencies (Hz) Difference (dR)
(ms) Pulses (pulses/s) (ms) lower (DFI) higher (DF2) DFI-DP2
V148 7 5.43 Yay 2140 2660 0
L063 qT 5.716 10.35 2100 2680 0
1262 8 3.67 8,59 2140 none _
L281 8 5.56 8.40) 2120 2740 2
LL7R 8. 6.00 S40 2140 none =
1009 7 6,08 7.23 2140 2680 i)
A NEW GENUS OF FROG FROM SOUTHERN TASMANIA ie
TABLE 2, belies for Brdvertisernent calls of several males
of Bryobatrachus nimbus gen, ef sp. ney, recorded by BB.
Brown at the type lawality an 30,47, 1992.
Dominant
Call Pulse rate frequency
(ns) Duranon = Pulses (pulses/s) (He}
| 16st & 4 20)
2 Q25 7 6,58 1740
3 VROO i) ART 1720
4 2069 it) 4.49 1540
5 1Y1Y Ww 4.74 (sdge*
6 2037 Lu 4.46 —
7 1756 10 §.20 —
i 1356 7 4.46 IBOO
q O34 5 O53 TRO)
1a 1169 T SAY 1960*
i bos 5 5.493 1s00)
12 gad 4 A Ad 1aO0
3 A22N 10 4.09 1620
4 2256 i! 4.47 fA20
is Jidg {0 4.94 1600
16 2100 1 4.53 Ta0)
\7 Wz Ww 534 1X00)
Ls 2394 i 4.21 1a00)
* midpoint of band: ** second peak at 1820 Hz,
vegetation dried out. Individuals called in cool
conditions during the early wiorning and in the evening
(19:00-21:00) at Mt La Perouse on 1i1,1994 (S,
Corbet! pers. comm. ). Frogs were not heard at night,
Zicgeler (1994) observed culling in air temperatures
of 4.5-12°C and relative humidities of 62-94% _ The
frequency of calls heard increased when there was
precipitation.
Behaviour
The species is cryptozoic, and was collected during
spring and summer. Calling aides Were collected from
the surface of the peat beneath course, low vegetation,
from within breeding chambers or nests. in cushions
of sphagnum or other similar plants, beneath rocks
(Ziegeler 1994), or under branches lying of or
amongst vegetation, Fernales were found in breeding
chambers with a male or. when males were chorusing
in spring. On the surfuee of the vegetation, Mules
continued to call even when females occupied the sume
chamber.
Atthe type locality where a huge population exists
individual frogs are regularly spaced, remiuining
hidden, They appear nol to aggregate or lo use aperi
Fig &. A. Nest of Brvebutruchus nimbus gen. et sp aov-; B- subalpine moorland habitat at Hartz Mis; type locality in
middle distance viewed from the track, (Photo. J, E. & A, Wapstra); C. subalpine moorland habitat al 800- LOO tn altitude
at Mt Hesperus; D. Implicate rainforest habitat ut over 750 m altitude at Mt Bobs,
IKQ) D. E. ROUNSEVELL, DB. ZIFGELER, PB, BROWN, M. DAVIES & M. J.
Onin td
A. dorsal yew
Fu 9 Brwhettreetas nrghus pen. eb sp, nov:
of eyes: B. dorsal view of embryo at stage 4 (Townsend
& Stewart 85); ©, dopal views of traglers
LITTLEIOHN
surtiace water. None was collected during autumn or
winter.
Locomotion is by crawling or walking. In the open,
frogs usually crouched snd ferained sull until
disturbed and [hen sought cover. They jumped or swan
only when forced. In captivity. both sexes displayed
thigmotropism by forming individual cavities in which
to shelter th loose sphagnum. They entered the
sphagnum backwards.
Development
The species breeds in spring, laying eggs in nests
m moss ar lichen. Nests are concealed spherical
ewvities (3.5-4.0 em in diameter) in clumps of moss
dpproximalely 2-5 ¢m below the surface (Pig. 8A)
They can be exposed by purting the stems of moss.
Single males, male-female pairs (sometimes in
implexus), eggs, or froglets were found in ten nests
ina small area of the type locality on three visits
(24. xi1.1993, 11.1.1994 and 281.1994), On the first visit,
tive nests were found. One conuned 6-10 eges, another
vontained five froglets (Pig. 9C), the third held a single
male frog, and the remaining two nests each held a
male-female pair of adult frogs (one pair included a
gravid female and a male in amplexus). On the second
Visil three nests containing eggs were found. Two
contained 12 and I4 eggs. Those in the remaining nest
could not be readily counted as the jelly surrounding
(hem had begun to merge. The last visit produced two
hew fests, one containing Y eggs, und another
containing 5-6 freglets. Each nest was found in a
separate cushion of moss, Other empty and possibly
disused nests were found in adjacent patches of moss
m the same ares,
Eggs containing embryos (Fig. 9B) at stage 5 of
Townsend & Stewart (1985) were located on 10,911.93
but not in nests. Two groups of eggs were found on
The sufface of the yepetation and contiined three and
4iX exes respectively. The latter included four dead
embryos infected with fungus.
The mean diameter of four of the newly laid egus
found on 11-94 was 349 mm (3.33-3,65), and the
mean capsule ciaineter Was 13.57 mm (13,02-14.14),
They were surrounded by a single jelly membrine (Fig
9A) These eggs cannot be ascribed to. a stage as for
the direct developing Aleutheredactyluy coani
(Townsend & Stewart 1985)
The mean S-Y Jength of five metamorphs was
6.0 mm (5,4-6.35) und the mean total length, 14S) mr
(13,.97-19,.05), Gal length varied considerably (Mig. 9C).
Tail muscle is well developed and, tail tins narrow,
equal in width dorsally and ventrally. Tail fins do not
appear to be yasculamsed. The body is heavily
pigmented: that on tail muscle being finer and that on
tail fins patchy. In life the body and tail are covered
with fine white dots. There is no cloacal tail piece
The mouth extends beyond the eye and the fingers and
loes appear to be fringed,
A NEW GENUS OF FROG FROM SOL THERN TASMANIA {st
Habjrat
Brvobatrachus nimbuy is tound in subalpine
moorland (Fig &C) and implicate muntorest in soulhem
Tasmania (Fig, 8D), It is restricted to poorly-drained
sites from lowland to subalpine localities (Ziegeler
1994), The soil type 1s peal overlying sandy or stony
substrate (Ziegeler 1994), Lr occurs on Pre-Cambrian
metamorphics, Upper Curbonilerous-Permian
sedimentary deposits. Triassic sandstone and
Pleistocene glacial deposils.
The subalpine moorland comprises (diverse cinge
af plant communities. Five recorded communities are
Fpaeris serpilifolia-Empodisma ininus-Gleichenia
alpiaw Sernland. Leptospermum nitidum-Gahitta
yrandis-E. minus tall shrubland (both occur at Hartz
Mts [Fig 8B), EL serpilifolie-Richea scoparia low
shrubland with sielia alpina, G. alpina and E. nurs
(Aduinsons Peak [46749 43°21'S}), E. serpilifelia-
Melaleuca \quamea open heath (Mt Sprent) and
Carpha alpina-Isophysis. wxmanica herbtield (Mt
Sprent. Mt Hesperus [146°4/E, 43°07/20"S] [Fig.
BC|) (Ziegeler 1994).
The implicate rainforest. cormmunities are flopistiwally
complex. The shrub layers aré dense and species
diverse at all aliitudes, At higher altitudes the dominant
trees are: Nothojaeuy cunninghenui, Eucrypliia
inifliganit, Atirotaxis seluginoides, Nothofagun eurnl
and Phyllocladus ayplentifolins and tie ground cover
is dense moss and/or slsiedia alpina (Pig, 8D» 750 m.
ML Bobs 146%36'E. 43°18'S), At low altitudes NV.
cunninghanii, Melaleuca squarrava and P.
asplentifolius are dominant and round cover is dense
mass and Blech wars) (Alexdnder Creek
146°0S'E. 43°26'S) (Ziegeler 1944),
The climate is in the perhumid cool zone of Gennlli
(1972) but is cool and consistently wet, The meun
annual preeipinttion iy 2500 mm in large parts of Uris
region und is reveived us niin, snow, hail, fog, mist
and frost evenly distributed across the seasons (Bureau
of Meteorology 1991),
Distriburion and abundance
Bryvbatrachus nimbus occurs in mountains ip
southern Tasmania soath of 42°46" latitude (Fig, 10),
li is known from 15 localities. mostly desenbed by
Ziegeler (1994). that range from sea leyel ty HOO m
in an area approximately $0 km = 50 km, The
localities are hounded by Me Sprent in the north-west.
Hartz Mis in the north-east, Mt La Perouse rm the
south-east and Bathurst Harbour area in the south west
(Fig. 10), All localities are south of the Huon River
und Serpentine River and none is over 40 km from the
sea. At the northern and eastern edges of (his
distribution, B. nits ts confined fo sites above SOO mi
altitude whilst in the south-west, its range descends
to near sea level,
Hundreds of mules.were heard m chorus at the type
lovalily during spring 1992, Other large popoladons
oveur at known locations (Ziegeler 1994), Although
geographically restricted. the species is abundant
within availible habitat, and calling ntales were
estimated (9 occur at densities of O.L-L.0/m--
Searches conducted further north at Philps Peak.
Tyndall Range, Mt Field, Mt Anne. Mr Weld and Mt
Picton failed to locate the species Uespite the presence
of suitable habitat and weather conditions (Ztegeler
1944),
Compervation stalus
The species issecure. All locations where it has been
recorded are in reserves. us is most of the surtable
known habitat. [ts habit shows evidence of no fire
or only very low frequencies of fire.
Erymalogy
From the Latin nimbus (= rain cloud),
reference to the habitat of the species.
wilh
Common name
The name “moss lroglet” has been coined and useel
for Ums species (Ziegeler 1994),
L. Pedder
Huon Res
145'30!
14600!
146°30'
k
MY Sprent
Harte Mt
ML La Perouse
Mt Hesperus
Mt Norold
6 Federetnan Fh
7 Mi Bats
6 Adanisons As
§ Magnhant fidar
10 Breeiptous Blin
17 |ranbound Aa.
12 Louiba P_
13 Bathurst Ra,
14 Mt Melaleucu
15 Alexander Ck
Fig. 10, Distribution of Bryobar rachis mimbys gen, et sp, now,
in southern Tasmania, A, Southeastern Austratia, and B.
Jowalised distribution fy Tasmania. Closed cipeles ure
localities from which spevimens have heen collected: npery
(numbered) cireles are call records
1X2 DOR ROUNSEVELL. Do WIECBLER. PB, BROWN. M. DAVIES & Mo J. LITTLRIOHN
Comparivon with other species
Bryehatrachis timbus is distinguished trom other
rayobalrachine frogs in Tasmania in the following. ways.
The species is not associated with badies of open
surface water, This most likely to be confused with
symputne Ranidelia fasmuniensts trom which W ca
be distinguished by relkitively-short, unfringed toes and
by the Jack of brilliant carmime patches usually presenc
on the Hanks and the concealed surface of the thighs.
The call of R. hasmianiensis ts a quavering “blear’,
slowly and irregularly repeated (Litileyohn 1970) and
its nerolabiat includes yegenition an the margins oF
Open and running water. R, sigeifera hus relatively
Jong. fringed tocs and is found in the vegetation on
the margins oF permunent or temporary water bodies,
The call is a series of short rapidly repeated notes
“erik eviek eriek erick” (Martin & Litthejohn }982)
Georrinia laeviy his a rounded snout with & sooth
ventral skin which is usually boldly marbled with dark
brown imurkings. The species os found im dry
selerophyll fopen) forests af low altirudes (Martin &
Liulejohn 1982). These authors describe une cull as
a harsh grating “cra-d-a-d-ack cra-i-a-a-ick, crack.
erick, crack”, The other myobatrachine species in
Tasmania is Preudsplrvme semimarmorata which has
asmooth ventral surtace strongly marbled in black and
whit, The head is rounded and the undersurface of
the thighs are bright yellowish orange to Orange in Life!
(he call is a short, harsh “creek” repeated slowly and
irregularly (Maclin & Lilthyohn 1982)
Comparutive meterial examined
Arenophryne ronda: UAZ B531, B54, B762,
ASTY-81, ATQ0-L. Assu derlingioni: UAZ Al33, Crinte
veonnanar UAZ BIS4, A406. Geacrinia laevisy SAM
R4260B. RAQR2ALE, G. feaics SAM RS7S7TA. G.
Wietoridna: SAM R9425 (2), Mybhatrachus venildil:
UAZ B48l, B534-5, B757. ATSY. Paracrinic fueswvelli,
UAZ B75), Runidtella bilingua: UAZ BI927, BYAS, R,
deserricala: UAZ BI930-31. RL elauertic UAE AVL,
RR oonyimifera> UAZ AI92. AOS BY2D RK,
parinsignifera: UAL AIS, BIG32-33.R, rented: UAL
BIS28-9, R. riparia: UAZ AlB4S, AISO. AlOS. R-
siphifera: UAZ AIW3, AIST, BROS ODR R
vubinsignifera: UAZ AIS), RK. taxnranionsiy; UAL
AIRG, Preuduphrvne sertimnarmmata: UNF B36 P
Dibroniz UNE AST? AI72. BS32, BS4dQ. FL ceniacea,
LAY A200. BS37, B764. FB wxenthert: UAZ ALO,
653.9, BIAS. PB acvidentaliy' SAM R17522.
Taydactylus acutirosiis: UAZ BISY, ABNS-6. 7
eungellensis: UAZ A392, Ty checphilus; UNE ATTS-6,
BIS2, Uperdleie altiysinu; SAM RAAB. U, arenteret,
SAM RI7347. U aspera, UA AST2-&. Uo borealis:
UAZ A863, B862, A226. LL capinilaras SAM R29586.
UL crassa: UAZ A869-70, B871, B483, B4k6. LI flevce.
SAM R29603, K296067 L/ vlanidnloxe> SAM
R77082. UL guindeta: UAZ ASI8-816, B3Y7, Ba7y.
laevigata: UA AGNI, BRIT U lithemrenda, UAE
Alé7-8, BR96, BRI2, A74o-R06, LU, littlejoints UA
AITI2, AVTU. BITIZ-1716. LD martin NMV D2363f.
UL anieromedess UAZ AIWW22. Lo mimilas SAM
R29642-3. R29645-6, Of myeberei. UAL A582,
ABSO-9, EF rugosa; UAZ 1012, AIN13, A816, Bald. UL
taipas UAZ A594, BS64, UO rrarhvlerma UA2
A622), ABYT-4, AS9S-6. UL) wleris NMV 123634, SAM
R29659 R29652,
Discussion
Brvohatrachus nimbus exhibits a number of urtusuil
featlires in its morphology and reproductive biology,
Many of (he chitracters contabuting to the recognition
of the wenus and species are influenced by helerochrony
in other myobatrachine genera (Davies 1989). The
presence of teeth on the niaadlary arch, reducuon of
the vomers and ubsence of vomerine teeth, presence
ofa columellit and reduction of the palatines, laterally,
are all presumed labile teatures that vary
intragenenvally in ¢.g.. Uperoleia and Crinta tvensie
Hever eral (982). However, nove ot these featares
eilher singly or in wombination is delinittye o]
Rrveburrachias. AU are coupled with the unusual
feature of fusion of ihe posterior portion of the vertebral
columo with the serum,
Verichral fusions tend to be in an dnterior/posterior
sequence (Trueb 1973), und lusions pf presacrals Vil
and VIL with the sucrum are unusual, The nature of
the fusion cannor be described as a presacral shield,
as known in a few taxa including Brachvwephelte
eppiphium (Brachycephalidae), and thought to be
protectlye on the basis of lerrestrialisna exhibited by
such taxu (Trueh 1974).
Fusions approaching thutsbowa by & immbus have
been recorded in the Buforidae (e.g , Didvnemiipies,
see Gjandison ISL), u) Denedrebates (Trueh 1973), ard
in the Pipidae; wllhouyh the fusions reported 19
Pyendhymenuchirus aod Aynenachiras by Coummatelhi
& ‘Troet (1988) ure inferred. and not identified hy
femhant vertebrae or recorded Spinal foraniina,
Vertebral fusion in Pipe miversi mmre closely
approxiimtes that in Didynanipts (Troeh W984). None
of these faimlies ts native to Australra,
Although conturming to the myubatrachine pattern).
the width of rhe junction of the alary processes with
(he hyoid plare 15 notas broud asan most other genera
and approaches that shown by the enigmatic genus
Rieohatrachus (Davies & Burton $982) Mahony etal
1984), the subfanilial placement of which renyaits
comentious (Tyler 1980) Littlejohn et a, 199A),
Terresinial Gey deposilion and direet development
have evolved a number of tintes in Austratan frogs
(Martin 1967). Reproductive modes include complete
iMracapsulur developnient (ea. in the onerohylids
Sphenophevne and Caphivalas), through hatching, al
ANEW GENUS OD TROG TROM SOLTHERN TASMANIA 183
a relatively late Stage an onlogeny (Gan. Une
myobatraching Pseadophryne) to non-swinning, mon
feeding larvae (hal are subject Lo brzarte tops of
parental care (eg. the myobatrachine Awe and the
rheohatrachine Kheoburrachas). Allof these gradarions
of direct development are found within” the
Myobatrachinac, amd reproductive mode can
somelines vary inrayenerically, In Pyeadopliryne, P
douglas: differs from congeners in laying eggs mn water
and thus not relying upon lauding of tests to iia
hitching of larvae at Tater stages (Main 964) Bradford
& Seymour 65), whilstan Geecrinfa, Go raved and
G. luiea do not have free-swiniming larval stages (Main
er ad. JOS. Main 1964),
Tolracapyular development ts known for the
monotypic genet Myobatrachiy and Arenupliryne
(Roberts 981, 984). and is interred for Meracrinin
(Main eral. 959), The larvae of Geoeritia rasea are
inactive in, broken-dewn egg capsules which lll
shallow depressions in damp soil or in the rotten
cemres of hollow jogs (Main er al 1959), Larval bite
ob G. lutea resembles that of G. rasea (Maryn 163)
Such larvae are highly modified, having ne niwuth disc,
large yolk sue, and ain elongate tail (Watson & Martin
1974), The structure of the tail repoimed for G, rosea
is similar to that observed in Bo nimbas froglets,
Although deails.of the lite history of B. rimbity have
pot yet been described, our Hryited data medicare that
development ts intracapsular sanitil the Cina statyres of
metmorphosss, and thatthe Froglets stay in the nest
with the broken down ege capsules antil the tatis fully
resorbed dnd metamorphic climax is reached.
The diamelers of the eggs al 33 3,7 mm are smaller
than ovarian diameters recorded for Arencuplirvne
rotdnada and eye diameters in Mvhatrachay goutiii
(Koberts IKI, 1984), but within the range recorded
lor Pseudephrvne spp. dnd lor Ceeocrinia vjetortara
und G. Juews (see Tyler 1989 for compilation of data).
AlL Gl these Species extibit forms of direc
development. Capsule diameter is partoularly large,
being almost double that recarded in the field for
Myobatrachus gouldii, and LS times Unal of eges.
hydrated inthe laboratory (Roberts 8h) Clutelr size
corpures Closely wath those Of other direct-developing
species (Tyler 1989).
Four other Tasmanian species (Ranidella scenifera,
KR, daxmaniensis, Generinia laevis, Pseadephryvne
seminarnorad) have advertisemen( calls composes
ottrains af pulses, The short pulse trains of Bo adn
ure sumilur to those of AL viweit/ere, but the ell
repetition nites ind pulse repetition: rites and daminant
frequenci¢s are higher iy this latter species (Littlejohn
W6d, WO). ‘The call ol AL fasmaniensix is more
complex, being composed of a group of pulse trains,
or notes. of high and regular pulse rites (Litthejahn
W70). so That has a heating quality, The call of G
(aes commsisis ab series of pulse Unis in which tre
pulse intervals decrease through each note (Littlepohn
& Martin 64) Harrison & Lirdeyoho 1985). The call
of P semrnarmorata consists oa single complex pulse
trun, usually with biphasic structure (MeDonnell cz
al 1978). Henee, the advertisement call ol Be yurahiry
is sulticiendy unlike those of all other species oF wnnran
occurring in ‘Tasmania to be a reliable miateator for
identification, Possibly, the most similar advertisement
call ob Southern myobutrachicl to that af Bo nirbiss
is that of Ranidella elavert of south-western Western
Austraha, which consists of a slow pulse trarn and very
short pulse durations (Littlejohn 1959, W6l Moo
Litleyohn & PG) Lithleyolin unpubl. ).
Brychatrackas ninbus is most similar im external
morphology and cranial Osteology to Ranidella
laxmantjenyiy (Dayies unpubl.) i ats reproductive
higlogy lo Geecrinia rayew and CG. ludea, and i the
structure of the hyoid ta Rheobarrachus, It ie not
possible. therclore, to idemtily dhe sister Liaxgn to the
2cnus on the basis of the phenetic comparison
presemed here. Such ain ientlicatian must awatt a
detailed unalysis of, at teast. the subfamily
Myobatrachinae, incorporating all available data,
8. nimbus appears to be confined to southern
Tasmania despite (he presence oF apparently suilable
habrat in highlands beyond, tt has been sought north
of the Serpentine and Huow River systents without
success, The distribution might be explained by the
exlentol Plastocene glaciation in the central highlands
of the island, and the severity of the cold_ dry climate
inte free areas (Galloway 1986) making the centrad
highlands unsuituble forit in the past, coupled with
g lack of northward dispersal since,
The region currently oecupiced by the spevies has o
continually wet eqoaable climate produced by your
round high rainfall within the perhumid cool climatic
qone (Gentilly 972). All loyalties are less than 40. ker
from the seu, and are subjeet to coastal climatic
influences, paruculatly inereased precipitation trot
nropraphic interaction with moist prevailing west tr
south-west winds, The direet development of the
juvenile stages requires a climate free from the
exirenes oF desicomion. This most unlikely that the
known habitat pes could support the speeies until a
very loag Line alter a wildfire (Brown & Podger 192)
A ontimbus can be regarded ay belouginy: to tire
inlolerant Cormmunities, and the ogeurrence of fire
represents [hy wreatest polenta! Uiredt oats survival.
During the present work, several elutehes of
developing eggs Were found ou the surface of moss at
Narty, Mountains. ‘The site had been covered by at
sanowdritt tor sometime, Egglaying beneath the snow
rather than ji nest could aecount for their location and
Ialet exposure alter the thaw. Phe erubryes would be
prone 16 desiccation belore completing wher
development, Altematively. stepping on patches at
moss when walking across the site could eject eps
ld 0. TE. ROUNSPRVELL. DL ZIGGELER, PB BROWN, M DAVIFS.& M, 1, LITTLEJO}IN
fron nest! Chambers, Whichever is the ease, caution
is advisuble when working al breeding locations.
Acknowledgements
Dr Graeine Watson recorded the call used in the
analysis. Dr Dale Roberts located the first nese chamber
and helped to clarify the breeding biolagy Dr Roy
Swain and Mr J. Lim provided ussistance in the
laburatory. and Nikki Mitchell helped im the field.
Burbara und Peter Willson, Jayne Baliner, Sib Corbett
and Michael Askey-Doran reported the cally frou
Mi Norold, Bathurst Range, Moonlight Ridge and
Mt Mekieuca, We thank staff ofthe Parks and Wildlile
Service who conttbuted to this study und Assoc. Prats
A, A, Martin and M, §, Tyler for constructive reviews
of the mantiseript,
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THE DISTRIBUTION OF NEMATODE PARASITES WITHIN
THE STOMACH OF THE WESTERN GREY KANGAROO,
MACROPUS FULIGINOSUS
By D, PAMMENT, I. BEVERIDGE & R. B. GASSER
Summary
Pamment, D., Beveridge, I. & Gasser, R. B. (1994) The distribution of nematode
parasites within the stomach of the western grey kangaroo, Macropus fuliginosus.
Trans. R. Soc. S. Aust. 118(3) 187-196, 30 November, 1994.
The distribution of strongylid nematodes in the stomachs of ten western grey
kangaroos (Macropus fuliginosus), from Hattah, Victoria indicated that each genus
encountered, Cloacina, Rugopharynx, Labiostrongylus, Popovastrongylus and
Filarinema, occupied a specific region within the stomach. Their distribution bore no
relationship to histologically defined regions of gastric mucosa, and the pathological
changes detected in the mucosa were related more closely with mucosal type than
with local nematode densities. The anatomical and histological features of the
stomach of M. fuliginosus are described.
Key Words: Nematoda, Strongyloidea, Macropus, Macropodidae, distribution, gastric
anatomy.
Jrnsactions of the Raval Suciery af 3. Aust, (94), MBCA), TST,
THE DISTRIBUTION OF NEMATODE PARASITES WITHIN THE STOMACH
OF THE WESTERN GREY KANGAROO, MACROPUS FULIGINOSUS
by D. PAMMENT, |. BEVERIDGE & R. B, GASSER
Sammary
Pavimenr. D. Bevermnor, 1 & Gasske, RB. (994). The distribulion of nematode parusites within the stomach
of the western grey kingurou. Macropus filiginasus, Trans, Ro Sac 5. Aust T8(4) 187-196, 30 Nayember, 4
The distribution of stroneyhd nernatedes in the stomachs of ten western grey kangaroos UMacreps flings),
ion) Hattah, Vietorul miicated that each genus encountered, Cloacing, Rugapheryns, Labtosirngylis,
Popovastrongylus aud Flarinema, occupied w specific region within the stomach, Their distribution bore av
relationshipete bistolagically defined regions of wastrie mucosi and the pathological changes detected ir the mucosa
were related more closely with mucosal type thin with Jocal neniatode densities. The anatomical and hystological
features of the stommieh of ML Juliyinosus are described.
Key Worps Nematoda, Strongyloides. Macrepuy, Macropodidae, distriburion, gastric anttory,
fntroduction
The differenual localisation of muluple, closely-
relited parasitic nematode species within the
gastrointestinal Lract of their hosts has been investigated
extensively in the case of the oxyuroid parasites of
tortoises (Petter 1963, 1966; Schad 1963). Niche
diversification has been demonstrated on the basis of
differences: in the linear and radial distribution of
nematode species und in their ceding behaviour Inglis
(1968) sugested that because of the complex anatomy
of their stomachs and the diversity of nematode species
harboured within them, the kangaroos represented an
equally suitable group of hosts for studying the
coexistence of congeneric and conflamilial nematode
species within the same host. Studies undertaken to
date support [nylis’ (968) suggestion, but have been
limited to the red kangaroo, Macropus rufus (see
Mykytowyez 1964; Dudzinski & Mykytowyer 1965;
Arundel eral. 1979) and the tammiur wallaby, Maeropus
eugenit (see Sniales & Mawsen 19784). Tn (his study.
the distribution of nematodes within the stomach of
the western grey Kangaroo, Macropus filiginosus. was
investigaled and relationships between this distribution.
and gastric mucosal histology examined, Pathological
changes in the gastric mucosa were also investigated.
Examination of M. fuliginosuy allowed analysts of the
relationships of four contanmiial genera. and in the cuse
ol one genus, Cloucina, the comparative distributions
of several congeneric species,
Methods
Collection of material
Stomachs were obtained from ten western prey
kangaroos, Macropus filiginosus, culled at Hattah
Lakes National Park (34°45'S, 142°IS'E). Victoria in
November 1990. Animals were shot. weighed and sex
and body measurements were recorded, ‘The stomach
was removed rapidly from the carcass, the besophiagus
Department of Veterinary Scienve, University of Melbourne.
Parkville, Vie. 3052.
Fig, 1 Stomach of Moeropiey fidtginayus, unopened, 1.ines
und numbers indicate (he positions of ligatures und the order
in which they were applied
Amitomical landmarks shown are the pesopliwzus (or), rhe
proximal diverticulum (i). tte gastric poueh (2) and the
pylorus (p). Seale bar: 5 cm.
Fi, 2. Stonmeh of Macropis fidiginesis, opened to show
distribution of the principal Ivpes of mucosal surfices,
delineated by while broken fines. the ghindulur mucosa at
jhe proximal diverticulum (dj, the squamous epithelium
of the saveular and tubular forestomach fy), proximal and
distal to he oesophagus (o) and the gastric sulcus (gs), the
glandular mucosa of the tububie forestormach (a. the
Mocwsa of the gastric PAUCh (QO) aril pylorusi py Seale hat
Soc.
IKK D. PAMMENT. |
and pylorus were tied with string, numerous stall
punctures were made in the stomuch wall and the entire
stormach was immersed in 50 litres of neutral buflered
15% lormol saline (6% formaldehyde),
In the laboratory, stomiehs were washed in water
ju remove formalin. photographed and weighed, then
subdivided with ligatures and opened. Anatomical
nomenchiture for the regions of the stomach lollows
Hume (1982). The ligatures were placed in the
following order (Fig. > at the level of the oesophagesl
opening (1), midway belween the oesophageal opening
and the extremity of the saccular forestumuch (2) ut
the junetion of the tubular forestomach with the gastric
pouch (3) Andhat its junetion with the pyloric antrum
(4). Subsequently, the tubular foresturnach was
subdivided by three ligatures (5-7) placed equidistantly.
The content from each seehon was removed and
weighed, Within each section of the stomach, a
(ransverse strip of lissue was removed, embedded in
parallin, sectioned at a thickness of Aum, and stained
with haematoxylin and eosin for histological
examination. The extent of the different epithelial
regions of the stomach was determined by cutting oul
the entire squamous, mucus-seereling, glandular
(proximal diverticulum). gastric and pyloric regions
and weighing them The average weight per cn of
each mucosal type was determined by weighing 2 cm’
portions of each mucosal type, and the anea occupied
nie each stomach was calculated by dividing the two,
Asa control for histopathological examination, a
single adule Ad. fuliudneses front Healesville Sanctuary,
killed for other reasons, was examined. This animal
had been treated regularly with anthelmintics to remove
nematodes prior to dealh by intravenous injection of
a barbilurate Portions of gastric mucosa were fixed
immediately in LOM neutral hullered formol suline (4%
formaldehyde) and were processed for histological
cxurimlions us deseribed ahoye.
farasirological absérvarians
The total number of nematodes in each section of
(he ten stornachs was estimated by a dilution rechnique
BEVIERIDGE & R. B GASSER
(Clark ef al: 1971) and the number of each species
present delermined by clearing all nematodes in the
appropriate subsample in lactophenol and identity ing
thei to species under a compound microscope:
‘To establish whether ornor parasite distribution was
affected by this ntethod of Fixation. twa additional
stomachs collected were ligated with string at sh points
along their length before being immersed in
formaldehyde solution,
Representative specimens of each nematode species
have been deposited 1 the South Australian Museo
(SAM). Adelaide. Nomenclature of species. of the
genera Lahtosrranuviis and Clodciner is currently unde
revision, Por this study, the species names applied ane
the same as those used by Beveridge & Arundel (1979).
Results
Gross anatomy of the sremeach
The gross anatomy of the stomach af M, fdistroxts
(Figs |. 2) resembles thatot M, eigenteus (Langer er
al (980, Dellaw 1982). Wet weight of stomach
contents (Table 1) in individual sections demonstrated
that sections | and 2 (= the saccular forestoniaeh
|Langer ef al, 1980)]) constituted 22.3% of Jotul
stomach weizht, sections 3-6 constituted 73.5% of towl
stomach weigh (= tubular fonestonach), and sections
7 and 8 constituted 4.4% uf total stomach weight (=
hind stomach). The pastri¢ sulcus was prominent and
extended halfway along (he tubule forestomach,
Relative surlitce areas occupied by different epithelial
types (Table 2) demonstrated Utat squamnous and
mucus-secereting epithelia were the two predominant
types. The proximal diverticulum) of the sacculir
forestomach was lined by a distinctive, glandular
epitheliam, The remainder of section | and section 2
of the stamach is lined by squamous epithelium. Apurt
from the gastric sulcus which is vovered with @
squamous epithelium, sections 3 to 6 are lined with
curdiae epithelium, Section 7 has un avid-secreting
fundic epithelium. while section & is lined by a
disuociye pyloric epithelium
Tanth | Wer welehiy of content af eight different reatons of the stomach of ten Maccopus tuliginosus fron Hatal Lakes
Neirional Park, Wiotertt
Proportion of
Stomach Meun Range Stundynl Tutal Weight Predonnunt
Section No. weight (4) (g) Deviation (%) Fpithelal Type
1+ ag 46-151 32 42 ws (equa ETON)
g AAT 55-638 322 18.1 5
3 14§ 10b-5-44 134 1p 3 m (MUCOUS)
4 405 145-(208 299 21.8 m
5 AZA 128-834 335 QW m
4 320 T1774 292 19.4 mn
7 78 8-1 hv 5) 17 & (wustric)
8 \2 as 12 0.6 p (pylons
Tou! 2136 712-La7iy — rou
* Por key io stomach sections, see Fig, 1.
DISTRIBUTION 6 NEMALODES IN RANGA RODS Is
TABLE 2. 4rea, av a percentage of the tote. Cevupied tn
differed epithetial surfaces in the stomach ef ten Macnpus
fulizinosus frat Heath Likes Nasional Rieck, Pletarin:
Pervemtage of
Lpithelual Total Stomach Standura
ype dren Oecupied Deviattan
Squamous 3) "
Mucus seereting
(tubular
foresionmch) nes Ww
Glandular
(anterior sustric
diverticulum) S |
Crasinic 5 4
Pylone 5 i
Histolowieal feqrures. of gastric epithelia
Squamous epithelium
This epithelium ranged from 80-320 yr in
thickness. Basal cells in this region were irregularly
cuboidal in shape. nucler were relatively simul,
basophilic and had a prontinent dark nucleolus. Che
cytoplasm was compact, eosinophilic.and cell miargins
were indistinct In the mid-region of the epithelium,
velly Were enlarged. cell margins were readily
distinguishable, irregularly cuboidal or polygonal in
shape with an enlarged, pale nucleus and promineni
nucleolus, Murgination of chromatin was evident in
most nuclei. Towards the lumen of the stomach. cells
hecume squamous and the evioplasm more densely
eosinaphilic, The quele: were indisuner, On the surtace
isell’, flauened keratinised cells. were visible sloughing
into the lurnen, and in several sections, a distinet layer
of adherent bacteria was evident, closely applicd to
the superficial keratinised layer. The squamous
epitheliany was folded. with interdigitations of (he
Lami propria projecting into the base of the folds
Cardlae epithelium
This epithelium was up to 600 pin thick and was
composed of elongate, parallel glands 45 jan in width.
The cells lining the glands were cuboidal to columnar,
25 pm in length and 10 um wide. The glands gonsisted
of wo disunet cellular components. The cytoplasm ot
cells atthe base of the glands was faintly cosinophilie
and. had yt Joumy appearance; cell boundaries were
distinct. Nuclei were situated at the bases of the cells;
they were slender and. elongated, and usually
demonstrated emargination of chromatin and a
prominent. small nucleolus. Cells in the mid-region
of the glands were gosinuphilic and the cytuplasn had
4 granular appearance. Nuclei were large and rounded
with w prominent nucleolus and punctate, emarginate
chromatin. Surface epithelial cells were longer and
more slender than cells within the glunds. ‘The lari
propria was very nurrow and was bounded titerhally
by a broad lamina musculins mucosae-
Epithelium of the proximal diverticulum
This epithelium of the proximal diverticulum of the
saccitorm lorestomach dilféred (rom thar of the cardiac
region (Fig. 3). The epitheliuin was-extremely thick,
up to 1.8 mm, and was composed of elongate glands,
up to. 1.55 mu long and 0.4 mm wide, which became
gintions rwards the base. Cells futing the glinds were
cubordal in shape, approximately yan by WOjan in size
with w highly cosinuphilic cytoplasm. The tuclet were
sniall and were Situated at the base of the cell wilh
emargination of chronvatin ar with chromatin
distributed ina purictate pattem, and a smiall nucleolus.
The turnina of glands were dilated und contuined a
highly eosinophilic Muid, presumably secreted by the
ghands.
Fundie epithelium
This epitheliam was 155 mm thick and was
composed of parallel, elongute ghinds. The histological
appearance of the glands wassimilar to that found in
olber mammals, with elongate surface cells, Cubuidal
mucous neck cells at the anterior ends of the glinds,
and chief cells and parietitl cells towards the base,
Pylorie epitheliun
This cpithelium was 12 non thick and consisted of
vroups ol very long slender glands curving inward
fowards the stomach fumen, with slightly sinuous
bases, Cells at the basal region were cuboidal to
columnar, with pale. foamy. eosinophilic cytoplasm,
prominent nuclei with emargination of chromatin and
alargenucleolus. In the mubreginn of the glands. cells
tended to be jow cuboidal in shape with & more
eosinophilic cytoplasm than cells at the base, The
lamina propna was narrow and a prominent lamina
muscularis mucosae was present uninedintely below
the base of the glands.
Histopathologiea! changes in the epitheha
In spite of the large numbers af nematode parasites
present in the stomachs of kangaroos. no gross
pathological changes were observed in the mucosa
The squamous epithelium demonstrated fow
histopathological changes, Occasional small clusters
of lymphocytes and macrophages were evident jn the
mid-region of the epitheburn. or in the lamina propria
(Pig 4). Eosinophils were occasionally prominent it
the lamina propria. Accurmulations of inflammatory
cells were visible, Most frequently close to the junction
of the squamous and cardiac epithelia.
The glands of the cardiac region were intact histo
logically and no developing nemutades were seen within
glands. The lamina proptia. however. was diffusely
D. PAMMENT, |. BEVERIDGE & R, B. GASSER
190
Bght OF oP Ae Te!
ws
HISTRIBL:- TION OF NEMATODES IN KANGAROOS
infiltrated with mononuelear cells and granulocytes,
principally eosinophils. with oecasional large, local
accumulations of cells (Figs 5, 6), The submucosit was
also-uniformly infiltrated with the same inflammatory
cells.
The proximal diverticulum) invariably exhibited
prominent infiltrations of the lamina propria with
mononuclear cells, predominantly plasma cells, and
lymphocytes. with a few eosinophils, sometimes to the
extent that iflammatory cell accumulations obliterated
individual glands (Fig. 3). In spite of the sometimes
intense but chronic inflammatory reaction, very few
nematodes were scen within glands, Those encountered
were wilhin ar just below the epithelium and were
surrounded by large accumulations of inflammatory
cells. The fundie mucosa exhibited nu signilicsnt
pathological changes. The pyloric mucosa was
histologically normal in most animals, but in several
kangaroos, the rmegalo-schizonts of a coccidium
developing towards the base of the glands obliterated
numerous glands in the vicinity, As gommon us the
schizonts were areas of neerosis and inflammatory cell
debris indicating uw site where a schizont had ruptured
to release merozoites. An intense inflammatory
reaction surrounded megalo-schizonts, characterised
by large numbers of neutrophils and cosinophils,
19]
Numbers and distribution of nematodes
The numbers of nematode parasites found in each
of the 10 kangaroos examined ure shown in Table 3.
The distribution of nematode species and genera in
terms of absolute numbers in each section wre shown
in Figs 7-8. Expressing the results as densities (worms
per g of stomach content) produced similar results. with
the sole exeeption of section | of the stomach, in which
densities of Cloucing spp. were higher than in other
sections. Distributions Of the genera differed. Cleeeine:
spp. occupied the anterior sections (1-4) of the stornach,
Luhiosirongvlus kings and Lo et. bipapillosus (sensu
Beveridge & Arundel 1979) ovcupied a relatively
restricted region in the third segment, and A, ausctraliy
and P pearson’ occurred in sections 3-6.0f the tubular
forestomach, Very few nematodes were detected in
regions 5-6 of the tubular forestomachs. Ip the gastric
pouch (section 7), the only nematode species
encountered was Ailarimema ausirdlis, This is & new
host record. A wustreliy was found in iwounimals, with
Iwo nemitodes in cach animal, No nematodes were
found in section $8. Comparison of data from these 10
kangaroos wilh those front the Qvo animals whose
stomachs had been ligated prior to Tixuation revealed
no differenves in the distribution of parasites within
the stomach,
Taney 2 Nembery of helainth parasites inthe stones of ten Maccopus tuliwinonus (vn Marah Lakes Natenal Park,
Victoria,
Labjustrangvlus Revupharvis Popovastronystlias
Kangaroo No, Clouciia spp: spp. australis pearsoni
I 480, 15 AT YU) 0
2 4.300 560) T.00 1.60
3 i400 1S 80,870 5.380
4 1310 Su 15,850) 25
5 2.200 0 (84,100 2,500
6 100 40 | 360 ail)
7 2,200 25 66,840 10)
8 &. 300 50 (21.10) 4AKW)
) 20 400 170.600 1,500
10 | 580 35 XL S00 125
Meun 2.415 20 $3,330) 1,600
Standard Error
of Mean 755 62 19,125 586
Figs 3-6. Histopathological changes in different regions of the stomich of Macropus fidiginosus associated with nematode
parasitism, 3, glandular epitheluin of the parietal diverticuluy showing dilated lumina (1), filled with eosinophilic secretion,
and massive infiltrations of lyimphoeytes and plas cells between vlinds; 4, squamous epithelium: (5) of the saccular
forestomach with minimal inflammatory change: 5. base of glands of cardite mucosa from tubular forestomach showing
diffuse infiltration of lamina propria with mononuclear cells (m, muscularis mucosae); 6, distal region of glands of cirdiae
mucosa showing diffuse infiltration of kimina propria wiih mononuelear velly, Seale bars: Pig. 3, lOaimy Pig. 4. 100 gan,
Figs 5.6. 50 pam.
192 D. PAMMENT, I. BEVERIDGE & R. B. GASSER
Cloacina spp. Labiostrongylus spp.
1200
1100
1000
900
wo
E —£ 800
5 o 700
3 = 600
; Oo 500
fe)
< = 400
300
200
100
6)
12 8 4 56 6 7 2B 1 2 3 4 5 6 7 8
Stomach section Stomach section
Popovastrongylus pearsoni Rugopharynx australis
1200 80000
ihe 70000
1000
” 900 ” 60000
£ =
= 800 c
fe) © 59000
= 700 =
6 800 s 40000
= one 30000
400
300 20000
aon 10000
100
6) 0
{ 2 BS & & 6 7 «8 jek oe 456. 7 +8
Stomach section Stomach section
Fig. 7. Distribution of genera of strongyloid nematodes in eight regions of the stomachs of ten Macropus fuliginosus from
Hattah, Victoria. (Bars represent standard errors of means.)
DISTRIBUTION OF NEMATODES IN KANGAROOS 193
Cloacina expansa
No. worms
12 3 4 5 6
Stomach section
Cloacina cf. elegans
No. worms
1 2 3 4 5 6
Stomach section
Cloacina cf. magnipapillata
240
220
200
180
160
140
120
100
80
No. worms
1 2 3 4 5 6
Stomach section
Cloacina hydriformis
1000
900
800
700
600
500
400
No. worms
12 3 4 5 6
Stomach section
Cloacina obtusa
No. worms
12 3 4 5 6
Stomach section
Cloacina sp.
30
No. worms
12 3 4 5 6
Stomach section
Fig. 8. Distribution of species of Cloacina in the eight divisions of the stomach of ten Macropus fuliginosus from Hattah,
Victoria. (Bars represent standard errors of means.)
Wh OD PAMMENT. |. BPVERIDNGE & ROB GASSER
Disvassin
The gastrit unutormy of pmacropodils Was reviewed
by Hume (82). The andtomy and listalagy of the
stomiucth af the eastern prey Kangaroo, M4, wfenters,
has heen desermbeal previusly (Schuler & Willian
876. Langer eral J980. Delluw & Hue 1982).
However thene 1s 10 comparable informatiun for the
elosely relaied spevies M. faliyinosus. In addiion,
While the bistologig and ultrastictaral features of
the stomach of (he kammar wallaby, Manreapas engent
have been described indetail (Gemmell & Enwelhardt
1977), dati for the castern grey kangaroo ure much
more litted. We examined patholoweal changes in
the mucosa assoviaed with nemalode parisitisny and
the ahstribulion of nemutodes in melationship to
cpilhehal cypes and therefore an exuniinaritin of both
(he gross aiid micruscape features of the stomach of
M, (uliminesns was necessary. In this study, (he gross
anatrny ofthe stumach of MM fiadiginesns did ae ditter
sienificantly from choral Mo ervariteds— Proportions
by wenghitof the three princupal stomach regians in Ad
filiginosns, sacculat forestomuch, (Ubular forestomuach
and hind stornach (22%, 74% and 4%), resembled that
ol M. givantewy Linsely (23% 70% and 7%). The
histological results fron this study provide uddigional
iMbrmation on gastic anatomy of grey kuagunwy. The
most significant difterence from earher works is the
recognition thal the mucosa of the proximal
diverticulum of Mo fulizimosus is distincily diferent
Histolaically from) that of the cardiue region and
consists of elongate purallel glunds producing an
eosinophilic secrenon. The same is true tor M,
ureeneus, although the histological differences have
mt teen reperted in the leeratupe and both stemuach
regions have been conmidercd. to have an identical
miueosal type (Hume 1982), a ennclusion which would
uppear lo be erponegus, Although anatomically
distinefive. the funelion of this isolated area of
glandular mucosa is unclear
Anatomical terms apphed fx the stomachs of
kugguroas ure net consistent (see Hume (982) The
syslenr ol nomenclature suggested hy Richardson (Yk)
as Conforming most vlosely to Nemine Anaroniia
literinerree iy suitable fur descriptive purposes, but is
not well suited to Minetional studies, The terminology
used by Hume (982) Un which (he stomieh 15 divided
inte Suceulae lorestomach, (ubulur forestommich and
Jundstomach ws more appropriate when dealing with
the distribution of nematodes anid al epithelial types
and has therelirre hewn utrlised im this study
Determining the positron of nernalades witht an
organ cleurly requires very rupid fiaurion or
imunemilisatiun of the nenuiides fnllawine the dearb
ol the host. With small hist species, Unis hers been
velieved using rapid freezing (Sehad 1965) Bush &
Hohiries 1986) or rinid dmmersion in Axatie (ohede
980), Wilh large hosts such as kangaroos. the technical
difficulties wre greater. The methods used in the past
have been ligation of the stomach jrnediately
fallowing the death of the host in the laboratory
(Smales & Mawson 1978b) or ligation following host
death tn the Meld (Dudzinski & Mykytuwyer Wo;
Arundel etd 1979), Dilfieulties oceur inaccurately:
identifying (inarotileal “haidmarks” and placing
Jigaiures on stomachs under feld conditions and
therelore |he approwh taken here was to capidly
immerse the entire stomach m i large volume of
locimaldehyde solution and carry out its subdivision io
the laboratory, That this did produce an wecurate
represenuion of the longitudinal disteibuuon of
nematodes was eGnfirmed by comparisun with data
from the stomachs of two kangaroos which were ligated
in the freld privet fixation.
Inspite of technical limitutions, cach of the nemidtode
genera exhibited a restricted localisation wathin (he
Stomach, Data have been presented (Fig. 7) as absolute
numbers of worms jneach stomach secuon rather than
number of worms per gram of stomach content
(density), The only genus for which the distribution
PACE Is alered in Compuring density rather than
absolute qumbers i Claacina because of the high
densily ol mernutodes in seetion | of the stamach.
Clowciia spp. were located primarily in the saccular
forestommuch with maximum densities in Section 1.
Lathlastroie vies spp. were algo present in the saccular
Jorestomach, bul maximum ninmbers occurred in
section 3 atthe anterior end ef the wbtlur forestamuch.
Both Kasopharwnn australis and Pepovastronyylus
pearyont occurred principally im the tubular
forestomach and all strongyloid nematodes were ubsent
from (he hindstomactr (sections 7-8). an area of the
Stomach characterised i macropodids by a low pH
(Dudzinski & Mykytowyez 1965; Smales and Mawson
W7#h), and Inhabited only by the trichostrongyloul
venus Pilarinenit.
The distribution of fenjdtodes within the stomach
OM. fuligidesus is similar (o that reported for other
kangaroo species. Syales de Mawson (1978b) found
that Cloecine spp. were most abundant in the saccular
forestomach of ML eugent while R. cestraliy was most
vbundant in the tubular forestomach. Likewise,
Arundel et al. (1979) tound that maximum numbers
oF R, australis occurred ip the tubular forestomach ol
Mo rufus. The distribution of Labiostrongvliw spp. was
restricted in M. fidlfginesus (mainly insection 3) when
compared with data fram other fests (Dudzinski &
Mykytowyez 1965. Sales & Mawson 1978b) but unis
may be mlated to parasite density, The number of
Lahiwsrremeylus spp in the present study was low
(mean = 120, fable 3), Dudzinshi & Mykylowyer
(1965) divided their specimens of M. ruin infected
with L. Jongispraduris into “heavily infected” (> 200
worms) und “lightly infected” (<200 worms) hosts.
DISTRIBU TION OF NEMATODES IN KANGAROOS 5
They Showed that the masinun ourober op,
loneispicularis always becurred in the tobwlar
fores(gmacl but that With increasing abandance, the
numbers of nematodes in the succuku forestamacch
rose, Hsintilair mechanisms operated in Mi filtsieseey.
then higher parasite densiues would be expected ty
result io at Significant jnerease in parasite numbers uy
seclinns 1 aml 2 of the stomach, 2 pearseni in our
study oecurred prectominanily ii the tubiforn
lorestomach (seertous 4 and 5) but in MM. waweniy, the
same species oceurs primary tt the suceiform
lorestomuett (Simales & Mawson 978) The reasons
for this difference are oot kogwer, but it may reflect
dufercoces in the anatomy and physiwlosy of the
stonmchs of the grey Kangaroos (WW, greanrens and M,
filivinosiny When compared with thar of AD eager
(sce Richardson 1980) Langer eral, 1980; Detlow
JYR2; Dellow & Hume 82) or may be a function oF
host associations. Po pearson ts a common ind
abundant parasite in Ad eygend (see Soles & Mawson
978A), [ts distribution in M. fieiginosiy is much more
restricted, bem Kiown only from Kangaroo Teland,
Soath Australia (Bevendee 986). Kepbrook, South
Australia (Wiesner, inpublished') aad Hata,
Victoria. Jt was not encountered ma survey ol the
parasites of 40M. filiginosus by Beverndge & Arunue!
(970), M, filreinosus pay be unsbnormal or arusiual
host lor (he parasite and (his may be reflected ma
differing distribution of the nematode io the stomachs
The distribution of congeneric’ parasite species was
Hot Investigated in (he cise of the wo species ol
Laliostrenuylas because of their relatively restricted.
distribution within the somach, Ot the several species
of Clave present. only three, © eypantay ©
Awderforniis and Cool, elegy oecurred at sufficiently
high densities to permit Comparisons. Tn euch) case.
there was no evidence of differing distributions, Eueh
parasite had ad maximum density inthe bhnd sae ol)
the saveulur lorestomach. One Cleaele species
deeurred in sections | te Bot the stomach, but in-very
low qumbers only, The dita therelore suggest thut there
is my or very te refional separauon within the
stomach when severa) congeneric specres are presecit,
The jack of dillerenees in the distribution of
Congeneric species Ol Cloaema contrasts with studies
on the oxyoroid nematodes of turtomes (Schad |963,
Petter 9601 in Which each nematode species was
Shown to have a restricted aiche, The results obtained
here with no apparent niche segregation. are consistent
with the hypotheses proposed by Rohude (980) for
mignogenean parusites ob fsb. im whieh resricted
' oD. Wiesner, KS Coripanson of helminth parasine
burdens in western grey kargsroos, Maeriupes filly inoxcey
(Destnarest, 1622) eraving matin), impyuved and depragded
postin, Revewteny Avneulrural College, unpubl isfed
WysPs
niches were absent but aperegations oF Conspecttic
parasites were assumed to oecur in Order Wy Tacilitace
reproduction (Rohde 1977), Conmipetition was nba
consmered to be a significant element in distabuttans
as niches were Super ubundant wid were nor saturated
In the specimens of M, filiginoxes exumoed, the
numbers of nermalodes present per kangaroo were low
when compared with previous studies (Beveridye d&
Arundel 1979: Wiesnet. unpublished!) in which total
Nematode numbers ranging From 300,000 ta 500,000
were encountered in some Kangaroos, These dite imply
that the numbers of niches availuble for most species
of easter nematodes in ML felginosus are lurge und
i this is the case, Competition would not be expected
fo be amor factoraffecting nematode distributions.
A tur larger sample of kangaroos should be examined
10 test for evidence of interactions between component
parasite taxa (Hoste & Cabaret 1992). However, littl:
or no evidence of competition has been found in
Stomacheinhabiting nematodes mn M, rufias, MM
viodateus or Perragale spp. (see Hoste & Beveridge
1994).
Histopatholovical changes in the gastric mucosa were
restricted to diffuse infiltraljans of monunuelear cells
inthe lamina propria between glands, Changes in the
squameus epithelium were oeglipible, The
monouuclear cellutar yatiuwations were diffuse ancl
there was mo apparent relationstup with fennatode
Wistribution, Cellular changes m the lanvina propria
were mosi marked ja the tubular forestommeh, and i
Was here thatthe gremest nombers of nematodes were
found, However, in the sacculut forestomach where
Cleacina spp. were dontinant. the inmurked changes J
the glandular epitheliun compared with the lagk of
change in the adjacent squamous epithelium suggested
that the type of epithelium presen significantly
influences the extent Ol pathological chames seen. Le
the tundus. a wea essentially devoid of nematodes,
no significant patholopical chunpes were found.
In the pylone antrum. tueal lestons detected were
caused by schizonts of a species of Eomeria Although
( muimber of species Ob this Germs are known to oecur
ut M. fulivinesus (see Barker-eral, 1989), the species
present in the pylorus could not be Wenttied, because
noly the oocysts exereted in lees Iave been described.
Iu ute study of the pathological changes induced in
the stomach of ML viganeus by nematodes, Arunded
etal, (990) commented on Microscopic lesions caused
by Labiestranevlus spp,. by Svoneyleides sp, and by
Rugapharyia roxemuriae. None of the lesjons they
reported was lound in MW. fidliginesuy at Halal,
Uthough ios evident thar the presence of C/oacine spp,
and A. uwastrafiy in the stemaech resulted inva diffuse
gastrins Similar io the anthimmiutory climes induced
by strongyloid aematodes in the hinge intestines: ol
equids (Barker & vain Dreumel |85),
We 1. PAMMEN'!
“The mean talensity of infection of Kungaroos wilh
gusiric nematodes wt Frattah (Table 3) (mean 87.200
nematodes) Was simu to that (93,200) reported (rev
40 free-ranging M. filiginasis collected in western
Vicloria and in South Australia by Beveridge &
Arundel (979). Given the high density of Kangaroos
ar Hattah at the ime of the collection of 78 per squure
kilometre (Morgan 1990), it was considered possible
that panisite abundance might alse be higher, as occurs
in popululions OF M. wiverieus (Arundel ef al 1990),
bul this was not the case. A moredetailed study of
the epidemioliey of pematude infeenuns in Ad.
fidizinosts In western Vietoria is. required before the
sigmificunee of We current data can be assessed.
Acknowledgements
We wish fo thank the Vietoriin Departmen vl
Conservation and Environment for (he oivitalion to
culleet kangaroos ul Huta and for assistance with the
collection. We are particularly indebted to B, Waltets
und G. Coulson tor their help.
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GROWTH OF THE SEAGRASS POSIDONIA SINUOSA
CAMBRIDGE ET KUO AT LOCATIONS NEAR TO, AND
REMOTE FROM, A POWER STATION THERMAL OUTFALL IN
NORTHERN SPENCER GULF, SOUTH AUSTRALIA
By R. C. AINSLIE, D. A. JOHNSTON & E. W. OF FLER*
Summary
Ainslie, R. C., Johnston, D. A., & Offler, E. W. (1994) Growth of the seagrass
Posidonia sinuosa Cambridge et Kuo at locations near to, and remote from, a power
station thermal outfall in northern Spencer Gulf, South Australia. Trans. R. Soc. S.
Aust. 118(3), 197-216, 30 November, 1994.
The growth of the seagrass Posidonia sinuosa was monitored in northern Spencer
Gulf, South Australia between late 1986 and 1990 at a site within the influence of the
thermal discharge from the Northern Power Station, and at a Gulf ambient site. P.
sinuosa growth was also monitored in Pt. Paterson, a large shallow bay surrounded by
extensive mudflats, immediately to the south of the power station, but beyond the
influence of the thermal plume.
Key Words: seagrass growth, water temperature, Spencer Gulf.
Transactions af the Royal Soewey ofS. dase (994), URES), 97206,
GROWTH OF THE SEAGRASS POSIDONIA SINUOSA CAMBRIDGE ET KUO
AT LOCATIONS NEAR TO, AND REMOTE FROM, A POWER STATION
THERMAL OUTFALL IN NORTHERN SPENCER GULF, SOUTH AUSTRALIA
by R, C. AINSLIE, D. A, Jotinston & E. W, OFFLER*
Summary
Aywscin, ROC. JOHNSTON, DD AW. & OFRUER. BW. 1194). Growth of the seagmiss Posydoniq sinuesa Cambridge
er Koo ut locations near to, and rene from. a power stilion thermal outfall in neérthera Spender Gulf. South
Australia. Tring Ro See, 5. esp, 118(3), 197-216, 30 November, 1994,
The prowih ol the seagmis Posed sintesa sis Wonitaced m qurthern Speacer Gulf, South Austethit, bebween
late 986 and 990 ata site within the infuence ot the thermal dtsehitrge pom the Northern Power Station, and
ata Gall anibicat site, 2 sinwese growth was also monitored in Pt Paterson, a large shallow bay sorrounded
by extensive mud flils, immediately to the south of the power stition, but beyond the influence of the therumal plume,
P sintwye theadoWs adjacent ti the power Stulton Show relatively minor reducuons ty growth characterrstics
flea! blude biomass, productivity, und Teal growth) compared to (hose jn Gulf ambient conditions, despite the
fact thal summer water lemperatures adjacent 16 the discharges ure consistently slightly higher than Gulf ambient
and have reached 28°C, a temperature comparable lo the highest feld temperatures previously recorded lor Pusrduna.
However. in the naturally warn waters of Port Paterson, where average summer witler (eniperurdves are marginally
higher than these within the influence of the thermal plume and Where Intermittent) pedk feaiperaiures exeeed
20°C, P sinmose has signtheantly reduced produvtiviry, skinding biomass. ble length and blade growth vate,
(ypicul of Fosdonia species in marginal envirgaments,
Despite the minimal effects of the current discharge on (he seagrasses, the evidence from Pt Paterson suggests
thal in northern Spencer Gulf where sulimer temperatures ure more typical oF i Subtropical thant a Lemperare
Wlarine environment. ~ sindose is near the upper limits of ity empenuure wlerunce during the hollest lime of
the year. Should there be Jocalised incredses in maxymum water (eraperatures in the Gult from future thermal
discharges (> a0°C), [here is the potential for more widespread Odeurrence OF stimlwd secyrnisses 4UCH ts Toute
inthe shallow Waters of TP Paterson, One passihle consequence vl this could be localised ingrease in the movement
OF Sediments on The sloping banks of rhe Gulf channel,
Kev Wokba: sedirass growth. water lenypeniure, Spencer Gull
Introduction
Spencer Gulf. South Australia, extends about 200
km inland to the arid mid north of Sourty Australia (Fig.
1). Northern Spencer Gulf ts defined as that purtion
of the Gulf north of 33°8. The waters of the northern
Gull are characterised by high summer salinities, 48
(Nines & Lennon 1986), and suinmer temperatures
in the mid to high 20s°C (Ainslie ev al. 1989),
Shepherd (1983) found that the subtidal benthic
communities were “impoverished interns of the overall
species richness” and concluded that this anuy be
juceative ofa stressed hypersaline environment, He
supeests that itis critical (hat the component party of
the biological system of (he northern Gulf “sliould
receive very detailed study to determine its capacity
ty receive additional stresses”,
* Environment & Technology Department. The Blectricity
Trust ot South Australia, PO Box 77, Adelaide's, Aust. 500]
Hains, I. R. (982) Submarine geology. sediment
trinsport, fydradynamies and quateniey history of nurtherh
Spencer Gulf, Sunth Australia. Semimer oa the Research
etd for Management of the South Australian Gulfs.
Austrailin Murine Svignees und Technologies Committee,
Adelaide, 10 November 1982, 45-52. Linpubl
The extensive seagrass meadows are an important
component ob the biological system of the gull asa
hubitat and nursery region. for their intrinsic
conservation value, and also for the cole they play in
stabilising sediments, particularly (he sloping banks
of the Galf channet!.
In 1985 a 90 Megawatt (MW) thermal power station.
Playford Power Station. was established on the easter
shore of northern Spencer Gulf, south of the city at
Port Augusta (Fig, 1: by the 1960s the capacity had
heen increased to 330 MW. The cooling water
discharge trom this developnient resulted in surface
water fmnperalures about 69C above auibient near the
power station. with only oecasional incursions of the
water 10 the intertidal regions LAinslie era/. 1989),
In the lave 1970s the deeisinn was made to proceed
with the development of a new power stanion of up ta
a possible 750 MW (Fig. Ll), with the ultimate potential
W discharge 4 million cubic metres of warmed seawaler
a day to the Gull, more than doubling (he volume of
water discharged al 6°C above Gull ambient (Ainstie
eral. |989), and with predictions of localised but
naticeable jnureases in temperatures of gull waters
198 R.C. AINSLIE, D. A. JOHNSTON & E. W. OFFLER
SITE 1, SEAGRASS DISTRIBUTION
ON THE SLOPE LW
PORT AUGUSTA Inshore Seaqrass
Verge
Deep Limit of _
Seayrass Fo
50 40 3 2 0 0
DISTANCE (m)
STIRLING NORTH
PLAYFORD
POWER STATION — MANGROVES
a“
Moe Cooling water inlet
DiScHagor SITE U6 wm vontucen
_- POWER STATION
COOL SITE 2
HIGH WATER
MARK
LOW WATER
MARK = —
Fig. |. Northern Spencer Gult South Australia, Seagrass monitoring sites: (1) power station thermal discharge (2) Gulf ambient
site remote from the discharge, and (3) Pt Paterson. Inset: seagrass distribution from low water mark to the Gulf channel
al site |
SEAGRASS GROWTH IS SPENCER GULE SOUTH AUSTRAT IA We
Linder some tidal conditions short term muaxiind were
predicted to exceed 30°C in the viguntty of the power
suutiets Ourfalls*.
The reported effects of thermal discharges on
scturasses Tange front large svale devastation resulting
root lemperature ingreases only a lew degrees
Centigrade ahove summer ambient (Thorhaug er ul.
(978), ty reduction in seagrass density (Robinson (987)
and changes in specific growth Charucteristes such as
leaf thickness and biomass (Vicente 1977)-
Circuitistantial evidenge suggests.that relatively sasall
changes in temperature of inshore waters may also have
indirect ctleets on seagrasses through increased
bucterml and fungal atck (Rasomssen 1977),
Onee seagrass meadows begin to detenorate,
emsion, sillaiton and reduction in water clarity may
lead ta a deeline in the seagrasses beyond the direct
influence ol the discharge (Shepherd 1986). Other
secondary effeets such as increased impacts of grazing
Organisms may also accelenite the decline of seag/asses
under siress (Shepherd er a/. J989).
Both Posidonia australis and Po sinrasea oevur in
nonhern Spencer Gulf bur the latter species ts dominant
in (he inshore meadows near the power stations.
This pauper desenbes the result of a monitoring
programme conducted ty assess the effects of the
increased thermal effluent front the first S00 MW
development of Northern Power Station on the growth
of the seagrass Posse, in the ght of predictions
of louilised water temperature extremes apprwching,
oreven exceeding. the upper known field tenyperatures
for this spegies”
Materials and Methods
Preliminiry field work was undertaken in the
9864987 sunnier season to establish sapling
tychoiques. The results of fhe seagriss monitoring
programme for the period [987-1989 are Teperted in
this paper
Water temperatures und growth of Posideiiva Siesa
wore measured at three sites i dortherg Spencer Gulf
(Pin 1). The Gull sites were established on tunsect
lines, the elevation and location of avhich were
confirmed relutive to an established Port Augusta
Power Station datum aS patt oF a contiguous study uF
the benthic intaung of the northern Gulf (Ainshe er
al. 1989). Site | is. within 200 m of the power station
and within the influence of the thermal plume of the
new Northern Power Slation: Site 2 is on the easteri
side of the Gulf, 3.5 km south of the power station
* Blevtricity Trust of South Austalia (983) Northern Power
Station Brvigoninentul Lpaet Stateniest. Auyiiat 185
Prepared by Kighill Stearns, Unpubl
and beyond the predicted influenge of tie discharuc
plume*. Site 3 is in Port Paterson. u large shallow
embayment, also beyond the predicted influence ot the
thermal plunie, The naturally high water temperatures
which have been recorded ir Port Paterson? can be
attributed to the inselation of the extensive surrounding
mudlMats.
2) im trupsects Were established wt each site about
5 m seaward of, and parallel to the inshore seaerness
verge. ull ata deprlt of abour 0.3-0.5 m below LWT)
(4 depth of up to 3.3 mm during the highest udes)-
Aveach site in sive water lemperatures were recorded
within the leaf blade cunopy with portable daha hyegers
in custom-built PVC waterproof housimrs. and fitted
with [5 k thermistor detectors.
Air temperatures were recorded at the South
Australian Bureau of Meieorntogy weather station
located on the Northern Power Stibon site,
Our object was lo estimate several iieusures of the
biomass and prowth rate of seagrass.
A wide range of sampling quadrat areus is chled tn
the literature for seagrass. studies, depending on the
leaf blade densify, und the need lo ensure that ier
adequine nuinber of teaf blades is tagged tir
measurement. Shepherd (983) used sub-samples ut
50 Jeaf blades for detailed measurements of Povidenia
austratiy leat blade lengths and widths in Spencer Gull.
More recently Pollard & Greenway (1993) used
simples of berween 20 and 60 shoots in a leuf marking
study of the productivity ol three species of seayrasysen
in the warny waters of Cayrns Harbour, Queensland
Tn this study 1S em> steel frayed quadrats were used
lo define measurement anus for each sampling site,
Pretuninary collections in the summers of [985 and
ke vielded information on the leaf blade nunibers
for the quadrat wired, Sixty tine quadrits, camlouily
placed along the (ransect lines, were siempled al all
locations: The mean number of leaf blades/quadran was
34.5 £17 Adecision was tnade to proceed with che
feld studies using live qumdnits per site (averaging 170
blades/site), The sampling area’site was 125 cm’,
compunible to that of a muriber of other productyy ily
studies of seagnisses with a similar growdr habit
(Ziemun 1974. Thorkauy ev al. 1978: Walker & Me
Comb (988).
Within the quadrats. all seagrass blades were ragged
and harvested using inetous oullined (im Zigmiua (1974).
Hurvesting was carded out at high ude, apprauimately
every six weeks depending on weather conditions and
underwater visibility.
Total blade lengths, incremental growth (length) and
dry biomass ol whole blades apd incremental growl
(productivity) were recone for euch harvest.
Numbers of pew shoots, Le shoops which grew
subsequent to the Hoval agate. and their growth and
dry biermass were also recorded. A simple “Shout
index” as derived Avhich 1s the olmber of new
200
shoots/m- (of seafloor area)/day expressed as a
percentage of the original number of leaves (agged/m
for each sumphing sile. Blade widths were also recorded
but were only used as a verification of the species
collections.
Dry biomass was determined by drying freshly
harvested blades to constant weight at 105°C. after
removal of epiphytes with g stainless steel scraper and
treatment in a 5% hydrochloric acid solution.
Results
Figure 2 presents a comparison of the Port Augusta
SUMMEF vir lemperiture and water temperatures at sites
| to 3 during the sunimer of 1987-1988 with w 500 MW
power Station operating.
In the shallow seagrass beds of northern Spencer
GuIF, air temperatures strongly influence the patterns
of variation in the water temperatures at all three sites.
In mid-summer average water temperatures al all
sites are between 20 and 25°C (Inset Fig.2). Short rer
fluctuations are most pronounced at site 3 in the
shallow bay of Pt Paterson, and least evident at (he
Gulf ambient site 2. Overall, summer water
temperatures are marginally higher at site | neur the
thermal discharge than at the Gulf ambient site, but
consistently highest at site 3-in Pt Paterson (Fig. 2).
WATER TEMPERATURE SITE 1
3a
25 |
za
14
10 —
JAN FEB MAR APR MAY
1988
JAN FEG
TEMPERATURE °C
JAN FEB MAR APR
MAR APR May
KC. AINSLIE. BD. A. JONNSTON & E,W OFFLER
Atall sites, despite the high summer temperatures,
seasonal temperature vanation is typical of temperate
Tegions with winter water temperature dropping below
53°C, even ut site | adjacent to the power stanon
outfall,
The summer relativity of the water temperatures
between the sites does not persist throughout the year.
During the cooler months, Gulf ambient temperatures
fall significantly below those of Port Paterson, which
in turn is cooler than site 1 adjacent to the power station
outalls.
Productivity (inean growth, dry weight/m*/d) ol
Posidonia sinuesa at sites | to 3, from November 1986.
ro August 1990, is shown on Fig. 3.
At the Gulf ambient site 2, and al sile | near the
power station cooling water outfall productivity shows
4 pronounced seasonal cycle with peaks between
October and April, Although the seasonal cycle is less
obvious tn the seagrasses of the nuturally-warmed site
in Port Paterson, highest productivity values at this site
were recorded in summer (Fig.3).
Productvily wis consistently highest at the Gull
ambient site 2 and least at site 3 in Port Paterson, with
highest recorded productivity being 3.7 + 1.1 g/m*/d,
3.1 + OS wime/d, and L6 + 0.3 g/m?/d at sites 2, 1,
and 3 respectively (Fig, 2),
AIR TEMPERATURE - PT AUGUSTA (DAILY MAXIMUM)
MAR APR MAY
“May
Fig. 2) Maximum daily air temperatures at the S.A. Bureau of Meteorology Station Port Augusta, and three hourly maximum
waler tomperatures in the seagrass canopy at sites 1, 2.and 9 Inset waler (emperatures, lines of best fit (polynemial
least squares regression) at sites | to 3.
SEAGRASS GROWTH IN SPENCER GULF, SOUTH AUSTRALIA 201
Productivity data for each site were compared, no _ the greatest difference being between sites 3 and | and
a priori attempt being made to discern seasonal “cut- between sites 3 and 2 (Fig. 3).
offs”, Variances were not homogeneous (Bartlett's test, Maximum mean standing biomass also occurs in
P<0.01), and analysis of variance was therefore not summer (Fig. 4). As with productivity, the seasonal
used. However Kruskal-Wallis analysis showed — cycle is most pronounced at the cool Gulf ambient site
significant differences between all three sites (P<0.01), 2, and least pronounced at site 3. The highest standing
Growth (Dry wat) g/m2/d
0 ;
Jan-87 Jun-87 Nov-87 Apr-88 Sep-88 Feb-89 Jul-89 Dec-89 May-90
—#— Site1 —°— Site2 ——*— Site3
Fig, 3, Seagrass productivity, leaf blades, g/m*/d (dry weight) at sites 1, 2 and 3.
400
300
200
100
Mean Dry Biomass (g/m2)
Fig. 4. Seasonal variation in mean standing biomass, leaf blades, g/m? (dry weight) at sites 1, 2 and 3.
42 RC, APNSLIL, B A. JOHNSTON & E,W. OPPLER
biomass was recorded at site 2 in November 1989,
352 g/m. The highest summer mean standing
biomass recorded at site | nearest the outfall was 231
gin. in November 1988, while the mean standing
biomass is consistently lower at site 3 in Port Paterson
than wt (he other [wo sites with a summer maximum
of OF g/ne (Pig. 4). in November 1988. Analysis of
variance using transformed data (log,,) lndicated
significant differences between the standing biomass
dala ol cach of the three transects (P<O0.01). An 4
Peserfory Mulople comparison test (Sokal & Rohlf
1969) indicated that although the differences between
siles 3 and 2 und 3 and | were significant, sites Land
2 were not significantly different.
Comparisons were made of total blade lengths and
blade growth rates for sites |, 2 and 3 after combining
the data collected lor these measurements. for January
IYSS8 and January 1989,
Mean blade lengths were 438.9 + 161.3 min, 316.0
+ 95.0 mm, and B58 + 50.3 mm at sites 2, 1 and
3 respectively (significantly different, ANOVA,
F = 367.6, F aang = 3,0).
Mean summer growth rates (incremental blade
length) were 4.7 % 2.8 min/d, 3.6 + 1.8 mm/d and
19+ Ll mm/d at-sites 2. | and 3 respectively, As
with blade lengths there were siynificant differences
between the sites (ANOVA, F = 254.0,
fF ayy > 3.0),
Wws*
ium"
Aosy, W. M, (1977) Marine Biological Studies in Relation
Jo the Operation uf the Torrens Island Power Station. MSc.
‘Thesis. Adelaide University, Unpubl.
4
3
x
ch
= *
B 2
°
o
=
n
1 \,
: *~™
0
Dec Mar Jun Sep Dec Mar
86 87 87 87 87 88 88
—
Site 1
—*—— Site2
Shoot production occurs all year round; the shoot
index is Highest at all sites during summer. Shoot
indices are similar at sites | and 2, while the shoot
index at site 3 is Consistently much higher (about 2
to 3 times as high) than at the other two sites (Fig. 5)
Although no measurements were taken, feld
observations suggest that the epiphytic growth was
consisteny higher at site | near the outfall than al the
eulf-ambient site 2, and higher again at site 3 in Pt
Paterson. This was particularly evident lor the serpulid
work) Euloypira convexis, a species which has
previously been demonstrated to thrive in the thermal
outtall at Torrens [sland Power Sttion in the Pt. River
estuary, South Australia’,
Discussion
A number of studies have indicuted the susceptibility
of seagrasses in subtropical regions tw water
temperature increases above ambient (Thorhaug ef el.
1978; GESAMP 1984). [In particular, there is evidence
that Many organisms, including seagrasses, are living
close 10 the upper limits of theie thermal tolerance
during the warniest part of the year, and even short
periods above {he summer maxima ean have signiticant
adverse impacts (Zieman 1974; GESAMP 984),
Larkum & Den Hartog (1989) hypothesise thal the
current distribution of species of the genus Posidenicr
in temperate regions, and the lack of evidence thatany
Posidonia species has ever occurred in wopicul
condilions inthe past. may indicate that species of this
genus have more limited temperature toleratice than
Species which occur in the tropics.
Jun Sep Dec Mar Jun Sep Dec Mar Jun
88 8& 89 89 89 89 90 90
° Site 3
Pig. 5, Seasonal variation in shoot index (number of new shoots/m7/d as a percentage of the original number of leal blades
per av ul ihe Cirne of tagging) av sites 1, 2 und 3,
SEAGRASS GROWTH IN SPENCER GULE. SOUTH AUSTRALIS
In northern Spencer Gull, although seasonal water
temperatures range fron as low as IC rm owinter fo
the Wud to high 20s. during Samimer (Auslie er al.
1989), the bigh summier water Lemperitures are more
wypival of subiropical conditions than those neuially
ussociied With temperate coastal waters of the soulher
Australian coastline. (Subtropical conditions are
defined as those where water temperatures generally
ringe from 20-25°C. atid do not exceed 30S!
IGESAMP 1984}]),
During vd summer When oir temperatures ninge
Wy 45°C. the waler temperature ui Me scugnasses at the
ourull (site 1) is consistently about 0.5 15°C higher
than at Gulfambient site 2 Griset Fig. 2). The difficulty
in define local ambient water temperatures is,
however, iHustrtted by the difference berween site 2
and site 3 onthe southern shores ol the large shallow
cribaymnent of Port Paterson. The latter site is also
beyond the influenve of the thermal plume, but exhibits
consistently High average suiimer temperatures Ginset
Pig. 2). up to 22C warmer thin Gull ambient (set
Fig, 2)
Comparison al summer leniperatiire regimes
belween sites as Complicated by the shor tern
occurrence of extreme temperatures, Short ler
XIN) Temperatures at site 7 (28°C on Two
ocuusions, Fig. 2) are ubout 1C higher than these
recorded at the Gulf ambient site, and as high vs the
maximum field temperatures recorded priar to tliys
study for Australian Posidonia species (Walker Ae
McComb (988). At site 3 which hay the most
pronounced diurnal vanations the Short term muexsuna
recorded ducing summer exceeded 30°C on a number
of oveasions. more than 3°C higher than the peiatk
temperature at the Gulf ambient site 2 (Fig. 2), and
the highest field temperatures at which Paytdene
species have been recorded.
Although seasonal date have been unavailable for
Posidonia sinvost, iitormation en the produeny ity ob
a noinber of diferent species Of searassen his been
reviewed by Walker & McComb (1988) und Hillman
ef etl (W989). These authors lourid that scaurass species
in Temperate waters in general tend to show a
Pronouneed seasonal varuiiion wih a distinct growing
season in summer However Walker & McComb cosa)
found no clear scasanal pattern in the productivity of
Posidomic auytratty in the seon-enclosed bays of Shark
Bay. Western Australia, over a one year period. The
water temperatures in Shark Bay ranged from about
18°C (o nrore (han 26°C) wad maxi lemperatures
up to about 28°C have been recorded uy this arcu
(Walker & McComb 1988), Despite the tuct that at ull
three sites an fhe present sttuly Sumter walter
temperatures resembled the semis tropical conditions
of Shark Bay, the P sinvesa growing al Gulfamibient
(site 2) and in the marginally warmer conditions
adjacent to the power station thermal discharge (site
TUG
1), shaw seasonal Muctuacon ie both producnvity and
standing bromass, With muximd lor bor measores te
the warmer simmer montis. Byer at site 3 with the
inost extreme conditions, summer ‘peaks are evialeut
in these features, although seasonal patterns are nat
w8 pronounced as atthe other Ivo sles, This persistance
of summer peaks in productivity and standing biomiss,
particularly acsites {and 2. supyessts thal the 2? yisteayer
of northern Speneer Gulf may be more tolerant ob
extreme sunimer conditions ian. lorexaniple. the #
auatealiy ot Shark Bay.
Atall sites there wus sonie varvation between the
valttes recurded from year to year lor summer peaks
of both productivity and stagditas biomass. ‘This
vatlariow was. however, ne more pronounced than
mhitural yeur lo year Variations recorded ina number
of recent studies Ol @ eanye of temperate Australian
seugfusses (Walker & McComb: 1988; Hillman wr al,
989). Although the growth ofthe seavrass Bo ydicecnyer
differs between the three sites, the interannual
comparisons indicate that. at any given site. the growth
iy comparable from year to year, There is ne evidence
of “deteriumlion” al site | adjacent ty the power slahion,
orat either of the twe control sites. during the course
of the current study.
Although pumerous studies hive documented the
range of effects (hat artiieiilly ioposcd temperaitire
fegunes May have on seagrasses. few have attempied
to deseribe the progressive effects on yrowth
chaructersiics whieh occur with incremental
lemperature mercusey above wpbient. “The exception
is that of Thorhauy ev af (Y78) who recorded detsthel
changes in prowth ef teal blades. productivity and
suinding, bionass (standing crop) tn trapical ane
subtropical Tiassa stands subjected 16 incremental
increases in wiler temperature above vubient,
Accepting thal diflerent species nay bave quite
different temperature Wlerances, it is nevertheless of
interest to examine the similarity berween TWreldssta
species wilh a subtropical distribution, ancl 2 ycereeaser
subjected to above-wmbient temperdiures un an
enuyiroomest where summer temperatures are already
high an terms of the geographic distribunon of this
Iefiperile species,
Thorhaog erat (M28) report that. with ime reases
wh Uide as 15°C above summer ambient water
lemperalure, growth per blade, productivity, and
standings bionss of Thalassia declined to 64%, GO%
am 82% vespectively, of thar ay ambient temperatures
und continued to dechne sharply with aeremental
increase inubove ambient remperature, At $°C above
utnment the seayruss “disappeared frooy the ares
(Thorhaty ee al. 178).
‘The Masini mean produenvity recorded tor &
Sinus ab site 1. with avejage summer water
lemperatures O.S-L5°C above Crutl ambyent and short
term peak lemperitures up to 1% higher than short
204 R. C, AINSLIE, D, A. JOHNSTON & E. W. OFFLER
term Gulf ambient peak temperatures, was 84% of that
recorded at site 2. A contributing factor to this lower
productivity is the lower rate of blade growth at site
1, mean growth per blade being only about 77% of
that of blades at site 2 during the warmest time of the
year. Despite the fact that over the period December
1988 to August 1990 standing biomass of P sinuosa
al site 1 was not statistically significantly lower than
at site 2, the maximum mean standing biomass
recorded during this period at site | was only about
70% of the maximum recorded during the same period
at site 2 (Fig, 4).
At site 3 in Port Paterson with average temperatures
consistently 2°C higher than Gulf ambient and peak
temperatures 3°C higher than ambient peaks,
productivity, standing biomass, total blade length, and
blade growth are all significantly lower than at the Gulf
ambient site (only 30-40% of Gulf ambient values for
maximum mean productivity (Fig. 3), maximun mean
standing biomass (Fig. 4), and mean summer blade
lengths and growth rates.
There is a lack of published information on P
sinuosa with which to compare the growth information
from the present study. Neverauskas (1988) examined
the effects of shading on a mixed P. sinuosa and P
angustifolia stand at a depth of 11-12 m, in an area
removed from any land based discharge. In winter, at
the commencement of the shading experiments
Neverauskas recorded a mean standing biomass of 100
g/m’. In this study the winter standing biomass of P
sinuosa at both sites | and 2 ranged from about 125
g/m to 175 g/m’, while the winter standing biomass
at site 3 in Pt Paterson was about 50 g/m’ (Fig.4).
Accepting that comparisons made between seagrasses
from different localities and depths can be misleading
(Ainslie 1989) these biomass values nevertheless
suggest that the seagrass stands at both site | and site
2 are al least as vigorous as in a location deliberately
chosen for its apparent “health”. The P. sinuosa at site
3 in Pt Paterson, on the other hand, is less vigorous
not only in comparison with the other sites in the
northern Gulf, but also in comparison to that examined
by Neverauskas (1988).
This paper does not rule out the possibility that, in
the higher water temperatures at site 1, and particularly
at site 3, indirect factors lead to the reductions in
seagrass growth, For example Neverauskas (1988) has
shown that shading of a mixed Posidonia sinuosa and
P. angustifolia stand with shade cloth (to simulate
epifaunal growth on the leaf blades) resulted in
significant decline in standing biomass (standing crop),
and leaf blade length. It is possible that indirect effects
from shading caused by relatively greater epiphytic
growth at site 1, and particularly site 3, may have
contributed to the reduced productivity, leaf blade
length and growth, and dry biomass at these sites
compared to site 2.
Neverauskas (1988) also found that with progressive
periods of shading, the shoot density declined.
suggesting a reduction in the appearance and growth
of new shoots, In the present study new shoot
production (with respect to leaf blade density) was very
similar at sites | and 2 throughout the year. Particularly
during the summer, however, new shoot production
as a proportion of the leaf blade population was 2 to
3 times higher at site 3 in Port Paterson than at the
other two sites. If shading by epiphytic growth is a
contributing factor to the observed reduction of
standing biomass and productivity in this study the
comparison to the results of Neverauskas (1988) does
not extend to new shoot production. However, while
in Neverauskas’s study whole shoots were shaded, in
this study the epiphytic growth shaded individual
leaves, with more shading of older leaf blades.
Although it seems likely that this shading by epiphytes
contributes to a reduction in standing biomass and
productivity, the consistently higher relative production
of new shoots at site 3 suggests a response similar to
that of terrestrial grasses, where cutting back to reduce
shading by old leaves can result in increased leaf
production, tillering and branching (Hendrick & Black
1986). The stunted growth of older leaf blades at site
3 may expose the new blades to more light, One result
of this higher shooting frequency in Port Paterson is
that. despite the stunted growth of the P. sinuosa, the
shoot density does not decline; there is no ongoing
deterioration of the stands as observed by Neverauskas
(1988) in artificial shading experiments.
Salinities at all three sites are high, as they are
throughout the northern Spencer Gulf, in summer
reaching about 48 (Nunes & Lennon 1986). Although
information on the salinity tolerance of P. sinuosa is
not well documented, P. australis survives in a wide
range of estuarine salinities up to 57 (Cambridge &
Kuo 1979). Tyerman et al, (1984) have shown that the
salinity tolerance of P. australis is, in part, due to the
physical shielding of the sheaths of older leaf blades
surrounding the bases of the youngest lamina. The leaf
sheaths of P. sinuosa are at least as robust as that those
of P. australis (Cambridge & Kuo 1979) suggesting that
the same mechanism for salinity tolerance operates in
this species. Although this paper does not rule out the
fact that a combination of factors (high temperature
and high salinity) may lead to the stunting of the
seagrasses in Port Paterson, the high water
temperatures would appear to be the key variable
between the sites,
From their studies of thermal effects on Thalassia
Thorhaug e7 al. (1978) suggest tentatively that increases
above ambient water temperatures of about 1.5°C may
be considered “rational” with respect to the Jimited
impact on seagrass and associated communities. In the
current study, the maximum summer temperatures at
site 1, near the thermal discharge, fall within this
SEAGRASS GROWTH IN SPENCER GULF, SOUTH AUSTRALIA
category, The results of this study also support the
contention that the overall change in P sinaosa growth
is relatively minor wilh this magnitude of ternperature
increase above ambient, less than the order of change
considered acceptable by Tharhaug er al. (1978) for
Thalassia. In Pt Paterson, in consistently higher above-
ambient summer water temperatures than recorded
near the thermal outfall, 2 sinvosa, although exhibiting
stunted growth characteristics of this species in
marginal environments (Cambridge 1974; Cambridge
& Kuo 1979), persists with no indication of ongoing
decline, in terms of biomass, productivity, leaf blade
growth and length, or areal extent of local distribution,
It has been proposed that the sediment stabilising
role of seagrasses may be particularly important in
maintaining the integrity of the sloping banks of the
chunnel of northern Spencer Gulf!. While the stunted
seagrasses on the relatively protected shares of Pt
Paterson survive without any evidence of ongoing
205
decline. if the seagrasses if the more exposed Gull
channel (lor example adjacent (o the thermal outfall)
were to be subjected to further, small summer water
leniperature increases, resultant stunted growth could
Jead to increased localised mobilisation of the shoreline
sediments. This. in turn. could lead to longer term
effects on the spatial distribution of seagrasses in this
area, Given this possibility, any considerauion. of future
development of the power station should recognise the
need to ensure that there is no increase in the maximum
temperature of the discharged cooling water,
Acknowledgments
This work was funded and supported by The
Electricity Trust of South Australia as part of its
ongoing programme of environmental research, We are
griteful to Dr Alan Buuler for reviewing the
THANUSCHIN
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TRANSACTIONS OF THE
ROYAL SOCIETY
OF SOUTH AUSTRALIA
INCORPORATED
VOL. 118, PART 4
POPULATION AND REPRODUCTIVE ECOLOGY OF THE
SMALL-MOUTHED HARDYHEAD ATHERINOSOMA
MICROSTOMA (GUNTHER) (PISCES: ATHERINIDAE)
ALONG A SALINITY GRADIENT IN THE COORONG,
SOUTH AUSTRALIA
By RoBYN L. MOLSHER*, MICHAEL C. GEDDES* & DAVID C. PATON*
Summary
Molsher, R. L., Geddes, M. C. & Paton, D. C. (1994) Population and reproductive
ecology of the small-mouthed hardyhead Atherinosoma microstoma (Giinther)
(Pisces: Atherinidae) along a salinity gradient in the Coorong, South Australia. Trans.
R. Soc. S. Aust. 118(4), 207-216, 30 November, 1994,
Atherinosoma microstoma in the Coorong exhibited a one-year life cycle with
multiple spawning over a four month breeding season from September to December.
Large numbers of larval and juvenile fish (5-15 mm long) appeared in samples during
October and November and grew rapidly over summer, most reaching lengths of 26-
35 mm by autumn and 36-45 mm by the following spring. Only one ovary developed
in females and this began to enlarge during August when batches of eggs began
maturing. Gonosomatic indices also began to increase at this time. Female fish with
spent ovaries were first caught in November. The numbers of large hardyheads (>35
mm) in samples declined in December and January, reflecting post-breeding
mortality.
Key Words: Atherinosoma microstoma, Coorong, hardyhead, salinity, reproduction,
population biology, fecundity.
Transactions af the Royal Sactety of 8. Aas (1994), W8(4), 207-216
POPULATION AND REPRODUCTIVE ECOLOGY OF THE SMALL-MOUTHED
HARDYHEAD ATHERINOSOMA MICROSTOMA (GUNTHER) (PISCES: ATHERINIDAE)
ALONG A SALINITY GRADIENT IN THE COORONG, SOUTH AUSTRALIA
by ROBYN L. MOLSHER® MICHAEL C, GEDpks* & Davip C, PATON*
Summary
Mrisuer, Ry L,, Genomes, M. C. & Patox, DC. (1994) Population and reproductive ecolugy of the small-
mouthed hardyhead Athertnivome microstema (Giinther) (Pisees: Atherinidae) along a salinity gradient in the
Coorong, South Australia. Trans. R. Soo. So Awe W8(4), 207216, 30 Novernber 1994,
Arherinaveme micrastoma inthe Courong vatibited a one-yeur life eyele with mulliple spawning over a four
month breeding season from Septemher to December. Large numbers of larval and juvenile lish (5-15 men long)
appeared. in sainples during October und Noyerber ind grew rapidly ayer sumone. most reaching lengthy oF
26-35 mm by auumn and 36-45 pin by the following spring. Only one ovary developed jn females and (his began
} enlarge during August when batches of eggs began maturing, Gonosomatic indices also began to meredse at
iis ime, Female fish with spent ovaries were tifst caught in November. The numbers ol karge handyheads (> 45, mun)
in samples declined in December and January, reflecting post-breeding mortality.
Saliniuies ranged from 9 Wy 67 g Lal Noonameena, the most northerly sampling ane and from 34 to 94 g
Lal Tea Tree Crossing in the southern end of the Coorong lagoon system. High salinities did aot have 4 pirked
effvecon the population ceology Or reproduenve prieatial of Atherinaspina mmierastome. Hardyheads were cauzht
wall sites on all sampling gecasions and ny marked differences were found in fish size, geoWwih mate, condition
Wp relative bute fecundity for fish caught fram different localities along the salinity gradient Hardyheads were
coping well with the high and fuetuaiting salinities thar exist in the Coerong, Significant differences. in batch
fecundity were found between years, wilh those in spring (990 being aboul hall those of 199], perfaps reflecting
diflerences in food availability, The possible eflects of future proposed reductions tn sulinity far the Coorong
onthe biology of 4. microstontar are discassed. Chahyes in salinity ure wolikely to lint ihe distriburinn of hardy heads
except thrangh possible wiluenves on their food supply,
Khy Worts: Arhéihoseiida diierastoma, Coorong, hardybead. salinity. reproduction. population hwology.
hecunidity,
Introduction
The Atherinidie (hardyheads) js.a Widespread family
of small fishes thar ure commonly found in calm,
shallow ‘waters aad ofien have short life cyeles (e.g.
Gon & Ben-Tuvia 1983: Prinee er uf (982), Prince and
Potier 1983; Potter ef a/. 1983, 1986). Atherinids are
offer. curyhaline but high salinity may affeet their
population and reproductive biology. For example,
suspected dwarfing in the Mediterranean atherinid,
Atherina boyert, may be associated with high sulinives
(Gon & Ben-Tuvis (983). The Narth American
alherinid. Atherinopy affinis affiniy, spawned in the
field at salinities of 72 p.p.t, but the young died within
four months (Carpelan 1955), Most fish species in the
Laguna Madre of Texas (up to 80 p.pt.) do nol spawn
it Salinities. greater than 45 p.p.t (Hedgpetlr 1967).
Generally, high salinities and Auctkations in salinity
restrict reproduction in many aqoatic animals, affecting
both the number af offspring produced and the tining
and length of ihe breeding season (Kine 64d),
* Department of Zoology, University of Adelaide, South
Australia SOS,
' ivantsan-, Vo (STR) Taxonomic and systematic review ot
the Australiin fish species of the family Atherinidie: with
references to reluted species of the olitwyorld, Unpublished
PhD Thesis, Macquane University,
The small-moulhed herdyhead, Aterinasome
microstoniy (Giinther). 's found jn abundance over it
wide range of salinities including estuaries, tnarine
enbay ments and hypermarine lagoons in south-eastern
Australia from the Tuggerah Lakes in New South
Wales, southwards and westwards lo Spencer Gulf in
South Australia (Lvantsoff 1978'; McDowall 1980; R.
Connolly & GK, Jones pers. comm, ), The maximum
sizes approach 90 fim (total length) in Tasmania and
80 mm in Victoria (Cadwallader & Backhouse 1983),
The life cvele of A. pricrestomer tus been studied in
Dee Why Layoon) N.SW_, where salinities ranged
from 3 to [oppt. (Patter ef al. 1986). In these
estuarme Condilinns, 4 yiitresroma exhibited a one
year hfe eycle with a foursmonth breeding season
during spring (August 6 November). Fry entered the
Lrappuble populalian in October, while larger adults
rapidly declined in abundance aller November; Growih
effectively ceased pver autunin and winter (Aprilia
Augiist)
A, omicrosiona has been found throughout the
Coorong where conditions range from estuarine in (he
nerth io hypersaline in the south (Geddes & Buble
1984), Although hardybeads have heen cauyhl an the
Coorong ip excess oF IDO pipet. PDS (total dissolved
solids), equivulent to a salinuy of YL pop... Geddes
()987) has suggested phat extremely high salinities
20 RL. MOLSHER, M. C, GEDDES & PD C. PATON
(>100 pp. TDS) during late summer and wutumn
May restnel the Southerly distribution of these fish in
(he Coorong in some years, This study takes advantage
of the longitudinal gradient of salinity: that exists in
the Coorong to study the effect of high salinity onthe
reproductive performance and population structure of
A, npierostomu-
Materials and Methods
The population structure and reproductive hinlogy
of hardyheads were studied at tive sites within the
Couotung: Noonumecna, Villa dei Yunipa, Policemans
Point, Salt Creek and Tea Tree Crossing (Fig. 1), These
sites spanned a 65 km length of the Coorong lagoon
system. with Noonameemt in the north usually
experiencing a lower range of salinities than the other
four sites that were further south (Geddes 1987). The
southernmost site, Tea Tree Crossing, was at the
= |
BOs Lake Alexandrina |
River a
Murray : aad Ty
Mouth = aN on
, q Wo
_ \
—__— \
Younghusband & \
Paninsula 2 “Fr n \
ane <
Albert 7
Nott Cageon “Sd MENINGIE
|
[— ae
South |
Australia |
|, Noonameesna
The Narrows —
2. Villa dal Yumpa
South Lagoon
3, Polleamans Point—
4. Salr Creak
‘5. Tea Tree Crossing
== Coorong Lagoon
Fig | The Codrone Lagoon showin the five sampling siles,
southern extremily of permanent water im the Coorong
where the highest sulinities in the Coorong had been
reported
The salinities and temperatures ol the water at each
of the sites were measured at monthly intervals between
September 1990 atid January 1992, Water temperatures
were recorded al a depth of 40 ent between 0900 h
and 1800 h. Water samples. collected from a depth-ol
40 em, were returned to the luboratory und sulimities
were estimated by measuring electrical conductiviries
(conductivity meter CDM3) and converting these
values [0 salinities using tables from Williams (986),
Samples with conduetvities over 100 mS env! were
dilutect and the calculated salinities multiplied hy the
dilution factor,
Each month. fish were caught fron) each site using
a5,5 m long, 1.75 m deep seine net with a mesh size
oF 1.9 mm. AIL seuiing was conducted ut a depth of
0.2-0.8 mowilh hauls over a distance of 40 m, On most
Oceasions a single haul was taken but when the sample
contained low numbers of fish (< 30). additional seines
were undertaken. Successive sejnés in the one urea
showed no siznificunt difference in (he mean length
of fish caught (t = 1,348. N — 200, P = 0.179), Fish
traps (63 by 36 hy 46 em, mesh sivé 1.9 mm) bailed
with meat (usually chicken) and set overnight in water
0.4-0.8 m deep, were used ia the first six months of
the study. Traps were expected to cateh larger Gish,
whose superior swimining ability may have cnabled
them to escape the seine net. und thus provided
information on the larger adults in the populution, They
also provided additional fish for assessing reproductive
condition. In addition, a plankton net (350 pm mesis.
60 em diameter) was hauled through the water (depth
0.2-0.6 m) for iwo minutes to check for the presence
of larval A, mivrostuma. Fish were immediutely
preserved im 10% bullered formalin and returned to
the laboratory where their length and reproductive
condition were measured.
The total Jength (TL-tip of sneut te end ot caudal
fin) of each fish from) both the seine und trap samples
was recorded to the nearest millimetre using dial
calipers. Where fhe number of individuals was large,
4 random subsample of approximately 150 individuals
was measured. The stundand length (SL-tip of snout
to posterior edge of the last lateral line scale) of 4
subsample of fish was measured to determine the
relationship bewween TL and SL, thereby wlowing
comparisons with other studies. Lengths and body
weights of hardyheads caught in Oetoher 1990,
December 1990 und March 1991 at cach of three sites
(Noonameena. Policermans Point and Tea Tree
Crossing) were medsured so that length weighr
regressiins vould be calenlated. Fish condition was
assessed by comparing these lenyth- weight regressions
hetween sites. All Jength and weight measurements
were Oblained tron fish that had been preserved i 10%
ECOLOGY OF ATMERINOSOMA MICKOSTOMA IN THE COORONG a)
buffered formalin, Sex ratios were calculuted for up
to 50 hardyheads (>30 mn length) caught in each
seine sumple from October 1990 to May 199]. The sex
of the fish was determined by macrosvopic examination
of the gonads. Gonads in fish <30 mm had not
differentiated clearly and so these fish could not be
sexed reliably. Orange coloration of the mid-lateral line
and eyes was observed in some fish and the sex of 95
fish that exhibited this coloration was determined,
Seasonal patterns in the reproductive eyele of
burdybeads were determined [rom changes in
gonusomatic indices (GSI) and the examination of
ooeyles in the ovaries of female fish, Male fish were
nol iivesuigated us preliminary examinations showed
no marked changes in gonadal weight, The
gonosomatic index (GSD was calculated by expressing
gonad weight as a percentage of body weight (De
Viamning eal, 1982). Gonad weight and body weight
were measured to the nearest milligram for up (yo 20
female fish (TL>40 mm) in each sample. Only female
fish that were at least 40 mov in length were used in
the analysis as the ovaries of smaller fish did) noe
contin maturing oogyles. Ovaries of female fish trom
three sites (Noonameens. Policernans Point and Tea
Tree Crossing) were excised and preserved in modified
Gilson’s fluid (Puckridge 1988"), These ovaries were
then teased apart. shaken vigorously, and stored for
tw weeks to separate the oocytes from the ovariin
tissuc. Preliminary microscopic examination revealed
three different cgy types, which parallels the situwnon
found for Menidia menidia (Conover O85). The three
veneral exe types were Classified by size (oueyte
diameter) and appearance us follows:
|. immature oveytes: 0.05-0.70 omni in diumeter, The
smaller poeytes in this group: (0.05-0.22 mm) had
a clear cytoplasm and large nucleus, while the lurger
oocytes (0.23-0.70 mm) were white. opaque und
olien irregular in shape.
maturing oocytes: 071-160 mm in diameter.
Spherical and dark yellow in colour.
3. ripe oocytes; |.61-2.50 mm in diameter. Spherical
and hydrated with a yellowish yolk centre.
Diameters of oocytes were determined from a Single
measurement on uw random orientation basis (West
1990) using uw microscope-video atltchment and at
digitiser pad downloaded to 4 computer, Subsequent
categorizations of ooeytes were based only on
appearance, Numbers of muturing and ripe ooeyles 10
ovaries from samples over the two breeding seasons
were coutted using a stereo-dissecting microscope,
immature oocytes were difficult to count and were only
counted for samples of fish caught between September
rh
* PUCKKIDGE, J, T. (1988) The hfe history of gizzard shad,
the bony bream, Nematalosa evebh (Glinther)
(Dernsomatinae, Teleosty) in the Lower River Murray,
Seuth Austraha; MSc Thesis, Unversity ob Adelaide.
Unpubl.
50
Immature fish
immature
n 40 oocytes
S
8
o 30
i)
a
2 20
E
=)
= 410
6)
0 05 10 15 20 25
immature
50 oocytes
Mature fish
% 40
S
3
© 30
Ko)
2
® 20
a
Ee
3
= 10 maturing
oocyles
0
0 05 10 15 20 2.5
50
Ripe fish
40
Ww
a ‘
= immature
2 30 oocytes
io)
P 20 maturing
2 oocytes
5
= i0
0
2.0 2.5
Oo O05 10 1.5
Oocyte diameter (mm)
Fip. 2. Frequency of oocyte diameters from the gonads of an
immature fish. a mature fish and a ripe lish.
210 ROL. MOLSHER. M,C GEDDES & C PATON
W90 and January 991. Fish were classitied inte tour
stages depending on the type of oocytes present (Fig, 2);
. immature fish: immature oocytes only, firm ovary.
mature fish: immature and maturing aneytes
ripe fish; immature, maturing and ripe gocyles.
_ spent fish; maturing or ripe oocytes absent, Maccid
OviLl'y.
“Batch fecundity” was defined as the nutnber of
maruring oMeyies in the ovary and presumably
represented (he maximum number of oocytes that could
be ripened and subsequently spawned af one line.
“Relative batch fecundity” was declined as the number
Ol THaluritys oocytes per araly of ovary-tree body weivhyt
(Conover 1985), Differences in the fecundity af female
lish at diferent sites in che Coorong were assessed by
compuring relative bateh lecundities.
whe
i
Results
Physical Characteristicy of the Coorong — Septertber
1990 ta January 1992
A longitudinal gradieot in salinity persisted ih (he
Coorong with salinities increasing front Noonameeni
in the North Lagoon to Tea Tree Crossing at the
southernmost end of the Coorong (Fig. 3). Hypersaline
conditions (35 2 1!) were maintained at all sites in
the South Lagoon, except briefly at Villa det Yurmpa
in Seprember 199]. At Noonameena, conditions were
estuarine during, winter and spring (9-32 2 14) and
generully hypermarine during summer, Seasonal trends
wete also evident atall sites with salinities high in
suromer, falling inauturnn and rising again in spring,
In Seprember 1990, salinities ranged from 32 2.1. ' al
uTea Tree Crossing
100 *Salt Creak
® Polisamans Point
+ Villa dei Yurnps.
ao it yee a
San | a" cf:
B 60 \~ \ jj i
iM a d ri
= \ a.
= BS
= 40-
a Wey a
5 ‘
wo ey a? \
- a
‘
ES Gaee Sme Seee See eet? Ce et 2 i rot Loa,
$ 30 i
& .
on 30 2=r ’ Pn. atid
e 4 | ed * 4
2 \o +
-— TR ett
SOND UFMAMSA JV ASOND Gl
1990 age 1892
Fig, 3. Seasonal Muctualions in salinity and temperature uf
five sites fete Coorong Lagoon, Salinities are show for
each sila and lempeniures dre shown da ire eu and mange
fan the five sites combined.
Noonameend to 57 gL! at Tea Tree Crossing, while
in Bebruary 1991 salinities had risen to 60 v E! and
944 L.' at these two sites respectively. Seasonal
vhunges in sulinity were similar in the Pwo years.
However, salinities reached their minima earlier in 199!
(August/September) and were lower than in (he
previous year Water temperature varied seasonally
wih maxinunt Water feniperatures if excess oF 24°C"
recorded during late summer and minimur
lemperalures OF 11°C in winter (Pig. 3),
Distribution of Fish iw the Coorons
Hanlyheads were collected from all sites throughout
the 17 month period. The highest salimity at which they
were found was 94p L! (Tea Tree Crossing,
February 1991), Five other species of fish were caught
commonly, yelloweye mullet (Aldrichella forsteri),
vongolli (Pyeudephritis urvilli), tiver wurtish
(Hyporhamphus regvlariy), greenback flounder
(Rhombosolea tapirina) aod blue spat goby
(Preudogobius oloruni), All species were caught iat
Noonameena and Villa dei Yumpa where salinities
were lower. However. yelluweye orullet was the only
species caught with 40 mlerostoma ut Tea ‘Tree
Crossing, and then only during winter and spring,
Yelloweye mulle(, congolli and blue spot goby were
caught at salinities up tw 64, 83 and 87 2° LL!
respectively. Flounder and gurfish were cuught only
at relatively low salinities (<36 g 1!) and only on
(hree occasions, 4. micreastame far ouinumbered other
fish species in eweh sarple:
Population Seucture ef Atherinosoma microstoma it
the Courony
The abundange of fish collected at the five sites on
(he 17 sampling Occasions was analysed by Iwo way
analysis of variance, The number of hardyheads caught
in the first seine for each sample was used in the
analysis. ‘There was no significant difference between
sites (F = O87, da, = 79, p>O0.05) but there was ou
highly siznificant difference between sample dales
(Fh = 6.03, d,f, = 79, p<0,001). Greater numbers al
hardyheads were caught during spring and summer
The length frequencies of fish in euch sample were
inspected and no consistent differences between sites
were noted so the sites were pooled for length
frequeney analysis, The sizes of hardyheads were
highly variable wl any one (ime in the year, bul there
were distinct seasonal patierns (Fig, 4). Larvae and
fry (5-15 min: length class bof Fig. 4) were prominent
in October, November and December (spring-carly
summer) of 199) and 199]. At this time of the year,
length-frequency distributions were often distinetly
bimodal, consisting of small fish (<25 Mun) and larger
fish (>35 mm), The 1990 cohort of smaller fish
showed o yritdual increase in length during summer
reaching lengths ef 276-35 min (length class 3) by Ma
ECOLOGY OF ATHERINOSOMA MICROSTOMA IN THE COORONG ai]
1991 and 36-45 mm (length class 4) by the following
spring (September 1991) at all sites (Fig. 4).
The length-weight relationships for fish collected
from Noonameena, Policemans Point and Tea Tree
Crossing are shown in Table 1. Statishcal comparisons
of the slapes by ANCOVA revealed non-significant
differences between the sites in December 1990 and
March 1991 (Tukey HSD, p>0.05). In October 1990),
significant differences were found between each of the
three siles (Tukey HSD, p<0.05), however the
ussummpuion of homogeneity of slopes was violated in
this month (F = 5.66, d.f. = 271, p = 0.004). The
slopes of the regressions, which indicate relative
M ver [Gare BM yarn [=e] 8) 5503 rr BO aby) foal
“4 as 45 45
“ x» w a
4) s 5 4
D Wah bree ' ae
B05 \ ney [ay * iran [ar a yay aa] wh [ne
asd a5 ds 45
ao & 10 w
E aa J 1 a
S koa! pom) Sah my ry ys [ze
i Ab 45 aa ah
a
wh Ww 3 ll M0
“ "i 4 Is
Se, RE eee eee
Fa) 774 ETI A ye Car] Bl 9151 60 pba
AS 45 bet 4s
Ay ™ \ Ai 26
is is 8 18
Cae OS
hQsyeg Saye wane wHayaged
E iene ded eeRaaae
Lennie 1h)
Fiy. 4 Relative léngth-class frequencies of Ailerinosome
microsioma cuught inseine net hauls trom Octoher 1990
unt] January 1992 (two samples [4 days apart-were laken
in Oetober 1991), Catohes from the five sample sites have
been pooled ay there were no consistent differences be(ween
sites in length-class frequencies, The number of fish
measured in each sample ts indicated at the top leff af each
distribution,
weights and thus may be interpreted as a “condition
factor’, showed no consistent relationship with the
salinity of the sites (Table 1).
The length frequencies of male and female fish in
the trap and Seine samples from all sites combined
taken in October 1990, December 1990, March 199]
and May 199] are shown in Fig. 5. Females were
significantly larger than males (independent t-tests.
P<(.05) in each month, The lurgest female fish caught
an the trap samples was 85 mm (November 1990) while
mules were below 67 mm (although a single male fish
of 75 mim was collected in May 1991). Sex ratios (fish
caught in the seine-net only) usually favoured females
with significantly more females for all months
combined (x7 = 37.82, N = 413, P<0.01). Orange
coloration of (he eyes and mid-lateral line was only
20 + Oci 90) 20
on=56 an=49
15 | eneise 15 §n=167
104 40 r
2 | |
EAM, JNU)
oOo " ¥ | 1
3 oJ aM Lo WINE
2 i EnnnaaAaie ep
o =
= 207 [Margi] 20 on=37
Fi Mar 91 9 n2o4 Ma 91
= on=24 -
@ 154 Pn=31 15
2
jo4 | 10
| "|
+ | i} nil I
of UMMM) TAM
1s ae A Pe A a mrt
39 =< atratoao ao asd yoo
mootronoore ans PF aRAoe
oawbonrow ow * Sranandm *
Onm7FrmMmMwoOw onrrTon Ooo
Lenath (mm)
CO Female OC Male
Fig. 5, Relative length-class frequeneies for female and male
Atherinoxoma microstiama in October 1990, December
1990, March 199! and May 1991. For each month fish fron
trap and seine-net samples were combjned.
TABLE 1. Length Weight regressions of Atherinosoma ticrastoma from Neonameena (NM), Palicemany Point (PP) ane ‘Ten
Tree Crossing (TIC) in October 1990, December 1990 and March 1991 ‘
Regressions are of the form ¥ = ax" wheres ¥ = weight, X = length, a = intercept and b = slope. - = coefficient af
dererminarion, a = santple size and ** — Pe <0.001
Sample date Site a b r n
October 1990 NM 9.12 « 10% 2,97 O.972*2% 69
PP 1.55 x 107 2,79 O,978##* 125
TIC 3.54 « 10" 3.12 ORO TF R3
December 1990 NM 4.36% 10° 3.12 O.975*** 100
PP 8.75 % 10% 3.04 0.98944 100
TIC 8.13 % 10% 2.96 0.966*** 100
March 1991 NM 6.03 % Lt 3.03 0.93q%% 100
Pp 8.32 x 10% 2.94 0.977#** 100
TRC 7.08 x 10° 2.0% 0.96 1? 97
212 R. L, MOLSHER. M. C GEDDES & D. C. PATON
found in mule fish over 36 mm TL und only during
the breeding season (SeptemberJanuary), Indicating
that iL is related to reproductive behaviour,
The relaionship between total length (TL) and
standard length (SL) was expressed by the following
equahon:
SL(mm) =0.61+0.85TLUmm)
(°=0,99, N=34, P<O001).
Annual Reproductive Cycle of Atherinosama
microstoma jm the Cooreng
Only one ovary developed in female A. microstama
and this remained small (<2% of body weight or
GSI<2) through autumn and winter in the Coorong
(Fig, 6), in spring, the ovaries showed a marked
increase in size before declining over summer with
similar seasonal patterns occurring at all sites (Fig. 6).
The GSIs were highest during September and October
in 1991, This annual cycle jn repreduction was alse
reflected in (he proportion of immature, mature, ripe
8 [ h 1
A !Noonameena | \
AYN ' —-+ cd \
44 / \ \
. —_- 43-77% * \
O-—1 ot tos a
B- ————— S {,
ba *\ [Villa der Yurnpa | } y
4 i : /
4 .
es =e 4
_ 0 —{———+ = =
c
=» 6
a Policemans Point| Pz
5 ;
bets tt 4 I a
a ra] ry
ca] ,
i — el »
3 my aa
8 /,
n A Sall Creek bX
i - = = 7 \
id } — \
Son) a .
0 i 7 Y oT
a! f — ,
|Tea Tree crossing] \
fe x ral
7 t ¢
i --p r Fo, '
o+—— + 17 Tr a a a |
AS ONDJIFMAMJI JASON Ded
1990 1991 1992
Fig. 6. Seasonal changes in the gonosomatic index of female
Atherinesema micrestome at five sites in the Coorung
Lagoon. Points represent mean values of Up to 20 fish and
vertical lmes are one slundard deviation euch side of the
mein, No standard deviation is given when sample size
was less thin 10.
and spent fish in the samples (Fig. 7). Female fish
possessed only immature oocytes trom May to July,
Maluring oocytes were first detected in August with
large numbers of muture and ripe fish present
throughout spring. Spent fish were first detected in
November. Some immature eggs remained in the ovary
indicating that nol all oocytes were matured and shed
Batch Fecundity of Atherinosoma microstoma it the
Coorong
The numbers of oocytes, classified as immature,
maturing or ripe inthe ovary of individual fish collected
from the two breeding seasons are shown in Fig. 8.
The large variation in numbers of immature oocytes
muy relate to fish size and lo losses due to maturation,
There were smaller numbers of maturing and ripe
vocytes with most fish having from 5 to 40 maturing
oocytes und 5 to 30 ripe eggs in the ovaries.
Batch fecundity (numbers of maturing oocytes) was
strongly correlated with ovary-free body weight
(r= 0,68, n = 101) in spring 199) at the time of peak
vonad weight (Fig. 9a). Relative batch fecundity
(numbers of inaturing eggs per gram of ovary free body
weight) for fish was independent of ovary free body
weighl (1° = 0.04, n = 101) (Fig. 9b), indicating that
100%
80%
o
ie)
x
Cc
o 60%
@
oO
i+)
=> 40%
5
®
ia
20%
0%
a + @ «ap DT I- it -L ¥
SES RRS SE SES
e2 °° SAg gO §
= < o *3
Month
Spent [| Mature
[-] Ripe [i Immature
Fig. 7 Relative proportion of female Atherinosome
microsiamm characterised by (he most advanced clutch
present in the ovary, Data from all five sites combined,
ECOLOGY OF A4THERINOSOMA MICROSTOMA IN THE COORONG 213
fish of different sizes were allocating a similar
proportion of their resources to egg maturation,
Relative batch fecundities (Fig. 10) were not
significantly different between sites in spring 1990
(F = 0.16, df. = 2,45, P = 0.851) or 199] (F = 3,23,
d.t. = 2.98, P>0.05). However, significant differences
were found between years. with relative batch
fecundiuies in 1990 being about half those of 1991
(F = 51.67, d.f. = 1147, P = 0.001). The number of
maturing eggs per ovary tended to decline after the peak
spawning period in October 1991,
Discussion
Influence of Salinity on Distribution, Growth, Size and
Condition of Atherinosoma microstoma
Over the summers of 1990-1991 and 1991-1992 the
distribution of Atherinesama microstoma in the
Coorong was not restricted by salinity. All other lish
species appeared to be limited by salinity during the
high-salinity summer season in a manner similar to
that reported in Geddes (1987) and Geddes & Hall
(1990), The highest salinity at which A. micraytoma
was collected was 94 ¢ L!, This record exceeds the
maximum field salinity for this species from coastal
salt ponds in Victoria (82 p.p.t., Chessman & Williams
1974), In March 1985, a few individuals of 4.
microstoma were trapped at 149 p,p.t, TDS (salinity
approximately 130 g L') at Tea Tree Crossing in the
Coorong (Geddes 1987; D. C. Paton unpublished). A.
microstoma ranks along with Cyprinedon variegatus.
| ripe cocyies — |
hon
Dt 7
a
Number ol fish
= re
o.6°86
dl Tool. a
o- a
10 4 | immature oocytes
gt Sd ' ) Lr ell eteone 9 2
— —
Q 100 200 300 >400
Number of oocytes
Fig. 8. Frequency distributions of the numbers of immature,
maturing and ripe oocytes in the ovaries of female
Atherinasoma microstoma caught in the Coorong during
the (990 and 199] breeding seasons.
which has been recorded at 142 p.p.t. TDS (Simpson
& Gunter 1956), as one of the jest salt-tolerant fish
species in the world. This ability allows A. micraytome
lo survive at (he southern end of the Coorong during
extreme hypersaline conditions. Abundance dita
suggested that hardyhead numbers ut the southern sites
Batch fecundity
. Qa a
Oo Oo Cc
Relative batch fecundity
~
=)
0 1 2 3 4
Ovary free weight (g)
Fig. 9, Relationships between (a) batch fecundity and ovary
free body weight (rm = 0,68. n = 101) and (b) relative
batch fecundity and ovary free body weight (r- = 0.04,
n=10L) for female Arherinasema microstoma, Points
represent fish collected in September, October and
November 1941.
ld RL. MOLSHER. M.-C. GEDDES & D C PATON
were not affected. but high variability, perhaps
associated with schooling behaviour of the fish, and
low numbers of samples were confounding factors in
this analysis.
No mujor dillerences were found between sites in
growth, size or condition of hardyheads in the
Coorong, Growth rates implied from the progression
Of length modes suggest that.4. microstome can achieve
a total length of 35 mo in four months, followed. by
a period of litte growth over winter, reaching lengths
of 45 om the tollowing spring, und thereby becoming
stkually mature within the first-year of life. There was
ny indication of dwarfism related to high salinity in
this population.. Maximum sizes for both sexes were
similar at all sites in (he Coorong and comparable to
those found in the Dee Why Lagoon population (Potter
etal, 1986) and tor (his species in general (Cadwallader
& Backhouse 1983). Maximum total lengths recorded
for males were 67 mm (present study) and 66,5 mm
(Dee Why Lagoon) and for females 85 mim (present
study) and 86.4 mun (Dee Why Lagoon), Fish
condition in the Coorong did pot differ significantly
between sites in March when salinities were high and
tahged from 58 at Noonameena to 92 ¢ 1! at Tea
Tree Crossing,
Life History, Annual Cycle and Population Structure
of Atherinosoma microstoma
‘The post-breeding decline in larger fish at all sites
Irom December indicates a one-year lite cyele. The
population showed a numerical dominance of females,
und mules were also significantly smaller than females.
Similar patterns of life history aud sexual dimorphism
* Noonamesena
* Policemans Point
U Tea Tree Crossing
50-
|
Dr 0
2
2m
8 2 40
BS ”
oo
2a
aw © 20 a
BS “wi
ca 7 q hs oe
a= ~
= Ww ~
“a
L
0 ] r 1
Oct Nov Dec Jan
1990
have been found in the population at Dee Why Lagoon
New South Wales (Potter ef a/. J986) and in atherinids
from the Northern Hemisphere (e.g. Gon & Ben-Tuvii
1983). The proportion of female fish in this population
appears to decline over the breeding season und a sex
ratio of close to one is reached in December. This may
be duc to differential mortality during the breeding
season, of fo an increase in the numbers of male fish
reaching maturity (> 30 mm) as the season progressed,
The orange coloration found in male fish appears to
be related to reproductive activity and may be under
hormonal control.
The marked difference in size between sexes, the
prolonged breeding season and the short life eyele of
A, micrastama may be indicative ofa species where
sex is determined after conception by environmental
factors, such as temperature (Conover 1984), Conover
& Kynard (1981) conclusively showed that temperature
exerted a direct influence on primary sex
differentiation, rather than causing sex specific
mortality in the atherinid, Menidia menidia. Thus, il
exgs spawned in the cooler spring: waters develop into
females, and eggs spawned in the warmer waters of
late summer develop inta males, then females would
have a longer growing season which would account
for their significantly larger size.
Spawning occurred [rom September to December
siving a protracted four month spawning season. This
18 supported by the large numbers of larvae collected
from October to December from all sites, The absence
of larvae in the samples, the small GSTs, and the
absence of maturing and ripe oocytes in the ovaries
during aulummn and winter J991, indicates that this
Jan
1992
Dec
Nov
Aug Sept
1991
Fiz 10. Moan relative batch fecundity of female Arhkerinosome mrerosiome from ures sites in the Coorong Lagoon over
the 990-9) and 199-92 breeding seasons, Points represent mean values of up to 20 fish and vertical lines are one slandard
deViatinn cach side of the mean, No standard deviation is viven when the sample size was Jess than (0, Only fish with
niluonge or ripe oveyles were included in rhe analysis.
ECOLUGY OF ATWERINOSOMA MICKOSTOMA [TT COORONG 2S
species has only one breeding season per year. Mean
ponosomune indivey and relitive batch fecundities were
sreatest in November in 90 and October in 1991.
indiculing (hat spawning probably peaked during these
months. The later onset of spawning if 1990 js rerlected
by the Jater drop in salinities in (hat year Thus i
reduction ia salinity may be one of che envinanmenial
variables involved in triggering spawning.
A protricted spawning season may reflect repewtca
“batches” of egas being spawned or a Tack ol popalanan
synchrony in gonadal development (DeViaming 1983),
Analysis of eee types shows that docyte developinent
in A, iicrayiome is “eroupesynchranaus’, Ws att lowst
two size groups of oneyles were present 10 the Wary
Of an individual at some toe during the reproduc ve
eydle (DeViaming 1983). Thin type af onevie
development iiiplies that the whole clutch of ripe
oocytes will be shed over a short period (West (98).
Mulrinwodul frequeney distributions of oneyle
diameters, as found (ord. gareresteone in the Coorong.
are Charneterisuie of multiple-spawning fishes (Henipe!
1979). The presence of several nodes of developing
upeyles. the presence of remnant ripe eggs in-some
ovaries and the decline in the number of maturing eges
per fish us the Spawning season. progressed indicates
thal A. yiferastoita is a multiplespawriing fish
Multiple bulches of egus spawned salccessively Within
une spawoing period huve been ound in other
athermids including Menai menitdia (Conover |954)
and M> dudens (Hubbs 1976). Protracted spawniag ts
common aioli athicrinids.in Australia (e.g, Ivantsoll
78; Prince & Pouer |983; Potter eral 1986) sunil
in the Northern Hemisphere (e.g. Curpetun 1955, Gon
& Ben-Tuviu 83: Conover M984: Midduugh &
Hemmer 1992). A four month protracted breeding
season over spring parallels that found far this speeres
from. Dee Why Lagoon, New Soith Wales, svhele
spawning ovcurred from August ter November (Poller
eral \986),
Influence of Salininy on the Reproductive Peectormianee
of Athermosoma microstome
4. nacrestomea is ihe only recorded Australian
alherinid. to reproduce in hypersaline waters. with the
possible exception of Crateravephidusy peice tradraiys
(Lenanton 1977). The gradient in saliniies along the
Coomng did nol reduce (he length of the spavalig
scuson in 4. miicrostonia as fish were in spawning
condition from Seprember to December in bolh 1Y9o
and 199) ut ull sites. Salinides during these perivds
ranged from 32 .¢ 1! ut Noonaneent in September
1990 to 74g Lt at Tea Tree Crossing in Deveinber
1990, Salinilies are usually at their lowest i the
Coorong during spring which may account for the lack
ol uny cleat influence nf salinity on reproduechon in
the present study.
Gomad development commences in tile winler/eatly
spring at 4 titne when day leegrh and temperature are
increasing and these ire likely w be the environmental
vues for gonad recrudescence. Breeding in spring muy
be pimed to take sudvantage uf seasonal peaks in food
availibility. ta the Corny, burdybeuds feed mainly
on zooplankton, in particular astrucods und copepods.
dnd these ure most abundant during winter and spring
in the Coorone when sulinies are pelitively low
(Geddes 87). Female hurdyheads only develop a
portion of their eggs alone (ime. suggesting thaleither
u physiulogical or ecological factor (e.g, insufficient
fond) limits huteh fecundity, Relalive batch fecundities
and gonosamatic indices were lower in 1990 when the
growth and performance of Ruppia tnberase,a key
aquulic plant in the suuther Coorong, was reducer
compared with I991, Other resources including
aooplankton amay also have been lower Mulliple
Spawning is 4 cunMion reproductive strategy aniong
small fish species und (aaiinises the jumbers oF eggs
aosmall fish ean produce (Potter pers, comin.) In
adlitinn. 4. scrasxema may be uble to adjust batch
sive TH response to environmental cunditions. Such a
strategy would allow hardyheads to exploit both longer
reproductive seasons und better quality seasons. This
stmategy would require the inifalion of more oocytes
than would be expected to be shed in ninst veurs.
The Role of Stherinosoma micrastoma iH the Cauniny
Atherinids ave an dnperiant component of the
biomass o) small fish of mahy aquatic ccasvstemts,
inchiding Bandawil Lagoon (Ben-Tuvas 1984), Laguna
Madre (Hedypetl 1967). Hamelin Pool (Lenanton
1977). Peel Harvey estuary (Purtee ered. 1983) and the
Swan-Ayou River syste (Prince & Patter 1983). A.
inicrasiome is a prominent component of (he Coorong
eeosystent. ii particular as fo major laod tem for
gelected pisciveraus water birds (Paton 1982). High
and fluctuating: Saliniies along rhe Coorong did not
haye a marked effect on the population ecology or
reproductive hehayiour of AL ynierestama and no
significunt differences im relative batch fecundities were
found between sites that diftered jo saliniry within p
year, Thus, 4. micréstvina appears Wy be Well adapted
16 livpersaling conditions und luture smull changes 109
sulinily Would pot be expected 8 arfeet hardvheads in
the Coorong. However, their toou supply miy be
uffeeted by salinity. Future studies should be directes!
toward the diet ol A. viferosmme i the Coorone ane
the factors that mipht limit these resources,
The ugriculluril areas of the Upper South Bust of
Sout Austialia suffer from drevland salinization and
seasonal mundation of lowlying areas with surplus
surface Water, There ire proposals to drvin some af
thiy Surface water wad possibly groundwater into the
Soult) Lagoon of the Coorong (Upper South bast
Dryland Sabmity und Flood Management Plan Swering
“6 R. L, MOLSHER, M. C_GRDDES & D.C. PATON
Committee, (993*). Such inputs of fresh or brackish
witer may discupt the seasonal Huetuations in salinity
whieh are an important feature of the Coorone and
which may act as a partial cue lo spawning in 4
microstome. Other consequences of whe inputs, such
as an inerease in Water depth and consequent reduction
in the exposure of the highly productive mud tats. or
a fise in nutrient Or heavy metal concentrations may
also aflect the food chains in the Coorong supporting
4. micrestoma and in-turn piseivorous birds,
Considerahon needs ro be given to the role of
hurdyheads in the Coorong ecosystem in Hihire
management proposals.
Acknowledyments
We thunk Damon Oliver, Colin Molsher, Alex
Weissmann dnd David Peake-Jones for field assistatice
and Dr Keith Jones tor helpful comments on the
manuscript. This research was supported by @ zrant
foy the Murray Darling Basin Commission, The
Department of Zoology in the University of Adelaide
provided laboratory space and logistical support.
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REDESCRIPTION AND ECOLOGICAL NOTES ON THE
PYGMY BLUETONGUE, TILIQUA ADELAIDENSIS
(SQUAMATA: SCINCIDAE)
By MARK N. HUTCHINSON*, TIM MILNEF & T1M CROFT#
Summary
Hutchinson, M. N., Milne, T. & Croft, T. (1994) Redescription and ecological notes
on the pygmy bluetongue, Tiliqua adelaidensis (Squamata: Scincidae). Trans. R. Soc.
S. Aust. 118(4), 217-226, 30 November, 1994.
A preliminary study of a population of the pygmy bluetongue, Tiliqua adelaidensis
(Peters, 1863) has enabled us to redescribe the species and give a preliminary account
of its natural history. Life colouring, intrapopulation variation, sexual dimorphism and
general morphology of the skull and mandible are described. Pygmy bluetongues at
the study site are diurnal inhabitants of open tussock grassland and use spider holes
for shelter. Males had enlarged turgid testes during spring and a female examined at
this time had yolked ovarian follicles. Males were more active and trappable than
females during spring, but both sexes were sedentary during late summer-autumn.
Litters of 1-4 live young were born in the maternal burrow during February-March.
We suggest that the lack of sightings of T. adelaidensis this century has been due
partly to its specialised ecology and partly to a real decline attributable to habitat
destruction,
Key Words: Tiliqua, Scincidae, lizards, morphology, natural history, conservation.
Transavtions of the Royal Society of 8. Aust, (994), VIB(4), 247-226
REDESCRIPTION AND ECOLOGICAL NOTES ON THE PYGMY BLUETONGUE,
TILIQUA ADELAIDENSIS (SQUAMATA: SCINCIDAE)
hy MARK N. HUTCHINSON* Tim MILNET & TIM CROFTE
Summary
Hutcuinsow, M,N... Mirwe, T..& Croer, T. (1994) Redescription and ecological notes on the pygmy bluetongue,
Tiliqua adelaidensis (Squamats; Scincidae). Trans. R, Sov, 8, Aust. W8(4), 217-226, 30 November, 994
A preliminary study of a population of the pygmy bluetongue, Tiliqua adcluidensis (Pelers, 1863) has enabled
us ti redeseribe the species and give a preliminary account of ils natural history, Lite colouring, intrapopulation
syriation, sexual dimorphism and general morphology of the skull and mandible are deseribed. Pygmy bluetongues
al (he study site are diurnal inhabitants of open tussock grassland and use spider holes tor shelter, Males bad
enlarged wurgid testes during spring und a female examined at this time had -yolked ovarian follicles, Males were
more active and trappuble than feniales during spring, but both sexes were sedentary during late summer-autumn,
Litters of 1-4 live young were born in the maternal burrow during February-March. We suggest that the lack
of sightings of 7! adelaidensis this century has heen due partly to ite specialised ecology and partly to a real
decline aunbutable io habitat destruction,
Key Wokts; Jiligua, Scmeidac, lizards, morphology, natural history. conservation.
Introduction
“kommt nur auf sandigem, steinigem terrain vor"
(Richard Schomburgk. quoted by Peters, 1863)
Schomburgk’s remark (“found only tn sandy, stosiy
terrain’) is the only published first-hand information
available on the ecology of Tiliqua adelaidensiy, he
pyemy bluetongue lizard, a species which has been
regarded as one of the most senously endangered of
Australia’s reptile species. if not actually extinct
(Cogger 1992; Ehmunn 1992; Hutchinson 1992). The
discovery of a population of the species near Burra,
S.A. (Armstrong & Reid 1993; Armstrong ee.a/. 1993),
following the first sighting of the species for 33 years,
has presented an opportunity for urgently needed study
of the species which had previously been known from
only 20 museum specimens, mostly collected last
cenlury (Ehmann 1982; Shea 1992),
The original description (Peters 1863) was brief,
based on two syntypes probably collected in the vicinity:
of Gawler, SA. Mitchell (1950) redescribed the species
based on SA Museum specimens and figured the head
shields and whole animal for the first time, No further
formal descriptions have appeared jn print, save for
those of Cogger (e.g. 1975, 1992), bused on the old
* Mark Hutchinson, South Australian Muscum, PO Box 239.
Adeliride, SA, SOOL,
+ View Milne, School of Biological Sciences, Flinders
Iiniversity, GPO Box 2K), Adelaide, SA, 500).
| Tim Ceott, Biological Conservation Branch, Department
of Environment and Natural Resources. GPO Bos 007,
Adelaide. SA. SOL,
' Sima, G. M, (1992) The systematics and repraduetian vt
blueangue lizards of the genus Tilia (Squurmats:
Scincidae), Unpub. PhD thesis, Dept ot Veterinary
Anaony, University of Sydacy,
and faded museum material. Shea (1990) described a
number of sealation and osieological character states
lor T. adeluidensis in order to establish the validity
of the penera Tiliqua and Cycledcmarphys. Shea's
unpublished thesis (1992!) gives a thorough
description of the Species’ scalation, morphometrics,
colow pattern and osteology based on the twenty
specimens then extant. Shea & Hutchinson (1992)
illustrated and commented on the dentary and dentition
of T adelardensis.
Ehmann (1982) summarised available data on diet
und external morphology and Jrew together the meagre
dala bearing on the provenance of the specimens known
to him. He also attempted to reconstruct the species’
ecology, using analogies with related or physically
similar reptile species. These speculations Were cited
several Umes subsequently. sometimes in such a way
that it was not clear that there were no direct
observational data on the subject (Ehmann 1992).
Field work has begun, ainied at determining the best
methods for locating To udelaidensiy, gathering
preliminary dats on its ecology and making a first
attempt to determine the number and size of surviving
populations. This article gives # sumumary of the
morphological variation that we have observed within
wsingle populition of 7) adelaiderisis and provides a
preliminary account of its natural history al this site.
Methods
Current research on the bivlogy and conservation
of this species is concentrated on one site located in
the Burra area (33°41'S, 138956"). approximately
160 km north of Adelaide, South Australia. The site
was the first of several found to support 7. adelaidensis,
following the initiation ol fieldwork in October, 982
NK
(Armstrong era! (993). The data presented here stem
from the first season of feld work on this population,
curried out from mid-October 1992 io mikMay 1993.
Specimen Collectinn
Live Po adelaidensiy were volleeted by three
ritethous.
Pitfall traps. 30 metre stretches of fly Wine deity
liencing Were erected, along which were placed sia
25 cm dengths of PYC pipe dug vertically iato the
ground. Four iraplines were setatihe swdy sie durin
the period 4 November-20 Devember, (892, An
wditional line was in place during 30 Novernber-20
Decemberund a further Lwo were set on 4 December.
The five most successful spring traplines were reset
during 2-2) February 1993, ‘Total trap nights by this
method were 1,956,
The other two methods relicd on the species tole-
dwelling hubils (see Habit seetion)
Hand collection, This method relied on opportunistic
mghtings of free ranging, animals. or on surprising
animals at the ontrances to burrows, Most individuals
cuught by hand were juveniles which were extracted
from burrow entrances Usiiy long forceps or clamps,
We also ised a method we termed “fishing” enticing
Iyeards Wy seize aun insect bail lied Lo a cottin theead.
A grasshopperion.a picee of cotton ved tu the end al
a three metre banibou pole was held outside the
entrance of the Tizard’s burrow; lizards seizing the
arasstiopper could be pulled inty the air and caprired
ws Hey dropped to the ground and teied tu return te
the burrow (Strong er af, 1993 desembe a sinilur
eallection leehmque). Lizards had tw be hired
completely out of the burrow before being allowed to
seize the grasshopper since the lizard’s hind limbs
could hold the burrow rim. preventing its dislodgment,
Their exireme wariness made the use ol a long pole
necessary to distance the collector from the lizard.
Trapping using baited (peanut butter or sardines)
aluminiuin “Sherman” traps was tried but without
success (approximately 2,500 (rap nightsy.
The time of day of collection Was noted, as Was the
datly maxim temperature, Lizards were measured
und sex was delermined by the presence of bulyes al
(he base of the jail and/or lempenes an mules, with
a lack of these foatures indicating a ternale The lizards
Were then marked by toe clipping, photographed «apa
releused. Only one or two wes were remeved, this
lined number ar conjvoetion with colour patiern
variations being sufficeent to dently each individual,
Limited ase Was made Of radiotrackiny using a Small
extertial unit supplied by Transceiver Services.
Adelaide, atiaehed using supergtue and surgical tape
ww the lizard’s shoulder The device wax intended for
shor term tse, and wits cun successfully on a single
lizard, located every rwo hours, for six days. A second
MON HUTCHINSON.
1
MILNE & T. CkOFT
tril resulted in the death of the lizard within the first
two-hour period and use of this approach was
discontinued,
In the descriptive section, scale fealures ovcurring
bilaterally were counted on both sides, Paravertebral
scales were vounted using Greer’s (1982) method,
Verelalion was quantitatively assessed around the
most successful pitfall line using (wo methods:
a) We revorded all plant species ina 30m * 30m
quadral, 13 m either side of caeh pitfall line, taking
collecrions of plunts in cases Where identity was not
vlear. These collections were subsequenrly identified
ani ladged with the State Herbarium,
6b) We assessed veyetalive cover within the
30 m >» 30 m quadrats by the simple transect mether!
know as “Step-point” (Cunpinghat 975). At every
stop in set directions across the quadrat we recorded
the phint species encountered at the Lip of the shoe
scoring approximately 300 points inthe quadrat. This
method vives estinutes of percentage of ground cover
ol phint species, bare ground, rook and litter,
The behavioucal data we present ster mainly fron
field observations, bul we obtained confirmatory duta
inmmany cases from observations of captive specimens
at Adelaide Zoo (T Morley und R. Ainsley pers,
cumin.
Description
Specunens
The following descriptive section js mended to show
varration within a single population. Earlier auitiors
(Ehmann 1983; Milchell 1950) give additional data
front some ol the alder specimens. and all of the
aviiluble data on this older material is discussed in
Shea's. thesis,
“A total Of 63 specimens Was collected during the
survey period, and an wdditional four were bern in
caplivily, Sis ofthese specimens were collected dead.
Killed by the vlapid snake Psewdonaja rexrifis (owe
specimens) ar birds of prey (two specimens). A
seventh, R4838, died during trials of an external radio
transmitter. Four specimens, an jmmiuture, an adult
male and two adult fimales, were retained and are on
Joun lo Adelaide Zoo. The remainder were murked ail
released, The success of collection methods varied
according t time of year Pittil| rrappine was
successtul only during Novenber-December, while
ham collection became much more successtil in
February-April (Table 1),
All predator-killed specimens are damaged, although
in two the damuge was minor and confined to restricted
ures of fhe body: R40687 and R40744 hud some
necroue patches an ihe dorsal and ventral body surface.
R40687 hud o sagittal freetuire of the skull curining trons
the restral Seale to tie level of the eye, R4AD689
hid jhe left peek und posterolateral region of the
skull deeply gouged and partly removed and a deep
REDESCRIPTION AND NOTES ON TILIQUA ADELAIDENSIS NG
wound in the right side of the neck, R40728 lacked
the head and right forelimb, as well as the liver, heart,
lungs and stomach, R40738 and R40745 were partly
digested, with [he skin on the body sloughing away
and much of the internal sofl anatomy missing, Two
skulls and associated mandibles were prepared trom
snuke-killed specimens, one articulated (SAMA
R40738) and one partly disarticulated (R40745).
Tarhe t Relatiw success of Capture techniques,
Number caught
Navembe r- February -
Method Decembet May
Pitfalls 7 nil
Hand 4 ar
“Fishing” not used 4
* All but one neonates
General Appearance
Tiliqua adelaidensis is'a moderate sized skink wih
short extremities, a relatively heavy body snd large
head. The body and tail are soft and flexible, but the
head is heavily armoured by the well developed head
shield asteaderms. The toes are short, the third ancl
fourth toes of the hind foot being equul in length. The
tail tapers rapidly from the base and is thin and slightly
laterally compressed over the distal two-thirds.
Colour
The dorsal surface of the head, body. linnbs and tail
is light grey brown, yellowish brown, orange. lan or
chocolate brown, the distal portions of the extremities.
especially the forelimbs, being a paler yellowish hue.
Dorsal and lateral scales usually have narrow darker
edges producing fine, longitudinal lines along the back
and sides. The lower lateral surfaces are pale greyish
becoming off-white ventrally, The dorsal surface ts
unmarked. or shows variable development of blackish
spotting, including a vertebral series of irregular small
blotehes (which may coalesce into 4 ragged vertebral
stripe) trom the nape to the base of the tail, and several
laterodorsal and upper lateral series of small black
flecks; these may be crudely aligned to form weak
transverse bars, The midlateral region often has
scattered grey-white flecks, The venter is immaculate.
or with slightly greyer marging lo (he seales forming
narrow longitudinal lines, The iris of [he eye ts bright
orange, The tongue is pale rose pink. with no trace
of melanic pigmentation, The rool of the mouth und
buccal commissures are mauve. The abdominal
peritoneum is black, Juveniles are consistently greenish
arey to mid-brown, becoming reddish tan on the Lail
and Jimbs. The range of variation in black pigmentalion
is similar to that seen in adults, but many juveniles
have more extensive and obyious white spotting on the
body,
Fig. |. Head shields of Tiliqua adeloidensis (SAMA R40838). A_ dorsal view. B, left lateral view. C, detail of chin shields
D. detail of rivht side showing asymmetric variation in circumocular sealition, Abbreviations for cireumocular scales;
po. postoculir: prsb. presubocular, psb, postsuboculars, sh, suboculars; sc. supraciliaries, Scale bar = 10 nim.
220 M. N. HUTCHINSON, T. MILNE & T, CROFT
REDESCRIPTION AND NOTES ON TMIGUd ADEE AIENSTS til
Selatan
This 1s based on eight specimens, four held in the
S.A, Museuit (R40687, R40689, R40744 and R40838),
plus alive adull male. two adult females and sutracdult
field at Adelie Zoo, The head shields are shown in
Vip. |
The species has the scalation gharaecteristios listed
by Shea (1990) for Tiga. The last supralabial is
horizontally divided, and the temporal scallion
postenor to the secondary temporals is irregatur and
hitle diierenuated from the beady sculahion. The
supraciliary seule count fs reduced (mode 3) A
complete subecular scale row, consisting of a laree
presuboculur, a Jarger anterior and smaller posterior
subocular ind Gne ot two postsuboculars, runs from
the anterior supruciliary to the postocular, Nuchals are
absent or a single Slightly enlarged pair may: be present,
olen separated by wu median oecipilal or internuchal
swale.
Dorsal scales smmboth, in 33-36 (mean 34.6) rows
al dudbody; paravertebral seules 77-86 finewh 85.0),
piravertebral scales belween piriclal and the level al
anterior edge of hind limb 70-81 (mean 7.6); subd igital
faniellae under fourth toe unpaired. 10-13 (new 11.6),
Huss iy point contaet or narrowly Separated (U.2);
strong postmarial groove present, prefrontals in broad
comuck, frontoparietals paired; interparictal about same
Size as frontal, much larger than frontoparietals and
separating paretals:; each parietal in contact
wityrolaterully wath the postocular and posterior
supracihary and bordered posterolaterally by four or
live enlarged scales: ua median oecipilal scale present
of abseot (U4) postenor to interpanetal: supranculars
3/3, the first (wo Contacting the frontal, supraviliaries
3/5 (5/6 in one specimen). the second the longest;
complete subocular scale row present; supralabials 4.
ihe eighth only balfas high us the seventh: infrulabials
79 (meun 7.9), a single primary temporal 1s followed
by three secondary temporals; postmental contact frst
two infralabials; 4 single large, quadrangular anterior
car lobule,
The dorsal head shields, frontonasal. prefrontals.
frontal, rontoparictals and interpartetal, are thickened,
with Weakly corrugated surfaces and deeply. incised
sutures: This rugasity increases with sie. 1 large
dais te suboculat scales also become thickened urd
slivhily overhane the suborbital suprilabials,
Skull and mandihle
The characternstics noted for ine species by Shea
(1990; 1992!) are coufirmed in the SAMA specimens.
The species has moderatcly nayrow separation of the
pre-and posilronials on the medial orbrlal margin,
binger-like nasal processes of the frontals are presem
1) give a W-shaped lrontal-nasal contact. The jugal 1s
broad and fateened and contacts the posttrontal, A
jwetimal bone ts absent, There is a well-developed
medial palatine process of the ectopterygard which
excludes the pleryguid from the edye ot the infraorbital
vacuity, The coranoid provess oF the dentary is enlarged
and swept back to cover the lateral tice of the dorsal
process of the coronmd. Dentition is heterodent, with
the cheek teeth markedly lamer than the unter teeth,
In venerul aspect (Fig. 2). the skull and mandible
of J) adelaidensts ure very like those af other Tiligue
species. The proportions of the snout, lapering und
pointed. rather than blunt and rounded, and the parietal
region, constricted, father than laterally expanded,
resemble thase of adults of the other species of Tage,
The dorsal head shield osteoidernis are initrimately bused
with the bones of the skull root, tog greater degree
than is usual inmost other Zi/gua. Thos, eved though
fo adelatlensis ts as small as or smaller than
cogeneric neonates, ibis ontogenetivally advanced in
8 proporhions and degree of ossification, In these
features 7 adelaidensis is progenetic. not neotenic as
1s Frequently the case in miniaturised lizurds (Rieppel
1984), Even so, 7, udelaidensix also shows some
neyienic features in retaining relatively large upper
temporal fenestrac and in some aspeets of braincase
anatomy (Shea pers, comm.).
Unique Or unusual features of the skull are few bal
include the very closely apposed, almost parallel
Palatal ramy of the plerygoids, In most other Tifique
(7. vigas iy an exception) the pterygoul margins are
more widely separated und diverging. The closely
“pposed plerygdids may be correlated with a nirrowing
of the skull posteriorly as J) udefeidensis bas the
ereates! skull width across the jugals, rather (han al
the level of the quadrates. The maxillary process of
the jugal tapers rapidly and fails to contace the
prefrontal. This character state 1s shured with 7
niultifasciaw and most TL occipitelis, Whereis ocher
Tilfqna species have the jugul extending anteriorly to
centact the prefrontal adjacent to the lacrimal loramer.
Pye. 2. Skull nd mandible of 7iligua adelamensts (SAMA RAOTIS). AL skull, right hileral view, showing conker beryeen
dugal Guy anc squaniosal (sq) bones, By skull, densa) view, showing soert inayillary process ef the jugal (p, mas) and
open supratemporud fenestra (st) Mote peesmtent head shield osteoderms whieh obscure sutures of prefrymtal und pastlpontial
bones: other skull rool sutures are visible through the osigoderms. , ventral View, Showing contipurauton of the palin!
complex (ec. ectopleryyoud: pl palatine: ply, pian. Do right mandibaler rivius, hibtal view, showing Very well developod
co
coroniid process of the dentity (po cond. E,
Voruimhbulitr nimus, lingual view. Seule hue = 10 nin,
WW
M,N, HUTCHINSON
The upper temporal fenestrae are relatively larger than
in adults ot other Ziligua species, in which they are
Slielike or ubsent,
Dimensinns
Based on 6 specimens. Snout-vent length (SVL)
38-107 mm (mean 73.2). Axilla-groin length (AGL)
23-67 mim (mean 44.0. 0 = 58). Sooutuxillu distance
(SAD) 16-41 mm (mean 28.3, = 38). Forelimb length
U2) mm. Hind limb length 1-21 mn. Tail length (TL)
22-79 nim,
Sexual Dimerphism
Adult males und females (arbilrarily defined as
>85 mm SVL) differ markedly in head and body
proportions. Adult males on average are shorter than
females (SVL males range 87-106, mean 93,4, n = 17;
females range $8-107, mean 98.3, 1 = 14) bul head size
does nol overlap (SAD/SVL. males 0.36-0.42, mean
0.38, n = 17; fermales 0.30-0.35, mean 0.33. n = 14).
The relatively large head size of males is a frequent
phenomenon in shinks (e.g. Simbotwe 1985;
Hutchinson ef al. 1989; Hutchinson & Donnellan !992)
but is rarely as marked as il is in large adult male 7
adelatdensis. Male combat has been reeorded tn other
Tiliqna species (reviewed by Greer |989), 4 selective
pressure which might lead to their large head to body
eH IOS
Ecological Notes
Habitat
The main study sile lies al an elevation of about
500 mon undulating terrain cut by small, intermittent
Stream courses. The ground is stony in places, with
underlying shale and sandstone bedrock just exposed
op the surtiee. The soil ig hard-packing clay-sandy
loam (red-brown earth, French ef al, WOR). The
vegetation of the main trap site is charavterisuc of
degraded remnant native grassland, with the area
around the site being bure of trees and Shrubs (Table
2) A lull list of plant species is provided in the
Appendix The exact original nature of thjs grassland
is unclear. as copper mining and farming near the site
Takin 2) Groged cover ai Tiliqua adelajdensis syeedy sire
faveraned over three quadrats).
Ue Cover No. Species
Vepetaluni
Native Armuial an
Perennial 2 At
Tntrodueedt
Annual 56 Ve
Perennial 14 4
Bure Grown 3
Rock 4
Litter <1
‘T. MILNE & T. CROFT
since 1845 may have resulted in the removal of trees
and shrubs for both industrial and domestic use,
Similar hilly areas in the district support Allocasuaring
vertivilfara (drooping she-eak) low open woodland
over similar geound cover of native grasses (Stipa spp..
Danthonia spp.) and mat-rush or “irongrass”
(Lomandra spp, as recorded in the survey area,
Jessup (1948) concluded that at least some of the area
had probably been essentially treeless priar to
European seulement.
The relative abundance and species composition of
native grass species varies within Lhe study area. The
immediate vicinity of the most productive trapline was
dominated by one species of spear grass (Stipa,
tentatively identified as 8. eremaphila), but elsewhere
in the same paddock, pygmy bluetongues were found
where §, nodosa was the common spear grass. and
other grasses, notably wallaby grasses. Deathonia
spp., and wire grass, Aristida behriana, were loeally
common, Thus the precise species composition of the
understorey may be less important for |he survival of
pygmy bluetongues than the tussocky siructure which
provides ground cover throughout the year. Other
areas surveyed in nearby paddocks which do not
appear ) Support pygmy bluctongues showed
increased pround cover by intraduced plant species,
and hence a decrease in the amount of caver during
Jate suminer-autumn. Feyen at the least disturbed part
of the study site, anly 50% of the ground cover js
perennial yegetanon, so thal the surlace of the ground
is far more exposed in wuturnn than in spring Adjacent
areas inal have been ploughed al any stage show
bhinimul recolonisauon by native plants, in particular
Lomandra spp.
Aside tron. a single juvenile found under a stone,
the only mierohabitats in which 7) adelaidensis have
been found sheltering are vertical or near vertical holes.
We suggest that the lizards are using spider holes, not
digeing their own burrows. The holes are perfeetly
circular, up to about 20 mm in diameter, lack any sign
of excavated soil at the entrances and are
indistinguishable trom holes al the study site inhabjted
by lycosid and mygalomorph spiders In at least one
case uw lizard inhabued & hole to which the lid of a
trapdoor spider was shill attached. Two large species
of Wolf spiders, Lycuse stirlinut, which makes a lid for
ifs burrow. and Lyeasa gilbert, which does nol, have
been identified af the study sile as has a species of the
trapdoor spider genus Bloktstonia (probably B. aurea)
Lycosids und their burrows are very common ia the
urea, The lizards appear to make no obvious external
modilieutions to the holes, save for a slight bevelling
of the edges caused by their coming and going,
Home ranve-and movements
We obtained few recaptures, and made only limited
use ol padio-tracking, se that our result. are
preliminary.
REDESCRIPTION AND NOTES ON JALIGLE SUELAIDENSIN par
The urea covered by a male that was radiotracked
during spring encompassed 70m- and two burrows
over a period of 4)X days, Home range overlap occurs,
us there were ah least Iwo other occupied burrows
within the area that the lizard covered. A second,
sinutler male avas fued with the cracker buc wheo
relocated’ after the first (Wochour period it wais
monbund, apparently dirough exhaustion as it
struggled to force iself into a hole bul was prevented
by the bulge of the transmitter Tt was striking that,
although shelwr in che farin of dense prass and
Lomandra tussocks was all around, the hazard put all
of its ellorts into finding Shelter ina hole,
The same ole successfully tracked in Noverniber
was caught again ducing March, It was still within the
sdine area as the previous spring but in yet amother
hole. In another case, a burrow containing o fenule
and young was abandoned and the female was
discovered. in another hole same Sm from the orignal.
Only one ol 22) caplures during Noveniber
December was an adult female. All animals captured
during spring/early Samimer were caught either by
pitfall trapping or by hand, and se cequired the
individual to be actively doying away: brome the burrow,
However, of the adults captured during Pebruary-April
one (a female) was captured by hand and i4 by “fishing”
bul pital trapping caught none, despite in excess of
Uiree weeks’ intensive pittalling tn areas kKnowi to
support a significant number of pygmy bluetonpues
Of the adults caught during summer/autunin, were
females. and (hree were niles, a reversal ol the trenet
shown ib spring/early summer,
The lop-sided sex ratio in our spring sample (only
a single adult female caught) suggests much greater
levels of mile wetivity during the spring mating seasort.
This is supported by the six predator kills collected
during Qetnber November, of which five were sexually
mature males, while only one was an adult female.
Males of other species ob Titigua Wave increased
activity Jevels. relative to females daring this time of
year (Bull et al. 1991). The results so far indicate that
both males and females may Dave relatively [rmited
home ranges throughout most of the-year, but on spring
inales appear to Wander more Widely (or at least, more
often),
During November-December 1992, yainfall Was
above average and daily mixin ranged between TS°C
and 36°C. Days when lizards were caught had maxinw
between 20°C and 36°C. and the time of vapture varied
from O8:1S to 19:30, AU of the days on which pyyrmny
bluctongues were caught were supny at some stage,
and so ground telnpesatures ofica would have exceeded
the air leimperstures, ‘Traps were monitored after dusk
on warm nights lo cheek for nocturnal activity, but ne
lizards were captured at this time: Diurnal behavsour
was alsa observed in captives. which sliawed tee sin
al nocturnal activity.
Diet
R40687 contuned wo late lepidopteran kurv (a
hadenine noctuid, probubly Perseenitia, G. Brown
pers. Comin); Ra40689 had the reuwuay of iui
apparently identical larva, plos several leaves ano
Howers of the herb Medicago. RA0744 conuined three
aero grasshoppers and the remains ol a small
beetle, Ehinann (1982), after examining speciinens then
inthe SAMA collechon, recorded cockroaches, wats,
a spider, grasshopper and beetle, and some plaints
(Dianella seed, possible chenopod material). Wild
lizards accepted grasshoppers. offered us bilit, und
caplive wnimalsy eat an ormmiverous diet, meting
Mealwomns, crickets, chopped truttund vegetables and
raw ego. Thus the evidence to date meheutes that 7
widelaidensis.cuts a wide variety of invertebrate unindls,
but also includes plants in its dict,
The relative dack of movement away trom the
burrows, at least in Jate summer-aurumo, suggests that
at this time ol year 2 adeleidenyiy ts probably a sit
and-wait foriger, Burrow entrances dre used as vantage
points rom which lizards would be wble to make short
forays afier any prey deteeted nearby. The presence of
leaves and flowers mi the diet sugeests wider forays
at east it Spring. The study site Suffered an Outhreak
of plague locusts (Cherteicertes rermondfera) durity the
period of observation which may have influenced the
livards’ behaviour. ‘The sedentary belaviour may alse
be related tw avoiding predation at the time at yeir
when ground cover is spursest.
Repraduetion
All of the predator killed mules (colleetcd between
MAO/92 wd 30/1/92) had testes that were enlarged
and turgid’ Gr starting to regress One of the tripped
males had strings of dried seminal material protruding
trom the vent, The single dead lemale (R4A0744,
coUceted 7/1/92) had four enlarged ovanan follicles
(iwi left, Qwo right), Together these data indicate a
spring testicular maximum, with spermyogenesis aid
tating behaviodr comeing with avalanon, (her
Tiliqgue species synlopie with Lo adelaidensis, 1 ragesa
and 7. sclveoides, are Known to be spring brecders
(Bourne e7 al, 1986) Stew 1993),
Neorites were first observed at the field site on 28
Pebruary, 1993, when four young were observed i the
burrow of ah adult female, Iispection of other burrows
known to harbour lemales revealed other tillers, liter
size ranging | Lmeay 2.3). AM juveniles examined ut
this time had row pink umbilical sears indicating very
revent birth. Two females which pave birth at Adelaide
Zoo during mid-March had liter sizes of three and one,
Within two weeks of ther discovery, juyeniies and
sommehmes adults abandoned the birth burrow, Juvertiles
were observed from late March olone in holes.
sometimes. rouch smaller than thase used by adults
The sinallest animals caught i Novether were
66-70 mm SVL. while dhe targest younp witht divdinus
224 MON, HUTCHINSON, ||) MILNE & T CROFT
umbilical scars in May were 56-60 tum SVL, This
mmphes that the 7 adelaidensis born at the end ot
summer reach about 70 mm SVL by the end of spring
of that year, This SVL is considerably smaller than
the smallest breeding female recorded (SVL. 96 nim),
so thar first breeding (ust occur po earlier than the
second spring (approx. ZO months of ape),
Beheviinar
Tiliqua adeluidensis appears to be extremely
sensitive to both movement and nose. mukiny a
difficult ta observe lizards basking outside their
burrows, Lizards bask with the back legs or tip of the
tail remaining in the entrance of the burrow. From this
position. they can back rapidly into their burrows. iL
disturbed. Once inside, the lizards increase their
security by turning the head sideways to force the snout
and peciput against the sides of the burrow. Thus
wedged tn place. the lizards present only {he armoured
head to an intruder und seem almost invulnerable: any
ereature small enough to enter the hole is unlikely to
be strong enough lo harm or distodwe the lizard. ‘The
sioniticwnee of the heavy osteodermal armour on (he
head seems to be associated with the usc of ie head
4s a combination anchor and doorstep.
Several behavioural attributes of captive \jzatds
sugges! why detection of the species in the field os
difficuh. The first t erypsis, When disturbed the
lizards freéze: such stationary animals are well
cnopfaged against the reddish brawn soil of the aret,
muking them bard to see, The second isa reluctance
iy emerge into open spaces, In caplive condihons the
lizards spend almost all of their active time in holes
or among liter. Thirdly. the lizards have a well-
developed tind unusual ability t move in confined
spaces, Both un the field and in captivity they tive been
observed moving directly backwards into cover or
dowtie burrow. avoiding a Ftum with is coneomitant
greater level of disturbance. The small, Una uly may
taeilitate this manoeavre by not getting: in the way as
ley might af longer or more massive. The body is
unusually flexible, an-athibute which probably assists
in negoliling the confined spaces of burriws oy
tussocks. On several occasions, an unimal was kaw
to jae entered i tile head first, turned around withia
the hole und emerged head first-
Wher disturbed or handled, this species has not yey
heen observed if exhibit the exagyerated defensive
display employed by ils larger rekitives (Cunpenter &
Murphy 1978; Greer 1989), When handled the liars
will iwist with great strength and agiity. often gaping
ind endeaveuring to bite in a siimiar fushien to oder
mehum-sized skinks (e.g. Bxertia spp.
Possible Reasuns for Decline
Tiligha adefaidensiy remained undetected ina well
lrequemed part of South Australia for over three
decades in spire of diligent searching by herpetaloensts.
Based on our experienee with the species, we sliggest
that byo factors Combined to hamper searchers ~ lick
otintormation on its habits and habilat, and a probable
real decline tn the number of populuuons.
Ehmann (1982), based on his inferpretation of the
specimens and histerjcal data. speculated that 7
ddeluidensis had inhabited a fimestone-chenopod-
mallee association, While the speculitive nature of this
ussessment wats noted by Ehmann himself. it
nevertheless exerted & bias on many of the altermpls
to locute the species. The grasshand habitar in whieh
We have found the species 1s difficult to search casually
for reptiles, and consequently has received litle
aliention, This is especially so because must grassy
terrains in this pan of South Australia are assumed to
be heavily disturbed. supporting only the most wide-
ranging and ecologically tolerunt reptiles. The
distinction between fative versus introduced grasslands
has been important in locating populations ol cts
specicn.
Even When one knows (he correer habitat, (he shy
nature of (he lizards makes (hem very cillicull to tingl
unless is known exactly tiow und where to seareh.
The crucial piece of information that allowed us cy
locate additional colonies was the discovery of their
dependence on spider holes, information that was only
acquired ater we had located the initial population
through good luck. The holes are difficult to see unless
the observer is right next to them, so that any lizard!
occupant has plenty of warning ef human approach ane
is oul of sight by the time the hole has been policed,
Intensive field work had been going on tor six weeks
atthe site before we observed any 7) avefaidensfs under
natural cenditions. However, having discovered the
combination of habital and rojerohabital, we have been
able to locate further populations of pygmy bluetonyzues
in nearby areas, The species is now known to occur
ui six Sues running from south of Burra Wo north ol
Hallet. the extremes separated by about 50 kin.
The apparent rarity, caused by lack Of allention {u
grasslands as a habjtal, ts probably coupled to a drastic
reduction in abundance caused by pasture improvement
und cropping. Native prassland similar co tht an which
the species oceurs at Burra once extended south on
lor the Adelaide plains, but as pritie agricultural land,
was ane af ihe first jnajor habitats in South Austra}
ts be cleared und ploughed. Ploughing permunently
aliers the vegetarion and ground cover, converting @
hulive, largely perennial Nora into en introduced.
lurgely annual one. In addition, ploughing would
destriy the burrows, killing lizards directly and leaving
Hie survivers without shelter and at the merey of
predators, The fact (hat most speenmens were colleetea
lust century may simply be correlated with the fact thal
this was when most of their habit was being convenes
lo agrieulure,
REVESCRIPTION AND NOTES ON TILIQUA ADELAIDENSIS
Reimaming native grassland ts now extrenely patchy,
and the remnants have generally been heavily invaded
by introduced annuals. However, populations of pygmy
bluctongues have now been tound in some of these
remaining pockets, and provided that land use is nut
changed, these colonies may be relatively secure.
Further work is focussing on finding the extentol the
current range of the pygmy bluetongue, approximating
population size within these areas, and confirming
muny of the subjective eas developed during the
course-af this first season's field work. This will lead
(ot aecurate assessment of the status of the species.
Acknowledgments
We are very grateful ty property owners, particularly
Barry and Judith Eberhard, lorallowing: access to their
lund, and to Steve Vogel for arranging aecommodution.
225
This work was made possible by grant support from
the Endangered Species Programme, Austmalian Nature
Conservation Agency. and we thank S. Stephens,
B, Male and J, Hicks of ANCA for their support.
P; Copley of the Biological Conservation Branch, Dept
of Environment and Natural Resources, assisted with
advice and ideas, and DENR also provided a vehicle
and assisted with financial support and held equipment,
For assistance in the Held we thank D. Armstrong.
G. Armstrong, R. Foster, B. Miller, T. Morley.
D_ Niejyalke and A. Yates. P Lang, Native Vegetation
Management Branch. helped with plant, especially
grass, identificulion, and R. Davies and M. Hyde
kindly shared their dat on sites where native grassland
remnams still occurred. D. Hirst (SAMA) identified
spiders and provided background information on
them. We thank C. M, Bull and G. M. Shea for
their comments which considerably improved the
THUNUSCTIPL.
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at Australian Wildlife, Sydney),
Preven, Ry J, Marneson, W, BR. & Carne, Ao L. 19648)
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(Luvertilit: Semeidae) trom Western Apstralia, with notes
oo the blotegy and rlatiiaships at other Australiin species
Ree Aust Muy 44, SAQ-ST3
HuteHivson, M,N, (1992) Theeatened reptiles in South
Australia. Section 7 Ar lay, S-P (Ed) “Threstened species
und hubitats in South Australia. A cutubyst lor community
uction”” fSouth Austdian Advisury Committee on
Threatened Species, Adelaidet
d& Downtd | As. S.C (1992) Taxonomy and genetic
yanialion in the Australian Haards of the genus Pyeudemoia
(Scieidaes Lygosominag). J Nur Hise, 26. 25-264
. Ronertsus, PB & Rawiinsunw, PA. (OND)
Taxonomic revision and ecology of the endemic Tasmiainnarn
scincid lizants Leilapisna microlepidoton and 1
pretrosain. Pap. Pree R Soe Kasiiania 123%, 257-274.
Jnssor, J. PL (1989))A list of the vascular plants of South
Australi (eeition 3), J) ddelaide Bor Gard. 12, 1-163,
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Counues Eyre. Burra und Kimberley, South Australia
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Phypes, Wo (1804) Eine Unersicht der van Hr. Richare
Schomburgk un dais zoologische Museum cvingesandlen
Amphbien, dus Buchslelde bel Adelaide in Stdaustratieo
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(Lacertlit: Seineidae): generic diagnoses und systematic
relulionships. Menr Od Mus. 29, 495-520.
(1993) The mate reproductive. cycle of the eastern blue
tongued lizard Tilfgua scrncoides scincoides (Squamiata:
Scincidae). pp, 397-403. Jn Luaney, D..& Ayres, Do (buts)
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Zoological Sociery of New South Wales with Surmey Beatty
& Sons. Sydney}.
a & Hutcuinsos. M,N. (992) A new species of
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reproduction of Levnproplolis. enichenoti (Lacertilia:
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and Remtiles” (Surrey Beaty & Sons. Sydney)
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Two new simple methods far catching siall fast Tigards.
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226 M. N. HUTCHINSON, T. MILNE & T. CROFT
Appendix I
Plany diversity at Tiliqua adelaidensis srdy site. Species noted during the general search but not recorded during step-pointing are indicated
hy (*), Txxonomy modified from Jessap (1989).
Annual/ % Ground
Species Common Name Perennial Cover
Native species
Chenopodiaceae
Atriplex semibuccat . 00.00.2002 ee ee berry saltbush |, oh, P *
Enchylaena tomentosa 0.20. ee ruby saltbush... . . P ti
Matreana aphylla o.oo. 00.0 ee eee cotton-bush ee ee eee P *
M. enchyvlaencides. . . does oe wingless bluebush —.. 2.2.2.2 2.22.22... P *
M. trichoptera soo... Ebetp eyed te alo ».+. Mallee bluebush - 2-222... - Sets P *
Salsola kali ........ Moab gd up tela de apa hoki ifs" 2 ee roly-poly 20.22... A "
Amaranthaceae
Prilotus spathulams .-. 2-0. 2.22. ee ee es pussytail cee ey ee apie (eg P <I
Oxaliaceac
Oxalis Pperennans (005 ce ee SHelebetpe, Hate sorte. tye yy terse eter P <I
Geranlaceae
Brodit Crinitum coc cece eee cece eee eee blue storks bi... 66, P +
Euphorbiaceae
Euphorbia drummondii. cece ee eee Coste Weed 222 ee P *
Rhamnaceae
Cryptandra amara. ccc. cee eee ecae ca. long-fower cryptandra ... 2 222.22... P °
Thymeliaceae
Pimelete micrantha... cc cc eens veeees 9 Silky rleeflower 00.0.6 ceva ee ti P *
Rubiaceue
Asperula conperta occ c cece cece vee ee seeeees = COMMON WOOdrUET.. 222.2222 222 - P =
Convolvulaceae
Convolvulus erubeseens .. 2.22.22 2 ee Australian bindweed 000... dre detols P <i
Campanulaceae
Wahlenbergia Inteola 22.22.2262 ee eee Bluebell te ee ees P +
Goudeniiceae
Goodenia pinnatifida.. 2.22.2 2.2 202202 002-.224- tut-leal goodenia, ~~... ee, P *
Asteraceae
Leptorhynchos Squamatus oo... 2.55 ide : scaly DUMONS 20 ce eee : P +
Minuria leptaphylla ..o.c cece eee .. minnie daisy , ae 1 , P *
Vittadinia cuneate... zesicd.t a - New Holland daisy —....-. he P #
Ve orabiiss «fe ht tc. pat bl chi as cskiecasBefder woolly New Holland daisy...) 2.0.0... P 2
Liltaceae
Lomandra efftsd veces rye cree ees ers eevee eee © SCONTed Mat-rush 2.6.2 Lecce eee P |
EL. multiflora oo cee es cranes es SUID Materdsl ee eee cee P 4
Juneaceae
Hearieus PufaMins oo eee toad USN eee eee P <I
SORPAUSSIT ee ce eee S154 bbeSe thstisa gt big TUM fuos eh pa emilee eres P <i
Poaceae
Aristida behriane.... brush wire-grass wivwit es rye se P *
Danthonia caespitesa white-top ..-0. 0.22.22. 2 2.2 2.28 P 1
D. pilosa... 4 pret velvet wallaby prass .. 20.0.0... P 3
D. racemosa... 1.6... wallaby grass... .-_..- P <l
Stipa Blacktie eee » rested spear grass 6. .....5., P <l
S. eremophila (possibly S. puberula) .. . desert spear grass 2... ee eve P 8
Se PUI ths tts Lp pages gebat ots bebe ttaieiedbe SPCR TENS Cup mei cree c aa cals bias P 3
Themeda triandra o.. 000 c cece eee eee eee ees Kangaroo grass leon Ne = P <l
Introduced
Polygonaceae
Rumer dyumosus 5... - 0 oe em ee WOU TIGER ee deisel oel adeeb deel nde fopee fol wena P =
Fabaceae
Medicage litteralis —. 2.2242 0 2s ee eee strand medic --.- 20-2. ee A 6
M. minima ...... seeceees WOOlly burr medic ._. . A a
Trifolium angu. aot » farrow-leaf clover... ., : aeons A 3
T. arvense ... veeeees hares fool clover oo... ee ee A 2
Boraginaceae
Echium plantagineum ~~~. -- 622 eee salvation Jane 0.0...) . 2... eee A 8
Nearastema apulun. cc ccccrverre cre rsercersees hairy Sheepweed ... 00.0.0... 22.22 ee A 7
Lamiaceae
Marrubium vileare oo... ce ee veesey. horehound .,..,.. tis ees oe A i
Salvia verbenaca ...-..-....--..-.--.--.--.-- . Wild sage —, Wdemet ister ddadea bebideeba teebnde P 14
Asteraceae
Arctotheca calenidula-.......- 2. -. 2-0-2 - 2 es Cape weed eee ead 4 A ‘4
Carthamus lanarus saffron thistle... 0. 0c. ce jee yuuyee A id
Hypockoeris glabra smooth catsear. =... ee i A <1
Sonphus yleraceusiy es. Soe ee ee ee tee ge common sow-thistle 2.20.2... - _ A *
[ridaceac
Gynandriris setifolia 00.0660 o es es ss opted veces thread iris... _ sb! --Bt6-. 2593 A <i
Poaceae
Avena BUrbata oo ccc eens bearded oat 22.22.0262 ee ee ee A 23
Brachypodium distachyon .. false brante: Hs c tite ete th oi A sa
Bromus rubeny.., . red brome ...,..,.., fecsee A ]
Hordeum glaacum . . northern barley grass . otter A *
Lolinn perenne... perenmal ryegrass... hot 4a lat fone . A <u
Malta muaraliy o..- Tats-tail fescue 2... ee ee A 2
MOOROWIPORA CHAMBERENSIS, A CORAL FROM THE
EARLY CAMBRIAN MOOROWIE FORMATION,
FLINDERS RANGES, SOUTH AUSTRALIA
By MARGARET K. FULLER* & RICHARD J. F. JENKINS*
Summary
Fuller, M. K. & Jenkins, R. J. F. (1994) Moorowipora chamberensis, a coral from the
Early Cambrian Moorowie Formation, Flinders Ranges, South Australia. Trans. R.
Soc. S. Aust. 118(4), 227-235, 30 November, 1994.
Moorowipora chamberensis gen. et. sp. nov., an Early Cambrian coral with a
morphology close to tabulates, occurs in the Moorowie Formation of the eastern
Flinders Ranges. The oldest accepted tabulate coral Lichenaria Winchell & Schuchert
first appeared in the Early Ordovician. However, it is possible that Moorowipora
chamberensis is an early representative of the tabulates, extending their time range
down to the Lower Cambrian. The tabulate-like characteristics of Moorowipora
chamberensis include the form of the corallum, which varies between cerioid and
fasciculate, the wedge-shaped to spine-like septa and generally complete, well-
formed, convex-upward or undulating horizontal tabulae. Moorowipora chamberensis
has some skeletal structures in common with the Early Cambrian species
Flindersipora bowmani Lafuste, and Tabulaconus kordae Handfield, but is unlike
other previously described Cambrian coralomorph.
Key Words: Moorowipora chamberensis, Early Cambrian, Moorowie Formation,
tabulate coral, Flinders Ranges, South Australia.
Transactions of the Royal Society of S. Aust. (1994), 118(4), 227-235.
MOOROWIPORA CHAMBERENSIS, A CORAL FROM THE EARLY CAMBRIAN
MOOROWIE FORMATION, FLINDERS RANGES, SOUTH AUSTRALIA
by MARGARET K. FULLER* & RICHARD J. F, JENKINS*
Summary
FULLER, M, K. & JENKINS, R. J. FE. (1994) Moorowipora chamberensis, a coral from the Early Cambrian
Moorowie Formation, Flinders Ranges, South Australia. Trans. R. Soc, S. Aust. 118(4), 227-235, 30 November, 1994.
Moorowipora chamberensis gen. et sp. nov,, an Early Cambrian coral with a morphology close to tabulates,
occurs in the Moorowie Formation of the eastern Flinders Ranges. The oldest accepted tabulate coral Lichenaria
Winchell & Schuchert first appeared in the Early Ordovician. However, it is possible that Moorowipora chamberensis
is an early representative of the tabulates, extending their time range down to the Lower Cambrian. The tabulate-
like characteristics of Moorowipora chamberensis include the form of the corallum, which varies between cerioid
and fasciculate, the wedge-shaped to spine-like septa and generally complete, well-formed, convex-upward or
undulating horizontal tabulae, Moorowipora chamberensis has some skeletal structures in common with the Early
Cambrian species Flindersipora bowmani Lafuste, and Tabulaconus kordae Handfield, but is unlike any other
previously described Cambrian coralomorph.
Key Worps: Moorowipora chamberensis, Early Cambrian, Moorowie Formation, tabulate coral, Flinders
Ranges, South Australia.
Introduction
The Early Cambrian coral Moorowipora
chamberensis gen. et sp. nov., is found in slumped
reefal blocks within a megabreccia forming part of the
Moorowie Formation in the eastern Flinders Ranges
of South Australia. Moorowipora chamberensis occurs
with Flindersipera bowmani Lafuste 1991, and at least
three other previously underscribed corals at a site
close to the disused Moorowie Mine (Fig. 1).
Most of the several known Cambrian coralomorphs
have been regarded as doubtful early representatives
of tabulate corals. They show unusual aspects in their
morphology; a significant gap in time (~20 my)
occurs between the Early Cambrian and the oldest
accepted Early Ordovician tabulate coral, Lichenaria
Winchell & Schuchert 1895 (Scrutton 1979; 1984;
1992). Moorowipora chamberensis has structural
characteristics suggesting its affinity with the tabulates,
and is possibly an early representative of this group.
Stratigraphy and associations
The Moorowie Formation and its lateral equivalents,
the Wilkawillina Limestone, Mernmerna Formation
{= Parara Limestone, Dalgarno & Johnson (1962)| and
Oraparinna Shale, form part of the Early Cambrian
Hawker Group (Dalgarno 1964). These units are
mainly limestones, calcareous shales and mudstones,
* Department of Geology and Geophysics, University of
Adelaide, Adelaide, South Australia 5005, Australia.
' Mount, T. J. (1970) Geology of the Mount Chambers
Gorge region. B.Sc. Hons. thesis The University of
Adelaide (unpubl).
Harr, J. (1989) Lower Cambrian corals from
Archaeocyathan-Epiphyton clasts within the Moorowie
Formation megabreccia, northern Flinders Ranges, South
Australia. B.Sc. Hons thesis, The University of Adelaide
(unpubl. ).
i)
and siltstones, with minimal siliciclastic arenites, and
variously reflect shallow marine, reefal, shelf-margin,
slope and basinal environments of deposition.
The corals occur within reefal boundstones that have
tumbled as large talus blocks to form a megabreccia,
which comprises the middle part of the Moorowie
Formation (Mount 1970!;, Hart 1989?; Lafuste e¢ al.
1991; Savarese et al, 1993). This stratigraphic level
represents part of a suggested third transgressive/
highstand phase of the Early Cambrian (Gravestock
& Hibburt 1991),
The talus blocks of the Moorowie Formation,
analagous to those in contemporaneous reefs, comprise
Type 5 shelf margin build-ups (James & Gravestock
1990). The reefal system was established in a high
energy marine environment encroaching on a marginal
fan (Savarese ef al. 1993). The fan comprises coarse
breccia and is suggested to have formed as a result of
local diapiric activity, However. we have not observed
any reef structures in their original placement.
Coral colonies, together with archaeocyaths and the
calcimicrobes Renalcis Vologdin 1932, Girvanella
Nicholson & Etheridge 1878 and Epiphyton Borneman
1886 occur in transported reefal blocks, which vary
from cobble size to about JO m in maximum
dimension. Within individual blocks, the organisms
are commonly preserved in life position, The dominant
faunal elements vary markedly between blocks, from
archaeocyaths, to stromatolites and more rarely corals.
These differences probably reflect mass collapse of
different parts of a zoned reef complex of reasonably
wide areal extent (the distinctive biofacies represented
surely formed in areas some hundreds of metres broad
implying that the main reef front had a fringing
geometry). The rapid slumping of the talus into
deeper water (presumably the fore-reef) probably
protected the carbonate frameworks from vadose
228 Mi
diagenesis, fesulting m thé remarkably pristine
preservation of the fauna, The corals commonly form
encrusting lo uprigh(, tall colonies. the latter up to
6-70 cm high, Individual colonics tend to be widely
spaced,
Flindersipora bowmant has. bee found in bioherms
tthe lower Qrupaniana Shale at Ten Mile Creek (Fig.
I), as well as near the Moorowie Mine (Lafuste er al.
1991), Al Ten Mile Creck, trilobite and echinoderm
fragnients foun hash beds in the Oraparinna Shale, and
assowlated archueocyalhs huve been correlated wilh
Hin Assernblagze 9 (Daily 1956), whieh equates with
Whe Pararaia juneue Zone of Sell (1990; Lufuste er al.
W491). This stratigraphic correlation indicates that the
Moorawie Formation is Boturmiun in age
Preservation
Although coniplete recrystallization of the colonies
has oceurred, an indication of the primary micro-
WILPENA
PouND
F)] MIDDLE CAMBRIAN
Wi] Lower Camanian
a wir
nm ‘ei wt ’ > rT 4
fi — ye
Afr asics 7 7)
20k: = 12!) Atelnide
—te NX
*QUORN
Fig 1. Location tdap shows fossil aceurrence near the
Maoorowie Mine and (he distribution of Early and Middle
Cumbrian outeraps in ihe Minders Ranges wf Seah
Australie.
Fig. 3. Longitudinal sections of holotype SAM. P34165; iMustrating parts of the colony (see Fig. 2), A & B
K. PULLER & R. J.
bh JENKINS
structure of (he skeleton can be seen ii seme patchy
domains. Edges of the possible primary hioeryvstal
Platelets ure distinguished hy crenate lo Wavy lines ot
Minute inclusions. Within the much coarser crystals
comprising the recrystallised fabric. Narrow, Jath-like
zones showing irregular extinetion under polarised light
occur al same places approximately wansverse lo
skeletal elements where secondary carbonale crystals
penetrate (he coral skeleton. This suggests a residual
overprint Of the orginal rhineralogy disturbing the
optical continuity of the subsequent recrystallisation
Apart trom recrystallised spar, cavities surrounding
corallites and within the calices may be filled with very
fine sund or sill. Tt appears that during life, part or
whole colonies may have been teniporarily covered
with a thin layer of sediment causing the death of some
vooids. The survivors rejuvenated pew parts of the
coralluny, Some colonies seem to have been eroded
hy rupid, energetic influxes of coarse sand (also noted
on caleimicrobe enerustations and drchaeneyaths),
allowing only.o lew corallites to contiue their srowllr,
Fractures post-date growth ind are often filled with
very tine Sediment (Figs 2 and 3C),
a
Fi, 2. Inlerpretive sketch (longitudinal section) of holotype
SAM P34165) luitrangy cone to fan shaped colony: the
form of individual corullites: protocorallite Ga); tabulae (b):
sop (ce): calice (d): lateral increase (top centre and righr)
and peripheral intracalicular merease (x 2),
Top of the
colony With Walls extendilig above the coma}, methods of increase and coralline swuctre: © — base of colony (x 4):
an Do Aivber magnifiewion of (B) showing normal and thickened tabulae (lower leit); and oblique prajections of the
ae
vuler wall (oenire, right) (x 1)
A CAMBRIAN CORAL. FROM THE FLINDERS RANGES 229
230) M. K. PULLER & R_ J. F JENKINS
PY
Fig. 4, Transverse sections and interpretive sketches of holotype SAM P34165: (A & B) cerivid (x 10) und (C & D) fasciculate
(x 10,5) areas of the corallum. Diflerences can be seen in corallite shape. septa, and new walls across corallites formed
during offset Jormation, Rejuvenation of corallites (C & D) is evident top right and bottom centre.
A CAMBRIAN CORAL FROM THE TLINDERS RANtIES ps
The term ‘platelet’ (Lafuste er al. 1991). 15 used to
describe relic microstructural elements of the skeletor,
Systematic palaeontology
Phylum; CNIDARIA
Class: ANTHOZOA
Subclass! 7TABULATA
Family: uncertain
Genus: Moorowipera gen. nov,
‘Lype spevies: Moorowtparnt chamberensis sf. ov
Epynalogy: For the type locality near the Moorowie
Mine 19 the eastern Flinders Ranges of South Auscralta,
Diaenesis, Corallum snail, varying from niassive
cerioid lo fasciculste, comprising polygonal, oval or
rounded corallites. Coralliles are Ing. tuberoid to
irregularly cylindrical, Walls are thin, wavy to crenale,
rarely straight. Tabulae are irregularly spaced. mostly
complete, concave upwards to undulating horizontal.
Septa absent, or number up ty 20 in each corallite.
Where present, septa are randomly spaved. short and
form wedge to spine-like projections mo coralliles,
arising from inward angulations of the wall. Pores
appeur to be absent,
Moorowipora chamberensis. sp. nov,
FIGS 27
Etymology: For nearby Mt. Chambers.
Didgnusis, As for genus,
Type specimens; The specimens described in this paper
gre held at the Seurh Australian Museum (SAM),
Holotype SAM P34165, four thin sections, SAM
Pad165-1, SAM P34165-2; Paratypes SAM 34166-1:
SAM P34166-2,
Material: The holotype, paratypes as well as several
other colonies come from one rock sample measuring
270 mam long, 230 mm wide and 120 mm thock: taken
from 2 large boulder within the Moorowie Formation.
near the Moorowic Mine in the eastern Flinders
Ranges.
Description: In wansverse section (Figs 4. 5), corallites
Show gradation into two distinct habits, massive cerioid
and tasciculate dendroid, In fasciculale habit, cocallues
rarely touch, are circular to slightly aval in shape and
vary i diameter from 0.95 to 3.75 mm. Corallites with
massive habit ure rather irregularly shaped 5 or 6 sided
polygons, sometimes oval of reetangular, rarely
ciroulary they vary individually in diameter frony 077
(oO 3.5 nim,
Walls are thin, varying bebween O.E fam and 0.15 1mm
in thickness and show a relic fibrous structure (Figs
4-6).
A midline is apparent between many adjoining
corullites. Walls are wavy to slightly crenate, being
rounded tn isolation or adjacent to-small spaces in the
corallum, with a tendency to become straighter and
jess distinct where they merge with the walls of
adjoming corallites.
in longitudinal Section (Figs 2, 3). each colony ts
yenerally small, numbering [rom 2 or 3 tu about 16
corallites, Colonies are cither cone- to fan-shaped,
diverging Outward front a single protocorallite, or more
rectangular, where they appear to anise from several
adjacent corallites. Increase is both lateral and
peripheral intracalicular, producing 1.2 or more offsets
(Figs 2, 3A).
Individual corallites are tuberoid to irregularly
cylindrival in shape and vary greatly in width and
length (up to 19.5 mm). priur to increase (furmiution
of a new corallite). ‘The base 1s rounded and blunt. the
protecorallite producing 3 or 4 short septa pnor to the
formation of an initial tabula, The calice js prominent
extending between 2.5 and 4.75 mm past the last tabula
(Figs 3B, 43D).
Tubulae are mostly complete, mainly convex-upwarl
or undulating horizontal, often down-turned where they
meet the wall (Figs 2, 3), They are irregularly spaced ,
but commonly oceur at the same level in adjacent
corallttes. The distance between them is highly vartable
(0.35 to.2.1 mm), while the thickness of tabulae varies
from 0.002 mm to OA mm.
Septa number up to 20 in isciculate corallites: are
very short (0.01-0,2 jan), generally equal in length,
triangular to wedge-shaped, often indistinct, They form
protrusions of the wall of the cordllite at sites of inward
creasing (Figs 4, 5). ly massive, cerioid eoralliies in
the main body of the-corallum, there may be up to KM)
sepi or septa may be absent, Sept are randomly
spaced; long and short sepia may alternate, or only
Jong of short septa may be present. Sepla are wedge
to spine-like in Shape. Sept are generally longer than
inthe fasciculate corallites. In longitudinal section septa
ure observed as continuous vertical Jumingar plates
intersecting nermally with tabulac.
Microsiratture: The miserostructure was studicd 3
diagnifications up to x 200, and photographs were
taken under polarised light.
In transverse section at low magnification (Figs 4,
5) relic fibrous elements which form the sclerenchyme
and apparently represent indications of original
biocrystals appear as Continuous lineae across the wall
und into the septa, At higher magnification (Fig. 6)
the interlocking fibrous elements form triangular ly
rectangular bundles. conyposed of narrow parallel-sidex!
nu M, kK
and hlude-like geniculate structures up to 164 gm lone
wd 47 jam wides ‘These are angled towards and away
front the centre of each corallam. The bundles have
the appearance of crossing, or bein Stacked over
underlying layers. Near the midlineof walls the fibres
ure often less oblique and have a slightly different
prienttion, uppewring to be broader and more
randonily oriented.
In longitudinal section at rmagnifications of * 100
to & 200, the midplane ol the wall seems ta bave been
composed of irregularly shaped, crenate, Interlocking
platelets which individually represent the fibrous lineae
OF (ransverse cUts, Platelets may be alrnost rectangular,
vertical to shghuly inclined, oevasionally almost
horizontal in the middle of the wall. They commonly
oveur diverging outward towards the top of corallites
(Fig. 7), Wall platelets are more clongate and wider
than the fibre hindles seen im (runsverse section, being
up) to J90 jam to length and 138 pm in width.
Tabulae are continuous with the inner edge of the
wall, which converges slightly ground them. The
stricture of the labulae ty similar to the sepla and wall
Hu 5 ‘Trtisverse section of holotype SAM P4dtS: part of
conilum showing both cero and fasewulate wreus (x 323),
PULLER & R, J, & JENKINS
in Untnaverse section, with bundles of fibres being
mainly triangular, or irregularly shaped. Trungular
bundles of fibres are up to 360 jam in length, and
308 yim in width, In tabulae of riormal thickness,
adjoining triangular bundles interlock forming 4 crenate
je 6. Transverse section and interpretive sketch af holotype
SAM P34]6S: illustrating triangular and rectangular bundles
af fibres extending across the walls of the corallite. xx!
and y-yl define boundaries of HMustriiion. Wis mudline
OF tho wall bx ety
A CAMBRIAN CORAL FROM THE FLINDERS RASHES 257
upper and lower surface, A more complicated
interlocking pattern is formed in thickened tabulige,
Remarks: Moorowipora chamberensis is dimorphic in
both the morphology of the colony yd the method of
inerease. Dimerphism often occurs in labulates and
may be shown as differences m corallite size, shupe
and internal structures (Oliver 196%, 1975). Many
intraspecific variations are probably environmentally
controlled, being influenced by factors nreluding the
adjacent sediment, and the position of corallites within
i eolony (Oliver 19686). The two different
morphological forms observed in M. chumiberensty
probubly reflect the mode of increase, whieh also
uppears to be dimorphic. bemg related to the postion
of individual corallites within (he corallum, Corallites
within the fasciculate area. whieh generally ovcurs al
tlie outer edges of the colony, appear to have resulted
trom lateral increase. Oltsets branch away frony the
main colony producing isolated corallites which rarely
(ouch and are therefore unnaffected by crowding (Figs
2, 5) Such corallites are thus round to slightly oval
hig 7. Longitudinal sevtion (cartoon sketeh) showing hurge
oryotals of the recrystallisation fabric (a) incorporating the
wall of compound corallines. (by Relic bioerystal fihees
diverge towards (he coral lun,
in transverse seeuion, Peripheral intracalicular increase
is most common in the niasive. eeriod purrs, und
where it solitary corillite has beconte established (Pigs
2,3), Usually one, Iwo or more offsets are produced
at the sume time, with new walls growing fron) sites
of septal insertion across the calive. Both methods af
increase Conmionly occur at different slupes of growth
within the same corallile, and are probubly related to
the amount of space surrounding tf,
The variuble distance hetween pibulae and the
presence Or absence of septs do not uppear tobe linked
to any particular sayge of growlh, @ Churacteristus which
hus been suggested as possibly oecurnmy ty some
tabalutes (Hi IRI. Sepa primarily oecur in the
protocurallite und inmature corallites.. while ar other
stages of growth they may or muy not be present,
Diseussivg and Conclusion
When compared with other Cambrian coruls
supeested to have labulate affinities (Serutlon 1979),
Muoranvipard Chamberensis is closest: in is
inorphalogy to Mihulaeomns kardae Huadheld 1964,
trom the Early Cambrian (Botomfan)y of east central
Alaska and Brilish Columbia. Mn vertical section,
chamberensis and T kordteae ditfer in the shape of the
coruiites, which are more tubular in appearance in the
former, Height and width wary, with mature corallites
being up to 1¥.5 mm long and 5.0 pi Wide th M
chamberensis. while corallites of To Aurdae are up le
65 nom long and 27 nuvin width inthe colonial torm
(Debrenne eral (O87). The tubule also differ. being
undulating Horizontal to concave upward in M,
chaniberensts and either horizontal oy slightly concave
downward in 7 kerdae. Invemplete labulae are more
dissepiment-like und walls are generally thicker in the
Natter (Handfield (969). The microstructure of both
corals is significantly. diterent if trinsverse section,
being yeniculaie fibres jm M. chenrberenois, wad
vongentic light and dark wavy laminations in 7
horec. “Vabulue also differ, being formed from bundles
of fibres extending upwitrd and downward from ia
medial Jine in the former, unlike the wo fayered light
and dark zones of TD kordae, However, platclets
tlongitudinal section) in (he walls ol ML chembereniyis
are of similar shape. hut generally laraer, MW,
chitmberensiy may belong wi the family Tabulacondac.
hut, the above differences, likely preclude it fram thes
division.
M. chamberensis is distinguished tram Lipepara
livye and L. daseia Jel) & Jell 1976, fron the Buurly
Middle Cambrian of western New South Wales, by the
presence of tabulae and the shape und arrangement ol
septa, although the corallites of L. /issa une al similar
length and width. CumArouyjpa mantanenss Frite &
Howell 1955, from) the Middle Cambrian of Brush
234
Columbus, is distinguished from M. chanierensis by
the more slender corallites and te absence ol labulie
and sepla. Most of the Early Cambrian vroraiunrorphs
deseribed by Korde (963, 1984a, b, 1986, 1990), are
partly synonyms of already described khasaktiids o1
hydroconozoans and partly nent dubia or auller
(Ahuravley fel, 1993, p. 369).
M. chamberensis also differs trom Flirudersipara
deawnon’ Lafuste 1991, although there are some
similarities in nierostructure. M chaniberersis. \s
verioid (o faseiculate-and has wedge to spine-shuped
sepa up to 0.2 mm in length arising from continuoys
walls O1-O0.15 mm thick. Ln contrast & fae is
mieatidroid Lo cetoid, and has 6-16 strongly developed ,
striigbt to slightly carved sepia up to D8 min in length,
with the edges of septa bearing very short blunt spines,
Walls form very shart segments between the septa and
are O15-0.25 mm in thickness (Lafuste ev al. 1991).
In BE bowen tabulac are mostly coneave-downward.
and closely spaced (0.2-0.3 fam), but are andulating
horizontal fo concave and more regularly spaced in M-
Chumberenasis. The mode of increase is by longitudinal
fissfon in the focmer while both lateral and intacalicular
peripheral increase oceurs in the fatter,
In transverse section. the microstructure of both
corals is similar, with walls consisting of geniculate
fibres which diverge and couverge in two directions.
In vertical section platelets in M_ chamberensts are less
clongate and broader When compared with F bonymunt.
IL ts considered (hat the genus Lichvearia which
has a time range from the basal to the eurly/lare
Ordovician is the most ancient tabulate caval. ts
colomal form is cerioid, it has a simple morphology
M K. PULLER & R. LT. UINKINS
and tabulae, is aseptate, and may have rare mural pores
(Bassler (950: Flower 196): McLeod 1979: Scrutton
1984. Laub 1984). M. chwnberensi¥ has structural
characteristics which demonstrate its affinity ty the
tabuliales Gncluding septa, which sre not present in
lichenarids). These are (1) the ceriond to fascieulaw
form of the eotony; (2) the spine-like to wedge-shaped
septa; (3) ts mode of increase; (4) the generally
complete well-formed tabulae. The observed relic
icrostructure appears to be similar lo the pinnately
(clinogonally) fibrous structure of some tabulates (sec
Hill 1981. p. F452)_ including lichenarids, though the
extent of the modilying influence of diagenesis js
uncertain,
M, chamberensix with its tabulatetike characteristics
may be either an early representative of the tabulates.
or belong lo a new group ol corals with convergent
evolution contributing to their similarities. These
alternatives have been susvested by Latuste er al. (1991)
tor F) bewnent, whereas Scrutton (1992). regards the
latter possibility as most likely. The addition of AZ.
chamberensiy to the group of known early
cormlomorphs, provides further evidence that tabulate
corals jnay. have their origin in the Karly Cambrian,
Acknowledgments
The authors gratefully acknowledge Brent Bowman
and John Hart who collected the studied material. Dr
David Gravestock is thanked for additional information
and his constructive criicism of a draft of this
manuscript. We would also like to thank Mr Gino
Snideto for his photographic work,
References
Bossier, R, 5, (1950) Baunal tists tnd deseripion pf
Palueovoiwe Corals. Geol, Sor, Any Meni 44, 1-305
Dany, B. t1956) The Cambrian in South Australia 20eh
Iniern Geel, Coneres,s Mexicvis 2, OF 47
DalLoarne, C. Ry (964) Lower Cambrian siratigmphy of the
Plinders Ranves. Trans. R. See, 8, Aus 8M. 129444.
& Johnson, J.B, (1962) Cambritn sequence of the
western Flinders Ranges, Query, geel. Norwes, Geel, Surv.
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Denresat, F My, Ganororr RooA, & Larusre, J G
(987) Jahulvenias, Handfield, Microstructure and its
ImMplicanon th the lasnony of primitive corals for, Fal
fl, 1-9
bhowie. RH. Ol) Moareya and related colemal corals.
New Mew (rts feet Ment, 7 1-97,
berry, MOA, & Howni BR (1935) An Upper Cambrian
coral from Manna, Jor Pal, 2% 18h 183.
Grwestock, D. 1, & Hipawer, J EB. (199]) Sequence
Sirtivraphy of ihe eastern Officer and Arrowie Baxins:
uw Hatiework for Cambnan oil search. APEd foe 3,
7-100)
HanpreLD, RoC. (1969) Early Cambriun coml-like fossils
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Hitt, DO G98) Rugosi itnd Tabulats. pp. 1762. i Teicher,
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Kansas, Boulder, Coloradey }
Jamis N. PF. & Gravisracn, Doi. (1990) Lower Cambrai
shelf and shelf niavgin burldups, Flinders Ranges, South
Australia. Sedimentology 37, 455-480.
dri PA, (1890) Biostralizgmiphy. de Bengtsen, 5., Conway
Morris. S.. Cooper, B, J. Jell, PA. & Runnepar. 8. N_
(ds) Early Cambrian fossils from South Australia. alssec:
Atastsn, Pal Mem. 9, 14-19.
& Je dS. (1976) Barly Middle Cambrian corals
from western New South Wsles. dleheringa t 181-195,
Konbe K. B. (1959) Problematic fossils from the Cambrian
deposits of the south-eastern Siberian Platform. Trans! ,
Dakt. Acad: Sct. USSR, Eurth Sev Sect. DS, 358-360,
Larustt. J. Deskinnr, F, Ganon. A, & GkavesTick, D
(199); ‘The oldest tabulate coral and the associated
Arvhuwocyatha, Lower Cambrian, Flinders Ranges, South
Nustealia. Caio, 24, 697-718.
A CAMBRIAN CORAL FROM THE FLINDERS RANGES 235
Laus, R. S. (1984) Lichenaria Winchell & Schuchert 1895?
Lamottia Raymond 1924, and the early history of the
tabulate corals. [V International symposium on fossil
Cnidaria. Paleontographica Americana 54, 159-163.
McLeop, J. D. (1979) A Lower Ordovician (Canadian)
lichenarid coral from the Ozark uplift area. Jour. Pal. 53,
505-506.
Ouiver, W. A. (Jr.) (1968) Some aspects of colony
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(1975) Dimorphism in two genera of Devonian tabulate
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(1993) Paleobiological and paleoenvironmental context of
coral bearing Early Cambrian reefs: implications for
Phanerozoic reef development. Geology 21, 917-920.
ScruTton, C, T, (1979) Early fossil Cnidarians, pp. 161-207
In House, M. R. (Ed.) “The origin of major invertebrate
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(1984) Origin and early evolution of tabulate corals.
IV International symposium on fossil Cnidaria.
Paleontographica Amer. 54, 0-118.
(1992) Flindersipora bowmani Lafuste and the early
evolution of the tabulate corals. Fossil Cnid. & Porifera
21, 29-33.
WINCHELL, N. H. & SCHUCHERT, C. (1895) Sponges,
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Early Cambrian microstructural diversification of Cnidaria.
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NOTES ON THE MORPHOLOGY AND BIOLOGY OF
CTENOPHORUS McKENZIEI
(STORR, 1981) (SQUAMATA: AGAMIDAE)
By M. PETERSON*, G. M. SHEAT, G. R. JOHNSTONE & B. MILLERS$
Summary
Peterson, M., Shea, G. M., Johnston, G. R. & Miller, B. (1994) Notes on the
morphology and biology of Ctenophorus mckenziei (Storr, 1981) (Squamata:
Agamidae). Trans. R. Soc. S$. Aust. 118(4), 237-244, November, 1994.
The systematic status of Ctenophorus mckenziei is reviewed, on the basis of
additional material from Western Australia and South Australia. Habitat data for
known localities, stomach contents, one record of clutch size, and behavioural
observations and thermal preferences of captive individuals are provided. C.
mckenziei is morphologically very similar to C. scutulatus, differing mainly in adult
size and subtle modifications of colour pattern. C. mckenziei and C. scutulatus are
phenetically most similar to C. cristatus.
Key Words: Squamata, Agamidae, Ctenophorus, morphology, biology, habitat, diet,
thermal biology.
Tronsucrims of the Royal Soetety of S$, tusi, (994), WIRE), 237 244,
NOTES ON THE MORPHOLOGY AND BLOLOGY OF CrENOPHORUS MCKENZIEI
(STORR, 1981) (SQUAMATA; AGAMIDAE)
by M. PETERSON”, G. M. SREAT. G. R. JOHNSTONE & B. MILLER§:
Summary
PEreRson, M., Sura, G: M., Jonnsion, G, R, & Milter. B. (1994) Notes on the morphology and bielogy of
Crenophoruy mokencier (Storr, 981) (Squamala: Agamidae). Trans, R See, 8, Aust, 8(4) 237-244, November,
1994,
‘The systematic status of Crevophorus mekenziet is reviewed, on (he basis of additional inuterial trom ‘Western
Australia and South Australia. Habital dats for known localities, stomach contents. one record of clutch siae,
and behavioural observations and thermal prederences of cuptive individuals are provided, CG mckenzie ts
morphologically very similar to Co scuelaius, differing mainly in adult size and subtle modifications. of colour
panier, ©, mekenzied and GC sentuaiyy ure pheneteally most-siinilan to CL erivneaniis.
Key Worps: Squamats, Agatnidac, Cretephorus, morphology, biology, habitat, diet, thermal hiolegy.
Introduction
The agamid Armphibolurus mekenziet was described
from two specimens (Western Australian Museum
[WAM| R59753-54), collected atS km and 8 km SW
of Ponier Rock, Western Australia (Storr 1981). The
species was subsequently transferred, With a number
of other agamid species, to Clenophoruy (Storr 1982).
Additional specimens of C. mekenzic? have since been
reported from Western Australia and South Australia
by Schwarier & Tyler (985), McKenzie et al. (1987).
Anon, (1988), Wilson & Knowles (1988), Armstrong
(1992) and Ehmann (1992), This additional material.
previously uncharacterised, considerably extends our
knowledge of the morphology. distribution and habitat
preferences for this species and provided the basis for
this paper.
Materials and Methods
Five specimens from Western Australia, additional
to the types, have been collected: South Australian
Museum (SAM) R2832, Western Australia; WAM
R9|764-65, 9km E Kilidwerima Granite Rock,
3204'S 124°0H)’E, 29.111, 1984, WAM R91842, ROIS52,
Is kin E Kilidwerinia Granite Rock, 32°03'S
1249°05'E. 5.x.1984. Eleven specimens have been
collected from South Australia: SAM R25572. R26162,
12.6. km (air distance) NE “Colona”, 31°31/8 132°09'Th,
SAM R26160-61, 16.2 km (airdistance) NE "Colona
3193'S 132°1'R; SAM R26890, 15.2 km (road
69 Alvah Street, St James, Western Australia 6102.
Departinent of Veterinary Anatomy, University, af Sydney,
New South Wales 2006.
School of Biological Serences, Flinders University, Bedford
Park, South Australia 5042
Department of Animal Services, Adelaide University, Suuth
Australia 5005,
or Da ad + oF
distanve) NE “Colona”, 31°31'S 132909'T, 13.x7.1984:
SAM R32264-65. R32268-69, 0.5 km S Mitcherie
Rockhole, Yumburra Conservation Park, 31°27'S
132°449'E, 17.x.1987. SAM R32266-67, Skm S
Mitcherie Rockhole, Yumburra Conservation Park.
31929’§ 132°S0'T, 18.x.J987 (Pig. 1)-
Measurements taken on preserved material follow
those used by Shea & Peterson (1985). Log-transformed
morphometric data were statistically analysed with
analysis of covariance, using the SYSTAT statistical
package (Wilkinson 1987).
Dietary data were obtained by examining stomach
contents of six specimens: SAM R2832, R25572.
R26160-62. WAM R59753.
Four specimens were maintained alive for sone ume
following collection. The lizards were housed in a
Vivarium with a heat lamp and fluorescent lighting,
both turned on at 630 h and off at 1930 h,
r RRR ed LAA Pe watt tte sfotemntctea dae)
of wen
yer a a \
" x “a i \ \
i]
\vay
( ye
ik’ ¥
i]
| | f
f r
Chermiivin ov henares Me 2 i
a
tame] yen cpm gemt ie pentmngerin eyes teegepet ie cpepemepe| tom sympey ofeyenrqey
for bite we 1 vat
Fig. 1 Distribution of ©. mekenciel,
Dan M. PETERSON. G. M. SHEA, Gi OR
corresponding to the local nattiral photoperiod at (hat
lime. Water, mealworms and chopped fruit were
available ad (ibitum. However. ane ineividual (R32266)
hecume emaciated and was euthanased At 1800 h ov
|Exi/ 1987 (ie, 54-55 d post captire) the remaining
three lizards were placed in a thermal prutient
(8-60°C). and allowed to aecliniate, Calibrated copper-
Constantin thermocouples were placed atteast 20 mm
into the cloaca and held in position with surpical tape
Cloacul temperatures were clectronieally revordéd
every WU qin, for each animal from H30 hh on
12.40.1987 to 130 hon 13-xi1987, Any temperatures
taken Wlule lizards were tungled in the thermocouples
were excluded from the analysis,
Thermal dala were uwnalysed using @ one-way
ANOVA to test for differences in thermal preferences
between individuals. In comparing thernial preferences
during scotophase and photophase, data fron) all
individuals Were pooled.
Results
Morphology
The South Australian sample (snout-vent length
|SVL] 41-76 nim, ¥ = 63.5 mm, n = (ly is similar in
Morphology and colourabon t the Wester Australian
sample (SVL 3L37h5 mm, * = 584 mm. n= 7)
There are minor, but statistically significant diflerences
beeen the two (Table |) in the relauionship between
tail length (TL) and snout-vent Jength (slopes,
Fj, = 5477, p =< 0.05; overall, TL/SVL
215,0-252.9%, ¥ — IRS 1 = 1 vw 206,3-236.9%
% = 222.5%. n = 6 respectively) and between head
depth (HD) and head length (HL) (slopes:
Fy = 9.900, p < 0.0 overall, HD/HL 57.7-67,9%,
¥= 46.4%, nil vy 62.0-67.5%. ¥ = 64.0%,
n= 7 respectively). with the eastern juveniles having
longer tail and the eastern aduley a more depressed
head. ltrs possible that the apparent long tail on eastern
juveniles may be an artefiet due to the Lick of very
small juveniles m the sample.
The affinities of Co rickevsred have liek previously
been adequately defined, Storr (1981) assumed a close
allinity with © yeurilarus (Stirling & Zein. II).
but provided no argument for this. The CL mekenzier
maternal now al hand clearly indicates that this species
is very close to C seutulatus. Head, body and limit
proportions are similar, the arrangemenn of spine
clusters, rows of enlarged scales and skin folds oa (he
head, geek and body. is identical, and most elements
of (he colour pattern are shared. Both species have the
following colouration elemems (Fig. 2):
1. a pale vertebral stripe, bordered by
2 a dark paravertehral stripe. bordered faterally by
JOHNSTON & B MILLER
& a longitudinal series of pale blotehes, vouleseip
anteriorly and ofien posteriorly to form an irregular
stripe from hehind eve, over the ear und upper
postauricular spines, to the fail base, where Ihe two
stripes ollen fuse. This clement is bordered laterally
by
4+. an uppee hier zone from below the eye, through
the ear, over the forelimb and hindlimb te the
proximal part of the tar, hordered ventrally. by
5. abroad pale midlateral stripe from axilla to groin,
then along itd edge of thigh along the proximal
parl of the tail, dark margined ventrally:
6. on nape and forebody, a Series of dark fransverse
bars conneeting the dark paravertebral stripes and
umterrupling the pale vertebral stripes;
7. narrow pule caadal rings usually present, overlying
other elements. of tail pattern;
8. dark ventral markings in males aud some females
consisting of al least a broad median stripe trom
chin to gular fold. broadest just rostral to the gulut
fold. and a rhomboidal to kite-shaped dark breast
patch, extending caudally a variable distiace, 1m
seine causes Wo the grom, Some specimens of both
species have additional dark ventral nurkings.
including streaks and spots laterally on the thnoat.
and dark anterior margins tu the thighs (Piz. 3).
Store (981) differentiated CL mckenciet from ©.
veuludatus hy four characters: smaller size, deeper
head, fewer subdigital lamellae onthe fourth toe anc
colouration (back blackish brown wilh whilish
yerlebral sinpe and transverse Hines vy pale brawa with
paired dark brows bletthes merging on forebuck
to form ceossbunds). The additional miaterial
now uvuilablé supports the size difference
(SVL = 315-765 mm yy 3-5 mm; Stor erul. 1983)
The smallest male C yackenaies with dilated, was-filled
femoral and preanul pores and turgid testes hus
SVL = 48 mm (SAM R25572), Whilea female of the
sume size (SAM R2ZO890) has well-developed ovaries
and oviduels: One female (SAM R2K42-
SVL = 715 mm) is gravid, with three oviducal epes
However, the remairting three characters enrployed are
non-diagansnc or require qualification.
The purportedly deeper head of © mekenzier is aot
Upparcar in our measurements (Table ), althoouh
ts Clear lrum the very much lower values given by Storr
(1981) that he used different measureiients to obtain
u head depth head length ratio (possibly head deptt
46 defined by Witte 119851), We haye measured heal
depth at ihe highest point of the skull. over the parietal
eye, and head length from tip ef siout to rostral onan
of-car. As noted above, the castera satnple of adult Oo
mekenztet had a shallower head than the western
sumple A difference of similar magnitude was also
present between the caster sample and C) setulae
(slopes: Fy oy = BSA, p<! UIT).
CTENOPHORUS MCKENZIE! BIOLOGY 239
Fig. 2. Dorsolateral views of A. male C. mckenziei (15.2 km NE “Colona”, SA) and B. male C. scutulatus (Hospital Rocks,
WA) in life (photographs M. Peterson).
214i)
The lower number of subdigittl lamellae in C.
mckenzie? is not an absolute difference, and is
geopraphivally variable. Western C. mckenziei had
counts. of 25-40 (% = 27.4, mode = 27, n = 14) while
eastern CL mekensiel had 25-32 (k = 29.6, nude = 30,
n= 18). The C. seatilatuy sample we examined had
28-41 (X = 34.9, n = 80) subdigital lamellae, only
7.5% fewer than 32, although Storr e7 al. (1983) give
a range of 31-44. All means are significantly different
(pairwise (-lests: castern vy western CL mekenicte/,
tay = 3.36, p < 0.05; western Co mekenziel vs C,
weutidatus, toys = O88, p < 0.005; eastern C,
inckenstet vs CL seutulatus, ty, = 8.33, p < 0.005).
As noted above. the colour pattern is composed of
the sume elements in cach species, The difference noted
by Storr (981) is due to paling of the dark dorsal
markings in C. seu/aletus, particularly the centres of
such markings, und enlargement of the pale dorsal
spots. The dark erossbunds on the foreback of C
seutilatis are also present in C, mckensiet, though
narrower, less contrasting and often tewer.
Sympatry 1s not yet known between C. mckenzie!
and C sewnlarns to confirma species-level difference,
M. PRTERSON. G, M SHEA. GR, JOHNSTON & B. MILLER
although the two species are Known to approach to
within &5 km of each other (WAM R91764-65 vy
RO5538, 0.5 km S Buningonia Spring, respectively)
However, the lower number of’ subdigital laniellae in
western C Mmckejidiel, Which are yeographically closest
oC. scutulaiuy, logether wilh the much smaller adult
size, and consistent evenly dark upper lateral zane (uy
usually light centred) suggest that C mekenziet ts
specifically distinct,
Habitat
The two Western Australian localities listed above,
and the two South Australian localities 12.6 kn NE
and 16.2 km NE “Colona” are respectively sites BAZB,
BAS, KO2 and KO4 of MeKenzie & Robinson (1987).
while the localiues 0.5 km S und.5 kim S Mitcheri
Rockhole are sites MIST and MI2I of Copley & Kemper
(1992). From the combination of phorographs of the
habitat and floristic lists at each locality provided by
McKenzie & Robinson (1987) and Copley & Kemper
(1992). specimen collection data from both museums
and observations by the senior author, it appears (hat
the primary Moristic components of the CL mckenzie
Piz. 3. Typical melanio ventral parterns ol A. CO mekensiel and B.C. seuuidaras, Scale bars = 2 en
CTENOPIIORUS MCKENZIE! BIOLOGY
habitats are Kuecalyprusy oleosa Ev.M. ex Miq. (giant
mallee). dcwcie asmeddli Fv.M. (umbrella wattle) and
A. papyreeurpa’ Benth. (western myall) over a
chenopod understorey of Arriplex vestcaria Heward ex
Benth. (saltbush), Maireana spp. (bluebush). Rhagedia
ypinesceny R.Br. and Sclerolaena spp. The chenopod
understorey is common to all localities. The substrate
type ut sites BA2B, BAS, KO2, KO4 and MISTI is
crusting sandy clay loam to loam (MeKenzie &
Robinson 1987, Copley & Kemper 1992), while the
specimen from 15.2 km NE “Colona” was found under
a bluebush on a non-crusting sandy substrate, similar
to the substrate recorded from site MI2L,
Al sites BA2B, BAS, KO2, KO4 and 15,2 km NE
“Colona. CL mekenzie/ was sympatric with C. pictus
(Peters 1866), and at the former three sites it was also
sympatric with the agamid Twupanecryptis lineata
Peters 1864. At site MI21, it was the only agamid
recorded, while at site MIST, C. fardi (Storr 1965) and
24
Pogona minor (Sternteld 919) were also recorded, The
latter species was.also found al [5.2 km NE “Colona”
Diet
Phe dominant prey tems in all specimens examined
were ants, particularly small Jridemyrmex, although
some larger ants (Cumpdnetus, Melophorus and other
unidentified genera) were present. Other prey items
recorded were Hemiptera (single individuals ol
Poecilometis sp. in R22835, R2832, two pentatomids
in R2832, and a small unidentified bug in R26162),
Coleoptera (one small curculionid prothorax and a
large scarab larva in R2832; three unidentified elytra
in R25572), Mecoptera (a larve abdomen in R2832),
Diptera (one smull fly wing in R2832), Homoptera
(one small wing in R25572), Orthoptera (one small
grasshopper head in R25572), Hymenoptera (one smal!
bee head in R25572) und tower parts (R26161).
TALE 1. Allametric equations and caleulater! valiws for cranial and somatic proportions in C. mckenzie, (ek), C- scutulatus
(yeu) and C, cristatus (eri), Eastern and western samples of C. mekeneici are kept seperate for tail lenuth and head depth,
hut pooled elsewhere, as The equations were nor significantly different, Values a and b solve the equation y = bx, se. by
Vlandard errorufia, C= correlation coefficient, al. < tirection uf significant allumetry, Cyr, Cog and Cyys-are caleulated
percentage proportions at SVL. = 32, 76 and 10S mim (approximate mininiim size for all spectes. meximim size of C. mckenzie
and upproximate large udult size for bath C. scutulatus and C. cristatus)-
a S.C, b i al. Cy Cr, Cus
AGL/SYL
mek 1.2128 0535 183 969K - a7 46 =
scu 1.0609 A284. 323 RGR + =a Ad 45
cri 1.0847 0381 295 O854 + 40 a) 4
TLISVL
mek(E) 9732 0648 2.006 9617 0 2.37 2,32
meki W) 1.1662 423 1.145 9948 4 2,04 2.35
seu 11020 0344 15h 9843 + 2.26 2.47 2.55
eri 11582 (282 1104 9030) \ Lol 2.19 2.31
FLLISVL
mek R450 O38 785 9677 - Ad 38
scu 9254 0348 573 9739 = Aa Al 0
cri CG 030K) 47h BRT () 43 42 42
HLLISVL
mek 8714 (0339 1.503 9763 - 6 46 =
scu 9166 ORL 1.275 9797 = OS 89 86
eri 9911 0196 1.097 9953 0 99 O8 8
HLISVL
nick 7939 (0458 530 9493 = 26 2 =
scu R563 (0223 421 ORT4 - 26 33 22
eri 7613 0174 631 9934 = 28 22 meal
HW/HL
mck R719 0231 1.234 9889 - 94 86 =
xCU 8950 O15 1.206 992K - 97 BY 87
eri BOOS 0193 1.217 9945 = 98 92 189
HD/HL
mek(s) 7185 0671 1.286 9272 = Jl 58 —
mek(W) 9809 0473 673 9BRS ) 65 64
scu YS12 0280 756 9831 _ 63 62 61
cri 9669 0287 726 9896 0 6% 66 66
142 M. PETRRSON. G M SHEA, GR JOHNSTON & BL MILLER
Behaviour
Two lypes of crrcumiduction and two types af hei
bob were seen. The circummduction types seem ta
correspoud to the clullenge wave and subinissive waive
described by Gratestrom (1971) in Pogone barbara
(Cuvier 1429). The head-bobs differed in cadence und
degree to which the head was moved, vided hy
extension of oe farelinihs. Ino dominant male the head
wis ved & prouter distance below und above te
Horna! plane at a faster cate (hin it was ina subordinate
Hine or twee lenibes.
Two captive mules were observed in an agonistic
display: Upon being, placed in the enclosure fur
jeclimation. they head-hobhed and cireurndueted
several nines before presenting. to each other ata
distance of about 15 cn. They were facing i the same
direetion and continued to head-bob, As the display
intensified they coiled their ails loosely and both did
several hind-lee push-ups, sinilir to those deseribed
tor the ©. dleerevii complex by Gibbons (1979), The
display ended when one Tizard bit the other on the nape.
They rolled vielently wbout the cage and then
soparkted,
Feuiperature preferences
The three C. piekensies siidied maintained body
feiperatures between 11°C and 434°C (x = 34,7°C.
wo = V4l) ino thermal gradient over a 24 bh period
(table 2), There were significant differences in thermal
prelerences between individual lizards (p = 0,04), This
hety have been due we the low variance of RI2267 with
respect oy the other two individuals. Differences
beiween flightlyae and daytime body temperatures
were barely stemfieantly different (p — (05).
However there was 4 large difference in voluntary
muninunr body temperature during photophase (I 1C)
tind scmaphase (287°C). This may reflect yreater
Taiht 2. Comperisen uf thermal prefereitves cimong tree
Co omekensiwiranin a thermal eradien? Measurements are
bya
ty PC,
Lizard n X a Runge
RI2767 129 35.5 Jas 29 24169
R32268 151 344 3b Th. 3¥0
RAD2AY 138 34.5 4.05 34rd 4
Totul 448 44.7 3,4) trei-444
Fagg — 22687, pp = O04.
Moony, SoM. 980) “Phylogenetic amd historpenl
Hugengraphicel relationships atthe genera in tie hinnily
Aguinidae (Repuilie Laeertilin). (PhD thesis. Uriversity
at Michigan).
Wierte. GJ. 982) “Contpurative morphology and
kurvalogy of the Australian micmnbers of the fanily
Agiimitac and reir phylogenetic implicatians® (EHD thesvts,
University i Sydacy)
"
activity during the day. Interestingly, the voluntary:
maximum body temperature (43.4°C) was achiever
during the night. indicating that some nocturnal activity
aecurred,
Discussion
Morphology and relationships
The affinities of C. piekenzied and C. scurnlanis to
other taxa remain uncertain. Pianka (1971) stated,
without providing evidence. that C. seudelelay and C.
rrisrams (Cray (841) were “obviously rather closely
related”, Muudy (980)! added the C. canedleimenis
group to the laticr conmplex, diagnosing Hon the busts
of shape of the medial process of the prearticular bone,
limb length, body size. and presence ofa nuchal crest
and keeled vertebral seale line. Storr (1982). in
resurrecting, Crenopharas, did not place C.
raudieinetus, CL. eristans, C. mekenziet or ©
seuludatay in any ot the species-eroups warhine Ube
venus, Subsequently, Storr ef al, (1983) placed all four
species. Lagether with the C. rereudarus species-group
of Storr (1966) and C. deeresi? species-group al
Houston (1978) in a single expanded Co derresif
species group, noting: thal this was merely an
assemblage, and associated CL nivkencied and ©
seniddanes in an undiagnosed informal subyraup, linked
by inference (Storr eta 83: 32) with C. erivratiuy
Witten (I982*, 1985) placed €. seatlatis with the €
mnaculatuy group und C, cristatuy with C, caudicincher
ina C onsins group, Both groups shared dark ventral
murkings and posterior interscalur femoral and prewnal
pores. The ©. crisratis group was differentiated from
the Co meetdelns group by the more widely spaced
pores, and smooth (vw usually keeled) ventral scales,
‘The presence of a vertebral seale cide and a nuchal
ridge was considered diagnostic tor the CL erivtatie
group, although the presenee of both dark ventral
patie and a vertebral scale ridge in Co seurudetis was
considered primitive within the C. maculatus group.
A vascular tissue block deep ta the vertebral and michal
ridges, possibly functioning in crest erection, was noted
for CL crivtaruy and CL candicinens, but was not found
inthe & macilaras group members examined (which
did not include C scurddarés), Crest erection was noted
for © cristuras, but nol tor C) caudicinenis, We haye
observed nuchal crest erection in both © raadicnecns
(G.M.S.) and Co seatalais (MP. Store eral. 1985_
Plate 2). However, Witten (1982) considered this
vasenlir tissue block to be plesiomorphie within
Crenophorus,
Body, head and limb proportions are similar in ©
rrivtaiia, Co mekeneietand ©, scululafus (Table ') ant
all Share similar male yeotral pattern, a nuchal crest,
and a venebral line of enlarged, strongly keeled scales
the latter unlike members ol the C. maeulathy species
eraup. Consequently, we believe that the Affinities ot
CTENOPHORUS MCKENZIEIT BIOLOGY 243
C. mekenziei and C. scutulatus are with C. cristatus,
although it is clear that a more rigorous cladistic
analysis of the phylogenetic relationships within the
Australian agamid radiation is sorely needed.
Diet
The ant-dominated diet of C. mckenziei is similar
to that reported for other small Crenophorus species
(Pianka 1986; Baverstock 1979; Mitchell 1973).
Although Jridomyrmex spp. were the most commonly
eaten ants, their dominance may simply reflect
availability rather than any selection by the lizards (E.
Matthews, pers. comm.).
Behaviour
The behaviour reported here for C. mckenziei has
been observed in other species of Australian agamids.
The hind-leg push-up display was thought by Gibbons
(1979) to be unique to the C. decresii complex. The
observations presented here show that this 1s clearly
not the case. Whether this behaviour is homologous
in C. mckenziet and the C. decresii group is unclear.
Thermal preferences
The mean body temperature of C. mckenziei reported
here is lower than that reported tor C. scutulatus in
a laboratory gradient by Licht er al. (1966). It is unclear
whether this is a real difference or an artefact of
conditions which the animals experience during
acclimation or while in the thermal gradient. It is
noteworthy that Licht ef al. (1966) used a thermal
gradient in which the minimum temperature available
was 25°C, considerably above the voluntary minimum
body temperatures experienced by two of the three C,
mekenziel tested.
While the observation that the voluntary maximum
was recorded at night may at first seem unusual in an
animal belonging to a group traditionally thought of
as diurnal heliotherms, this is not really so, Several
species of agamid are known to exhibit some nocturnal
activity when thermal conditions allow (Fyfe 1981;
Morley & Morley 1985; Bedford 199]; G.R.J., G.M.S.,
pers. obs.). A constant source of heat in a thermal
yradient probably presents as near optimal conditions
for nocturnal activity of agamids as possible. However,
the low variance exhibited by C. mckenziei at night
(1.86 vs 6.22) indicates that nocturnal activity is limited.
Comparative material examined
(all localities in Western Australia)
C. cristatus: WAM R41827, 2 mi SW Wahlyamoning
Rock; R68001-04, R68023-24, 4 km SW Lake Cronin;
R68005, R68029, Lake Cronin; R68006, 2.6 km SW
Lake Cronin; R68021-22, 5 km SW Lake Cronin;
R70707, Frank Hann National Park; R71833, 19.5 km
78° Toomey Hills.
C. scutulatus: SAM R1459a-j, R3024a-b, R4814a-!
(syntypes), between Fraser Range and Queen Victoria
Springs; WAM RI1235, RI761, Laverton; R2841, Gutha:
R5306, Wadgingarra; R8170-71, “Yuin”; R9352.
Malcolm; R9510, Morowa; RI1236, Carnarvon;
R12209, Shark Bay; R21865-67, Caron; R48385-89,
40 km N Beacon; R53551, 15 km E Point Sunday;
R59605-06, 20 km ENE “Meadow” HS: R86769,
14 km WNW Mallee Hen Rocks.
Acknowledgments
T. Schwaner, A. Edwards and L. A. Smith allowed
us access to specimens in their care, and in the latter
case provided locality data for C. scutulatus, E.
Matthews identified the stomach contents. B. T. Firth
permitted use of his thermal gradients. B. Coulson ts
thanked for field assistance.
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244 M. PETERSON, G. M. SHEA, G. R, JOHNSTON & B. MILLER
& Rorinson, A. C. (1987) Ibid.
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REPRODUCTION AND GROWTH OF THE SMOOTH PEBBLE
CRAB PHILYRA LAEVIS (BELL 1855) AT TWO SITES IN
SOUTH AUSTRALIA DURING 1990-91.
By STEPHEN C. MCKILLUP* & RuTH V. MCKILLuP*
Summary
McKillup, $. C. & McKillup, R. V. (1994) Reproduction and growth of the smooth
pebble crab Philyra laevis (Bell 1855) at two sites in South Australia during 1990-91.
Trans. R. Soc. S. Aust. 118(4), 245-251, 30 November, 1994.
The growth and life history of the smooth pebble crab Philyra laevis (Bell) was
inferred from collections made between July 1990 and December 1991 from Sultana
Point and Coobowie, South Australia. P. laevis appeared to be semelparous and to
reproduce twice a year at both sites. At Sultana Point ovigerous females were
common in winter (June-July) and from late spring to late summer (November-
February). Dead adults were common from late winter to early spring and also in
early summer. Distinct cohorts of new recruits were found in early spring and again in
early summer. Recruits grew rapidly, reaching sexual maturity within four months.
The mean size of adult females and the proportion which were ovigerous differed
amongst populations. A laboratory experiment showed that food availability affected
growth and whether females became ovigerous. We postulate that differences in size
and ovigery amongst populations of Philyra laevis may be caused by differences in
the availability of food.
Key Words: Philyra, pebble crab, life-history, intertidal, food, sandflat.
Trans tions vf the Reval Society of 8 Anse (294), HRC. 245-251
REPRODUCTION AND GROWTH OF THE SMOOTH PEBBLE CRAB PHILYRA LAEVIS
(Bell 1855) AT TWO SITES IN SOUTH AUSTRALIA DURING 1990-91.
by STEPHEN C. McKituup* & RUTH V. McKILLup»
Summary
Mekieuup, SC. & Mekituup, R, V, (1994) Reproduction and growth of the smooth pebble crab Philyea laevis
(Bell 1855) al two sites in South Australia during 1990-91, Tras. R. Soo. 8. Aust, 184), 245-251, 30 Novernber,
1994.
The growth jd He history of the smpoth pebble crab Pitre laevis (Bell) was interred trom volluchions made
between July (990 and December }99f from Sultana Point and Coobuwie, Suuth Australia. Po Jaeos appeared
t@ be semelparous und to reproduce tWice a year at both sites, At Sultana Point ovierous females were commen
in Woner June-July) sind from late spring to late summer (November February), Dead adults were common tron
lute winter to early spring and also in carly summer, Distinct cohorts of new recruits were found w curly spring
and again in vayly summer. Recruits grew rapidly, reaching sexual maturity within four months, The mean size
of adull fermiles and the proportion which were ovigerous differed amongst popalations A laborstory experiment
showed that food availability affected growth and whether females became ovigerous, We postuliite (huc differences
Wn size and ovigery amongst populations of Philyra aeviy may be caused by differences 1n the availability nf food,
Key Worps: Philyru. pebble crab, lifethistory, ineertigal, loud. sandflat
Introduction
The smooth pebble crab, Philyrir laevis (Bell 1855},
is common on sheltered intertidal sandflats in southem
Western Australia, South Australia, Tasmania and
Victoria (Phillips ez al. 1984). Hale (1976) describes
ihe feeding and courtship behaviour of Po laeviy, but
tne else is known aboul this crab. Tasimaman
populations of B laeviv have been reported to host a
herertean (genus Carcinonemertes) (Bell & Hickman
1985) ancl the trematode Microphallus paragrupst
Smith. 1983 (Bell 1988). From 1990-91 we studied the
feeding behaviour of Philyra laevis, tinding that
although individuals often ted upon other dead or
Jamaged animals. many were unwilling to feed upon
members of their own species and thal Water borne
coes from damaged P laevis inhibited feeding
(McKillup & McKillup 1992), During the study. we
collected P laevis over an J8-month period from twa
sites 10 kilometres apart in South Australia, and also
sampled 10 other southern Australian sites. These data.
logether with results of a laboratory teeding
experiment, are used to infer the Life history of P laevis.
Methods
Sampling of Philyrs laevis from Sultana Point wnrtd
Coahowie
Samples of Phi/vra laevis were collected from the
intertidal sandflat at Sultana Point. lower Yorke
Peninsula, South Australia (35.08°S 137.44°6) in July
1990, and then approximately every four weeks from
September 1990 to August 1991 and in December J991,
* Department of Zoology, University of Adelaide, South
Australia, S005 Australia,
* Present address: Department of Bivlogy, Central
Queenshund University, Rockhampton, Queensland 4702
Australia
The sandflat at Coobowie Bay, 10 kim north of Sultana
Point, was sampled inthe same way but less frequently
(November 1990 und January, Maroh. April. August
and December 1991)
P. laevis is active Whilst the sandflat is cavered by
water during ebb and rising tides, bul remains buried
in the substratum at high water or whea fhe sandfat
is completely exposed (Hale 1976). At least 30 and
usually more than 100 individuals were collected from
within. the intertidal zone where Po laevis occurred
(From low water tw about mid tidal level) between the
time of luw water and when the sanudiut was inundated
by the rising tide, At each site ut least 160m" of
substratum, consisting of several haphazardly chosen
0.5 metre wide strips from the water's edge to mid tide
level, were hand searched to a depth of 35 em, P lacvis
was found no deeper than 3.0 om in the substratum
(McKillup & McKillup unpubl). Crabs were frozery
and Jaler examined for sca, Whether females were
ovigerous, and the carapace widlh of all individuals
was measured to the nearest 0.05 mim.
Reproductive condition wid average size af uduels
femules at additional sites
During January and February 1991, at least 75
Philvra laevis were collected from each of J] sites in
South Australia and one in Victoria (Swan Bay within
Port Philip Buy), and examined for sex. size and
reproductive state ds described previously.
Laboratary experimen on sexual development anid
growth
Abdominal morphology i the Crustavea.can usually
be used to determine sex: adult males often have a
relatively narrow, concave sided and lapered abdomen,
whilst females have a broader and often circular
abdoinen almost as wide as the carapace (Hartnoll
Ty so
Dip 1. Abdominal morphatagy of Philera laevis: From lett
Wright male, intermediate ( = juvenile fermalerunel aul
tumale
(982). This was the case for all Pid deevis with
carapuces wider than B.5 mm, but three morphological
lypes Were found in crabs. 13,5 mm wide or smaller:
the two previously deseribed forms, plus andayiduals
with a ocomvex Sided abdomen whieh, at its widest
paint, Was approximately hall the width of the varapace
(Fig. Lo The last described individuals were named
“intermediates
The growth of 20 females, 20 intey mediates and 20
males. all of carapace width 10,5 mm ar less, was
Observed in the laboratury Considering that we alsa
folind differences in the size and proportion wl adil
lernales which were ovigerous amongst sites, and that
differences im adult size amongst (uaspeeified)
loealities were alsa reported by Hale (1976),
pbservutrons Were made as part of a Manipulative
luboradory experiment desrgned lt exumine the efleets
Of load availability upon growih and egg production
of Piva laevis. Crabs were placed mdividually in
30 x 50.4 30 nin deep plustic dishes, cach filled with
40 inl of seawater, All individuals were numbered on
the dorsal side of their carapace with non-taxic
waterproof ink. Six dishes, cautamning two mules. two
fitrules and two intermediates were placed within vuch
of 10 fidded 280 x 380 >< LIO mm deep plistic trays
conklining seawaler (oa depth of IO mim. Aeration was
not needed, The seawater in the trays mitintained a high
level oP homidity whieh reduced evaporition from the
dishes undalse provided.a marina environment for the
hiew crabs which climbed out during, the experiment.
Trays were Kept Ut roon) (emperatire aad meduiral day
iength
Five (rayé were assigned randomly toa “high food”
treatment and the 30 crabs within these were fed every
working day Gront Monday to Friday), whilst the 30
vrabs in the wither five trays were fed weekly, on
Wednesdays, as (“lew food” teeatiient. Dillerent
(requencres OF feeding provide different levels of tood
availabilty to Wivertebrates (cou. Calow 1973; Moriarty
78), For euch feeding the six crabs in ua tray were
removed, pluced in sik sepunue dishes used for feeding
only and offered ae ibirum crushed voukle. Autelvsia
Sealaring Luniarek. frond Sultana Point Bo faevis is
oftet. rund feccing on this bivalve ip the field
(McKillup & Melci)lup 1992), All crabs fed for L2 min
vir less Und Were returned to thea permanent dishes
McKIETUP & R, Vo MeRILLUP
afler 15 min. This method of feeding prevented the
seuwaler in the permanent dishes from beeomlae
fouled, it was replaced fortnighily: Crabs were
inspecled on every working day, individuals whieh had
tmoulted were examined und sexed as either male,
female or mtermediates, and uny which had eliimbedt
Aut were rephiced ip (hele dishes:
The experiment began on April 2nd (991 using new
recruits trom {he Deceniber 1990 - January 1991 cohort
it Coobowie and eontinued unui all erabs had moultedt
ul least onee.
Results
Sampling of Philyra laevis prom Sultana Point ame
Chabhawre
The size structure at {he population of Philyre laevis
wl Sullana Point From July 1990 co December 1991 ts
iMusteated im Figs 2. 3. In mid-winter (luly 1990), ull
mudles Were at leust RS mm wide, all temalesat least
TS mm wide and 93% of the later were ovigerous,
By early spring (September 1990), dead males and
lemules were extremely eommon and large live
individuals were nots almost half of the live mules
culleeted Were sinaller than 8.5 tm and appeared ta
be @ distingt cohort of recent recruits, Only two live
aduil females were found; both were more than 7.5 min
wide and ovigerous. The remainder of the sample
consisted of “intermediates” 7.5 mim wide var smaller,
which were not present in the July sample, A
laboratory experiment showed that intermediates were
juvenile fenrales (see below). In mid-spring (October
1990) only ope cohurlof miles was present, and these
males Were hirper than the recruits first scen in the
previous month, The juvenile tenales were also larger
and sore relanvely small adult females were present.
bul none was ovigerous, In late spring, (November
1990), no juvenile lemales were found; the sample
consisted ol adult femules ut lenst 9,5 min wide, 89%,
of Which Were Pvigerous, plus males from 4.5 to
15.5 mim wide, In early summer (December [990)
another cohort of relatively soil males was present.
together with a cohort of juvenile females up w-9.5: mun
wide. Dead adults were uguin common, but live sdules
were also found and 71% of adult females were
ovigerous. The small tales and juvenile females
continued to grow through summer and autumn
(January lo April 199]) and the number of ovigerous
adult females declined (30.6% in January. 14% in
February and 5.6% in Mureh), From mid- to late-
autumo (April and May 9) no juvenile females were
found and only LS% and 2% respectively of adult
femiles were ovigerdus, but inearly winter Gune 1991),
the percentage of oViperous feinales hud risen to 68%
“uid further inereased to 94% by late winter Guly 199)
The early spring (August 199]) sample was very similar
(y that of Seplember 1990: dead adult nades and
REPRODUCTION AND GROWTH OF PHILYRA LAEVIS 247
30
30
3th July 90 T9th Jan “91 15th June *91
30 30
7th Sep °90 23rd Feb °91
23rd July “91
0 is}
0 14.5 27.5 a
30
éth Oct *90 7th Mar '9) 18th Aug ‘91
1S
0
a
145 275
30
3rd Nov °90 13th Apr °9] 19th Dec ‘91
18
ie)
0 14,5 27.5
SSDIO OZIS UODDS UI SQDID JO IOQUUNN
8th Dec *90
25th May ‘91
0 14.5 27.5 ie} 145 27.5
Carapace width (mm)
Fig. 2. The size structure of female Philyra laevis collected from Sultana Point, South Australia on 14 occasions between
July 1990 and December 1991. Solid bars indicate juveniles; open bars, adults.
248 S.C. McKILLUP & R. V. McKILLUP
30
13th July '90 19th Jan “91 Sth June *9}
0 14.5 27.5 a 14.5 275
17th Sep ‘90 23rd Feb 9] 23rd July ‘91
0 14.5 275 0 14.5 27.5
6th Oct “90 7th Mar ‘91 18th Aug 91
e) 14.5 27.5
13th Apr ‘91 19th Dec 91
SSDIO SZIS YODS U! SQDID jo sequuNYA
8th Dec ‘90
0 14.5 27.5
Carapace width (mm)
Fig. 3. The size structure of male Philyra laevis collected from Sultana Point, South Australia on 14 occasions between
July 1990 and December 1991.
REPRODUCTION AND GROWTH OF PHILYRA LAEVIS
females were common and the sample of live Philvra
levyiy consisted entirely of juvenile females less than
8.5 mm wideand males which were almost all smaller
than those collected during the previous month. The
December 199] sample was similar to the one from
the previous December, consisting of adult males plus
adult and sonie juvenile females, with 65% of adult
females ovigerous, Dead adults were again common
on the sandflat at Sultana Point in December 199].
The data for Coobowie were consistent with those
from Sultana Point. In late spring (November 20th
1990) only males and adult females were found, but
in mid summer (January 19th 1991) juvenile lemitles
were also present and dead males and adult females
were common. The average carapace width of the
249
cohort of juvenile females present in early autumn
(March 17th 1991) was larger than in January and most
appeared lo have moulted to-adults by April 13th 1991,
In late winter (August 18th 1991) dead adalts were
common and juvenile females plus small males were
present. In carly summer (December 19th 1991) dead
adults were also found, very few live adult females
were present and juvenile females were common.
The mean carapace width of adult females was
always greater at Sultana Point than Coobowie (Table
1) and a greater proportion of adult fernales was
ovigerous at Sultana Point than Coobowie on five
occasions when comparison was possible (Table 2).
Males were not compared since it was impossible to
distinguish between juveniles and adults.
TABLE |. Coniparison of the mean carapace width (in nim) of adult female Philyra laevis from Coobowte and Sultana Paint.
The November collections were made 17 days uparl; on the other 5 vecasions crabs were collected from both sites on the
sume day, ¥ = mean, x, = standard deviation and n = sample size
Site
Coobowie Suliana Point
Date collected a x Sy n x Sy
November [990 33 9.23 2.37 125 13.17 2.02
19 January 1991 17 12,74 175 49 13.14 173
\7 March 1941 52 12.62 }.42 71 13.68 1.72
13 April 1991 85 13.18 i.75 65 14.19 1.77
IK August 199] 23 11.82 1.94 none found
IY December 1991 9 11,72 1.72 22 13.50 1.83
Tashi 2. The number of adult female Philyra laevis and those ovigerous in samples collected from Coubowie and Suliane
Paint on 6 accasions during the same month. The November collecuoms were made 17 days apart, on the other 5 aecasions
crabs were collected Jram bath sites on the same dity.
Site
Canobowie Suluina Point
Date collected Collected Ovigerous Collected Ovigerous
November 1990) 33 ! 128 119
19 January 1991 {7 0 49 15
7 March 1991 52 0 71 4
13 April 199] RQ 0 65 2
18 August 1991 22 {5 0 -
1) December IYI 9 0 22 {4
Tanie 3. The percentage of ovigerbus adult female Philyra laevis ar /2 sites. sampled during January and February 1991,
twuether with the meun carapace width of adult females from each site.
Number of Number and Mean varupace
mature females percentage width of
Dute sampled Site collected ovigerous females (nm)
19 January [991 Edithburgh Bay 9 0 (0) 12,67
19 January 1991 Sultana Point 49 15 (31) 13.19
19 January 199] Coobowie 7 0 (0) 12.74
19 January 1991 Hickey's Point 25 J (8) 13.10
19 January W9L Stansbury 33 7 (21) 14.35
20 January 194] Pout Turton 3 0 (Q) 14.83
20 January l991 Rogues Point hin 6 (7) 13.90
29 January 1991 James Well 20 ] (5) 12,65
29 January 199] Pine Point a] ] (4) 12.33
2 February 1991 Foul Bay 87 0 (0) 13.89
2 February 199] Sturt Bay 70 10) (14) 13.10
13 February 1991 Swan Bay (Vic.i 66 56 (85) 17.46
150 SC MiRILLUP& Ro Vv. McKILLUP
Reproductive condition and average size of females at
additiunal sites
Data Jor the number of adult females calleeted and
the number and percentage which were ovigerous lof
12 populations sampled in mid ty late summer 1941
are in Table 3, ‘The percentage of ovigerous females
decreaped wt Sultana Point from January i February,
so only samples collected on the 19-20th January 199]
were compared stutisticully. Nevertheless, the
propartion ovigerous diflered significantly amongst (he
seven sites sampled on lower Yorke Peninsula (2 x 7
voningeney table comparison: di = 6, Chi-syquared
statistic = 22.26 P< 0.005) The significant
heterogeneity amangs! sites was largely dae We the
greater proportion of ovigerous females at Sultana Poinr
(Table 3) Considering all sites sampled. the highest
perceniage of ovigerous adult females (85%) Was al
Swan Buy, Vietoria during early February (wheu only
4% were ayipetous al Sultana Point; see cartier
discussion of Figure 1), The Swan Bay population also
contained the largest Philyra laewis ound.
Furthermore, for the seven sites sampled trom the
9 20th Fanuary (excluding Pome Turton where only
three females were collected), ovigerous females were
found oly at sites Where the average carapace width
Of femules was greater than 13.00 mm (Table 3)-
Laboraiory experiment an sexual development
Male, adult temate and “intermediate” Phitvra laevis
ollered Foad on five af seven days per week moulted
sooner than those only fed once per week (Table 4),
At thear first moult in the laboratory all males moulted
to mates, all fernales (o females und all intermediates
Io females, except for the two smallest which remained
as the intermediite form unl they moulted again. In
all cases the Variance of days clapsing before moulting
was prealer in the low food treatinent, and by
imspeotion the distributions in this weatrnent were
skewed to the sight. None of the adult females in the
low Jood treatment, bucall in the high food treatment,
wer ovigerous by July 1991.
Diseussion
Philvra laevry reproduced Fwite a yeur at Sultana
Point and death of most adults during the breeding
Months sugpests Uns species ts largely semelparaus
Reeruits found in carly spring. (September 1990)
reached sexual maturity and reproduced trom. late
spring ta date summer (November 1990 to February
91). while those first found in early summer
(Decernber 1990) reached sexual maturity by nid
autumn (April 1991) and reproduced until early winter
(June (981). Data from Coobowie were consistent with
this pattern of recruitment, but suggest that many adult
P lawvis af Coobowie did not reproduce in the summer
of 1990-91, although dati were scanty, being only foe
November 1990 and January 1991,
Reeruits were found one mouth after reprodueiag
females were present in late spring and two months
after they were present in carly winter, Considering
(hat the sandflat was only sarmpled monthly. (hat the
smallest crab found was 2.8 nim (MeKillup &
McKillup unpubl.) and that smaller individuals were
likely io be overlooked amongst sand grains and
detritus, the larval stage of A laevis is likely to be of
telulively short duration (perhaps only 2 weeks),
Another member of the same genus, the purse crab
Philyri glohasa (Rabricius), has a larval stage lastine
I) days in the laboratory at an average temperature of
26°C (Krishnan & Kannupandi 1990),
The moult from juvenile to adult form ia females
appears to coincide with sexual maturity, sinee only
iwo ferniles with juvenile ahdomens of more than 300
examined Were ovigerous (McKillup & McKallup
unpubl.j. A relative (non-allometric’) inerease in
abdomen.compared to curapace width during the moult
to adulthood is common in bruchyurans (Hartnoll
1974),
Por sites sampled trom the 19-20th January 1991,
3 greater proportion of ovigerous adult fernales was
present where the mean carapace width of females was
relatively large. Also, there Was u greater proportion
of ovigerous females at Sultana Pojnt than Coobowie
on all oceasigns when comparison was possible,
Differences amongst sites were not caused simply by
larger females being more likely to be ovigeraus:
cxammmation af the data ised to compile Table 2 showed
thut in Novenrber, 60 of 69 adult females between ¥
dnd (3mm wide were ovigerous at Sultana Pornt, but
none of 19 gollected on the same diate und within the
sume size range was ovigerous al Coobowie.
Furthermore. on January (9th 199], pone of the (7 adule
females collected [rom Coobowie was ovigerous. even
Though the carapace widths of these mdividuals were
VABLE 4) The mean dayy elapsing befare Philyva laevis firs¢ multed in tiie and law food treamnents, 9 mean
5) 7 setnelerd deviititan, ae = sample size
Treaimeni
Plinth) Pood Low Fool
n x Sy tl = ty
Miles iT) 0 3.78 ti 29:10 17.4
Pemnales fo 23.5 7-38 1g 45.20 35:34)
Intermediates Itt 2m in 7a i 38.10 TKI
REPRODUCTION AND GROWTH OF PHILYR A Lahey |
within the size cange of the 1S ovigerous lenales
vollected frum Sultana Pot. Similarly, on December
17th 1991, none of Qadult females from Coobowie was
ovigerous. even though six were Within the size range
of ovigerous females from Sultana Point,
Results of the laboratory experiment were consistent
with food supply affecting the frequeney of moulting,
A sumple of crabs will contain individuals at different
stages Of the moult evele- In the low toad proup, erubs
uboul to moult would have done so svon after the
experiment began, but those which had moulted shortly
before being calleeted would have to prepare to moult
again under the laboratory conditions. of low foad
uvallability, In contrast, to the high food group, crabs
prepared to moult would do so. and others which had
tecently moantted would have adequate resourees
available to prow and moult again. ‘This ean explain
the greater mean, variance and positive skew of the
number of days before moulting in the low compured
io the high food treatment (Table 4).
We suggest, for the following reasons, thal
differences in size and the proportion of ovigerous adult
leniales adionyst sites were Caused. by differences in
food availability; Firstly. in the laboratory, crabs 1 the
high tood treatment moulted sooner (and therefore
grew faster) than those in the low food treatment, und
only females ti the high food treatinent produced eggs.
Secondly, adult female Po laevis al Coobowie were
consistendy sinaller than those at Sultana Point ‘The
intertidal scavenger Newseriuy pauperatus (Lanvarek),
4 prosobranch snail whieh oecupies a similar niche and
i Offen found fveding with Philyra laevis, is also
smaller (and hungrier) at Coobowie than al Sultana
Point. and we have postuluted there is less food
ivallable oN, panperaius at Coobowile (banat Sultans
Point (Mek illup & Butler 1979, 1983; MeKillup 1983)
Furthermore. we have more recently postuliuted thut
intertidal scavengers nay penerally be short of foou
(McKallup & Mekidlip 1994). All South Austrilion
populations of Po laevis sampled contained smaller
females on average than the populatian at Swan Buy
Victoria, sugvesting that load availability tay be
limiting the growth and reproduction of many
populations. of this scavenger, Notably, females of
another feucositd erat. Abadia laevie( Bell, 1835) were
only found ovigerous during December January, May
and Aupust at Wellington, New Zealand, bul records
tram othec logabitles sugeest the duration of he
reproductive season yartes amongst siles (Wear de
Fielder 1985).
The hypotheses that differences in fd availability
are limiting the duration of the reproductive pertad of
Pirilvra fereviy and (hat individuals im natural
populations of this species are short of food could be
tested by frequently feeding marked individuals in the
field and comparing their growth und reproductive
Output With the rest of the population, The results of
these experiments may explain why Phifyra laevis has
arelatively short Hiespan and two seasonally opposed
breeding seasons in South Australi.
Acknowledgments
We wish ta think Alun Buller und Keith Walker for
theirencouragement during the study, and Alan Butler,
Michael Coates, Don Flelder and Alice Wells for their
erideal and constructive coulments on a draft of (he
manuseript.
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laryal development pf the purse crab Philyra glebpa
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Wa, 171-182.
Mesitue, 50. (983) A behavinural palynyorphisny in the
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THE HISTORY OF THE DEVELOPMENT OF THE
PACIFIC OYSTER, CRASSOSTREA GIGAS (THUNBERG)
INDUSTRY IN SOUTH AUSTRALIA
By A. M. OLSEN*
Summary
Olsen, A. M. (1994) The history of the development of the Pacific oyster Crassostrea
gigas (Thunberg) industry in South Australia. Trans. R. Soc. S. Aust. 118(4) 253-259,
30 November, 1994.
The dredge fishery for the native mud oyster Ostrea angasi Sowerby in South
Australia had a chequered history before finally collapsing in 1945. Attempts to
cultivate the native oyster on leases were only partially successful. The importation of
seed oysters of the Pacific oyster Crassostrea gigas from Japan in April 1970 by a
private company began the successful aquaculture of this rock oyster. The aquaculture
of the Pacific oyster has filled the market niche left by the loss of the mud oyster
fishery.
The background history of the several introductions of seed oysters (spat) from Japan,
Tasmania and Scotland is described.
Key Words: Pacific oyster, Crassostrea gigas, history, introduction, South Australia.
Tranvoiions af the Raval Sacien af S Aus W994), URE), 289 259
THE HISTORY OF THE DEVELOPMENT OF THE PACIFIC OYSTER,
CRASSOSTREA GIGAS (THUNBERG) INDUSTRY IN SOUTH AUSTRALIA
by A. M. OLSEN*
Summary
Oisen, A.M, (1994) The history of the deveiopment of the Pacific oysler Chassuyired pigas (Thunberg) nidustry
im South Austraha Trumy. R. Sae 3. Aust. VB(4) 253-259, 30 November, 1994,
‘The dredge fishery for the native mud oyster Oserew engaat Sowerby in South Austaidtia haul a chequered listory
before fmally collapsing in 1945. Atremipts to cultivate Wie native oyster on leases were Only partially Successful,
The importation of seed oysters of the Pavific oysler Crassastreu gigas from Japan in Apal 1970 by a private
vompany hegan the successful aquaculture of this rock oyster, Phe aquaculture of (he Pavifie oyster has filled
ihe market niche left by the Joss of the mud oyster fishery
The back#round history of the several introductions of seed Oysters (ypat) from Japan. Tasmania and Seotkand
is described
KrY Words: Paeifi¢ oyster, Crassasnea gigas, history. introduction. Seat Australia
Introduction
From the early days of (he colony, oysters were being
markeled from beds on the western side of Gull St
Vincent and the northern side of Kangaroo Island. By
!¥7t the dredge fishery was concentrating on oyster
beds in Kellidie Bay where 30 sailing vessels employing
80 men were dredging 60 thousand bushels of oysters
a year, Overtishing of the stocks occurred and by the
late 1880s the feet maved to new beds near Stansbury.
Withura few years these beds were also depleted and
the few remaining vessels moved back t the Eyre
Peninsulit beds so that by IY0S the nud oyster Hshery
had virtually collapsed, Over Ihe next 25 years, when
old beds showed signs of recovery boats entered the
fishery only io leave Liter. In 1945 the mud oyster
fishery Ceased to exist (Olsen and Priest 1971; Wallace-
Carter 1987),
Attempts to cultivate the native oyster at Stansbury
and Kellidie Bay were only partially successtul with
the -ourput limited by low recruitment. Mud
(incubatory) oysters have a low fecundity compared
with that of the mass spawnings and spal settlement
of the rock (nen-incubatory) oysters such as Succesrea
cnmmercialis (Iredale and Roughley), the
commercially important Sydney rock oyster und the
introduced Pacific oyster Crassosrrea givas.
Early attempts to grow rock oysters
in southern Australia
An attempt in 1886-7 to acclimatise the Sydney rock
oyster at Hobart, Tasmania was unsuecesstul (Saville-
Kent 1887). A trial shipment of the same species spread
#1 Orchard Grove, Newton, South Australia. 5074
Amon, W934) Large seule farming of Sydney rock Uysiom
in Port River. "The Saturday Mail” {4 Juty, (93d.
out on trays ul Kelso. River Tamar, in northern
‘Tasmania in 1948 by a Sydney rock oyster grower failed
too (Olsen 1965),
In 1933-4 altlempts were made to cultivate young
Sydney Rock oysters on a commercial scale in South
Australian waters, 228,000 oysters were hid gut om
trays off the west bank of the Port River below the
Osborne Power Station where am earlier experimental
consignment had reportedly grown very fast (Anon,
1934'; Wallace-Carter 1987),, Other attempts to
acclimutise the Sydney rock oysters were made in 1937
at Mt Dutton and Kellidie Bays without success,
A-small consignment of juvenile Sydney rock oysters
from a warm water environment at a solar saltfield at
Port Alma, Queensland was transterred in May 1976
int) seawaler of approxiniately similar salinity and
temperature of the primary pond of the solar saltfield
at Dry Creek. Within 3° months 95% of the
expetunental consignment died, Later the same year
a second experimental Consignment was transferred
from the same source and these oysters suffered 70%
mortality within a month. As a result of these
experiments turher transloecauion experiments into
quarantine areas. at Dry Creek ceased (Melvin per
comm.},
So far us the wuthor is joware the last atlempl ty
acclimatise Sydney rock oysters in South Australia was
in 1977 when aysters put into the commercial prawa
farm ponds at Port Broughton failed.
Thomson (1952) remarked that conditions of the
waters of the soulbern stales are guiside the natural
range of the Sydney rock ayster,
When CSIRO Division of Fisheries and
Qcéanography wnported Pacitie oysters trom Japan
between 1947 and 1952, the author assisted with their
establishment-and supervision and. in September 1953,
the transfer of the majority of the surviving stoek to
Port Sorell. [t was on the basis of this experience that
25¢4 A M OLSEN
the author m 1968 considered the aquaculture of the
Pacific oyster would be successful in South Australia,
The Pacific oyster wits already established as a viable
reproducing stock on the rocky foreshore of the River
Tamar near Sidmouth. in Tasmania (Thomson 1959).
By 1964 the Tasmanian authorities were granting leases
lor tls commercial production, (Fig. 1)
AL its January 24, 1968 meeting, the Flora and Fauna
Advisory Committee. a body responsible to the
Minister for Agriculture, South Australia, examined
a proposal from the Director of Fisheries and Fauna
Conservation Department for Pacific oysters to be
introduced from Tasmania for aquaculture in South
Australian waters. [It was argued that the ‘Tasmanian
stock was disease-free and its introduction could fill
a niche no longer occupied by the native mud oysters.
The Committee did not raise any objections to the
recommendation. The Department itself did not have
the staff or funds to undertake the import and
establishment of the Pacific oyster but wanted to be
able to encourage private enterprise to do so.
In June 1968, J. T. Belling of Balaklava asked the
Department whether un oyster lease, currently held by
W. Lee of Kellidie Bay, could be transferred to another
company of seven shareholders wha wished lo expand
production of the native mud oyster. There was. no
objection and subsequently Oyster Farmers Coffin Bay
Pty Ltd was registered as lessee of the Kellidie Bay
oyster lease,
Later in the year two Adelaide men, C, J, Mack and
R. C. Sprigg, also approached the Department about
obtaining leases for experimental oyster produchion.
The author suggested (hey meet J. T. Belling to discuss
the problems associated with cultivation of inud oysters.
He also suggested that they might like to combine and
import a consignment of Pacific oysters from Tasmania,
The group was advised to visit Wivell Bros’ Tasmanian
Rock Oyster Company at Sidmouth, River Tamar, 1
see the techniques used there and to find out about the
transport of oysters to Adelaide. After a visit by the
men to Wivell Bros in March 1969, an order was placed
with the company for 50 bags, each containing about
120 adult oysters for delivery to South Australia ut
September 1969.
After their return, arrangements were made with
R. C, Sprigg’s organisation, Geosurveys Ply Ltd for
its workshop to make and tar oyster trays. Tarred trays
were taken to Sprigg’s site at Coobowie and Mack's
at Kangaroo Island which had been preselected so that
the trays could be attached to stakes, above the sea
floor, bul situated below water level at low tide.
The 15 bags purchased for Mack’s lease were set out
above u seagrass bed (Zostera muelleri) at 0, | and 2m
below low water near Pienic Point, American River,
Kangaroo Island and Sprigg’s were set out below low
water off the Coobowie Marine Research Station
Yorke Peninsula, The remaining 20 bays Were taken
by track and laid out on the Oyster Farmers Coffin
Bay Pty Ltd lease at Kellidie Bay, Eyre Peninsula.
The adult oysters all acclimatised and grew well,
Al(hough they spawned three months afler being set
out no spat settled at or near any of the three leases.
Buss Strait
Pon Sarelt™ VK
Sidmouth \ River Tamar
LT
Scale (km)
\
is
Launeedston
Tasmania
Fig. 1. Locations in Tasmania of spawning wild populations of Pacific oysters.
EARLY DEVELOPMENT OF PACIFIC OYSTER INDUSTRY 255
Environmental conditions including High salinities
(35 ppt) must have been immical to the survival and
aclemen! of faryae of the Pacific oyster Many
commercial hatcheries operale ul salinities of 3244 ppt
of even 36 ppt and sulinilies of 30-34 ppt are reported
from sced- producing areas in Japan and fram areas of
Franee where C. gigs reproduces suceessiully
(Coleman, 986). However, continuous exposure (a
salinities grealer than 42 ppt is lethal to Paciiie oyster
larvae und aceording to Medcof and Wolf (975) this
isthe reason there has been no Spat fall in Coffin Bay,
South Australia,
The sustained growth and excellent conditjon of the
imported adults on the Oyster Purmers lease prompled
the Company to consider importing Pacific oyster seed
(spat) hy dir from Japan. One of the shareholders, who
wrote and spoke Japanese fluently, flew to Japan ter
examine cultivation techniques first hand. While at
Sendai. Myagi Province. be sought details of air-
freighung oyster Spal Adelaide, Pull detuils of queility.
quantity and costs from. Sendai to Adelaide were
reeeryeu! on Febriary 23, 1970,
[portation of seed oysters (spat)
In early March an application ta import a ural
shipment of spat by air from Japan was lodged wath
the Chief Quarantine Officer (Animals), South
Australia. Permission ty import the spat was rece Ved
on Murch 24, sabjeet lo conpliance with al) quarantine
requirements. A 19 kg hox containing 6000 Grade vA
unbroken oyster sput was ordered tu be wu-Treighted
to Adelaide and was received on April 12. Some miiumir
problems wath quarantine requircments were resolved
quickly and the spat was released in puwod condition
from quarantine on April 28 und set out on the Kellidie
Bay leuse.
A second application from Oyster Farmers Cottin
Bay to import 100,000 certilted cisease-tree oyster spat
hy air from Mitsui and Co. Lid. Sendai. was lodged
with the Department of Health, Canbenrs in Aueust
(970 (Pig, 2), This was cheaper than buying and
(transporting the spat by sea from Tasmania,
In the second application the company wrote “Ihe
growth cate oF this 970 seed is proving ro be excellent
und we. therefore. wish to further Gurexpenments wilh
the rate of growth for production with view. to sale.
I is anticipated that 100,000 seed be imported in. 107)
with follow-up quantities as and when considered
appropriate, Our next consigninent will be by air,
however, we request permits lo coyer both ATR and
SEA Cruispart so that seed mortality during transit may
be compared.”
The upplication was refused by the Cammunwealth-
Sates Advisory Conunittee in Febriary 197) The
“Rescue. W. 1972) The ovstermen at Colin Bay. ~Sutdiy
Mail> 32 Apel. 172.
grounds lor rellsal were qever made Known despite
a sequess for such from the South Australtian
Department. The intraduction of cerited disease-free
oyster spat direct from Japan to South Australia ced
nul pose w disease threat ty the NSW oyster bur the
development of un oyster industry did pose a potential
competitor lor the SSW conmnerciul oyster monapaly.
Aller the refusal to alloy §pat lar be jmported trom
Japan, Oyster Farmers Cotlin Bay ordered 2000 “sticks”
from Wivell Bros. These “slicks” ure milled siakes 2
metres lung and 25 millimetres square. I was indecd
fortuiiate that when the miss spawning of Pacific
oysters from the higher trays occurred between January
26 and February 12. 197), Wivell Bros had pul our
§7,000 tarred and cured “sticks”, Consequently they had
ample “sticks” to supply the arder A quantity of scallop
shells (Pacten alba) Were set out also as wy altemative
cultch mualerial (Wivell pers, comm, ).
In April. 1971, Oyster Faroees Coffin Bay reeeived
2000 “sticks” holding (20,000 spat oysters along wath
a quantity oF scallop shells averaging 15 spat/shell, The
shell sample was included to sce if cransport Costs could
be reduced by using this allernytwve culteh ivaterial.
The flowing year another consignment of KOUQ00
spal on scallop shells was peceived on April 20, The
1U7| spat oysters coatinued to flourish and by
December 1972, the spat had reuched conmere ial
OVsleT size and Were sold (Resehke 1972*)- There was
a ready focal marke for these [Le month old oysters
and Pucific oysters from Melntyre’s Stansbury lease
were. ws being murketed. He had obtained his 171
spat from Wivell Bros, Silmouth.
In early 1972 9 tentative order lor Pacific oysler Spal
was reecived by Wivell [rom a Welstipuot (Vieneria)
hover. However, the order was cancelled later when
permission to (part inte Vietoria was refused on the
arounds of possible transmission of diseases CWivell,
pers. comm.) although na disease had appeared tn 1955
when a large number of Pacilie oysiers hud heen
Lrafsterred by the Victorian fisheries authorities from
Piltwater (Tasmania) to Mallacoota Inlet, Alter ttiree
years 74,000 oysters were still alive but there was nu
evidence of any spat fll (Thomsun 54).
Lhsuster however. struck the Tamar River oystet
imlustry in 1974 when there was virtually ny spat fall
aLany of the reehgnised settlemenr areas in Tunuary
February that year Wis beled tab Higher salinities
ut the lewses resulted in poor gonad developmen
followiny 4 prolonyed dry spell — the longeston recon
(Wivell pers. comin) The luck of freshwater Hows
if the Esk Uibutaries of the River Tumar allowed
inarine water from Bass Strfit wo penetrate further inte
the estuary ducing the long dry spell, The developity:
South Ausiratiin uyster rndusiry, dependent on spit
From Tasmunic, was balted as the 2 million spal ariher
ter April 1973 could mor be filled
256
Oyster Farmers Coltin Bay attempted to produce
their own spat for on-going operations with a small
scale industry operation but were unsuccessful because
of difficulties in providing suitable algal food for the
developing larvae.
About the same time that advice was received that
no Tasmanian spat were ayailable, the SA Department
of Fisheries received an enquiry from Pacific
Aquaculture Pty Ltd, a company holding an oyster
lease in southern Tasmania, about the feasibility of
growing Pacific oysters on long lines in South
Australian waters. It also raised the possibility of
establishing a co-operative venture hatchery in South
Australia to remove the dependency on spat derived
from a wild population in the River Tamar. The
Department arranged a meeting between representa-
Spencer
Gulf
al
, Mt Dutton Bay ;
/ . x 7 Kellidie Bay (
) Eottin Bay it Boston Island
“Port Lincoln")
to wf
\, ‘oper Bay
Vf a
1
0 100
i a
Scale (km)
A, M. OLSEN
lives of this company and Oyster Farmers Coffin Bay.
A beneficial outcome of this meeting was that Pacific
Aquaculture had been granted permission to put down
two experimental long lines seeded with oysters from
their Tasmanian lease. One long line was located off
Streaky Bay near Boston Island and the other in Proper
Bay, west of Horse Rock, Port Lincoln. The subsequent
growth of these oysters was excellent and by October,
1974. they averaged 7 cm in length. As the oysters grew
their added weight dragged longline floats underwater
until the bottom section lay on the sea floor, Starfish
then attacked the oysters and about 10% were lost.
Arrangements for additional flotation to be addded.
as needed, had broken down and the interstate company
did not proceed any further with its Jongline
experiments.
x
‘Middle Beach
\ Dry Creek
Osborne \_ Salt Field
Power Station s
Stansbury /
}
Goobowie .
,
C
Eastem Cove
Ambagen- River aa)! :
J
Kangaroo Island
Fig. 2. Locations where attempts were made to grow introduced rock oysters.
LARLY DEVELOPMENT OF PACIFIC OYSTER INDUSTRY
Fron WOX the Departiient of Fisheries had been
fovelved in Surtive and ecological studies and more
reeendy st hid carried out similar studies under
contract (y other government departments and agericies.
Ty date 973M. V. Melvin, Semor Production Officer,
Alkili and Chemical Group, ICL, approwched the
Department ty undertake for Oba contract cenlogical
survey of the large 3 ken’ primary pond of tie solar
suit evaporation coniplex at Dry Creek. We wis
exurmining the feasibility of a multipurpese use of this
pond for aquaculture of the western king. prawn,
Peraens larisuleans or any suitable fish speeies
(Resehke 1977"). This primary pond ranges frou L5
to 2.5 metres depth, has nonetidal and hypersaline
finnual mean salinity of 40,9 ppt),
An cenlopical study lasting wo months undertaken
in January-February 974. showed thet the pend could
be Suitable for uo aquaculture use (sing 1974) The
duthor advised ICT agyinst the uquaculnire of prays
or fish because of predabon by bards but ratlier te
consider the culiyation ol the Pacilic oyster C) efges,
The recommendation was based on the successtul
establishment of reproduce stocks of the mussel
Modiolas inconstans and the cockles Karel vsia spp. ies
well its tie veneral high productivity of olber biota m
(he pon. The recornmendation was accepted by the
Board of HCL Mo VY. Melvin ICL and M. G. Karu.
Department of Pisherics, were to be responsible for
A do-eperitive study of grow mg Pacific oysters in the
provary pond. Preliminary expenments were stiriect
with oysters from Oyster Farmers Coffin Bay while
uwarting delivery Ob spat ordered from ‘Tasmimia. ‘There
wus, however a liilure of the lanuary-Pebruary 1974
spel seiement bul wath aia oul-oleseasonm March
spawning, Wivell Bros were able to till the ICL order
with spat serled on scullap shell eulich, The sput ot
IY nin tiean sive were delivered and set out in the
pond on fuly 4, 1974
Astall central hole hid been punched in each cultel
shell. enabliay iio he threaded on a length of 2 nim
diameter galvanised steel wire (ren). Nine to f2 eulieh
Shells each carrying one to nine oysters were strung
op cach wire ren and separated by 1 cra lengths: ot
plastic hose, These rens were hung fromm fixed wooden
tacks al Wiree positions in (he pond, By May 1975
erowlh averaged 15 g/month, The spat hace mean net
weivht of 2.1 whe placed in the pond aod grew te
a mean wer Weight ol97.3 a in 9 touts. King (977)
reported ain acceptable market sive oF 70 2 was attained
afler being in the pond for & qnonths of withia
upproximately 1 year ot settlement.
Because of the success of the initial experiment with
srowing Pacific oysters in the primary pond, ICT in
Y Resciikh. W. W977) Here’s news to rock those Sydney
oysters. “Sunday Mail’) 6 February. 1977.
Trmrook, RO WUT Bowron the oyster lum, "Adwerteser™,
24 Kanoary, (997
a7
inid 1975 approved a subroysston lor funds tor yin on
soing program for Pacili¢ ovsters in the primary pond
at the Dry Creek sal fields. Asvorder for 30,000 seallon
shells averaging 8 sput/shell and SOQ) “sticks” averaging
40 spar/shell was placed with Wivell Bros. The
consignment shipped from Burnie, Tasmania, arrived
in Adelaide on July 25. 1975 and was platted que at
selected positions in (he pond (Melvin 1977)
By now there Was a growing awWwarcogss of a
developing South Austrahin ovster incustry, Ar
Stansbury. J. Melneyre, who had been cullivating the
mud oyser, Quine eorgas’, On bis lease Since about
46. switched some of his cultivation to Pacific oysters
in 71, In April, 1975 B, Rrankham, Wenley Beach,
Adelaide. put ont scallop culteh shells with spat from
Wivell Bros on Jonghines in his deepwater lease al
bastern Cove. Kangaroo Istand (Tilbrook (9774), As
Frankham was being advised by M. G. King,
Departiient of Pisheries. he did not repeal (he mistakes
with flotation experieneed carlier by Pacifie Oystee
Aquaculture at Pert Lincoln,
Che fighleht of 1975 regarding the cultivation of
Pacific oysters in South Australia was the success of
an application by Director of Fisheries to the
Goverment for the appointnent of an oyster biologist
experienced in oyster hatchery production, Dr. B.
O'Sullivan from Carna Research Station, Galway Bay,
Ireland was appointed to this position and arrived it
Adelaide in Marteh 1977.
ti Jinuary L976 TCT, onthe recommendation ot M,
Vo Melvin, brought Dr DL BL Quayle. a Canadian
oyster bioloyist and world authority on the cullivation
of Pacific oysters to report on the aew developments
at Dry Creek sultfields. He was impressed by the
growily of the Paeiic oysters am the unusual
environment of the primary pend of a solar salt field.
2 CNW LFONMENE Hot previously considered to be saitible
for oyster cultivation. Dr Quayle provided pravtical
advice on cultivation ad htrvesting lechiques at Dry
Crevk,
There had bee a number of approaches in the
previous two years ty the SA Government to support
the building of @ local hatchery for Pacitie oysters lu
remove Lhe depenlency on spat derived front wild
populabons ia Tasmania, There were two oveasions,
1973 and 1974, when there was no spat settlement trom
the Tasmanian January-February spawnings.
Becuuse of the anreliability of supplies of spat from
Tusmania. M, Vo Melvin had considered slternitive
sources of spat. On October 20, 1976 ICT submittect
in uppheation to import 50,000 cultehless spat of
Pacifie oyster from Seortish Sea Barns, Connell.
Argyle. Scotland througl the Minister ol Fisheries, SA,
This Scottish Compiny was supplying certified disease-
tree cultchless oyster spat to buyers it Europe and
South Afiiea. The parents of this source of spat were
OG adult Poeific oysters fron) Pendrll Sound,
JAR.
British Columbia, Canada. imported by the Ministry
al Agrieultire und Fisheries Shellfish Culiure Unie at
Conway, Wales Ui dune 964, A yearaflecdhe Minister
tad forwarded the (CL application, he was advised that
the Advisory Conmuittee ore fportind Export of Live
Fish of the Austrafan Fisheries Cruneil hod
recommended that the proposal be approved, Another
12 months elapsed before formal Wiitten-approval Was
received, There had been a rigorous examination by
Conunonwealth Quarantine and Departinent of Health
authoritics before permission. oo import culichless spat
from Seotland was jranted, OV arrival che spal were
subjected to detailed quarantine inspection provedures
and a peciod in quarantine. During the 2 year waiting
perind for upproval to import spat, the 1977 Riyer
Timur spat fall was a commercial failure with only 2
spal/scallop shell and 18 spat/*stick” being caught, This
was the third failure in four years and highlighted the
twed Jor ullerbauve sources of spat.
Tiriv in 1977. 4 depuration url wits constructed and
installed ac Dry Creek. This apparatus uscd recirculated
sel waiter, sterilised by irrachated WV light. to lush
ihe wut af oysters, Gysters of marketable sive were
removed from the pond, separated fram cach other,
cleaned externally with a jet of Sea waler and placed
inthe depuration unit where they remained far we
days This prodess was destined to prevent outbreaks
Of lool poisoning sueh as had occurred in Vieborigs 1
1975 and which had been allribuled bo infected NSW
oysturs. A. trial marketing survey conducted in 1978
indicated 4 ready ueveplance of the treated oysters.
There were no requirements for depuration treatment
ol NSW. commercial oysters unit 197879 when a
beenme mandatory following gn extensive outbreak of
(und. poisoning tt that state from local oysters.
As a consequence uf a favourable report from Dr
Quayle ulier his Australian visit and the suceessful
marketing of Pacific aysters Tram the primuiry pond,
Ihe Round of Ch approved M. VO Melvins
recommendation tty build an oyster Hatchery for Pacific
oysters de Dry Creek A drall agreement covering
joint participation by [Chand SA Fisheries Research
Branch in the design, commissioning and operation
ofan oyster hatchery at Dry Creek was received from
the Rowrd of ICT an March 20, 1978. Shortly after the
dereement Was signed on May 2, 1978, ICTadvised that
(hey hud appointed Coho Palm. a Project Engineer with
(Cl, tbe responsible for (he design and construction
of the hatehery through to commissioning, He wits to
be ussisted tircugheaur by his colleague. MV Melvin
and B. O'Sullivan, the latter represenmne the SA
Fisheries Research Branch af the now amalodriated
Deparurient of Agriculture and Fisheries, These three
men brought chemical engincering und biological
expertise fo lhe design of the hatchery. Each aspect
of the design and operation was carefully studied and
uperntonal difficulties were identified wnd where
AM, OLSEN
possible either elintinated or reduced (Anon 80).
Copper, zine, lead and their ullovs were excluded Fron
why Contech With hirvae or spat in the batchery
operalions.
The 50,000 cultchless sput from Scudand. purked
in esh bags surtounded by damp newspaper to
mainitin at humid environment, arnyed in Adelaide on
September |4, 978 im a@ polystyrene container, The
spat were immersed in chlorinated fresh water for a
few ninutes und thea finsed in Sresh water under
supervision of Quarantine-officers before being set Out
on trays in the madilied depuration uni, Water from
the primary pond was pumped dircetly through the unit
und discharged back into the pond for the quarantine
period. The polystyrene Contéiner and newspupers
Were burnt under supervision,
Beenuse of the failure of the 1977 Pacitic oysterspat
fall in Tasmania and the desire to maintain continuity
af supply of adult oysters the Tasmanian fisheries
authorities requested a permit to tinpert one rmiTHow
culichless oysler spat from Scottish Sea Farms
Pertnissian was refused on the grounds that quarantine
facilities in Tusmiania were inadequate compared with
the stringent conditions imposed at Dry Creek, Sauth
Australia.
ILtook only six months Jrom the signing of the joine
igrédment in May until the designs of the building and
equipment and the layoul were approved in Noyernber,
1978. Seven months later the building was erected
eyuipment instulled, tested und ready ter
commissioning. A fortnight liter adults were brought
into the hatchery from the primary pond thy
conditioning for spawning when required. AL the same
time the culturing. of the algal food species for feeding
the larvae and spat began. The first batch cf one million
ayster spat (limm in size) was placed in the pond on
October 2 (Melyin 1979), They were expected lo be
ready for sale a year laler after the depuration
freatment. Batches in excess of one iillium spat were
produced regularly thereafter (Olsen 1981)
In 1977 ICL Saltflelds begal an eXparsioh prograiye
of the solr evaporatiwe pond complex with the
construction of 4 arge impoundment at Middle Beach
north of ils original tease, New and more powerlul
pumps were installed 1 draw water from Gulf St
Vincent. It was planned i incurporate the primary
pond into the chain of ponds of jnéreasing salinities.
By 1980 purnping Wf seawater into the primary pond
from Chapman Creek was reduced and the new Middle
Beach pond became the primary pond of the sular-salt
evaporative pond camplex. Salinities. in the Jormer
primary pond.rose slowly to the required production
level and comsequently altered the long-established
equilibsium vavironnent there. Much of the biota
identitied m the January-February 1974 ecologicul
survey Jailed to adapl to the water quality and sulinity
regimes, Thus the food ot the oyster spat hegan te
EARLY DEVELOPMENT OF PACIFIC OYSTER INDUSTRY
decline us did the rate of growth of the oysters.
Coincidentally with this expansion of the production
of solar salt was the widespread Australian drought,
As a consequence of the drought conditions and
lowered demand for agricultural chemicals and other
products manufactured by ICT, the Board of ICI
decided on # restructuring program of its Australian
activiies and lo concentrate On production of its core
products. It decided to withdraw from and close down
its many diversified projects whether profitable or not.
Tn 1983, the South Australian solar saltfield complex
at Dry Creck was sold. The production of oyster spit
ceased and the hatchery equipment was sold.
Fortunately for the developing South Australian
oyster industry a new hatchery at Bicheno, Tasmania
had Come “on stream” in June 981 so that the industry
‘as no longer dependent on spat from the wild
population of Pacific oysters in (he River Tamar.
The success of the early aquaculture of Pacific
oysters On leases in Kellidie Bay, Kangaroo Island,
159
Stansbury and in the onginal primary pond of the solar
saltfield complex at Dry Creek stimulated the demand
lor Jeases for aquaculture of Pacific oysters. Initially
there were 98 approved leases but when fees were
introduced in 1992 this number dropped to 82. The
present active, vigorous aquaculture of Pacific oysters
in SA stems from the efforts and close co-operation
between the fishery authority, three private individuals
and one of the staff'and resources of a large industrial
manufacturing Organisation,
Acknowledgements
Tam extremely grateful to Dr 8. J. Edmonds.
Dr M. G. King and Mrs Jean Bird, Editor for their
valuable constructive comments on an earlier draft of
this manuscript. Thanks are due to Gavin Wright,
South Australian Aquatic Sciences Centre for the
preparauon. of the maps und Ms. 8. Arnold for typing
the manuscript.
References
Anos (1980) Oyster industry development underway. SAFC
(Adelaide) 4, 3-5.
CoLemMans. N, (986) A review of the introductions of Pacilie
oyster (Crassostrea giges) around the world and a
discussion ol the possible ceologival consequences of
introducing the species into Victoria, Austria, Mar. Ser
Lab,, Technical Report No. 36.
King. M. G, (1974) Ecology and aquacultare prospects of the
sea water holding ponds at Dry Creek. Report of the
Department of Fisheries, Adelaide, SA 33 pp.
(1977) Cultivation of the Pactlic oyster (Crassastrea
gigas) in-a non-lidal hypersaline pond. Aguaculiure I,
123-136.
Meveor, |. C. & Wore. P. H. (1975) Spread of Pacific oyster
worries NSW culturists. Australian Piyheriey 34, 1-7,
Mervin, M. ¥. (1977) ICL oysters au naturel. /C/ Maguzine
55, 175-177
______ (1979) Dry Creek hatchery to supply oyster farm, 767
Cirele 26, | & 6.
Osen, A. M, (1965) Fisheries, pp. 74-75, In Davies, J, T..
(EL) “Athis of Tasmania (Lands & Surveys Dept-.
Hobart, Tasmania),
(1981) The ICL oyster — goes well down under, /C?
Mavazine 59, 46-48.
___ & Priest, J, M. (1971) A history of (he South
Australian oyster industry, Report of the Department of
Fisheries. Adelaide. SA 3 pp.
SAVILLIe KENT, W. (L887) Report of Inspector of Fisheries
1887, Tasmanian Parliamentary Paper No. 31,
THomsom, SM (1952) The acclimatisation and growth of
the Pacilic oyster (Gryphaee gigas) in Australia. Anse. J.
Mat. Freshw. Res. 3. 64-73.
(1959) The naturalization of the Pacifie oyster iy
Australia, /htd, 1, 144-149.
War pace-Carrer, E, (1987) “For They Were Fishers”.
(Amphrite Publishing House, Adelaide, SA).
STUDIES ON APRUTIDES GUIDETTII
(NEMATODA: SEINURIDAE) ISOLATED FROM SOIL
AT NORTHFIELD, SOUTH AUSTRALIA
By ALAN F. BIRD* & GREGOR W. YEATEST
Summary
Bird, A. F. & Yeates, G. W. (1994) Studies on Aprutides guidettii (Nematoda:
Seinuridae) isolated from soil at Northfield, South Australia. Trans. R. Soc. S. Aust.
118(4), 261-266, 30 November, 1994.
Aprutides guidettii, originally described from Italy and subsequently from Turkey,
was isolated from a heavy clay soil from Northfield, South Australia, where it
comprises a major part of the soil nematode population. Only females have ever been
detected and these have been examined and photographed alive under the light
microscope. They have also been examined and measured under the light microscope
after fixation and, after sectioning, under the electron microscope. This is the first
record of this nematode in Australia.
Key Words: Aprutides guidettii, Seinuridae, soil nematodes, structure, taxonomy.
‘Transactions «if the Royal Society af S. Aust, (994), VIK(A), 2hb 266.
STUDIES ON APRUTIDES GUIDETTH (NEMATODA: SEINURIDAE)
ISOLATED FROM SOIL AT NORTHFIELD, SOUTH AUSTRALIA
by ALAN F BiRD* & GREGOR W. YEATES?
Summary:
Biro. A, FP o& Yeats, G, W. (1994) Studies on Apratides eiideril (Nemiivdas Seinuridive) wold (rom sor
ut Narthfield, South Australia. Trans, Ro Spe 8. Asi UIRC4), 261-266, 30 Noveiiber 1994.
Apratidey guidetti, originally desctibed trom tly and subsequently from Turkey, was isetated froma heavy
clay soil from Northfield, South Australi. where it Comprises a ajor partot the soi] nenaode population,
Only femiles have ever heen detected and these huye been examined and pholowriphyd ative uper the light
microscope, They have also been examined and measured under the leht micnseope after fixation id. aller
sectioning, under the electron microscope. This ts the first record of this nematode im Australia
Kiev WorRps: Apruttdeys cufdetiii, Seinuridae, soil nematodes, structure, laxanciny,
Introduction
During an examination of the number and diversity
of nematodes from three different soil types in South
Australia in the spring of 1992 (Yeates & Bird 1994)
it was observed that u clavale-tailed aphelenchordl wits
the most common nematode in the black earth
(Vertisol) of Northfield making up 19% of the total
nematode population, This nematude was not observed
in the red brown earths (Alfisol) of Kapunda and
represented only 0.4% of the solonised brown carth
(Enusol) of Avot and then only from land occupied
hy native shrubs, Because of its relatively lorge
quijbers in the Northfield site we have attempted (6
identify it using light and electron microscopes,
In this pauper we show that the nematade from
Northtield closely resembles the aphelenehoid
Aprugides yuidertii first deseribed by Scognamigho
(1974) in Ttaly and Tater by Saltukaglu er af, (976) in
soil sumples taken front grassland in the Wartal
Maltepe, Camlica and Gostepe areas around Istanbul
in Turkey.
We are confident thal the clavate-tailed nematode
from Northfield is Apraides waidernt and have
provided further information to establish its identity
Materials and Methods
Nematode
A nematode subsequently identified as ¢lpridides
guidedtii was isolated by means of 4 misting Epparatus
from soil collected at Northlield, South Australia, from
Which this nématode hud previously been extracted
(Yeats & Bird, 1994).
* CSIRO Division of Soils. Prryate Bag 2, Glea Osinond,
South Australia S064, Australia.
+ Landcare Research, Private Bag HOS2. Palmerston North.
New Zeulund-
Ligh) microscapy
Nematodes were examined under bright (eld, phase
contrast and differential interference contrast oplics
using a Vanox Olympus AHBT research microscope,
Living nematodes were examined im distilled water
under a coverslip sealed at ils edges with nail varnish
and photographed using Mlord Delt 400° fil.
Specimens were fixed by adding un equal volume wl
builing double strength PA 42) (20 ml 40%
formaldehyde und 2 mil glacial acetic ueid in 78 ml
distilled Water) to a shaken suspension of the neniatodes
in distilled water, These specimens were processed to
pure glycerol by Seinhorst’s (1959) method ane
mounicd inunhydraus glycerol on sides sealed to a
coverslip by molten paraffin ay deseribed by De
Maesencer & D’'Herdle (1963), Fixed nematodes ip
glycerol were photographed with IHord Pan & film,
Fleeirun mloruscopy
Specivens were picked out with a mounted eyelash
and fixed in cold (49°C) phosphate buffered (pH 7.3.
01M) 4% paraformuldehyde, washed, fixed and stained
in 2% aqueous Osmium tetroxide. washed. dehydrated
in-ethanol and embedded and polymerized at 60°C in
TK3 resin as. previously described in more detail (Bird
& Ryder 1993),
Seetions were cul witha diamond knee ina Rewhert-
Juny Ultracut B altranticrotome. mounted on uncoated
gnds and stumed with uqueous 2% uranyl avenue
followed, afer washing. by 0.3% lead ertrate in O.1M
sodium hydroxide as. described previously (Burd &
Ryder 1993), These sections were exumined. und
phologriphed in a Philips RM 400° transmission
electron microscope (TEM),
Feeding experiments
Attempts to determine the feeding hubtits of 4.
euidertii were tade using two possible loud
sources, derobeloides fants growe on Preudoamenay
corrugake on mall extrack agar (Bink & Ryder
1993) and Rhlzgenmia solani grown on a diluted
Czupek-Dox + yeast ugar (Warcup 1955),
262 A. F BIRD & G. W. YEATES
APRUTIDES GUIDETTH FROM NORTHFIELD, SOUTH AUSTRALIA ahd
Results
Description
Aprulides guidetiii Seognamighio, 1974
IGS 1-8
Adult feniales: Small nematodes, 0.5 mon in length,
that exhibit normal coiling and uncoiling in distilled
water (Figs 1, 2), When fixed with gentle heat they
shrink (abou! 6% in length and 20% in width) and
become slightly curved to C-shaped. Under bright
field, photographs of the same living specimen which
became coiled after a few seconds (Fig. 2) show the
delicale stylet with small thickenings at its base und
the lip region bearing a distinct cap characteristic for
this species. ‘The latter is seen more clearly in fixed
material in glycerol and under interference contrast
optics at higher resolution (Fig. 3). This photograph
also illustrates the prominent metacorpus with the
intestinal junetion and the opening of the secretory~
excretory pore immediately behind. ‘This opening
oecurs 57-64 um from the anterior end (Table 1). Also
clearly visible under interference contrast optics are
the vulva and vagina (Fig. 4), Transverse sections cut
through the mid-intestinal region and observed with
the TEM show that the cuticle is 0.2 pm thiek,
becoming 0.5-0,6 yam thick at the lateral alae (Figs 5,
6), The literal alae (lateral fields) are distinct, although
not pronounced and are 2 ~m wide with three distine|
striations, incisures or furrows (lines). The cuticular
annulutions are fine and spaced about | jum apart, The
pharyngeal gland lobe is obscure and extends about
50 jam posterior to the metacorpus. The intestine ts
2.5 ju in width (Fig. 7) und in the specimens
exumined showed titde evidence of tbod material other
than a number of multilamellar bodies that occupied
ihe Jumen of the intestine (Fig. 8). The lamellue of
these structures ure approximately 4 Ti apart,
The rectum is about (7 ym in lengih and so ts
ipproximately twice the width of Ure mentatade at the
anus, The tail is clavate, tapering und then widening
inly the characteristic club shape (Fig. |). Tt ranges
from 39-47 pam in length and is 4-5 times the width
of the nematode at its anus. The vulva protrudes slightly
and the vagina is about 7 jam an length and usually
clearly visible at tight angles or slightly oblique to the
body axis (Fig. 4), The gonad is single and Gutstretehed
and consists of ovary with a single row of oocytes,
oviduel, spermatheca und uterus all having a total
length of 83-103 jan. The post-uterine se is collapsed
and up to 30 jam in length. Males were not found.
In an ultempt to resolve Whether this nematode is
ua predator ora fungal feeder small numbers were added
10 cultures of either Aerabelaides nuns or the fungus
Rhisaeromin solani. However, no prowth oF
development of 4, wididerli took place in either of these
cultures und ng feeding wis observed,
Discussion
There appears to be no doubt that the nematode we
have isolated Prom the heavy clay soll at Northfield
is Apruritles guider, Scognamiglio ef ul, (1970) used
the generic name of 4pruddes after the Latin name
Aprutian for the Abruzzi region in Italy where this
venus was first found, The specific name was aller
Beniamino Guidetti, a professor of neurosurgery at the
University of Rome. Measurements of the nematode
found in South Australia are comparable to similar
TABLE Lo Measurements of adult females of Aprutides guidetint jan).
n Range Mean +8D (SE)
1, 20 395-518 454 434.4 77
Maximum body: with 20 1-17 16.3 (5 {|
Pharyax length 20 54-62 S75 2 5
Tail length 20 39-47 44.4 2.4 Ot
Body width at anus 20 9-10 i) A (0,1
Stylet length 17 }2-|4 133 7 (}>
Anterior end to secretory—
excretory pore ty) 57-64 60,3 i4 7
Anterior end to vulva ath 291.339 204 238 5.3
De Man's indices V 20 63-64 66 18 04
De Man's indices a 20 25-31 28 20 O4
De Man's indices. b 20 6.9-9 4 74 07 0.2
De Mars indices ¢ 20 9,411.3 10,5 6 Ou
De Man's indices ¢ 20 3.93.2 4.6 H4 at
Figs | and 2, Two photographs of a living specimen of adult female ot dprufides xuidentii taken within a few seconds
ofeach other, Bright field optics showin (he anus (a), Hips (1), fetacerpus (m). stylet (s) and vulva ty), Seale bar = 50 pr,
164 A. F BIRD & G. Wo YEATES
-—
APRUTIDES GUIDETTH FROM NORTHEIELD, SOUTH AUSTRALTA 265
measurements made in Italy by Seognamiglio (1974)
und in Turkey by Saltukoylu ev e/. (1976) (Table 2).
Ti can be readily recognised under the light microscupe
by ils small size (only 0.5 min in length), pronounced
clavate wil und distinctive cap on the lip region,
The high concentration of this nematode in the heuvy
vlay soil at Northfield (Yeates & Bird 1994), where
it made up almost One fitth of the (otal nematode
population, suggests that it is particularly adapted to
this type of soil. It is tempting to speculate thal 4.
enidettii might have arrived in Australia on the wheels
of the Viekers-Vimy bomber that made the first fight
from London to Australia as this aircraft landed en
route at Pisa, Rome and Taranto in Ttaly before finally
landing al Northhelu, This nematode has the capacity
lo withstand considerable environmental fuetuations
in the soil during storage and could have survived in
an unhydrobiotie state on the undercarriage of the
pline, However, it is unlikely that it was introduced
ly Northfield in this fashion because the plane lended
many times before landing at Northfield. A similar
clavate-tailed nematode has also been detected, albeil
in very small numbers (one in 250 counted). tn the
sandy loam at Avon on land occupied by native shrubs
(Yeates & Bird 1904), However, this clavate-tailed form
from Avon has not been detected in re-examination of
the original fixed material and sa no. precise
measurements (6 establish its identity have been
possible. [itis 4. guider’, (his would indicate that
itis capable of surviving in a range of soils and that
it has probably existed in South Australia for some
ume,
Many species of the family Seinuridae, inta which
the genus Aprutides has been placed, are considered
to he predacious (Hunt 1993). This has been abserved
especially inthe genus Sedmura which has been shown
to prey on bacterial feeders such us members of the
genus Acrobeloides (Hunt 1993), However, the genera
Apniides, Setiura, Aphelenchoides and others are
included in the fumily Aphelenchoididue by Andriissy
(1976) and thus this author considers that dprundes and
Aphelenchoides have characteristics in common. The
TABLE 2, Conipurisen of measurements of different populations of females of Aprutides guidertii,
Pescara — aly
(Scognamigho 1974)
Istanbul — ‘Turkey
(Salttukoglu ev a/. 1976)
Northfield — South Australia
(Present stucly)
n=I15 n=9 n= 20
Parts Measured lange mean range mean range mean
Length (jum) 379.469 423 AWO-S10 - 395-518 454
Pharynx (jum) - — 47-58 — 54-62 458
Stylet quan) 11.9-14.6 13 12.5-13 5 ~ |214 13
Tail (urn) 32.3 37.1 as 34 42 - A9-47 43
Head to sec-exc.
pore (wm) 31 -o4 57-64 Oo)
De Man's indices Vo 66, 170.5 68 - - 3-69 Oy
De Man's indices a 28-3 a 2432 2531 28
De Man's indices b 5.4-6.8 6 7.1-4_6 - 69-45 4 8
De Man’s indices e¢ L1L.0-14.5 119 O.0-12.2 - 94-113 10.5
De Man’s indices © - ~ 4.24.7 _ 349-52 +6
Pig. 3. Fixed specimens of female dpridides guidetti in alycetol, Interference contrast opties. Antero (egion showing
lips () bearing a vharacteristic cup, the prominent meticorpus (mn) and (he secretory-excretory duct and pore (se), Seale
bar ~ LO jam,
Fig, +. As lor Fig 3 but showing the vagina (va) amd vulva (¥) Seale bur = 10 yan.
Bip 5 Tringaission electron microvraph of a section through Apridides vuidetii, Showing the cuticle (c), Seale bar
0.2 jim.
Mig. 6. As har Pig. 5 but showing a lateral ala (la) dnd the three distiner indisures (black arrows). Scale bar = O.A pn.
Fig, 7, As tor Fin, 5 bul showing w section through the intestine (i) und the intestinal villi (iv), Seale bar = 0.5 am
Fin, §, As for Mig, 3 bur showing part obo section throvgh the intestine sawing tarestinal villi Gy) and multiimedllar body
(mlb in the intestinal lumen. Seale har = (2 pam,
266 Ab
teeding and growth of Aphelenchoides Aylurgr om
various fungi have been observed and its ability lo prow
on these different species compared (Bird eral 1989).
This would sugpest that Apraddes euiderti) may also
be @ fingal feeder. In field stuclies i wits assumed (har
the chivate aphelenchoid, the name given lo A) gauderu
hefore it was identified (Yeates & Bird 1994), was a
fungal lveder, We have not, however, been able bo
resolve Whether this nematode is a predator or a fungal
feeder TL is possible that storage of the nematode in
soil al low teniperatirce. the misting procedire tsed
to separate the nematodes from the soil and their
subsequent handling and duration tm water in Petri
Wishes prior to Ganster to the food source, may have
inhibited their desire or ubility to feed. They appeared
wenve but TEM photographs of ‘transverse seetions
through the intestine showed that the lumen was wniastly
oceluded and the only contents were a number of
mulliamellaied hadies (Fig. 8). 1 was not clear if these
were the breakduwn produets of either nematode prey
or fungi, Because the divestive and reproductive
systems of 4. glider’ are Similar to those af the genus
Aphelenchoides (Sallakoglu ef al. 1976) it seems more
likely that they are fungal feeders rather (han predatory
nematudes and the assumption that they feed ina
Similar fashion to 4. Aylare? seems reasonable until
evidenee to the contrary is forthcoming.
The improved resolution of the TEM hus revealed
thatthe lateral alae of this nematode, being, 2 jim, are
narrower than the previously-reported 2.5-4,5 aim
(Saltukoght er al 1976) and have three distinet incisures
fulier than two as reported by these workers, In
addition, (he cuticle ys only 0,2 am thick eather (han
BIRD & 4G. W YEATES
U7 jar although it thickens to .0,5-0.6 jam in the region
of the literal alae. The original measurements of these
structures by Saltukoglu er a/. (1976) were made with
the light microscope and are close to the 0.25 yin hint
of resolunion of this instrument so that precise
measurements al these extremes of resolution would
have been difficult.
Tt has been shown that shrinkage in seetions of
embedded nematodes is about 10% greater than that
of unembedded fixed material (Stynes & Bird 1980),
However, this discrepaney would only account for a
small fraction of the differences that we have reeurded
in the thickness of the cuticle and. lateral alae.
Shrinkage similar to that which we have observed die
to fixation in females of A. vudderrit has been recorded
in both larvae and males of Aneuina agrosris (Stynes
& Bird 1980) where shrinkage in width was 17%
compared to 20% ind, guider, However, similar
heal-fixation and processing (through to glycerol ot
Anguina agrastis laryae led to greater shrinkage in
length than ind. guider lemales (13% compared to
6%), Although it is acknowledged that fixation and
processing lead W shrinkage, very few measurements
are made on living nematodes because, until recently,
it has nol been possible to use a suitably last fine-grain
film lo obtain them in sharp focus, Tis still necessary
to Use conventional hot fixation, dehydration and
mountiog in anhydrous glycerol to enable accurate
vomparisons to be made because this technique his
been used to measure the majority of free living and
plant parasitic nematodes whose dimensions have beer
recorded ,
References
Aspwassy, L, U976) “Byolition as a bass for the
sysiematizatian of nematodes: (Pitman Publishing,
Limden),
Biren, A Ck. Bik, JL, Forrusne, R.& Monn, R. ULYsY)
Observanons of Aphaleicholdes Avieg Mussey, i974
fecding On Fong pathogens oF wheat i Australia, Revie
Nernany, 12, 2734,
— & Ryprn, M. M, (1993) Feeding of (he nemitode
Aoribeloides. sanus on bacteria A. Nemutal, 28, 493-490,
Db MAgsexpit. J & DHeRDE. 1 (1963) Methoues utilises
peur etude des anguillules: libres di sol, Revue de
(Avricditane. Brixellés 16, 441-447,
Hest, D £ (998) “Aphelenchida, Longidomdne and
Trichodoridue, Their systematics aod bonoques” (CAB
International, Wallingford, CK.)
Sanmpnoubu, Mo &B,, Geka, EB, d& Coomanms, A (1976)
Some Tylenchida trom the tstanbul-area (Turkey). Nemearot,
medit, & 189-153)
ScoGnamiahto, A, (974) Apruridys guider uu. sp.
(Nematoda: Apheleavhowleae). Ball Lab, Bae que Pilippne
Silvestri. WW. 17-2,
_, TALAMI, M. & Stacob, 1 J. (970) Aprundes
marticol (Nematoda: Aphelenchoididae) no. gy... sp. Abel.
28, 4b
SpemHorsy, J. W. (1959) A rapid method (or Ure transter at
nemalodes from fixative to anhydrous plyeerin
Noernatilueica 4, 67-69,
Syvnes, BA & Bir, Ab (YAO) Effects of methods of
killing. fixing and mounting On measurements Ol dapieine
agnosis, (bid. 26, 467-474.
Woareur. JH. (955) fsolation of fungi trom hyphae present
in soll Manire 178, 953-954,
Yeares, GL Wo & Rirn, ALF (1994) Sonie observations on
the fifluence of -agrivullural practices: on the neniatode
faunwe of some South Australian soils. Fudan, appl,
Nematol UP, (43-145,
THE HYLID FROG LITORIA ALBOGUTTATA (GUNTHER)
IN THE NORTHERN TERRITORY
BRIEF COMMUNICATION
Summary
Knowledge of the diversity and biology of the anuran fauna of the Northern Territory
has increased substantially over the past 30 years. In 1961 15 of the currently
recognised species had been reported’ but by 1986 the number had tripled to 45”,
During taxonomic studies in the late 1960s** that contributed to the resolution of the
N.T. anuran fauna, a specimen from Alexandria Station identified as Cyclorana
alboguttata in 1935” was re-examined because it represented the only record of the
species in the Northern Territory, and its specific identity has been the subject of
dispute*’. The specimen was demonstrated to be a misidentified topotypic individual
of C. cultripes Parker, whilst C. alboguttata was shown to possess intercalary
cartilages, causing it to be referred to Litoria Tschudi*. The geographic distribution of
L. alboguttata was plotted from all known museum voucher specimens and
demonstrated to be confined to Queensland and northern New South Wales*.
Transactions of the Reval Seciety ef SL Aust. (199d), UBId), 267-268.
BRIEF COMMUNICATION
THE HYLID FROG LITORIA ALBOGUTTATA (GUNTHER) IN THE
NORTHERN TERRITORY
Knowledge of the diversity und biology of the anuran tiunit
of the Northern Territory has increased substantially over the
past 30 years. In 1961 15 of the currently recognised species
had been reported! bul by 1986 the number had (ripled to
45,
During taxonoinie studies in the Iate 1960%°7 that
contributed to the resolution of the N.T, anuran fauna, a
specimen troat Alexandria Station identified as Cyclarana
albouuttata in 1935" was re-examined because it represented
the only record of the species in the Northern Territory, and
its specific identity had been (he subject of dispute 7’. The
specimen Was demonstrated to be a misidentified topotypie
individual ef C. ewliripes Parker, whilst C. alboyutiaia wax
shown lo possess inlercalary cartilages, causing it to be
referred to Liroria Tsehudi®, The geographic distribution of
1. ulhogutiaia was plotted trom all knowe muscu voucher
specimens and demonstrated (o be confined to Queensland
amd northern New South Wales®.
In recent years L. alboguttiata has been vollected on a
Humber of occasions i the Northern Terttlory at sites heat
the mouth of the McArthur River in the vicinity of Borrolooka.
leading to the inclusion ofa portion of the western boundary
of the N.T. ina revised distribution map’. ff hus also been
reported approximately 20 km northeast of Devil's Marbles.
considerably further south than the other records",
Because the places of lodgement of the specimens on which
these locality records are based have nut heen reported. Wwe
provide this information bere The spectinens examined are
deposited in the Australian Museum, Sydney, (AM), the
National Wildlife Colleetion, CSIRO, Canberra (NWC), the
Northern Territory Museum & Art Gallery, Darwin (NTM)
and the South Australian Museum. Adelaide (SAM), Methods
OF Measurement follow Tyler!
All of the specimens of 4, ad/bowuiraie that were examined
were taken ut two sites. The first is an aren of seasonally
inundated grussland extending for 1 kin flanking the
Finn. Atari alhaenmitia collected on Carpentaria Highway, 9 kim west of Borroloola, NT. (Photos M Davies!
268
Curpentaria Highway from its termination | km south of the
Borroloola township, and 2 kim east of the causeway crossing
of the McArthur River. (Floodwaters flow north into the Rocky
Creek catchment, and then east to discharge into the McArthur
River - a distance of 8 km). The floodplains extend for only
10 km west, Where the Curpentaria Highway rises to pass to
the north of the Bukalara Range, From various points along
this section 20 specimens haye been collected: AM R53214,
53650-52, 53654-62, 54490, 56923, NWC A536, 538; NTM
R17749; SAM R33907. $0749, One specimen collected in the
Borroloola area by W. J. Freeland (pers. comm.) has been
seen by us but its whereabouts is unknown,
The second site is the Minngarda Waterhole on Batten
Creek. This waterway arises west of the Carpentana Highway
and the Bukalara Range approximately 50 km from
Borroloola. Minngarda Waterhole is near ils termination
upproximately 30 kin northeast of Borroloola at the edge of
the saline coastal flats, Specimens from that site are AM
R453532 and NWC AS&6-90,
The specimens reported from near the Devil's Marbles"
(NTM 30446-7) have been examined by one of us (M.ST.)
ind reidentified as juvenile C australis.
‘Moore, J. A. (961) Bull, Amer Mus. Nut. Hist. 121,
149-386,
Tyler, M. J. & Davies, M. (1986) “Frogs of the Northern
Territory’ (Conservation Commission of the Northern
Territory. Alice Springs),
“Tyler, My J. (1968) Rec. S. Aust. Mus. 15(4), 711727,
*byler, M. J. (1969) Thid. 16(1). 1-11.
“Loveridge, A. (1935) Bull. Mus. comp. Zool, Harv, 78,
1-60,
"Loveridge, A. (1949) Trans. R. Soc. 8. Aust, 72, 208-215,
In the key to frogs of the northeast of the N.T.*, Litaria
albogutiata keys oul to Cyclorana australis. Both are fossorial
species with size ranges that overlap: L. alboxurtane males
54-67 mm snout to vent length, females 61-83 mm, ©.
australixy males 71-79 mm, females 71-102. Each species has
an enlarged inner metatarsal tubercle employed for burrowing,
but L. alboguttata is distinguished externally by its
exceptionally rugose skin with short, broad, disrupted plicae
on the dorso-lateral surfaces (Fig. 1). In contrast the skin of
C. australis is smooth apart from narrow continuous dorso-
lateral folds, In addition L. alboguttata possesses a very
narrow vertebral stripe not exhibited by C. australis.
We are indebted to the Conservation Commission of the
Northern Territory for permits enabling us tq collect
specimens in the vicinity of Borroloola and to Bill Freeland
for advice about his collections there. Out field studies were
funded by the Australian Research Council and by Mount [sit
Mines Ltd. We thank David Hughes of M.[.M_ for logistic
support and David Williams (University of Adelaide) for help
in the field. For the loan of specimens we thank Ross Sadlier
(Australian Museum) and John Wombey (CSIRO). The
inanuscript was typed by Catherine Pascoe.
Parker, H. W. (1940) Novit. Zool. 42, 1-106,
‘Tyler, M. J. (974) Proc. R. Soe. Qld 8S, 27-32.
"Cogger, H. G. (1975) “Reptiles and amphibians of
Australia’ (Reed, Sydney).
“Fleming, M., Gillam, M., Hewitt, M. & Latz, P. (1983)
A preliminary biological survey of portions of the McLaren
Creek Pastoral Lease. (Wildlife Division, Arid Zone Research
Institute, Alice Springs). (Unpublished),
"Tyler, M. J. (1968) Zool, Verhand. (96), 1-203.
MICHAEL J. TYLER, MARGARET DAVIES, Department of Zoology, University of Adelaide, S. Aust. 5005
and GRAEME F. WATSON, Department of Zoology. University of Melbourne, Parkville, Vie 3052.
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